ATTWATER'S PRAIRIE-CHICKEN (Tympanuchus cupido attwateri ...

ATTWATER’S PRAIRIE-CHICKEN 

(Tympanuchus cupido attwateri) 

DRAFT RECOVERY PLAN 

Second Revision 

Photo by George Levandoski. 

Southwest Region 

U.S. Fish and Wildlife Service 

Albuquerque, New Mexico

DRAFT Attwater’s Prairie-Chicken Recovery Plan – September 2007 

ATTWATER’S PRAIRIE-CHICKEN RECOVERY PLAN 

(Tympanuchus cupido attwateri) 

Second Revision 

Original Approval: December 1983 

First revision approved: February 1993 

Southwest Region 

U. S. Fish and Wildlife Service 

Albuquerque, New Mexico 

Approved: 

Draft 

______________________________________ Date: __________________ 

Regional Director, U.S. Fish and Wildlife Service 

Concurrence: 

______________________________________ Date: __________________ 

Director, Texas Parks and Wildlife Department


DISCLAIMER 

Recovery plans delineate reasonable actions believed necessary to recover and/or protect 

listed species. Plans are published by the U.S. Fish and Wildlife Service, sometimes 

prepared with the assistance of recovery teams, contractors, state agencies, and others. 

Objectives will be attained and any necessary funds made available subject to budgetary 

constraints affecting the parties involved, as well as the need to address other priorities. 

Recovery plans do not necessarily represent the views nor the official position or 

approval of any individuals or agencies involved in the plan formulation, other than the 

U.S. Fish and Wildlife Service. They represent the official position of the agencies 

mentioned only after they have been signed as approved by appropriate personnel and 

posted on the public registry. Approved recovery plans are subject to modification as 

dictated by new findings, changes in species status, and the completion of recovery 

actions. 

ACKNOWLEDGEMENTS 

Principal authors of this document were Dr. Jim Bergan, Dr. Mike Morrow, and Terry 

Rossignol. Dr. Jeff Johnson provided feedback and expanded discussion of genetic 

issues. Dr. Wade Harrell and Tim Anderson provided data for updating maps that were 

developed by Donna Roach and T. J. Shultz. The document was reviewed and edited by 

Attwater’s Prairie Chicken Recovery Team members. Wendy Brown, and Kathy 

Granillo provided initial agency review of the document for the U.S. Fish and Wildlife 

Service. Recovery Team members provided overall recovery objectives and strategies. 

Members included Dr. Jim Bergan (The Nature Conservancy of Texas), Royce Jurries 

(Texas Parks and Wildlife Department), Bobby McCan (McCan Ranch), Dr. Mike 

Morrow (Attwater Prairie Chicken National Wildlife Refuge), Stan Reinke (U.S. Natural 

Resources Conservation Service), Terry Rossignol (Attwater Prairie Chicken National 

Wildlife Refuge), Dr. Steve Sherrod (Sutton Avian Research Center), Dr. Nova Silvy 

(Texas A&M University), Dr. John Toepfer (Society of Tympanuchus Cupido Pinnatus, 

Ltd.), and Bruce Williams (Fossil Rim Wildlife Center). Additional advisors to the 

Recovery Team included Hannah Bailey (Houston Zoo, Inc.), Dr. Joe Flanagan (Houston 

Zoo, Inc.), and Dr. Brent Ortego (Texas Parks and Wildlife Department). 

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EXECUTIVE SUMMARY 

Current Status and Distribution: The Attwater’s prairie-chicken (APC) (Tympanuchus 

cupido attwateri) was listed as endangered with extinction in 1967. This listing was 

“grandfathered” into the Endangered Species Act of 1973. The APC represents the 

southern-most subspecies of T. cupido, and currently occurs in the wild at only two 

locations - the Attwater Prairie Chicken National Wildlife Refuge (Colorado County, 

Texas) and the Texas City Prairie Preserve (Galveston County, Texas). Approximately 

50 birds remained in these two populations as of March 2006. In addition, 160 breeding 

individuals were held in captivity at the Abilene Zoo (Abilene, Texas), Caldwell Zoo 

(Tyler, Texas), Fossil Rim Wildlife Center (Glen Rose, Texas), Houston Zoo (Houston, 

Texas), San Antonio Zoo (San Antonio, Texas), Sea World of Texas (San Antonio, 

Texas), and Texas A&M University (College Station, Texas). 

Habitat Requirements: Lehmann (1941) described APC habitat requirements as follows: 

“Optimum prairie chicken range apparently consists of well-drained 

grassland supporting some weeds or shrubs as well as grasses, the cover 

varying in density from light to heavy; and with supplies of surface water 

available in summer. In short, diversification within the grassland type is 

essential.” 

Lehmann (1939) succinctly summarized habitat needed by APC: 

“It is therefore upon the existence of adequate prairie habitat that the 

welfare of the prairie chicken depends.” 

Reasons for Listing and Limiting Factors: The APC once occupied expansive prairie 

grasslands of coastal Texas and Louisiana. Habitat destruction and degradation, and to a 

lesser extent overharvesting, are the primary factors contributing to historic population 

declines. Current threats include extremely small populations, habitat and population 

fragmentation resulting in genetic isolation, diseases and parasites in both the wild and 

captive setting, inability of captive breeding facilities to produce large numbers of 

captive-reared birds that are capable of survival and reproduction in wild habitats, and 

poor brood survival in wild populations. 

Recovery Goal: The goal of this plan and recovery effort is to protect and ensure the 

survival of the APC and its habitat, allowing the population to reach a measurable level 

of ecological and genetic stability so that it can be reclassified to threatened status 

(downlisted) and ultimately removed from the endangered species list (delisted). 

Recovery Strategy: APC recovery must be focused on 3 primary areas: (1) habitat 

management, (2) captive and wild population management, and (3) public outreach. It is 

imperative that habitat management, enhancement, and restoration be carried out to 

maintain existing grasslands currently suitable as habitat and to restore degraded 

grasslands. These grasslands must be provided at a landscape scale so that multiple areas 

>25,000 acres (ac) (10,120 hectares (ha)) are available to support viable APC populations 

and provide for gene flow between them. Population management consists of actions 

iii


required to manage captive and wild populations. If viable populations are to be 

established in presently unoccupied but suitable habitat, large (>100) numbers of birds at 

multiple release sites will be required. It is clear the captive program must be retooled in 

dramatic fashion to achieve APC recovery. Numerous challenges face the wild APC 

population. Predation (raptors, mesocarnivores, snakes), red imported fire ants 

(Solenopsis wagneri), disease, ectoparasites, accidents (flying into fences, wires), 

flooding, incompatible grazing, altered fire regimes, and countless other factors are 

collectively suppressing optimal recruitment of the two remaining wild populations. 

Additional applied research efforts are essential to identify factors limiting recruitment in 

free-ranging populations, which currently depend heavily on release of captive-reared 

birds. However, conducting meaningful research with broad ranging applicability is very 

challenging given the low population numbers and varied grassland habitats at these two 

sites. An ongoing challenge to recovery has been difficulty in attracting a large 

constituency engaged in APC conservation. A broader support base is critical for timely 

implementation of actions required for APC recovery. 

Recovery Objectives and Criteria: 

1. Downlist to threatened status when the overall population maintains a minimum 

of 3,000 breeding adults annually over a 5-year period. These birds should be 

distributed along a linear distance of no less than 50 miles (80 km) to mitigate for 

environmental stochasticity (e.g., hurricanes) while maintaining gene flow. 

2. Delist when the overall population reaches a minimum of 6,000 breeding adults 

annually over a 10-year period, and occupies habitats along a linear distance of no 

less than 100 miles. 

Specific objectives and criteria for habitat management, captive and wild population 

management, and public outreach necessary to accomplish these recovery goals are: 

Objective 1: Maintain and improve 300,000 ac (121,457 ha) of coastal prairie habitat for 

APC throughout the bird’s historical range on both private and public lands. APC 

recovery will require a network of large, high quality coastal prairie habitats containing 

multiple core areas distributed along at least 100 linear miles (160 km). A core area is 

defined as an area of habitat capable of supporting a population of 500 (250 displaying 

males), or approximately 25,000 ac (10,121 ha) (assuming a carrying capacity of 1 

bird/50 ac (20 ha) (Lehmann1941). 

Objective 2: Enhance propagation and release efforts to boost wild populations to viable 

levels and re-establish physically and behaviorally healthy birds to their former range, as 

measured by the following criteria: 

(a) Maintain 90% of original gene diversity for 20 years with a minimum of 200 

birds in the captive flock. 

(b) Produce enough chicks annually to release at multiple sites (approximately 

100 birds per release site). 

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• Increase capacity of breeding pairs to a minimum of 100 pairs within 

two years. 

• By 2008, increase survival in the captive environment so that 50% of 

eggs produced survive to 8 weeks of age. 

(c) When number of young available for release exceeds 100, pilot releases of no 

fewer than 30 should be considered on private lands. 

Objective 3: Establish populations of at least 500 birds in multiple core areas, providing 

for gene flow between populations (see Objective 1). 

Objective 4: Broaden public support and partner in efforts to conserve the APC and its 

coastal prairie ecosystem. 

Estimated Date and Cost of Recovery: Because the APC is an r-selected species, it is 

capable of explosive population growth. Assuming exponential population growth, and a 

maximum growth rate of r = 1.1 (the maximum metapopulation growth rate observed 

during 1971−1996 for any one year; M. Morrow, APCNWR, unpublished data), the 

threshold population size of 6,000 required for delisting could be achieved within 5 years. 

However, a more realistic sustained r of 0.1 (1971−1975 average for statewide population 

during last recorded 5-year interval of population growth – see Appendix 1) would 

require 48 years to achieve the population threshold for delisting. An average r of 0.08 

was observed from 1972-1987 on the Attwater Prairie Chicken National Wildlife Refuge 

(APCNWR) during a period of general population increase (M. Morrow, APCNWR, 

unpublished data). Estimated costs for implementation of tasks described in the 

implementation schedule (Section III) over a 50-year recovery period are provided in 

Table 1. 

Table 1. Cost estimates ($1,000) for implementation of Attwater’s prairie-chicken 

recovery actions over a projected 50-year recovery period. 

Year 

Habitat 

(Action # 1) 

Captive 

Population 

Management 

(Action # 2) 

Wild 

Population 

Management 

(Action #3) 

Public 

Outreach 

(Action # 4) 

Total 

1 3,862 1,715 400 225 6,202 

2 3,687 1,357 325 50 5,419 

3 3,612 957 348 50 4,967 

4 3,612 797 350 50 4,809 

5 3,612 797 350 50 4,809 

50 102,600 13,363 7,698 425 124,086 

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TABLE OF CONTENTS 

DISCLAIMER................................................................................................................... ii 

ACKNOWLEDGEMENTS ............................................................................................. ii 

EXECUTIVE SUMMARY .............................................................................................iii 

LIST OF TABLES AND FIGURES.............................................................................. vii 

BACKGROUND INFORMATION ................................................................................ 1 

BRIEF OVERVIEW..................................................................................................... 1 

TAXONOMY AND DESCRIPTION.......................................................................... 3 

DISTRIBUTION AND ABUNDANCE....................................................................... 5 

HABITAT/ECOSYSTEM............................................................................................ 9 

LIFE HISTORY/ECOLOGY .................................................................................... 11 

Reproduction ............................................................................................................ 11 

Booming Behavior ............................................................................................... 11 

Nesting.................................................................................................................. 13 

Brood rearing ....................................................................................................... 15 

Habitats Used Outside the Breeding Season........................................................... 18 

Food Habits.............................................................................................................. 18 

Survival and Mortality Factors................................................................................ 19 

Home Range and Movements.................................................................................. 21 

Habitat Management ............................................................................................... 23 

CRITICAL HABITAT ............................................................................................... 25 

ON-GOING CONSERVATION EFFORTS ............................................................ 25 

Research ................................................................................................................... 25 

Habitat Management ............................................................................................... 25 

Captive Breeding...................................................................................................... 29 

Population Supplementation ................................................................................... 34 

REASONS FOR LISTING/CURRENT THREATS ............................................... 39 

RECOVERY.................................................................................................................... 44 

RECOVERY STRATEGY......................................................................................... 44 

GOALS, OBJECTIVES, AND CRITERIA.............................................................. 46 

NARRATIVE OUTLINE OF RECOVERY ACTIONS ......................................... 49 

REDUCTION OR ALLEVIATION OF THREATS............................................... 53 

IMPLEMENTATION SCHEDULE ............................................................................. 55 

LITERATURE CITED .................................................................................................. 68 

APPENDIX 1. Attwater’s prairie-chicken 1937−2006 population estimates by Texas 

county............................................................................................................................... 81 

APPENDIX 2. LIST OF ABBREVIATIONS AND ACRONYMS ........................... 84 

APPENDIX 3. GLOSSARY OF TERMS .................................................................... 86 

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LIST OF TABLES AND FIGURES 

Table 1. Cost estimates ($1,000) for implementation of Attwater’s prairie-chicken 

recovery actions over a projected 50-year recovery period................................................ v 

Figure 1. Approximate distribution of Attwater’s prairie-chicken in southeast Texas 

historically, 1937, 1963, and 2007 (from Morrow et. al 2004 with modification)............. 2 

Figure 2. Attwater’s prairie-chicken population trends in southeast Texas, 1937–2006... 6 

Figure 3. Land use (1984) and its relationship to Attwater’s prairie-chicken priority 

management zones.............................................................................................................. 7 

Figure 4. Land use within the Austin-Colorado County, Texas priority management 

zone................................................................................................................................... 27 

Figure 5. Land use within the Refugio-Goliad County, Texas priority management zone. 

........................................................................................................................................... 28 

Figure 6. Projected Spring 2007 distribution of captive Attwater’s prairie-chickens by 

location (n = 164).............................................................................................................. 31 

Figure 7. Comparison of Attwater’s and greater prairie-chicken mass propagation efforts. 

........................................................................................................................................... 32 

Figure 8. Comparison of Attwater’s and greater prairie-chicken captive egg production. 

........................................................................................................................................... 33 

Figure 9. Captive Attwater’s prairie-chicken mortality during the first month post-hatch. 

........................................................................................................................................... 35 

Figure 10. Captive Attwater’s prairie-chickens (n = 1,005) released at the Attwater 

Prairie Chicken National Wildlife Refuge (APCWR), Colorado County, Texas and the 

Texas City Prairie Preserve (TCPP), Galveston County, Texas from 1995−2006........... 36 

vii

DRAFT Attwater’s Prairie-Chicken Recovery Plan –September 2007 

I. BACKGROUND INFORMATION 

A. BRIEF OVERVIEW 

The Attwater’s prairie-chicken (Tympanuchus cupido attwateri) (APC) is a 

subspecies of prairie-chicken endemic to prairies along the Gulf of Mexico (Bendire 

1894). Historically, APC populations approached 1 million individuals on an estimated 6 

million acres (2.4 million ha) of prairie habitat (Lehmann 1968). By 1937, populations 

had declined to an estimated 8,700 individuals and have continued to decline. As of 

spring 2006, approximately 50 remained in two free-ranging populations (Figure 1). The 

APC was listed as endangered in March 1967 under the Endangered Species Preservation 

Act (ESA) of 1966 (32 FR 4001). It is currently listed as endangered by the U.S. Fish 

and Wildlife Service (USFWS) under provisions of the ESA of 1973 (50 CFR 17.11) and 

by Texas Parks and Wildlife Department (TPWD) (31 TAC 65.181-184). The APC has a 

recovery priority ranking of 3 on a scale from 1 (high priority) –18. This ranking reflects 

a high degree of threat, high potential for recovery, and the APC’s taxonomic status as a 

subspecies. Priority 3 is the highest ranking assigned to subspecies (48 FR 43104). 

Loss and fragmentation of its coastal prairie ecosystem and associated isolation of 

sub-populations brought about by agricultural conversion, urban and industrial 

expansion, overgrazing, and invasion of prairies by woody species have been the ultimate 

factors responsible for the APC’s decline (Lehmann 1941, Jurries 1979, Lawrence and 

Silvy 1980, McKinney 1996, Morrow et al. 1996). Probable proximate contributors to 

range-wide population declines in recent history include stochastic weather events 

(Morrow et al. 1996), reduced genetic variability (Osterndorff 1995), parasites (Peterson 

1994, Purvis 1995), disease (Peterson et al. 1998), and red imported fire ants (Mueller et 

al. 1999). These and possibly other factors have contributed to reduced survival and 

reproductive output (Peterson 1994, Peterson and Silvy 1994). 

A captive breeding program was initiated for the APC in 1992. This program had 

two primary goals: (1) preserve as much genetic variability as possible, and (2) provide 

birds for supplementation of remaining populations and the re-establishment of extirpated 

populations. From 1995−2006, a total of 1,005 captive-reared birds has been released in 

an effort to buoy failing populations at the Attwater Prairie Chicken National Wildlife 

Refuge (APCNWR) and at The Nature Conservancy’s (TNC) Texas City Prairie Preserve 

(TCPP) (APCNWR, unpublished data). As of October 2006, 166 APC were held in 

breeding facilities at the Abilene, Caldwell, Houston, and San Antonio Zoos, and at 

Fossil Rim Wildlife Center, Sea World of Texas, and Texas A&M University (TAMU) 

(H. Bailey, APC Species Survival Plan (SSP) Coordinator, Houston Zoo, Inc.). 

1


Figure 1. Approximate distribution of Attwater’s prairie-chicken in southeast Texas historically, 1937, 1963, and 2007 (from Morrow et. al 

2004 with modification). 

2


B. TAXONOMY AND DESCRIPTION 

Tympanuchus cupido attwateri is a member of the class Aves, family Phasianidae, 

and subfamily Tetraoninae (American Ornithologists’ Union 1998). It shares its 

subspecies status with the greater prairie-chicken (GPC) (T. c. pinnatus), which occupies 

grasslands of the North American great plains, and the extinct heath hen (T. c. cupido), 

which once occupied grasslands of the northeastern United States (Aldrich 1963, Silvy 

and Hagen 2004). The APC was described by Bendire (1894): 

“Smaller than T. americanus [greater prairie chicken], darker in color, more 

tawny above, usually with more pronounced chestnut on the neck; smaller 

and more tawny light colored spots on the wing coverts, and much more 

scantily feathered tarsus, the latter never feathered down to base of toes, 

even in front; a broad posterior strip of bare skin being always exposed, 

even in winter, while in summer much of the greater part of the tarsus is 

naked.” 

Physical differences between the APC and the GPC are minor. Smaller 

measurements of wing, tail, bill, and total length and differences in general ruddiness and 

buffiness of the underparts are characteristic and can be used to separate the APC as a 

subspecies (Lehmann 1941). Oberholser (1974) described the APC subspecies as similar 

to the GPC, 

“…but smaller; feathering of tarsus somewhat shorter and sometimes 

leaving lower half of leg bare; coloration somewhat more rufescent and 

buffy, particularly on flanks and other lower parts; dark bars on lower 

surface usually narrower.” 

Lack of feathering extending onto the feet of the APC, less feathering on the tarsus, and 

lack of well-developed pectinae on the toes of the APC during winter are probably the 

most concrete phenotypic differences between the APC and GPC subspecies. Other 

differences, such as plumage coloration and size, are more subtle. 

Svedarsky (1979) reported average breeding season weights for Minnesota GPC 

females (hens) of 919 grams (2.0 lbs) (n = 16). Breeding season weights for combined 

GPC age classes from Wisconsin and Minnesota, weighted by sample size from Toepfer 

(1988:68), were 1,033 grams (2.3 lbs) for males (cocks) (n = 89) and 929 grams (2.0 lbs) 

for females (n = 55). APC weights reported by Lehmann (1941) averaged 1,112 grams 

(2.5 lbs) (n = 5) for males and 737 grams (1.6 lbs) for females. Morrow and Silvy 

(unpublished data) observed average breeding season weights of 986 grams (2.2 lbs) for 

male (n = 26) and 874 grams (1.9 lbs) for female (n = 28) APC captured on APCNWR 

booming grounds from 1983–1985. Toepfer (1988) observed that maximum weights of 

wild GPC cocks from Wisconsin and Minnesota occurred approximately 3–4 weeks 

before the breeding peak, while female weights peaked just before egg laying. Toepfer 

(1988) observed that minimum weights for GPCs occurred during August in Wisconsin, 

with both sexes losing an average of 14% from their peak April weights. Similarly, 

Svedarsky (1979), also working with the GPC subspecies, observed a 14.6% weight loss 

during summer for Minnesota hens. This summer weight loss was attributed primarily to 

3


high energy demands of the annual molt (Toepfer 1988). Immature GPCs do not reach 

full size until late fall or early winter of their second year, with males gaining an average 

9% and females 6% compared to their hatch-year weights (Toepfer 1988). 

