Chapter FOURTEEN

Grasslands of Western Kansas, North of the Arkansas River*

David K. Dahlgren, Randy D. Rodgers, R. Dwayne Elmore, and Matthew R. Bain

Abstract. Lesser Prairie-Chickens (Tympanuchus pal- Prairie-Chickens in the region will be best accom- lidicinctus) occur in short-grass and mixed-grass plished by maintaining current habitat and provide prairies and associated grasslands restored management tools, guidelines, etc. and imple- through the Conservation Reserve Program (CRP) ment, recommend management practices, such north of the Arkansas River in Kansas. The Short- as grazing, prescribed fire, herbicide application, Grass Prairie/CRP Mosaic Ecoregion currently and prairie restoration, to improve habitat quality supports ~65% of the range-wide population of at smaller spatial scales. The ecoregion has been Lesser Prairie-Chickens. CRP lands provide impor- only recently occupied by substantial numbers of tant grassland habitats, especially nesting and Lesser Prairie-Chickens, and new data are needed brood-rearing areas for breeding Lesser Prairie- to develop conservation and management plans. Chickens. A combination of implementation of Current knowledge gaps include information on CRP grasslands at a landscape scale and favorable population demographics, limiting factors, habi- environmental conditions is thought to have led tat use and seasonal movements at various scales, to a significant increase in the occupied range habitat management techniques, energy develop- and population density of Lesser Prairie-Chickens ment impacts, and climate change. Additionally, in the ecoregion. Spring lek surveys since 1999 improved land use policies are needed for long- have documented the northern expansion of the term protection of habitat within the region, observed distribution of Lesser Prairie-Chickens beyond the typical duration of 10–15 years for in 2008, 2011, and 2012. An expanding distribu- CRP contracts. If conservation goals are met, the tion has led to the development of a contact zone ecoregion north of the Arkansas River in Kansas of sympatry between Lesser Prairie-Chickens and could continue to remain a stronghold for the Greater Prairie-Chickens (T. cupido). Hybridization Lesser Prairie-Chicken in the future. between the two congeneric species has been doc- umented and is currently estimated to occur at a Key Words: Conservation Reserve Program, habi- rate of ~5%. The potential effects of ­hybridization tat management, hybrid, Kansas, Short-Grass on the genetic structure of Lesser Prairie-Chickens Prairie/CRP Mosaic Ecoregion, sympatric range, are poorly understood. Conservation of Lesser Tympanuchus pallidicinctus.

* Dahlgren, D. K., R. D. Rodgers, R. D. Elmore, and M. R. Bain. 2016. Grasslands of Western Kansas, North of the Arkansas River. Pp. 259–279 in D. A. Haukos and C. W. Boal (editors), Ecology and conservation of Lesser Prairie-Chickens. Studies in Avian Biology (no. 48), CRC Press, Boca Raton, FL.

259 he Short-Grass Prairie/Conservation Before European settlement, the ecoregion was Reserve Program (CRP) Mosaic Ecoregion a landscape of generally flat short-grass prairie T lies on the eastern extent of the short-grass interspersed with mixed-grass prairie and small prairie and the transition to mixed-grass prairie tracts of sand sagebrush (Artemisia filifolia) prai- in western Kansas, north of the Arkansas River rie along some drainages and unique soil types. (Figure 14.1). The short-grass prairie is unique Playas or small ephemeral wetlands were histori- to the western edge of the Great Plains abutting cally a common feature across this area, espe- the eastern front range of the Rocky Mountains cially within the large expanse of flat “table” (Samson and Knopf 1996). Short-grass prairie lands (Haukos and Smith 1994). Following extends into the western quarter­ of Kansas, and European settlement, much of the extant prai- the area is often referred to as the northern High ries were cultivated and playas were incorporated Plains (Shiflet 1994). Mixed-grass prairie also into larger crop fields. Center-pivot irrigation occurs in the region, especially along the east- systems became widespread in the 1960s and ern edge of the distribution of Lesser Prairie- 1970s, allowing farmers to tap into the Ogallala Chickens (Tympanuchus pallidicinctus) in Kansas, but Aquifer to provide season-long water supply for also as inclusions further west (Kansas Native increased crop production. The use of ground- Plant Society 2014). McDonald et al. (2014) and water revolutionized agriculture in the area, even Van Pelt et al. (2013) have described the area as during periods of drought. During the 1960s to the Short-Grass Prairie/CRP Mosaic Ecoregion for the 1980s, large areas of prairies were again bro- Lesser Prairie-Chickens, and the name describes ken and plowed as a result of this advancement in the gradient of grassland types found in the agricultural technology (Waddell and Hanzlick ecoregion. 1978, Sexson 1980).

N Nebraska

Arkansas River LEPC distribution 2000 LEPC distribution 2008 LEPC distribution 2011 Current LEPC distribution States

Colorado Kansas Downloaded by [David Dahlgren] at 13:51 03 March 2016

New Mexico Oklahoma

Figure 14.1. Changes in the distribution of Lesser Prairie-Chickens (LEPC) in Kansas and Colorado since the early 2000s. Spring lek surveys were conducted by the Colorado Parks and Wildlife, Kansas Department of Wildlife, Parks, and Tourism, and partners and resulted in multiple northward changes in distribution boundaries during the past decade (also see Figure 14.3). Distributional limits and dates represent where and when biologists officially moved the known boundar- ies for the species distribution, but not necessarily how Lesser Prairie-Chicken were using the landscape in real time.

260 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal Plant cover and vegetative structure of native Kansas differed from other states by requir- short-grass prairie are not generally consid- ing seed mixes to resemble native mixed-grass ered suitable habitat for the life cycle of Lesser and tall-grass communities, which increased Prairie-Chickens (Hagen et al. 2004). For potential habitat, reduced landscape frag- example, Lesser Prairie-Chickens tend to select mentation, and resulted in increased popu- plant heights and visual obstruction for nesting lation abundance and occupancy of Lesser cover much greater than the habitats provided Prairie-Chickens (Rodgers 1999, Fields 2004, by typical short-grass communities (Hagen Rodgers and Hoffman 2005, Fields et al. 2006; et al. 2004). In other parts of their range, Lesser Figure 14.2). Prairie-Chickens often use shrub-dominated The objectives of our chapter are to (1) pro- landscapes that provide needed vegetation vide a synthesis of known ecological informa- structure in semiarid environments (Chapters tion regarding Lesser Prairie-Chickens and their 15 and 17, this volume). Historically, the sand habitat requirements in the short-grass, mixed- sagebrush and mixed-grass prairies along the grass, and CRP prairie complex of the ecoregion; larger drainages in this region may have pro- (2) consider distribution changes and northern vided habitat for Lesser Prairie-Chickens. It expansion in the ecoregion since the early 2000s; is unknown what proportion of the eastern (3) provide insights regarding the sympatric dis- short-grass and western mixed-grass regions tributions of Lesser and Greater Prairie-Chickens in Kansas were historically occupied by Lesser (Tympanuchus cupido) and evidence for hybridiza- Prairie-Chickens or how densities varied among tion; (4) provide management recommendations vegetation types (Hagen 2003). However, vol- specific to the ecoregion; and (5) describe the untary conversion of cropland into perennial most important research and information needs grass cover through the Conservation Reserve that are still needed for Lesser Prairie-Chickens Program of the U.S. Department of Agriculture within the Short-Grass Prairie/CRP Mosaic has improved habitat conditions. The state of Ecoregion.

Nebraska Current LEPC distribution

Kansas Downloaded by [David Dahlgren] at 13:51 03 March 2016

Figure 14.2. Vegetative land cover at the northern extent of the range of Lesser Prairie-Chickens (LEPC) in Kansas. Light gray represents grasslands, black represents Conservation Reserve Program (CRP) cover in 2005, and white represents cropland. The interspersion of CRP and grassland seems to be important to prairie chickens in this ecoregion. The area includes a contact zone where Lesser and Greater Prairie-Chickens are sympatric (see Figure 14.3).

