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_3|6by ,:. ::: MichaelA. Schroeder and .:. .^. :%9: M:g9tW;x3&90Wt93FiXti-:::: LET:: lvlcllaruD* h 2 1x.r Davuacsn ,1 1
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ESe'E ,.,.,::. :::>':E ': .::ji :E:in: :: 1 .X.-72 , t0Ws:: :, x --: :piaper E ::: ii8.:.g ,:.E: examines the importanceof predationin the life cycles of sage grouse (Centrocercus urophasianus), sharp-tailedgrouse (Tympanuchus phasianellus), greaterprairie-chicken (17 cupido), and lesser prairie-chicken(l: pallidicinctus). Most individualprairie grouse eventuallysuccumb to predation,with substantial effects on nest success, juvenilesurvival, and adult survival. Predatorcontrol has occasionallybeen used as a managementtool with the belief that reducingpreda- tor numberscan enhance viabilityof game populationsin generaland prairie grouse in particular.Although some experimentalresearch has shown that direct reduction of predatornumbers can increase grouse recruitment,most currentman- agement plans recommendindirect management of the grouse-predatorrelation- ship by manipulatinghabitats. However,as habitatsbecome more fragmentedand alteredand populationsof prairiegrouse become more threatenedand endan- gered,it is importantto reconsiderpredator control as a managementoption and to evaluateits viabilitythrough experimentation. Key Words Centrocercus urophasianus, greaterprairie-chicken, grouse management,lesser prairie-chicken,population regulation, predator-prey relationships, sage grouse, sharp-tailedgrouse, Tympanuchus cupido, Tympanuchus phasianellus, Tympan- uchus pallidicinctus
ife historiesof sage grouse,sharp-tailed grouse, greater We review availableinformation on the importanceof prairie-chicken,and lesser prairie-chickenhave been stud- predationin the life cycles of prairiegrouse and place iedjhroughout their ranges. Despite variationin behav- thatinformation into a managementcontext. Because abitat, and status,populations of prairiegrouse are predationpressure can be manipulateddirectly by con- similarregarding extent, timing, and significanceof mor- trollingpredator numbers (Batterson and Morse 1948) talityfrom predation (Schroeder and Robb 1993, and indirectlyby manipulatinghabitats (Hamerstrom et Connellyet al. 1998, Giesen 1998, Schroederet al. 1999). al. 1957), an additionalobjective of this paperwas to The consequencesof predationon the populationdynam- evaluatethe viabilityof alternativestrategies. ics of prairiegrouse are often consideredto be substantial enoughthat the effectivemanagement of prairiegrouse requiresthe directand indirectmanipulation of predation Descriptionof prairiegrouse predation pressure(Batterson and Morse 1948, Hamerstromet al. Sage grouse 1957, Lawrence1982, Riley et al. 1992). Averageclutch size for sage grousevaries from 6.6 to
Addressfor MichaelA. Schroeder:Washington Department of Fishand Wildlife,P.O. Box 1077, Bridgeport,WA 98813, USA;e-mail: grouse@ televar.com.Address for RichardK. Baydack:Natural Resources Institute, University of Manitoba,Winnipeg, Manitoba, R3T 2N2, Canada.
Wildlife Society Bulletin 2001, 29(1):24-32 Peer refereed Predationand prairiegrouse * Schroederand Baydack 25
9.1 eggs, and the nest success ratevaries from 15 to 86% raven,and American crow (Corvusbrachyrynchos). (Wallestadand Pyrah1974, Connellyet al. 1993, Gregg Becausenest success is positivelycorrelated with the et al. 1994, Schroeder1997). Nest predatorsinclude presenceof relativelythick grass cover, activities such as groundsquirrel (Spermophilus spp.), badger(Taxidea grazingor hayingmay negativelyimpact nest success taxus),coyote (Canis latrans),and commonraven (Kohn 1976, Kirschet al. 1978, Marksand Marks1987, (Corvuscorax, Battersonand Morse 1948, Patterson Giesen and Connelly 1993) 1952, DeLong et al. 1995). Nest success is positively Primarypredators of sharp-tailedgrouse include coy- correlatedwith the presenceof big sagebrush(Artemisia ote, northerngoshawk (Accipiter gentilis), gyrfalcon tridentata)and relativelythick grass and forb cover (Falco rusticolus),rough-legged hawk (Buteo lagopus), (Connellyet al. 1991, Gregg et al. 1994, DeLong et al. Predator controllLs are rarely recommended for North 