From a genetics perspective, the APC does not represent a phylogenetically 

distinguishable group based on criteria of monophyly when compared to all recognized 

species within the genus Tympanuchus, based on analysis of mitochondrial DNA 

(mtDNA) control region sequence data (Palkovacs et al. 2004, Johnson and Dunn 2006. 

Johnson et al. 2007). In fact, with the exception of the heath hen (Palkovacs et al. 2004, 

Johnson and Dunn 2006), studies utilizing a number of different molecular markers 

including allozymes, mtDNA, and nuclear intron sequence data (Ellsworth et al. 1994, 

Dimcheff et al. 2002, Drovetski 2002, Palkovacs et al. 2004, Johnson and Dunn 2006, 

Spaulding et al. 2006) with traditional gene-tree approximations found no clear 

phylogenetic differentiation among any species of this genus, which also includes sharptailed 

grouse (T. phasianellus) and lesser prairie-chicken (T. pallidicinctus). However, 

recent analyses by J. Johnson (University of Michigan, unpublished data) using a 

coalescent approach to investigate the demographic history associated with each taxon 

based on mtDNA sequence data indicate that despite morphological and behavioral 

similarities between APC and GPC, these two taxa are as genetically divergent from each 

other as either is from morphologically distinct lesser prairie-chickens and sharp-tailed 

grouse. In a recent unpublished study using nuclear microsatellite DNA allele frequency 

data, J. Johnson (Univ. of Michigan) was also able to identify significant population 

genetic differentiation between all Tympanuchus taxa, including the APC population, 

suggesting that no contemporary gene flow exists between sampled populations. 

The apparent lack of reciprocal monophyly among these taxa based on traditional 

phylogenetic methods is due to incomplete lineage sorting rather than contemporary gene 

flow (Johnson et al. 2007). Ellsworth et al. (1994), Drovetski (2002), and Spaulding 

(2007) suggested that morphological and behavioral differentiation among species in this 

genus, particularly for those with overlapping geographic distributions, is largely driven 

by sexual selection and has progressed more rapidly than mtDNA or allozyme 

differentiation. This suggestion is consistent with the significant amount of time that 

would be required for attainment of reciprocal monophyly due to the recent 

diversification within this genus and its large ancestral effective population size 

associated with this genus and its recent ancestry (e.g., Hudson and Coyne 2002, Johnson 

et al. 2007). 

In summary, most molecular approaches to date have not been able to identify distinct 

groups associated with commonly accepted species taxonomy within Tympanuchus (i.e., 

lesser prairie-chicken, GPC, and sharp-tailed grouse) suggesting that this genus 

experienced a rapid diversification within the past 10,000 years. However, more recent 

genetic analyses have suggested that APC and GPC are as differentiated from each other 

as either is from other recognized species within the genus (J. Johnson, University of 

Michigan, unpublished data). Therefore, APC and GPC may warrant separate species 

status despite any observable behavioral or morphological differences. 

4


C. DISTRIBUTION AND ABUNDANCE 

The APC represents the southernmost extension of the genus Tympanuchus. 

Historically, APCs ranged from southwest Louisiana to possibly near Brownsville, Texas 

(Lehmann 1941, Oberholser 1974, Peterson 1994, Silvy et al. 2004). However, in 

reviewing historical accounts Lehmann and Mauermann (1963) concluded: 

“…Attwater’s prairie chickens have almost certainly never been abundant 

in any part of the southern coastal prairie south of the Nueces River from 

the mid-1800’s to the present.” 

These authors suggested the propensity for severe droughts along the lower Texas coast 

and the Rio Grande River plain limited establishment of long-term populations in those 

areas. Lehmann (1941) reported the northern distribution of the APC was limited by the 

northern edge of the coastal prairie. Oberholser (1974) reported data that suggest APCs 

may have ranged as far north as Bastrop and Travis counties in Texas, but Lehmann 

(1941) considered records from these two counties as being questionable. Silvy et al. 

(2004), citing data from Oberholser (1974), reported a 2-county overlap in the historic 

distribution of APC and GPCs in Texas. GPCs were extirpated from Texas by 1920, with 

the last records occurring in northeast Texas near Marshall (Oberholser 1974). 

APCs were extirpated from Louisiana by 1919 (St. Amant 1959, Oberholser 1974), 

although St. Amant (1959) reported huntable populations existed in 12 parishes by as late 

as 1890. Only an estimated 8,700 individuals remained in 19 Texas counties by 1937 

(Lehmann 1941), down from a historic distribution of up to 48 Texas counties (Silvy et 

al. 2004) which may have supported numbers approaching 1 million in peak years 

(Lehmann 1941). Population declines continued and by 1999, APC remained in only two 

counties (Morrow et al. 2004, Silvy et al. 2004) (Figure 1, Appendix 1). Populations in 

these two counties have been buoyed since 1996 by supplementation with birds reared in 

captivity (Morrow et al. 2004, Silvy et al. 2004). In spring 2006, approximately 50 APC 

remained in free-ranging populations at the APCNWR (Colorado County, Texas) and 

TCPP (Galveston County, Texas) (APCNWR, unpublished data) (Figure 2, Appendix 1) 

Loss of its prairie grassland habitat was the primary cause for the APC decline 

(Figure 3). Lehmann (1941) indicated that 93% of the 6 million acres (2.4 million ha) of 

coastal prairie that once supported APC had been lost by 1937. Coastal prairie loss 

continued through the remainder of the 20 th century. From 1952–1990, grassland acreage 

containing the two largest remaining APC populations declined by 67% in Austin and 

Colorado counties and by 46% in Aransas, Goliad, and Refugio counties (McKinney 

1996). Smeins et al. (1991) estimated that


Figure 2. Attwater’s prairie-chicken population trends in southeast Texas, 1937–2006. 

10000 

Population Estimate 

8000 

6000 

4000 

2000 

0 

1937 

1942 

1947 

1952 

1957 

1962 

1967 

1972 

Year 

1977 

1982 

1987 

1992 

1997 

2002 

6


Figure 3. Land use (1984) and its relationship to Attwater’s prairie-chicken priority 

management zones. 

7


supported maximum densities of 1 bird/10 acres (4 ha) and poorly drained areas (30% of 

former range) supported not more than 1 bird/50 acres (20 ha). However, densities 

suggested by Lehmann (1941) are high, at least for the best range, compared to those 

reported for the GPC subspecies. Hamerstrom et al. (1957) reported that observed 

densities for the best habitat within each state or Canadian province ranged from less than 

1 cock/1,000 acres (405 ha) in Ontario to 1 cock/16 acres (6 ha) in Kansas. Maximum 

population densities reported for GPCs on areas of managed habitat (i.e., considering 

only the area under management in density estimates) ranged from 1 cock/6−8 acres (2−3 

ha) (1 bird/3−4 acres, assuming 1:1 sex ratio) of ecologically-patterned habitat (Missouri 

Department of Conservation 1984). Arthaud (1970) observed a density of 1 males/3.5 

acres (1.4 ha) on a 1,680-acre (680 ha) management area in Missouri. 

However, Toepfer (2003) cautioned that one must evaluate a population’s health 

based on range-wide density estimates, and suggested that for prairie-chickens, a 

population’s range be determined by a minimum convex polygon which connects the 

outer-most booming ground locations. Taking this approach, Toepfer (2003) reported a 

range-wide density of approximately 1 bird/section for Wisconsin and 1.5 birds/section 

for Minnesota. Using booming ground location data described by McKinney (1996) and 

census data compiled by APCNWR, 1979 density estimates calculated with this approach 

for the two geographically separated APC populations described by McKinney (1996) 

were 3.3 and 4.5 birds/section for the Aransas-Goliad-Refugio (254 mi 2 , 659 km 2 ) and 

the Austin-Colorado (113 mi 2 , 292 km 2 ) County populations, respectively (Figures 4, 5). 

The Austin-Colorado County population contains the APCNWR. However, because 

APC were traditionally surveyed on fixed routes to monitor population trends, these 

estimates are likely very conservative. 

Toepfer (2003) suggested that high densities of wildlife are not necessarily 

reflective of healthy populations, especially from a genetic perspective. He suggested 

that for prairie-chickens, a key objective must be to maintain gene flow among 

populations within relatively large areas (see Johnson et al. 2004). Toepfer (2003) holds 

the Minnesota population, with an average density of roughly 1.5 birds/section, up as “the 

real prairie chicken success story in the U.S.” Despite going through several population 

bottlenecks within its roughly 3,000 mi 2 (7,770 km 2 ) range, by 2003 the Minnesota 

population had recovered to the point that it was able to sustain its first hunting season 

since 1942 (Toepfer 2003). 

However one reports population-level data, it is clear from a review of the literature 

that as an r-selected species, prairie-chicken populations of both subspecies are subject to 

sudden, catastrophic population declines. For example, Oberholser (1974) suggested that 

GPC populations in north-central Texas may have numbered 500,000 circa 1850. 

Oberholser (1974) continued: 

“Between 1870 and 1890, these birds were shot by the wagon load for 

meat and blood sport; even worse was the plowing up or overgrazing of 

their grassland habitat…. the last small flock disappeared after June 

1920… Along the Texas coast, the Attwater’s Greater Prairie Chicken, T. 

c. attwateri, was slaughtered with customary frontier abandon between 

1840 and 1900…. For attwateri, as for other races, plowing under of the 

8


native sod was even more damaging than gunshots. Where overgrazing by 

livestock weakened the turf, huisache (Acacia farnesiana), mesquite 

(Prospis), retama (Parkinsonia aculeata), and other brush species 

encroached, reducing suitable acreage for this prairie chicken. As of about 

1880, the Attwater’s was still generally distributed over the Texas coastal 

prairie… some 810,000 birds… By 1937, the occupied area had shrunk to 

disjunct colonies…at this time Lehmann determined there were only about 

8,700 attwateri in Texas – and the world….” 

Smaller, isolated populations can disappear with even greater rapidity. For 

example, Walk (2004) reported that in 1962 approximately 2,000 GPCs remained in 

Illinois. By 1966, fewer than 400 remained. Since 1987, range-wide APC populations 

declined from an estimated 1,108 individuals to 42 in 1996 (Figure 2). Remaining APC 

populations have been supplemented with captive-reared birds since 1996. In the 

absence of this supplementation, APCs populations would have undoubtedly become 

extinct in the wild (Morrow et al. 2004). Toepfer (2003) stated that historical GPC data 

suggests isolated populations generally disappear once they fall below 100 cocks unless 

intensive management is implemented. APC population trends support that hypothesis, 

and suggest that populations dropping below 250 cocks for more than 3 years in 

succession have a high probability of ultimately going extinct (Appendix 1). 

D. HABITAT/ECOSYSTEM 

Description of habitat required by the T. cupido species in general, and the APC 

subspecies in particular, is relatively simple: they require lots of grass and open space 

(e.g., Lehmann 1939, 1941; Schwartz 1945; Baker 1953; Hamerstrom et al. 1957; Cogar 

et al. 1977; Toepfer 1988, 2003; Johnson et al. 2003, 2004; Silvy et al. 2004; Niemuth 

2005). Lehmann (1939) summarizes APC habitat requirements: 

“True to its name, the Attwater [sic] prairie chicken is a bird of the prairie. 

Woodland, brushland, fallow land, and cultivated land furnish some food 

and cover at certain times and under certain conditions, but use of these 

types by prairie chickens is optional, not vital. Individually or in 

combination, these types of land furnish little or nothing in the way of 

critically necessary courtship grounds and nesting cover. Moderately 

grazed and moderately burned grassland, on the other hand, provides 

prairie chickens with everything they need at all seasons. It is therefore 

upon the existence of adequate prairie habitat that the welfare of the 

prairie chicken depends.” 

While native prairie is most often identified as a habitat requirement for the APC 

subspecies (Lehmann 1939, 1941; Cogar et al. 1977, Horkel 1979), Toepfer (2003) stated 

there is no evidence GPCs prefer or require native grasses. However, both Hamerstrom 

et al. (1957) and Toepfer (2003) stressed the importance of permanent grassland as GPC 

habitat, especially for nesting, brood rearing, and year-round night roosting. Hamerstrom 

et al. (1957) indicated that total grassland appeared to be a rough index of GPC habitat 

quality. 

9


Although there is general agreement that quantity of grassland is directly related 

to prairie-chicken population levels (Hamerstrom et al. 1957, Newell et al. 1987, Toepfer 

2003), there is no consensus on the size and composition of management areas required. 

Hamerstrom et al. (1957) found minimum populations of GPCs in 10-40% permanent 

grassland, while areas with more than 40% permanent grassland supported 

proportionately larger populations. These authors observed the densest population on 

record at that time (38.8 cocks/640 ac) in an area of 77% permanent native prairie, and 

“low lingering” populations in 10-15% relatively undisturbed grassland. Based on these 

observations, Hamerstrom et al. (1957) stated that as a rule of thumb, GPCs occurred on a 

sustainable basis in areas which were at least 33% grassland, but were abundant only 

where grass comprised 50-75% of the area. 

Minimum areas required to support a viable population of T. cupido range from 

several hundred to several thousand acres (Niemuth 2000, Toepfer 2003). However, 

Toepfer (2003) stated that previous estimates of minimum management area size for 

prairie-chickens “…all are much too small.” Toepfer (2003) observed the approximately 

3,000 mi 2 (7,770-km 2 ) Minnesota GPC range has undergone several population 

bottlenecks and still maintained its genetic viability, whereas prairie-chickens occupying 

smaller ranges have declined, become extirpated, or undergone substantial declines in 

genetic diversity (see Johnson et al 2003, 2004). Toepfer (2003:55) concludes: 

“At this point in time we still do not know the minimum size necessary to 

sustain a viable greater prairie chicken population….We should all now 

understand that to be successful, management would have to spread 

thousands of acres of grassland habitat over a landscape of several 

thousand square miles and maintain connectivity.” 

While grass has long been recognized as an important component of prairiechicken 

habitat, open space has been given less detailed attention (Toepfer 1988). 

Prairie-chickens occasionally use trees for food, roosting, or loafing (Lehmann 1941, 

Hamerstrom et al. 1957, Toepfer 1988), but in general have an aversion to being closed in 

by woodland or overhanging cover (Hamerstrom et al. 1957, Toepfer 2003). 

Hamerstrom et al. (1957) indicated good prairie-chicken cover should contain less than 

20-25% woodland cover where woody cover is distributed in scattered blocks, whereas 

Ammann (1957) observed GPCs in Michigan survived best with 10-25% woody cover. 

Toepfer (1988) reported the mean size of treeless areas and open space was positively 

associated with the number of cocks attending booming grounds in Wisconsin. Toepfer 

(1988) and Niemuth (2003, 2005) found landscapes surrounding GPC booming grounds 

in Wisconsin contained more grass and less forest than unused random points. Similarly, 

Merrill et al. (1999) found Minnesota booming grounds occurred in landscapes 

containing less residential farmstead, smaller amounts and smaller patches of forest, and 

greater amounts of Conservation Reserve Program (CRP) lands, which provide suitable 

grasslands for prairie-chickens (Toepfer 2003). Hamerstrom et al. (1957), Anderson 

(1969), and Toepfer (2003) have reported movement or abandonment of booming 

grounds in response to natural or artificial structures near booming grounds. Toepfer 

(2003) reported that increasing the treeless area from 140−540 acres (57–219 ha) around 

a Wisconsin booming ground by removal of scattered trees increased annual survival of 

10


cocks from this booming ground by at least 20% compared to males from control 

booming grounds. 

Hamerstrom et al. (1957) indicated the distribution of woods and openings is 

probably more important than total acreage. Specifically, Hamerstrom et al. (1957) 

stated nesting habitat should have a tree-free area of at least 0.5 mi (0.8 km) in one, and 

preferably two dimensions (i.e., length and width), and Keenlance (1998) found distance 

to nearest woodline was greater at nests compared to random points. McKee et al. (1998) 

found nest success decreased substantially when more than 5% woody cover was present 

at nest sites. Merrill et al. (1999) found no traditional booming grounds within 1 mi (1.6 

km) of any town or forest patch greater than 75 ac (30 ha), although Toepfer (STCP, 

personal communication) has since observed several booming grounds within 1 mi (1.6 

km) of towns in the Merrill et al. (1999) study area following a substantial GPC 

population increase. 

Hamerstrom et al. (1957) succinctly summarized the foregoing discussion on T. 

cupido habitat requirements: 

“Grassland is of vital importance to prairie chickens, the keystone in 

prairie chicken ecology….Wherever one looks, the answer is the same: to 

save the prairie chicken, grasslands must be preserved and managed for 

them. There are no substitutes.” 

E. LIFE HISTORY/ECOLOGY 

Reproduction 

Booming Behavior. The most conspicuous phase of the APC life cycle occurs on 

communal display areas known as booming grounds, named after the resonant 

vocalizations made by displaying males. Courtship behavior of the APC and GPC 

subspecies is similar (Bent 1963). Several studies have pointed to the importance of 

booming grounds as focal points for prairie-chicken ecology (Schwartz 1945; 

Hamerstrom et al. 1957; Toepfer 1988, 2003). Toepfer (2003) states: 

“The booming ground is the social center of prairie chicken ecology. 

Movements are best characterized as being associated with the habitat 

within and surrounding a complex of booming grounds…The role of the 

booming ground in prairie chicken ecology cannot be overstated as most, 

if not all, of the life history of individual birds occurs within a mile of a 

booming ground. This concept goes back to Schwartz (1945) who 

believed that each booming ground had its own ‘sphere of influence’ with 

its own group of cocks and hens. This idea is supported by years of radio 

tracking that indicate the majority of radio-tagged regular adult cocks of 

adjacent booming grounds rarely come together, and that areas used by 

these adult cocks show little overlap unless an individual shifts booming 

grounds.” 

11


Booming grounds vary in size from about one-eighth to several acres (Jurries 

1979). They may be naturally occurring short grass flats or artificially maintained areas 

such as roads, airport runways, oil well pads, plowed fields and drainage ditches (Jurries 

1979, Horkel 1979). Numerous studies have observed that active booming grounds are 

usually in close proximity to grass suitable for nesting and night roosting (e.g., Lehmann 

1941; Horkel 1979; Niemuth 2000, 2003; Toepfer 1988, 2003). Due to the large number 

of artificially maintained areas currently available within the APC range, sufficient 

booming areas are generally available to all males (Horkel 1979). However, booming 

grounds found on artificial areas are sometimes less stable than ancestral booming 

grounds. For example, Lehmann (1941), Kessler (1978) and Jurries (1979) observed 

recently established booming grounds on fallow rice fields had poor territorial hierarchy 

when compared to ancestral grounds. Similarly, Horkel and Silvy (1980) found booming 

grounds on narrow, linear areas such as roads and pipeline rights-of-way were less stable 

than more typical circular-shaped leks. Stable GPC booming grounds appear to have 

greater male visitation on average than unstable booming grounds (Hamerstrom and 

Hamerstrom 1973, Schroeder and Braun 1992, Merrill et al. 1999). Merrill et al. (1999) 

found traditional GPC booming grounds were surrounded by proportionately less forest 

and cropland (i.e., more grass) than were temporary booming grounds. Schroeder and 

Braun (1992) noted, that on average, 22.9% of GPC booming grounds in their Colorado 

study disappeared each year. Jurries (1979) observed temporary booming grounds 

usually resulted from increased populations, which were often abandoned when 

populations decline. Cover changes also may influence location and attendance of 

booming grounds (Lehmann 1941, Hamerstrom et al. 1957, Anderson 1969, Toepfer 

2003). 

Males gather on booming grounds in early morning and late evening to establish 

individual territories and to attract females, although attendance in the morning is more 

regular (Schwartz 1945). Jurries (1979) reported the number of cocks on an APC 

booming ground ranged from 3–40, but averaged 6–15. Hamerstrom and Hamerstrom 

(1973) observed annual average numbers of GPC cocks/booming ground during their 22- 

year study from 6.4–13.5 (range 1–45, n = 529) for stable booming grounds, and 1–4.5 

for booming grounds of uncertain status (range 1–7, n = 82). Attendance by APC males 

is sporadic in fall (October–December), but both attendance and intensity of territorial 

defense increases by late January to early February (Lehmann 1941, Jurries 1979). 