Grasslands of Western Kansas, North of the Arkansas River 261 CHARACTERISTICS OF THE tall- and midgrass species, including the follow- GRASSLAND–CRP MOSAIC ing: big bluestem (Andropogon gerardii), little blue- stem, Indian grass (Sorghastrum nutans), switchgrass The Short-Grass Prairie/CRP Mosaic Ecoregion (Panicum virgatum), and sideoats grama, with occa- is in a rain shadow of the Rocky Mountains and sional additions of western wheatgrass, blue far removed from the moist influence of the grama, and buffalograss. Forbs were included in Gulf of Mexico. The environment is typically CRP seeding mixes for new contracts beginning semiarid with most precipitation falling as rain with the 1996 Federal Agriculture Improvement during the warm growing season. The average and Reform Act (Farm Bill), and most mixes annual precipitation ranges from 28 to 51 cm, commonly included a variety of native forbs: with amounts increasing west to east. Short-grass Maximillian sunflower Helianthus( maximiliani), prairie is dominated by grasses such as buffalo- purple prairie clover (Petalostemon pupureum), prai- grass (Buchloë dactyloides) and blue grama (Bouteloua rie coneflower Ratibida( columnifera), and Illinois ­gracilis). Midgrasses usually present in mixed-grass bundleflower Desmanthus( illinoensis). Introduced prairies include sideoats grama (B. curtipendula), forbs such as alfalfa (Medicago sativa), white sweet little bluestem (Schizachyrium scoparium), sand drop- clover (Melilotus alba), and yellow sweet clover (M. seed (Sporobolus cryptandrus), and western wheatgrass officinalis) were also permitted in the ecoregion (Pascopyrum smithii). Woody species are generally (Fields 2004). not abundant, although some small inclusions of The CRP seed mixes produced grasslands, sand sagebrush occur along a few drainage cor- which provided similar structure to mixed- and ridors (Küchler 1974). tall-grass prairies. The resultant stands were inter- Estimates can vary but ~60% of short-grass and spersed among native prairies and cropland, pro- mixed-grass prairies remain in North America viding nesting cover for Lesser Prairie-Chickens (Bragg and Steuter 1996, Samson and Knopf 1996, that was adjacent to shorter, more open grass- Weaver et al. 1996). However, within the Short- land habitats preferred for brood rearing (Fields Grass Prairie/CRP Mosaic Ecoregion, at least 73% 2004, Hagen et al. 2004; Figure 14.2). Both nest of the landscape has been converted to cropland, success and chick survival are critical factors in with ~7% of the area currently in CRP (M. Houts, the population growth of Lesser Prairie-Chickens Kansas Biological Survey, unpubl. data). Areas with (Hagen et al. 2009). Existing evidence suggests less productive soils, steeper slopes, or insuffi- the interactive effects of “newly” available habitat cient precipitation or groundwater resources have in CRP cover and favorable environmental condi- remained grasslands. Livestock grazing is the pre- tions likely contributed to significant expansion dominant land use for the remaining grasslands. and growth of Lesser Prairie-Chicken populations Much of the High Plains that is not dominated by in the ecoregion in the 1990s and early 2000s large expanses of cropland is currently a complex (Channell 2010). of grazed short- and mixed-grass prairies, seeded native grasses in CRP fields, and intermixed crop-

Downloaded by [David Dahlgren] at 13:51 03 March 2016 land (Figure 14.2). CHANGES IN THE DISTRIBUTION The Conservation Reserve Program is currently OF LESSER PRAIRIE-CHICKENS administered by the Farm Service Agency (FSA) of the U.S. Department of Agriculture (USDA). As recently as 2000, published distributions gen- The program was created in 1986 and provided erally delineated the Arkansas River as the north- funds for the conversion of marginal croplands ern extent for the contemporary range of Lesser to perennial grasslands (Figure 14.2). In this Prairie-Chickens, and associated with sand sage- particular region, nearly all of the seed mixes brush vegetation in Colorado and Kansas (Bailey for establishing CRP consisted of native grasses. and Niedrach 1965, Andrews and Righter 1992, However, the species of grass seeded were rarely Jensen et al. 2000; Figure 14.1). However, there the historically dominant grass species for a were numerous reports of Lesser Prairie-Chicken given ecological site. In Kansas, the USDA, in occurring in areas north of the Arkansas River cooperation with the state wildlife agency, pro- prior to 2000 (Jensen et al. 2000, Hagen 2003). vided technical guidance for grass seed mixes Lek surveys for Lesser Prairie-Chickens in the in CRP plantings primarily consisting of native ecoregion began in earnest from 1999 to 2004

262 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal (R. Rodgers, unpubl. data). The estimated distri- range-wide populations occurred in the ecore- bution of Lesser Prairie-Chickens was expanded gion (McDonald et al. 2014). Based on the best to incorporate more northern locations in 2008 available data, it appears that populations of Lesser (Figure 14.1). Some detected leks were only Prairie-Chickens have grown substantially in the attended by Lesser Prairie-Chickens, but mixed grasslands north of the Arkansas River since the leks of Lesser Prairie-Chickens and Greater Prairie- mid-1980s and were associated with introduc- Chickens were also located in an area of sympat- tion of CRP plantings (Rodgers 1999, Rodgers and ric distribution (Figure 14.3). From 2010 to 2013, Hoffman 2005). intensive lek searches along the northern Lesser Prairie-Chicken distribution boundary were com- SYMPATRIC RANGE AND HYBRIDIZATION pleted (M. Bain and D. Dahlgren, unpubl. data). Based on information gathered, the known bound- The grasslands north of the Arkansas River rep- ary of Lesser Prairie-Chicken distribution was resent the only portion of the range of Lesser again moved north in 2011 and 2012 (Figures 14.1 Prairie-Chickens where the species is sympatric and 14.3). with Greater Prairie-Chickens (Figure 14.3). Most In recent years, the highest densities of Lesser of the known leks in the ecoregion are either Prairie-Chickens within Kansas and range-wide Lesser Prairie-Chickens only or mixed-species were in the Short-Grass Prairie/CRP Mosaic leks (Figure 14.3). Hybridization has been docu- Ecoregion north of the Arkansas River in Kansas mented between Lesser Prairie-Chickens and (Pitman 2013, McDonald et al. 2014). In fact, Greater Prairie-Chickens, and hybrid vocaliza- during the recent range-wide survey and sub- tions have been found to be intermediate between sequent population analysis for Lesser Prairie- the two species (Bain and Farley 2002). Further, Chickens, an estimated 65% of the remaining some vocalizations in the area of sympatry are

N Nebraska

Current LEPC distribution States Prairie Chicken leks Species GRPC LEPC Mixed (both spp.)

Colorado Kansas Downloaded by [David Dahlgren] at 13:51 03 March 2016

New Mexico Oklahoma

Figure 14.3. Lek locations of Greater Prairie-Chickens (GRPC), Lesser Prairie-Chickens (LEPC), and mixed-species leks in a contact zone where both species are sympatric in west-central and northwest Kansas. Data on lek locations were provided by the Kansas Department of Wildlife, Parks, and Tourism. The map includes all historic lek locations, but some leks may not be currently active. These data represent only known locations, but do not account for variation in abundance or areas without surveys. Two mixed-species leks located north of the current range of Lesser Prairie-Chicken were occasional occurrences of a single Lesser Prairie-Chicken at leks of Greater Prairie-Chicken and were omitted from delineation of the current distributions.

