YELLOWSTONE NATIONAL PARK

Transcription

1 YELLOWSTONE NATIONAL PARK MONITORING PLAN FOR YELLOWSTONE BISON Prepared by: P. J. White, Chief, Wildlife and Aquatic Resurces Jhn Treanr, Yellwstne Wildlife Health Prgram Rick Wallen, Bisn Eclgy and Management Prgram Apprved August 2008 Updated Nvember 2012 TABLE OF CONTENTS PURPOSE AND RATIONALE FOR MONITORING 2 MONITORING ACTIVITIES 2 IMPLEMENTATION AND EXPECTED OUTCOMES 3 MONITORING SYSTEM EVALUATION 4 MONITORING FINDINGS 5 LITERATURE CITED 24

2 PURPOSE AND RATIONALE FOR MONITORING The successful cnservatin f plains bisn (Bisn bisn) in Yellwstne Natinal Park frm abut tw dzen animals in 1901 t 5,000 animals in 2005 has led t an enduring series f disagreements amng varius publics and management entities regarding bisn abundance and distributin, and the ptential transmissin f the Brucella pathgen t dmestic cattle (Plumb et al. 2009). Als, since the State f Mntana and the Natinal Park Service (NPS) agreed t the curt-mediated Interagency Bisn Management Plan (IBMP; United States Department f the Interir [USDI] and United States Department f Agriculture [USDA] 2000a, b), prgress has been slw at cmpleting the plan s successive management steps. Thus, the Gvernment Accuntability Office (2008) recmmended that the IBMP agencies develp specific management bjectives, cnduct mnitring t evaluate the effects and effectiveness f management actins, and develp methds fr adjusting the IBMP based n these assessments. These recmmendatins were implemented by the IBMP agencies thrugh an adaptive management plan in 2008 (USDI et al. 2008). Als, under prvisins f the IBMP, the NPS is cnsidering implementatin f a lng-term, remte delivery, vaccinatin prgram fr brucellsis in free-ranging bisn inside Yellwstne Natinal Park (USDI 2010). Thus, there is a need t estimate key parameters f bisn and brucellsis dynamics, and evaluate the likely effects and effectiveness f a variety f management activities. This plan identifies and reprts n a suite f lng-term mnitring and research activities fr Yellwstne bisn that meet the missin f the NPS and infrm adaptive management. MONITORING ACTIVITIES The varius types f actins in the IBMP t ensure the cnservatin f a viable, free-ranging bisn ppulatin while safely and effectively reducing infectin frm, and transmissin f, the nn-native Brucella bacteria can be gruped int three general categries: 1) managing brucellsis transmissin risk; 2) reducing the prevalence and transmissin f brucellsis; and 3) cnserving a viable ppulatin f wild bisn and the eclgical prcesses that sustain them (Figure 1). Thus, we develped management and research bjectives fr these desired cnditins that are multidimensinal and invlve trade-ffs, whereby imprving an utcme assciated with ne bjective affects utcmes assciated with ther bjectives (Williams et al. 2007). We als develped ne r mre sampling bjectives fr each mnitring activity (White et al. 2008). DESIRED CONDITIONS Bisn abundance averages 3,000-3,500 per decade, while maintaining 95% f existing genetic diversity. Increased tlerance fr bisn utside Yellwstne, while maintaining bisn separatin with cattle. Mre than 50% decrease in brucellsis prevalence in bisn. Manage Brucellsis Transmissin Risk Cnserve a Wild Bisn Ppulatin Brucellsis Suppressin Separatin t prevent bisn-cattle mixing Cattle management Management culls and harvests Adaptive management Research: disease dynamics/transmissin Migratry behavir Eclgical rle and functin in ecsystem Natural selectin/evlutinary ptential Demgraphic health Vaccinatin Culling infectius bisn Disease surveillance Adaptive management Brucellsis research Figure 1. Cnceptual mdel f cnservatin and disease management fr Yellwstne bisn. 2