Lehmann (1941) stated that courtship activity was at its peak in March, while Horkel 

(1979) indicated the “height of the booming season” occurred in late February to early 

March. Counts of cocks (minimum of 3 recommended), taken 45 minutes before sunrise 

to one hour after sunrise during the 2–3-week period of peak display, are recommended 

for use as population indices (Hamerstrom and Hamerstrom 1973, Svedarsky 1983). 

APC booming activity typically ends by the third week in May (Lehmann 1941). 

Largest groups of females are generally observed on booming grounds a few days 

prior to the peak in breeding (Hamerstrom and Hamerstrom 1973, Robel and Ballard 

1974). Booming intensity increases when hens are present. Several males may follow 

the hen(s) as they walk across the booming ground, resulting in a temporary break down 

of cock territorial boundaries (Jurries 1979). Copulations begin to occur in late February, 

peak in early March, and gradually decrease through April and early May (Jurries 1979, 

Lutz 1979). Secondary peaks in breeding occur in April resulting from hens attempting 

12


to re-nest after initial attempts fail (Jurries 1979). Hens may copulate with more than one 

male (Lehmann 1941). 

GPC studies have shown males occupying territories near the center of booming 

grounds are generally the most dominant, and usually perform the majority of copulations 

(Robel 1970). Robel (1970) reported that only approximately 10% of the male GPC 

population was directly involved with breeding. However, Hamerstrom and Hamerstrom 

(1973) observed 18% of copulations by known-age GPC cocks (n = 506) were by 

juveniles, and 31% (n = 555) were by cocks with exterior territories. Rates of booming 

ground visitation appear to be similar for adult and juvenile cocks, although juveniles 

visit more booming grounds during the breeding season than adults (Schroeder and Braun 

1992). However once established, males maintain strong fidelity to booming grounds. 

Toepfer (1988) observed 84.6% of surviving GPC cocks returned to the booming ground 

on which they displayed the previous year (n = 66), and 80.0% of those that shifted to a 

new ground were between their first and second booming season. All juveniles (both 

sexes) attempt to breed during their first booming season (Schroeder and Robb 1993). A 

detailed description of behaviors and vocalizations associated with APC booming 

grounds is provided by Lehmann (1941). 

Nesting. After the female has mated, she leaves the booming ground to initiate egg 

laying within approximately four days (Svedarsky 1983). The earliest date observed for 

initiation of incubation for APC was before March 21, based on observations of a hen 

with young chicks at the TCPP on 16 April 1999 (M. Morrow, APCNWR, personal 

observation). The latest initiation of incubation recorded was May 29 (Lehmann 1941). 

Eggs pip approximately 23-24 days after the onset of incubation, and hatch 

approximately 48 hours later (Lehmann 1941). Hatching dates ranged from April 16 (M. 

Morrow, APCNWR, personal observation) to the third week in June (Morrow 1986). 

Hens lose approximately 15–20% of their body mass during incubation (Rumble et al. 

1987, Toepfer 1988). 

A summary by Peterson and Silvy (1996) of several APC nesting studies indicated 

clutch size ranged from 7–16 eggs, averaging 12.1 for initial attempts (n = 106), and 9.5 

for renesting attempts (n = 25). The 11.6-egg average for all attempts was not 

statistically different from clutch sizes reported for GPCs (Peterson and Silvy 1996). 

Peterson and Silvy (1996) observed an average APC egg hatchability of 87.3% (n = 648), 

which was not statistically different than the 88.7% reported for GPCs. Peterson and 

Silvy (1996) found that APC nest success averaged 32.2% for 143 nests observed during 

studies conducted 1937–1985, significantly lower (P = 0.0087) than the average 49.5% 

reported for 480 GPC nests. Lehmann (1941) observed an average nest success of 

31.5%, so the observed difference between APC and GPC nesting success is not a 

phenomenon that has developed in recent history. If a female’s first nest is destroyed, 

she may re-nest (Lehmann 1941, Jurries 1979), with egg-laying in the second clutch 

beginning as soon as 8–9 days later (Schroeder and Robb 1993). McKee et al. (1998) 

reported when all nesting attempts were considered, 56% of Missouri GPC hens hatched 

chicks even though average nest success was only 35%. Newell (1987) observed 57.9% 

of radio-tagged GPC hens in North Dakota successfully hatched chicks. Toepfer (1988) 

found re-nesting by Wisconsin GPCs provided 38% of the production for a year. 

Similarly, Newell et al. (1987) observed 36% of North Dakota GPC chicks came from re- 

13


nesting attempts. Newell (1987) found 28% of subadult and 88% of adult hens re-nested, 

respectively. Fields et al. (2006) observed the survival of Kansas prairie-chicken nests 

declined as the nest aged and as the nesting season progressed. They found survival 

probability of early-, mid-, and late-season nests was 0.77, 0.61, and 0.19, respectively. 

Later ring-necked pheasant (Phasianus colchicus) hatches resulted in smaller clutch 

sizes, lower chick weights, and reduced chick survival (Riley et al. 1998) 

APC nest predators include skunks (Mephitis mephitis, Spilogale putorius), 

opossum (Didelphis virginianus), raccoon (Procyon lotor), coyote (Canis latrans), 

armadillos (Dasypus novemcinctus), snakes, and domestic dogs and cats (Jurries 1979). 

Abandonment caused by human disturbance, nest flooding, or unknown causes also has 

been reported (Lehmann 1941, Horkel 1979, Lutz 1979). Nest flooding has been 

observed in several studies (Lehmann 1941; Jurries 1979; Lawrence 1982; M. Morrow, 

APCNWR, unpublished data). Lehmann (1941) recounts observations recorded in his 

1938 field notes: 

“The prairie has been transformed into a miniature ocean dotted by tiny 

islands that previously had been the tops of knolls and ridges. On these 

islands sit wet and bedraggled prairie chickens and other birds that seem 

as confused and astounded as I by the sudden change in their environment. 

About a 5-inch depth of water covers the sites of nests 14 and 17, and 

former nest 15. Nest 16 has escaped by a hair’s breadth, but the lining is 

very soggy. Problems due to hawks, skunks, and other predators seem so 

petty when excessive rain destroys virtually everything at a single stroke.” 

Poor surface and internal drainage is a characteristic feature of the APC’s coastal prairie 

ecosystem resulting from low relief and dense clay subsoils (Smeins et al. 1991). In 

some years, nest losses from flooding can be substantial. For example, Lehmann (1941) 

observed a 33% loss due to flooding (n = 6) during 1938. Lawrence (1982) attributed 

abandonment of 1 of 3 active nests to flooding in 1981. Morrow (APCNWR, 

unpublished data) attributed abandonment of 4 of 15 nests during 2004 to nest flooding at 

APCNWR. Water partially covered eggs in two of these nests, and nesting materials in 

the other two nests were water soaked. One of 3 nests observed at TCPP during 2004 

was flooded (B. Crawford, TCPP, personal communication). 

Most nests are located in grasslands within 1 mile (1.6 km) of a booming ground 

(Lehmann 1941, Horkel 1979, Toepfer 1988), although nests may not be nearest to the 

booming ground on which the hen mated (Toepfer 1988). Females display fidelity to 

general nesting areas between years, although this is not the case for all hens (Toepfer 

2003). While most nests are located in grasslands, Kessler (1978) and Jurries (1979) 

found a small number of nests in fallow rice fields. These nests were generally 

unsuccessful. Ryan et al. (1998) also observed substantially decreased success for nests 

located in agricultural habitats. 

Plant species composition at nest sites varies by region and even from location to 

location within regions, but in general T. cupido requires grass for nesting habitat (e.g., 

Lehmann 1941, Hamerstrom et al. 1957, Hamerstrom and Hamerstrom 1973, Toepfer 

1988, Toepfer 2003). Toepfer (1988) indicated that nesting GPC hens seek out 

14


undisturbed residual grass cover 6-20 inches (15–50 cm) in height. Toepfer (1988) found 

63% of GPC nests in residual grass from 10−20 inches (26–50 cm), and found these nests 

to be more successful than those found in shorter vegetation. Newell (1987) and Golner 

(1997) also observed lower effective heights of vegetation at unsuccessful GPC nests. 

Buhnerkempe et al. (1984) recommended that T. cupido nesting habitat should have 90% 

of standing vegetation distributed below 16 inches (40 cm), with vegetation vertically 

dense to that point. Lutz et al. (1994), working on private ranches, found obstruction of 

vision (OV) (Robel et al. 1970) values at APC nests averaged 9 inches (23 cm), and were 

higher (P = 0.04) at successful (10 inches, 25 cm) than at unsuccessful nests (9 inches, 22 

cm). Morrow (1986), working on the APCNWR, also observed an average OV of 9 

inches (23 cm) at APC nest sites (n = 26). It should be noted that Buhnerkempe et 

al.1984, Lutz et al. 1994, and Morrow (1986) did not obtain OV data until after 

completion of the nesting attempt. Lehmann (1941) observed that rapid plant growth in 

April and May provided cover for nests that may have been relatively exposed when 

found. However, Svedarsky (1979), who determined OV measures at nests when found, 

also recommended that nesting habitat should have residual vegetation with 100% OV at 

10 inches (25 cm), and 50% OV at 14 inches (35 cm). 

Vegetation can become too tall and dense for nesting T. cupido (Westemeier 1972, 

Svedarsky 1979, Buhnerkempe et al. 1984). Speaking of Illinois GPCs, Westemeier 

(1972) stated “We have not found prairie chicken nests in any rank vegetative cover…” 

Westemeier (1972) described rank vegetation such as big bluestem (Andropogon 

gerardi), Indiangrass (Sorghastrum nutans), and switchgrass (Panicum virgatum) which 

when undisturbed “…develop a rank impenetrable layer of cane-like stems and residual 

cover.” Supporting this observation, McKee et al. (1998) found horizontal litter cover 

was the best single variable predictor of GPC nests. These authors indicated that nests 

with >25% litter cover had a failure rate twice that of nests with less litter cover. 

Svedarsky (1979) found greater litter depths at unsuccessful than at successful GPC 

nests. Morrow et al. (1996) discussed the importance of variability in grassland structure 

for nesting and brood rearing. Grasslands with tall (>39 inches, >1 m) vegetation appear 

to be avoided by nesting hens (Svedarsky 1979, Toepfer 1988), although such vegetation 

may be important for night roosting during periods of inclement weather (J. Toepfer, 

personal communication). Other nest-site vegetation threshold values identified by 

McKee et al. (1998) beyond which GPC nest success declined substantially include >5% 

woody cover, ≤5% forb cover, and ≤25% grass cover. 

Brood rearing. Hens leave the nest with their chicks after the last egg has hatched 

(Lehmann 1941). T. cupido chick weights at hatch average approximately 16–17 grams 

(0.6 oz) (Gross 1932:255; O. Dorris, Fossil Rim Wildlife Center, unpublished data). 

Griffin (1998) reported average chick weights for captive APC, GPC, and APC x GPC 

hybrids as 15.4–17.8 grams (0.5−0.6 oz), 15.4–20.4 grams (0.5−0.7 oz), and 14.6–17.3 

grams (0.5−0.6 oz), respectively. Chicks are not capable of thermoregulation until 

approximately 10–14 days (Toepfer 2003) and are brooded by the hen approximately 

50% of the daylight hours during the first week (Lehmann 1941). By two weeks of age, 

Lehmann (1941) observed that APC chicks were brooded little except early in the 

morning, during inclement weather, and at night. Toepfer (2003) stated that GPC hens 

brood their chicks for up to five weeks post-hatch. Contrary to behavior of some 

gallinaceous species, there is no evidence that prairie-chicken brood hens feed chicks or 

15


show them what to eat (Schroeder and Robb 1993). Lehmann (1941) described the 

behavior of hens leaving nests with their broods: 

“When leading chicks from the nest, old birds traveled through the lightest 

cover or followed trails, probably because heavy matted vegetation 

impeded progress and increased the chance of chicks getting lost. Cow 

trails were favorite travel ways. Chicks ranged in front, behind, and on 

both sides of the hen over an area of 1 to 5 yards in radius. Interruptions 

for sporadic feeding and for frequent brooding, which was probably more 

necessary for assembling than for warming the young, made progress 

slow.” 

Lehmann (1941) suggested this loose feeding formation resulted in chicks becoming 

separated from the brood unit. Chicks can perform weak flights by two weeks of age, 

and can fly >120 feet (36 m) by three weeks (Lehmann 1941). 

APC broods spend the first weeks after hatching in grasslands near the nest, 

typically moving less than 900 ft/day (274 m/day) (Lehmann 1941). Lehmann (1941) 

observed two broods within 0.5 mi (0.8 km) of their nests until they were 7 and 12 days 

of age. Similarly, Morrow (1986) found APC broods moved approximately 0.4 mi (0.7 

km) by 7–10 days post-hatch, and Svedarsky (1979) observed GPC broods in Minnesota 

moving an average 0.6 mi (1 km) from the nest site within two weeks. Toepfer (1988) 

observed movements by GPC broods in Wisconsin of just over 330 ft/day (100 m/day) 

during the first week and approximately 990 ft/day (300 m/day) by 14 weeks. However, 

broods are capable of movements of 1.8−2.4 mi (3–4 km) during the first week of life 

(Cebula 1966, Viers 1967, Silvy 1968, Svedarsky 1979). Newell et al. (1987) observed 5 

of 22 North Dakota GPC broods moved 1.2−9 mi (2–15 km) within 34 days of hatch. 

Newell et al. (1987) observed brood home ranges during the first 2–3 months posthatch 

averaging 1,205 ac (488 ha) (range 54−5,553 ac, 22–2,248 ha), but small areas 

within these home ranges averaging 99.8 ac (40.4 ha) were used more intensively. 

Newell (1987) observed home ranges for broods hatching from re-nests and adult hens 

were smaller than those from initial nests and sub-adult hens, respectively. Distance 

between siblings increases among brood members, progressing toward brood break-up 

(Schroeder and Robb 1993). For the APC, brood break-up begins at 6–8 weeks, although 

some chicks may remain with the hen until late October or November (Lehmann 1939, 

1941). 

Cogar et al. (1977) and Morrow (1986) found broods less than 5–6 weeks old used 

grasslands types similar to those used for nesting. Broods move away from the dense 

residual cover associated with nesting cover to less dense cover which facilitates 

movement by the hen and chicks (Lehmann 1941, Toepfer 1988). Kessler (1978) and 

Svedarsky (1979) recommended that APC and GPC brood cover respectively, should 

have sufficient canopy cover to provide shade during the summer, but be open enough at 

ground level to allow uninhibited chick movement. Jones (1963) noted the importance of 

areas dominated by forbs in supporting high insect populations, which form a large 

proportion of the chick’s diet (e.g., Lehmann 1941, Savory 1989). 

16


Toepfer (1988) found Wisconsin GPC brood rearing habitat consisted of grass or 

mixed grass in the 10−39 inch (25–100 cm) range that was undisturbed during the season 

of use, but disturbed within the past 6–24 months. Golner (1997) observed 77% of 

Wisconsin GPC brood hen locations were in grass or grass/forb cover. Night roosts for 

North Dakota GPC broods had 4-10 inch (10-25 cm) OV ( = 4 inches, 10 cm) with 

vegetation heights of 10-20 inches (25–50 cm) for 86% of observed locations (Newell et 

al. 1987). Toepfer (1988) stated that non-grass vegetation types were relatively 

unimportant to GPC hens with broods, and Newell et al. (1987) found brood hens 

avoided cash crops, especially row crops during summer. Toepfer (2003) found little 

evidence for disturbed areas being a habitat requirement for GPC broods in Minnesota, 

North Dakota, or Wisconsin. 

Starting about 4–6 weeks post-hatch, APC broods use more open habitats 

associated with mid-grass nesting cover (Cogar et al. 1977, Horkel 1979, Morrow 1986). 

Lehmann (1941) attributed this shift in habitat use to movements to areas of shade and 

surface water. Lehmann (1941) stated: 

“More than 95 percent of the more than 500 Attwater’s prairie chickens 

observed from June 24 through September 4, 1937, were in heavy cover 

within a mile, generally within less than half a mile of surface 

water….Prairie chickens require abundant shade in summer, for birds that 

were herded from such cover at midday panted vigorously, drooped their 

wings, and showed other signs of discomfort.” 

Toepfer (2003) observed that Minnesota GPC broods began feeding in agricultural fields 

(wheat and soybeans) at about six weeks of age during the day while usually night 

roosting in adjacent grasslands. 

Mortality of broods is typically high during the first four weeks after hatch 

(Lehmann 1939, 1941; Jurries 1979). Lehmann (1941) observed 50% mortality by 4−6 

weeks, and Morrow (1986) observed 62% mortality of APC brood units by 8 weeks. 

Newell et al. (1987) observed that 62.8% of North Dakota GPC chick losses occurred 

during the first 2.5 weeks. Lehmann (1939) observed that APC brood mortality was 

approximately 12% after four weeks post-hatch. Toepfer (2003) noted that once GPC 

chicks reach six weeks of age, survival to 12–16 weeks is 75-85%. Peterson and Silvy 

(1996) observed the mean number of chicks per brood reported for APCs was less (P = 

0.0001) than observed for GPCs. 

Heavy or persistent rain during the brooding season, predation, and separation from 

the brood are the most commonly reported sources of mortality for APC chicks 

(Lehmann 1939, 1941; Jurries 1979). Egg quality as influenced by nutrition of the hen, 

ability of the hen to care for chicks as influenced by hen condition, and the quality of 

brood rearing habitat relative to the abundance of insects required by chicks may 

contribute to high chick mortality during the first weeks of life (Peterson and Silvy 1996, 

Riley et al. 1998, Toepfer 2003). Fields et al. (2006) found that daily survival rates of 

prairie-chicken broods increased as broods aged, and decreased as the season progressed 

(i.e., late broods were less successful). Age of brood hens was also an important 

17


indicator of brood survival. Survival probability to 60 days was 0.49 and 0.05 for broods 

reared by adults and sub-adults, respectively (Fields et al 2006). 

Habitats Used Outside the Breeding Season 

During summer, males and hens without broods use areas where shade is available 

in the form of weeds, tall grasses, and shrubs (Lehmann 1941, Yeatter 1943, Baker 

1953). Svedarsky (1979) observed that vegetation providing dense canopy cover, 

understory openness, and forb abundance were important for GPCs during summer. 

However, Toepfer (1988) stated the greatest difference in habitat use by adults without 

broods compared to those with broods was greater use of shorter vegetation by broodless 

adults during the day, with both groups using predominantly grass or mixed grass cover. 

Kessler (1978) found APCs in the rice belt region of Texas dispersed from native prairie 

cover to surrounding forb-dominated fallow rice fields during summer months. Jones 

(1963) indicated that mid-forb communities were important for GPC day loafing cover in 

Oklahoma. Morrow (1986) observed APCs in the rice belt region using a wide variety of 

cover types during the summer–fall months. Jurries (1979) described summer months as 

a time of wandering for the APC, although Lehmann (1941) observed that once APCs 

found suitable summer cover, they moved little until fall. 

Beginning in late August–early September, flocks begin to form which move as a 

unit in their daily activities (Yeatter 1943, Schwartz, 1945, Baker 1953, Kessler 1978, 

Jurries 1979). Jurries (1979) noted APC males showed a pronounced movement back to 

booming grounds in September–early October. By approximately November 15, 

Lehmann (1941) observed APCs moved to pastures 

“….where food and cover conditions are adequate. Having found such an 

area, they remain until spring. Probably the best way to attract a good 

breeding population, therefore, is to provide suitable food and cover 

conditions during the preceding winter.” 

Morrow (1986) found selection of vegetation types by the APC during the winter was 

correlated with vegetation density. Moderate–heavy cover at least 6 inches (15.2 cm) in 

height is generally adequate to provide protection from weather and predators (Schwartz 

1945). The range of flock movements during fall and winter depended on the relative 

proximity of booming grounds, feeding areas, roost sites, and loafing areas (Schwartz 

1945). 