Grasslands of Western Kansas, North of the Arkansas River 263 intermediate between hybrid and parental forms, attendance attributes, or display behaviors that suggesting possible backcrossing (i.e., F2 or F3 could minimize the occurrence of hybridization. offspring, M. Bain and D. Dahlgren, pers. obs.). The effects of genetic introgression on recruit- Intermediate vocalizations suggest that hybrids ment of Lesser Prairie-Chickens are unknown, can produce viable offspring, but this observa- as are the possible negative effects of deleterious tion has not been confirmed with molecular alleles, or the positive consequences of hybrid methods and the extent of genetic introgression vigor and adaptation to a dynamic landscape. remains unknown. Distinguishing a first-gen- Evidence at different spatial scales suggests eration hybrid (F1) or hybrid offspring (F2, F3) that some degree of lek segregation occurs and from either parent species due to morphology the hybridization rate is <5%, which has not or plumage characteristic is difficult. However, changed along lek survey routes in this ecore- like vocalizations, some plumage characteristics gion during the last decade (Bain 2002, Pitman seem to be intermediate between the two species 2013). Bain (2002) found that hybrid males had (Figure 14.4). high attendance rates, agonistic behavioral traits, If hybridization reduces fitness, isolating mech- and lek territories similar to males of the parent anisms are likely to evolve. However, postzy- species. However, copulation by hybrid males gotic isolation evolves slowly and low levels of was never detected despite a large number of hybridization may occur with no loss of fitness observation periods. Female choice might pre- (Grant and Grant 1996). Reproductive isola- vent further genetic introgression by avoiding tion between Lesser Prairie-Chicken and Greater hybrid males as mates. Therefore, it is likely that Prairie-Chicken appears to be weak in this region, hybrid genetics would introgress at a greater particularly gametic and postzygotic isolating rate through hybrid females. Characteristics mechanisms. However, prezygotic behavioral iso- involved in attracting females are likely the first lating mechanisms could include female choice, to diverge (Ellsworth et al. 1994). Acoustic mask- male competition, lek segregation and other lek ing of display vocalizations might discourage Downloaded by [David Dahlgren] at 13:51 03 March 2016

Figure 14.4. Hybrid Lesser-Greater Prairie-Chicken on a lek in western Kansas. The eye combs are characteristic of Lesser Prairie-Chicken and the air sacs are intermediate in color between the two parental species. Lower pictures are rump feathers of Lesser, hybrid, and Greater Prairie-Chickens (left to right). Note the contrast in the amount of dark and light brown for both upper and lower barring, with the hybrid intermediate between the two parental species.

264 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal mixed-species leks (Gibson et al. 1991), because both species in their sympatric range. Because the optimization of sound windows might have ecoregion currently supports the highest densi- contributed to the divergence of Tympanuchus ties of birds reported for Lesser Prairie-Chickens, (Sparling 1983). Relatively, minor differences >65% of the extant range-wide population may in display behaviors such as boom duration can be exposed to potential hybridization with affect antiphonal booming, other display char- Greater Prairie-Chickens (McDonald et al. 2014). acteristics, and lek segregation (Bain 2002). Hybridization has critical implications associ- Therefore, it is likely that even the most different ated with systematics, and the potential impacts traits, such as behaviors associated with breed- of genetic introgression or “dilution” of the ing displays, could have diverged very recently. genes of Lesser Prairie-Chickens. However, little The issues of hybridization and the possibility is known about the consequences of hybridiza- of viable offspring and potential backcrossing to tion between these two species or implications parent species warrant consideration, as does the for management or conservation measures in the degree of speciation between Tympanuchus species. future. Understanding hybridization between the The Lesser Prairie-Chicken was initially classified two species remains a future research need. as a unique species in 1885 (Ridgway 1885). Due The systematics literature suggests that Lesser to the fact that genetic information was not avail- Prairie-Chickens and Greater Prairie-Chickens able at the time, the distinction was clearly based are genetically similar and are species that have on morphological, behavioral, and distributional only recently diverged. However, we are not differentiation. Moreover, there are differing spe- suggesting that Lesser Prairie-Chickens are not cies definitions in the scientific literature for the unique or on significantly divergent evolution- two species of prairie chickens. Jones (1964) and ary trajectories, nor that any area that supports Crawford (1978) considered Lesser and Greater Tympanuchus spp. is not critical to their conserva- Prairie-Chickens to be separate species, but sug- tion. Rather, we argue that clear articulation is gested that reproductive isolation and species dis- needed regarding genetic and functional popu- tinction could be tested in a zone of sympatry. lation goals for both Lesser Prairie-Chickens and Aldrich and Duvall (1955) and Johnsgard (1983) Greater Prairie-Chickens. We also suggest that considered Lesser ­Prairie-Chicken to be a subspe- sympatric zones, distributional shifts, and pat- cies of Greater Prairie-Chicken, and the American terns of speciation and introgression need to be Ornithologists’ Union (1998) considered them better understood and considered as conserva- to be a single superspecies. More recent analysis tion efforts move forward and objectives for using modern genetic methodology amplifies the recovery must be clearly stated (Chapter 5, this lack of evidence for definitive genetic speciation volume). (Chapter 5, this volume). Luchinni et al. (2001) considered the Lesser Prairie-Chicken and Greater Prairie-Chicken “nominal” species; meaning ECOLOGICAL DRIVERS phenotypically, but not necessarily genetically

Downloaded by [David Dahlgren] at 13:51 03 March 2016 Drought different. The authors have stated that “Nominal species of Tympanuchus hybridize extensively where The northern High Plains is subject to periodic they are in contact…; their mtDNA haplotypes drought (Samson and Knopf 1996). In fact, it may are not fixed among species… and show shallow be more appropriate to describe climate patterns in genetic distances, suggesting that speciation has the region as regularly in drought conditions, but been recent and perhaps incomplete” (Luchinni periodically interrupted by wetter periods. Similar et al. 2001:159). Additionally, Oyler-McCance to most of the Great Plains, climatic conditions are et al. (2010) provide little evidence for genetic highly variable. Timing, frequency, and amount of divergence among Tympanuchus spp. If molecular precipitation, in relation to soil type, growing sea- data indicate that the Lesser Prairie-Chicken and son, and temperature, are major drivers of plant Greater Prairie-Chicken are not yet fully geneti- species composition, annual growth, and produc- cally divergent, current distribution boundaries tion (Holecheck et al. 2000), as well as associated might be considered less taxonomically impor- invertebrate and vertebrate animal communities. tant. For practical purposes, it is likely that the Most precipitation comes during the growing sea- application of conservation practices will benefit son, generally favoring warm-season grasses in

