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258 Is the sun rising for the prairie-chicken in Wisconsin . . .

Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

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Page 1: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

258

Is the sun rising for the prairie-chicken in Wisconsin . . .

Page 2: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

INTRODUCTION

The purpose of this paper is to re-port on the past and present status ofthe Greater Prairie-Chicken (Tympa-nuchus cupido pinnatus) in Wisconsin.Much of this paper is an update of thereport, Prairie Chickens & Grasslands:2000 and Beyond [PCG2&B] (Toepfer2003), funded and published by theSociety of Tympanuchus Cupido Pin-natus, Ltd. (STCP). Significant por-tions have been taken directly fromthat report and a proposal titled: APlan for the Genetic Restoration ofthe Greater Prairie-Chicken in Wis-consin, Capture, Release and Evalua-tion 2006–2010 written by the authorfor STCP (Toepfer 2005a). The his-tory of the Greater Prairie-Chicken inWisconsin is one of expansion, con-traction, and more contraction, and isa microcosm of the history of thespecies in North America. If theGreater Prairie-Chicken is to have afuture as a viable species in Wisconsinand in North America then range con-traction will have to cease and expan-

sion will have to occur—history willindeed have to repeat itself.

Originally native only to the south-ern prairies of Wisconsin, GreaterPrairie-Chickens responded to Euro-pean settlement in the 1800s and ex-panded their range throughout thestate as suitable habitat was created byagricultural and timbering practices(Schorger 1944). Grange (1948) indi-cated that as late as 1941 prairie chick-ens were still present in every countyin Wisconsin after which natural suc-cession and the expansion of modernagriculture drastically contracted theirrange. Today just five subpopulationsexist in five counties associated withthe grasslands of four Wildlife Man-agement Areas [WMAs] (Fig. 1) incentral Wisconsin (Toepfer 2003,Anonymous 1993, Robbins 1991,Hamerstrom and Hamerstrom 1973,Grange 1948).

The Buena Vista/Leola subpopula-tion has been maintained through themanagement of state owned and man-aged “scatter pattern” of permanentgrassland reserves. Most of these re-

259

Status and Management of the GreaterPrairie-Chicken in Wisconsin—2006

John E. Toepfer

Research ConsultantSociety of Tympanuchus Cupido Pinnatus, Ltd.

3755 Jackson Ave.Plover, WI 54467715. 342. 4071

[email protected]

Page 3: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

260 Status and Management of the Greater Prairie-Chicken in Wisconsin—2006

Figure 1. Contraction of Greater Prairie-Chicken Range Central Wisconsin, 1990, 2003, and 2006and Wildlife Management Areas Central Wisconsin, 2006. Updated from Toepfer (2003).

Page 4: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

serves (10,777 acres) were purchasedspecifically for Greater Prairie-Chick-ens as recommended by the Hamer-stroms and Mattson in A Guide toPrairie Chicken Management between1954 and 1969 (Berkhahn 1973).These lands were purchased by privateorganizations such as the WisconsinConservation League (WCL), STCP,Dane County Conservation League(DCCL), Wisconsin Society for Or-nithology (WSO), and also the state ofWisconsin Department of Natural Re-sources (WDNR). An additional 1,220acres have been purchased since 1969,mostly by the WDNR for a total of ap-proximately 14,000 acres of perma-nent grassland reserves on BuenaVista (12,000 acres) and Leola(1,860). A small subpopulation atGeorge W. Mead Wildlife Manage-ment Area has probably survived onlythrough compatible management forwaterfowl. The Paul Olson WMA andthe Northern Range subpopulationsare not managed for prairie-chickens(approximately 2,000 acres of re-serves) and grasslands still exist onprivate lands there only because thewet clay soils restrict intense agricul-ture and haying (Warnke 2004,Anonymous 1993).

In the “Guide to Prairie ChickenManagement,” Hamerstrom, Hamer-strom, and Mattson (1957) stated:“Grassland is of vital importance toprairie chickens, the keystone inprairie chicken ecology. . . . Whereverone looks, the answer is the same: tosave prairie chickens grasslands mustbe preserved and managed; there areno substitutes.” The loss of suitablegrassland habitat from most of theiroriginal range has resulted in the ex-tirpation of the species throughoutmuch of its range. Today only two

small isolated remnant GreaterPrairie-Chicken populations remaineast of the Mississippi River; one insouthern Illinois, and the second incentral Wisconsin.

In summarizing the current statusof the prairie grouse in North Amer-ica, Robel (2004) indicated that “anaggressive plan” needs to be imple-mented to reverse the downwardtrend if there are to be “large viablepopulations 50 years from now.” Hewas referring to populations with a“huntable surplus” not just a series of“minimum viable populations so birdwatchers can observe males cavortingon leks in the spring.” He emphasizedthat “unless aggressive managementprograms are implemented soon,”Sage Grouse and Lesser Prairie-Chick-ens will follow the path of the Attwa-ter’s Prairie-Chicken, and the GreaterPrairie-Chicken may not be far be-hind. Robel (2004) referring to Silvy’sremarks (Silvy et al. 2004) stated,“Simply put, prairie grouse requireprairie, lots of it.” The “lots of it” wasdefined as “millions of hectares.”

The current distribution of GreaterPrairie-Chickens in North America isassociated with grassland habitat thatexists by an accident of geography oras the result of the development ofpermanent grassland reservesthrough active management. The acci-dents of geography I refer to typicallyconsist of permanent grassland on pri-vate land and exist on the poorer soilsthat typically cannot be used for cropor hay production. When grazed,these lands have to be grazed carefullyso that residual cover remains forgrassland wildlife, especially prairie-chickens.

Prairie-chickens will not likely everbe associated with higher quality soils

The Passenger Pigeon, Vol. 69, No. 3, 2007 261

Page 5: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

because of their high value as crop-land so they will through default belimited to the poorer soils that at thistime are not profitable for agriculturalproduction. Schwartz (1945) put allthis in perspective over 60 years ago:“It is this grassland (permanent) whenproperly managed that furnishes theirnesting, roosting, and other require-ments. Thus the basic structure of thesoil, its fertility, and topography setthe pattern of land uses which in turndefines the present and future distri-bution and to a great extent the num-bers of the prairie-chicken inMissouri.”

Over fifty years ago Grange (1948)proposed: “If Wisconsin were to clas-sify and dedicate from 1,000,000 to5,000,000 acres of northern and cen-tral Wisconsin for the major land usepurpose of producing prairie chickensand sharptails and accompanied thiswith adequate management, the fu-ture of these species would be bright”.

CENSUS

Booming Ground Survey

The population information pre-sented in this paper was obtainedfrom annual booming ground surveysand cock counts conducted, coordi-nated, and/or compiled by various in-dividuals 1950–2006 (Toepfer 2003).My methods follow the survey proto-col established by Hamerstrom andHamerstrom (1973) where the num-ber of cocks per booming ground wasdetermined from at least three morn-ing counts per booming ground madejust before and during the hen/breed-ing peak in April. Cocks displaying bythemselves or singles were not in-cluded in the total by Hamerstrom

and Hamerstrom (1973) because bydefinition a single cock does not con-stitute a booming ground or “lek”(Toepfer 2004). Nor does a singlecount of cocks observed booming atone location necessarily constitute abooming ground.

It is the norm to get variation in thenumber of cocks on a boomingground on different days and at differ-ent times of the day especially “early”versus “late” morning counts. Hencefor many grounds the exact numberof cocks cannot be determined andrealistically and practically, one canonly get a range of the number of“regular” cocks. The Hamerstrom’s ac-counted for this by reporting a rangeof counts for each ground then pro-vided three numbers/ground for eachyear, a range, maximum, and averageof the range. The rounded average ofthe ranges was used to determine theannual changes in the number ofcocks. (See Hamerstrom and Hamer-strom 1973, Table 1:11). The differ-ence between the maximum andaverage number of cocks per year re-ported by Hamerstrom and Hamer-strom (1973) from 1950 to 1971 was15.6% with a range of 6.9% to 29.3%.

In an effort to maintain continuitywith the historical counts made from1950-71 by Hamerstrom and Hamer-strom (1973), and those of R. K. An-derson (Anderson and Toepfer, 1994,1999) from 1972–1993, the authorand assistants on behalf of STCP havesince 1996 conducted independentbooming ground surveys and cockcounts on the two main wildlife areas,Buena Vista and Paul Olson. Thesetwo areas contain over 78% of theprairie-chickens in Wisconsin. Wehave located booming grounds usingGPS equipment to provide more accu-

262 Status and Management of the Greater Prairie-Chicken in Wisconsin—2006

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rate locations than the forty-acre orquarter section locations used in thepast or by other surveys. The areas re-ported for range changes were deter-mined by measuring the size of apolygon created for the range by con-necting the outermost points (Mohr1947) created by the boominggrounds. The counts presented hereare consistent with the historicalcounts above and provide an ongoingoverview of population cycles andtrends as well as range contraction.

