Impacts of Predation on Northern Bobwhite and Scaled Quail Dale Rollins Texas A&M University, D-Rollins@Tamu.Edu

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Papers in Natural Resources Natural Resources, School of

2001 Impacts of predation on northern bobwhite and scaled quail Dale Rollins Texas A&M University, [email protected]

John P. Carroll University of Georgia, [email protected]

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Rollins, Dale and Carroll, John P., "Impacts of predation on northern bobwhite and scaled quail" (2001). Papers in Natural Resources. 651. http://digitalcommons.unl.edu/natrespapers/651

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mpacts of predation on

,! orthern bobwhite and caled quail

Nrthen bobwhite (Colinus virginianus) and scaled quail (Callipepla squamata) populations have declined throughout most of their distribution, and these declines have become more dramatic in recent years. In this review, we examine the role of predation in quail management. Predation is the major source of nest loss and of mortality for young and adult quail. Mean nest success across studies reviewed was 28%. Estimates of annual survival rates have varied from 5 to 26% for radiotelemetry studies and from 15 to 30% based on age-ratio studies. Breeding season survival estimates ranged from 13 to 51%in telemetry studies reviewed. Brood survival is the least studied aspect of quail survival;estimates ranged from 13 to 47%. Mammalian predators most often implicated in nest predation include striped skunks (Mephitis mephitis), raccoons (Procyon lotor), opossums (Didelphis virginianus), foxes (Urocyon cinereoargenteus and Vulpes vulpes), coyotes (Canis latrans), and feral hogs (Sus scrofa). Accipiters (Accipiter spp.) and northern harriers (Circus cyaneus) are the most common avian predators of quail. Less information is available to assess impact of predation on scaled quail, but observations from areas where bobwhites and scaled quail are sympatric suggested that scaled quail are less vulnerable to predation than bobwhites. Although quail have adapted to cope with high predation rates (e.g., renesting, large clutches), populations in some areas may be suppressed by predation. Changes in land use, management practices, and predator communities interact to depress quail populations over much of the bobwhite's range. Additional studies are needed to assess the role of predation and predation management in light of these landscape-level changes. A variation of the Integrated Pest Management philosophy used in crop production is proposed as an appropriate model to address predation management for quail. Key Words avian recruitment, Callipepla squamata, Colinus virginianus, game birds, Integrated Pest Management, mesomammals, northern bobwhite, population regula- tion, predation, raptors, scaled quail

ecse of theirrelatively small size and the fact thatthey ment suggestthat predators are rarelya managementcon- | ."-'' theirentire lives on the ground,various species of cern and thatpredation should be managedonly indirectly R qui s(Odontophoridae)are extremelyvulnerable to pre- (e.g., habitatmanagement; Errington 1934). Hereinwe ii mn.However, prevailing paradigms in quailmanage- reviewthe evidencerelative to the impactsof predation

Address for Dale Rollins: Texas Agricultural Extension Service, Department of Wildlife and Fisheries Sciences, Texas A&M University, 7887 U.S. Highway 87, San Angelo, TX 76901-9714, USA; e-mail: [email protected]. Address for John P. Carroll: Daniel B. Warnell School of Forest Resources, University of Georgia, Athens, GA 30602, USA; e-mail: [email protected].

Wildlife Society Bulletin 2001, 29(1):39-51 Peer refereed ......

40 Wildlife Society, Bulletin 2001, 29(1):39-51

on northern bobwhite (Colinus virginianus, hereafter quail trends vary among ecoregions (Peterson and Perez bobwhite) and scaled quail (Callipepla squamata) as 2000). Bobwhite populations in the Rolling Plains and models for North American species. For bobwhites, we South Texas Plains ecoregions have remained relatively focus our discussion primarily on the eastern and western stable, but roadside counts in 2000 were the lowest since peripheries of the range, i.e., the southeastern United counts began for the Gulf Prairies and Marshes, Cross States and Texas, respectively, because bobwhite abun- Timbers and Prairies, and Edwards Plateau ecoregions

x Parks and Wildlife (Texas 2000).J- We do not discount the current managemtent paradigm of indirect predator control (i.e., halbitat man- Scaled quail However, the issue of ion as it Scaled quail range over most of agement).... predati the Chihuahuandesert, including relates to quail must be evaluated in a moire contem- portions of Arizona, Colorado, porary context of an increasingly fragmenited land- Kansas, New Mexico, Oklahoma, in and Texas. Scaled quail populations scape...and temporal changes predator populations. ^havedeclined significantly (from -3.8%/yr from 1966 to 1991 to dance typically declines along a west-to-east gradient. -8.2%/yr from 1982 to 1996) throughouttheir range, We also identify areas where additional research is need- especially during the last 15 years (Brennan 1993; Church ed and offer a more contemporary philosophy of preda- et al. 1993; Schemnitz 1993, 1994; Peterson and Perez tion management for quail, which is based on the philos- 2000; Rollins 2000; Sauer et al. 2000). Scaled quail ophy of Integrated Pest Management. populations experienced a drastic, inexplicable decline about 1989 over much of their range in Oklahoma and Current status of bobwhite north Texas (Rollins 2000). Populations in the Oklahoma panhandle declined 50% from 1956 to 1991 (Schemnitz and scaled quail 1993), and scaled quail essentially disappearedalong the Bobwhites eastern periphery of its range, where they were common The decline of the bobwhite in the southeastern United to abundantin 1987 (Rollins 2000). Data from the Breed- States (U.S.) is well documented (reviewed by Brennan ing Bird Survey (Sauer et al. 2000) and the Texas Parks 1999). Bobwhite populations declined -2.8%/year from and Wildlife Department (2000) documented this demise 1966 to 1999 (P<0.01) across its range according to (Figure 2). Relative to the Southeast, land-use changes Breeding Bird Survey data (Sauer et al. 2000, Figure 1). have been less dramatic in scaled quail range, which is The decline of bobwhites is correlated generally with dominated largely by livestock grazing. dramatic changes in land use throughout the region over the last 80 years. The shift away from a landscape domi- Factors to nated rather diverse and in the contributing quail by low-impact agriculture declines early twentieth century to landscapes dominated by hard- population wood forests and intensive pine silviculture in the latter Aside from the possible impacts of predation and twentieth century reduced habitability for bobwhites. In land-use changes, other factors may be involved in the more recent dramatic in times, changes agriculturalprac- 50 - tices (e.g., clean farming, increased use of pesticides) also x may have contributed to poorer quality or quantity 0