Grasslands of Western Kansas, North of the Arkansas River 265 both short-grass and mixed-grass communities Prairie-Chickens (Shaughnessy 2014). If landown- (Samson and Knopf 1996). Recent drought years ers choose to graze CRP fields with domestic live- of 2010–2013 have been moderate to severe in the stock, they are required to have a stocking rate of ecoregion. Residual grass and shrub cover associ- no greater than 75% of NRCS established rates. ated with CRP and lightly stocked grasslands are of Again, landowners must retain a stubble height that particular importance for Lesser Prairie-Chickens averages 25.4 cm (10 in.) within the known dis- during drought, as annual biomass production is tribution of Lesser Prairie-Chickens (Shaughnessy minimal and grazing pressure removes much of 2014). In recent drought years, most counties in the the cover outside of CRP fields. ecoregion have authorized emergency haying and grazing in multiple consecutive years (Shaughnessy 2014). In 2013, the Kansas FSA required that any Grazing portion of CRP fields within the range of Lesser Short-grass and mixed-grass prairies coevolved Prairie-Chickens can only be emergency hayed with periodic seasonal grazing by large ungu- or grazed in 1 of 3 consecutive years. In 2014, the lates such as American bison (Bison bison), prong- policy was adopted as part of the biological opinion horn (Antilocapra americana), elk (Cervus canadensis), in the federal listing of the Lesser Prairie-Chicken mule deer (Odocoileus hemionus), and white-tailed under the Endangered Species Act (Shaughnessy deer (O. virginianus). Other important herbivores 2014). have included smaller organisms such as insects, songbirds, and black-tailed prairie dogs (Cynomys Fire ludovicianus). Prairie dogs, in particular, played key roles in soil disturbance, affecting the distribution Fire is an important ecological process influenc- of large ungulates (Coppock et al. 1983, Krueger ing grassland systems in the Great Plains. Prior to 1986, Whicker and Detling 1988). Short-grass European settlement, Native Americans used fire prairie can be particularly resilient to grazing to influence grassland and animal communities pressure (Shiflet 1994). Current grazing prac- (Moore 1972, Frost 1998). Fire-return intervals have tices in the ecoregion are primarily cow-calf and been largely determined by climate, physiographic, stocker cattle operations. High grazing intensity edaphic, and vegetation conditions and resiliency has generally reduced or completely removed (Daubenmire 1968, Wright and Bailey 1982). Since midgrass species such as little bluestem and European settlement, natural fire has largely been sideoats grama from many ecological sites where suppressed across most of the Great Plains, includ- these plants were once common. However, in ing the Short-Grass Prairie/CRP Mosaic Ecoregion. recent years, some producers are utilizing more Both short-grass and mixed-grass prairies respond moderate stocking rates and intensities. Changes well to fire if moisture is available following a in grazing management are especially true for burning event (Frost 1998, Brockway et al. 2002). landowners involved in contracts with the Lesser In many native grasslands in this ecoregion, sideo- Prairie-Chicken Initiative (LPCI) administered ats grama may be the only species capable of pro- Downloaded by [David Dahlgren] at 13:51 03 March 2016 by the Natural Resources Conservation Service ducing adequate structure for nesting. If fire is (USDA NRCS 2013). Data are currently lacking on followed by intensive grazing or drought, sideoats the potential responses of Lesser Prairie-Chickens grama can take on a lower growing, sod-forming to different grazing practices in the ecoregion. structure and competition appears to favor buf- The CRP fields, based on policy of the FSA, are falograss and blue grama; thus, fire can dimin- generally not subject to grazing during the con- ish nesting habitat in some cases (M. Bain, pers. tract period. However, FSA can permit haying and obs.; Archer and Smeins 1991, McPherson 1995). grazing of CRP stands during emergency drought However, in stands of grass that have not received conditions within a county during a given grow- disturbance, such as some CRP fields, or where ing season, which allows producers to hay or woody plant invasion has occurred, fire can be graze their own CRP contracted fields. According a cost-efficient tool for improving nesting and to Kansas FSA policy, if landowners choose to brood-rearing habitats for Lesser Prairie-Chickens. hay, they must leave at least 50% of the field in When grasslands in the ecoregion were less frag- standing cover and a stubble height on average of mented, intact, contiguous, and comprised of a 25.4 cm (10 in.) within the distribution of Lesser greater proportion of ­midgrass species, fire likely

266 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal played an important role in maintaining habitat for plains, prairie chickens may have benefited from Lesser Prairie-Chickens. In recent years, there has waste grain as a novel food source (Chapter 2, this been an increased effort by multiple agencies and volume). Fields (2004) demonstrated that the use nongovernmental organizations to implement fire of forb-rich areas across vegetation types by broods for grassland and CRP management in this area. suggests that forbs and associated invertebrates However, landowners in the ecoregion are often were important food items for chicks as reported reluctant to use fire as a management tool due to in the literature from other regions (Taylor et al. fear of losing control of a prescribed fire, lack of 1980, Hagen et al. 2005). training, lack of equipment, and social or cultural constraints (Elmore et al. 2009). Lekking Lek sites in the region are predominantly in HABITAT REQUIREMENTS grasslands, but sometimes in croplands such Few scientific studies specifically addressing as winter wheat or fallow fields. Few leks are the ecology and habitat use of Lesser Prairie- found within CRP fields due to habitat condi- Chickens have been conducted in this ecoregion. tions with taller vegetation height and greater The knowledge gaps are largely due to the recent stem density. However, when the vegetation recognition of expanded distribution boundaries is removed, such as after an emergency hay- for the species in the ecoregion (Figure 14.1), as ing event, leks have been documented in these well as the recent realization of the large popula- areas (R. Plumb, Kansas State University, pers. tions of Lesser Prairie-Chickens in the ecoregion obs.). The region has Greater Prairie-Chicken, (McDonald et al. 2014). Many research needs exist Lesser Prairie-Chicken, and mixed-species leks, for this area, with relatively little scientific infor- with a species dominance gradient ranging from mation to date. Fields (2004) was the first wildlife Greater Prairie-Chickens in the north to Lesser ecologist to radio-mark individual Lesser Prairie- Prairie-Chickens in the south (Figure 14.3). Chickens (along with Greater Prairie-Chickens) Additionally, hybrid prairie chickens have been and monitor habitat use and survival in the Short- detected at multiple lek sites (Bain and Farely Grass Prairie/CRP Mosaic Ecoregion (see also 2002). Jarnevich and Laubhan (2011) used maxi- Fields et al. 2006). Researchers from Kansas State mum entropy modeling and known lek loca- University began a field study in 2013 with the tions to produce a probability map for lekking objectives of collecting new data on patterns of habitat of Lesser Prairie-Chickens that included habitat use, space use and movements, and sur- the Short-Grass Prairie/CRP Mosaic Ecoregion. vival (R. Plumb and D. Haukos, unpubl. data). Lek and prairie chicken densities can be rela- tively high in some areas, with as many as 12 leks along a 16.1 km route (10-mile) in an area Diet of 51.8 km2 (20 mile2), and up to 6.18 birds per No diet studies of Lesser Prairie-Chickens have km2 (16 birds per mile2, see Gove Route, Pitman Downloaded by [David Dahlgren] at 13:51 03 March 2016 been conducted in the ecoregion. Grasslands 2011). The Kansas Department of Wildlife, Parks, are highly fragmented by cropland throughout and Tourism (KDWPT) conducts lek surveys the ecoregion (Figure 14.2), and Lesser Prairie- each spring across Kansas, and currently has four Chicken use has been documented in croplands, lek routes within the ecoregion (Pitman 2013). especially during late fall and winter months A recent study was conducted in the northeast- (Fields 2004). Ingested contents in the digestive ern portion of the Lesser Prairie-Chicken range in tract of harvested birds have shown significant a four-county area of Kansas, including Graham, use of croplands for food sources during this sea- Rooks, Trego, and Ellis counties. V. Cikanek sonal period (Dahlgren et al., pers. obs.). The use (unpubl. data) investigated lek sites of both prairie of grains as a food source, especially during the chicken species in relation to surrounding land- potential resource bottleneck of winter, may be scape characteristics at three spatial scales: close an important diet consideration, but relative con- to the lek, a 1.5 km radius from leks, and a 3 km tributions to the diet of Lesser Prairie-Chickens radius from leks. At the smallest scale, lek sites are unknown. It has been suggested that when were further from paved roads and higher in ele- cropland first became part of the landscape on the vation than random locations. At the larger scales

