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Guidelines for Managing Lesser -Chicken Populations and Their Habitats Author(s): Christian A. Hagen, Brent E. Jamison, Kenneth M. Giesen, Terry Z. Riley Reviewed work(s): Source: Wildlife Society Bulletin, Vol. 32, No. 1 (Spring, 2004), pp. 69-82 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3784543 . Accessed: 24/04/2012 17:07

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http://www.jstor.org SPECIALCOVERAGE 69

...y Christian A. Hagen, Brent E.Jamison, Kenneth M. Giesen, and Terry Z. Riley

Abstract Lesser prairie-chicken ( paidicinctus) populations have declined by >90% since the 1800s. These declines have concerned both biologists and private conservation groups and led to a petition to list the lesser prairie-chicken as threat- ened under the Endangered Act. Most of the land in the current range of the lesser prairie-chicken is privately owned, and declines have been primarily attributed to anthropogenic factors. Conversion of native to cropland and excessive have been implicated as leading causes in the species' decline. Periodic drought probably has exacerbated these problems. Little research on habitat requirements was conducted prior to 1970. Despite recent advances in the knowledge of lesser prairie-chicken ecology, no comprehensive guidelines for management of the species have been published. In these guide- lines, we provide a synopsis of our current knowledge of lesser prairie-chicken habitat requirements and suggest management strategies to monitor, maintain, and enhance lesser prairie-chicken populations. Key Words filifolia, guidelines, lesser prairie-chicken, management zone, mixed-grass prairie, , sand sagebrush, shinnery oak, Tympanuchus pal- lidicinctus

he distribution of lesser prairie-chicken (Tympanuchus occur in , , , , and pallidicinctus) populations and their occupied habitats (Figure 1). Concern for this species has escalated have declined by >90% since the beginning of the twenti- in the twenty-first century as lesser prairie-chicken eth century (Crawford and Bolen 1976a, Taylor and (LPCH) population indices still suggest long-term Guthery 1980b, Giesen 1998). Populations currently declines (Bailey and Williams 2000, Giesen 2000, Horton

Address for Christian A. Hagen: Division of Biology, Kansas State University, Manhattan, KS 66502, USA; present address: Oregon Department of Fish and Wildlife, 237 5. Hines Blvd., RO. Box 8, Hines, OR 97738, USA; e-mail: [email protected]. Address for Brent E. Jamison: United States Geological Survey, Northern PrairieWildlife Research Center, 8711 37th St. SE, Jamestown, ND 58401-7317, USA; present address: Yakama Nation, Wildlife Resource Management, P.O. Box 151, Toppenish, WA 98948, USA. Address for Kenneth M. Giesen: Wildlife Research Center, Colorado Division of Wildlife, 317 W Prospect Rd., Fort Collins, CO 80526, USA; present address: 4200 North Shields, Fort Collins, CO, 80524, USA. Address for TerryZ. Riley: Wildlife Management Institute, 1 146 19th St. NW, Suite 700, Washington, DC 20036, USA.

Wildlife Society Bulletin 2004, 32(1):69-82 Peer refereed 70 Wildlife Society Bulletin 2004, 32(1):69-82

2000, Jensen et al. 2000, Sullivan et al. 2000). Because Kansas. This semi-arid region receives <60 cm of annual of recent declines, sport-hunting seasons were closed in precipitation and sustains periodic drought that can affect New Mexico (1995) and Oklahoma (1998). Currently, short-term population trends (Brown 1978, Giesen 2000). LPCHs are state-listed as threatened by Colorado and Numerous anthropogenic factors have been implicated considered a "warrantedbut precluded" threatened in the decline of LPCHs. Conversion of native species by the United States Fish and Wildlife Service communities to cropland, fire suppression leading to (USFWS) (Giesen 1998). Concern for LPCHs has led to juniper (Juniperus spp.) encroachment, excessive grazing research in all 5 states where the species occurs to better by livestock, suburban developments, and fossil-fuel understand population trends and habitat requirements. drilling and exploring are suspected to be the primary Mixed-grass prairie communities that historically sup- factors (Giesen 1998, Woodward et al. 2001). These fac- ported LPCHs were thought to be comprised of regions tors, coupled with periodic drought, may have driven patchily dominated by sand sagebrush (Artemisia long-term population lows to critical levels. Continued filifolia) and sand shinnery oak (Quercus havardii). human encroachment and overgrazing likely will exacer- bate population declines. Although the rate of habitatloss has slowed over the Management guidelines have been past 20 years, LPCH populationtrends continue to published for sage (Centrocercusspp.) and Columbian decline, suggesting that it is the quality (i.e., human- sharp-tailed grouse (T. phasianellus induced or drought)of the habitatthat may t>e limiting columbianus), species of conservation the of these concern (Braun et al. 1977, Giesen recovery populations. and Connelly 1993, Connelly et al. 2000), but not for LPCHs. We pro- However, northern and eastern fringe areas of their range vide a synopsis of the current state of knowledge and primarily were mixed-grass interspersed with provide guidelines to monitor, maintain, and enhance including sand sagebrush, plum (Prunus spp.), sumac LPCH populations and the habitats upon which they (Rhus spp.), and yucca (Yucca spp.) (Baker 1953, depend. Lesser prairie-chickens occupy 3 distinct vege- Copelin 1963, Horak 1985). Currently,LPCHs occupy tative communities; however, data are available primarily sand sagebrush and mixed-grass communities in Kansas for populations inhabiting sand shinnery oak and sand (south of the Arkansas River) and Colorado, sand shin- sagebrush communities, and our guidelines reflect the nery oak and mixed-grass communities in New Mexico available information. and southwest Texas panhandle, an interspersion of these types in Oklahoma and northeast Texas panhandle, and mixed-grass prairie north of the Arkansas River in Population biology Population monitoring r"^, Lesser prairie-chicken populations are surveyed annually during the spring VJ\% ^> when males congregate at lek sites. AJr X (J Surveys of displaying males have been '/\f \t used to provide long-term monitoring of population trends (Giesen 1998). Survey methods and effort vary by state. Harvest statistics from check sta- tions indices ••f ••f ~ provided production (age ratios) and sex ratios in New Mexico ?.'-fr ~ (1958-1968, 1988-1989, 1995) (New /~2 ~ Mexico Department of Game and Fish, unpublished data) and Texas (1967-1988) (Texas Parks and Wildlife X\~ ~ Department, unpublished data). Kansas w~J0 ~ and Texas currently monitor harvest by Figure 1. Current (2003) distribution of the lesser prairie-chicken sha ded in gray(After using surveys mailed to random sam- Jamison et al. 2002a). ples of hunters. These data are used to