of remaining habitats. In any event, a landscape that i A-AA A and of is 'A -A A. supported large widespread populations quail 0 it A A now gone. Presently, bobwhite populations over most of AA.A A-A ,..., A A A 0 .A', il~~~~Cr'd A. A.A the Southeast seem to be generally at much lesser levels "3 AA.h - and very fragmented (Brennan 1999). Although bobwhite densities are generally greater at 0 ...... I I. . I I I . . . . I I. . I I I . . . . I I . I I , ~~, the western periphery of their range (e.g., Texas), their 67 71 75 79 83 87 91 95 abundance there has declined at a rate of since -4.7/year Year 1981 (Sauer et al. 2000). The Texas population's trend is 1. Northern bobwhite abundance in the southeastern U.S. (dot- essentially parallel to that of populations in the Southeast Figure ted line) and Texas (solid line) according to Breeding Bird Survey data, (Figure 1). Bobwhites occur over most of Texas, but 1966-99 (Sauer et al. 2000). Predators and quail * Rollins and Carroll 41

declineof bobwhiteand scaled Table1. Nest success rates(%) and percentageof mortalitieslost to predatorsof bobwhitenests at various locations. quailpopulations. Other factors believedto be to contributing Nest Methodof Lostto the declineof quail in the Location n success (%) monitoring predators(%) Reference Southeast fromfire ants range Illinois 863 34 Nest searching 37 Roseberryand Klimstra(1975) (Solenopsis spp.) to acid rain Florida 601 36 Nest searching 64 Stoddard(1931) (Brennan1999). However,there Florida 51 45 Telemetry 89 DeVosand Mueller(1993) also is ampleevidence that bob- Georgia 1,725 18 Nest searching - Simpson(1976) white populationsin some areas Missouri 157 44 Telemetry 68 Burgeret al. (1995b) of the Southeastare being main- NorthCarolina 35 34 Telemetry 73 Puckettet al. (1995) Oklahoma 161 50 76 et al. tainedat densities, Telemetry Peoples (1996) high espe- Tennessee 766 23 Nest search - Dimmick(1974) in the cially plantationregions Texas(north) 34 12 Nest search 88 Jackson(1947) of southernGeorgia and north- Texas(north) 81 46 Telemetry 91 Hernandez(1999) ern Florida. In these areassuit- Texas(south) 32 45 Nest search 84 Lehmann(1984) able landscapeshave been main- Weightedmean 28 tainedand quail are managed intensively(Burger et al. 1998). This suggeststhat landscape changes are importantat the ConservationReserve Program) were responsiblein the regionalscale, but thatadditional mechanisms may be Oklahomapanhandle. Rollins (2000) providedanecdotal operatingat the local scale. informationsuggesting that disease was the initialfactor Presentlythere is much speculationabout the role involvedand thathigh nest depredationrates (>80%) predatorsplay in the long-termdeclines of quailpopula- may have keptpopulations suppressed. Bobwhites, tions at a local scale, despitehabitat management (Hurst which are sympatricwith scaledquail over muchof the et al. 1996). Thereis little evidenceto suggestthat pred- Rolling Plainsecoregion, declined in aboutthe same time atorsare suppressingbobwhite populations at a regional period(1989-90), but have since reboundedand exhibit- level. However,anecdotal observations of predator ed irruptivepopulation changes typical of the species in removalwhere habitat management is being practiced this area(Jackson 1962, Petersonand Perez 2000, Sauer suggestthat some predatorsmay be suppressinglocal et al. 2000). Scaledquail remainedabsent or occurredat quailpopulations. only remnantlevels over much of theirformer range in How predatorsinteract with quail populationsmay be Texasfrom 1989 to 1999. affectednot only by the way landscapechanges have impactedhabitat, but likewise predatorpopulations, com- Predatorsof bobwhites munities,and searchefficiencies. For example,recent changesin land use may have madequail more vulnera- Predationis the primarysource of mortalityfor bob- ble to predation(Hurst et al. 1996, Rollins 1999a). Such whitesat all life stages. Thereare numerous studies, dat- changesmay operateat landscapeand local levels. ing backto Stoddard(1931), documentingthe impactsof The causes of the scaled quail decline are unknown. predatorson bobwhites,especially in the southeasternU.S. Schemnitz(1993) speculatedthat land-use changes (e.g., Nest depredation Rollins(1999a) estimatedthat only about4 of 100 eggs - 50 resultin a bobwhiteeventually being addedto the breed- 8 - A -40 x x cm ing populationin Texas. Nest depredationon bobwhites 0. 6- -30 0 has been studiedin variouslocations using a rangeof C) 4- - 20 techniquesfrom anecdotal observations to video monitor- coi 0- A k. I *A. A A ' L A I of hens. Estimatesof rateson 2 A , 'p,' t: 2 - 10 ing radio-tagged predation I II tI i I I I I nests are andhatch success rates 0 0 quail typicallyhigh, vary 67 71 75 79 83 87 91 95 99 from 12 to 50%(weighted x=28%, Table 1). Hatchrates of 36%, 18%,and 34% were reported from long-term Year studies in Florida, Illinois, and Georgia, respectively Figure 2. Scaled quail abundance in Texas according to Breeding Bird (Stoddard1931, Klimstraand Roseberry 1975, Simpson Survey data (solid line), 1996-99 (Sauer et al. 2000) and Texas Parks 1976). Hatchrates in Texas rangedfrom 12 to 46% and Wildlife Department roadside counts (dotted line), 1968-2000 (Texas Parksand Wildlife Department 2000). (Jackson1947, Lehmann1984, Hernandez1999). Peoples 42 Wildlife Society Bulletin 2001, 29(1):39-51