RESULTS AND DISCUSSION

Isolation and Range Contraction

The Greater Prairie-Chicken popu-lation in central Wisconsin has de-clined 30.6% (709–492 cocks) since1990 and their range has contractedby half from 2,202 to 1,077 squaremiles (Toepfer 2003). Since 2003 andwith the recent loss of the northern-most booming ground near Sports-man’s Lake (Michele Windsor,personal communication) and thebooming ground at Dewey Marsh(Dave Halfmann, personal communi-cation), the range has continued tocontract to the WMAs and in 2006 therange of booming grounds encom-passed only 663 square miles.

The booming grounds at Neilsville(Jonas, personal communication) aregone and the Dewey booming groundwhich had 16 cocks in 1993 disap-peared in 2003. Also gone is theSearles Cranberry ground (DavidHalfmann, personal communication),which had as many as 14 cocks, andthe Mosinee airport booming groundwhich disappeared in 1993 (TomMeier, personal communication). Fig-ure 1 shows the Greater Prairie-

Chicken range in central Wisconsin1990, 2003, and 2006.

As indicated above the prairie-chicken range in Wisconsin has con-tracted from birds present in everycounty as late as 1941, to five subpop-ulations in just five counties in centralWisconsin. These contemporary sub-populations are now associated withfour WDNR Wildlife ManagementAreas [Buena Vista/Leola and PaulOlson/Mead (Toepfer 2003, Ander-son and Toepfer 1999, Hamerstromand Hamerstrom 1973)] and with theprivate grasslands in the NorthernRange near Unity. The WisconsinGreater Prairie-Chicken population istotally isolated from the rest of theU.S. range and the nearest major pop-ulation is about 350 miles away innorthwestern Minnesota.

This contraction has not only oc-curred on a range-wide basis but alsoon all of the WMAs except Paul Olsonwhere the booming ground range ac-tually increased from 17 to 76 squaremiles 1962–2006. The boomingground range on Buena Vista has de-clined from a maximum of 72 squaremiles in 1951 to 37 square miles in2006. Range contraction has been es-pecially obvious on Leola WMA wherethe range has declined from 12 to 3square miles 1950–2006 and the Meaddeclined from 27 to 17 square miles1962–2006. The number of boominggrounds on Buena Vista has also de-clined with contraction from 44 in1951 to 25 in 2006. Today all of thebooming grounds occur on or withina quarter mile of the state managedpermanent grassland reserves. Boom-ing ground habitat has also changedwith 96% occurring on grasslandcover in 1972 in contrast to 2006where 48% were on plowed ground.

The Passenger Pigeon, Vol. 69, No. 3, 2007 263

Page 7: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

This change reflects the major loss ofgrassland on private land to plowedland.

The status of prairie-chicken in Wis-consin has reached the point wherethe subpopulations are now threat-ened, having become ecologically andgenetically isolated from each other(Johnson et al. 2003, 2004, Toepfer2003, Halfmann 2002, Toepfer 1988).This range contraction and isolationhave resulted in a decline in numbersand a 29% reduction in genetic diver-sity since the early 1950s (Bellinger2001, Bellinger et al. 2003). It was ini-tially recommended by Palkovacs et al.(2004) and Toepfer (2003) that birdsbe translocated into the Wisconsinpopulation to increase genetic diver-sity. A Wisconsin Conservation Genet-ics Advisory Committee subsequentlycorroborated these conclusions andindicated that “translocations are nec-essary for the long-term persistence ofthe Greater Prairie-Chickens in Wis-consin and given the overall reductionin genetic diversity throughout theWisconsin Greater Prairie-Chickenrange, interstate translocations shouldoccur as soon as feasible.” This com-mittee of geneticists recommendedthat 30 to 40 hens from outside Wis-consin be translocated into the statefor a number of years and that a con-trol group of radio-marked birds beestablished to monitor survival and re-production (Bouzat et al. 2005a,2005b). They also indicated “TheWDNR goal to acquire 15,000 acres ofhabitat over the next 10 years is in theright direction. However, it seems lim-ited given the decline in numbers ofGreater Prairie-Chickens over the lastdecades and the rates of habitat lossand fragmentation.”

The fact that prairie-chickens have

to be translocated into Wisconsinfrom the outside indicates that man-agement has not been successful andthat we as managers have failed tomaintain a viable population in thestate. The prairie-chicken in Wiscon-sin is currently considered a threat-ened species by the WDNR Bureau ofEndangered Resources (BER)(Anonymous 1979). However, basedon its isolation, small disconnectedrange, rapid range contraction, ge-netic state, and limited amount ofgrassland habitat, I believe it now war-rants endangered species status inWisconsin.

All of the contemporary GreaterPrairie-Chicken samples used for Wis-consin genetic analysis were collectedand provided by me throughPCG2&B. The historic samples ana-lyzed to compare genetic changesover time consisted of wings collectedby the Hamerstroms during the lastWisconsin hunting seasons 1952–54.Quite contrary to what has been re-ported, these wings were not justfound in their attic. The readershould know that they were very care-fully archived in sealed wine boxesand entrusted to the author. Thewings were cataloged by the author,hybrids and sharptails removed andindividual wings segregated to town-ship and range so they would be accu-rately classified and analyzedaccording to subpopulation by Dr.Peter Dunn and his students at UW-Milwaukee.

Population Trends

There have been booming groundcounts of Buena Vista and Paul Olsonreported by various individuals withUW-Stevens Point and the WDNR.

264 Status and Management of the Greater Prairie-Chicken in Wisconsin—2006

Page 8: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

These reports have not included anymethods and have provided only a sin-gle number of cocks per boomingground. In addition, likely because ofchanges in personnel and methods,these censuses have regularly missedbooming grounds. In 2000, threebooming grounds totaling 14 cockswere missed on Paul Olson. In 2001,ten booming grounds were missed;five on Buena Vista totaling 26 cocks,four on Paul Olson (8 cocks), and onein the Northern Range (3 cocks). In2003, five booming grounds totaling30 cocks were missed on the BuenaVista; two were major boominggrounds with 10 and 11 cocks. It hasrecently been determined that thereare no daily records for the boomingground cock counts provided byWDNR for Leola for at least the past10 years and hence these data areweak. The author has at one time oranother during the course ofPCG2&B personally visited everybooming ground in central Wiscon-sin. These discrepancies in census pro-tocol and missed booming groundsultimately fail to provide an accurateaccounting of the state of the prairie-chicken in Wisconsin and are whySTCP and the author have been con-ducting our own booming ground sur-veys and counts to maintainconsistency and accuracy.

The general trend since 1990 hasbeen for the Buena Vista to remainstable with an average of about 270cocks. This population is down only6.4% from 1990 but 52% below thelast peaks of 550 cocks in 1981 and1950 (Anderson and Toepfer 1999,Toepfer unpublished data). The MeadWMA population has declined 60.5%and Leola WMA 60% since 1990. TheNorthern Range population is all but

gone, declining just over 90% from ahigh of 134 cocks in 1991 to just 12 in2006 (Anderson and Toepfer 1999,Michele Windsor, Tom Meir, KenRosenthal, personal communication).The Paul Olson WMA, most of whichis unmanaged, has been the excep-tion. This subpopulation actually in-creased 20% between 1990 and 2006,and is the only population in Wiscon-sin with a statistically significant up-ward trend. Figures 2 and 3 showGreater Prairie-Chicken populationtrends in central Wisconsin for 1950-2006 and 1996–2006.

A series of Weighed Least SquaresLinear Regression lines for theGreater Prairie-Chicken subpopula-tions in central Wisconsin (Toepfer2005b) showed that the long-term(1950–2005) Buena Vista/Leola WMAstrend has been downward but not sig-nificantly (P=0.571). Paul Olson andMead WMAs trends also are down-ward during the common period1969–2005 but not significantly(P=0.409). Long-term the BuenaVista’s trend is upward but is not sig-nificant (P=0.718) and as reported hasto be considered stable. Both Leola(1950–2005) and Mead (1969–2005)have been in long-term decline andthis trend is significant for Leola(P=0.002) but not for Mead(P=0.222). Population trends forMcMillian and the Outlying Area(Northern Range) are separatelydownward (P= 0.001), and obviouslywhen combined are significantlydownward (P= 0.001), not positive asindicated by WDNR Warnke and Advi-sors (2004, Figure 7).