Food Habits 

Lehmann (1941) summarized the food habits of the APC (scientific names of 

plants have been added to Lehmann’s text): 

“The food of adult prairie chickens is about 85 percent vegetable matter 

and 15 percent animal. With young birds the ratio of vegetable to animal 

is approximately reversed. Favorite sources of plant food are ruellia 

(Ruellia spp.), perennial ragweed (Ambrosia psilostachya), blackberry 

(Rubus spp.), doveweed (Croton capitatus), and sensitive briar (Schranka 

18


spp.). Leading animal foods are grasshoppers and beetles. Greens 

(leaves, flowers, buds) are lowest in the diet in November and December; 

seeds are taken in the smallest proportions in January, February, and 

March. Insects are least frequently captured in November, December, and 

January.” 

Lehmann (1941) observed 50 species of plants and more than 65 species of insects being 

consumed by the APC. He indicated native plants were the most important source of 

food, with ruellia being the most important single food (Lehmann 1941). Lehmann 

(1941) also noted APCs used cultivated crops such as corn, peanuts, and rice as food 

sources. Kessler (1978) found APC diets consisted of more than 50% forbs in all seasons 

except fall when peanuts and rice were heavily used. Grasses and grass-like plants also 

were used heavily, but less than forbs. Kessler (1978) also observed seasonal use of 

insects, with greatest use occurring in the summer when insects were most available. 

Cogar (1980) found 74% of the APC annual diet in a predominantly rangeland ecosystem 

was composed of foliage, 18% by seeds, and 8% insects. Forbs were the primary source 

of foliage and seeds in this study. 

Lehmann (1941) observed APCs consuming water only once, despite close 

observation of birds near water and thorough examination of soft mud bordering ponds in 

inhabited prairie chicken-range. However, Lehmann (1941) writes: 

“The summer movements of prairie chickens to heavy cover near water 

are not satisfactorily explainable on the basis of cover, water, and food, 

but these habitat conditions must be provided where stable populations are 

desired….The balanced prairie chicken habitat should offer a generous 

supply of surface water throughout the year. Although Attwater’s prairie 

chickens may not be dependent on free water for survival during normal 

years…it has been established that their favorite summer range is rather 

well watered.” 

Survival and Mortality Factors 

Toepfer (1988) estimated survival of GPCs from hatch to the following May of 

approximately 25%. Horkel (1979) and Lutz (1979) observed 57% and 77% mortality, 

respectively, for color-banded APCs captured during the breeding season in 

predominantly rangeland habitats on private property. Unpublished data from APCNWR 

on the relationship between productivity and annual population change from 1988–1993 

suggested that mortality on the refuge averaged 43% in those years (M. Morrow, personal 

communication). Hamerstrom and Hamerstrom (1973) reported an average mortality of 

54% for banded GPCs in Wisconsin (n = 942), while Toepfer (1988) also working in 

Wisconsin with banded birds reported an average 51% mortality (n = 270). Therefore, 

Toepfer (2003:31) concluded that GPC survival in central Wisconsin averaged 

approximately 50%, with survival slightly higher for hens than cocks. Toepfer (2003) 

reported that preliminary analysis of survival data from radio-marked GPCs was also 

49%, comparable to that of banded birds. Morrow (1986:28) reported 36% survival of 

known-fate radioed APCs, although this value was not statistically different from 50% (P 

< 0.05). 

19


Factors which contribute to APC mortality or otherwise limit their populations 

include “natural” factors such as unfavorable weather, predators, and disease; and 

“artificial” factors such as cultivation, heavy grazing, burning, and overshooting 

(Lehmann 1941). The last APC hunting season occurred in 1936 (Jurries 1979). Except 

for hunting, the “artificial” factors primarily result in reduction of the grassland habitats 

required for prairie-chickens, although ill-timed agricultural operations may cause 

mortality of nesting hens and broods (e.g., Jurries 1979, Morrow 1986:58, Toepfer 2003). 

Encroachment of woody vegetation in the APC range has resulted in dramatic declines in 

habitat (Lehmann 1941, McKinney 1996). Lehmann (1941) observed: 

“The encroachment of mesquite, live oak, various acacias, and other kinds 

of brush onto open prairie land has been an extremely important factor in 

reducing the range and doubtless the numbers of Attwater’s prairie 

chickens….Within the memory of living men extensive prairies have been 

transformed into brush jungles.” 

A number of studies have identified adverse weather as a direct mortality factor 

for nests and young broods (Lehmann 1939, 1941; Schwartz 1945; Jurries 1979; 

Svedarsky 1979; Lawrence 1982; Svedarsky 1988; Morrow et al. 1996) although 

Peterson and Silvy (1994) found no relationship between spring precipitation variables 

they examined and proportional changes in APC populations. Lehmann (1939, 1941) 

suggested, based on examination of rainfall records, that on average 2 of 5 years were 

favorable for prairie-chicken production, 2 of 5 years were fair, and 1of 5 years was poor. 

Other adverse weather conditions that tend to have a more local impact include 

hurricanes and tropical storms, hail, and drought (Lehmann 1939, 1941, 1968). 

Predators of APC include red-tailed hawks (Buteo jamaiciencis), white-tailed 

hawks (B. albicaudatus), peregrine falcons (Falco peregrinus), Cooper’s hawks 

(Accipiter cooperi), great-horned owls (Bubo virginianus), coyotes, skunks, raccoons, 

bobcats, and domestic dogs and cats (Lehmann 1941; Jurries 1979; M. Morrow, 

APCNWR, unpublished data). Toepfer (2003) reported 69.9% of observed GPC 

predation was by raptors and 30.4% by mammals. Toepfer (1988) indicated only 

perching raptors could be considered serious predators of wild adult GPCs, and noted that 

the presence of trees in prairie-chicken habitat provided perching raptors with hunting 

opportunities. Toepfer (2003) observed 78.9% of GPC mortality in Wisconsin was 

attributed to predation, unknown causes 13.2%, electric wire collisions 6.5%, auto 

collisions 1.0%, and fence collisions 0.6%. 

Peterson (2004) conducted an extensive review of information available on 

parasites and infections diseases of prairie grouse. Peterson (2004) concluded that 

macroparasites Dispharynx nasuta and Trichostrongylus cramae; the microparasites 

Eimeria dispersa, E. angusta, Leucocytozoon bonasae, and Plasmodium pedioecetii; and 

infectious bronchitis and reticuloendotheliosis viruses (REV) have the potential to 

regulate prairie grouse populations. E. dispersa and E. angusta are coccidia while L. 

bonasae and P. pedioecetii are malarial agents (Peterson 2004). Peterson (2004) also 

indicated Histomonas meleagridis (causative agent for blackhead), Pasteurella multocida 

(causative agent for avian cholera), E. dispersa, E. angusta, and other microparasites 

20


which result in high mortality have the potential to extirpate small, isolated prairie grouse 

populations. 

Peterson et al. (1998) observed 4 of 27 (14.8%) APCs sampled were serologically 

positive for P. multocida antibodies. These four birds came from two of three remaining 

APC populations. Purvis et al. (1998) also observed specific antibodies to P. multocida 

in 3 of 53 (5.7%) northern bobwhites (Colinus virginianus) collected from the APCNWR. 

Periodic outbreaks of avian cholera have occurred in wintering waterfowl in coastal 

Texas (Peterson et al. 1998). How easily avian cholera can be transmitted from 

waterfowl to other species, including prairie-chickens is not known (Peterson et al. 1998, 

Purvis et al. 1998). Peterson et al. (1998) found T. cramae in eight of nine suitable 

samples from APC, representing the first report of this parasite in prairie grouse. Infected 

individuals, which came from all three remaining APC populations, had T. cramae 

infection intensities averaging 1,019.3, similar to that seen for T. tenuis in red grouse 

(Lagopus lagopus scoticus) (Peterson et al. 1998). T. tenuis has been experimentally 

shown to affect red grouse body condition, productivity, and survival (Peterson et al. 

1998). Purvis et al. (1998) found T. cramae in 97% of northern bobwhites collected from 

APCNWR. Peterson et al. (1998), Purvis et al. (1998), and Peterson (2004) all stressed 

the importance of determining whether T. cramae limits or regulates APC populations. 

Peterson et al. (1998) also found that one of three APC samples contained D. 

nasuta. They hypothesized that because D. nasuta is particularly pathogenic for chicks 

of other grouse species, its presence in APC populations could explain the low number of 

juvenile APCs surviving per brood as compared to GPCs (Peterson and Silvy 1996). 

Peterson (2004) also stated that while ectoparasites are relatively common on prairie 

grouse, their population-level significance is not known. Routine surveillance of APC 

blood samples from captive and free-ranging birds has yielded several positive antibody 

titers for West Nile virus (WNV), indicating exposure to this virus has occurred (J. 

Flanagan, Houston Zoo, Inc., unpublished data). Neither active disease nor mortalities 

attributed to WNV have been observed for APC or GPC in Wisconsin and Minnesota (J. 

Flanagan, Houston Zoo, Inc., J. Paul-Murphy, University of Wisconsin-Madison, and J. 

Toepfer, STCP, unpublished data). 

Disease caused by mycotoxins, pesticides, and toxic compounds also could lead 

to the extirpation of small populations (Peterson 2004). Lehmann and Mauermann 

(1963) described an account of several hundred dead APCs near a cotton field that had 

been dusted aerially with arsenic (no longer in use) as a defoliant. Flickinger and 

Swineford (1983) observed that organochlorine, polychlorinated biphenyl (PCB), and 

metal residues in APCs and northern bobwhites from cropland areas were low. 

Home Range and Movements 

Annual home range size reported by Morrow (1986) for APC hens averaged 

1,470 ac (595 ha) while those of males averaged 889 ac (360 ha). Jurries (1979) 

observed a median home range of 726 and 456 ac (294 and 185 ha), respectively, for hens 

and cocks in the ricebelt region and 1,490 and 1,796 ac (603 and 727 ha), respectively, in 

the native prairie region, where the primary land use was ranching. Morrow’s (1986) 

study was conducted at the APCNWR, and is included in Jurries’ (1979) ricebelt region. 

21


Although Lehmann (1939) stated “…Attwater [sic] prairie chickens frequently 

travel several miles in a day….”, radio telemetry data indicate that movements are 

generally more local in nature. Of 49 radioed APCs, Morrow (1986) observed only 10 

movements that were classified as “extensive”. Eight of these were by hens and averaged 

2.3 mi (3.8 km); the two “extensive” movements by males averaged 1.8 mi (3.0 km). 

Maximum cumulative APC movements observed by Morrow (1986) were 3.5 mi (5.8 

km). Lawrence and Silvy (1987) observed a maximum movement of 4.4 mi (7.3 km) for 

a translocated APC. Daily movements observed by Horkel (1979) in the native prairie 

region were greatest for cocks in December (1,914 ft, 580 m) and lowest in August (396 

ft, 120 m), while female movements ranged from 429 ft (130 m)/day in June to 1,518 ft 

(460 m)/day in November. Morrow (1986) also observed mean movements by male APC 

were greatest during December (2,845 ft, 862 m) and lowest in August (657 ft, 199 m), 

while non-reproductive (without nests or broods) female movements were greatest in 

March–May corresponding with the nesting season. Average daily movements for all 

females were least during July (700 ft, 212 m). Jurries (1979) described the breeding 

season as a period of limited daily movement. After the booming season ended, the APC 

began summer movements which Jurries (1979:23) described as “wandering”. Daily 

movements during summer were 300−500 yards (300–500 m), but cumulative 

movements of 5−10 mi (8–16 km) were observed. During September–October birds 

moved back to the vicinity of booming grounds (Jurries 1979). 

Maximum movements observed for GPCs are larger than observed for APCs. 

Hamerstrom and Hamerstrom (1973) reported 49% of cock (n = 588) and 85% of 

observed hen movements (n = 59) were at least 2 mi (3.2 km) from their “home” 

booming ground. In general, juveniles were more mobile than adults, and hens more 

mobile than males (Hamerstrom and Hamerstrom 1973, Toepfer 2003). Hamerstrom and 

Hamerstrom (1973:34) reported 38% (n = 156) and 17% (n = 222) of observed moves 

by juvenile and adult hens, respectively were greater than 5 mi (8 km). Halfmann (2002) 

observed 14% of hens (n = 88) dispersed more than 7.5 mi (12 km) from their natal areas 

prior to their first breeding season, whereas only 3% of immature cocks (n = 71) did so. 

Natal dispersal distance for hens ranged from


readily colonize unoccupied habitat. He also stated that range expansion was not likely 

to occur in the absence of increased competition associated with population increases. It 

should be noted that comparable data on dispersal distances for juvenile APCs are not 

available. 

Habitat Management 

As stressed in the foregoing sections, T. cupido requires grass and open space. 

Therefore, management for this species must be focused on providing these life requisites 

at a landscape level. Hamerstrom et al. (1957) summarized management for T. cupido: 

“Prairie chicken management is primarily grassland management: no grass, no 

chickens.” In general, Lehmann (1941) described ideal APC habitat: 

“Optimum prairie chicken range apparently consists of well-drained 

grassland supporting some weeds or shrubs as well as grasses, the cover 

varying in density from light to heavy; and with supplies of surface water 

available in summer. In short, diversification within the grassland type is 

essential.” 

A variety of management tools has been used to maintain or enhance grasslands 

for APCs and GPCs including prescribed burning, grazing, haying, mowing, herbicide 

application (for brush management), tree cutting, and cultivation (for food plots) (e.g., 

Lehmann 1941; Chamrad and Dodd 1972; Westemeier 1972; Kessler 1978; Jurries 1979; 

Morrow 1986; Svedarsky 1979, 1988; Toepfer 1988). However, because there are so 

many differences in soils, climatic conditions, and management histories that occur 

geographically and temporally across the range of T. cupido in general, and the APC in 

particular, the focus of grassland management for this species must be on the end results 

desired (i.e., habitat objectives) – not on the details of how the tools should be applied. 

Each situation will dictate how the “bag of tools” should be employed, recognizing there 

may be more than one approach to accomplish management objectives. However, clear 

elucidation of habitat objectives is essential for proper and consistent application of 

management tools. The following habitat management objectives for T. cupido in 

general, and T. c. attwateri in particular, were identified through review of the literature: 

• T. cupido management areas should be ≥33%, and preferably ≥50% grassland 

(Hamerstrom et al. 1957). 

• Priority for management should be given to habitats within 1mile (1.6 km) of 

existing and historical booming grounds (Toepfer 1988, 2003). 

• Mowing in APC habitat should not occur before 1 July (Lehmann 1941). 

• Prescribed burning should be completed in APC habitat by 1 February (Lehmann 

1941:56). 

• Availability of grasslands for nesting and brood rearing cover most often limit T. 

cupido populations (Hamerstrom et al. 1957:18, Morrow 1986:91, Toepfer 

1988:482). As such, 

o No more than 33% (Toepfer 1988) to 60% (Lehmann 1941) of grassland 

habitat managed for T. cupido should be burned on an annual basis. 

23


o Patches of unburned cover should be as large as possible, but at least 80 

(Toepfer 1988) to 618 ac (32−250 ha) (Kessler 1978) (but see Lehmann 

1941). 

o More than 50% of grassland residual cover (still standing from growth of 

previous seasons) should be 10−39 inches (25–100 cm) in height during 

spring (Toepfer 1988). Cover with OV values averaging 10 inches (25 

cm) should be readily available and well distributed within grasslands as 

nesting sites (Cogar et al. 1977, Svedarsky 1979, Morrow 1986, Lutz et al. 

1994). 

o Cover which becomes rank [>39 inches (1 m) tall (Buhnerkempe et al. 

1984), >25% horizontal litter cover (McKee et al. 1998)] should be 

disturbed by burning, grazing, or mowing (Westemeier 1972, Toepfer 

1988, McKee et al. 1998). 

o Brush must be carefully controlled to prevent excessive encroachment into 

grassland habitats (Hamerstrom et al. 1957, Toepfer 1988, DeHart 2003), 

and trees, especially near booming grounds should be cut down 

(Svedarsky 1979, Toepfer 2003). Less than 25% of the landscape should 

be wooded, with woodlands in scattered blocks (Hamerstrom et al. 1957). 

In order to support booming grounds, open grasslands must be as large as 

possible. At a minimum, open grasslands of >1,480 ac (600 ha), and 

preferably >2,175 ac (880 ha) should be maintained (Toepfer 1988). 

• Although some disagreement exists regarding the value of food plots in APC 

management, agricultural crops are probably not essential for APCs because of 

their southern distribution (Lehmann 1937, 1941). Although Cogar (1980) 

observed only slight use by APCs of available grain sorghum, agricultural crops 

are usually readily used when available (Lehmann 1941, Kessler 1978, Jurries 

1979, Morrow 1986). Therefore, 10−15 ac (4–6 ha) food plots distributed at a 

density of approximately one for every three booming grounds may be provided 

(Toepfer 1988). Food plots should be carefully monitored for aflatoxin 

development in crops such as peanuts, soybeans, corn, and cereal grains (Fraser et 

al. 1991). While Peterson (2004) found no records of prairie grouse mortality 

caused by mycotoxins, hot, humid conditions characteristic of APC range are 

ideal for growth of aflatoxin-producing Aspergillus fungi (Reddy and Waliyar 

2000, Larson 2002). Aflatoxicosis associated with waste corn has caused 

wintering goose mortality in Colorado County, Texas, within 20 mi (32 km) of 

APCNWR (M. Morrow, APCNWR, unpublished data). Birds appear to be more 

susceptible than mammals to aflatoxicosis (Davidson and Nettles 1988). 

Lehmann (1939) summarizes habitat management for APC: 

“Moderately grazed and moderately burned grassland…provides prairie 

chickens with everything they need in all seasons. It is therefore upon the 

existence of adequate prairie habitat that the welfare of the prairie chicken 

depends.” 

24


F. CRITICAL HABITAT 

Critical habitat has not been designated for the APC. 

G. ON-GOING CONSERVATION EFFORTS 

Research 

Prior to the late 1960s, APC conservation efforts consisted of life-history research 

(Lehmann 1941), periodic population surveys (Lehmann 1941, Lehmann and Mauermann 

1963, Lehmann 1968), and protection from hunting (since 1937) (Lehmann 1941, Jurries 

1979). Beginning in 1967 through the present, a multitude of research projects has been 

conducted primarily at TAMU on topics including life history, habitat management, 

predator management, genetics, limiting factors, captive breeding, and population 

supplementation (Morrow et al. 2004). Silvy et al. (1999) provided a review of much of 

this research. In 1969, TPWD initiated a series of research projects that addressed a 

range of basic life history and population inventory issues. This research culminated in a 

monographic work on the APC (Jurries 1979). 

Habitat Management 

The APCNWR was established in 1972 to protect and enhance the APC’s 

severely diminished prairie habitat. This refuge contains 10,538 ac (4,265 ha), including 

2,500 ac (1,027 ha) added since 1998. Most of the recently acquired lands were formerly 

in rice production, and need restoration to provide optimal prairie-chicken habitat. APC 

populations on the refuge have ranged from an estimated 25 when the refuge was 

established to 222 in 1987 (APCNWR unpublished data). The refuge population has 

declined since 1987, corresponding to range-wide population declines (Figure 1). 

Morrow et al. (1996) discussed factors affecting the refuge decline. They observed 

acreage burned within the APC’s core habitat, variability in grassland structure, offrefuge 

APC population changes, and several climate variables were correlated with APC 

population changes on APCWNR. 

Even though recovery plans (USFWS 1983, 1993) emphasized the need for 

habitat protection and restoration in geographically separate areas, little habitat protection 

or management was accomplished other than at APCNWR until approximately 1990. 

Since then, considerable effort and funds have been spent in cooperative private-lands 

projects. Initially, these efforts were spear-headed by TPWD with federal aid to states 

dollars made available through Section 6 of the ESA. Beginning in 1995, an initiative 

was undertaken with the primary mission of restoring native prairie grasslands within the 

APC’s former range. This effort, now known as the Coastal Prairie Conservation 

Initiative (CPCI) is a diverse partnership effort involving private landowners, local soil 

and water conservation districts, the USFWS, the Sam Houston Resource Conservation 

and Development Board, the U.S. Natural Resources Conservation Service (NRCS), 

TNC, TPWD, and the Grazing Lands Conservation Initiative (GLCI). Integral to the 

CPCI has been incorporation of Safe Harbor Agreements into management plans where 

desired by cooperators. The purpose of Safe Harbor Agreements is to promote voluntary 

25


management for federally listed species on private property while giving assurances to 

landowners that no additional future regulatory restrictions will be imposed if these 

species colonize or increase in numbers as a result of management activities. As of 

October, 2006, approximately 76,681 ac (31,045 ha) have been enrolled under Safe 

Harbor Agreements for APC management, with cost-share assistance provided on 

approximately 66,626 ac (26,974 ha) (T. Anderson, USFWS, personal communication). 