I Lesser prairie-chicken management * Hagen et al. 71

estimatethe numberof birdsharvested, but demographic Attwater'sprairie-chicken (T. c. attwateri)(Peterson et al. dataare not collected. 1998). Survival Seasonal movementsand home range Survivalestimates for LPCHsare basedon short-term Lesserprairie-chicken populations are considerednon- studiesof selectedpopulations using radiotelemetryand migratory,and seasonalranges and movementsof indi- bandingdata. Annualsurvival of adultLPCHs is similar vidualstypically are restrictedto suitablehabitats within to thatof otherprairie grouse species (Hagen2003). an individual'sannual range (Giesen 1998). However, Campbell(1972) estimatedannual survival of banded movementsof LPCHsin southwestKansas resembled males in New Mexico to be 35%,and Merchant(1982) partialmigrations; approximately 5% of radiomarked estimatedsurvival of radiomarkedfemales for a 5-month individualsmoved 30-50 km fromtheir capture areas period(April-August) at 59%,extrapolated to 12 months duringnesting or postnestingand returnedto the capture =31%. In Kansas,Jamison (2000) estimatedannual sur- areaduring winter (Hagen 2003). Prenesting(spring) vival of radiomarkedmales to be 57%. Male and female rangestended to be largerfor females (355-596 ha) than 6-monthsurvival curves (April-September) that included springranges for males (120-211 ha) in Coloradoand breedingand nestingwere 74%, and this estimateextrap- Kansas(Giesen 1998, Jamison2000, Walker2000). In olated to 12 months=55%. Hagen(2003) documented Coloradothe combinedannual ranges (95% ellipses) of age-specificannual survival of live recapturedata for all ,males and females,from a lek were bandedmales from 1998-2002 in Kansas,and rankings 2,450-5,130 ha (24.5-51.3 km2)(Giesen 1998). In were: yearling=62%,adult=49%, and older adults=35%. Kansassummer ranges (95% fixed-kernel) generally Annualsurvival (April-April) of radiomarkedfemales were smallerthan spring ranges for both males (120 ha) was estimatedat 53 and 38%for yearlingsand adults, and females (150 ha) (Jamison2000). Lesserprairie- respectively(Hagen 2003). Femalesurvival rates were chickensin New Mexico and Oklahomamoved consider- lowest in the monthsof May andJune, but second-lowest able distancesin yearsof drought,but prenesting,nest- ratesoccurred during winter months. Twenty-nineper- ing, and postnestingranges of femaleshad patternssimi- cent of all female mortalitywas directlyrelated to nest- lar to those of Coloradoand Kansas,although they were ing activities(Hagen 2003). slightly smaller(Copelin 1963, Riley et al. 1994). Winter rangesof male LPCHsin Kansaswere 300-700 ha Reproduction (Jamison2000) and 331-1,945 ha in Texas(Taylor and Most LPCHfemales attemptto nest, with 81-100% of Guthery1980a). These enlargedranges have been attrib- radiomarkedbirds initiating at least one nest (Riley et al. uted to foragingin grainfields, which may be some dis- 1992, Giesen 1994, Pitman2003). Renestingattempts tancefrom winterloafing androosting sites in native may varywith yearlyhabitat conditions and conditionof rangeland(Taylor and Guthery1980a). females,but 10-30% of females lose theirinitial clutches 1994, Pitman success (Giesen 2003). Nesting (percent Habitat clutcheshatching >1 egg) averaged28% for 10 telemetry requirements studies(Giesen 1998), and hatchability(number of eggs Food habits hatchedper successfulnest) averaged>90% (Copelin Insects,seeds, ,buds, and cultivatedgrains domi- 1963, Merchant1982, Giesen 1998, Pitman2003). natethe annualdiet of LPCHsthroughout the range Clutchsize is similarrangewide (Giesen 1998), with an (Schwilling1955, Jones 1963a,Donaldson 1969, Riley et overallmean of 11.3 (SE=0.51) and 8 (SE=0.82) eggs al. 1993). Greenleafy forageand insectswere the domi- for first and secondclutches, respectively (Giesen 1998, nantcomponents of LPCHdiet fromNovember to April Pitman2003). Chick survivalis highly variablebut aver- andMay to October,respectively (Jones 1963a). Adult ages 39%rangewide (Davison 1940, Schwilling 1955, LPCHsrelied heavily on sites of mixed-grassspecies of Copelin 1963, Merchant1982, Jamison2000, Pitman 25-80 cm in height,and seeds of 6-weeksfescue (Vulpia 2003). Survivalof chicks (54%)from fledgingto first octoflora), western ragweed (Ambrosiapsilostachya), and breeding(Aug to Apr) was similarto thatof adultsdur- fragrantsumac (Rhus aromatica) were importantfood ing the same periodin Kansas(Pitman 2003). Annual items (Jones 1963a). Wheat,western ragweed, and blue variationin chick survivaland nest success may have the grama( gracilis) were commonat foragingsites largestimpact on LPCHpopulation growth rates (Hagen (Donaldson1969) throughoutthe yearin Oklahoma. 2003), as had been documentedfor greaterprairie-chick- In New Mexico LPCHsforaged nearly exclusively in ens (T cupidopinnatus) (Wisdom and Mills 1997) and shinneryoak-tallgrass communities during autumn and 72 Wildlife Society Bulletin 2004, 32(1):69-82

A lesser prairie-chicken nest in a predominantly grass pasture. A Kansas pasture with a sand sagebrush density of 2,588 per ha.