Ironically it is these least disturbedgrasslands in the Paul Olson WMA ofcentral Wisconsin, which are not man-aged for prairie-chickens, that have

The Passenger Pigeon, Vol. 69, No. 3, 2007 265

Page 9: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

been more successful than the inten-sively managed areas. From 1997 to2006 the Paul Olson WMA GreaterPrairie-Chicken population has in-creased eight out of ten years and theMead has increased or remained thesame seven times, while the intensivelymanaged core Buena Vista and LeolaWMAs have increased only five andfour of the last ten years respectively.Only 4 miles separate the Buena Vistaand Leola WMAs. The annual in-creases on Buena Vista/Leola since 1990 have never exceeded 18%(Low –23.1 to 17.7%) while PaulOlson/Mead have experienced in-creases as high as 45.8% and 66.1%(Range –40.5 to 66.1%). GreaterPrairie-Chicken populations in Min-nesota during this same period haveincreased by as much as 54.3%(Low–45.7%) (Wolfe 2004; Toepferunpublished data), the Illinois popu-

lation increased 56% in 2005 (ScottSimpson, personal communication),and the reestablished North Dakotapopulation doubled between 2001and 2002 even though 17 inches ofrain which fell in 24 hours in June2000 devastated that population(Toepfer 2003).

Since the largest increases of 62.5%and 71.4% in 1977 and 1978, annualincreases on the Buena Vista since1990 have not exceeded 20%. Andduring the last ten years the inten-sively managed Buena Vista and Leolasubpopulations have increased in suc-cessive years only once and not at all,respectively. The Paul Olson andMead subpopulations have increasedin successive years five times and infour successive years 1997–2000, sug-gesting a possible correlation withlocal management activities. Differ-ences in weather conditions are a pos-

266 Status and Management of the Greater Prairie-Chicken in Wisconsin—2006

Figure 2. Greater Prairie-Chicken population trends (cocks) by area Central Wisconsin, 1950–2006.

Page 10: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

sible explanation for these differencesbut Paul Olson is only 12 miles fromthe Buena Vista.

These differences in populationtrends and patterns suggest that re-cent management practices on BuenaVista/Leola are having a negative im-pact and may even be suppressing re-production and recruitment. In 1987there was one more cock counted onthe unmanaged Paul Olson thanBuena Vista (194 versus 193). This re-sulted in concerns being raised byHamerstroms and R. K. Anderson re-garding too much disturbance andclaims that the Buena Vista was beingover-managed. Others even withinWDNR have privately raised similarconcerns that were ignored. The im-pact of “too much” management be-came more obvious recently when the

grazing/mowing management regime

to control brush between 2004–06 re-

moved almost all of the cover in the

grassland reserves surrounding three

booming grounds (Meil, Zielinski,

and South Bertottoo). These three

grounds declined 97% from 27 cocks

in 2004 when cover was eliminated to

just two cocks in 2006 and the Meils

and Zielinski booming grounds disap-

peared altogether in 2006. This is in

total contrast to the overall popula-

tion on Buena Vista, which declined

only 5% from 2004 to 2006. The re-

sponse to this was a STCP meeting

with WDNR in August 2005 that re-

sulted in a moratorium of all manage-

ment disturbances (burning and

grazing) except for food plots and

spot brush mowing in the northeast-

The Passenger Pigeon, Vol. 69, No. 3, 2007 267

Figure 3. Greater Prairie-Chicken populations trends (cocks) by area Central Wisconsin, 1996-2006.

Page 11: Is the sun rising for the prairie-chicken in WisconsinIn the “Guide to Prairie Chicken Management,” Hamerstrom, Hamer-strom, and Mattson (1957) stated: “Grassland is of vital

ern half of the Buena Vista starting in2006.

Nesting

The fact that nest success varies be-tween management areas and de-creases with the intensity ofmanagement gives one cause for con-cern. The nesting information pre-sented here comes from Toepfer(2003). Nest success for the combinedmanagement areas showed that suc-cess was about 10% higher for the lessintensively managed areas; 58.9% for119 nests at Paul Olson and Mead ver-sus 49.0% for 251 nests for BuenaVista and Leola or a difference ofabout 10%. Success was lower for theareas with the most state-managedpermanent grassland reserves andmost active annual disturbanceregime (Toepfer 2003). Residual grasscover was present at all initial GreaterPrairie-Chicken nests and it is impor-tant in nest site selection and success.Residual cover takes time to accumu-late especially in the drier better-drained wetland soils on BuenaVista/Leola.

The Bottleneck

Bellinger et al. (2003) indicatedthat Wisconsin’s Greater Prairie-Chicken population experienced apopulation “bottleneck” in the 1950s.This is not accurate as this is when thedecline to the actual “bottleneck”began (Fig. 2). There have been twomajor population lows on the BuenaVista since 1950, one in 1961 (135cocks) and a second in 1969 (104cocks). There was also a low periodfrom 1959–1977 where the populationon the Buena Vista remained below250 cocks (range 104–234 cocks)

(Hamerstrom and Hamerstrom 1973,Anderson and Toepfer 1994, 1999).This long 19 year low period no doubtled to the decrease in genetic varia-tion documented in the contempo-rary populations (Bellinger et al.2003). The decline in genetic diversitywas also precipitated by the loss ofconnectivity between subpopulations,especially between Buena Vista/Leolaand Paul Olson/Mead, Halfmann(2002). Johnson et al. (2004) indi-cated that there was allelic/geneticexchange between Wisconsin subpop-ulations in the early 1950s but nottoday.

Cycles

Leopold (1931), Grange (1948),Keith (1963), and Hamerstrom andHamerstrom (1973) all felt thatprairie chickens populations werecyclic. In Figure 1 it is possible to de-tect four higher peaks approximatelyten years apart from 1972 to present.Three peaks can also be seen in theMinnesota population 1980–2003(Fig. 4). Whether the Wisconsin popu-lation is cyclic or not is moot becausefrom a population security viewpointone will be more concerned aboutmanagement keeping the populationlows as high as possible and well aboveany minimum number.

Minimum Population Size

How many prairie-chickens will beenough? The number of individualsrealistically necessary to offset geneticdrift and inbreeding in an isolatedpopulation varies and has been esti-mated at between 500–5,000 breedingindividuals (Franklin and Frankham1998). In prairie-chickens with theirlek breeding system where only 10%

268 Status and Management of the Greater Prairie-Chicken in Wisconsin–2006

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of the cocks breed (Robel 1970), theminimum number needed for main-taining 500 breeding birds would beabout 2,500 individuals (Walk 2004).

Minimum numbers can be danger-ous because they become “minimum”fixed goals for administrators andpoliticians that provide little securityfor a population. Total local recruit-ment failures have been documentedin Greater Prairie-Chicken popula-tions. In Minnesota in 1992 heavyrains caused significant declines of45–50% the following year, and in1996-97 the population declined byhalf due to severe winter weather andsubsequent flooding.

Catastrophic weather also occurredin North Dakota in 2000 when 17inches of rain fell in 24 hours result-ing in a 50% population decline in

2001. Real or optimal management se-curity, both genetic and catastrophic,for prairie-chickens will only be pro-vided with a population greater thanany calculated minimum number.

There are only two ways to managepopulations for a natural catastrophe:hope for good luck, or, through man-agement, spread the habitat and pop-ulation out over a large enough rangeso the whole population will not beexposed to a single devastating natu-ral or weather event (earthquake, tor-nado, blizzards, heavy rains,hurricanes, or prolonged drought) ordisease. Annual Greater Prairie-Chicken mortality is approximately50% so if a disease or weather catas-trophe occurred that eliminated ayear’s recruitment the populationwould decline by at least 50%, and if

The Passenger Pigeon, Vol. 69, No. 3, 2007 269

Figure 4. Greater Prairie-Chicken population trends (cocks) northwestern Minnesota andWisconsin 1981–2006.

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there were no production for twoyears in a row there would be a 75%decline, etc. Therefore, populationseven in excellent habitat could quicklygo well below any minimum in just ayear or two.

Population security for Wisconsin’sprairie-chickens will be accomplishedonly with more habitat and more birdsspread out in a connected populationover a larger area, like what we see inMinnesota, Kansas, South Dakota, andNebraska. Of these, Minnesota has byfar the smallest numbers of prairie-chickens and size of range (approxi-mately 2,500 square miles), howeverits long narrow north to south distri-bution (approximately 150 by 5–25miles) provides at least a degree ofprotection from catastrophic weathermoving east to west. The Minnesotapopulation also went through severalshort bottlenecks of approximately500 cocks (Svedarsky et al. 1997). Forreference, the Nebraska sandhills, oneof the strongholds for prairie-chickensin the U.S. is 20,000 square miles or12.1 million acres in size (Vodehnal1999). The Buena Vista by contrast isonly 73 square miles or 46,770 acres insize.

MANAGEMENT

The sportsmen of Wisconsin havealways been very supportive of theGreater Prarie-Chicken, as much ofthe funding for management of theWMAs especially for Buena Vista/Leola comes from federal PittmanRobertson funds paid for primarily bysportsmen. However, it is obvious thatthis effort has not been enough, norwill the current support level be

enough, to sustain a genetically viablepopulation.