et al. (1996) recorded a 50% hatch rate in western Oklahoma, with predatorsaccounting for 81% of the losses. Accounting for predationon nests and adults, DeVos and Mueller (1993) suggested that 81% of nest losses were caused by some type of predation, with 52% of the losses attributedto mesomammals (i.e., medium-sized car- nivores). Other locally importantnest predatorsmay include corvids (Slater 1996), rodents (Stoddard 1931), armadillos (Dasypus novemcinctus; E. Staller, Tall Timbers Research Station and University of Georgia, unpublished data), and various snakes (Stoddard 1931). Bobwhites are persistent renesters, resulting in much greater percentages of hens actually producing chicks than would be suggested by low hatch rates (Burger et al. 1995b, 1995, Brennan 1999). Rollins (1999a) Figure 3. Rattlesnakes, although not a major predator of quail, have Guthery been documented on radio-marked in and Texas. a hatch rate of no hen mortali- preying quail Georgia estimated that, given 30%, The bulge in this western diamondback rattlesnake represents the ty, and 2 renesting attempts, 66% of hens would eventu- remains of a scaled quail wearing this telemeter. ally hatch a clutch of eggs. However, number of success- ful clutches decreased to 49 and 33% when hen mortality Brood survival was 20 and 40%, respectively. Chick survival is the least understood aspect of quail Mesomammals are the most important group of nest mortality (DeVos and Mueller 1993, Hurst et al. 1996). predators. In Virginia, Fies and Puckett (2000), using Researchers have attempted to assess mortality of chicks simulated ground nests containing quail eggs, found that after hatching, but logistical constraints have complicated 41% of nest predators photographed by motion-sensing such attempts (Carver et al. 1999). DeMaso et al. (1997) 37% cameras were striped skunks (Mephitis mephitis), reported a survival rate of 36% from hatching to 39 days were were opossums (Didelphis virginianus), 8% gray post-hatch in western Oklahoma. DeVos and Mueller foxes (Urocyon cinereoargenteus), and 4% were raccoons (1993) estimated 29% survival to 1 month post-hatch. (Procyon lotor). Hernandez et al. (1997) used similar Roseberry and Klimstra (1984) reported chick survival equipment to study nest depredation in west Texas and rates of 25-47% in southern Illinois. In Iowa, Suchy and reported that raccoons (82% of all nests destroyed) were Munkel (2000) reported survival rates of 81% for chicks the primary predator of simulated quail nests. Less com- 21-56 days post-hatch. They reported a cause-specific mon predators included striped skunks, bobcats (Lynx mortality rate of 14% for mammalian predation and 6% rufus), gray foxes, armadillos, and opossums. for avian predation. Stoddard (1931) identified several species of snakes Fire ants also may impact chick survival (Allen et al. that were important nest predators. Snake depredation of 1995, Mueller et al. 1999). Allen et al. (1995) found that bobwhite nests has been confirmed by recent camera-sur- bobwhite declines in southeastern Texas were correlated still cam- veillance studies in Virginia (passive infrared with a particularcounty's invasion by imported red fire eras, Fies and Puckett 2000) and in Georgia (continuous ants (S. invicta). Mueller et al. (1999) reported that 38% infrared video cameras; E. Staller, C. Sisson, W. Palmer, of all chick mortality up to 21 days post-hatch was attrib- and J. Carroll, University of Georgia, Auburn University, utable to fire ants. and Tall Timbers Research Station, unpublished data). Often snakes are diagnosed as the cause of nest depreda- Post-brood survival tion when no eggshells are found. For example, Peoples Adult survival also varies widely by season and caus- et al. (1996) implicated snakes in 55% of the nest losses es. Taylor et al. (2000) found that breeding season sur- recorded in western Oklahoma. However, Hernandez et vival over 4 years in Mississippi ranged from 17 to 51%. al. (1997) cautioned that snakes may be overly maligned Predators accounted for most of the mortality. During as an egg predator when diagnoses are based on lack of this study, avian mortality showed characteristics of eggshell evidence. Aside from depredating nests, rat being density-dependent; however, mammalian predation snakes (Elaphus sp.) and rattlesnakes (Crotalus spp.) continued to increase despite declining quail populations. have been documented preying on bobwhites in Florida Over 5 years on a plantation in Georgia where predators and bobwhite and scaled quail in Texas (Stoddard 1931; were actively controlled, Burger et al. (1998) found that Carter 1995; D. Rollins, unpublished data; Figure 3). breeding season survival was 41%. Mammalian (25%) Predators and quail * Rollins and Carroll 43

and avianpredators (20%) accounted for most of the 1973) cited predationas a managementconcern; in fact, mortality.Other studies documenting breeding season predationwas hardlymentioned as a sourceof mortality. survivalsuggested generally lower survivalrates, includ- Wallmo's(1957) only mentionof predationwas that ing 33% in Missouri(Burger et al. 1995a), 40% in which occurredat quail trappingsites; species involved Florida(Curtis et al. 1988), 34%in Mississippi(Taylor et were gray foxes, stripedskunks, ringtails (Bassaricus al. 2000), and 33%in NorthCarolina (Curtis et al. 1988, astutus), raccoons, and Cooper's hawks (Accipiter Puckettet al. 1995). Carter(1995) monitoredthe fate of cooperii). Schemnitz(1961) mentionedpredation only in 131 radio-markedbobwhites in west Texasduring that7 of 36 scaledquail nests (19%)were destroyedby 1994-95 and reporteda February-Julysurvival rate of predators.Campbell et al. (1973) calculatedannual sur- 13%. Mammalswere responsiblefor 56% of the kills, vival ratesof scaled quail fromband recoveries in south- whereasraptors caused 25%. Burgeret al. (1998) also easternNew Mexico as 6% on huntedareas and 10%on suggestedthat nonbreeding season mortalityof adults nonhuntedareas, but madeno mentionof whatproximate was attributedmainly to predationfrom mammals(25%) mortalityfactors were involvedother than hunting. and avianpredators (16%); overall adult survival aver- aged 49%. Pollock et al. (1989) reportedannual survival Relative of ratesof leg-bandedbobwhites from 10 to 24% (x= 17%) vulnerability sympatric over a 13-yearperiod in a huntedpopulation in Florida. bobwhite and scaled quail Huntingaccounted for 30%of the annualmortality, but Bobwhiteand scaledquail occur sympatricallyin por- cause-specificrates of naturalmortality (e.g., predation) tions of Texas,and some studieshave mentionedthe rela- could not be determined. tive vulnerabilityof these species to predators.Jackson (1947) suggestedthat predation was the proximatecause Predatorsof scaled of a catastrophicdecline in bobwhitesin northwestTexas quail winter1943. Bobwhitesand scaled were Nest during quail depredation sympatricon Jackson'sstudy site, and scaledquail Nest success for scaled is low quail typically (<25%, accountedfor about65% of the totalquail during Wallmo and has been cited as a 1957), depredation 1941-43. Totalquail density(bobwhite + scaledquail) if not cause of nest failure. Nest major, primary, preda- was high (estimatedat 1.6 birds/ha). Jackson(1947: tors commonin scaled include quailrange coyotes, 514) detailedhow bobwhitepopulations on his studyarea skunks, foxes, corvids various striped gray (Slater1996), crashed"with explosive suddenness and all but remnants snakes,and, feral et al. increasingly, hogs (Tolleson were lost to predation"between 7 and 15 January1943. 1993). Jackson(1942) reportedthat 10 of 13 scaled He conductedtransects to estimateamount of nests failed in the Texas In a more mortality quail panhandle. thathad occurredand concludedthat "everywhere the detailedreport from the same area,Jackson (1947) groundwas litteredwith evidencethat predation had that 30 of 34 bobwhitenests (88%)failed. He reported been recentand terrific"(Jackson 1947: 514). Northern attributedthe losses to snakes coyotes (11 nests), (6 harriers red-tailed hawks (Buteo and small mammals Schemnitz (Circus cyaneus), nests), (5 nests). (1961) and hawkswere the that 6 of 42 nests hatched. The jamaicensis), Cooper's raptors reported only (14%) pri- involved,but Jacksonconcluded that northern harriers marycause of nest failurewas humandisturbance (e.g., were the only raptorspecies abundantenough in thatarea farmingpractices and mowing),and predatorswere to have killed so manyquail. Fourof 18 northernharri- implicatedin only 19%of the nest losses. Recentstudies ers examinedhad consumedbobwhites. Jacksonreport- (Hernandezet al. 1997, Fies and Puckett2000) suggested ed thatevidence of predationon scaled quail was "light" thatthe accuracyof assigningspecies-specific causes of and thatscaled quail were apparentlyless vulnerableto quail nest depredationis tenuousat best. avianpredation than were bobwhites. Broodsurvival P. S. Carter(Angelo StateUniversity, unpublished We foundno publishedreports on chick or broodsur- data)radio-marked 27 scaled quailin west-centralTexas and vival for scaled quail. As scaled quailpopulation "busts" (IrionCounty) reportedhigher survival (70%) from are characterizedby poor recruitment,information on Februaryto July thanfor sympatricbobwhites (18%, n= In Carter's chick survivaland broodecology is sorelyneeded. 54). study(for both species of quail)60% of mortalitieswere attributedto mammalianpredators. He Post-broodsurvival documented9 scaled quailmortalities, 5 from mammals, None of the 3 majorautoecological studies on scaled and 2 fromunknown raptors. One scaled quail was quail (Wallmo1957, Schemnitz1961, Campbellet al. killed by a greathorned owl (Bubovirginianus) and one 44 Wildllife Societl' Bullletil 20)(01,29(1):39-51