Grasslands of Western Kansas, North of the Arkansas River 267 of 1.5–3 km, leks were associated with larger and Brood-Rearing Ecology more contiguous patches of grasslands (including Habitats used by broods of prairie chickens have CRP), less oil structure development, and more mainly been native grasslands, followed by CRP, CRP compared to random locations. and then rarely in cropland (Fields et al. 2006). Previous research demonstrated that grassland Nesting Ecology sites have relatively high forb cover and open areas for the movement of small chicks (Fields 2004). Nests of prairie chickens in the region have been While CRP provides nesting habitat, the vegeta- found in CRP (70%; n = 42 of 60), grassland tive structure can be too dense to provide good (27%; n = 16 of 60), and only rarely in cropland brood-rearing habitat in the absence of ecological (3%; n = 2 of 60, nest sites of Lesser and Greater disturbances from grazing, haying, or fire. CRP that Prairie-Chickens combined, Fields 2004). In the contains significant forb cover has been shown to vegetation types, nest sites were found in west- provide important brooding habitat (Fields 2004). ern wheatgrass, little bluestem, big bluestem, and From field observations, it appears that the transi- switchgrass (Fields 2004). CRP fields that were tional edge between CRP and grasslands can be an interseeded with forbs after grass establishment important interface for prairie chicken broods (R. or not treated were used for nest sites in greater Rodgers, pers. obs.). proportion than their relative availability (Fields In their 2-year study, Fields et al. (2006) reported 2004). Fields et al. (2006) reported apparent nest that only 7 of 25 (28%) monitored broods had success of 48.3% for nests where ≥1 egg hatched ≥1 chick survive more than 60 days after hatch. (n = 29 of 59). In this study, vegetation cover in Brood success from hatch to 60 days was much CRP, grassland, or cropland did not influence daily greater for broods attended by adult females (0.49, nest survival (Fields et al. 2006). Age of the nest SE = 0.19) compared to broods attended by sub- and seasonal timing had the greatest influence adult females (0.05, SE = 0.03, Fields et al. 2006). on daily nest survival. There was a progressive The 2-year study was conducted in years with decline in nest survival among early-, mid-, and above-average temperatures and one of the years late-season nests (Fields et al. 2006). Fields et al. was extremely dry during the nesting and brood- (2006) also found that increasing temperature led rearing period. to decreased nest survival. About 50% of com- plete nest losses were attributed to depredation by Water mammals (Fields 2004). Lek searches and monitoring were not initiated Lesser Prairie-Chickens have been documented until after CRP plantings were established across using free water sources in other ecoregions (Boal much of the ecoregion. However, anecdotal evi- et al. 2014, Grisham et al. 2014). However, it is dence suggests that Lesser Prairie-Chickens may unknown whether they benefit from free water, or have been present in the ecoregion, but abundance if they simply use these water sources when avail- and occupancy expanded following the initia- able. There are multiple natural and artificial water Downloaded by [David Dahlgren] at 13:51 03 March 2016 tion of CRP (Pitman 2013, McDonald et al. 2014). sources in the ecoregion. Livestock grazing and Many biologists have concluded that the popula- associated water facilities have increased the avail- tion increase was largely due to the increase in ability of surface water sources and distribution available nesting habitat in the form of CRP plant- across the landscape well beyond historic condi- ings that were conducted at a large landscape scale tions. Additionally, water facilities for wildlife or within the region (Figure 14.2). Nest survival has “wildlife guzzlers” have been constructed within been shown to be the most important vital rate CRP fields as part of the USDA contracts. Kansas impacting population dynamics of Lesser Prairie- NRCS guidelines specifically state that guzzlers Chickens (Hagen et al. 2009). However, in their within CRP fields are intended for upland game 2-year study with a relatively small sample of nests, birds. However, the impact of guzzlers on game Fields et al. (2006) did not find a difference in nest birds and other wildlife is poorly understood survival rates between vegetation types. The result (Boal et al. 2014). suggests that the increase in the quantity of nesting In a study during the summers of 2011 and habitat in the region may have been most influen- 2012, cameras associated with guzzlers were tial in increasing population abundance and range. used to obtain data for occupancy modeling of

268 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal DLCcovert.com 04–11–2012 16:39:54

Figure 14.5. Female Lesser Prairie-Chickens at a CRP guzzler site in Gove County, Kansas. The guzzler was monitored from late March to early May 2012 with a motion-sensing camera (DLC Covert). All photos during this pilot study ­indicated attendance by females only. The open ground-level water tank allowed access to prairie-chickens without entry under the guzzler roof. The design was unique compared to other monitored 23 guzzlers that had 50 gal (189.3 L) drums situated under the guzzler roof, where only one visit was recorded for a Lesser Prairie-Chicken. Each 50 gal drum had a small opening cut out with a wildlife ramp leading down into the water source.

upland game birds, mesopredators, and other (Figure 14.5). Notably, the guzzler with the most wildlife (B. Calderon, unpubl. data). The study prairie chicken use had a different design than area had multiple guzzler and paired nonguzzler others monitored with a ground-level open water sites in CRP fields in a five-county area of Kansas tank compared to all other guzzlers, which had in Logan, Gove, Trego, Ellis, and Russell coun- 50 gal (189.3 L) drums with small access openings ties. Sites spanned an east-to-west precipitation and wildlife escape ramps (Figure 14.5). The use gradient across the northern extent of the range of specifically designed water sources has been of Lesser Prairie-Chicken. During the study, no similarly documented in other ecoregion (Boal detections of Lesser or Greater Prairie-Chickens et al. 2014). Mesopredators were also detected at were recorded at any guzzler sites. However, guzzler sites during the pilot study. preliminary data indicated that raccoons (Procyon lotor) and other mesopredators had higher detec- POPULATION DYNAMICS tion and occupancy rates at guzzler compared to nonguzzler sites. Little information is available regarding popula- Downloaded by [David Dahlgren] at 13:51 03 March 2016 In a pilot study conducted during the spring tion dynamics of Lesser Prairie-Chickens within of 2012, motion-sensing cameras were placed at the Short-Grass Prairie/CRP Mosaic Ecoregion. guzzlers in CRP fields in Gove County in an area The KDWPT currently conducts spring lek surveys of relatively high lek densities of both species of for both species of prairie chickens in March– prairie chickens and in the same focal study area April along four 16.1 km routes (10-mile). Routes used by Fields (2004; D. Dahlgren, unpubl. data). in the region occur in Hodgeman, Ness, Gove, and Twenty-four cameras and guzzler locations were Logan counties (Pitman 2013; Chapter 4, this vol- used. Monitoring occurred from late March to ume). Each spring, biologists conduct two samples mid-May during nest initiation and start of the with 11 stops that are 1.6 km apart (1 mile) along incubation period. Prairie chickens were detected the route listening and looking for leks of prairie at only 2 of 24 guzzler sites during this period chicken. All detected leks within 1.6 km (1 mile) (8%) and only once at 1 of the 2 sites. The remain- on either side of the route are located and flushed ing site had multiple and regular detections at least once, and preferably twice during the sur- of female Lesser Prairie-Chickens until water vey period. Lek survey data provide both regional levels in the tank dropped below accessibility and statewide trends for populations of prairie

Grasslands of Western Kansas, North of the Arkansas River 269 chickens. For the Short-Grass Prairie/CRP Mosaic CONSERVATION AND MANAGEMENT OF Ecoregion, Lesser and Greater Prairie-Chickens LESSER PRAIRIE-CHICKENS IN GRASSLANDS are combined for trend information. The Logan NORTH OF THE ARKANSAS RIVER route was tested in 2012 and established in 2013, but not used here because of a lack of trend infor- Large blocks of native grasslands provide the most mation. Based on three of the four current routes, valuable habitat in other three ecoregions in the lek densities and numbers of birds per lek have range of the Lesser Prairie-Chicken. Populations of been relatively stable since the inception of these the species in the Short-Grass Prairie/CRP Mosaic routes in the early 2000s (Figure 14.6). However, Ecoregion, by necessity, depend more on the the trend has been slightly down in recent years, interspersion of native grassland and CRP tracts likely due to implications of region-wide drought (Figure 14.2). CRP grasslands provide suitable conditions on recruitment (Figure 14.6). nesting cover while native grasslands can often

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Figure 14.6. Lek survey trends for Lesser and Greater Prairie-Chickens with both species combined for routes in the Short- Grass Prairie/CRP Mosaic Ecoregion. Logan County route was not included due to establishment in 2013 and lack of trend information. Gove and Ness routes have an * to indicate that both Lesser and Greater Prairie-Chickens are observed during annual surveys along each route. Data were provided by the Kansas Department of Wildlife, Parks, and Tourism, and the most recent reports for lek surveys of prairie chickens are available at the agency website. (From Kansas Department of Wildlife, Parks, and Tourism 2014.)