Robel (1980) over twenty-five yearsago indicated, “Much research needsto be done before we can scientificallymanage prairie grouse populations.Quantitative aspects of habitat are im-portant and must continue to receivethe attention of biologists workingwith prairie grouse. Continued effortsmust be made to census grouse popu-lations accurately and/or precisely. Ofutmost importance, however is that re-search horizons must be expandedand efforts must be increased to initi-ate basic studies on grouse biologyand grouse-habitat relationships.” Hewarned, “that when remnant popula-tions exist, little basic biologically ori-ented research is initiated by stateagencies because of the low probabil-ity of the grouse populations develop-ing into a huntable resource.”Currently sportsmen and state wildlifemanagement agencies have more in-terest in species that are abundantand can easily be hunted such as theWild Turkey (Meleagris gallopavo),Ruffed Grouse (Bonasa umbellus), andthe non-native Ring-necked Pheasant(Phasianus colchicus).

Much of the history of prairie-chicken management in Wisconsinhas been outlined in other venues,most recently by Anderson andToepfer (1999). At this time far moresupport for grassland restoration andmanagement comes from federal pro-grams within the U.S. Department ofAgriculture (USDA), which arefunded by taxes from all citizens. Theover 36 million acres of grasslands inthe Conservation Reserve Program(CRP) that have benefited grasslandwildlife exists as a temporary scatterpattern of grassland intermixed with

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prime or better agricultural land. InWisconsin there were 666,844 acres ofland enrolled in CRP in 1997 (Anony-mous 1997) but very little within thecurrent Greater Prairie-Chickenrange. This grassland habitat willlikely last only so long as these landsare not economically viable for agri-cultural or energy production—prob-ably 10 or 20 years or until the GNP ofChina approaches that of the US andtheir demands for food and energy in-crease. Federal and state laws, andeven leases on “permanent” conserva-tion easements on private land can bequickly changed by legislation if thereis a perceived greater national eco-nomic need for food, energy, and/orjobs. We tend to forget that CRP is aconservation program to conserve soiland water, not a wildlife habitatrestoration program. So we here inWisconsin and nationwide have a lim-ited window of opportunity and obli-gation now to develop a national“permanent” grassland program com-parable to the national wetland pro-gram. History tells us that habitatsecurity for prairie-chickens and othergrassland wildlife unfortunately willonly be created through the establish-ment of permanent grassland re-serves.

One of the biggest obstacles tograssland conservation has been andwill continue to be the public’s per-ception that if it is not growing a crop,trees, or contains houses it is justwasteland. We have had it poundedinto us since we were very young thatplanting trees is conservation. Wetend to forget that the millions andmillions of acres of quality soils thatwe use to grow most of our crops werecreated by grasslands.

In the future the answer to success-

ful wildlife habitat management andespecially prairie grouse managementlies in the Hamerstroms’ concept ofecological patterning (EP) or a scatterpattern of essential/critical habitats;sage brush for Sage Grouse, wetlandand grassland for waterfowl, and for-est land for turkeys scattered throughan agricultural landscape. This is be-cause people along with wildlife haveto live on the same land and that anylandscape will have to provide food,water, cover, and energy for both. Thiswill be an inescapable paradox “ascomplete competitors cannot coexist.”Establishing large contiguous blocksof wildlife habitat, especially perma-nent grasslands necessary to sustain vi-able prairie-chicken populations, willnot occur often if at all.

The Hamerstrom et al. (1957) habi-tat model shows that minimum popu-lations of Greater Prairie-Chickens areassociated with 10–40% permanentgrassland with areas containing morethan 40% permanent grassland sup-porting increasingly larger popula-tions. Greater Prairie-Chickenpopulations occurred on a sustainablebasis in areas with a minimum of 33% grassland, were considered abun-dant only where grass comprised50–75% of the area, and had “low lin-gering” populations in 10–15% rela-tively undisturbed grassland. A prairie-chicken area should be no more than10–25% scattered woodland. Prairie-chickens require grass and openspace. They use trees for feeding andloafing but tend to avoid woodlandpatches. Booming grounds havemoved and/or been abandoned dueto woodland encroachment (Hamer-strom et al. 1957, Anderson 1969,Toepfer 1988). Likewise “new” boom-ing grounds have shown up when

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trees have been removed and moreopen space created (Toepfer 2003).The cutting of trees and creatinglarger treeless areas increased annualsurvival of cocks on one boomingground by 20% (Toepfer 2003). Thelatter information and its implicationsled to the cutting of almost all thelarge trees or potential raptor perches(Peterson 1979) on state managedland on the Buena Vista.

Prairie-chicken nesting, brood rear-ing, and year round night roostinghabitat on the Buena Vista/Leola areprovided almost exclusively by thepermanent grassland reserves. Exceptduring the summer (mid-May to mid-September) feeding occurs in foodplots and private agricultural fields ad-jacent to the permanent grassland re-serves managed by the state ofWisconsin (Toepfer 1988). The typicaldaily pattern for Greater Prairie-Chicken mid-September to late Aprilis to feed twice a day in the privateagricultural fields and fly to and fromnight roosts in the undisturbed tallergrass or grass/forb cover provided bythe undisturbed grassland reserves.The exception is the cocks which reg-ularly visit the booming grounds be-fore they feed during the morningduring autumn and winter and thenduring both the morning and eveningduring the breeding season. This pat-tern persists today and has been welldocumented in numerous othertelemetry studies (Robel et al. 1970,Toepfer and Eng 1988, Toepfer 1988)and documented in Wisconsin back toGrange (1948) and Hamerstorm andHamerstrom (1949). The movementsof prairie-chickens are best character-ized as being associated with the habi-tat within and surrounding a complexof booming grounds (Toepfer 2003).

The booming ground is the year-round center of prairie-chicken ecol-ogy; they nest, night roost, feed, andraise young all within a mile of abooming ground. What all this meansis that the booming grounds and habi-tat surrounding them should be thefocal point of management.

Not all grassland conservation pro-grams benefit prairie-chickens. TheGreater Prairie-Chicken was touted bythe state as the “flagship” species forthe Conservation Reserve Enhance-ment Program (CREP) in central Wis-consin (Anonymous 2004 and Warnke2004). Unfortunately, grasslands en-rolled in CREP are not highly benefi-cial to prairie-chickens because CREPrules required that the grasslands bewithin 1,000 feet of a riparian area,which in central Wisconsin meanstrees and shrubs. The presence oftrees and shrubs has been shown tonegatively affect nesting success(Keenlance 1998, McKee et al. 1998)and survival in prairie-chickens(Toepfer 2003). A review of over 150CREP parcels totaling over 4,000 acresenrolled within the central Wisconsinprairie-chicken range indicated thatthe average size of the parcels was 24acres and few of the parcels were nearexisting or historical boominggrounds. Hamerstrom et al. (1957) in-dicated that in order to benefitprairie-chickens 40 acres should bethe minimum size of a grassland re-serve; today it is believed that the min-imum parcel size should be 80 acres.

At this time 25% of the Buena VistaWMA consists of permanent grasslandreserves (Warnke 2004). However,based on the distribution of boominggrounds, the permanent grassland re-serves make up just over 50% of therange of booming grounds. In 2006,

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based on the current boomingground distribution, only about halfof the Buena Vista proper is occupiedby prairie-chickens in contrast to over90% occupancy in 1951. This highgrassland ratio on the Buena Vistamakes it good habitat for prairie-chickens and based on the Hamer-strom model they should be abundantand they are. However the BuenaVista as we have seen is too small andhas become a zoo population, or whatJohnsgard (2002) refers to as a “mu-seum population.” Sample and Moss-man (1997) provide no actual data orreferences but indicated that in theiropinion, the Buena Vista was an idealmanagement unit for grassland birds,yet it has proven too small to sustaingenetic diversity for the resident key-stone species, the Greater Prairie-Chicken. These grasslands provideideal habitat for grassland birds butonly during the breeding season, asmost of these birds are migratory andspend less than half of their lives onthe breeding range.

Bergerud and Gratson (1988) haveoften been quoted for their descrip-tion of the Greater Prairie-Chicken inWisconsin as “probably the most in-tensively managed grouse in NorthAmerica.” However this is very mis-leading, and should instead state thatthe prairie-chickens on the BuenaVista and Leola are the most inten-sively managed grouse in North Amer-ica, as over 78% of grassland reservesand almost all of the funding for man-agement occurs on the BuenaVista/Leola WMAs. As we are now wellaware neither the Buena Vista/Leolanor the Paul Olson/Mead WMAs werelarge enough to sustain populationswith a healthy genetic diversity and wenow have to translocate birds from

outside Wisconsin to increase geneticdiversity.