3 We then develped the fllwing mnitring activities t prvide timely and useful infrmatin t help develp adaptive management adjustments. Cnservatin (Preserve a Functinal, Free-Ranging Bisn Ppulatin) 1. Estimate the abundance, demgraphy, and limiting factrs fr the verall bisn ppulatin and tw primary subppulatins (i.e., central and nrthern breeding herds). 2. Describe migratry and dispersal mvements by bisn at a variety f tempral and spatial scales in and utside the park. 3. Estimate the existing heterzygsity, allelic diversity, and lng-term prbabilities f genetic cnservatin fr the verall bisn ppulatin and identified subppulatins. 4. Prmte cperative cnservatin in bisn management by partnering with states, Tribes, and thers interested in bisn health and recvery. Risk Management (Prevent Brucellsis Transmissin frm Bisn t Livestck) 5. Estimate the prbabilities (i.e., risks) f brucellsis transmissin amng bisn, cattle, and elk, and between the elk feed grunds in Wyming and nrthern Yellwstne. 6. Estimate age-specific rates f bisn testing serpsitive and sernegative fr brucellsis that are als culture psitive, and the prtin f serpsitive bisn that react psitively n serlgic tests due t expsure t crss-reactive agents ther than B. abrtus (e.g., Yersinia). 7. Estimate the timing and prtin f remvals frm the central and nrthern herds each winter, including the prtin f remvals frm each age and sex class and calf-cw pairs. 8. Dcument bisn use f risk management znes utside the nrthern and western bundaries f Yellwstne and cmmingling with livestck during the likely brucellsis-induced abrtin perid fr bisn each spring. 9. Estimate the effects f hazing r temprarily hlding bisn in capture pens at the bundary f Yellwstne (fr spring release back int the park) n subsequent bisn mvements r pssible habituatin t feeding. Brucellsis Suppressin (Reduce Disease Prevalence) 10. Determine the strength and duratin f the immune respnse in bisn fllwing parenteral (e.g. syringe delivery) vaccinatin fr brucellsis. 11. Determine the strength and duratin f immune respnse in bisn fllwing remte delivery (e.g. bi-bullet) vaccinatin fr brucellsis. 12. Dcument lng-term trends in the prevalence f brucellsis in bisn, and the underpinning effects f remte and/r parenteral vaccinatin, ther risk management actins (e.g., harvest, culling), and prevailing eclgical cnditins (e.g. winter-kill, predatin) n these trends. IMPLEMENTATION AND EXPECTED OUTCOMES T accmplish this suite f mnitring activities, NPS staff wrk with the ther IBMP agencies (Animal and Plant Health Inspectin Service, Cnfederated Salish and Ktenai Tribes, InterTribal Buffal Cuncil, Mntana Department f Livestck, Mntana Department f Fish, Wildlife, and Parks, Nez Perce Tribe, U.S. Frest Service), and ther scientists and stakehlders t implement field, cntrlled, and labratry studies t cllect empirical data fr evaluating prgress. The data are used t develp and parameterize mdels that serve as analytical tls fr evaluating hw bisn and brucellsis may respnd t management actins within specified cnfidence bunds. The IBMP agencies prduce an annual reprt that describes mnitring activities, the status f Yellwstne bisn, and relevant brucellsis management issues. This reprt is disseminated at < The Wildlife Bilgist leading the Bisn Eclgy and Management Prgram at Yellwstne is respnsible fr managing the mnitring system and prducing the mnitring prtin f the annual IBMP reprt. This mnitring plan will als be psted at < and analyses, and as apprpriate, articles resulting frm mnitring will be subject t peer review by ther scientists frm the NPS, agency partners, and/r 3

4 annymus reviewers selected by editrs f scientific jurnals. Pursuant t Bulletin M issued by the Office f Management and Budget n December 16, 2004, the intensity f peer review will be cmmensurate with the significance f the infrmatin being disseminated. Success in adaptive management ultimately depends n effectively linking mnitring and assessment t bjective-driven decisin making (Williams et al. 2007). Thugh different philsphies exist regarding hw adaptive management shuld be implemented, certain characteristics transcend them, including: 1) linkages amng key steps such as identifying bjectives, implementing mnitring, and adjusting management actins based n what is learned; 2) cllabrating with agency partners; and 3) cmmunicating with and engaging key stakehlders (U.S. Gvernment Accuntability Office 2008). This mnitring prgram will prvide timely and useful infrmatin t help develp adaptive management adjustments needed t cnserve Yellwstne bisn, reduce the risk f brucellsis transmissin frm bisn t cattle, and reduce the prevalence f brucellsis in the bisn ppulatin. It will als allw IBMP managers t track system respnses t these management actins thrugh cntinuatin f mnitring. Examples f actins by the