1995, Sveumet al. 1998). Americanprairie grouse, even for increasinglythreatened Consequently,improper man- agementof habitathas often and endangeredpopulations living in altered,isolated, or been implicatedin declinesin fragmentedhabitats. This is due to...factorsincluding the nest success (Trueblood lack of informationabout the long-termconsequences of 1954, Klebenow1969, Braun et al. 1977, Fischeret al. predatorcontrol, the relativelyhigh cost of predatorcon- 1996). trol,the protected status of many potential predators,and Primarypredators of sage grouseinclude golden eagle concerns about public attitudestoward predatorcontrol (Aquilachrysaetos), red- | tailed hawk (Buteojamaicensis), Swainson's hawk (B. northernharrier, red-tailed hawk, and greathorned owl swainsoni),ferruginous hawk (B. regalis),northern harri- (Bubovirginianus, Lano 1912, Gross 1930, Marshalland er (Circuscyaneus), common raven, weasel (A8ustela Jensen 1937, Blus 1967). Althoughmortality of juve- spp.), and coyote (Rasmussenand Griner1938, Scott niles is poorlydocumented, it appearsto be substantially 1942, Patterson1952, Dunkle 1977). Mortalityfor juve- greaterthan for breeding-agebirds (Ammann 1957, niles has been estimatedto be 63% duringthe first few Hillmanand Jackson1973, Connellyet al. 1998). weeks afterhatch (Wallestad 1975). Habitatalteration Annualmortality of breeding-agesharp-tailed grouse associatedwith grazing,drought, and wildfiremay variesfrom 17 to 55% (Robelet al. 1972, Moyles and increasethe rateof predationon juveniles,but the rela- Boag 1981, Giesen 1987, McDonald1998). Predation tionshipsare not clear (Battersonand Morse 1948, appearsto be particularlyimportant for breeding-age Klebenow1972). birdsduring the breedingseason (Bergerud1988, Annualmortality of breeding-agesage grousevaries McDonald1998). Predationalso may be substantialdur- from 55 to 75%for femalesand 38 to 60%for males; ing severewinters, particularly from avianpredators mortalityappears to be greaterin huntedpopulations (Ulliman1995, Connellyet al. 1998). (June 1963, Zablan1993, Connellyet al. 1994). Rela- tively little mortalityis causedby f1re,weather, and colli- Greaterprairie-chicken sions with wires, fences, andvehicles (Patterson1952, Averageclutch size for greaterprairie-chickens varies Dalke et al. 1963, Schroederet al. l999). Predation from 8.2 to 12.9 eggs and nest success ratefrom 22 to appearsto be particularlyimportant for femalesduring 65% (Yeatter1943, Ammann1957, Vanceand the incubationand brood-rearing seasons and for males Westemeier1979, Petersonand Silvy 1996). Nest preda- duringthe breedingseason (Patterson 1952, Schroederet tors includeground squirrel, badger, striped skunk, opos- al. 1999). Predationrate on breeding-agebirds may not sum (Didelphisvirginiana), raccoon (Procyon lotor), be influencedby harvestrate or habitat(Braun 1998). coyote,American crow, and fire ants (Solenopsisspp., Gross 1930, Lehmann1941, Bowen et al. 1976, Sharp-tailedgrouse Svedarsky1988). Nest success is greaterin areaswith Averageclutch size for sharp-tailedgrouse varies from relativelythick grassand forb cover (Yeatter1963, 10.9 to 12.3 eggs and nest success ratefrom 50 to 72Wo Bowen et al. 1976, Buhnerkempeet al. 1984, Lutzet al. (Hamerstrom1939, Hartet al. 1950, Ammann1957, 1994). Managementof habitatthat results in loss of Meints 1991). Nest predatorsinclude coyote, striped residualvegetation may resultin reducednest success skunk(Mephitis mephitis), ground squirrel, common (Lehmann1941, Arthaud1970, Kirschet al. 1973). 26 Wildlife Society Bulletin 2001, 29(1):24-32
red-tailedhawk, rough-legged hawk, ferruginoushawk, prairiefalcon (Falco mexicanus), greathorned owl, gold- en eagle, and northernharrier (Campbell 1950, Copelin 1963, Merchant1982, Haukos 1988, Giesen 1998). Juvenilemortality appears to be substantialduring the first few weeks afterhatch (Giesen 1998). Annualmor- tality of adultmale lesser prairie-chickensranges between35 and 45% (Campbell1972). Relativelysmall amountsof mortalityare causedby drowningand colli- sions with wires, fences, and vehicles (Campbell1972, Sell 1979, Merchant1982). Thereis little informationon the annualdistribution of predationpressure (Giesen 1998).