In recent years it has been indicatedby some Wisconsin biologists that “AGuide to Prairie Chicken Manage-ment” by Hamerstrom, Hamerstrom,and Mattson (1957) is outdated. It hasalso been implied that due to chang-ing land uses on Buena Vista moregrassland reserves will be necessary.The main concern was that theHamerstroms and Mattson recom-mended only 3,200 acres of grasslandand that the 40-acre parcels were toosmall. This is true, but the Hamer-stroms and Mattson emphasized that“flexibility was key” for any such planto be successful, and that “if the areashould be as thoroughly changed toplowed land as southern Wisconsinnow is, the plan will fail in its presentobjective unless it were modified.”One of these modifications was “toconsolidate into fewer, larger, self-sup-porting units with a smaller total pop-ulation.” And that is exactly what theHamerstroms and Mattson did. Theydesignated the necessary additionalgrassland reserves for purchasethrough STCP, DCCL, WSO, andWDNR while they were employees ofthe WDNR, and managing the BuenaVista and Leola. In the process theprairie-chicken in Wisconsin was tem-porarily saved from extinction. Onceone such grassland complex was de-veloped (Buena Vista) then it wouldbe necessary for management to ex-pand into other areas such as the PaulOlson, Mead, and outlying areas to in-crease numbers to provide securityand sustain connectivity.

When the recommendations ofHamerstrom and Hamerstrom (1973)to “immediately” expand prairie-chicken management into the outly-

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ing areas away from BuenaVista/Leola were not implemented, it,by default, established a core manage-ment philosophy for the prairie-chicken in Wisconsin. This coremanagement philosophy of the focusof management on one primary areabecame further entrenched in the1990s when a proposal was preparedto establish a Habitat Restoration Area(HRA) with 20,600 acres of grasslandreserves to make up 25% of theNorthern Range (Stetsonville; 4,000acres, Unity: 7,000 acres) and Mead(9,600 acres) (Keir and Meier 1990).This HRA was not approved, nor was asmaller proposed HRA at Unity, asoutlined in the Wisconsin GPC Man-agement Plan (Anonymous, 1995).This same core management philoso-phy, which focused on one area or sev-eral areas with permanent grasslandreserves, was also used to manage or“save” prairie-chickens in Texas, Illi-nois, Missouri, Oklahoma, and NorthDakota. This approach obviously didnot work, as chickens have not faredwell in any of these states either.

One question that lingers is whywere the Hamerstrom’s recommenda-tions to expand management awayfrom the Buena Vista/Leola ignoredfor so many years once a grassland re-serve complex with over 10,000 acreswas established? There would seem tobe two main reasons. The first was thatthe population returned to 1950 levelsin 1981 and there were highs on allthe WMAs (Fig. 2) and everybody washappy that management seemed tohave been successful. However, this inhindsight was an unusually high, tem-porary cyclic peak. Second the man-agement plan in 1983 allocated noadditional acreages to Paul Olson soall of it was allocated to the core area

Buena Vista/Leola. Everyone was ledto believe that this allocation re-stricted land purchases only to theBuena Vista/Leola and that no landcould be purchased on Paul Olson un-less the Wisconsin Natural ResourcesBoard (NRB) changed the allocation.It should be noted that this manage-ment plan was actually for all threeWMAs because the total allocatedacreages were managed as a complexby one manager. Hence the acreageallocations for Buena Vista/Leolacould have been shifted to Paul Olsonadministratively without NRB ap-proval. Such a request was nevermade. This misled many, includingSTCP and the author, into thinkingfor years that land just could not bepurchased on Paul Olson unless themanagement plan was rewritten andapproved by NRB.

The Tale of Two States

A core management philosophyand fate of the Greater Prairie-Chicken in Wisconsin since 1973 is ap-parent in the land purchases andmanagement focus. From 1973–2003approximately 2,800 acres of grass-land reserves or less than 100 acresper year were purchased with almostall of it on the Buena Vista/Leola.Since this time, and especially since1981, the prairie-chiken population inWisconsin has declined and con-tracted to the grasslands on theWMAs.

Minnesota by contrast, throughleadership of Minnesota Departmentof Natural Resources (MDNR) andthe Minnesota Prairie Chicken Society(MPCS), during this same time framepurchased a total of at least 56,000acres of grassland habitat or 1,890

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acres/year within their prairie-chicken range. These lands were pur-chased by MDNR, The NatureConservancy (TNC), and US Fish andWildlife Service (USFWS). In additionapproximately 170,000 acres of grass-land habitat in CRP were added, asthe state of Minnesota, unlike Wiscon-sin, designated their prairie-chickenrange as a CRP priority area. All thisgrassland has resulted in an increasefrom about 500 cocks in 1985 to 3,200in 2003, a six-fold increase (Fig. 4).The Greater Prairie-Chicken popula-tion data for Minnesota come fromthe MPCS Prairie Chicken Inventory(Wolfe 2004) (Also see Svedarsky et al.1999). However the Minnesota popu-lation despite the large amount ofgrassland is not as secure as one wouldthink. This is because most of thegrassland habitat has been createdthrough CRP and if this program werelost or modified it would negatively in-fluence Greater Prairie-Chicken num-bers and distribution. The loss of CRPin Minnesota could create large gapsin grassland distribution especially be-tween Polk and Norman County andconnectivity would be lost creating asituation similar to what occurred inWisconsin that resulted in a signifi-cant loss in genetic diversity.

Hamerstrom et al. (1957) state“Prairie-chicken management is pri-marily grassland management, nograss, no chickens” and now, as wehave seen in Minnesota and otherstates, more grass means more chick-ens. Since 1973 Minnesota took toheart the Hamerstroms’ recommen-dations to expand management awayfrom the core areas and applied theconcept of ecological patterning ex-actly as intended. For now at least,Minnesota has become the real suc-

cess story in grassland managementfor prairie-chickens.

This is exactly what Hamerstrom etal. (1957) indicated was required herein Wisconsin because managementcould not purchase the whole BuenaVista for economic and political rea-sons. They felt that it would be just asecologically and cost effective to pur-chase and manage portions of the crit-ical limiting habitat type (permanentgrass, used for nesting, brood rearing,and night roosting) rather than thewhole area. With this done, manage-ment could then let the adjacent pri-vate land provide the rest (open spaceand food in the agricultural fields) atno cost to management. This is incontrast to purchasing the whole areaor one large block and leaving man-agement to provide everything forGreater Prairie- Chickens. When man-agement in Wisconsin failed to ex-pand grassland management into theoutlying areas it adapted a core man-agement philosophy by default whichfocused land purchases, funding, andmanagement activities on the BuenaVista/Leola so by the late 1970s, theGreater Prairie-Chicken in Wisconsinwas destined to become what Johns-gard (2002) referred to as a “mu-seum” population.

None of this has come as a surpriseto those who were paying attention,especially STCP, to declining prairie-chicken populations in the late 1980sin other states such as Illinois, Okla-homa, Missouri, and Texas and to theincreases that were occurring in Col-orado, Minnesota, and even Kansas,Nebraska, and South Dakota wheretens of thousands of acres of grasslandwere being added through CRP(Svedarsky et al. 2000). Some man-agers in Wisconsin in the early 1990s

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said everything in the state was finedespite concerns raised by the Hamer-stroms and R. K. Anderson about toomuch disturbance on the Buena Vista.In the early 1990s I met with the STCPCouncil and indicated that indeed theprairie-chicken in Wisconsin was introuble and the Hamerstroms and R.K. Anderson supported this con-tention. This concern was presentedto the WDNR, and STCP was criticizedat the time and were told that every-thing was fine and all that was neededwas to “fine tune” management on theBuena Vista and further research andconcerns regarding genetics were onlyof “academic” interest. This was thegeneral belief despite the transloca-tion occurring in Illinois at the sametime to offset inbreeding (Bouzat etal. 1998; Westemeier et al. 1998a,1999). I began working with STCP inApril 1996 and developed the re-search project Prairie Chickens andGrasslands; 2000 and Beyond to docu-ment the status of Greater Prairie-Chicken in Wisconsin (Toepfer 2003).

Pheasants and Prairie-Chickens

There are other areas in Wisconsinwith goals for the development of sig-nificant amounts of grassland habitatthat could support Greater Prairie-Chickens: The WDNR Glacial RidgeHabitat Restoration Area (38,000acres); the Western Prairie HabitatRestoration Area (20,000 acres); andthe Blue Mounds-BlanchardvillePrairie and Savanna Area. Althoughthese areas may, or will, have grass-lands, they have limited potential forprairie-chickens because they are iso-lated and have relatively high densitiesof Ring-necked Pheasants, which nega-tively impact prairie-chicken produc-

tion via nest parasitism (Kimmel 1988,Vance and Westemeier 1979, Weste-meier et al. 1998b, Toepfer et al.2005a). At certain densities hen pheas-ants lay their eggs in prairie-chickennests and pheasant eggs hatch at 23days and chickens at 25 days. In west-central Minnesota in 2006, 40% of 19chicken nests contained pheasant eggs,11 nests hatched and none fledgedchicken chicks. The pheasant has to beconsidered a serious threat to theGreater Prairie-Chickens in Wisconsinand Minnesota, as the competition be-tween the two species is not only bio-logical but also political. GreaterPrairie-Chickens are not hunted inWisconsin and are no longer a populargame bird nationally because they arenot abundant and are difficult to hunt.