NPS that mnitring may trigger based n the infrmatin cllected include: Deciding whether r nt t implement remte vaccinatin based n vaccine efficacy (i.e., stimulatin f cellular immunity) and the develpment f adequate delivery ptins t btain the desired reductins in serprevalence and infectin; Discntinuing vaccinatin in its implemented frm if there is n indicatin f prgress ver a reasnable perid; Implementing cnservatin measures t decrease mrtality and increase the grwth rate f the ppulatin if estimated bisn abundance decreases twards 2,500; Altering culling r harvest strategies if significant and bilgically imprtant effects t age, genetics, herd, and/r sex structure are detected; and Discntinuing brucellsis cntainment r suppressin actins if estimated bisn abundance decreases belw 2,500 and agency partners d nt strictly implement cnservatin measures t abate further reductins in abundance. MONITORING SYSTEM EVALUATION The mnitring prgram will be cnsidered successful if it prvides data t: 1) evaluate prgress twards achieving bjectives; 2) determine resurce status t identify apprpriate management actins and adjust management decisins; 3) increase understanding f resurce dynamics via the cmparisn f predictins against survey data; and 4) enhance and develp mdels f resurce dynamics as needed and apprpriate. The fllwing questins were adapted frm Williams et al. (2007) and will be cnsidered thrughut the duratin f this plan t increase the likelihd f successful mnitring and evaluate prgress in achieving bjectives: Stakehlder Invlvement: Are stakehlders cmmitted t and invlved in the mnitring and assessment prgram? Is the mnitring prcess able t adapt t changes in stakehlder and public viewpints? Objectives: Are the mnitring bjectives achievable and sustainable? Is the mnitring prgram prviding infrmatin that can be used t track prgress in meeting management bjectives and better understand trade-ffs amng bjectives? Management Actins: Des the mnitring plan prvide infrmatin that can be used t assess the ptential effects and effectiveness f the range f feasible management actins (e.g., n actin, hunting, vaccinatin, selective culling, quarantine) and trade-ffs amng them? Is prgress being made twards achieving management bjectives? Has the set f management alternatives r implemented actins been adjusted ver time based n infrmatin btained during mnitring? Mdels: Are the hyptheses underlying the strategies fr resurce management expressed as testable mdels? Have explicit links between management actins and resurce dynamics been incrprated 4

5 int the mdels? Has the mnitring plan cntributed t a better understanding f the eclgical prcesses that drive resurce dynamics? Are the relevant envirnmental factrs incrprated int the mdels? Are the mdels calibrated with available mnitring infrmatin? Mnitring Plan: Des the plan supprt the testing f alternate mdels and measurement f prgress twards accmplishing management bjectives? Des the plan mnitr the metrics necessary t estimate relevant resurce and disease attributes? Have the necessary levels f accuracy been attained? Have cmmitments amng managers, scientists, and ther stakehlders been sustained during the mnitring prgram? Des the plan prvide meaningful and useful data and infrmatin within timeframes that allw fr adaptive decisin making? Decisin Making: Are decisins based n the understanding and status f the resurce derived frm mnitring data? Are decisins being guided by management bjectives and mnitring infrmatin regarding these bjectives? Are stakehlders infrmed and given the pprtunity t cmment befre decisins are made r changed? Fllw-up Mnitring: Are analysis needs understd and being met? Is mnitring cnducted n a timely basis? Is mnitring targeted t system attributes that are useful fr evaluatin and learning? Are mnitring data cllected and managed s they are available and easy t access? Can the mnitring data be used t update measures f mdel cnfidence? Assessment: Have mnitring data been used t evaluate the expected effects f alternate management strategies and update predictins? Have changes in management been implemented when mnitring data indicate management bjectives are r are nt being met? Iteratin: Are management actins and decisins reviewed frequently based n mnitring and