In addition, TPWD and NRCS landowner assistance agreements have been implemented 

on several thousand acres for the purpose of restoring coastal prairie habitat within the 

APC’s former range. 

TNC took ownership of the TCPP in 1995 through a donation from Mobil Oil 

Corporation. Since 1985, the APC population on this site has numbered fewer than 50 

individuals (Morrow et al. 2004). TCPP and APCNWR currently contain the last freeranging 

APC populations. Both of these populations have been supplemented with 

releases of captive-reared birds since 1996, although net additions from released birds to 

the TCPP population have been minimal since 1999. 

Currently, APC restoration efforts are focused in three priority management zones 

(Figure 3). The 2,396-ac (970-ha) TCPP supports the only population still containing 

wild-hatched birds. Although remnant prairies exist in the area that could be restored 

through removal of woody species [primarily Chinese tallow, (Sapium sebiferum)], 

management potential for this area is limited because of rapid urbanization. Current 

land-use data for this area is lacking. The Austin-Colorado County priority management 

zone, which historically supported relatively large APC populations (Lehmann 1941, 

1968; Appendix 1), contains the 10,538-ac (4,265-ha) APCNWR. The boundary for this 

zone represents the 58,193-ac (23,560-ha) priority acquisition area for the APCNWR 

(Figure 4). This does not reflect plans for future refuge boundaries, but delineates an area 

where land acquisition should be focused. Currently, the approved target acquisition size 

for this refuge is 30,000 ac (12,145 ha). The Austin-Colorado County priority 

management zone contains 80% of the minimum convex polygon that encompassed the 

occupied range defined by booming ground distributions from 1979-1992 (McKinney 

1996). Based on most recent land-use data available (1990 data from McKinney 1996), 

the Austin-Colorado County priority management zone contains approximately 17,806 ac 

(7,209 ha) of grass (31% of total area). A total of approximately 15,525 ac (6,285 ha) 

(27% of the total area) are currently under grassland management or restoration in this 

area (Figure 4). 

The 663,670-ac (268,690-ha) Refugio-Goliad County priority management zone 

also historically supported large prairie-chicken populations (Lehmann 1941, 1968; 

Appendix 1) and contains the largest contiguous blocks of coastal prairie remaining in 

Texas (Figure 3). The boundary for this zone was delineated by TNC (Miller and 

Halstead 2003). The minimum convex polygon delineating 1979–1992 booming ground 

distributions for the APC population in this vicinity (McKinney 1996) is almost entirely 

contained within the zone (Figure 5). This 162,785-acre (65,905-ha) former booming 

ground range currently contains three relatively contiguous grassland blocks totaling 

approximately 50,550 acres (20,445 ha), or 31% of the historic booming ground range 

(Figure 5; W. Harrell, TNC, unpublished data). Currently, 58,948 ac (23,865 ha) (8.9%) 

26


Figure 4. Land use within the Austin-Colorado County, Texas priority management zone. 

27


Figure 5. Land use within the Refugio-Goliad County, Texas priority management zone. 

28


of the Refugio-Goliad priority management zone are under prairie restoration through the 

CPCI (T. Anderson, USFWS, personal communication) (Figure 5). 

Captive Breeding 

Mass propagation of most grouse in captivity has proven particularly difficult 

(McEwen et al. 1969, Johnson and Boyce 1991). The first documented attempt at 

breeding and rearing APC in captivity occurred at the TAMU Poultry Science 

Department (Watkins 1971). Thirteen male and 13 female APCs were trapped from the 

wild during December 1967–January 1968. Two hens laid 10 eggs in spring 1968. Two 

of these eggs hatched, four were infertile, and four died as embryos. During spring 1969, 

one hen produced seven eggs. Three of these eggs were infertile, two died as embryos, 

and two hatched. No data were available on chick survival, but Watkins (1971:3) 

observed that none of the chicks were as strong and active as expected of precocial 

young. An additional 10 female and 8 male APCs were trapped from the wild in 

February–March 1970. In addition, 21 eggs were collected from wild nests that year. Of 

the 18 adults taken into captivity, six died within three weeks. The remaining birds 

produced 18 eggs, thought to have been produced by two hens. All eggs were infertile. 

Eight of the remaining adults eventually succumbed to Newcastle’s disease, fowl pox, 

and injury. Nineteen of the 21 wild-collected eggs hatched, but one chick drowned in a 

water pan of the incubator. Sixteen of the remaining 18 died within one week post-hatch. 

APC captive breeding was not attempted again until 1992. By that time, the wild 

population had already declined to an alarmingly low level of 456 birds, and the need for 

an aggressive captive breeding program was established to (1) preserve as much of the 

genetic representation of the wild stock remaining as possible, and (2) provide stock to 

re-populate depleted or extirpated populations. Fossil Rim Wildlife Center, after working 

with GPCs during the previous year, received 4 clutches totaling 49 eggs (1 clutch from 

APCNWR and 3 from what is now TCPP). While hatchability of the eggs was good 

(86%), the chicks proved very difficult to rear. Only 5 (3 females, two males) of the 42 

hatched chicks survived to the following breeding season. While previous rearing 

attempts with GPC chicks reported reduced survivability related to feeding problems 

during the first few days of life (Kruse 1984), most problems experienced at Fossil Rim 

during this first year were related to development of enteritis. Various attempts at 

antibiotic therapy were ineffective. 

More wild collected eggs (29) were transported to Fossil Rim in 1993. However, 

15 eggs were infertile, including 1 entire clutch from APCNWR. Of the 14 remaining 

eggs, 12 hatched. Additionally, 50 eggs (30 viable) produced by chicks reared in 1992 

resulted in 26 chicks. However, despite aggressive treatment by Fossil Rim staff, 

enteritis continued to be problematic with respect to chick survival. 

In part due to recommendations resulting from a population and habitat viability 

analysis conducted for the APC (Seal 1994) to substantially expand the captive breeding 

effort, two additional facilities began rearing APCs in 1994. Twenty-three wild collected 

eggs were provided to the TAMU Department of Wildlife and Fisheries Sciences and 26 

to the Houston Zoo. TAMU had been working with GPCs since 1991. The Houston Zoo 

also had previous experience with GPCs. In addition, Fossil Rim continued to rear APC 

29


from their captive flock. The addition of more insects to the chick diet appeared to 

improve chick survival. Overall, 36 chicks were reared to 8 weeks from 52 hatched eggs. 

The San Antonio Zoo was added as a breeding facility in 1996. They received 

breeding stock from other facilities, two wild-caught males, and one clutch of 12 eggs 

from APCNWR. Five chicks were reared to eight weeks of age from these eggs. San 

Antonio experienced periodic mortalities of older chicks and adults over the next two 

years from a Clostridial enteritus. After intensive research, it was concluded that soil in 

the breeding pens was contaminated with the Clostridium bacterium causing the observed 

enteritis deaths. Therefore in 1998, all APC were removed from the San Antonio facility 

to allow for complete sterilization of pen infrastructure and substrate. No mortalities 

from Clostridial enteritis have been observed at that facility since. Sea World of Texas, 

the Abilene Zoo, and the Caldwell Zoo were added as breeding facilities in 1999, 2000, 

and 2002, respectively. As of the spring 2007 breeding season, approximately 50% of 

the captive flock will be housed at Fossil Rim Wildlife Center (H. Bailey, APC SSP 

Coordinator, Houston Zoo, Inc., personal communication) (Figure 6). Additional 

breeding facilities are being sought to dilute the risk of a potential catastrophe to the 

breeding program and to increase the capacity of the program to produce more birds for 

release into suitable habitats. 

In summary, a total of 175 eggs (representing 14 clutches) and 9 males were 

collected from wild populations for inclusion in the captive breeding program during 

1992–1998. Because not all of these founders survived to contribute offspring, and 

conservative assumptions were made about the relatedness of founder individuals, the 

captive population was derived from 19 founders representing 8.5 founder genome 

equivalents. 

Comparing the survivorship at various stages of the APC captive breeding process 

with two GPC mass propagation efforts, the APC program has performed comparably to 

Kruse (1984) and substantially better than McEwen et al. (1969) with respect to egg 

viability, hatchability, and chick survival (Figure 7). However, the average 11 eggs/hen 

observed in the APC program is substantially less than the average 23 eggs/hen observed 

by Kruse (1984) (Figure 8). 

Medical issues most commonly encountered by the APC breeding program 

include poor chick survival during the first days post-hatch, enteritis (particularly among 

young chicks), wryneck in newly hatched chicks, dispharynxiasis, capillariasis, leg 

rotations among growing chicks, curled toes, self-induced trauma from collision with pen 

structures, gastrointestinal tract obstruction by impacted vegetation, and REV (J. 

Flanagan, DVM, APC Recovery Team Veterinary Advisor, Houston, Zoo, Inc., personal 

communication). Development of protocols for prophylactic treatment of macroparasites 

has largely minimized their impacts on the captive flock. Although both enteritis and leg 

rotations may have multiple causes, research currently focused on nutritional quality of 

breeder and chick diets will hopefully lead to reductions in the incidence of these 

maladies. Wryneck also results from various causes, but analysis of historic captivebreeding 

records strongly suggests a genetic influence (K. Willis, APC SSP Small 

Populations Advisor, Minnesota Zoo, personal communication). 

30


Figure 6. Projected Spring 2007 distribution of captive Attwater’s prairie-chickens by location (n = 164). 

San Antonio 

6% 

Sea World 

5% 

TAMU 

5% 

Houston 

20% 

Fossil Rim 

50% 

Abilene 

9% 

Caldwell 

5% 

31


Figure 7. Comparison of Attwater’s and greater prairie-chicken mass propagation efforts. 

80% 

70% 

60% 

50% 

40% 

30% 

APC (1996-2005) 

GPC - Kruse (1984) 

GPC - McEw en et al. (1969) 

20% 

10% 

0% 

% Eggs Viable % Hatched % Chicks to 8 wks 

32


Figure 8. Comparison of Attwater’s (1996−2005) and greater prairie-chicken (1972−75) (Kruse 1984) captive egg production. 

35 

30 

25 

20 

15 

APC (1996-2005) 

GPC - Kruse (1984) 

10 

5 

0 

1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 

33


Approximately 34% of hatched APC chicks die during the first 10 days of life 

(Figure 9) (K. Willis, APC SSP Small Populations Advisor, Minnesota Zoo, unpublished 

data). Cause of death for these chicks is most commonly attributed to enteritis or “failure 

to thrive”. Johnson and Boyce (1991) observed 85% mortality of sage grouse 

(Centrocercus urophasianus) chicks produced by captive hens within a few days of 

hatch. Most of these deaths were attributed to peritonitis (Johnson and Boyce 1991). In 

contrast, 84% of chicks collected from the wild, or produced from wild-collected eggs 

survived 7−9 weeks. These authors speculated that inadequate hen diets in captivity may 

have contributed to low chick viability. Kruse (1984:12) observed that 78% of mortality 

observed in their GPC propagation program occurred during the first week of life, and 

was usually associated with the chick’s reluctance to eat. Drake (1994) and Griffin 

(1998) stressed the importance of insects in the diet to survival of captive APC chicks. 

The importance of insects in the diet of other captive-reared gallinaceous species, 

especially early in chick growth, has also been observed (e.g., Thomas 1987, Johnson and 

Boyce 1990). Johnson and Boyce (1990) observed that 25 sage grouse chicks hatched in 

captivity not provided with insects all died within 4−10 days, whereas all chicks provided 

with insects survived the initial 10 days. 

The inability to effectively manage major outbreaks of REV which have occurred 

in recent years at San Antonio Zoo and Fossil Rim Wildlife Center has severely 

hampered APC propagation efforts. TAMU and University of Georgia continue to 

research the etiology, testing protocol, and management of REV. Researchers at TAMU 

believe they are very close to developing a vaccine for this disease (E. Collison, 

Department of Veterinary Pathology, TAMU, personal communication). 

REV was first diagnosed in GPCs at TAMU in September 1993 (Drew et al. 

1998). Initial testing of captive GPCs and APCs at TAMU in December 1994 revealed 

that >50% of the flock were viremic (Drew et al. 1998). Although REV was ultimately 

detected at the Houston Zoo and Fossil Rim a year or two later, monitoring efforts 

precipitated by experiences and research efforts at TAMU prevented the disease from 

spreading beyond a few birds at each facility until 2002. In 2002, an REV outbreak 

coupled with an outbreak of avian pox at San Antonio Zoo ultimately required their flock 

be euthanized to affect control. Fossil Rim experienced an outbreak of REV and avian 

pox in November, 2003, which also proved very difficult to contain using the standard 

test and cull protocol. 

Population Supplementation 

By 1995, the captive flock had grown to a point that 13 excess males were 

available for a pilot release at APCNWR. From 1995−2006, a total of 1,005 captivereared 

APCs has been released at APCNWR and TCPP (Figure 10). Most of these birds 

have been fitted with radio transmitters (


Figure 9. Captive Attwater’s prairie-chicken mortality during the first month post-hatch (K. Willis, APC SSP Small Populations Advisor, 

Minnesota Zoo, unpublished data). 

0.07 

0.06 

0.05 

Proportion Dying 

0.04 

0.03 

0.02 

0.01 

0 

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 

Days Post-Hatch 

35


Figure 10. Captive Attwater’s prairie-chickens (n = 1,005) released at the Attwater Prairie Chicken National Wildlife Refuge (APCWR), 

Colorado County, Texas and the Texas City Prairie Preserve (TCPP), Galveston County, Texas from 1995−2006. 

160 

140 

Number Released 

120 

100 

80 

60 

40 

20 

0 

1995 

1997 

1999 

2001 

2003 

2005 

TCPP 

APCNWR 

Year 

36


(www.isis.org) and Population Management 2000 (Pollak et al. 2002) software, were 

placed in 30 X 50 ft (9.1 X 15.2 m) holding pens at the release site to recover from the 

stress of transport and to acclimate to release site surroundings. At the end of the 

acclimation period, pen doors were opened and birds generally allowed to exit at their 

own pace. Food and water were provided outside the release pens for up to 30 days postrelease 

to allow for a gradual transition to a natural diet. Findings to date suggest (1) 

mortality during the first 30 days was more than five times higher in birds acclimated for 

3 days versus 14 days (P < 0.005); (2) birds released during seasons when migrant 

raptors were present (1 October–19 April) experienced 1.9 times higher mortality during 

the first 30 days than those released when migrant raptors were absent (P < 0.005); no 

difference (P > 0.05) has been detected in post-release survival attributable to the age of 

birds at release (after hatch-year versus hatch-year) (APCNWR, unpublished data). 

Lockwood et al. (2005) observed movements and monthly ranges of released 

APCs were similar to those of wild APCs. Lockwood (1998) also observed use of habitat 

structure by released birds was comparable to wild birds observed by Morrow (1986) and 

Horkel (1979). Hess (2004:10) observed no difference in flight speed between penreared 

APCs released at APCNWR and wild GPCs from Kansas and Minnesota, but 

observed that wild GPCs flew farther when flushed than pen-reared APCs ( ≥ 250 m 

versus ≤ 97 for GPCs and APCs, respectively). Hess (2004) observed wild GPCs and 

APCs >365 days post-release flushed at greater distances from humans than those


close. The pen environment conditions prairie-chickens to run rather than flush (Toepfer 

1988). 

Kaplan-Meier estimates of annual post-release survival have ranged from 9% 

(1999) to 35% (1996) ( = 19%) (Morrow et al. 2004). Since 2000, annual survival 

estimates have been more consistent (13–18%, = 15%) (APCNWR unpublished data). 

Survival of released pen-reared stock has typically been low for many species (e.g., 

Roseberry et al. 1987, Toepfer 1988, Hernandez et al. 2006). Toepfer et al. (1990) noted 

that of upland game, prairie grouse have the poorest record with regard to establishment 

of populations with translocated birds. Toepfer (1988) reported that 90% of released penreared 

GPCs were dead within 90 days, and none survived 120 days. Survival of released 

APCs to 120 days through 2004 averaged 51%, ranging from 76% in 1996 to a low of 17 

and 18% in 2000 and 1999, respectively (APCNWR, unpublished data). Survival to 120 

days for all birds released during 2001-2004 averaged 58%, ranging from 49–71%. 

Despite higher mortality observed in released pen-reared birds compared to wild 

cohorts, enough have survived to produce viable nests during the spring following 

release. Number of nests from released birds has ranged from two in 2001 to 19 in 2004 

(APCNWR, unpublished data). However, documented survival of offspring from these 

nests has been extremely poor to non-existent (Lockwood et al. 2005; APCNWR, 

unpublished data). Nesting success has been enhanced by installation of predator 

deterrent fences around most nests since 2000 (J. Toepfer, Society of Tympanuchus 

Cupido Pinnatus, and M. Morrow, APCNWR, unpublished data). Nesting success at 

APCNWR from 2000–2004 averaged 64% for fenced nests (n = 36) and 0% for unfenced 

nests (n = 9) (APCNWR, unpublished data). In 2003, intensive observations on eight 

broods at the APCNWR found no chicks survived past 11 days post-hatch (APCNWR, 

unpublished data). Several chicks were found dead or dying at night roosts, suggesting 

that predation was not the sole cause of chick mortality. 

This type of mortality has also been observed in wild GPCs, but with much lower 

incidence (J. Toepfer, STCP, personal communication). Previous attempts to allow 

captive APC hens to parent-rear their chicks have ended in failure unless the brood was 

confined to a relatively small space (Drake 1994, J. Carviotis, Houston Zoo, Inc., and N. 

Silvy, Department of Wildlife and Fisheries Sciences, TAMU, personal communication). 

Increased, although still limited, chick survival was achieved at APCNWR during 2003- 

2006 by confining hens with their broods to an approximately 4 x 8 x 2-ft (1.2 x 2.4 x 

0.6–m) pen immediately after hatch. These broods were provided insects (collected from 

area herbaceous vegetation) and water (in most cases) ad libitum, and released with the 

hen at approximately 2−4 weeks post-hatch. A limited number of similarly-aged chicks 

hatched in captivity were also added to confined APCNWR broods to evaluate the 

efficacy of artificially increasing the number of chicks/brood, especially for smaller 

broods. The limited evaluation of this technique to date suggests that this technique may 

be useful in future recovery activities (APCNWR, unpublished data). Poor survival of 

chicks produced by released captive-reared APCs is currently the single-most factor 

limiting significant progress toward recovery. 

Poor productivity in other species of released captive-reared birds has been 

observed (Leopold 1944). Leopold (1944) observed differences in the size of the brain, 

38


pituitary, and adrenals of domestic-strain and wild turkey poults, and Liukkonen-Anttila 

(2001) observed differences in gut dimensions and liver weights of wild and captive grey 

partridge (Perdix perdix) and capercallie (Tetrao urogallus). Liukkonen-Anttila 

(2001:37) stated that “According to Dahlgren (1987) the effects of an insufficient diet 

during growth may last until the first breeding season.” Grindstaff et al. (2005) suggested 

that resource limitation may have transgenerational effects on immune function. 

Lochmiller et al. (1993) found that deficiencies in northern bobwhite (Colinus 

virginianus) chick diets can lead to atrophy or suppressed development of primary and 

secondary lymphoid organs. Thomas (1987) speculated that survival of captive-reared 

birds in the wild may depend on a digestive system conditioned anatomically and 

physiologically to bulky, low quality diets as compared to commercial diets designed for 

maximum digestibility. Thomas (1987) also hypothesized that providing monotypic, 

nutrient-dense commercial rations may preclude development of optimal foraging and 

food selection behaviors needed for survival in the wild. These observations suggest that 

anatomical, physiological, and behavioral changes attributable to the captive environment 

may contribute to poorer survival and reproduction of released pen-reared birds 

compared to wild birds. 