winter (Riley et al. 1993). Fall diets were comprised of Breeding habitats seeds (43%), vegetative material (39%), and insects Lekking. Leks (breeding and display sites) generally (15%); winter diets were acorns (69%) and wild buck- are in open areas of short grasses surroundedby sand wheat (Eriogonum annuum; 14%). Vegetation at forag- sagebrush or sand shinnery oak grassland (Giesen 1998). ing sites was dominated by grass and shinnery oak, with Other such sites include, but are not limited to, aban- shinnery oak more prevalent at winter sites than at doned oil-drilling sites (oil pads) with little or no vegeta- autumn sites (Riley et al. 1993). Autumn foraging sites tion, unimproved roads with little traffic, areas treated contained more grass and fewer shrubs than did winter with shrub-specific herbicide, recently burned areas, sites, reflecting the potential importance of shinnery oak heavily grazed areas (e.g., stock tanks, mineral licks), mast and oak insect galls to LPCHs in winter (Davis et and cultivated fields adjacent to grassland (Jamison et al. al. 1979). 2002a). These sites may be located on ridgetops or Small grains, short-horned grasshoppers (Acrididae), knolls that are higher than surrounding topography. beetles (Coleoptera), and green wheat were the 4 most Nesting. Lesser prairie-chicken females tend to select common food items found in LPCH diets of southwest- nest sites with high visual obstruction readings (VOR) ern Kansas (Schwilling 1955). Similar to other regions, (Robel et al. 1970) and horizontal cover (Giesen 1998), invertebrates and small grains comprised >90% of LPCH with most females selecting sites within 3.5 km of the diets in summer and winter, respectively. Lesser prairie- nearest lek. High VORs can result from selecting tall chickens were documented foraging on budding willows residual grass cover (Riley et al. 1992) or selecting vari- and cottonwoods (Populus spp.) in riparian areas during ous shrub species under which to nest (Table 1). In winter months (Schwilling 1955). either case, average vegetation height above the nest was The birds commonly forage in grain sorghum, corn, and 43-81 cm (Riley et al. 1992). Vegetation height at nest other grain fields adjacentto native pasture from late Table 1. Nest-site characteristics of lesser prairie-chickens from five studies in Texas, New Mexico, autumn throughearly spring Colorado, and Kansas. (Jamison et al. 2002a). Alfalfa was an food source Veg ht Shrubb Shrub Grass Forb important State n VORa (cm) % ht (cm) Grass % ht (cm) Forb % ht (cm) Studyc for females and prenesting Tex. 13 70 45 1 males in southwestern lekking N.M. 36 51 46 46 8 2 Kansas (Jamison 2000). Diets Colo. 29 61 7.2 48 29.4 36 1.4 21 3 of juveniles <10 weeks old are Kans. 174 48 15.2 43.8 37.2 19.2 8.4 16.3 4 thought to consist primarilyof a insects, specifically short- VOR = visual obstruction reading. b horned grasshoppers(Jones Note methods for estimating percent cover varied across studies; thus, percentages are not directly comparable. 1963a,b; Davis et al. 1979; c References for measurements: 1, Haukos and Smith (1989); 2, Riley et al. (1992); 3, Giesen (1994); 4, Jamison et al. 2002b). Pitman (2003). Lesser prairie-chicken management * Hagen et al. 73

Table 2. Habitat characteristics of successful and unsuccessful lesser prairie-chicken nests from 4 habitat (approximately4,000 in 2 studies in New Mexico and Kansas. types plants/ha)coupled with high invertebratebiomass A. Successful Unsuccessful (C. State n VORc Shrub % Grass % Forb % n VOR Shrub % Grass % Forb % Hagen,unpublished data). N.M.a (1) 5 87.4 32.5 64.0 3.5 3 36.6 31.3 49.6 19.1 Fall and winter N.M.a 4 55.9 41.8 55.1 3.1 17 39.5 48.1 44.5 7.4 (2) LPCHwin- N.M.a (3) 1 50.0 66.2 23.8 10.0 6 31.2 54.7 37.9 7.4 Historically, Kans.b 42 2.7 18.4 37.6 8.9 113 2.2 13.7 38.9 8.4 ter habitatincluded riparian corridorscomprised of a Riley et al. (1992) 1 = High Plains Bluestem Subtype (HPBS-1), 2 = HBPS-2, and 3 = HBPS-3 in south- deciduousshrubs and young eastern New Mexico. trees in the sand b sagebrush Pitman (2003) quantified vegetation in the sand sagebrush of southwestern Kansas. c Visual obstruction was measured the nearest to the nest et regions(Schwilling 1955), readings (VOR) (cm) by growing (Riley and al. 1992) and using a VOR pole (dm) by Pitman (2003). they foragedon acorns of shinneryoaks in southern partsof the range(Copelin sites was tallerthan adjacent rangeland sites (Davis et al. 1963, Davis et al. 1979). (Cultivation and availabilityof 1979, Haukosand Smith 1989, Riley et al. 1992, Giesen small grainschanged their foraging habits rangewide in 1994, Pitman2003). In sand sagebrushhabitat, success- the twentiethcentury (Gies sen 1998). LPCHsbegin for- ful nests often were locatedin areaswith relativelyhigh aging in grainfields (where available)once harvesthas shrubdensities (>5,000 plants/ha)(Giesen 1994, Pitman begun,usually in OctoberI(Schwilling 1955, Taylorand 2003). Successfulnesting generally required taller and Guthery1980a, Jamison2( 000). Despitethis shift away denserstands of vegetation(Table 2). Nest-site selection from nativeprairie, rangel and is importantfor roosting may be affectedby the proximityof humandisturbance, and loafinghabitat (Baker 1953, Taylorand Guthery as most LPCHnests were placedfarther from prairie 1980a, Jamison2000). In OklahomaLPCHs occupied edge and anthropogenicfeatures (e.g., powerlines, nativevegetative communi ties comprisedof grasses>80 pump-jacks,improved roads, and buildings)than expect- cm tall and were documenlted using wheat,western rag- ed at random(Pitman 2003). weed, andblue grama(Do: naldson 1969). In New Broodrearing. Lesserprairie-chicken broods occupy Mexico LPCHsforaged all most exclusively in shinnery habitatswith moderatestands of cover (Table3). oak-tallgrasscommunities during autumn and winter Specifically,they foragefor invertebratesin areaswith (Riley et al. 1993). abundantbare ground and approximately25% canopy cover of shrubs,forbs, or grasses20-30 cm in height Effects of (Jones 1963a, Donaldson1969, Davis et al. 1979, management Ahlborn1980, Riley and Davis 1993). In Kansas,brood Habitat management habitatwas selectedbased on vegetationthat provided Burning. In shinneryo< ak grasslands, spring burning high invertebratebiomass with moderateescape cover may resultin increasedcot unts of displayingmales and (Jamisonet al. 2002a, C. A. Hagen,unpublished data). relocationof leks to recent:ly burned areas (Cannon and Broodfemales selectedhabitats consisting of 15%forb Knopf 1979). The number*of displayingmales on one cover, sagebrushcover of 20% and moderatedensities areaincreased from a preburn total of 26 to a postbur total of 39. A 2-yearstudy yon the effects of fire on vege- tationtat in shinneryoak rang;elands of Oklahomasuggested Table3. Broodhabitat characteristics of lesserprairie-chickens from 5 p res rie rning o studies in Oklahoma, New Mexico, and Kansas. that prescrbed burning COIuld benefit LPCHs by provid- ing foragingareas, but the immediateeffects of fire on Veg nestingcover were negative, particularlywhen burns State Study ht (cm) Shrub %a Grass % Forb % were conductedin spring( Boyd 1999, Boyd and Bidwell Okla. Jones (1963a,b) 23 8 16 2001). A 3-yearstudy on 1greater prairie-chickens nesting Okla. Donaldson (1964) 14 51 35 in burnedand unburnedlar ndscapes in Oklahomafound N.M. Ahlborn (1980) 30 30 50 20 that 80%of all nests occurred in unburnedsites, and N.M. and Davis (1993) 25 43 43 15 Riley those nests >200 m from tlhe burn edge had the greatest Kans. Hagen (unpublished data) 30 17 26 11 probabilityof successfullyhatching (Wolfe et al. 2001). a documen Note methods for estimating percent cover varied across studies, Synder(1997) ltedimpacts of fire on sand thus percentages are not directly comparable. sagebrushcommunities. V 'isual obstruction and canopy 74 Wildlife Society Bulletin 2004, 32(1):69-82