assessment infrmatin? Have resurce management alternatives been revisited r mdified ver time? Has uncertainty related t resurce and disease dynamics and the effects f management actins been reduced thrugh mnitring and learning ver time? Are the mnitring bjectives likely t be achieved within specified r reasnable timeframes? Each year thrugh regularly scheduled IBMP meetings, cmpletin f the annual IBMP reprt, and update f this mnitring plan, we slicit review, cmment, and discussin by ur agency partners and key stakehlders in the refinement f bjectives, design f mnitring, and assessment t build supprt fr the legitimate prcess and prvide a fundatin fr learning-based resurce management. Public infrmatin staff share the results f mnitring activities with key stakehlders thrugh timely press releases and web-mails, and reprts and articles will be made available n-line at the website fr the IBMP agencies (< MONITORING FINDINGS The fllwing paragraphs summarize findings f mnitring and research since the plan was initiated in These findings were reprted at IBMP meetings and cnsidered by the IBMP agencies in develping the annual reprts and recmmendatins fr adaptive management adjustments (White et al. 2009, Zaluski et al. 2010, Canfield et al. 2011). Cnservatin (Preserve a Functinal, Free-Ranging Bisn Ppulatin) 1. Estimate the abundance, demgraphy, and limiting factrs fr the verall bisn ppulatin and tw primary subppulatins (i.e., central and nrthern breeding herds). Bisn abundance, age and sex structure, and recruitment are estimated each summer fr the central and nrthern breeding herds. Results are dcumented in an annual cunt reprt that is psted n the website fr the Interagency Bisn Management Plan ( A sample f 45 t 60 radi-cllared bisn is maintained annually t estimate distributin, grup 5

6 sizes, habitat use, mvements, pregnancy, and survival. These findings are released peridically in published articles (see belw). NPS staff cllabrated with clleagues at Mntana State University t estimate demgraphic rates frm 80 adult female bisn in the central herd during (Geremia et al. 2009). Animals testing psitive fr expsure t brucellsis had significantly lwer pregnancy rates acrss all age classes cmpared t sernegative bisn. Birth rates were high and cnsistent fr sernegative animals, but lwer fr yunger, serpsitive bisn. Sernegative bisn that cnverted t serpsitive while pregnant were likely t abrt their 1 st and 2 nd pregnancies. There was a prnunced decrease in survival fr animals >12 years ld. Als, brucellsis expsure indirectly lwered bisn survival because mre bisn were culled ver cncerns abut transmissin t cattle when bisn attempted t mve t lwer-elevatin areas utside the park. There was a significant decrease in adult female survival when the number f bisn in the central herd exceeded 2,000-2,500 animals, which was exacerbated during winters with severe snw pack because mre bisn mved utside the park. Except during , the vast majrity f radi-marked bisn culled at the nrth and west bundaries during came frm the central herd. The effects f brucellsis n survival, pregnancy, and birth rates lwered the grwth rate in the central herd. Ppulatin grwth rates will likely increase by mre than 15% if vaccinatin plans are implemented and successful. NPS staff cllabrated with clleagues at Clrad State University t synthesize available infrmatin and interpreted results f a spatially explicit mdel (Cughenur 2005) f the Yellwstne system (Plumb et al. 2009). Bisn abundance has nt exceeded the theretical fd-limited carrying capacity f 6,200 in Yellwstne. Mre bisn migrate earlier t lwer-elevatin winter ranges as numbers increase and climatic factrs interact with density t limit nutritinal intake and fraging efficiency. A gradual expansin f the winter range as bisn numbers increased enabled relatively cnstant ppulatin grwth and increased fd-limited carrying capacity. Current management actins shuld attempt t preserve bisn migratin t essential winter range areas within and adjacent t the park, while actively preventing dispersal and range expansin t utlying areas via hazing, translcatins, and culls. A ppulatin f 2,500-4,500 bisn shuld satisfy cllective interests cncerning the park s frage base, bisn mvement eclgy, retentin f genetic