Greaterprairie-chicken nest in northeasternColorado. Researchon prairiegrouse has demonstrateda relationshipbetween nest success Predationand life history and habitatquality. Most prairiegrouse are eventuallykilled by predators (Bergerud1988). Predationhas the potentialto affect the Primarypredators of greaterprairie-chickens include annuallife cycle in 3 primaryways: 1) success of nests, red-tailedhawk, northern goshawk, rough-legged hawk, 2) survivalof juveniles duringthe f1rstfew weeks after broad-wingedhawk (Buteo platypterus), northernharrier, hatch,and 3) annualsurvival of breeding-agebirds. The greathorned owl, and coyote (Yeatter1943, Bergeret al. relativeimportance of predationon populationviability 1963, Hamerstromet al. 1965, Sparlingand Svedarsky remainslargely unstudied (Peterson and Silvy 1996). 1978). Juvenilemortality appears to be greaterthan for breeding-agebirds, particularly during the first few Nest success weeks afterhatch (Bowman and Robel 1977, Peterson Nest success, which is the proportionof nests where and Silvy 1996). Annualmortality of breeding-age >1 egg hatches,is usuallyconsidered the most significant greaterprairie-chickens is estimatedto be 45% for males featureinfluencing the populationdynamics of prairie and49% for females;mortality tends to be greaterin grouse (Angelstam1986, Bergerud1988, Petersonand huntedpopulations (Hamerstrom and Hamerstrom1973). Silvy 1996). This is primarilydue to the fact thatnest Relativelysmall amountsof mortalityare causedby fire, success is extremelyvariable and differencesin nest suc- weather,and collisions with wires, fences, and vehicles cess can be attributedto variationin habitatcharacteris- (Gross 1930, Hamerstrom1939, Lehmann1941, tics or predationpressure associated with year,area, pop- Ammann1957, Svedarsky1988). Thereis little informa- ulationdensity, or managementstrategy (Peterson and tion on the annualdistribution of predationpressure Silvy 1994, Connellyet al. 1998, Giesen 1998, Schroeder (Schroederand Robb 1993). et al. 1999). Althoughprairie grouse may respondto high ratesof nest predationby renestingwithin the same
LesserP rairie chicken breedingseason (Svedarsky1988, Schroeder1997, Averageclutch size for lesser prairie-chickensis 10.4 Connellyet al. 1998, Giesen 1998), predationpressure on eggs and nest success ratevaries from 0 to 67Wo(Copelin nests also can be mitigatedby providinghabitat in suffi- 1963, Sutton 1968, Donaldson1969, Riley 1978, cient qualityand quantity(Hamerstrom et al. 1957, Merchant1982, Haukos1988, Giesen 1998). Nest pred- Kirsch 1974, Connellyet al. 1991, Riley et al. 1992). atorsinclude Chihuahuan raven (Corvus cryptoleucus), coyote, badger,striped skunk, ground squirrel, and bull Juvenilesurvival snake(Pituaphis melanoleucus, Davis et al. 1979, Survivalof juveniles has been diff1cultto assess Haukos 1988, Haukosand Broda 1989, Giesen 1998). becauseof problemsin obtainingaccurate information Nest success is positivelycorrelated with the presenceof abouttheir fate, particularlyduring the first 2 weeks after relativelythick grasscover (Riley 1978, Wisdom 1980, hatch(Ammann 1957, Christenson1970, Hillmanand Riley et al. 1992). Consequently,drought and grazingby Jackson1973, Bowmanand Robel 1977). Nevertheless, livestockcan reducenest success (Riley 1978, Wisdom juvenile survivalhas the potentialto dramaticallyaffect 1980, Merchant1982, Haukosand Smith 1989). populationviability (Peterson and Silvy 1996). Because Primarypredators of lesser prairie-chickensinclude variationin juvenile survivalmay be correlatedwith Predationand prairiegrotlse * Schroederand Baydack 27
variationin habitatcharacteristics, increased attention has been directedtoward managing habitat to increasesur- vival of juveniles (Petersonand Silvy 1996, Edelmannet al. 1998). Survivalof breeding-agebirds Annualsurvival of breeding-agebirds usually is not variableenough to permitcomparisons among different habitatsor amongareas with differentdensities of preda- tors (Schroederand Robb 1993, Connellyet al. 1998, Giesen 1998, Schroederet al. 1999). Consequently,adult survivalusually is consideredto be a relativelyunman- ageableperiod of the life cycle. The primaryexception to this considerationhas been with manipulationof har- vest. For example,survival of female sage grousewas estimatedas 67% in a harvestedpopulation in Wyoming (June1963), 55% in a harvestedpopulation in Colorado (Zablan1993), and 75% in an unharvestedpopulation in Idaho(Connelly et al. 1994). Althoughharvest may have significanteffects on local populations(Ammann 1963, Taylorand Guthery1980, Crawfordand Lutz 1985, Marksand Marks1987), harvestrates generally are con- sideredto be small when comparedwith predationrates (Dalkeet al. 1963, Hamerstromand Hamerstrom1973, Displayingmale sage grouse on lek in north-centralColorado. Male prairiegrouse may be vulnerableto predationduring the breedingsea- Hillmanand Jackson 1973, Johnsonand Braun1999). son. Photoby R. E. Bennetts.