cover of sagebrushwere reduced,and neitherhad recov- thatLPCHs used moderatelygrazed pastures more fre- ered to preburnlevels 7 yearspost-treatment. Forb quentlythan heavily grazed pastures. In New Mexico, recoverywas highly variablein post-treatment,but LPCHsused lightly grazedhabitats during drought years perennialforbs tendedto increasein the shortterm and but were able to use moreheavily grazed habitats in annualforbs respondedon a species-by-speciesbasis. years of near-averageprecipitation (Merchant 1982). Littonet al. (1994) and Snyder(1997) cautionedthat Habitatrestoration. Prairie restoration from agricul- burningin areasof loose, sandysoils shouldbe avoided turalland to grasslandhas had mixedresults on the becausea lack of adequateprecipitation and subsequent LPCH. ConservationReserve Program (CRP) lack of revegetationincreased the potentialfor wind ero- may providesuitable habitat, but few dataare available sion. thatquantify benefits of CRPto LPCHs(Rodgers 2000). Brush-beating and herbicides. Mechanical shrub Monocultureseedings have not providedadditional habi- removalis uncommonin habitatstypically occupied by tat to LPCHs. One studyis in progressto evaluatebene- LPCH;therefore, no dataare availableon the effects of fits of CRPfields as nestinghabitat, and preliminary brush-beatingon the species. Effectsof shrub-specific analysessuggest that a diversityof nativetall grassesand herbicideson LPCHsprobably are compoundedby inter- forbs in CRP seed mixes is important(T L. Fields, actionswith livestockgrazing, size of treatedarea, and ColoradoState University, personal communication). A resultingherbaceous cover (Jamisonet al. 2002a). new conservationpractice aims to restorerare and declin- Herbicidetreatment kills shrubsand allows an increasein ing habitats,but these plantingsare not yet well estab- grass cover if grass cover is not reducedby heavy graz- lished. ing (Donaldson1966, Doerrand Guthery1983, Olawsky Agriculture.Conversion of nativerange to cropland 1987, Olawskyand Smith 1991). Thereare no datathat probablyis most responsiblefor declinesin LPCHs,as it demonstratehow herbicidescan be used to createan has directlyimpacted available nesting habitat and interspersionof differentvegetation types or thatany her- reducednumbers of breedingbirds (Crawford 1974, bicide treatmentswill increasesurvival or recruitmentof Jamison2000, Hagen2003). Most agriculturalpractices LPCHs. Negativeeffects of herbicidetreatment on shrub in croplandare suspectedto affect LPCHpopulations cover may not become evidentuntil >3 yearsfollowing (Crawford1974). Maximumnumbers of LPCHswere herbicideapplications as the treatedshrubs fall and decay foundin areasin which 5-37% of the landscapewas (Rodgersand Sexson 1990, Jamisonet al. 2002a). plantedto grainsorghum using minimum-tillagetech- Loss of nativerangelands to woody cover is insidious niques(Crawford 1974). Recently,conversion of grass- as easternredcedar (Juniperus spp.), osage orange landsto agriculturehas slowed, as the numberof hectares (Maclurapomifera), and some exotics have transformed per yearconverted has not increased(Jensen et al. 2000, nativegrasslands into shrubor even forestedlandscapes. Woodwardet al. 2001). Woodwardet al. (2001) documenteda negativeassocia- Energydevelopment. Although abandoned oil-drilling tion betweenlandscapes with increasedwoody cover and sites frequentlyare used as lek sites, explorationand LPCHpopulation indices. developmentfor gas and oil productioneliminated use of Grazing. Overgrazingis a majorreason for declines 2 leks and disruptedactivity on a thirdlek in New in numbersof LPCHsbecause of degradationto nesting Mexico over a 3-yearperiod (Candelaria 1979, Davis et habitat(Taylor and Guthery1980b, Leslie et al. 1999, al. 1979). In Texasdisplaying males abandonedone lek Mote et al. 1999, Bailey et al. 2000). However,few data afteran elevatedroad was built acrossit (Crawfordand demonstratethe mechanismsby which grazingimpacts Bolen 1976b). Powerlines placednear leks may nega- LPCHdemography. In sandysoils heavy grazingmay tively affectbreeding activity of males (C. A. Hagen, resultin a shortageof the tall residualcover (Berg et al. unpublisheddata) as raptorsperching and huntingfrom 1997, Sims and Gillen 1999) requiredfor successfulnest- these poles may resultin reducedlekking activity. ing (Hoffman1963, Jacksonand DeArment1963, Litton Acousticaldisturbance (noise pollution)from oil or gas et al. 1994), and in firmersoils grazingmay resultin pumpsalso may affect lekkingdisplays. Proposedwind- conversionof tall-andmid-grass communities to a short- power-generationfarms also may increasevisual frag- grass-dominatedcommunity (Quinn and Walgenbach mentationof rangelandand cause abandonmentof 1990). Alternatively,moderate grazing in sandyregions lekkingsites. Pitman(2003) reportedthat females select- can yield greaterbasal cover of mid-grassesand forbs ed nest sites in southwesternKansas that were signifi- (Quinnand Walgenbach 1990, Sims and Gillen 1999) cantlyfarther from anthropogenicfeatures (e.g., power thatmay be beneficialto both nestingsuccess and brood lines, pump-jacks,improved roads, and buildings) associ- rearing,respectively. In Oklahoma,Copelin (1963) noted ated with energydevelopment than expected at random Lesser prairie-chicken management * Hagen et al. 75