by a westerndiamondback rattlesnake (C. atrox). in cover thatwas uniformwith the surroundings. These datasupport Jackson's (1947) observationthat Becausequail population "busts" are usuallyassociat- scaledquail may be less vulnerableto predationthan ed with droughtconditions in the southwesternU.S. and bobwhitesand Lehman's(1984: 225) opinionthat "blue often confoundedby overgrazing,suitable nesting cover (i.e., scaled) quail seem somewhatmore intelligentthan is often limitedin dry years. Slateret al. (2001) found bobwhites"in sympatricranges. thatnest success of simulatedquail nests in 8 countiesin It seems plausiblethat earlier investigations of scaled west Texaswas greateron sites thatprovided >760 quail (Wallmo1957, Schemnitz1961, Campbellet al. potentialnests sites/ha,a numbersimilar to Guthery's 1973) eitherwere unawareof or dismissedthe incidence (1986) recommendationof >500 suitablenest clumps/ha of predationbecause they lackedthe technologyto study for bobwhitesin Texas. Carter(1995) foundthat sym- it (i.e., radiotelemetry).Rollins (2000) dividedthe patricbobwhites and scaled quailfrequently used prickly knowledgeabout scaled quail ecology into 2 distinct pear(Opuntia spp.) for nestingsites. Subsequently,Slater eras: beforetelemetry and aftertelemetry. More com- et al. (2001) documentedthat nests situatedin prickly prehensivestudies involving radio-marked scaled quail pearsurvived at abouttwice the rateof moreconventional are neededto assess cause-specificmortality patterns. nest sites (i.e., bunchgrasses).Thus, prickly pear appears to providesome measureof protectionagainst nest predators,especially when traditionalnest sites are limitedby Role of predationin quail irruptions overgrazingor drought(Hernandez 1999). Bobwhiteand scaled quail exhibitirruptive population growthin Texas(Jackson 1962, Lehmann1984). Population"busts" are believedto be a resultof normal Temporalchanges in predator attritionbut below-normalreproduction (Wallmo 1957). populationsand communities Such buststend to be characterizedby droughtconditions Ourreview of publishedresearch suggests that bob- (Wallmo1957, Campbellet al. 1973, Giulianoand Lutz white and scaled quailpopulations have changedat local 1993), but scaled quail appearto be moreproductive dur- andregional scales. Whatabout their predators? ing droughtyears than are sympatricbobwhites Comparingearlier studies (e.g., Stoddard1931) to more (Schemnitz1964, Lehmann1984, Rollins 2000). contemporarystudies suggests that changes have Irruptionsappear to be relatedindirectly to rainfall,pos- occurredwithin populations and communitiesof various sibly throughsome plant-relatedstimulus (e.g., nutri- predatorsthat are often implicatedin the decline of quail tion). Variousinvestigators have proposedvitamin A populations.Such temporalchanges in predatorpopula- deficiencies(Nestler 1946, Lehman1953), phytoestro- tions may be important,especially in light of landscape gens (Cainet al. 1987), and waterdeprivation (Koerth changesthat may makequail morevulnerable to preda- and Guthery1991) as possible explanationsfor reproduction (Rollins 1999a). tive failuresin quail in the southwesternU.S. An alternatehypothesis is thatprecipitation increases Mesomammaltrends nestingcover acrossthe landscape,i.e., "usablespace" Thereis generalconsensus that mesomammal popula- (Guthery1997: 294), and subsequentlyincreases nesting tions (e.g., raccoons)have increasedover the last 20 success by complicatingthe predators'search efficiency yearsin the Southeast. Rollins (1999a) identifieda num- (Rollins 1999a). Quailirruptions in the RollingPlains ber of mechanismsthat may be contributingto greater ecoregionof Texasare characterizedby landscapesdomi- mesomammalpopulations or otherwiseaccentuating pre- nated by common broomweed (Xanthocephalum dracun- dationon quail and theirnests. These mechanisms culoides,Jackson 1962, Rollins 1999b). Dense canopies include 1) demise of the fur marketin the mid-1980s,2) of commonbroomweed effectively "insulate" quail from increasedsupplemental feeding of deer (Odocoileus predators(avian and mammalian)and hence increase spp.), 3) increasinglyfragmented habitats, and 4) a pro- "usablespace." liferationof farmponds on the landscape. Otherauthors Predatorsearch efficiency may decline as abundanceof have speculatedon the impactsof variousof these factors suitablenest sites or habitatheterogeneity increases on recruitmentin galliformes(Palmer et al. 1993, Hurst acrossthe landscape(Bowman and Harris1980). It is et al. 1996). Raccoons,a primarynest predatorin Texas moredifficult for predatorsto locate groundnests in areas (Hernandezet al. 1997), probablybenefit by an increas- supportingabundant bunchgrasses compared to areaswith ingly popularpractice of providingsupplemental feed to few bunchgrasses(Jackson 1947). Lehmann(1984) noted deer in Texas. As manyas 12 raccoonshave been pho- greaternest survivalin areaswhere the nest was situated tographedat a single free-choiceprotein feeder (D. Predators and quail * Rollins and Carroll 45