H. REASONS FOR LISTING/CURRENT THREATS 

Section 4(a)(1) of the 1973 ESA (50 CFR 17.11) requires a 5-factor analysis of 

threats to endangered species. A general discussion of threats facing APC within this 5- 

factor framework is presented in the following sections. This 5-factor analysis is also 

used in Section II.D (Reduction or Alleviation of Threats) of this plan as a framework for 

evaluating how proposed recovery actions address recognized threats. 

Factor 1: The present or threatened destruction, modification, or curtailment of 

habitat or range. 

Remaining populations have become geographically fragmented and genetically isolated 

(Toepfer 2003; Johnson et al. 2003, 2004). This scenario, observed at the continental 

scale, was manifested regionally with regard to the APC (Lehmann 1941, Jurries 1979, 

Lawrence and Silvy 1980, McKinney 1996, Morrow et al. 2004, Silvy et al. 2004). 

Smeins et al. (1991) estimated


Currently, illegal take by dove, quail, and waterfowl hunters is a possibility, but no 

evidence exists that this has occurred in >10 years. 

Factor 3: Disease or predation. 

Disease. Peterson (2004) provided an extensive overview of information 

available on parasites and infectious diseases of prairie grouse, including the APC. This 

overview was summarized previously in Section I.D (Survival and Mortality Factors). 

Briefly, Peterson (2004) concluded that macroparasites Dispharynx nasuta and 

Trichostrongylus cramae; microparasites Eimeria dispersa, E. angusta, Leucocytozoon 

bonasae, and Plasmodium pedioecetii; and infectious bronchitis and reticuloendotheliosis 

viruses (REV) have the potential to regulate prairie grouse populations. Peterson (2004) 

also indicated Histomonas meleagridis (causative agent for blackhead), Pasteurella 

multocida, E. dispersa, E. angusta, and other microparasites which result in high 

mortality have the potential to extirpate small, isolated prairie grouse populations. 

Positive serologic tests for P. multocida, and infestations of T. cramae and D. 

nasuta have been documented from wild APC samples (Peterson et al. 1998). Diligent 

prophylaxis in the captive setting is required to prevent significant mortality from D. 

nasuta (J. Flanagan, Houston Zoo, Inc., personal communication). REV remains 

problematic in captivity (see Section I.G. Captive Breeding). REV has been documented 

in free-ranging APC populations (Drew et al 1998; M. Morrow, APCNWR, unpublished 

data), but its population-level impacts are unknown. Insects are possible sources of REV 

infection through mechanical transmission of the virus (Davidson and Braverman (2005). 

Peterson (2004) stressed that wildlife managers need ecologically-based studies which 

address the potential significance of parasites and diseases to prairie grouse populations, 

especially for small isolated populations subject to stochastic extinction. Hudson et al. 

(2006) speculated that increased temperatures and climatic disruption brought about by 

global warming will result in increased frequency and intensity of outbreaks of some 

parasite populations like Trichostrongylus tenuis. 

Predation. APC predation, including nests, has been discussed in previous sections. 

Predation of captively-reared released birds is heavy in some years, especially during the 

first weeks following release (Lockwood 1998, 2005; M. Morrow, APCNWR, 

unpublished data). However, survival of released APC has been higher than that seen 

with other similar efforts (e.g., Roseberry et al. 1987, Toepfer 1988). Survival of 

released birds still living one year post-release approximates the 50% survival expected 

from wild T. cupido populations. Pre-release conditioning and modification of rearing 

techniques to enhance post-release predator avoidance behaviors may offer promise for 

increasing post-release survival (e.g., see van Heezik et al. 1999, Hess 2004). However, 

survival of broods from released hens in the wild is currently the factor most limiting 

APC recovery, not post-release survival. Current thinking is this poor brood survival is 

related to physiological, behavioral, or habitat quality issues rather than predation (see 

discussion in Section I.G. Population Supplementation). Peterson and Silvy (2004) 

observed that spring breeding success, measured by summer juvenile:adult ratios, drives 

APC population changes. Such ratios are a composite not only of brood survival, but 

also of nesting success. Peterson and Silvy (1996) observed both APC nest success and 

brood survival were lower than those for GPC. Predation likely plays a role under some 

40


environmental conditions to APC’s relatively poor reproductive success. Therefore, due 

diligence is necessary in managing predation, especially that associated with reproductive 

success. 

Factor 4: Inadequacy of existing regulatory mechanisms. 

As was the case with many species, the APC become endangered primarily due to 

habitat loss that occurred prior to the enactment of current conservation legislation. In 

general, the current legal framework (e.g., ESA, National Environmental Policy Act) 

provides for adequate protection and conservation. However, in some cases agency 

programs may conflict with APC recovery. For example, the NRCS Environmental 

Quality Incentive Program (EQIP) not only provides cost share assistance to landowners 

for management practices that improve habitat for APC (e.g., brush and grazing 

management), but it also provides assistance to landowners for practices that degrade or 

destroy APC habitat (e.g., converting native grasslands to exotic tame pasture grasses). 

Factor 5: Other natural or manmade factors affecting its continued existence. 

Population fragmentation. Small, fragmented populations are generally at 

greater risk of extinction than large, contiguous ones (e.g., Shaffer 1987, Fahrig and 

Merriam 1994, Frankham et al. 2002, Toepfer 2003). Further, r-selected species like the 

prairie-chicken are subject to relatively high inherent mortality, resulting in volatile, and 

sometimes catastrophic changes in populations when faced with environmental 

perturbations (Pianka 1970, Odum 1971). Population and habitat viability analyses 

conducted for the APC (Seal 1994) have indicated a high probability of extinction within 

20 years (Seal 1994, Brooks et al. 2002) given the small, isolated nature of APC 

populations. 

Genetics. Genetic variability has been correlated with population fitness and 

viability; isolated small populations lose genetic variation through genetic drift which 

may then result in inbreeding depression or an increased susceptibility to diseases and 

parasites (Frankham et al. 2002; Reed and Frankham 2003; Spielman et al. 2004a, 2004b, 

Whiteman et al. 2006). Bouzat et al. (1998a, 1998b) and Westemeier et al. (1998) 

demonstrated a reduction in fitness associated with reduced genetic variation in a small 

Illinois GPC population. Reduced genetic variation has also been attributed to population 

isolation in Wisconsin GPCs (Bellinger et al. 2003; Johnson et al. 2003, 2004; Johnson 

and Dunn 2006). 

The genetic viability of remaining APC populations, both in the wild and captive 

setting is of interest for conservation and management purposes. Osterndorff (1995) 

assessed genetic variability in the three remaining wild populations during 1991–1994. It 

was during this time that many founders for the captive population were collected (see 

Section I.G. Captive Breeding). Osterndorff (1995) found the Galveston County (TCPP) 

and the Colorado County (APCNWR) populations exhibited genetic similarity indices, 

63% and 30% greater respectively, than observed for Kansas GPCs (i.e., Galveston and 

Colorado County APC populations had less genetic variability than Kansas GPCs) (P < 

0.01). The genetic similarity index for the Refugio County population was not different 

from the GPC sample. Osterndorff (1995) found the composite sample combining the 3 

41


extant APC populations had a higher genetic similarity index than did the Kansas GPC 

sample (P < 0.001). Stoley (2002) examined APC samples collected from wild 

populations during 1936–1965 and the captive flock from 2000 and found that observed 

levels of microsatellite DNA heterozygosity were less than expected, even in the historic 

sample of wild birds. However, the historic levels of observed heterozygosity may be 

higher than reported by Stoley (2002) because this study did not assess potential allelic 

dropout issues, which is a common problem with historic samples when amplifying 

nuclear loci (see Miller and Waits 2003). Johnson and Dunn (2006) observed high 

mtDNA control region haplotype diversity (h=0.912, n = 19) using Attwater’s samples 

from museum specimens collected between 1887-1942, while Palkovacs et al. (2004), 

also working with mtDNA control region, found that a sample (n = 8) of captive 

Attwater’s had the lowest values for three measures of molecular diversity when 

compared to heath hens and greater prairie-chickens. In fact, in a report of preliminary 

analyses, Jeff Johnson (Univ. of Michigan, unpublished data) used mitochondrial DNA 

and microsatellite techniques to assess genetic variability in APC samples collected from 

Colorado, Galveston, and Refugio counties during 1992–1994. He found that 

“… analyzed as three groups, levels of mitochondrial haplotype variability 

and diversity were quite low, with the exception of haplotype diversity for 

Colorado County… The microsatellite DNA analysis agrees with the 

above DNA results. Compared to a large number of surveyed GPC 

populations, the three counties sampled in 1992−1994 had significantly 

low levels of genetic variability based on mean number of alleles 

(P 0.05) in 

allelic richness (4.3 + 0.5 (SE) vs. 5.2 + 0.8 (SE) for captive vs. wild, respectively) or 

heterozygosity (0.696 + 0.034 (SE) vs. 0.642 + 0.067 (SE) for captive vs. wild, 

respectively) (J. Johnson, University of Michigan, unpublished data). 

An effective population (N e ) of 500-5,000 breeding individuals has been 

suggested to balance the effects of genetic drift (Franklin 1980, Lande and Barrowclough 

1987, Lande 1995, Franklin and Frankham 1998, Lynch and Lande 1998). Because as 

few as 10% of prairie-chicken males breed (Robel 1970), a prairie-chicken population of 

>2,750 individuals would be required to maintain a N e >500 assuming a 1:1 sex ratio. 

(Roughgarden 1979:68, Walk 2004:53). 

To date, there is no direct evidence that APC populations have experienced 

inbreeding depression. Although Peterson and Silvy (1996) reported APC reproductive 

parameters were substantially lower than observed for the GPC, these differences in 

productivity may be lineage specific because similar parameters were also observed when 

the first major APC life history work was conducted more than 70 years ago (Lehmann 

1941). Thus these differences, while important, may not likely be classified as recent 

inbreeding depression. Further, Griffin (1998) observed no substantial improvement in 

42


fertility, hatchability, or chick survival in captivity for GPC x APC hybrids when 

compared to comparable nonhybrid groups of GPCs and APCs, although outbreeding 

depression can not be ruled out in this case (e.g., Edmands 2007). A comparison of 

chicks from wild-collected GPC eggs (n = 27), taken from a Minnesota population that is 

increasing in size (J. Toepfer, STCP, personal communication) yet reared at Fossil Rim 

Wildlife Center using the same husbandry techniques used for the APC, initially showed 

better hatchability and chick survival, but they subsequently experienced the same 

reproductive problems observed in APCs during their first breeding season (low numbers 

of eggs/hen, reduced hatchability, poor chick survival) (O. Dorris, Fossil Rim Wildlife 

Center, personal communication). This experiment, designed to investigate the influence 

of husbandry and genetics with respect to fitness-related problems associated with the 

current APC captive breeding program, suggests that husbandry issues are playing a more 

significant role than genetics in limiting survivorship at this time. 

However, it must be noted that current levels of genetic variability in the captive 

population are low and comparable to those observed in the Illinois GPC population 

when Westmeier et al. (1998) documented a significant decline in hatching success 

following a reduction in levels of genetic variability. Therefore, concern still exists that 

genetics may be an issue in the future, and the captive population should be closely 

monitored for any signs that may be associated with inbreeding depression until its 

effective population size has increased to levels associated with a genetically viable 

population (i.e., N e >500). 

Husbandry issues. As discussed previously, major biological constraints facing 

the captive breeding program include issues contributing to poor productivity including 

husbandry methods, nutrition, and management of diseases and parasites, particularly 

REV. Additionally, having 50% of the captive flock currently at one location (Fossil 

Wildlife Center) (Figure 6), has the potential for resulting in disastrous consequences to 

the APC captive breeding and APC recovery programs. The major REV event at Fossil 

Rim in recent years has heightened the gravity of this situation. 

Poor brood survival. Significant progress toward recovery will not occur until 

factors influencing the poor survival observed for chicks produced by released captivelyreared 

APCs (see Section I.G. Population Supplementation) are identified and resolved. 

Factors hypothesized as possible contributors to poor brood survival include: (1) habitat 

quality, especially as it pertains to insect availability for foraging chicks, (2) genetics, (3) 

physiological changes attributable to the captive environment, (4) maladaptive parental 

behavior as influenced by the captive environment, (5) disease/parasites, and (6) stings by 

the exotic red imported fire ant. 

43


II. RECOVERY 

The following sections present a strategy to recover the species, including 

objective and measurable recovery criteria to achieve downlisting and delisting, and sitespecific 

management actions to monitor and reduce or remove threats to the APC, as 

required under section 4 of the ESA. The Recovery Plan also addresses the five statutory 

listing/recovery factors (section 4(a)(1) of the ESA) to demonstrate how the recovery 

criteria and actions will lead to removal of the APC from the lists of Threatened and 

Endangered Species. 

A. RECOVERY STRATEGY 

As described in previous sections, the APC is facing numerous threats at multiple 

ecological scales. Presently, the APC is functionally extinct in the wild, maintained by 

supplementation from captive-reared birds. Recovery efforts defined by strategies 

detailed in this plan must be multifaceted and managed concurrently. For example, while 

more investment is warranted for captive propagation programs, restoration and 

acquisition of potential APC habitat also must be continued. Other avian species 

recovery efforts have adopted such an integrated approach. The winter of 2004–2005 

marked a record setting wintering population of whooping cranes (Grus americana). On 

the surface, the recovery of the whooping crane faces more complex demographic and 

political challenges when compared to the APC. These include low reproductive rates 

(K-selected), historically low population numbers, wide ranging migratory patterns, and 

the vulnerability of wintering wetland habitat to pollution, fragmentation, and other 

threats. In contrast, the APC is an r-selected species with a historically large population. 

However, at such currently low population levels brought about by habitat loss and 

fragmentation, and other factors identified in previous sections, the APC cannot 

withstand stochastic, catastrophic events. Since the last revision of the APC Recovery 

Plan in 1993, a core population in Refugio County has disappeared, and the two 

remaining wild populations are dependent upon releases of captive-raised birds. A better 

understanding of why other recovery efforts are succeeding is necessary as we refine 

existing and develop new recovery strategies. 

If recovery of the APC is to be successful three primary action areas must be 

supported. They are (1) habitat management, (2) captive and wild population 

management, and (3) public outreach. Specific objectives and numeric goals will be 

identified in the following section. These three action areas cumulatively address the five 

listing/current threat factors identified in section I.H (Reasons for Listing/Current 

Threats) of this plan. Specific strategies for addressing these threat factors are identified 

in section II.D (Reduction or Alleviation of Threats). 

Although only 50 APC currently exist in the wild, habitat management, 

enhancement, and restoration must be carried out to maintain existing high quality 

grasslands and restore degraded grasslands that may serve as APC habitat in the future. 

If existing and expanded grassland conservation efforts (e.g., CPCI) at the right scale 

[multiple core areas >25,000 ac (10,120 ha)] and context (minimal fragmentation, allow 

for gene flow between core areas) succeed, then viability for future APC populations is 

44


possible. There are very few areas left in Texas that can provide the long-term 

requirements for APC survival. The relict population in Galveston County at the TCPP 

exists upon a small patch [100) numbers of birds at multiple release sites will be required. In 2006, 

a record high of only 160 birds was released. It is clear the captive population 

management program must be retooled and elevated in dramatic fashion if we are to 

recover the APC. 

Numerous challenges face the wild APC population. Predation, red imported fire 

ants, disease, ectoparasites, accidents (e.g., flying into fences and wires), flooding, 

incompatible grazing, altered fire regimes, and countless other factors are collectively 

suppressing optimal recruitment of the two wild populations. At APCNWR, no 

recruitment has occurred from captive-reared released birds without very intensive 

intervention by APCNWR staff. Evidence suggests limited recruitment occurring at 

TCPP however, few brood survival data are available to positively document 

reproduction from captive-reared released birds. Such evidence leads to a core question: 

Do captive-reared birds lack essential physiological and behavioral traits that prevent 

their survival once released, or are there vital habitat requirements lacking that limit 

brood survival and population recruitment? Additional research efforts are essential to 

address these questions. However, conducting meaningful research with broad ranging 

applicability is very challenging given the low population numbers and varied grassland 

habitats at these two sites. Research addressing these questions should be conducted on 

non-endangered GPC populations to the extent practicable. 

An ongoing challenge to recovery of the APC has been difficulty in attracting a 

large, engaged constituency to support conservation of the APC. A focused effort is 

needed to engage both public and private partners in funding opportunities that can be 

strategically used for recovery actions. 

This plan outlines current strategies believed to be critical to APC recovery. Goals, 

objectives, and criteria were derived from information and literature discussed in 

previous sections of this plan or were based on the professional judgment of recovery 

team members. Specific observations and assumptions that weighed heavily in the 

formulation of goals, objectives, and criteria include: 

45


• From a theoretical perspective, a N e >500 is needed to maintain genetic variation 

within a population (Lande and Barrowclough 1987:94). 

• Assuming the worst case scenario of only 10% of males breeding (Robel 1970) 

and a sex ratio of 1:1, a population of 2,750 would be required to produce an N e 

of 500. 

• A corollary to the N e requirement is that small fragmented populations must be 

situated on the landscape so that gene flow among populations is maintained 

(Johnson et al. 2003, 2004; Toepfer 2003) and recolonization following local 

extinction is facilitated (Fahrig and Merriam 1994, Hanski et al. 1996). 

• Prairie-chickens require lots of grass and open space (e.g., Lehmann 1939, 1941; 

Schwartz 1945; Baker 1953; Hamerstrom et al. 1957; Cogar et al. 1977; Toepfer 

1988, 2003; Johnson et al. 2004, Silvy et al. 2004). 

• T. cupido management areas should be ≥33%, and preferably ≥50% grassland 

(Hamerstrom et al. 1957). 

• Brush must be controlled to prevent excessive encroachment into grasslands 

(Hamerstrom et al. 1957, Svedarsky 1979:102, Toepfer 1988:484, Toepfer 2003). 

Less than 25% of the landscape should be wooded, with woodlands aggregated in 

scattered blocks (Hamerstrom et al. 1957). 

• The minimum management area size required to maintain a viable prairie-chicken 

population is unknown (Toepfer 2003). In general, prairie-chicken management 

areas should consist of thousands of acres of grass distributed over a landscape of 

several thousand square miles in a fashion that maintains connectivity among 

populations (Toepfer 2003, Johnson et al. 2004). 

B. GOALS, OBJECTIVES, AND CRITERIA 

The goal of this plan and recovery effort is to protect and ensure survival of the APC 

and its habitat, allowing the overall population to reach a measurable level of ecological 

and genetic stability so that it can be reclassified to threatened status (downlisted) and 

ultimately removed from the endangered species list (delisted). This goal can be 

achieved only if threats previously identified are sufficiently reduced or removed. 

Objective and measurable criteria for downlisting and delisting are as follows: 

1. Downlist to threatened status when the overall population maintains a minimum 

of 3,000 breeding adults annually over a 5-year period. These birds should be 

distributed along a linear distance of no less than 50 miles (80 km) to mitigate for 

environmental stochasticity (e.g., hurricanes) while maintaining genetic flow. 

2. Delist when there is a minimum overall population of 6,000 breeding adults 

annually over a 10-year period occupying habitats along a linear distance of no 

less than 100 miles. 

Specific objectives and criteria for habitat management, captive and wild population 

management, and public outreach necessary to accomplish these recovery goals are: 

Objective 1: Maintain and improve 300,000 ac (121,457 ha) of coastal prairie grasslands 

for the APC throughout the bird’s historic range on both private and public lands. 

46


Habitat is one of the major factors currently limiting APC populations. The 

APC’s prairie grassland habitat has been reduced by an estimated 99% of historic levels 

(Smeins et al. 1991). Remaining habitat has become highly fragmented, making isolated 

APC populations more susceptible to inbreeding, localized weather extremes, land use 

changes, predation, and disease. 

APC recovery will require a network of large, high quality coastal prairie 

grasslands containing multiple core areas distributed along at least 100 linear miles (160 

km). A core area is defined as an area of habitat capable of supporting a population of 

500 (250 displaying males), or approximately 25,000 ac (10,121 ha) (assuming a carrying 

capacity of 1 bird/50 ac (20 ha) (Lehmann 1941:7). 

Objective 2: Enhance propagation and release efforts to boost wild populations to viable 

levels and reintroduce physically and behaviorally healthy birds to their former range 

(a) Maintain 90% of original gene diversity for 20 years with about 200 birds in 

the captive flock. 