Table 4. Distances (m) to anthropogenic features from lesser prairie- these effortsfailed and resultsof the otherwere undeter- chicken nest sites, use-sites and areas not likely to be frequented by less- mined(Horton 2000). It is likely thatthe numbersof er prairie-chickens (non-use), and areas absent of prairie chickens. birdsreleased were too small and the qualityof habitatat areaswas unsuitableto sustainthese birds Mean distances (? SE) to anthropogenic featuresa release Nest sites Use Non-use Absent sites (Giesen2000). Feature (n = 187) (n = 44) (n = 38) (n = 46) Hunting. The impactof moder-era sporthunting on in the Power line 1,320 + 66 1,106 81 666 + 80 705 + 82 the LPCHis unknown,although market hunting Wellhead 564 + 22 435 31 446 +31 323 + 31 1800s may have been detrimentalto local populations Building 2,129 + 56 1,397 + 106 1,061 + 105 759 + 108 (Jacksonand DeArment1963, Horak1985, Johnsonand Unimproved roadb 214+14 193?18 178+18 184+19 Knue 1989). The lesser prairie-chickenonce was hunted Improved road 2,377 + 150 in all 5 stateswithin its range(Giesen 1998). However, Agricultural edge 1,049 + 47 concernsover low populationsled to closureof hunting 709 Nearest lek + 36 seasonsin the early 1900s in Colorado(Giesen 2000) and in the 1990s in New Mexico and Oklahoma and a Distances for nests taken from Pitman (2003) and other use sites (Bailey from Hagen (2003). Williams2000, Horton2000). Prairie-chickenharvests b Hagen (2003) did not differentiate between type of roads, and these (i.e., harvestedbirds-per-hunter ratios) have declinedover distances likely reflect those of unimproved roads. the long term(Figure 2). Currently,a 2-day season and a 2/4-birdbag/possession limit are allowedin Texasand a 2-monthseason and 1-birdbag and 4-birdpossession (Table4). Hagen(2003) foundthat areas used by limit are allowedin Kansas. radiomarkedmale and nonnesting-femaleLPCHs were significantlyfarther from these same featuresthan areas not used by LPCHs(Table 4). These studiesindicated 4.0 thatLPCHs likely would use less-disturbedareas even 3.5- 3.0- thoughvegetation composition or structuremay be simi- lar betweenused and unusedsites. Muchresearch is 2.5- 2.0-

neededto determinethe effects of energyexploration and 1.5-

developmenton LPCHs. 1.0 - 0.5 -

0.0 Population management 1965 1970 1975 1980 1985 1990 1995 2000 Texas Extrinsic factors 4.0 Weather.Impacts of weatheron LPCHpopulations cu 3.5- C are not well known. However,an associationappears to 3.0- V 2.5- exist betweenproduction (i.e., age ratiosin harvest)and 'a precipitation(Brown 1978, Giesen 2000). Driercondi- 2.0- tions resultin sparsenesting cover and less food for 1.5- 1.0 - chicks. Harvestlevels in New Mexico were correlated m with from the 0.5- positively precipitation previousyear 0.o (Brown 1978). Populationtrends in Colorado,as meas- 1965 1970 1975 1980 1985 1990 1995 2000 uredfrom lek counts,were correlatedpositively with the Kansas 4.0 levels from 2 to census (Giesen precipitation yearsprior 3.5-

2000). 3.0- Translocations. to restore Generally,transplants 2.5-

prairiegrouse in unoccupiedhabitats have been success- 2.0- ful (Synderet al. 1999). However,3 stateshave conduct- 1.5- ed translocationsof LPCHswith little success. Colorado 1.0- released155 birdsduring spring over a 6-yearreintroduc- 0.5- tion attempt,but the effortwas not successful(Giesen 0.0 1965 1970 1975 1980 1985 2000). Texas translocated46 LPCHsdur- 1990 1995 2000 unsuccessfully New Mexico ing 2 years to supplementan existingpopulation in a native Oklahomatranslocated Figure 2. Annual lesser prairie-chicken harvests in Kansas, New vegetativecommunity. Mexico, and Texas expressed as birds harvested per hunter. Note there LPCHsin an attemptto re-establish2 populations;one of are a number of years with missing data. 76 Wildlife Society Bulletin 2004, 32(1):69-82