Predationand population viability a population(Hannon 1986). Increasesin population Predationof prairiegrouse is often consideredto be a densitymay increasethe likelihoodof dispersaland, con- ramiElcationof habitatquality and distribution,popula- versely,may reducethe opportunitiesfor recruitment tion pressureand density,or predatorbehavior and (Keppie 1979). Subordinatemale greaterprairie-chick- dynamics(Christisen 1969, Miller and Graul1980, ens and lesser prairie-chickens,which are less likely to Taylorand Guthery1980, Braun1998). Inadequatequal- establishterritories, may have greaterrates of predation ity of habitatmay increasethe predationrisk for birds (Robel 1970, Campbell1972). Subordinatefemale attemptingto locate escape cover (Svedarsky1988, greaterprairie-chickens may be inhibitedby dominant Connellyet al. 1991, Riley et al. 1992, Gregget al. females from mating,resulting in delays in nestingwith 1994). Habitatdegradation that alters visibility at lek associateddeclines in productivity(Robel 1970, Robel sites may increasethe risk of predationto displaying and Ballard1974). Increaseddensities of sharp-tailed males (Hartzler1974, Baydackand Hein 1987, Berger grousenests may resultin an increasedrisk of predation and Baydack1993). The lack of adequatefeeding areas (Apa et al. 1997). may increasepredation by forcingbirds to feed longer,to The dynamicsof predatorpopulations are determined feed in riskierhabitats, or to travelfarther to feeding typicallyby the abundanceof theirprimary prey species, areas(Gregg et al. 1993, Fischeret al. 1996, Pyle and which usuallyare rodentsor lagomorphsrather than Crawford1996). Fragmentationof habitatmay increase grouse (Bumpet al. 1947, Angelstam1986, Marcstromet predationpressure by forcingnesting birds into marginal al. 1988, Myrberget1988). In situationswhere popula- habitats,by increasingtravel time throughunacceptable tions of the primaryprey species fluctuate,grouse num- habitats,and by increasingthe diversityand densityof bers can be influencedby the changingdensities of pred- predators(as shownfor Europeangrouse; Andren et al. atorsand the effects thatprey densitieshave on the 1985,Andren and Angelstam 1988, Bernard-Laurentand predator'sforaging behavior. For example,when preda- Magnani1994, Kurkiet al. 1997). tors are forcedto searchfor relativelyscarce prey, they Populationpressure is often consideredto be a density- are more likely to encountergrouse and grouse nests dependentmechanism controlling the size and growthof (Angelstam1983). 28 Wildlife Society Bulletis] 2001, 29(1):24-32
Predatormanagement In contrastto researchon NorthAmerican prairie grouse,predator control Habitatquality, prairie grouse density, and predator researchhas been relatively commonfor the numbersare all manageableto a certainextent. For managementof Europeangrouse, includingblack grouse example,prairie grouse density can be reducedby har- (Tetrao tetrix), capercaillie(E urogallus), hazel grouse vest, which has the potentialto reducethe predationrate (Bonasa bonasia), and willow ptarmigan(Lagopus on the remainingbirds (Ellison 1991). However,because lagopus). Most predatorcontrols in Europehave been shown to the goal of predatormanagement usually is to increase increasenest success, juve- nile survival,and populationsize the numbersof the targetspecies, reducingprairie grouse (Parker1984, Marcstromet al. 1988, Baines densityis not a viable managementoption to manipulate 1990, Moss 1994), with rareexceptions (Korsch predatorpopulations. In contrast,manipulation of habitat 1984). Europeanmanagement plans for grouseconsistently qualityor predatornumbers has the potentialto decrease includereferences to preda- tor control(Hudson and predationrate and ultimatelyto increasegrouse popula- Rands 1988; Bergmannand Klaus tions (Hamerstromet al. 1957). 