diversity, brucellsis risk management, and prevailing scial cnditins. NPS staff cntributed t a chapter n cnservatin guidelines fr ppulatin, genetic, and disease management f American bisn fr the IUCN (Gates et al. 2010). Overarching principles fr cnserving bisn were t (1) maximize the number f bisn in a ppulatin (i.e., maximum sustainable rather than a minimum viable ppulatin size) t better retain natural variatin and prvide mre resiliency t surprises r catastrphic events, (2) supprt and prmte wild cnditins and behaviurs in an envirnment where bisn are integral t cmmunity and ecsystem prcesses, expsed t natural selectin, and active management interventins are minimized, (3) preserve genetic integrity and health by maintaining bisn lineages and carefully evaluating all mvements f bisn between ppulatins, and (4) cnducting rutine mnitring and evaluatin f demgraphic prcesses, herd cmpsitin, habitat, and assciated eclgical prcesses that are central t evaluating herd health and management efficacy. 6

7 NPS staff develped a ppulatin mdel using data cllected frm Yellwstne bisn during and estimated the abundance, cmpsitin, and trends f each breeding herd t evaluate the relative impacts f harvests and ther types f management remvals (Geremia et al. 2011a, 2012). Demgraphic estimates were integrated with a mdel f bisn migratin (Geremia et al. (2011b) t predict the numbers f bisn mving t the park bundary each winter. These tls cmbined lng-term mnitring data with infrmatin gained frm radi-cllared bisn t draw cnclusins abut future cnditins f Yellwstne bisn. A decisin-making prcess was develped t advise the management f ppulatin abundance and trans-bundary mvements f bisn. During June and early July, NPS staff cnducted ppulatin cunts and age and gender classificatins f each breeding herd. They then used lng-term weather frecasts and the mdels described abve t predict herd abundances and cmpsitins at the end f the upcming winter, and the magnitude f numbers f bisn migrating t park bundaries. NPS staff established annual remval bjectives fr bisn based n abundance, disease, distributin, and demgraphic (age, herd, sex) gals t reduce bisn numbers twards an endf-winter guideline f 3,000, while prgressing twards equal abundance in each herd and sex ratis f 50% adult males and 50% adult females in each herd. A variety f management tls were cnsidered fr reducing bisn numbers including (1) public and treaty harvests in Mntana, (2) selective culling (shipment t slaughter) at bundary capture facilities t reduce the prprtin f infectius bisn, (3) selective culling (shting, shipment t slaughter) in Mntana t prevent brucellsis transmissin t nearby livestck r due t human safety r prperty damage cncerns, (4) transfer f bisn t American Indian tribes r ther rganizatins fr quarantine and eventual release, and (5) transfer bisn t research facilities. NPS cllabrated with Syracuse University (Dr. Duglas Frank) during t quantify frage prductin and cnsumptin at six study sites acrss the nrthern grasslands in Yellwstne Natinal Park. Five r six grazing exclsures were deplyed at each site. Prductin and percent cnsumptin estimates were made mnthly frm May t September. Data cllectin will cntinue in During the 1980s and 1990s, migratry ungulates n the nrthern grassland f Yellwstne had tight bigechemical linkages with plants and sil micrbes that dubled the rate f net nitrgen mineralizatin, stimulated abvegrund prductin by as much as 43%, and stimulated belwgrund prductivity by 35% (Frank and McNaughtn 1993). These bigechemical linkages were largely driven by high densities f elk that depsited large quantities f nitrgen, phsphrus, and ther nutrients via dung and urine. Hwever, rates f ungulate grazing intensity and grassland nitrgen mineralizatin were reduced by 25-53% by , partially as a result f 60% fewer elk. Since 2002, bisn numbers in nrthern Yellwstne have mre than tripled frm 813 t 2,600. Larger grups f grazing bisn culd ptentially have quite different effects than elk n nutrient redistributin and cycling n nrthern Yellwstne grasslands. This prject shuld help elucidate the influence f recent changes in elk and bisn numbers and distributins n ecsystem prcesses such as the spatial pattern and intensity f ungulate grazing and grassland energy and nutrient dynamics. 2. Describe migratry and dispersal mvements by bisn at a variety f tempral and spatial scales in and utside the park. NPS staff cllabrated with clleagues at Mntana State University t quantify annual variatins in the magnitude and timing f migratin by central herd bisn during and identify ptential factrs driving this variatin (Bruggeman et al. 2009c). 7