Intrinsicfactors Conservationstrategies Disease and parasites. There is a potential for intrin- We recommend that each state develop and implement sic factors to limit LPCH populations (see Peterson this conservation plans for LPCHs. These plans should use issue); however, most studies have not documented dele- local groups comprised of representativesfrom all interest- terious effects at the population level. Hagen et al. ed stakeholdersto identify and solve regional issues within (2002) found low levels (<5%) of Mycoplamsa spp. anti- ecological regions. Conservationplans should include 1) bodies in LPCH sera (n= 162) in Kansas and concluded quantity and quality of LPCH habitatremaining in the that such levels likely were not limiting to the popula- state, 2) common problems involved in conserving the tions. In Texas, Peterson et al. (2002) documented the LPCH, and 3) conditions needed to maintainhealthy pop- first incidence of infectious bronchitis (a coronavirus) ulations. Regional variationsin vegetative communities antibodies in LPCHs. If a coronavirus should become (e.g., sand sagebrush, shinnery oak, mixed shrub, or grass widespread in a population, the effects could be devastat- dominated), weather, or resource use that affect popula- ing, and Peterson et al. (2002) urged that further work be tions and their management need to be considered in con- conducted to assess the prevalence of coronaviruses in servation plans. To date, only New Mexico has developed LPCHs. Robel et al. (2003) reported the presence of and is implementing such a plan (Massey 2001). helminthic parasites of Tetrameresspp. (stomach worm), Because LPCH populations are nonmigratory or local Oxyspirurapetrowi (eye worm), and Subulura spp. (cae- migrants (i.e., migrations of <60 km), large ecological cal worm) in 92, 95, and 59% of LPCH carcasses (n= regions should be identified as LPCH management zones 93), respectively. Alternatively, incidence was lower in (Figure 3). We recommend that these zones cover eco- fecal samples of radiomarkedindividuals (n=46 para- logical regions (e.g., Arkansas River Sand Hills) within a sitized birds, n=52 nonparasitized birds), but the pres- state, and that target areas no smaller than 64 x 64 km be ence of helminths did not measurably affect body mass, identified in each zone (Figure 3), as this would encom- clutch size, nesting success, daily movements, or survival pass the longest known movements of individual birds of radiomarkedLPCHs (Robel et al. 2003). (Hagen 2003). Identifying and prioritizing target areas Genetics. Van den Bussche et al. (2003) found that will facilitate better field management and working rela- LPCHs in Oklahoma and New Mexico maintained high tionships with private landowners, thus creating a frame- levels of genetic diversity at both nuclear and mitochon- work for ensuring connectivity throughout a management drial loci, but there was some structuringbetween states. zone. Concentrating habitat restoration and management Hagen (2003) extended the mitochondrial work of Van efforts in target areas will ensure that funding and per- den Bussche et al. (2003) to include 5 populations from sonnel resources are used efficiently. Given the frag- Kansas and 1 from Colorado. There was substantial pop- mented nature of LPCH habitat, management zones will ulation structuringand genetic diversity across the 4 encompass large areas of unsuitable habitat (Figure 3). states; notably, New Mexico had the fewest haplotypes. Identifying target areas where habitat restoration, man- Much work is needed to identify genetically isolated pop- agement, and population monitoring are the most effica- ulations and to determine whether there has been a loss cious will facilitate protection and creation of large habi- of heterozygosity from population bottlenecks, as has tat blocks and maintain adequate levels of connectivity been documented in greater prairie-chickens (Bouzat et (Figure 3). These target areas should be ranked for man- al. 1998, Bellinger et al. 2003). agement action based on existing habitat quality and quantity, LPCH populations, and the potential for LPCH to expand into a target area. Management guidelines Macro-scale management specific to physiognomic The following guidelines reflect our current under- and ecological characteristics of LPCH management standing of LPCH population ecology and habitat zones could be conducted. A common set of recommen- requirements. Because experimental data on the effects dations that apply to the characteristics of a management of habitat alterations and other management activities on zone would specify grazing regimes, furbearerharvest, LPCH populations are lacking or absent, these guidelines and CRP plantings. Zone management should include should serve as working hypotheses to be tested using the LPCH population monitoring; inventory of habitat quan- scientific method under the principles of adaptive tity, quality, and connectivity; and harvest regulations for resource management (Walters and Holling 1990). Using states with open seasons. this approach, rigorous monitoring of management actions could provide practical information on effects of Habitatguidelines management on habitats and population responses. There is little question that prairie-chickens are area- Lesser prairie-chicken management * Hagen et al. 77

(C) Nesting. Provide dense (A) shrubs and residual bunch- grasses >40 cm tall that pro- vide >75% vertical screening in the first 33 cm above ground and 50% overhead cover for quality nesting habitat. In shinnery oak communities, sand bluestem (Andropogon spp.) that is >50 cm in height provides suitable nesting cover. Maintain or increase shrub cover in sand sagebrush com- munities (particularly in areas where taller species of grass have been reduced by grazing practices). Manage grazing to maintain adequate height (>25 ArkansasRiver Sand Hills cm) and density of residual ManagementZone and4 TargetAreas grasses and forbs. Consider Figure 3. An LPCHManagement Zone (gray box) should encompass an ecological region, as in the sand fencing areas (>1.6 km2) to x sagebrush of southwestern Kansas (A), target areas are identified based on 64 64-km subdivi- prevent grazing during the sions (4 black outlines) within the management zone (B), and specific habitat patches (habitat in gray, non-habitat in white) can be monitored and managed within a target area (C). The target area in Figure nesting season; because breed- 1C includes the Cimarron National Grassland along the Cimarron River, Kansas. ing females generally space their nests to avoid predation pressure, fencing small areas sensitive species, and large quantities of habitat are will not provide nesting cover for significant numbers of essential for population growth. However, habitat quality females. Evaluate the feasibility of predator removal is of equal importance. The population ecology of LPCH efforts to enhance nesting success of imperiled popula- (i.e., reproductive potential and relatively stable mortality tions in small areas (<2,000 ha) of habitat (Bergerud rates in specific habitats) strongly suggests that increas- 1988, Schroeder and Baydack 2001). Predator removal ing breeding success is the key to increasing numbers of can increase nest success of prairie grouse (Lawrence and birds (Bergerud 1988, Wisdom and Mills 1997, Jamison Silvy 1995) and other ground-nesting birds (Garretson et 2000, Hagen 2003). Thus, LPCH habitat management al. 1996, Witmer et al. 1995), but long-term intensive should focus on providing adequate cover for nesting and management may not be economical. Passive predator brood rearing, given the specific cover requirements of management, through relaxation of restrictions on har- certain habitats. Cover needs are greatly increased where vest, take, or opportunistic gunning of certain furbearers, predation pressure is high (Bergerud 1988). should be considered within the boundaries of manage- Breeding habitat. Protect, maintain, and restore ment zones. Nesting habitats should be interspersed with >2,000-ha tracts of native shinnery oak-tallgrass or sand brood habitats to facilitate easier movements of young sagebrush grassland within LPCH management zones. broods between habitats and reduce predation or starva- These areas must be large enough and close enough to tion rates (Bergerud 1988, Pitman 2003). other patches (<30 km) to support viable LPCH popula- Brood rearing. Provide habitat with 20-40% canopy tions during drought. However, tracts that are smaller of shrubs, forbs, or grasses that are 24-30 cm in height. (>500 and <2,000 ha) but with high connectivity (<10 In shinnery oak communities, provide vegetation domi- km spacing) also should be included in such efforts. nated by warm-season grasses and shinnery oak with Maintain >63% native grassland (Crawford 1974) and about 60% bare ground (Riley and Davis 1993). Provide stabilize land-use practices in these landscapes managed vegetation composed of about 43-60% grasses, 24-43% for LPCHs, and conserve shrub-dominated or grassland shrubs (primarily shinnery oak), and 13-26% forbs communities within >3.5 km of lek sites because most (Riley and Davis 1993). This diversity of vegetative nesting occurs within this radius (Giesen 1998, species and structure is important for providing the prop- Woodward et al. 2001, Pitman 2003). er substrate for insects needed by juvenile LPCHs (Riley 78 Wildlife Society Bulletin 2004, 32(1):69-82