Plains, typically has the least quail abundance of these 3 ecoregions. The fact that the Edwards Plateau has the lowest coyote densities in Texas (because of a history of sheep and goat ranching in this area [Nunley 1985]) suggests that coyote suppression may "release" mesomammals like raccoons, gray foxes, and feral cats. Additional studies are needed to document this relationship, however, as edaphic factors also differ among these 3 ecoregions. Other mammals When Stoddard (1931) undertook his studies on plantations in southern Georgia, striped and spotted skunks (Spirogale putorius) were important predators of quail nests. However, recent video data from several hundred Figure 4. The increasing popularity of supplemental feeding for white- nests on plantations in the same region found no evi- tailed deer be raccoon in Texas. may influencing populations dence of predation by either species (E. Staller, C. Sisson, W. Palmer, and J. Carroll, University of Georgia, Rollins, unpublished data; Figure 4). Cooper and Ginnett Auburn University, and Tall Timbers Research Station, (2000) found that simulated ground nests (e.g., chicken unpublished data). This video surveillance of nests also eggs) had lesser survival rates in 2 of 3 years on sites confirmed armadillos as a predator of bobwhite nests, where deer feeders were present. substantiating the findings of Hernandez et al. (1997) in Another example of temporal changes in a predator Texas. During Stoddard's era, there were no armadillos community is suggested by comparing 2 studies conduct- in that region, but today they are ubiquitous. ed in north-centralTexas (Wise and Parkercounties). Finally, the distribution and abundance of feral hogs Jackson (1952) removed potential quail predators (n= (Sus scrofa) has increased over much of the Southeast 574) from a 1,200-ha study site in Wise County, Texas, and Texas (Tolleson et al. 1993). Feral hogs were impli- over a 13-month period (1948-49), but dismissed the cated in 9 and 24% of the simulated nest losses in predatorremoval as having no impact on quail abundance. Shackelford and Foard counties (respectively) in Texas. Of particularnote, only 11 raccoons (2.0% of the preda- However, the impact of feral swine depredation on quail tors removed) were trapped during his study. Fifty years nests is unclear. Similar to coyotes, those areas of Texas later, E. Lyons (Angelo State University, unpublished with the greatest feral hog abundance (e.g., Rolling data) removed 21-40 raccoons from 2 study sites (each Plains, Rio Grande Plains) also support the greatest quail 260 ha) during only 30-day trapping efforts in an adjacent populations. county (Parker)during 1999 and 2000, respectively. In other words, Lyons removed about 3 times more raccoons Status of avian predators than Jackson did on study sites only 20% the size of Among common avian predators of bobwhites, popu- Jackson's sites and with only 10% of the trapping effort. lation increases of >2.0%/year have been observed using In Mississippi, hunter harvest data from 1980 to 1996 the Breeding Bird Survey (1966-99) over large areas of suggested that red foxes (Vulpes vulpes), gray foxes, and 4 - bobcats remained stable but that coyote (Canis latrans) populations increased 7-fold (Lovell et al. 1998). The A 03- -. .f between and is unclear. relationship coyotes quail .5 , .Ic-~~~~A h .AAh - Lehmann (1984) identified coyotes as perhaps the most a_ .A.. I2- common mammalian predatorof bobwhites in south ,AAA ~A..-A but concluded that . A. Texas, Guthery (1995) controlling coy- mli^ h * otes would not increase the . A --ti likely quail productivity given ^^^^^^_^_^_^_ bGA quail's ability to renest. Interestingly, the greatest bob- 01io . white populations are found typically in the Rolling 67 71 75 79 83 87 91 95 Plains and Rio Grande Plains and these are ecoregions, Year the same areas of Texas that typically harbor the greatest 5. hawk and hawk (dotted densities of the Edwards Plateau Figure Cooper's (solid line) sharp-shinned coyotes. Similarly, line) abundance in the U.S. according to Breeding Bird Survey data, ecoregion, located between the Rolling and Rio Grande 1966-99 (Sauer et al. 2000). 46 Wildlife Society Bulletin 2001, 29(1):39-51