8 Bisn frm the central herd were partially migratry, with a prtin f the animals migrating t the lwer-elevatin Madisn headwaters area during winter while sme remained yearrund in r near the Hayden and Pelican valleys. There was significant bisn migratin t the Madisn headwaters area befre the Hayden and Pelican valleys were fully ccupied and abundance apprached the fd-limiting carrying capacity f these valleys. After the central herd exceeded 2,350 animals, hwever, the number f bisn wintering in the Hayden and Pelican valleys appeared t stabilize, while bisn cntinued t migrate t the Madisn headwaters area. Als, mre bisn migrated earlier as density increased. Sme bisn migrated utside the west-central prtin f the park between the summer and winter cunts each year when the central herd exceeded 2,350 bisn, perhaps relcating t nrthern range. The timing and magnitude f bisn migratin were accentuated during years f severe snw pack that limited access t fd. NPS staff cllabrated with clleagues at Mntana State University t quantify hw snw, tpgraphy, habitat attributes, and rads influenced the travel patterns and nn-traveling activities f 30 radi-marked, adult, female bisn frm the central herd during three winters (Bruggeman et al. 2009a, b). Bisn were less likely t use a pint n the landscape fr traveling r feeding as snw pack increased. Hwever, bisn used lcal areas with deeper snw as the verall snw pack increased n the landscape. Distance t stream was the mst influential habitat cvariate, with the spatial travel netwrk f bisn being largely defined by streams cnnecting fraging areas. Distances t fraging areas and streams als significantly influenced nn-traveling activities, being negatively crrelated with the dds f bisn fraging r resting. Tpgraphy significantly affected bisn travel patterns, with the prbability f travel being higher in areas f variable tpgraphy that cnstrained mvements (e.g., canyns). Distance t rad had a significant, negative effect n bisn travel, but was nine times less influential cmpared t the impact f streams. Rad grming has a minimal influence n bisn travel and habitat use given the imprtance f natural dynamic and static landscape characteristics such as snw pack, tpgraphy, and habitat attributes n bisn chice f travel rutes and habitat use fr fraging and resting. NPS staff cllabrated with staff frm Clrad State University t analyze the relatinships between bisn ppulatin size, winter severity, and the number f bisn remved near the bundary f Yellwstne during (Geremia et al. 2011b). Migratin differed at the scale f herds, but a single unifying expnential mdel was useful fr predicting migratins by bth herds. Migratin beynd the nrthern park bundary was affected by herd size, accumulated snw water equivalent, and abvegrund dry bimass. Migratin beynd the western park bundary was less influenced by these predictrs, and mdel predictins since 2006 suggest additinal drivers (e.g., learning) f migratin were nt in the mdel. Simulatins f migratins ver the next decade suggest that a strategy f sliding tlerance where mre bisn are allwed beynd park bundaries during severe climate cnditins may be the nly means f aviding episdic, large-scale reductins t the Yellwstne bisn ppulatin in the freseeable future. 3. Estimate the existing heterzygsity, allelic diversity, and lng-term prbabilities f genetic cnservatin fr the verall bisn ppulatin and identified subppulatins. 8