(b) Produce enough chicks annually to release at multiple sites (approximately 

100 birds per release site). 

• Increase capacity of breeding pairs to a minimum of 100 pairs within 

two years, with no one facility containing more than 25% of the 

captive flock. 

• By 2008, increase survival in the captive environment so that 50% of 

eggs produced survive to eight weeks of age. 

(c) When number of young available for release exceeds 100, pilot releases of no 

fewer than 30 should be considered on private lands. 

Captive propagation and release efforts must be enhanced in order to boost wild 

populations to viable levels and reintroduce physically and behaviorally healthy birds to 

their former range. Maintaining the integrity of the captive APC flock is crucial for APC 

recovery. Without a healthy, genetically sound captive flock the APC is doomed to 

extinction. 

Objective 3: Establish populations of at least 500 birds in multiple core areas, providing 

for gene flow between populations (see Objective 1). 

Objective 4: Broaden public support and partner in efforts to conserve the APC and its 

coastal prairie ecosystem. 

A lack of understanding and awareness currently exists among the public and 

other groups concerning the perilous condition of APC populations and prairie grouse in 

general. As a result, public support for APC recovery efforts is lacking. There needs to 

be an increase in outreach activities to raise the public’s awareness of the plight of the 

APC and its endangered coastal prairie ecosystem. Explaining why the APC has 

47


declined, what challenges it faces for recovery, and why it is important to save this 

imperiled bird will increase support for recovery of the APC. 

48


C. NARRATIVE OUTLINE OF RECOVERY ACTIONS 

This section lists the site-specific actions necessary to accomplish goals and 

objectives outlined in Section II.B (Goals, Objectives, and Criteria). Priority 1 actions 

include activities that must be taken to prevent extinction or prevent irreversible 

population declines in the foreseeable future. Priority 2 actions include activities that 

must be taken to prevent a significant decline in APC populations or habitat quality, or to 

prevent some other significant negative impact short of extinction. Priority 3 actions 

include all other activities necessary to accomplish full recovery. 

1. Continue to maintain and improve at least 300,000 ac (121,457 ha) of coastal prairie 

habitat for the APC throughout the bird’s historic range through the CPCI and other 

habitat improvement projects on both private and public lands. 

1.1. Create a network of large, high quality coastal prairie habitats containing 

multiple core areas maintained compatibly with APC occupation and distributed 

along >100 linear miles (160 km). A core area is an area of suitable habitat 

capable of supporting a population of 500 (250 displaying males) [approximately 

25,000 ac (10,121 ha) assuming a carrying capacity of 1 bird/50 ac (20 ha) 

(Lehmann 1941:7)] (Priority 1). 

1.2. Coastal prairie habitats compatible with APC occupation should be 

interconnected through grassland corridors (1–3 miles (1.6-4.8 km) wide) within 

the APC’s historic range to allow for dispersal and genetic exchange and to 

hedge against environmental stochasticity (e.g., hurricanes) (Toepfer 2003). 

Areas such as national wildlife refuges (e.g., APC, Aransas, Brazoria), TNC 

preserves (e.g, Mad Island, TCPP), and private lands will be pivotal in making 

these grassland corridors a reality. To maximize benefits for APC recovery, 

management priority should be given to habitats in close proximity to existing 

populations or future release sites (Priority 1). 

1.2.1. Cultivate partnerships with managers of existing grasslands currently 

under public (e.g., Aransas, Texas Mid-Coast, and Texas Chenier Plains 

National Wildlife Refuges, TPWD Mad Island Wildlife Management Area 

or NGO (e.g., Mad Island Preserve) ownership to encourage management 

consistent with APC habitat requirements (see 1.3) (Priority 1). 

1.2.2. Cultivate partnerships with private landowners through mechanisms listed 

under 1.4 (Priority 1). 

1.3. Manage and monitor progress on public lands to maintain, improve, and/or 

restore native prairie grasslands as APC habitat by: 

1.3.1. grazing to maintain clumped grass structure (Priority 1). 

1.3.2. controlling brush and exotic plants (Priority 1). 

1.3.3. prescribed burning (Priority 1). 

1.3.4. maintaining and restoring natural hydrology to reduce nest flooding 

(Priority 1). 

1.3.5. planting food plots to provide supplemental winter foods and brood 

habitat (Priority 1). 

1.3.6. mowing as necessary to control vegetation density (Priority 2). 

49


1.3.7. restoring formerly farmed fields to native grass species (Priority 1). 

1.3.8. monitoring and managing sympatric wildlife species giving priority to 

APC (Priority 3). 

1.3.9. controlling exotic wildlife species (Priority 1). 

1.3.10. prohibiting introductions of exotic wildlife species on public lands (e.g., 

ring-necked pheasants, etc.) (Priority 2). 

1.3.11. managing waterfowl, especially geese, to minimize competition and 

potential for disease transmission (Priority 1). 

1.3.12. controlling public use to prevent APC disturbance (Priority 1). 

1.4. Secure additional habitat through: 

1.4.1. acquisition of additional refuge lands including at least 20,000 ac (8,097 

ha) to the existing APCNWR through a combination of fee simple and 

long-term easement acquisitions from willing sellers (Priority 1). 

1.4.2. private land easements (Priority 1). 

1.4.3. safe harbor agreements (Priority 1). 

1.4.4. habitat conservation plans (Priority 1). 

1.4.5. mitigation (Priority 1). 

1.4.6. partnerships with other governmental, non-governmental organizations 

(NGOs), and private land managers (Priority 1). 

1.4.7. pursuit of other mechanisms/programs for private lands work such as 

NRCS’s Grassland Reserve Program, EQIP, Farm Bill, TPWD’s 

Landowner Incentive Program (LIP), USFWS’s Partners for Fish and 

Wildlife program, etc. (Priority 1). 

1.4.8. technical assistance, economic incentives, and regulatory incentives on 

private lands through the CPCI program (Priority 1). 

1.4.9. coordination with NRCS and other governmental agencies to resolve 

conflicting programs that are detrimental to APC recovery (Priority 1). 

1.4.10. acquisition of grassland additions to existing national wildlife refuges 

within the APCs historic range (Aransas, Texas Mid-Coast, Texas Chenier 

Plains) (Priority 2). 

1.5. Survey status and trends of native grasslands every five years. Identify public 

and private release sites that maximize the probability for success (Priority 2). 

1.6. Cultivate market-driven financial incentives for private landowners that establish 

and maintain APC populations on their lands (e.g., eco-tourism, economic value 

of native grasses, etc.) (Priority 1). 

1.7. Focus recovery actions in priority management zones (Figures 3, 4, 5). Priority 

management zones are areas within the APC’s former range currently supporting 

APCs or that have been identified as having high potential for repatriation. Core 

areas will be located within priority management zones (Priority 1). 

1.8. Conduct research necessary to: 

1.8.1. Determine landscape scale habitat needs (Priority 2). 

1.8.2. Evaluate patch burning as a management tool (Priority 2). 

1.8.3. Evaluate current management practices on release sites (Priority 1). 

50


2. Continue propagation and release efforts to boost wild populations to viable levels 

and reintroduce physically and behaviorally healthy birds to their former range. 

2.1. Maximize effectiveness and production of current captive flock by: 

2.1.1. evaluating and implementing different rearing techniques (e.g., broody 

hens, APC hens, etc.) to generate physically and behaviorally healthy 

birds (Priority 2). 

2.1.2. implementing different strategies for maximizing production such as: 

2.1.2.1. photoperiod manipulation (Priority 3). 

2.1.2.2. using the expertise of a particular facility to their fullest extent 

(e.g., a facility that has problems getting birds to produce eggs 

receives eggs from a facility that does very well at producing 

eggs, etc.) (Priority 2). 

2.1.3. identifying and rectifying disturbance issues at breeding facilities 

(Priority 3). 

2.1.4. aggressively managing diseases (e.g., REV, pox, etc.) and other health 

issues (Priority 1). 

2.1.5. sharing resources and information among all captive flock facilities and 

recovery partners (Priority 3). 

2.1.6. evaluating diets for chicks and adults to help address health and 

production issues that may be related to nutritional issues (Priority 1). 

2.1.7. completing the husbandry manual within one year to standardize 

husbandry techniques and strategies, while continually evaluating the 

effectiveness of these techniques (Priority 2). 

2.1.8. evaluating screening pens which house the breeding flock to minimize risk 

of REV transmission by insects (Priority 2). 

2.2. Determine and continually monitor genetic health of the captive flock. Evaluate 

APC genetic variability to determine if hybridization with GPCs is warranted. 

Also, explore and consider gene banking (Priority 1). 

2.3. Increase production of birds through increased efficiency at current facilities and 

the addition (dedicated exclusively to APCs) or expansion of breeding facilities 

to allow for a capacity of a minimum of 100 pairs, with no one facility containing 

more than 25% of the captive flock population (Priority 1). 

2.4. Conduct research to provide information needed for captive APC management 

such as: 

2.4.1. REV (Priority 1). 

2.4.2. Factors affecting egg production and viability and chick survival 

(Priority 1). 

2.4.3. Rearing environment/methods (Priority 1). 

2.4.4. Factors affecting skeletal/muscular problems (e.g., leg rotations, 

wrynecks, etc.) (Priority 1). 

2.4.5. Management of parasites and diseases (Priority 1). 

3. Establish populations of >500 birds in multiple core areas within 10 years, providing 

for gene flow among core populations. 

51


3.1. Continually evaluate release techniques and implement changes as needed to 

improve survival (Priority 1). 

3.2. Survey APC numbers annually by conducting spring booming ground and brood 

survival counts when appropriate (Priority 1). 

3.3. Expand the release of captive-bred APCs where suitable habitat exists 

(Priority 1). 

3.4. Evaluate translocation of birds as a possible technique for repatriation and 

genetic management of existing populations (Priority 2). 

3.5. Develop regulatory procedures (e.g., safe harbor, habitat conservation plans 

(HCPs)) for establishing additional populations (Priority 1). 

3.6. Protect APC from take by enforcing current Federal and State legislation and 

regulations (Priority 1). 

3.7. Conduct research to provide information needed to determine: 

3.7.1. Factors affecting wild brood survival (Priority 1). 

3.7.2. Factors affecting post-release survival (Priority 1). 

4. Engage the public in efforts to conserve the APC and its coastal prairie ecosystem. 

4.1. Develop and implement a comprehensive public outreach plan that will 

incorporate specific outreach tasks to target groups. 

4.1.1. Contract with consulting group to develop specific marketing strategies to 

effectively and efficiently engage the public in APC recovery (Priority 3). 

Although numerous outreach activities have been conducted regarding the 

APC and recovery activities (e.g., annual APC festival, numerous 

newspaper and magazine articles, television and radio newsclips and short 

documentaries, presentations and development of curriculum materials for 

school groups), outreach remains non-focused. Therefore, these actions 

have likely not been as effective as they could have been in engaging 

public support for APC recovery. With limited funding and personnel to 

accomplish outreach actions, it is imperative that the right groups are 

targeted with the proper tools to maximize effectiveness of these actions. 

4.1.2. Assist in the establishment of an APC partner coalition hosted by an NGO 

and dedicated to supporting APC recovery objectives. A host organization 

with national focus and credibility is needed to liaise with corporate 

partners and manage potential contributions and grants (Priority 2). 

52


D. REDUCTION OR ALLEVIATION OF THREATS 

Factor 1: The present or threatened destruction, modification, or curtailment of 

habitat or range. 

Destruction, modification, or curtailment of APC habitat or range is addressed in 

Actions 1.1−1.8 which provide for creation of large interconnected grasslands that 

allow for dispersal and gene flow. This grassland habitat will result from 

management of existing public lands for optimal APC habitat (1.3); addition of 

habitat to the APCNWR (1.4.1); and partnerships with other agencies and NGOs 

through such programs as the CPCI, GLCI, EQIP, and LIP which provide landowner 

incentives for habitat restoration (1.4.2−1.4.8). Actions listed under 4.1 address the 

need for developing more support for APC recovery, including habitat restoration. 

Factor 2: Overutilization for commercial, recreational, scientific, or educational 

purposes. 

The hunting season for APCs has been closed since 1937. Actions 1.3.12 and 3.6 

address disturbance and take issues, respectively, that may still remain. Action 4.1 

provides for public outreach to raise awareness of the APC’s plight and its presence 

in the coastal prairie ecosystem. 

Factor 3: Disease or predation. 

Disease. Action 1.3.11 provides for management of waterfowl on public lands 

where APC are present to minimize potential for disease transmission. Action 1.8.3 

provides for evaluation of management practices at release sites. This action could 

include routine disease screening to assess potential interactions between habitat 

management and disease issues. Captive flock health issues are addressed in 2.1.1, 

2.1.4, 2.1.6, 2.1.7, and 2.4. 

Predation. Assessment and management of predation threats are addressed in 

actions 1.3.8, 1.3.9, 3.1, and 3.7. Action 1.8.3 provides for evaluation of management 

practices at release sites. This action could include evaluation of predator 

management. 

Factor 4: Inadequacy of existing regulations. 

In general, the current regulatory framework provides for adequate protection and 

conservation. Action 1.4.9 provides for coordination among government agencies to 

minimize the impact of programs that may be detrimental to APC recovery. 

Factor 5: Other natural or manmade factors affecting its continued existence. 

Population fragmentation. Reduction in the risk of extinction associated with the 

small, fragmented nature of APC populations is a major focus of this recovery plan. 

Practically the entire plan focuses on increasing the amount of usable APC habitat, 

providing for connectivity among habitat blocks to minimize isolation of populations, 

53


and the restoration of APC populations to viable levels through management of wild 

and captive populations. Actions 1.1, 1.2, 1.3, and 1.4 involve the broad-scale 

maintenance, creation, and acquisition of interconnected, suitable grassland on private 

and public lands to help promote dispersal and genetic exchange. Actions under the 

heading of 1.3 (1.3.1 – 1.3.12) cover specific management activities to maintain, 

improve, or restore contiguous, native habitat, while minimizing competition and 

human disturbance. Actions 1.8.1 and 1.8.2 call for research to determine habitat 

needs across the landscape and to evaluate patch burning as a management tool to 

form suitable habitat. 

Genetics. Maintenance of gene flow among population segments is addressed in 

action 1.2. Action 2.2 specifies determining and monitoring the genetic health of the 

captive population. This action also specifically directs the evaluation of APC 

genetic variability to determine if hybridization with GPCs is warranted as a recovery 

measure. Action 2.2 also directs consideration of APC gene banking. Action 3 

provides for establishment of at least two self-sustaining core populations with gene 

flow between the populations. Action 3.4 lists translocation as a tool for genetic 

management of populations. Action 3.7 provides for research to determine factors 

that may influence wild brood and post-release survival. This research should include 

an assessment of genetic factors that may affect survival of broods and released birds. 

Husbandry issues. Husbandry issues thought to be limiting APC recovery are 

addressed in all actions under the Action 2 heading (2.1 – 2.4.5). Specifically, these 

actions direct refinement of husbandry practices to increase production of physically 

and behaviorally healthy birds. Action 2.4 provides for research on issues currently 

limiting production including REV, egg production, chick survival, husbandry 

practices, skeletal/muscular problems, and parasite/disease management. 

Poor brood survival. Action 3.7.1 specifically directs that research be conducted 

to determine factors affecting wild brood survival. 

54


III. IMPLEMENTATION SCHEDULE 

The following implementation schedule outlines and prioritizes recovery tasks over 

the next five years. It will be used in monitoring recovery actions and will provide the 

basis for funding for these actions. Actions are identified under general categories, and 

all headings are derived from Section II.C (Narrative Outline of Recovery Actions). This 

implementation schedule ranks objectives and actions, identifies respective responsible 

agencies/groups, defines implementation time-frames, and estimates costs. Actions must 

be continually revised as plans move from implementation to completion. Each revision 

will identify additional actions and studies that will be needed during the recovery period. 

Recovery priorities are defined as follows: 

Priority 1: An action that must be taken to prevent extinction or prevent the APC from 

declining irreversibly in the foreseeable future. 

Priority 2: An action that must be taken to prevent a significant decline in APC 

populations or habitat quality or to prevent some other significant negative impact short 

of extinction. 

Priority 3: All other actions necessary to provide for full recovery (or reclassification). 