the U.S. (Saueret al. 2000). Trendsfor accipiters,e.g., duction. Gutheryand Beasom (1977) conducteda simi- the Cooper'shawk (6.7%/year,n=359, P<0.01) and lar studyof intensiveremoval of mammalianpredators sharp-shinnedhawk (A. striatus,2.8%/year, n=233, P= (e.g., coyotes, stripedskunks) from a 15-km2study area 0.03), have increasedsteadily over the last 30 years in the westernRio GrandePlains of Texas,but could not (Figure5). Factorsresponsible for the increaseof vari- demonstratea treatmenteffect on eitherbobwhite or ous avianpredators of quail are unknown,but could scaled quailpopulations. Their conclusion was thatif includethe dissipationof organochlorineinsecticides, predatorremoval was effectiveat all, the effect wouldbe increasedlaw enforcement,and educationalefforts on demonstratedby allowingquail populations on "poorer" raptorconservation. Accipiters are generallyconsidered areasto be similarto betterhabitats. the most efficientpredator of quail, and Stoddard(1931: Burgeret al. (1998) measuredbobwhite mortality and 212) characterizedCooper's hawks as "theoutstanding predationrates on intensivelymanaged plantations in naturalenemy of the bobwhite."Other raptors, such as Georgiaand foundthat 35% of mortalitieswere a result the greathomed owl, have not increasedover the whole of predationby mammals,even with intensivemeso- southeastern region (0.5%/year, n= 136, P=0.82), but in mammalcontrol. However,there were no comparative some states,such as Georgia,they have increased datawhere predators were not controlled. (5.5%/year,n=23, P=0.002). Errington(1934) consid- If an effect is to be realizedfrom reducingpredators, it ered greathorned owls an importantpredator of bob- will most likely resultfrom reducing potential mesomam- whites in Wisconsin. Greaterroadrunners (Geococcyx mals involvedin nest depredation(Rollins 1999a). californianus)are implicatedoften as a seriouspredator However,reducing the populationsof mammaliannest of quailnests and chicks in Texas. However,recent predatorsis laborintensive, costly, and will not necessar- researchin southTexas (C. Ruthven,Texas Parks and ily resultin an increasein quail abundance.Frost (1999) WildlifeDepartment, personal communication) found removedapproximately 1 mesomammalper 5 ha (mostly quail remains(2 chicks) in only 1 of 120 roadrunner raccoons)from 260-ha studyareas over a 30-dayperiod stomachs. Nevertheless,roadrunner abundance in the just priorto the 1998-99 nestingseasons in TomGreen ChihuahuanDesert has increased3.6%/year over the last County,Texas. Survivalof radio-markedbobwhites and 30 years (n=32, P=0.19, Figure 6). fate of simulatedquail nests were similaron trappedand nontrappedsites. Scent stationsindicated that at this Effectsof predatorreduction on quail scale and level of trapping(75 trapnights/ha), mesomam- populations mal abundancewas not reducedeven in the shortterm. Empiricalevidence of the impact(or lack thereof)of predatorremoval on quail abundanceis limited. Beasom (1974) studiedthe effects of intensivepredator control on Discussion I I I 1 I ,. .III I .I . X., It if . bobvvhltes and wild turkeys(Meleagris gallopavo) in the Althoughpredation is usuallythe primarysource of eastern Rio GrandePlains of Texas. He removed188 mortalityfor quail at all stages of theirlife cycle, preda- coyc tes, 120 bobcats,65 raccoons,46 stripedskunks, and tor controlhas historicallybeen dismissedas a manage- 38 other mammalian predators from a 23-km2study area mentrecommendation for quail. Errington's(1934) over a 2-yearperiod. He observedmoderate gains in long-termstudies of bobwhitesand predators in the bobvahite abundanceand strongincreases in turkeypro- upperMidwest suggested that habitat, not predators,limited bobwhites. His concept-i.e., managehabitat, not 4 predators-has been pervasivein the quail management literaturesince thattime. We do not discountthe current x lE 3 - X^ managementparadigm of indirectpredator control (i.e.,

c habitatmanagement) and especiallyas the "firstline of o2 defense." However,the issue of predationas it relatesto

0. quailmust be evaluatedin a morecontemporary context i 1 of an increasinglyfragmented landscape (Robel 1993) and temporalchanges in predatorpopulations. In light of 0 ' ' ' ' ' ' ' these changes, and the current rate of declines observed 67 71 75 79 83 87 91 95 . . . in quail in some regions, we concur with Hurst et al. Year (1996) thatthe issue of predatorcontrol relative to avian Figure 6. Greaterroadrunner abundance in the Chihuahuandesert recruitment in Galliformes should be revisited. accor to Bird 1966-99 et al. ding Breeding Surveydata, (Sauer 2000). If we compare the prevailing thoughts of predation on Predators and quail * Rollins and Carroll 47

quailwith those for a species for which predationand et al. 1996, Redpathand habitatmanagement have been studiedmore thoroughly Thirgood1997, Kauhala i b (e.g., gray partridge[Perdix perdix]), we find some inter- et al. 2000). How these esting parallelsand contradictions.Potts (1986) modeled resultstranslate to more the populationdynamics of gray partridgeusing a variety complexecosystems of datasources in the United Kingdom. Potts (1986) pre- remainsto be seen. sentedevidence of density-dependentrelationships for More diverse (i.e., com- survivalof gray partridgefor majorcomponents of their plex) systemsin the life history,including nesting, hunting, and over-winter Southeastare character- survival,as also was suggestedby Roseberryand ized by competing Klimstra(1984) for bobwhites. However,Potts (1986) risks;accordingly, a also demonstratedthat some aspectsof the life historyof reductionin one preda- graypartridge survival were impactedby density-inde- tor's abundance does Telemetrystudies of scaled quail have laggedbehind those of bobwhites. pendentfactors, chick survivalbeing the most important not necessarilyensure a factornot relatedto birddensities. He suggestedthat greaterhatch or recruit- predatorcontrol was very importantin determiningthe ment rate(Guthery 1995). relativeimportance of density-dependentmortality, espe- The difficultyencountered when tryingto understand cially nest mortality,in gray partridgeand thatthe rela- the efficacy of predatorcontrol in quail managementis tive shapeof mortalitycurves with predatorcontrol thatbiological and social (i.e., political)considerations resultedin lower mortalityrates at greaterpartridge den- are intertwined.In the examplesoutlined previously for sities. Potts'(1986) datasuggested there was still density- gray partridgein the UnitedKingdom, predator manage- dependentmortality in partridgenesting even with preda- ment has been demonstratedto be beneficialto produc- tor control,but thatthis relationshipwas much weaker. tivity and populationdensities of an importantgame bird. Tapperet al. (1996) went on to demonstratethe suppres- Thereare, however,a couple of key differencesbetween sive effect of nest predatorson gray partridgeproductivi- that system and attemptsto managequail and predators ty, recruitment,and populationswith a 6-yearpredator in the U.S. removalexperiment. First,the predator-preycommunity on farmlandin the Burgeret al. (1998) suggestedthat annual mortality United Kingdomis much simplerthan those found in the rateswere not a good measureto developan understand- Southeast. Much of the rangeof bobwhiteand scaled ing of the populationdynamics of bobwhites. They point quail harborsgreater numbers and diversityof predator out thatthe annualsurvival rates they measuredwere and prey species. Thus, thereare morecomplex interac- only slightlygreater than those for otherstudies in the tions and possible outcomesas we begin to alterthe com- Southeast,yet bobwhitedensities on theirstudy were munity. Second, managementof gray partridgeis occur- muchgreater than other studies in the region. Burgeret ring in a very intensivelymanaged agricultural landscape. al. (1998) suggestedthat predator removal changed the In effect, the argumentbecomes one in which a desire to dynamicsof mortalitywithin seasons in supportof Potts' have largernumbers of gray partridgehelps to mitigate (1986) theory. We believe thatBurger et al. (1998) are the intensiveland managementfor productionagricul- demonstratingthat suppressionof predator-mediated, ture. This can positivelyimpact land managementfor density-dependentmortality produces greater recruitment not only the desiredgame birdbut also for otherspecies, rates. This resultsin continuedhigh levels of productivi- includingthe predators.Predator management helps ty becausethe density-dependentmortality of nests also enhancehabitat management, thereby making the system is suppressedby predatorcontrol, thereby providing at more tenablefor those wantingto huntgray partridge. least a possible solutionto the annualmortality conun- The situationin the Southeastis one characterizedby drumpresented by Guthery(1997). a much greatervariety of ecosystemsmanaged at differ- ent intensities. If predatormanagement to enhancequail Predator control in simple versus complex populationsis done simply to increasethe numberof environments quail shot by hunters,then argumentsfrom a conserva- Among those studiesdemonstrating a positiveimpact tion perspectiveare diminished. However,arguments of predatorcontrol in one form or anotheron game bird demonstratingthe positiveimpact of intensivequail man- populations,most have occurredin simplerecosystems agementon otherspecies also could be made (Webband and with simplerpredator and prey communitiesthan Guthery1983, Harvesonet al. 2000). Withmore diverse those foundin the Southeast(Sargeant et al. 1995, Tapper predatorand prey communities,we are then requiredto 48 Wildlife Society Bulletin 2001, 29(1):39-51