270 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal offer better brood-rearing habitat (Fields 2004), be important for successful nesting. Small-scale allowing females to exploit the edges where patchiness might be more easily achieved in these two vegetation types occur adjacent to each endemic grasslands than in CRP, under current other. The addition of CRP grasslands has also CRP management policy. In summary, it is evi- been sufficient in many areas to shift the over- dent that Lesser Prairie-Chickens need adequate all ­grassland–cropland ratio from a landscape of concealment in a grassland stand that is neither mostly cropland to one that is mostly grassland, too dense nor too tall. In the Short-Grass Prairie/ likely benefiting Lesser Prairie-Chickens through- CRP Mosaic Ecoregion, bunchgrasses, such as lit- out their annual cycle. tle bluestem and sideoats grama, can best provide Management priorities have been created such grassland structure, particularly when com- within the Conservation Reserve Program that plimented with forbs. provides greater opportunity and incentives for Rodgers and Hoffman (2005) strongly dis- the establishment of CRP grasslands adjacent couraged the use of invasive exotic grasses such to or near extant native grasslands. Targeted as smooth brome (Bromus inermis) and Caucasian recruitment has been accomplished through the or yellow bluestem (Bothriochloa ischaemum) in establishment of carefully targeted Conservation the establishment of new stands of CRP fields. Priority Areas, which provide additional Environ­ Without a program policy that allows for graz- mental Benefit Index points for appropriate lands ing of CRP stands during the established nesting enrolled through the competitive General Signup season of birds (April 15–July 15), cool-season process of the CRP. The State Acres for Wildlife grasses such as western wheatgrass and smooth practice (CP38E) of the Continuous Signup pro- brome have shown a capacity to outcompete cess has also provided opportunity and incen- native warm-season species in CRP stands in this tives to target CRP grasslands toward key focal semiarid region, resulting in monocultures with areas where enrollments are most likely to little heterogeneous structure. If native warm-sea- benefit Lesser Prairie-Chickens. The Lesser son grasses have been heavily invaded by smooth Prairie-Chicken Range-wide Conservation Plan brome or western wheatgrass, the recommended developed by the Western Association of Fish and treatment is a November application of glypho- Wildlife Agencies (WAFWA) has also delineated sate (plus surfactant) active ingredient at 0.42– critical focal areas in the region and established 0.83 kg per ha (6–12 oz per acre) followed by 1–2 an avoidance, minimization, and mitigation pro- hard freezes at <-4°C when the temperature has cess whereby greater habitat protections, better warmed at least 12°C. The treatment has been habitat management, and new habitat establish- shown to be effective at killing smooth brome ment can best be targeted (Van Pelt et al. 2013). and would likely be effective on western wheat- Based on a broad assessment of where CRP grass while not harming the already-dormant grasslands had apparently benefited prairie grouse warm-season species (M. Bain, pers. obs.). populations in the western United States, Rodgers Prescribed fire is another useful tool in man- and Hoffman (2005) recommended desired CRP aging CRP stands for prairie chickens by provid- Downloaded by [David Dahlgren] at 13:51 03 March 2016 stand heights of 30–75 cm (12–30 in.) or roughly ing for grassland succession across the landscape, shin-to-thigh high. The authors also recommended reducing litter accumulation, and controlling that CRP stands be established using species that undesirable plant species. In this semiarid region, would produce a diverse, clumpy stand structure. new CRP stands with warm-season grasses gen- Such grasslands will provide three critical habi- erally do not reach maturity until at least the tat structural requirements for prairie chickens to fourth growing season after seeding (R. Rodgers be able to easily (1) hide and be concealed, (2) and M. Bain, pers. obs.). Excessive litter accu- move without obstruction, and (3) see approach- mulation typically will not occur until at least ing danger (G. Horak, pers. comm.). Fields (2004) 2 years after maturity is reached. Consequently, found that vegetation at the nest is generally much initial prescribed burns of new CRP stands typi- taller, and often twice as tall as the surrounding cally are not needed until at least six years after habitat conditions (e.g., 45.7 cm versus 22.9 cm), the stand was seeded, perhaps longer, depending and that vegetation at successful nests was often on stand density. Following the first burn, we taller than unsuccessful nests. The results sug- suggest that prescribed fire repeated at 4–6 year gest that habitat patchiness at small scales may intervals, depending on precipitation, would

Grasslands of Western Kansas, North of the Arkansas River 271 likely maintain suitable vegetation vigor and grazing (Van Pelt et al. 2013). The 3–5-year plans structure to satisfy the habitat needs of Lesser of the LPCI and the 5–10-year plans of WAFWA Prairie-Chickens. are relatively short term, with the frequency of The use of prescribed fire during the late sum- long-term adoption by landowners after con- mer months of mid-July to August has the poten- tract expiration unknown. In the arid region tial to benefit Lesser Prairie-Chickens as well. with common, prolonged drought, changes in Summer burns can reduce excessive height that species composition as a vegetative response to occurs in some CRP stands and have been shown management changes generally require decades to favor forb production in subsequent growing rather than years. Only long-term management seasons (Howe 1994), particularly if only portions programs over 15–30 years or in perpetuity can of a landscape are burned resulting in heteroge- ensure habitat improvements. Moderate stocking neity of vegetation structure and composition rates and rest rotations can be used to achieve at (Fuhlendorf and Engle 2004). The use of summer least a minimum amount and appropriate distri- burns also extends the potential days available in bution of quality nesting habitat. To efficiently the year for conducting prescribed fire and can target nesting habitat, management units that provide an enhanced margin of safety if surround- include grass species capable of producing struc- ing vegetation is still green and less flammable. ture for nesting must first be identified. The most Prescribed grazing can be an important CRP common species capable of producing suitable management tool. In addition to cool-season structure are sideoats grama, little bluestem, sand management, high-intensity grazing for short dropseed, and western wheatgrass, but some eco- duration during the dormant season may increase logical sites simply do not support these plant forb production and the vigor of warm-season species. The most effective grazing plans begin grasses, but this management tool needs more with identifying sites to target the development testing. Typical CRP grazing in July–November of nesting structure and then provide the stock- generally occurs too late in the growing season to ing rate or rest that best expedite the development effectively control cool-season species of grasses. of that structure. Grazing plans should comple- Dormant or early season high-intensity, short- ment producers’ long-term goals for their opera- term grazing may reduce current year nesting tions, ensuring benefits to both the producer and cover, but would at least improve nesting habitat wildlife habitat. Depending upon environmental in subsequent years if given adequate rest during conditions and existing species composition and the growing season. structure, deferment, rest, or light to moderate The habitat provided by much of the intact stocking rates with 25%–40% total utilization native grassland in this ecoregion has been should be used in these areas. Research is needed reduced in quality and quantity by high grazing to confirm and increase the efficacy of grazing intensity with little or no grazing rest for more recommendations. than a century. However, little bluestem, sideo- Haying CRP fields may also produce some ben- ats grama, and other grass species selected by efits to habitat similar to fire or grazing. However, Downloaded by [David Dahlgren] at 13:51 03 March 2016 Lesser Prairie-Chickens are sometimes still pres- the negative impact of long-term loss of cover, ent in such pastures but in a suppressed condition. especially if haying is followed by drought, may Reducing grazing pressure by using lighter stock- outweigh any benefits. Habitat loss may especially ing rates and periodic rest over time can improve be true if haying occurs at large scales. Generally, the quantity and quality of habitat available to we do not recommend haying to benefit Lesser Lesser Prairie-Chickens. Prairie-Chickens. Grassland grazing plans that address the More information is needed concerning the most common limiting factor of quality nest- impacts of energy development on Lesser Prairie- ing habitat are especially important in areas Chicken, but based on the negative impacts to lacking CRP. The USDA programs such as Lesser other species of prairie grouse (Hovick et al. Prairie-Chicken Initiative (LPCI) and the Lesser 2014), anthropogenic development should prob- Prairie-Chicken Range-wide Conservation Plan ably be minimized in priority Lesser Prairie- under the Western Association of Fish and Chicken habitat such as the WAFWA Lesser Wildlife Agencies (WAFWA) have provided land- Prairie-Chicken focal areas (Van Pelt et al. 2013). owners with incentive payments for prescribed Approaches for discouraging energy development