One of the main factors that haskept pheasants from moving northinto the Wisconsin and Minnesotaprairie-chicken range is winters withdeep snow and cold. Such cleansingwinters dramatically reduce pheasantnumbers and distribution. However, ifwarming trends continue (resulting inopen mild winters, i.e., global warm-ing) we can expect the pheasant toquickly move northward into the Wis-consin and Minnesota prairie-chickenrange. In recent years pheasants haveshown up at the Mead, the furthestnorth WMA with prairie-chickens.And pheasant numbers and distribu-tion are expanding (Tom Meier, per-sonal communication). The mix ofundisturbed grassland and agriculturein the Wisconsin prairie-chickenrange, especially on Buena Vista/Leola, will make for ideal pheasanthabitat. The indiscriminate release ofpen-reared pheasants by anyone atany place, will serve only to com-pound these problem. There will al-

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ways be a very serious and real threatof released pen-reared pheasants in-troducing disease and parasites intograssland bird populations. Over time,increases in pheasant numbers in theprairie-chicken range will becomemore and more of a threat. A decisionwill have to be made whether to keeppheasants out of the prairie-chickenrange, or even eliminate them as hasbeen done in Illinois Greater Prairie-Chicken management areas (ScottSimpson, personal communication).At this time research needs to acquirea better understanding of the nicheseparation between prairie grouse andpheasants.

FUTURE

The biggest threats to prairie-chick-ens in Wisconsin as well as in theUnited States are apathy and igno-rance. Ecologically the main threatwill be “unchecked human populationgrowth” resulting in the loss of evenmore grassland habitat, which willonly lead to further fragmentationand loss of connectivity. If GreaterPrairie-Chickens are to have any fu-ture as a genetically viable populationin Wisconsin it will be accomplishedonly through the addition of tens ofthousands of acres of interconnected,undisturbed, grassland reserves in ascatter pattern within the open agri-cultural landscape between PaulOlson and the Mead, at the Mead,and to the northwest of the Mead inwhat is called the Northern Range.This open agricultural landscape con-tains approximately 1,670 squaremiles or 1.06 million acres.

The connectivity aspect or move-ment of individuals and hence gene

flow between these grassland patchesthat Hamerstrom and Hamerstrom(1973) promoted for prairie-chickenswill have to be created as a scatter pat-tern in a gerrymandering fashionacross or within existing agriculturallandscapes. Large contiguous blocksof wildlife habitat, especially grass-lands that can sustain a viable prairie-chicken population, will not likelyoccur in Wisconsin because of thechecker-boarding of private land own-ership. Hence, grasslands createdthrough purchase, easement, or gov-ernment programs will have to be de-veloped, one parcel at a time aswilling sellers or program participantsare located. Failing this, the state willhave to periodically translocate birdsfrom larger populations to maintaingenetic diversity.

Management Plans—PCG2&B and WDNR

The recent Wisconsin GreaterPrairie-Chicken Management Plan2004-2014 (Warnke and Advisors2004) is definitely a step in the rightdirection. This plan calls for the addi-tional purchase of 15,000 acres ofgrassland reserves to the central Wis-consin prairie-chicken range. How-ever, 1,500 acres of this total (10%) isto be added to the Buena Vista leaving13,500 for the rest of the range. Un-fortunately the acreage goal of thisplan will maintain prairie-chickens inWisconsin away from the Buena Vistaonly at the low end or at “marginal orlingering levels” (Hamerstrom et al.1957). In addition, A Feasibility Studyand Environmental Analysis for theCentral Wisconsin Grassland Conser-vation Area (Anonymous 2004) callsfor a “portion of the 13,500 acres to

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be leased to local farmers for hay, pas-ture land, and cropland to maintainthe agricultural landscape.” Thiscould take a significant amount ofgrassland habitat away from grasslandbirds further reducing the percentageof permanent grassland reserves awayfrom the Buena Vista.

These plans also indicated thatthere are 7,300 acres of grassland forprairie-chickens on the Mead WMA. Ifthis were the case then the ratio ofgrassland would be higher than onthe Buena Vista and the Mead shouldhave more than just 50 cocks. Theamount of grassland on Mead in factis only 1,910 acres (Tom Meier, per-sonal communications) and is in linewith the 50 cocks counted there in2006. Much of the so-called grasslandson Mead are wetland sedge of little orno value to prairie-chickens for nest-ing and brood rearing, and only usedduring the winter for night roostingcover.

Ironically the goals of the CentralWisconsin Grassland ConservationArea (CWGA) (Anonymous 2004) toadd 13,500 acres of “grassland habi-tat” could indeed be met on paperand even greatly benefit certain grass-land birds yet have little impact onprairie-chickens. In order to benefitprairie-chickens grassland habitatparcels will have to be of a minimumsize, located near existing or historicalbooming grounds, and will have to bemanaged in a manner as to maintainundisturbed grassland cover for nest-ing, brood rearing, and night roostingcover.

The recommendations made by mein PCG2B essentially would haveadded 31,000 acres and created threegrassland complexes similar to theBuena Vista within the current

prairie-chicken range, one each atPaul Olson, Mead, and the NorthernRange. This would have increased thetotal grassland reserves from 10 to18%. However, more important it in-creases the acreage of permanentgrassland reserves in Wisconsin forGreater Prairie-Chickens from 4% to21.3% in the areas away from theBuena Vista/Leola. At this pointBuena Vista/Leola would account foronly 28% of the grassland reserveswithin the prairie-chicken range. Thisis in contrast to the plan by Warnkeand Advisors (2004) to add 15,000acres to the central Wisconsin range,which increases the amount of perma-nent grassland reserves from 10 to17%, with about 46% of the reserveslocated on Buena Vista/Leola.

In all likelihood we will not be ableto save or create enough grasslandhabitat on Paul Olson because of therapid increase in housing develop-ments and the prairie-chicken popula-tion will continue to contract to theBuena Vista/Leola where 78% of thegrassland reserves currently exist.Others, this author, and many withinWDNR cannot envision a grasslandcorridor of any significance being de-veloped in the agricultural landscapenorthwest of the Buena Vista and thenacross the Wisconsin River to recon-nect to Paul Olson. This means theonly realistic way the BuenaVista/Leola and Paul Olson/Meadpopulations can be biologically recon-nected today is by increasing produc-tivity on Buena Vista by modifying andimproving management to increasedensity to a threshold point where in-dividuals (primarily young of the yearhens) will naturally disperse betweenthese areas. In northwestern Min-nesota when populations have in-

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creased for several years, young of theyear radio-marked hens have dis-persed 10–32 miles.

When the goal of only 15,000 acresin Warnke and Advisors (2004) wasquestioned by STCP and the author, aWDNR administrator indicated that itrepresented the most they as anagency felt they could accomplish,given their very real constraints ondollars and personnel. Hence thesegrassland acreage goals in Warnkeand Advisors (2004) are not based onthe biological needs of the GreaterPrairie-Chicken and are contrary towhat was recommended by thePrairie-Chicken Committee estab-lished by WDNR to develop a GreaterPrairie-Chicken recovery plan.

In the three years since the recom-mendations in PCG2&B were made,we have gained a better understand-ing of the number of prairie-chickensnecessary to sustain a viable popula-tion. As indicated above, any mini-mum population number should takeinto account 2-3 years of reproductivefailure. Thus in order to maintain theminimum population size of 2,500birds necessary to maintain the ge-netic health of an isolated GreaterPrairie-Chicken population (Walk2004), a target population of 10,000individuals would be needed to with-stand 2 years of reproductive failure,assuming an average annual survivalof 50% (Toepfer 2003), an average of10 cocks per booming ground, andapproximately 450 acres of grass-land/booming ground as per theBuena Vista grassland ratio. Consider-ing the above, over 225,000 acres ofbiologically interconnected grasslandreserves would be necessary to sustaingenetic diversity of Greater Prairie-Chickens in Wisconsin. Morrow et al.

(2004) indicated that there are cur-rently 70,000 acres of coastal grasslandavailable for Attwater Prairie-Chickenand that the long-term recovery habi-tat goal for them in Texas is the man-agement of 300,000 acres of costalprairie (Mike Morrow, personal com-munication).

However, on the positive side forWisconsin and prairie-chickens ingeneral one should keep in mind themessage from Walk (2004) regardingprairie-chickens in Illinois: “The re-covery potential of the GreaterPrairie-Chicken in Illinois is very goodand is limited only by human motiva-tion to provide adequate habitat.”