makethese decisionsin a more complex system (e.g., we Messmeret al. (1999) surveyedpublic opinion in the are also likely to makemistakes and reducespecies that U.S. aboutmanaging predators to enhanceavian recruit- resultin no net benefitto quail). ment. Whengiven specific predatorcontrol scenarios, respondentssupported control to enhanceavian recruit- Predatorcontrol or predationmanagement? ment,except for controllingraptors to protectexotic As bobwhitepopulations continue to decline in the uplandgame birds. Theirresults suggested that public Southeast,there is increasingpressure to implement supportfor predatorcontrol is greaterwhen controlprac- predatorcontrol to increasebobwhite abundance. Some tices are applied"surgically" rather than applied broadly. conservationorganizations, e.g., QuailUnlimited, are Thus, an IPM model for predationmanagement on quail increasinglyquestioning the "if you buildit, they will could be supportedby the generalpublic. come"habitat paradigm as the sole meansof sustaining bobwhitepopulations. In calling for broad-spectrum predatorcontrol, some in and out of the wildlife profes- Conclusion sion may be actingprematurely. As Leopold(1953: 60) Changesin land managementover the last 30 years suggested,"the urge to comprehendmust precede the have resultedin conditionsthat make it more difficultto urge to reform."Waterfowl managers have done an maintainhigh densitiesof quail (especiallybobwhites) admirablejob in the quest to understandthe ecological over muchof theirdistribution. There is no doubtthat implicationsof predatorcontrol and subsequentlypreda- landfragmentation will continueand likely accelerate tion management.Quail managers may be well advised over the next 20 yearsin bobwhiteand scaledquail to study such examples. As was suggestedby Tapper ranges(e.g., Texas;Wilkins et al. 2000). At the same (1999) and Reynoldsand Tapper (1996), we mightnot time, thereis evidencethat some predatorsof quail may even need to see a net reductionin predatornumbers to have benefitedfrom these changes. How these land- have a positiveimpact on game birdpopulations. scape-levelchanges in landuse andpredator and prey As in the northernplains, some predatorsmight populationsimpact the interactionof quail and their adverselyimpact those predatorspecies thatprey heavily predatorsis unclearat this time. But our challengeas uponnesting game birds. For example,coyotes at low quail managersis apparent:how to maintain(or restore) densitieswill displacered foxes, therebyresulting in quailpopulations in an increasinglyfragmented habitat. greaterduck nest survival(Sovada et al. 1995). Similar We suggestthat appropriate predation management tech- correlationsbetween coyote densitiesand bobwhiteshave niques shouldbe one of the tools consideredin such been reportedin Texas(Rollins 1999a). Coyotesmay restorationefforts. suppresssmaller, more efficient nest predators(e.g., gray Althoughwe have discussedquail population dynam- foxes, raccoons),or at least restricttheir distribution on ics on a regionalscale, the interactionof habitat,predator the landscape. communities,and quail is likely to be playedout on a finer spatialscale. For example,the presenceor absence An integratedapproach to predation of a hardwooddrain in a fire-maintained,longleaf pine management (Pinuspalustris) community might have an enormous We suggestthe developmentof an IntegratedPest influenceon the dynamicsof a local predatorcommunity, Management(IPM) approach for managingpredators of regardlessof what mightbe happeningon a regional quail. The conceptof IPM was developedto enhance scale. Fartherwest, the interactionsof livestockgrazing, strategiccontrol of pests in crops (Pedigo 1989) and rec- droughtperiodicity, and subsequentlynest-site availabili- ognizes thata species of insect may be eithera "pest"or ty may increasethe vulnerabilityof quailto predation. "beneficial,"depending on the situationinvolved. The potentialrole of predationas a suppressingagent Further,IPM introducesthe idea of economicthresholds, in quailpopulations needs additionalstudy. It is crucial i.e., the level of pest damage that can be sustained before to understandhow landscape-levelchanges in landuse it becomes economicalto providea correctivetreatment. mightchange relationships between quail and theirpred- Most IPM strategiesinclude nonlethal (e.g., croprota- ators,as well as changepredator and prey communities. tions) and lethal (e.g., insecticides)control alternatives. Whatis neededis experimentalresearch to definemore The formeris appliedas the first line of defense,with the clearlythe relationshipsbetween quail and theirpreda- latterbeing appliedin the most "surgical"manner feasi- tors withinthe contextof currentland-use and habitat ble to reducetreatment costs and minimizerisks to the management.Leopold and Hurst(1994) outlinedstrate- environment.Appropriate parallels relative to predator gies to studyimpacts of predatorson game birdmanage- managementfor quail are numerous. ment. Specifically,the relationshipsamong precipitation, Predators and quail * Rollins and Carroll 49