272 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal are highly controversial, but the WAFWA Lesser at a landscape scale. At a landscape scale, infor- Prairie-Chicken Range-wide Conservation Plan mation on habitat requirements is critical for effi- currently offers a mechanism for accomplish- cient and targeted implementation of programs ing this goal. Recent evidence suggests that the such as CRP. Conversely, with the reduction of established design of guzzlers designed to provide the CRP acreage cap and competitive commodity open water sources for Lesser Prairie-Chickens prices, knowing how much grassland (including is not effective and could be benefiting meso- CRP) is needed in a given area to support popula- predators that are detrimental to grassland birds tions of Lesser Prairie-Chickens and the optimal (B. Calderon, unpubl. data). Guzzlers are no lon- size and configuration of available patches will be ger incentivized by USDA CRP signup programs. critically important. If Lesser Prairie-Chickens truly depend on CRP to persist in the Short-Grass Prairie/CRP Mosaic INFORMATION NEEDS AND GAPS Ecoregion, the current habitat availability is in a As with the three other ecoregions for Lesser precarious situation where >50% of the range- Prairie-Chickens, the most urgent research needs wide population depends upon the existence are related to identifying limiting factors of popu- of a relatively short-term (10–15-year) program lations and associated habitats at multiple spatial subject to political support and a dynamic finan- scales and how known threats influence these fac- cial market driven by commodity prices. If a tors (U.S. Fish and Wildlife Service 2012). To our process to develop long-term solutions for main- knowledge, the only published population and taining or increasing CRP and the subsequent habitat related research for Lesser Prairie-Chickens mosaic of vegetative structure in this landscape specifically within the Short-Grass Prairie/CRP is not found, we could eventually lose the ecore- Mosaic Ecoregion are presented in Fields (2004) gion as a remaining stronghold for the species. and Fields et al. (2006). An example of needed Identifying the process and creating long-term information for our area of interest is given in solutions based on market-based or conservation Hagen et al. (2009), where population modeling programs will require knowledge of financial was conducted to identify vital rates that may be drivers that influence land use decisions, such as drivers of population growth. Once population annual payments that are competitive with cur- and habitat information becomes readily avail- rent market values. In addition, policy develop- able, the development of meaningful population, ment and subsequent communication must focus habitat recovery, and long-term conservation on maintaining or increasing landowner partici- objectives will become possible. Economic incen- pation in CRP. The potential threatened status of tives necessary to obtain these objectives need to Lesser Prairie-Chickens has created landowner be determined, as well as the most appropriate fear of participation in CRP. If policy is not devel- mechanisms to implement these incentives. For a oped to alleviate this fear and that policy is not wide-ranging species that occurs primarily on pri- clearly communicated to landowners, a threat- vate lands, the effectiveness of the strategy using ened status could have a net negative effect on Downloaded by [David Dahlgren] at 13:51 03 March 2016 voluntary conservation is of primary concern. populations of Lesser Prairie-Chickens in the Perhaps the most urgent information needs region. The effort must ultimately identify the in this region are associated with understand- level of incentives required to guarantee at least ing the effects of CRP and adjacent grasslands on a minimum area of CRP and other grasslands in population persistence, occupancy, and growth. the landscape for longer periods than current Anecdotal observations indicate that areas within programs, and up to 30 or 50 years to perpetu- the region with the highest Lesser Prairie-Chicken ity. Conservation planning could include iden- densities are comprised of ~60% grassland, 20% tifying an incentive-based path for landowners CRP, and 20% cropland. Quantification of these to transition current CRP into permanent graz- land use attributes at landscape scales and their ing lands. For example, provisions in the “Farm functional influence on demographic rates are Bill” and other conservation programs, such as urgently needed. To efficiently target and imple- Agricultural Land Easements and the WAFWA ment conservation, the minimum amount of nest- Lesser Prairie-Chicken Range-wide Conservation ing structure that is required in a given area must Plan (Van Pelt et al. 2013), could be used to be identified for local property management and secure long-term easements, which then could

Grasslands of Western Kansas, North of the Arkansas River 273 be modified and targeted to focus conservation management needs for individual fields or measures for Lesser Prairie-Chickens. If such habitat patches and provide suitable options for solutions are not found, the owners of CRP and landowners. First, research must be conducted native grassland that currently provide Lesser that identifies the most cost-efficient manage- Prairie-Chicken habitat may ultimately succumb ment practices that adequately address needs for to economic pressures to convert to alternative nesting habitat, survival, or other limiting fac- types of land use. tors within the management unit. For example, Further investigation into the trade-off between research and subsequent policy could be devel- spatial extent and duration of conservation prac- oped related to the following list of manage- tices is needed to develop sound strategies for ment information questions: Is flash grazing conservation, particularly related to adaptive with high-intensity stocking for a short duration management. In our ecoregion, long-term trends during the dormant season an effective means suggest range expansion and increasing densi- of reducing grass biomass and improving veg- ties remain possible for Lesser Prairie-Chickens. etative structure and plant density? What are the In such areas, conservation priorities that focus optimal stocking rates to maximize the quality on securing long-term maintenance of intact, of nesting and brood-rearing habitat? Is summer quality habitat, such as perpetual easements on burning the most effective management tool for CRP-grassland complexes, may be more critical mature stands of warm-season grasses with few than producing additional habitat. Once a certain forbs that do not provide adequate brood rear- threshold of habitat is protected over the long ing? In more arid portions of the region, when term, or participation and demand for long-term is the initial burn appropriate for restored CRP conservation has been met, a shift toward restor- grasslands in terms of stand age or plant struc- ing potential habitat may be warranted. In areas ture? For the range of existing stand types, what where little habitat is available to conserve, con- is the optimal time to burn? Would short-term, servation priorities should focus on restoring as intensive grazing or disking have a greater distur- much habitat as possible, as quickly as possible. bance response and subsequently be preferable Once a certain threshold of habitat is restored, it to burning? Under what environmental condi- may be more efficient to shift strategies and begin tions does burning reduce nesting habitat and long-term conservation easements for some of the for how long? If cool-season grass dominance most functional habitat. The greatest information is the primary threat to the stand, are spring needs associated with these issues are related to burning, glyphosate application in autumn, or the appropriate proportions of short- and long- other management practices effective at control? term habitat at landscape scales and triggers that At what level of dominance by smooth brome, might be useful in optimizing cost-efficiency and western wheatgrass, or other cool-season grasses effectiveness over time. is a prescribed burn during nesting period worth Energy development is increasing in the Short- the benefit of habitat restoration versus potential Grass Prairie/CRP Mosaic Ecoregion. Specifically, loss of production by Lesser Prairie-Chickens? Downloaded by [David Dahlgren] at 13:51 03 March 2016 oil and natural gas development is creating new Is potential nest loss by spring burning offset roads and associated infrastructure throughout by renesting or long-term benefits in increased prairie chicken habitat. Additionally, pressure production? For the range of existing stand to develop wind energy resources is likely to types, what is the most effective means of seed- increase in the region. Similar to other ecoregions bed preparation for interseeding with forbs— within Lesser Prairie-Chicken range, we currently disking, burning, spraying, or mowing? For lack a full understanding of the potential impacts established stands, are there any scenarios where that energy developments could have on popula- seedbed preparation is not necessary? What is the tions of Lesser Prairie-Chickens. optimum seed mixture and rate or seeding for Management prescriptions currently in place interseeding of legumes or other priority forb for Lesser Prairie-Chickens within our ecoregion species? We recognize that waiting for consider- need more experiential and research informa- ation of a laundry list of applied research needs is tion to help guide and improve the methods in not an option, and managers must move forward the future. Policy and resulting management based on the best available science and their practices must be flexible enough to prescribe own knowledge base from personal experience.