and Davis 1993, Jamison et al. 2002b). tive framework would allocate different levels of live- Burning. Prescribed bums should be conducted with stock grazing based on the previous 12 months' precipita- great caution in LPCH habitats because the vegetative tion. This system would allow the rancher to manage the response to fire is not well understood in these xeric operation more effectively with few surprises. grasslands (Engle and Bidwell 2001). Some nesting habi- Habitat restoration. Conservation Reserve Program tats may require 7 years or more to recover before they grasslands should range from 30-75 cm in height, as provide adequate concealment following a fire (Synder stands <30 cm are generally inadequate for concealment 1997). Experimental spring bums could be conducted at cover and those >75 cm seem to be avoided (R. D. sites recently abandoned by LPCHs but adjacent to areas Rogers, Kansas Department of Wildlife and Parks still inhabited. Conserve shinnery oak motts and protect [KDWP], personal communication). Multispecies seed- oak bud, mast, and catkin production (Boyd et al. 2001) in ings create height and growth-form heterogeneity and relatively mesic shinnery oak communities by discing must include native bunchgrasses, forbs (particularly firebreaks around motts, and avoid annual burning of legumes, important for structureand as a food source), large areas to conserve residual nesting cover (Boyd and and native shrubs. Aggressive grasses that can crowd out Bidwell 2001, Harrell et al. 2001). Alternatively, burning other components of the mixture or grass monocultures may be an advantageous tool to reduce juniper (Wright must be avoided (R. D. Rogers, KDWP, personal commu- 1974) or mesquite (Prosopis spp.) encroachment (Harrell nication). Native plant communities should be restored et al. 2001). Given the conservative use of fire in LPCH based on USDA-NRCS Ecological Site Guides. range, we recommend managing juniper using mechanical Agriculture. Prevent further cultivation of grassland removal techniques when possible. surroundingleks and disturbance of lek or nest sites Grazing. A grazing system that maintains middle to because such activities directly cause habitat loss. late stages of plant succession interspersed with early However, planting small grains or corn in existing agri- stages of plant succession is optimal for LPCHs (Bidwell cultural fields adjacent to native prairie can provide addi- 2003). To achieve this heterogeneity, grazing systems tional winter food sources. Minimum tillage techniques must incorporate periods of rest to prevent excessive will reduce soil erosion and may benefit LPCHs that are grazing. Because excessive grazing yields lower-quality using the fields. concealment cover and reduces foraging habitat (particu- Brush treatments. Much research is needed to demon- larly for nesting and brood rearing) continuous grazing is strate effects of brush treatments on LPCHs. Minimize not recommended (Bidwell 2003). Alternatively, we rec- the use of herbicides, except to control invasive nonnative ommend light or moderate grazing that will ensure vegetation. However, if herbicides must be used, we rec- 60-70% of key herbaceous species (Holochek et al. ommend that such treatments not reduce sand sagebrush 1989) will be available as residual nesting cover (Berg et or shinnery oak to less than 25% of the canopy within al. 1997, Snyder 1997, Sims and Gillen 1999). At least one year after treatment. Shrub removal treatments 20-30% of a pasture should be rested completely in rota- should create a mosaic of treated and untreated areas to tions of about once every 3-5 years (Fuhlendorf and provide an interspersion of vegetative structuresdominat- Engle 2001, Bidwell 2003), and this vegetative response ed by grasses and shrubs for nesting cover and areas with can be maintained using patch-burningmethods in which a diversity of vegetation for brood rearing, foraging, and 20-30% of an area is burned annually (Bidwell 2003). adult autumn and winter cover. To create a mosaic of Livestock preferentially graze recently burned patches, vegetative structures, apply tebuthiuronin strips <10 m leaving the remaining 70-85% in various successional wide at rates of 0.2-0.4 kg/ha. Such treatments should stages. No data are available on the effects of deferred be applied with a tractor-mountedsprayer (Snyder 1997). grazing systems (which postpone grazing until grassland Preserve small (<10-ha) motts of tall shinnery oak that plants have matured) or rest-rotation grazing systems produce mast crops by excluding these areas from herbi- (which involve multiple pastures through which livestock cide applications. Treat shrubs in contour strips on a 10- are rotated) on LPCH populations, but appropriateuse of year rotation to provide suitable interspersion of nesting these systems probably would create suitable intersper- and brood-rearinghabitats while reducing wind erosion sion of different vegetation heights (Manely et al. 1997). of sandy soils (Jamison et al. 2002a). Avoid annual Large pastures and fewer livestock water sources also chemical brush treatment of large areas because this may will result in a diversity of grazing pressures at the land- reduce LPCH production (Wiedenfeld et al. 2001). scape level. We suggest an adaptive grazing strategy that Woody vegetation >3 m in height also should be thinned would adjust stocking rates and season of use based on using mechanical methods when possible. Similar proto- grazing system and annual precipitation. One such adap- cols should be followed with mechanical treatment of Lesser prairie-chicken management * Hagen et al. 79