1994; Klausand Bergmann1994a). This is in contrastto NorthAmerican Most directefforts to controlpredator populations managementplans for prairie grouse,even endangered have been regionalin natureand not motivatedby a species, which rarelyinclude referencesto predatorcontrol desireto increasenumbers of prairiegrouse (Willis et al. (Taylorand Guthery1980, Morrow1986, Giesen and 1993). Consequently,there are few examplesof predator Connelly 1993, Westemeier and Gough 1999). controlsin which populationsof prairiegrouse were Thereare fundamentalreasons monitored. Battersonand Morse (1948) removedcom- for the differencesin grousemanagement in mon ravenson one sage grousearea in Oregonand left Europeand NorthAmerica. First, the small and isolatednature anotherarea as a control. They found a 3% nest success of remaininghabitat in Europeis used as a ratein the untreatedarea and 35% in the treatedarea. justificationfor interventionin the predator-preyrelationship Mammalianpredators (striped skunk, opossum, raccoon) (Korsch1984, Andrenand Angelstam1988, Hudson were controlledon an experimentalarea in the rangeof and Dobson 1995). Second, grouserestoration efforts in Europeusually the Attwater'sprairie-chicken (T c. attwateri, Lawrence dependon birdsraised in captivity 1982). The subsequentrate of success for artificialnests thatare extremelysusceptible to predation(Schroth 1991, was 82%on the removalarea and 33% on the areawhere Starlingl 991, Makinenet al. 1997, Merker1997). Third, predatorswere not controlled. Unfortunately,the small the financialbenefits of grousehunting in Europe numberof predatorcontrol experiments has left a sub- for the landownersincrease the pressureto use predatorcontrol stantialvoid in the informationnecessary to evaluatethe to increasethe numberof birdsavailable for viabilityof predatorcontrol as a managementtool. For harvest(Jenkins et al. 1964, Jensen 1970, Hudsonand Rands example,there is essentiallyno informationon the long- 1988, Moss 1994). Predatorcontrols are rarely termimpacts of predatorcontrols on the behavior,genet- recommendedfor North American ics, and abundanceof prairiegrouse. prairiegrouse, even for increasinglythreatened andendangered populations living in altered,isolated, or fragmentedhabitats. This is due to numerousfactors includingthe lack of informationabout the long-term consequencesof predatorcontrol, the relativelyhigh cost of predatorcontrol, the protectedstatus of manypotential predators,and concernsabout public attitudestoward predatorcontrol (Messmer et al. 1999). Predatorman- agementfor NorthAmerican prairie grouse generally has beenaddressed by manipulatinghabitat, because it is believedto be the most economical,efficient, and viable long-termstrategy to enhancepopulations of prairie grouse(Hamerstrom et al. 1957, Dalke et al. 1963, Giesenand Connelly 1993, Edelmannet al. 1998). All 4 species of prairiegrouse in NorthAmerica have populationsor subspeciesthat are eitherfederally listed orbeing consideredfor federallisting as threatenedor Badgerin southwesternWyoming. Badgersare common nest predators endangered(Connelly and Braun1997, Connellyet al. in portionsof the rangesof all prairiegrouse. 1998,Giesen 1998, Silvy et al. 1999, Westemeierand Preclationand prairiegrotlse * Scllroederand Baydack 29
aboutthe relationshipsbetween predators and grouse. We specificallythank K. E. Church,L. A. Robb, F. A. Servello,and an anonymousreviewer for comments aboutearlier drafts of the manuscript.This researchwas supportedby the WashingtonDepartment of Fish and Wildlife,Federal Aid in WildlifeRestoration, and the NaturalResources Institute at the Universityof Manitoba.
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the Kansas Academy of Science 79: 141-147. a remote sensing/GIS-Basedhabitat-explicit population model for BowMAN,T J. ANDR. J. ROBEL.1977. Brood break-up,dispersal, mobility, sage grouse (Centrocercus urophasianus). Idaho Forest, Wildlife and mortality of juvenile prairie chickens. Journal of Wildlife Man- and Range Experiment Station Technical Report 25, Moscow, USA. agement 41: 27-34. ELLISON,L.N. 1991. Shooting and compensatory mortality in BRAIJN,C. E. 1998. Sage grouse declines in western North America: tetraonids. Ornis Scandinavica 22: 229-240. what are the problems? Proceedings of the Western Association of FISCHER,R A., K. P. REESE,AND J. W. CONNELLY. 1996. An investigation on State Fish and Wildlife Agencies 78: 139-156. fire effects within xeric sage grouse habitat. Journal of Range Man- BRA[JN,C. E.,T. BRITT,AND R O. WALLESTAD.1977. Guidelines for mainte- agement 49: 194-198. nance of sage grouse habitats. Wildlife Society Bulletin 5:99-106. GIESEN,K. M. 1987. 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