274 STUDIES IN AVIAN BIOLOGY NO. 48 Haukos and Boal However, a need for an urgent response does not for either species. Research should be designed preclude the need for research-based informa- to assist in the determination of whether genetic tion concerning management practices, espe- distinctness between Lesser Prairie-Chickens and cially if certain practices become popularized in Greater Prairie-Chickens should be a recovery management prescriptions. goal. Therefore, understanding hybridization Currently, livestock grazing is considered rates between these two species is an important a compatible use with habitat requirements research need. Vocalizations that are intermedi- (USFWS 2012). However, different grazing sys- ate to the parental species suggest that hybrids tems and stocking rates will have variable impacts. produce viable offspring; however, these obser- Identifying the most appropriate stocking rates vations have not been confirmed genetically. and grazing systems, including species of grazer, Reproductive isolation between Lesser and timing, and duration, that are most compatible Greater Prairie-Chickens appears to be weak in with requirements of Lesser Prairie-Chickens this region, but behavioral isolating mechanisms will be important to future conservation in this could include female choice, male competition, ecoregion. A focus on grazing would be espe- lek segregation, and other lek attendance attri- cially important if the management of CRP fields butes and display behaviors. In fact, within the could include livestock grazing as a future man- 96 leks described by Bain and Farley (2002), seg- agement practice. Although not specific to Lesser regation by species was greater than would be Prairie-Chickens, much guidance can be garnered predicted by chance alone. The effects of hybrid- from the existing literature on grazing impacts to ization on production by Lesser Prairie-Chickens, vegetation. extent, and direction of genetic introgression, and In addition to grazing, fire is another ecologi- other negative or positive effects are unknown cal driver that can be managed. Prescribed fire and require more research attention. has rarely been applied to arid grasslands in the Shifts in distribution and occupied range for region due to ongoing fire suppression and con- both Lesser and Greater Prairie-Chickens and a cerns of landowners, and consequently, effects subsequent increase in area of the sympatric zone of fire on habitats of Lesser Prairie-Chickens are will likely continue until new range limits are largely unknown. Prescribed fire on CRP lands reached or changes in land use, especially conver- is more common, and where undisturbed veg- sion of native grasslands or CRP to cropland, occur etation likely limits brood rearing, it is likely the at larger scales in this region. The dynamic nature most appropriate management tool. Additionally, of the distribution of Lesser Prairie-Chickens in where tree encroachment is an issue, prescribed relation to environmental conditions, particu- fire is often the most cost-effective method to larly in this region, clearly has been influenced maintain open grasslands. Information is needed by humans and suggests that historic distribution to determine the optimum timing and environ- distinctions may offer little toward recovery plan- mental conditions, and appropriate role of pre- ning. Rather, research needs to be developed that scribed fire in existing grassland fragments and identify current and projected functional popula- Downloaded by [David Dahlgren] at 13:51 03 March 2016 CRP for the purpose of Lesser Prairie-Chicken tions and the limiting effects of land use, climate conservation. change, or other factors in those areas. Therefore, Habitat requirements of prairie chickens understanding climate change and its potential within their sympatric range are likely to be impact on this species is warranted (Channell similar to allopatric populations, and habitat 2010; Chapter 12, this volume). Specifically, selection by Lesser Prairie-Chickens and Greater how will predicted increases in temperature and Prairie-Chickens is similar in the contact zone. evapotranspiration influence life history traits or However, if differences were detected, those demographic parameters? To help managers, cli- findings would likely shed light on important, mate models could be incorporated into recov- limiting factors related to habitat for each species. ery efforts for the Lesser Prairie-Chicken to help Research that detects similar habitat use for both direct resources to areas with a higher probabil- species would be just as insightful. Currently, lit- ity of persistence. Information on climate change tle is known about the potential role of hybridiza- and distribution shifts that includes potential tion between Lesser and Greater Prairie-Chickens hybridization with Greater Prairie-Chickens in or implications for future conservation measures northern reaches might help identify important

Grasslands of Western Kansas, North of the Arkansas River 275 recovery areas, perhaps beyond the recognized ACKNOWLEDGMENTS current distribution. We thank the two reviewers who helped to improve this chapter. We also appreciate our CONCLUSION two volume editors for their efforts herein. We The Short-Grass Prairie/CRP Mosaic Ecoregion thank M. Smith, KDWPT Farm Bill Biologist, for north of the Arkansas River in Kansas represents his expertise and contributions. We would like to a unique portion of the current distribution for thank all KDWPT personnel who participated in Lesser Prairie-Chickens in their five-state occu- lek surveys for prairie chickens and the reporting pied range. Based on current evidence, Lesser process over the years. We also thank the many Prairie-Chickens in this area use CRP and associ- landowners in this region who have participated ated grasslands to meet their seasonal life cycle in management and allowed access to private needs (Fields 2004, Fields et al. 2006; Figure 14.2). lands for the purpose of gathering new data on The introduction of CRP grasslands at landscape prairie chicken ecology. scales is believed to be the cause of population increases in this area over the past three decades, LITERATURE CITED but particularly in the late 1990s and early 2000s. However, it has been documented that Aldrich, J. W., and A. J. Duvall. 1955. Distribution of Lesser Prairie-Chickens occurred in this region, American gallinaceous game birds. Circular 34. but presumably at much lower population levels U.S. Fish and Wildlife Service, Washington, DC. before the Conservation Reserve Program was American Ornithologists’ Union. 1998. Check-list started (Jensen et al. 2000, Hagen 2003). The of North American birds, 7th edn. American CRP provides a short-term contract for grasslands Ornithologists’ Union, Washington, DC. on a field-by-field basis and does not currently Andrews, R., and R. Righter. 1992. Colorado birds: address long-term landscape-scale certainty for a reference to their distribution and abundance. persistence of the species in this area. Long-term Denver Museum of Natural History, Denver, CO. security is a significant need for the species, espe- Archer, S., and F. E. Smeins. 1991. Ecosystem-level cially considering increasing commodity prices processes. Pp. 109–139 in R. K. Heitschmidt, and and other competing issues for CRP contract J. W. Stuth (editors), Grazing management: an eco- renewal and expansion in the region. The rec- logical perspective, Timbers Press, Portland, OR. ognized distribution of Lesser Prairie-Chickens Bailey, A. M., and R. J. Niedrach. 1965. Birds of has been moving northward in recent years with Colorado. Denver Museum of Natural History, new data from spring lek searches conducted by Denver, CO. biologists since 1999. Much of this portion of Bain, M. R. 2002. Male-male competition and mating the range of Lesser Prairie-Chickens is sympat- success on leks attended by hybrid prairie chickens. ric with Greater Prairie-Chickens (Figure 14.3). M.S. thesis, Fort Hays State University, Hays, KS. Hybridization between Lesser and Greater Bain, M. R., and G. H. Farley. 2002. Display by appar-

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