OTHER CONCERNS

The annual mortality from prairie-chicken collisions with electric trans-mission lines in Wisconsin along roadscan be relatively high at 6–14%(Toepfer 1988, 2003). The ideal solu-tion to reduce such mortality wouldbe to bury the wires. This was donesuccessfully through STCP in 2004along a three-mile stretch of CountyHighway F along the west edge of theBuena Vista. Alternatives to buryingthe wires include placing food plots sobirds do not have to negotiate suchwires when flying between knownnight roosting cover and feeding areas(Toepfer 1988), or placing markerson the wires to make them more visi-ble to the birds.

Recent research has documentedsignificant mortality (40%) on LesserPrairie-Chickens colliding with barbedwire fences (Wolfe, 2006). This hasraised concern by some here in Wis-consin about fence-related mortality.This type of mortality in Wisconsin for

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radio-marked birds was very low at0.6% or just two birds from 1996–2003(Toepfer 2003). In addition, most ofthe fence wire currently used on thestate grassland reserves is electricfence and the WDNR technicians thathave put out and taken in hundreds ofmiles of wire during the last ten yearscan recall finding only one deadchicken near these fences (KenRosenthal, personal communication).The explanation for the dramatic dif-ference in the mortality rate due tofences between the two species is quitesimple; Lesser Prairie-Chickens flymuch lower than Greater Prairie-Chickens, making them more suscep-tible to colliding with barbed wirefences (Don Wolfe, personal commu-nication).

For the past several years, concernhas been raised by local residents andbiologists that dog training (not thedog trials) on Buena Vista is becom-ing pervasive, and that dog trainershave begun to share information onwhere broods are located and thenwork multiple dogs on the hens withbroods. This could be controlledthrough permits and by putting cer-tain areas off limits and/or allowingtraining on alternate days (seeToepfer 2003).

Pesticides are a concern. However,at this time based on cholinesterasedepression, we know that GreaterPrairie-Chickens both in Wisconsinand Minnesota are exposed to organo-phosphate and carbamate insecticidessprayed in agricultural fields. Cockshave a higher exposure rate than hens(Toepfer 2003). This is because cocksspend more time in agricultural fieldsduring periods of aerial spraying whilevisiting their booming grounds (May-June) and during the peak of the sum-

mer molt (July–September). It is notuncommon for radioed birds to feedand then night roost in potato or soy-bean fields during the summer. Sev-eral radio-marked hens have takenolder broods and fledged chicks toagricultural fields in Wisconsin. InMinnesota it is common for hens withbroods to leave the CRP grasslandsand take half-grown chicks to nearbyadjacent soybean fields and raisethem. Some hens have actually movedbroods to soybean fields immediatelyafter hatching and successfullyfledged young. We have found no di-rect mortality as a result of pesticidesbeing sprayed on prairie-chickenswhile in agricultural fields.

There has been concern raisedabout disease, especially West Nilevirus. However all disease surveys todate (1998–present) show no diseaseproblems in Wisconsin, Minnesota,North Dakota, and Illinois. The WestNile virus is present but the prairie-chickens that have tested positive werehealthy and, unlike Sage Grouse, allhave survived the exposure (Paul-Murphy et al. 2003). The incidence ofinternal parasites today is less than re-ported in past studies (Toepfer 2003).

TRANSLOCATIONS

The goal of any future prairie-chicken translocation should be to in-crease genetic diversity (geneticrescue/infusion) or to expand the ex-isting range, not to establish new iso-lated populations. Lewis et al. (1961)indicated that minimal movementaway from the release sites is key to asuccessful translocation. Historicallyprairie grouse have been very difficultto establish as the birds tend to wan-

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der away from their release sites(Toepfer et al. 1990, Moe 1999,Toepfer 2003). The first knowntranslocations occurred in the 1860swhen Greater Prairie-Chickens weretranslocated to the east coast to sup-plement declining Heath Hen num-bers (Phillips 1928). Briefly, prairie-chickens translocated during thebreeding season into unoccupiedhabitat, especially hens, tend to wan-der away from the release sites withonly 25–33% establishing within twomiles of their release site (Toepfer1988, 2003, Toepfer et al. 1990). Sev-eral radio-marked hens released dur-ing April have ended up 50, 80, andeven 90 miles from their release sites.However, when released during thebreeding season into occupied habitat(supplementation) they are morelikely to stay, with about 50% establish-ing within 2 miles of their release siteas the translocated birds are attractedto resident birds (Toepfer et al.2005b). The development of the sum-mer translocation, releasing birds dur-ing the summer molt when they areless mobile, has greatly improved ourability to establish prairie-chickens invacant and occupied habitat with 80-90% establishing within 2 miles of therelease site (Toepfer 2003).

Prior to 2006, the only GreaterPrairie-Chicken translocations oc-curred in Wisconsin between 1973–77(Toepfer 1976, 1988) using pen-reared bird and wild birds in an at-tempt to reestablish prairie-chickensat the Crex Meadows WMA. This proj-ect was evaluated and published inToepfer (1988), Toepfer et al. (1990),and summarized by Evrard (2004). Allof the telemetry data in this laterpaper come from Toepfer (1988).Evrard (2004) indicated that this rein-

troduction project likely failed be-cause the releases were not spreadout. This is partially true, as we havelearned that one has to spread birdsout over a large area to reestablish aprairie-chicken population (Toepfer2003). However the release arealacked upland grass and its shrubgrass habitat was used more by Sharp-tailed Grouse (Toepfer 1988).

Grassland was initially inflated by adrought at Crex Meadows in 1976–77to the point where the impoundmentswere bone dry in the refuge or releasearea. This almost doubled the amountof grassland for nesting and especiallybrood rearing. The 20 wild prairie-chicken hens translocated from Min-nesota in 1977 fledged 19 young. From1977 to 1978 the Crex population ex-perienced its only increase from 17 to25 cocks and from this point on it de-clined to two cocks in 1989, three in1991, and none in 1992—13 years afterthe last birds were released (Evrard2004). The Sharp-tailed Grouse popu-lation during this same time frame in-creased from 16 cocks in 1974 to 84 in1989 and 115 in 1996 (Evrard et al.2000). The real value of these Wiscon-sin translocations was that we con-ducted the first experiments with thesummer translocation and docu-mented some of the problems of usingpen-reared birds.

Ammann (1957) indicated thatonce Sharp-tailed Grouse (Tympa-nuchus phasianellus) became moreabundant than Greater Prairie-Chick-ens the latter usually disappeared infive to six years, and Toepfer et al.(1990) indicated that at least 12 iso-lated areas that historically had bothspecies were taken over by sharptails.Crex Meadows and the surroundingarea is better suited for Sharp-tailed

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Grouse and should be managed forsharptails as they are in just as muchtrouble in Wisconsin as prairie-chick-ens. In hindsight the presence ofprairie-chickens only complicated themanagement situation at Crex Mead-ows and realistically just created an-other isolated prairie grousepopulation susceptible to inbreeding.

FORTY HENS OF DESTINY—GENETIC RESCUE

Between 14–25 September 2006forty adult hen Greater Prairie-Chick-ens, all from northwestern Minnesotaand all radio-marked, were released atfour locations on the Buena VistaMarsh to increase genetic diversity.This project was initially based on aproposal written by Peter Dunn UW-Milwaukee and me and presented tothe Genetics Panel and WDNR totranslocate Greater Prairie-Chickensto increase genetic diversity (Dunnand Toepfer 2004). This plan was up-dated in 2005 with A Plan For The Ge-netic Restoration Of The GreaterPrairie-Chicken in Wisconsin, Cap-ture, Release and Evaluation 2006-2010 (Toepfer 2005a) for STCP andsubmitted to WDNR for a permit inDecember 2005. This was done be-cause STCP indicated, as it had for theprevious two years, that it would fullyfund and conduct the entire translo-cation and evaluation effort.

These forty hens were initially cap-tured on booming grounds (Toepfer etal. 1988) in April 2006, radio-marked,and then recaptured by night-lightingduring mid-September. All the birdsflew well at release and all were radio-marked. Survival and establishment sofar has been very good. As of the end

of April 2007, six and half months post-release, 36 of the 40 hens can be ac-counted for: 28 (70%) were alive, eightwere dead (most fed upon by raptors),and four were missing, one since its re-lease. Of the 28 alive, 24 (85.7%) wereon the Buena Vista and 4 were onLeola. The latter four had moved 5.0to 11.2 miles from their respective re-lease sites. Overall, 63% were within 2miles of their respective release sites(mean, 2.8 miles, range 0–11.2 miles).All of the radio-marked birds havebeen observed moving, feeding, andnight roosting with resident radioedbirds. The development of the summerGreater Prairie-Chicken translocationprotocol and translocation of GreaterPrairie-Chickens from Minnesota toWisconsin in 2006 was conducted bySTCP, the author, assistants, and part-ners with funding from STCP andWDNR.