LPCHhabitat to ensurethat a mosaic of habitatsis pro- information(sex and age ratios),lek surveysare the pri- vided. marymethod of estimatingminimum spring breeding Energydevelopment. Oil and natural-gasexploration populations.In stateswhere LPCHs are hunted,age drill pads may createareas that are suitablefor lek sites, ratiosare difficultto estimate,given the low harvestrates but lek sites generallyare not limitingto the species. and the trophystatus of the species (i.e., huntersprobably The disturbanceassociated with explorationand produc- are not willing to contributewings or otherparts). tion of petroleummay destroynesting habitat or cause Additionally,methods for evaluatingrecruitment rates lek or nest abandonmentand shouldbe discouraged.If are importanttools for monitoringmanagement actions. constructionis unavoidable,it shouldoccur outside of the We recommendthat brood survey methods be developed breeding,nesting, or earlybrood-rearing seasons. Thus thatcan be implementedand comparedwithin each of 3 we recommendthat all constructionand extractionbe habitatsof the LPCH. Ideally,such methodscould be avoidedfrom 15 March-15July. We also recommend comparedbetween habitat types. thatwind turbinesor otherlarge vertical structures be Hunting. Currentlevels of harvestin Texasand constructed>2 km fromknown or potentiallyoccupied Kansasappear to be low enoughthat populations should LPCHhabitat. If such structuresmust be placedin not be impacted. However,hunter surveys and lek counts knownLPCH habitat, they shouldbe positionedalong shouldensure that local areasare not overharvested.We the prairieedge or clusteredin sites with otherdistur- recommenda permitsystem (similarto thatimplemented bances. by Texas)specific to huntingLPCHs, which shouldfacil- itate moreefficient recording of harvests. Wherepossi- ble, the use of check stationswould allow evaluationof Populationguidelines huntersuccess, productionof juveniles,and monitoring Population monitoring. It is imperative that survey of sex ratios. methodologiesand effortbe standardizedfor estimating Translocations.Future translocations must firstquan- springpopulations and recruitmentacross the range. tify the qualityof the nestingand brood-rearing habitat at Improvementsto lek surveymethods are worthwhile the releasearea; then the success of translocationsshould (Schroederet al. 1992, Giesen 2000) but will require be evaluatedusing radiotelemetry,lek monitoring,and time and researchto develop. Specifically,future broodsurveys. Becausenumbers of meso-mammalpred- researchis neededto determinethe relationshipof lek atorsmay be elevatedin fragmentedlandscapes and surveysto 1) numberof nestingfemales, 2) variationin becausethose predatorsmay enjoy easieraccess to totalpopulation size, and 3) actualdensities of leks and femalesnesting near habitat edges, predator-control breedingbirds. In the interim,surveys that attempt to effortsimplemented immediately before and afterreleas- count all leks in areasknown to be occupied,where es of birdsmay proveadvantageous to newly supple- occurrenceis likely, and thatmay be restoredshould be mentedor translocatedpopulations (Lawrence and Silvy implementedby wildlife and naturalresource agencies 1995). Once populationsare established,predator-con- rangewide(Davison 1940, Hamerstromand Hamerstrom trol effortsmay no longerbe cost-effectiveor necessary. 1973). This shouldinclude surveying known leks and potentialsites at least 3 times duringthe peak of breeding (approximately21 March-21April) andbetween the 30 Conclusions minutesbefore and no laterthan 1 hourafter sunrise Althoughthe rateof habitatloss has slowed over the (Crawfordand Bolen 1975), on days with wind <16 past 20 years,LPCH population trends continue to km/hrand no precipitation. decline, suggestingthat it is the quality(i.e., human- Once methodshave been standardized,spring popula- inducedor drought)of the habitatthat may be limiting tions of LPCHsshould be monitoredusing lek survey the recoveryof these populations.Reliable knowledge is methodsto count all activeleks and searchpotential neededon restoration,the characteristicsof healthy areasfor new or satelliteleks withinLPCH management grasslandand steppeecosystems, and the relationshipof zones. Once breedingpopulations have been identified grazingand LPCHproduction. Field experimentsare in these managementzones, lek surveyscan be used to neededthat measure the effects of variouslevels of graz- monitorlong-term population trends. Lek surveysmay ing pressureon nest success, chick survival,and inverte- providean index to the size of the breedingpopulation brateabundance. Other management tools for habitat and may detectlong-term changes in policy or landuse maintenance(e.g., brushtreatments) and restoration (e.g., thatimpact LPCH populations (Beck and Braun1980). CRP) also requireexperimental manipulations to evaluate In the absenceof mark-recapturestudies and harvest the best managementpractices for LPCHs. 80 Wildlife Society Bulletin 2004, 32(1):69-82

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Prairie Grouse Technical ics and habitat use in ptarmigan, prairie grouse, mountain plovers, Council 24:39 (Abstract). and grassland passerines. Retired (2003). TerryZ. Riley (photo) is WOODWARD,A. J., S. D. FUHLENDORF,D. M. LESLIE,JR.,ANDJ. SHACKFORD.2001. Director of Conservation for the Wildlife Management Institute (WMI) in Influence of landscape composition and change on lesser prairie- in Tijeras, New Mexico. Before moving to New Mexico October he was Director of Conservation for WMI in D.C. chicken (Tympanuchus pallidicinctus) Populations. American 2003, Washington, He was a WMI field for the Midwest and Northern Midland Naturalist 145:261-274. representative states in Aberdeen, and Chariton, Iowa WRIGHT,H.A. 1974. Effects of fire on southern mixed grass prairies. from 1994-1999. Prior to working for WMI, Terrywas the upland Journal 27: 417-419. Range Management wildlife research biologist for the Iowa Department of Natural Resources in Chariton, Iowa (1989-1994) and a wildlife biologist for the United States Forest Service on the Black Hills National Forest in Christian A. is the Grouse Conservation Coordinator for Hagen Sage South Dakota (1981-1985) and on the Chequamegon the of Fish and Wildlife in Spearfish, Oregon Department Hines, Oregon. National Forest in Park Falls, Wisconsin (1978-1981). He received his Christian received his Ph.D. in wildlife from Kansas State biology B.S. in fish and wildlife from Kansas State University in 1976, a Master's of Natural Resource from biology University (2003), Management an M.S. in wildlife science from New Mexico State University in of Manitoba (1999), and a B.A. in science from University political 1978, an M.S. (1987) and Ph.D. (1989) in zoology from Ohio State Fort Lewis College (1993). He has been an active member of TWS is a certified wildlife and his activities in since as student to the Manitoba University. Terry biologist, 1997, serving representative Chapter The Wildlife include of the Kansas State University and on the Committee for the CMPS annual meet- Society president (1998-99) Planning Student (1975-1976) and the South Dakota Chapter (1985), he and co-chaired the at the 2002 Chapter ing (2000); organized symposium of the North Central Section (1994), board member of the TWS annual conference in Bismarck from which this was secretary paper pub- Central Mountains and Plains Section (1998-2001), and various com- lished. His interests include habitat and of population management mittee assignments. His professional interests include wildlife habitat birds, of human disturbance on wildlife, demo- upland game impacts management on agricultural, prairie wetlands, and grassland ecosys- and habitat selection Brent E.Jamison graphic modeling, analyses. tems. He also has extensive experience with legislative policy. currently is the upland game biologist for the Yakama Nation in south-central Washington, where he directs habitat restoration work and participates in regionwide bird conservation activities including sage-grouse management. Brent earned a B.S. degree in fisheries and wildlife from the University of Missouri-Columbia and an M.S. in wildlife biology from Kansas State University. His interests include management of upland game bird populations and habitat issues affecting grassland and shrub steppe birds. Kenneth M. Giesen earned his B.S. from the University of Minnesota (1972) and M.S. from Colorado State University (1977). He was a researcher for Colorado Division of Wildlife (1980-2003) and investigated population dynam- Special section associate editor: Silvy