vegetationdynamics (including nest-site availability [i.e., CARTER,P. S. 1995. Post-burn ecology of northern bobwhites in west land livestock Texas. Thesis,Angelo State University,San Angelo,Texas, USA. thresholds]), management(e.g., grazing, A. L.W ANDL.A. BRENNAN.1999. Passive and con- CARVER, V, BURGER,JR., integrated brushcontrol), predator searching effectiveness, transponders and patagial tag markers for northern bobwhite sequentnesting success and recruitmentneed additional chicks. Journal of Wildlife Management 63:162-166. investigation(Schemnitz 1994). CHURCH, K. E.,J. R. SAUER,AND S. DROEGE.1993. Population trends of in North America. 44-54 in K. E. Church andT.V Dai- An IPM-basedapproach to predatormanagement for quails Pages ley, editors. Quail III:national quail symposium. Kansas Department quailneeds to be developed. Informationis neededto Wildlife and Parks,14-17 July 1992, Pratt,USA. developeconomic thresholds and integratedpredation COOPER,S. M.,ANDT. E GINNETT.2000. 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Texas Agri- predator deterrent in nest site selection by northern bobwhite (Col- cultural Extension Service, 23 March 1996, Corpus Christi, USA. inus virginianus). Thesis,Angelo State University,San Angelo, PETERSON,M.J.,AND R. PEREZ.2000. Is quail hunting self-regulatory? Texas, USA. Northern bobwhite and scaled quail abundance and quail hunting SLATER,S. C., D. ROLLINS,R. C. DOWLER,AND C. B. SCOTT.2001. Opuntia: a in Texas. Pages 85-91 in L.A.Brennan,W. E. Palmer,L.W Burger,Jr., "pricklyparadigm" for quail management in west Texas. Wildlife and T. L. Pruden, editors. Quail IV:Proceedings of the Fourth Society Bulletin 29: in press. Predators and quail * Rollins and Carroll 51

GRIER. Differential effects of ' SOVADA,M.A.,A. B. SARGEANT,ANDJ.W. 1995. :.vi:st::c WeDaleRollins (photo) is a profes- coyotes and red foxes on duck nesting success. Journal of Wildlife sor and extensionwildlife spe- Management 59:1-9. a F cialist with the Department of Wildlife and Fisheries Sciences STODDARD,H. L. 1931. The bobwhite Its habits, preservation, and quail: at TexasA&M Sta- increase. Charles Scribner and Sons, New York,New York,USA. : i University. p tioned in San Angelo, he con- SUCHY,W. J., AND R.J. MUNKEL. 2000. Survival rates of northern bobwhite around ecolyr ducts outreach education across chicks in south-central Iowa. Pages 82-84 in L.A.Brennan,W. E. the western half of Texas. He and T. L. editors. IV:Proceed- Palmer,L.W. Burger, Jr., Pruden, Quail H~-e~obtai earned his B.S. degree in biology ings of the Fourth National Quail Symposium. TallTimbers s from Southwestern Oklahoma Research Station, 6-9 May 1997,Tallahassee, Florida,USA. : State University, his M.S. in wild- TAPPER,S., editor. 1999.A question of balance: Game animals and their life ecology fromOklahoma role in the British countryside. The Game Conservancy Trust, Ford- State University, and his Ph.D. in ingbridge, United Kingdom range science from Texas Tech l i He served as TAPPER,S. C., G. R. POTTS,AND M. BROCKLESS.1996. The effect of an University. presi- dent of the Texas Chapter of The reduction in on the breeding suc- experimental predation pressure Wildlife Society in 1997 and as cess and population density of grey partridges Perdix perdix. Jour- a director for the Texas Section nal Applied Ecology 33:965-978. of The Societyfor RangeMan- TAYLOR,J. D., III, L.W. BURGER,JR., S.W. MANLEY,AND L.A. BRENNAN. 2000. agement. Dale'sfondness for Seasonal survival and cause-specific mortality of northern bob- quail shows in his programsand whites in Mississippi. Pages 103-107 in L.A.Brennan,W. E. Palmer, popularwritings (he is a colum- L.W. Burger, Jr., and T. L.Pruden, editors. Quail IV:Proceedings of nist for Quail Unlimited maga- the Fourth National Quail Symposium. TallTimbers Research Sta- zine, Livestock Weekly, and the TexasFarmer-Stockman). He coordinatesthe BobwhiteBrigade, a tion, 6-9 1997,Tallahassee, Florida,USA. May wildlife leadershipprogram for high school youth, and has conceived TEXASPARKS AND WILDLIFEDEPARTMENT. 2000. forecast 2000-2001. Quail programslike Quail AppreciationDays. His researchinterests center (http://www.tpwd. state.tx.us/hunt/regs/quail/). aroundreproductive ecology of bobwhiteand scaled quail. John P. TOLLESON,D. R., D. ROLLINS,W.E. PINCHAK,M. IVY,ANDA. HEIRMAN.1993. Carrollis an assistantprofessor of wildlife ecology and management Impact of feral hogs on ground-nesting gamebirds. Pages 76-83 in in the WarnellSchool of ForestResources at the Universityof Georgia. C.W.Hanselka and J. E Cadenhead, editors. Feral swine: a compendi- He obtaineda B.S.degree in wildlife biology fromthe Universityof um for resource managers. Texas Agricultural Extension Service, Massachusetts,an M.S. in biology from EasternKentucky University, 24-25 March 1993, Kerrville,USA. and a Ph.D. in biology fromthe Universityof NorthDakota. His researchfocus is on land and of WALLMO,O. C. 1957. Ecology of scaled quail in west Texas. Disserta- management populationdynamics birds. He is also interestedin the and managementof tion,Texas A&M College Station, USA. game biology University, Galliformesin tropicalAmerica, Africa, and Asia. He cur- F S. GUTHERY. Avian to habitat endangered WEBB,W. M.,AND 1983. response manage- rentlyserves as chairof the Partridge,Quail, and FrancolinSpecialist ment for northern bobwhites in northwest Texas. Journal of Wild- Groupfor the IUCNSpecies SurvivalCommission. life Management 47: 220-222. WILKINS,N., R. D. BROWN,R.J. CONNER,J. ENGLE,C. GILLILAND,A.HAYS, R. D. SLACK,AND D.W STEINBACH.2000. Fragmented lands: changing land ownership in Texas. Texas Agricultural Extension Service, Publica- tion MKT-3443,College Station, USA. Associate editor: Conover