In anticipation of the translocationof hens from northwestern Minnesotawe trapped and radio-marked 142 res-ident Greater Prairie-Chickens on theBuena Vista to serve as a controlgroup in order to evaluate survivaland reproductive parameters to com-pare these with the translocated Min-nesota hens as outlined by Toepfer(2005a). Survival of resident controlbirds since mid-September has alsobeen good at 72% (adult cocks 65.4%,immature cocks 78.5%, adult hens73.1%, and immature hens 77.8%). Aswas done with the translocated birds,blood was taken for disease and ge-netic analysis. Blood for disease wasprovided to the WDNR and the ge-netic samples will be given to Dr. PeterDunn, UW-Milwaukee, for geneticanalysis (Dunn and Toepfer 2004). Atthe end of April 2007 there were 82radio-marked Greater Prairie-Chick-

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ens alive and accounted for on theBuena Vista (77) and Leola (5). Per-sonnel from WDNR and a graduatestudent from UW-Madison are now re-sponsible for the evaluation and havealso monitored the radio-markedbirds since January 2007 without sup-port from STCP.

The key to survival of transplantedbirds is successful establishment of in-dividuals, as extensive orientationmovements represent responses of in-dividuals, not groups, to being placedin an unfamiliar area. Thus, in atranslocation venture, most principlesof population dynamics are not oper-ating, and the population will not befunctional until the translocated indi-viduals establish themselves and re-produce (Toepfer et al. 1990). Thenext adjustment that the radio-marked hens from Minnesota had tomake occurred in mid-March 2007 asadult and immature hens began to es-tablish their territories. It is likely thatsome of the translocated hens will notbe able to establish themselves on theBuena Vista and will disperse to theoutlying areas to breed and nest.

I would sincerely hope everyone re-alizes that we have now foreverchanged the Greater Prairie-Chickenpopulation in Wisconsin. It will neverever be the same. The prairie-chickensthat the Hamerstroms, and our par-ents, and grandparents knew, and thatSTCP and the state of Wisconsin ini-tially attempted to save are gone for-ever, genetically extinct. This nowbecomes “a new beginning.” To putthis all in perspective, this is simplythe result of not creating enoughgrassland habitat to sustain a largeenough prairie-chicken population tomaintain a “healthy” genetic state.

However, that said, there is even a

much bigger fear in all this and that isthat conservationists and especiallythe general public will come to believethat all we have to do is translocate afew animals to fix the problems ofsmall declining populations. Translo-cating animals is a cheaper and easierfix than addressing the real problem,which is not enough habitat. We cannever lose sight of the fact that“enough” well-managed habitat will al-ways be the key factor in the survivalof any population. The recovery ofthe timber wolf (Canis lupus), BaldEagle (Haliaeetus leucocephalus), Ameri-can alligator (Alligator mississippiensis),and many other species was less com-plicated since it did not require therestoration of literally millions of acresof degraded habitat. We just had tostop killing them and remove DDT.

The translocation of a few prairie-chickens will indeed increase geneticdiversity temporarily, as we will soonsee in a paper about to be publishedon the results of the Greater Prairie-Chicken translocations the author ini-tiated in Illinois in 1992 and 1993(Westemeier et al. 1999, Rubin 1994,Toepfer 2003). However, withoutadding habitat, all this will only be atemporary easy fix to the symptomthat is “loss of genetic diversity” and itsassociated biological problems. Thisapproach can only result in the estab-lishment of a series of isolated GreaterPrairie-Chicken populations that willhave to be periodically supplementedforever with the release of birds fromlarger populations; essentially creatinga series of “zoo populations.” The onlylong-term solution will be to increasethe amount of grassland habitat andmaintain “connectivity” between exist-ing subpopulations as recommendedby Hamerstrom and Hamerstrom in

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1973. What Hamerstrom et al. (1957)stated almost 50 years ago warrants re-peating: “Grassland is of vital impor-tance to prairie chickens, the keystonein prairie-chicken ecology. . . Wher-ever one looks, the answer is the same:to save the prairie-chicken, grasslandsmust be preserved and managed forthem. There are no substitutes.”

The current WDNR Greater Prairie-Chicken Management Plan ten-yeargrassland acquisition goals will onlyincrease the range wide permanentgrassland reserve ratios for all theareas (Buena Vista, Leola, Paul Olson,Mead, and the Northern Range) from10 to 17%. The Buena Vista/Leolaarea will still contain 46% of the grass-land reserves dedicated to GreaterPrairie-Chickens in Wisconsin. In theareas away from the historical coremanagement area (Buena Vista/Leola) acquisition goals will only in-crease the ratio of grassland reservesfrom 4 to 13%. So at best, what we canexpect with the future addition ofthese 15,000 acres of grassland re-serves, at a cost of 15 to 25 milliondollars, a “low lingering” GreaterPrairie-Chicken population (Hamer-strom et al. 1957) in Wisconsin. Thissmall population however will have tobe periodically supplemented with re-leases from out-of-state birds fromlarger populations forever.

One only needs to review the con-servation status of grouse in Europe tosee where we are truly headed in Wis-consin and North America. In Europethe status of grouse has reached thepoint where the strategy from now onwill be to prevent some of the smallisolated populations from just goingextinct (Storch 2000). This will meanconservation genetics via transloca-tions similar to what was done in 1992

in Illinois and what STCP started lastSeptember in Wisconsin.

To the plight of the prairie-chickenwe must add all the prairie grouse andassociated grassland species but inWisconsin I would be remiss if I didnot mention the Sharp-tailed Grousewhich is currently just as threatened asthe Greater Prairie-Chicken, both eco-logically and genetically, and is head-ing into the “shadows” despiteapproximately 60,000 acres of federaland state managed fragmented “habi-tat.”

Unless there are dramatic changesin land use or a Herculean effort tosave and add 100,000 acres of grass-land in the “outlying areas” in centralWisconsin the current Greater Prairie-Chicken range will continue to con-tract like a vise to the BuenaVista/Leola. It is here where theGreater Prairie-Chicken will likelymake its last stand in Wisconsin, andsurvive, but only as an isolated rem-nant “zoo” population. Such a smallisolated population of only 500 birdswith a 125 square mile range of boom-ing grounds will be highly susceptibleto natural catastrophes and will onceagain have to be periodically supple-mented with the release of birds fromlarger populations to maintain ahealthy genetic diversity. Hence themost valuable prairie-chickens to thepeople of Wisconsin in 2006 nolonger reside on the Buena Vista butin Kansas, Nebraska, South Dakota,and Minnesota. To say anything elsewould just serve to mislead the public.

“A little repentance just before aspecies goes over the brink is enoughto make us feel virtuous. When aspecies is gone we have a good cry andrepeat the performance”—AldoLeopold.

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ACKNOWLEDGMENTS

I am grateful to the Society of Tym-panuchus Cupido Pinnatus, Ltd. forsupport both financial and political.Special thanks to current and pastpresidents Bernard Westfahl and RussSchallert, and especially Greg Septon,Chairman of the Projects and Re-search Committee for STCP, for themany hours of work on “Prairie Chick-ens & Grasslands: 2000 and Beyond.”Many groups (MPCS, MNDNR,NDGF, TNC, USFWS, WDNR) andcountless individuals have assistedwith field work in Wisconsin, Min-nesota, Nebraska, South Dakota,Texas, and North Dakota as STCP Fel-lows but a special note of thanks isdue to David Halfmann, Jacob Stich,and Mike Morrow for assisting innight-lighting hens in Minnesota fortranslocation to Wisconsin, and toDeann De La Ronde for her helptransporting these birds to Wisconsin.See also PCG2&B.

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A native of Wisconsin, John E. Toepferearned his B.S. and M.S. in 1972 and1976 at UW-Stevens Point where he devel-oped a life-long interest in prairie-chickensand a special friendship with Professor R.K. Anderson and Drs. Frederick andFrances Hamerstrom. As part of his PhDwork he made the first attempt to reintroduceprairie-chickens in Wisconsin at CrexMeadows Wildlife Area in northwest Wis-consin. Toepfer earned a PhD in BiologicalSciences at Montana State University withhis thesis on “The Ecology of the GreaterPrairie-Chicken as Related to Reintroduc-tions.” As a Professor at Little Hoop Com-munity College at Fort Totten IndianReservation in North Dakota he developedthe first Tribal College Native AmericanWildlife Program and was instrumental inthe development of the Inter-Tribal BisonCooperative. Since 1996, Toepfer has servedas Research Consultant with the Society ofTympanuchus Cupido Pinnatus, Ltd.(STCP). To address their concerns, and atthe request of STCP Council members, re-garding the future of prairie-chickens inWisconsin, he developed and carried outSTCP’s field research project: Prairie Chick-ens & Grasslands: 2000 and Beyond.Toepfer serves on the Attwater’s Prairie-Chicken Recovery Team and on the Boardof the North American Grouse Partnership.In 2003, he also received The HamerstromAward from the National Prairie GrouseTechnical Council for outstanding contribu-tions in the field of prairie grouse biology.

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or has it already set?

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Tree Swallow seen resting by Scott Franke.