55


Priority 

# 

Action 

# Action Description 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 

(* = lead) Year 1 Year 2 Year 3 Year 4 Year 5 Total Comments 

1 

1 

1 

1 

1.1 Create a network of 

coastal prairie habitats 

containing multiple core 

areas 

1.2 Create interconnected 

grassland corridors of 

coastal prairie habitats 

between core areas to 

allow for dispersal and 

genetic exchange and 

hedge against 

environmental 

stochasticity 

1.2.1 Cultivate partnerships 

on existing grasslands 

under public or NGO 

ownership 

1.2.2 Cultivate partnerships 

with private 

landowners 

Ongoing 

Ongoing 

Ongoing 

Ongoing 

FWS-ES*, 

FWS- 

APCNWR*, 

TNC, USDA- 

NRCS, 

TPWD, 

Private, 

SHRC&D, 

GLCI 

FWS-NWRS, 

FWS-ES*, 

TNC, TPWD, 

USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 

FWS- 

NWRS * , 

TNC, TPWD 

FWS-NWRS, 

FWS-ES*, 

TPWD, 

USDA- 

NRCS, TNC, 

SHRC&D, 

GLCI 

750 750 750 750 750 3,750 

--- --- --- --- --- --- See also 

Action # 1.1 

and 1.4.1 – 

1.4.9 

--- --- --- --- --- --- See also 

Action # 1.1 

and 1.4.1 – 

1.4.9 

--- --- --- --- --- --- See also 

Action # 

1.4.2 − 

1.4.9 

56


Priority 

# 

Action 


Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


1 

1.3.1 Manage and initiate 

grazing on public lands to 

maintain clumped grass 

structure 

Ongoing 

FWS- 

NWRS*, 

TPWD, 

USDA-NRCS 

300 50 50 50 50 500 

1 

1 

1 

1 

1 

1.3.2 Control brush and exotic 

plants on public lands 

1.3.3 Conduct prescribed 

burning on public lands 

1.3.4 Maintain/restore natural 

hydrology to reduce nest 

flooding on public lands 

1.3.5 Maintain food plots to 

provide supplemental 

winter foods and brood 

habitat on public lands 

1.3.7 Restore formerly farmed 

fields to native grasses on 

public lands 

Ongoing FWS- 

NWRS*, 

TPWD, 

USDA-NRCS 

150 150 150 150 150 750 

Ongoing FWS- 75 75 75 75 75 375 

NWRS*, 

TPWD, 

USDA-NRCS 

Ongoing FWS- 25 25 25 25 25 125 

NWRS*, 

TPWD, 

USDA-NRCS 

Ongoing FWS-NWRS* 5 5 5 5 5 25 

Ongoing 

FWS- 

NWRS*, 

USDA-NRCS 

150 150 150 150 150 750 

57


Priority 

# 

1 

1 

1 

1 

1 

1 

Action 


1.3.9 Control exotic wildlife 

species on public lands 

1.3.11 Manage waterfowl to 

minimize competition 

and disease 

1.3.12 Control public use on 

public lands to prevent 

disturbance to APCs 

1.4.1 Acquire at least 20,000 

acres adjacent to APC 

NWR through a 

combination of fee 

simple and long-term 

easements from willing 

sellers 

1.4.2 Secure additional habitat 

through private land 

easements 


through safe harbor 

agreements 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


Ongoing FWS- 

NWRS*, 

TPWD, 

USDA- 

NRCS, 

USDA-WS 

Ongoing FWS- 

NWRS*, 

TPWD 

Ongoing FWS- 

NWRS*, 

TPWD 

15 years FWS- 

APCNWR* 

15 years FWS-NWRS, 

FWS-ES*, 

TNC, TPWD, 

Private, 


FWS-ES*, 

Private, 

SHRC&D, 

GLCI 

50 50 50 50 50 250 

5 5 5 5 5 25 

5 5 5 5 5 25 

2,000 2,000 2,000 2,000 2,000 10,000 See also 

Action # 1.1 

--- --- --- --- --- --- See also 

Action # 1.1 

--- --- --- --- --- --- See also 

Action #1.1 

58


Priority 

# 

1 

1 

1 

1 

Action 



through Habitat 

Conservation Plans 


through mitigation 


through partnerships with 

other governmental, 

NGO, and private land 

managers 


by pursuing other 

programs for private 

lands 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 



FWS-ES*, 

TPWD, 

USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 

Ongoing FWS-NWRS, 

FWS-ES*, 

TPWD 


FWS-ES*, 

TPWD, TNC, 

USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 


FWS-ES*, 

TPWD, 

USDA- 

NRCS, TNC, 

SHRC&D, 

GLCI 

--- --- --- --- --- --- See also 

Action # 1.1 

20 20 20 20 20 100 See also 

Action # 1.1 

10 10 10 10 10 50 See also 

Action # 1.1 

10 10 10 10 10 50 See also 

Action # 1.1 

59


Priority 

# 

Action 


Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


1 

1 

1 


by providing technical 

assistance, economic 

incentives, and regulatory 

incentives on private 

lands through the CPCI 

program 

1.4.9 Coordinate with NRCS 

and other governmental 

agencies to resolve 

conflicting programs that 

are detrimental to APC 

recovery 

1.6 Cultivate market-driven 

financial incentives for 

private landowners that 

establish and maintain 

APC populations on their 

lands 

Ongoing 

Ongoing 

Ongoing 

FWS-NWRS, 

FWS-ES*, 

TNC, TPWD, 

USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 

FWS-NWRS, 

FWS-ES*, 

USDA- 

NRCS, 

TPWD, 

SHRC&D, 

GLCI 

FWS- 

APCNWR, 

FWS-ES, 

TNC, 

TPWD*, 

USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 

--- --- --- --- --- --- See also 

Action # 1.1 

10 10 10 10 10 50 See also 

Action # 1.1 

10 10 10 10 10 50 

60


Priority 

# 

1 

1 

1 

1 

1 

Action 


1.7 Focus recovery actions in 

priority management 

zones 

1.8.3 Conduct research to 

evaluate current 

management practices at 

release sites 

2.1.4 Aggressively manage 

diseases and other health 

issues 

2.1.6 Evaluate diet for chicks 

and adults to help address 

problems that may be 

arising from current diets 

2.2 Determine and 

continually monitor 

genetic health of the 

captive flock 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


Ongoing FWS- 

NWRS*, 

FWS-ES, 

TNC, TPWD, 

USDA- 

NRCS, 

SHRC&D, 

GLCI 

5 Years FWS- 

NWRS*, 

FWS-ES 

TNC, private 

Ongoing FWS- 

APCNWR*, 

BFs 

5 years FWS- 

APCNWR, 

BFs, 

FWZ*,SARC, 

1 year, 

ongoing 

STCP 

FWS- 

APCNWR*, 

STCP*, BFs 

--- --- --- --- --- --- 

75 75 75 75 75 375 

425 125 125 125 125 925 Critical 

Need 

25 25 25 25 25 125 Critical 

Need 

65 15 15 15 15 125 Critical 

Need 

61


Priority 

# 

1 

1 

1 

1 

1 

Action 


2.3 Increase production of 

birds through increased 

efficiency at current 

facilities and the 

addition/expansion of 

breeding facilities to 

allow for a capacity of 

100 pairs, with no facility 

containing more than 

25% of the captive flock 

population 


determine factors 

affecting captive 

breeding, such as REV 



affecting egg viability 

and chick survival 


determine best rearing 

environments/methods 



affecting 

skeletal/muscular 

problems 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


5 years FWS- 

APCNWR*, 

BFs 

5 years FWS- 

APCNWR, 

TAMU*, 

UofG 

3 years FWS- 

APCNWR*, 

BFs 

3 years FWS- 

APCNWR*, 

BFs 

3 years FWS- 

APCNWR*, 

BFs 

900 900 500 500 500 3,300 Critical 

Need 

50 50 50 50 50 250 Critical 

Need 

40 40 40 --- --- 120 

40 40 40 --- --- 120 

40 40 40 --- --- 120 

62


Priority 

# 

1 

1 

1 

1 

1 

1 

Action 



determine how to best 

manage for parasites in 

the captive setting 

3.1 Evaluate release 

techniques and 

implement changes as 

needed 

3.2 Survey APC numbers 

annually 

3.3 Expand the release of 

captive-bred APCs where 

suitable habitat exists 

3.5 Utilize procedures such 

as safe harbor, HCPs, etc. 

for establishing 

additional populations 

3.6 Protect APC from take by 

enforcing current Federal 

and State legislation and 

regulations 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


3 years FWS- 

APCNWR*, 

BFs 

10 years APCNWR * , 

TNC 

Ongoing 

Annually 

as captive 

production 

permits 

FWS- 

APCNWR*, 

TNC, TPWD 

FWS- 

APCNWR*, 

FWS-NWRS, 

TNC, TPWD, 

-USDA- 

NRCS, 

Private, 

SHRC&D, 

GLCI 

5 years FWS- 

APCNWR, 

FWS-ES* 

Ongoing 

FWS-NWRS, 

FWS-ES, 

FWS-LE*, 

TPWD 

40 40 40 --- --- 120 

100 100 100 100 100 500 

5 5 8 10 10 38 

150 75 75 75 75 450 See Action 

# 1.1 

25 25 25 25 25 125 

10 10 10 10 10 50 

63


Priority 

# 

1 

1 

2 

2 

2 

2 

2 

Action 




affecting wild brood 

survival 

3.7.2 Conduct post-release 

survival research 

1.3.6 Mow to control 

vegetation density on 

public lands 

1.3.10 Prohibit the introductions 

of exotic wildlife species 

on public lands 

1.4.10 Acquisition of 

grassland additions to 

existing national 

wildlife refuges 

1.5 Survey status and trends 

of native grasslands. 


determine landscape 

scale habitat needs 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


5 years FWS- 

APCNWR*, 

TNC 

60 60 60 60 60 300 Critical 

Need 

10 years FWS- 

APCNWR 

75 75 75 75 75 375 

As needed FWS- 15 15 15 15 15 75 

NWRS*, 

TPWD, 

USDA- 

NRCS 

As needed FWS- 2 2 2 2 2 10 

NWRS*, 

TPWD 

On-going FWS-NWRS * --- --- --- --- --- --- Costs 

included in 

Action # 1.1 

Once 

every 5 

years 

FWS- 

APCNWR, 

FWS-ES*, 

TNC, 

TPWD, 

USDA-NRCS 

5 years FWS- 

APCNWR 

-- 75 --- --- --- 75 

75 75 75 75 75 375 

64


Priority 

# 

2 

2 

2 

2 

2 

2 

Action 



evaluate patch burning as 

a management tool 

2.1.1 Evaluate and implement 

the use of different 

rearing techniques to 

generate physically and 

behaviorally healthy 

birds 

2.1.2.2 Use the expertise of a 

particular breeding 

facility to their fullest 

extent 

2.1.7 Complete husbandry 

manual to standardize 

husbandry techniques and 

strategies 

2.1.8 Evaluate screening pens 

which house the breeding 

flock to minimize risk of 

REV transmission by 

insects 

3.4 Evaluate translocation of 

birds as a possible 

technique for repatriation 

and genetic management 

of existing populations 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


5 years FWS- 

APCNWR*, 

USDA-NRCS 

5 years FWS- 

APCNWR*, 

BFs 

Ongoing 

1 year, 

ongoing 

FWS- 

APCNWR*, 

BFs 

FWS- 

APCNWR*, 

BFs 

5 years FWS- 

APCNWR * , 

BFs 

In 3 years 

as populations 

permit 

FWS- 

APCNWR*, 

TNC, STCP 

50 50 50 50 50 250 

25 25 25 25 25 125 

--- --- --- --- --- --- 

10 2 2 2 2 18 

25 25 25 25 25 125 

--- --- 20 20 20 60 

65


Priority 

# 

2 

3 

3 

3 

Action 


4.1.2 Assist in the 

establishment of an 

APC partner coalition 

hosted by an NGO and 

dedicated to supporting 

APC recovery 

objectives. 

1.3.8 Monitor and manage 

sympatric wildlife 

species on public lands 

2.1.2.1 Maximize effectiveness 

and production of current 

captive flock though the 

use of photo-period 

manipulation 

2.1.3 Identify and rectify 

disturbance issues at 

breeding facilities 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


5 FWS * , 

NFWF 

Ongoing 

Ongoing 

FWS- 

NWRS*, 

TPWD, 

USDA-NRCS 

FWS- 

APCNWR* , 

BFs 

5 years FWS- 

APCNWR*, 

BFs 

125 --- --- --- --- --- 

20 20 20 20 20 100 

10 10 10 10 10 50 

10 10 10 10 10 50 

66


Priority 

# 

3 

3 

Action 


2.1.5 Share resources and 

information among all 

captive breeding facilities 

and other partners 

involved with recovery 

4.1.1 Contract with 

consulting group to 

develop specific 

marketing strategies to 

effectively and 

efficiently engage the 

public in APC 

recovery. 

Action 

Duration 

(years) 

Responsible 

Parties 

Cost Estimate 

($1,000) 


Ongoing FWS- 

APCNWR*, 

FWS-NWRS, 

FWS-ES, 

FWS-LE, 

TAMU, 

STCP, 

TPWD, TNC, 

BFs, USDA- 

NRCS, FWZ, 

USDA-WS, 

UofG, 

Private, 

SHRC&D, 

GLCI, SARC 

5 Years FWS- 

APCNWR*, 

FWS-NWRS, 

FWS-ES, 

TPWD, TNC, 

USDA- 

NRCS, BFs, 

SHRC&D, 

GLCI 

10 10 10 10 10 50 

100 50 50 50 50 300 

67


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APPENDIX 1. Attwater’s prairie-chicken 1937−2006 population estimates by Texas county. Data are from Jurries (1979) 

(1937−1977) and Attwater Prairie Chicken National Wildlife Refuge and Texas Parks and Wildlife Department (unpublished data) 

(1978−present). 

Year 

County 1937 1950 1963 1967 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 

Aransas 1212 902 35 30 10 10 15 14 50 96 35 50 76 

Austin 339 200 200 200 284 332 364 330 292 576 498 186 308 326 

Brazoria 948 53 20 20 20 40 20 20 20 20 20 20 

Calhoun 25 40 15 10 

Chambers 220 15 10 2 6 8 8 

Colorado 926 350 200 175 186 378 166 144 150 422 324 178 206 186 

DeWitt 272 80 12 12 12 12 6 8 8 8 

Fort Bend 10 30 35 22 26 92 136 114 148 96 80 78 54 

Galveston 332 35 90 130 96 50 148 238 166 100 140 124 132 96 

Goliad 4 

23 75 216 402 290 260 486 188 a 164 80 56 34 

Harris 261 123 140 120 78 56 92 92 112 58 16 24 2 

Jackson 35 10 

Jefferson 220 85 10 

Lavaca 93 b 

Liberty 10 

Matagorda 15 5 

Refugio 3030 2100 412 175 310 440 166 192 356 336 530 550 742 726 

Victoria 620 200 100 90 112 234 166 224 242 342 218 110 126 64 

Waller 64 32 15 20 26 26 26 10 10 10 30 

Wharton 75 50 30 40 76 214 108 72 42 24 16 20 

Total 

8711 b 4200 1335 1070 1469 2220 1660 1782 2019 2254 2090 1506 1500 c 1718 1584 

81


Year 

County 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 

Aransas 20 19 22 24 16 22 10 12 6 2 2 

Austin 234 246 292 198 114 116 150 106 46 36 48 54 26 10 

Brazoria 20 20 

Calhoun 

Chamber s 

Colorado 184 280 320 218 246 228 242 162 90 90 70 50 34 20 

DeWitt 

Fort Bend 44 48 54 38 32 38 16 8 4 2 

Galveston 110 104 66 74 36 48 38 30 16 26 30 26 24 18 

Goliad 100 62 84 114 78 34 16 12 4 12 8 0 2 

Harris 4 

Jackson 

Jefferson 

Liberty 

Matagord a 

Refugio 658 438 646 838 810 340 582 562 246 292 310 330 370 110 

Victoria 64 64 112 116 94 48 54 34 20 10 8 2 

Waller 

Wharton 

Total 

1438 1281 1596 1620 1426 874 1108 926 432 470 476 462 456 158 

82


Year 

County 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 

Aransas 

Austin 2 

Brazoria 

Calhoun 

Chamber s 

Colorado 10 14 d 24 d 8 d 18 d 20 d 20 d 22 d 42 d 40 d 20 d 32 d 

DeWitt 

Fort Ben d 

Galveston 16 10 22 d 36 d 28 d 30 d 24 d 18 d 16 d 22 d 20 d 18 d 

Goliad 

Harris 

Jackson 

Jefferson 

Liberty 

Matagord a 

Refugio 40 18 12 12 

Victoria 

Waller 

Wharton 

Total 68 42 d 58 d 56 d 46 d 50 d 44 d 40 d 58 d 62 d 40 d 50 d 

a Incomplete survey 

b Jurries (1979) did not include Lavaca County which accounts for a 93 bird discrepancy between his 1937 state total and that reported 

by Lehmann (1941). 

c Estimate only - survey incomplete 

d Totals include birds released from captive breeding program 

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APPENDIX 2. LIST OF ABBREVIATIONS AND ACRONYMS 

ac 

APC 

APCNWR 

BFs 

CFR 

cm 

DNA 

DVM 

EQIP 

ESA 

FR 

ft 

FWS-APCNWR 

FWS-ES 

FWS-LE 

FWS-NWRS 

FWZ 

GLCI 

GPC 

ha 

km 

lbs 

LIP 

mi 

N e 

NFWF 

NGO 

NRCS 

oz 

PCR 

Private 

SSP 

TAC 

TAMU 

TCPP 

TNC 

TPWD 

SHRC&D 

acre 

Attwater’s prairie-chicken 

Attwater Prairie Chicken National Wildlife Refuge 

APC Captive Breeding Facilities (includes Fossil Rim Wildllife 

Center, Houston Zoo, Inc., San Antonio Zoo, Abilene Zoo, 

Caldwell Zoo, Sea World of Texas) 

Code of Federal Regulations 

centimeter 

deoxyribonucleic acid 

Doctor of Veterinary Medicine 

Environmental Quality Incentive Program 

Endangered Species Act 

Federal Register 

foot 

U. S. Fish and Wildlife Service – Attwater Prairie Chicken 

National Wildlife Refuge (NWR) 

U. S. Fish and Wildlife Service – Ecological Services 

U. S. Fish and Wildlife Service – Law Enforcement 

U. S. Fish and Wildlife Service – National Wildlife Refuge System 

(Includes Attwater Prairie Chicken NWR, Aransas Refuge 

Complex, and Texas Mid-Coast Refuge Complex) 

Fort Worth Zoo 

Grazing Lands Conservation Initiative 

Greater prairie-chicken 

hectare 

kilometer 

pounds 

Landowner incentive program 

mile 

effective population size 

National Fish and Wildlife Foundation 

Non-governmental organization 

U. S. Natural Resources Conservation Service 

ounces 

polymerase chain reaction 

Private landowners 

Species Survival Plan 

Texas Administrative Code 

Texas A&M University 

Texas City Prairie Preserve 

Nature Conservancy of Texas 

Texas Parks and Wildlife Department 

Sam Houston Resource Conservation and Development 

84


STCP 

UofG 

USDA- NRCS 

USDA-WS 

USFWS 

Society of Tympanuchus Cupido Pinnatus 

University of Georgia 

U. S. Department of Agriculture – Natural Resources Conservation 

Service 

U. S. Department of Agriculture – Wildlife Services 

U. S. Fish and Wildlife Service 

85


APPENDIX 3. GLOSSARY OF TERMS 

aflatoxin Toxin produced by certain fungi. Contaminated food products include cereal 

grains (e.g., corn, sorghum, millet, rice, wheat), oilseeds (e.g., soybeans, sunflowers, 

cotton), spices, nuts, and milk (Reddy and Waliyar (2000). 

allele 

Form of a gene at a particular locus (http://www.dna.gov/glossary). 

allelic dropout Failure to detect an allele within a sample or failure to amplify an allele 

during PCR (http://www.dna.gov/glossary). Basically, allelic dropout is a failure to 

detect alleles within a sample that are actually present. This may be caused by 

degraded DNA within the sample, or by problems with testing procedures. 

booming ground Areas where male Attwater’s and greater prairie-chickens gather to 

display in an attempt to attract females for breeding. 

capillariasis 

Parasitic disease caused by roundworms of the genus Capillaria. 

chromosome The biological structure by which DNA is transmitted from one 

generation to the next (http://www.dna.gov/glossary). 

DNA Genetic material present in the nucleus of cells which is inherited from each 

biological parent (http://www.dna.gov/glossary). 

dispharynxiasis 

nasuta). 

Parasitic disease caused by the spiral stomach worm (Dispharynx 

effective population size The number of adults in a population contributing offspring 

to the next generation. Generally, effective population size is smaller than the census 

size, and sometimes much smaller 

(http://wiki.cotch.net/index.php/Effective_population_size). 

etiology The assignment of a cause, origin, or reason for something. The cause of a 

disease or disorder as determined by medical diagnosis. 

gene The basic unit of heredity; a functional sequence of DNA in a single chromosome 

(http://www.dna.gov/glossary). 

genetic similarity indices 

Quantitative measures of genetic variability. 

genotype The genetic constitution of an organism as distinguished from its physical 

appearance (http://www.dna.gov/glossary). 

86


haplotype A way of denoting the collective genotype of a number of closely linked 

loci on a single chromosome (http://www.dna.gov/glossary). 

heterozygosity Heterozygosity refers to the state of being a heterozygote (i.e., 

possesses two different alleles at one locus) . Heterozygosity can also refer to the 

fraction of loci within an individual that are heterozygous. In population genetics, it is 

commonly extended to refer to the population as a whole, i.e. the fraction of 

individuals in a population that are heterozygous for a particular locus 

(http://en.wikipedia.org/wiki/Zygosity). 

inbreeding depression Inbreeding depression is reduced fitness in a given 

population as a result of breeding of related individuals 

(http://en.wikipedia.org/wiki/Inbreeding_depression). 

lineage sorting Term describing the development of monophyly by genetic drift 

during the speciation process. 

locus (plural loci) The specific physical location of gene on a chromosome 

(http://www.dna.gov/glossary). 

mesocarnivores Medium-sized carnivores (e.g., coyote, fox, raccoon, skunk) as 

contrasted to larger carnivores (e.g., wolf, bear, lion). 

mitochondrial DNA (mtDNA) The DNA found in the many mitochondria found in 

each cell of the body. The sequencing of mtDNA can link individuals descended 

from a common female ancestor (http://www.dna.gov/glossary). 

monophyly A term which refers to a taxon comprised of members derived from a 

common ancestor which includes all descendants 

(http://wiki.cotch.net/index.php/Monophyly). 

outbreeding depression Refers to cases when progeny from crosses between 

individuals from different populations have lower fitness than progeny from crosses 

between individuals from the same population 

(http://en.wikipedia.org/wiki/Outbreeding_depression). 

phylogenetics The study of evolutionary relatedness among various groups of 

organisms (http://en.wikipedia.org/wiki/Phylogenetics). 

polymerase chain reaction (PCR) Polymerase chain reaction (PCR) is a 

biochemistry and molecular biology technique for enzymatically replicating DNA 

without using a living organism 

(http://en.wikipedia.org/wiki/Polymerase_chain_reaction). 

prairie 

An extensive area of flat or rolling grassland. 

87


r-selected In ecological notation, “r” refers to the rate of population growth per 

individual for a given time period, whereas “K” refers to the upper limit (carrying 

capacity) of an environment to support a population of individuals. Therefore, r- 

selected species have evolved life history strategies for rapid growth (quanity) as 

contrasted with K-selected species, which have evolved mechanisms that allow them 

to compete for resources when populations are near K (quality). R-selected species 

typically have high reproductive rates with relatively short life cycles (i.e., high 

mortality rates) (Odum 1971). 

reciprocal monophyly 

Taxons of comparison are each monophyletic. 

stochastic 

Random or statistical in nature. 

taxon A group of organisms constituting one of the categories or formal units in 

taxonomic classification (e.g., phylum, order, family, genus, or species) and 

characterized by common characteristics in varying degrees of distinction. 

viremic 

The presence of viruses in the bloodstream. 

wryneck An unnatural condition in which the head leans to one side because the neck 

muscles on that side are contracted (http://www.thefreedictionary.com/wryneck). 

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