9 NPS staff cllabrated with clleagues at the University f Mntana t test the hypthesis that bisn frm the central and nrthern breeding herds wuld be genetically differentiated based n mitchndrial and micrsatellite DNA frm fecal samples. Based n mitchndrial DNA analyses, there was significant genetic differentiatin between bisn sampled frm the nrthern and central breeding herds, likely due t strng female fidelity t breeding areas (Gardipee 2007). NPS staff prvided infrmatin t the Department f Interir fr review by scientists frm gvernment agencies and nn-gvernmental rganizatins with prfessinal ppulatin geneticists and the develpment f guidance fr the genetic management f federal bisn ppulatins (Dratch and Ggan 2010). Parks and refuges that currently have bisn ppulatins, with the exceptin f Yellwstne Natinal Park, d nt have enugh land t supprt a ppulatin f mre than 1,000 bisn (i.e., minimum target t preserve genetic variatin ver centuries). Yellwstne bisn have relatively high allelic richness and heterzygsity cmpared t ther ppulatins managed by the Department f Interir. Yellwstne bisn are the nly ppulatin with n mlecular evidence (i.e., micrsatellite markers) r suggestin (i.e., SNPs) f ptential cattle ancestry (i.e., intrgressin f cattle genes). Thus, this ppulatin cnstitutes a genetic resurce that must be prtected frm inadvertent intrgressin. The Yellwstne and Wind Cave bisn ppulatins are genetically unique and the lineages are nt represented elsewhere within ppulatins managed by the Department f Interir. Thus, high pririty shuld be given t replicating these significant lineages via satellite herd establishment (Halbert and Derr 2008). The NPS reviewed a study by Pringle (2011) that cncluded that sme Yellwstne bisn have deleterius genetic mutatins and, as a result, are predicted significantly impaired in aerbic capacity, disrupting highly evlved cld tlerance, winter feeding behavirs, escape frm predatrs and cmpetitin fr breeding." Bisn with hapltype 6 in their mitchndrial genme carry a duble mutatin that affects tw genes: cytchrme b and ATP6. These bisn are primarily fund in the central breeding herd based n recent genetic sampling. This inherited mutatin culd affect their prductin f energy (i.e., ATP prduced by mitchndrial xidative phsphrylatin). Bisn with hapltype 8 in their mitchndrial genme d nt carry the duble mutatin and are primarily fund in the nrthern breeding herd. Even if the genetic sequences and analyses reprted by Pringle (2011) are crrect, genetic mutatin des nt autmatically equal genetic disease. There are multiple cmpensating mechanisms in bilgical systems that cmbine t vercme theretical metablic deficiencies. Als, there is direct evidence that even if Yellwstne bisn have sme srt f genetic deficiency, it has nt been manifested thrugh any bilgically significant effect n their ability t survive. Estimated annual survival rates and birth rates fr adult female bisn were quite high during ; especially given the severe, prlnged, high-elevatin winter cnditins and predatr-rich envirnment in and near Yellwstne Natinal Park. The NPS is taking steps t fllw-up n Dr. Pringle's wrk and recmmendatins. NPS staff cllabrated with clleagues at the University f Mntana and t cnduct a mathematical mdeling assessment that prvided predictive estimates f the prbability f preserving 90 and 95% f the current level f genetic diversity values (bth heterzygsity and allele diversity) in Yellwstne bisn (Pérez-Figuera et al. 2012). 9

10 Findings suggested that variatin in male reprductive success had the strngest influence n the lss f genetic variatin, while the number f alleles per lcus als had a strng influence n the lss f allelic diversity. Fluctuatins in ppulatin size did nt substantially increase the lss f genetic variatin when there were mre than 3,000 bisn in the ppulatin. Cnservatin f 95% f the current level f allelic diversity was likely during the first 100 years under mst scenaris cnsidered in the mdel, including mderate-t-high variatins in male reprductive success, ppulatin sizes greater than 2,000 bisn, and apprximately five alleles per lcus, regardless f whether culling strategies resulted in high r lw fluctuatins in abundance. Hwever, a stable ppulatin abundance f abut 2,000 bisn was nt likely t maintain 95% f initial allele diversity ver 200 years, even with nly mderate variatin in male reprductive success. Rather, maintenance f 95% f allelic diversity is likely t be achieved with a fluctuating ppulatin size that increases t greater than 3,500 bisn and averages arund 3,000 bisn. NPS staff cllabrated with clleagues at University f Mntana t cnduct DNA extractins with fecal samples cllected frm Yellwstne bisn in the nrthern and central breeding herds during 2006 and Mitchndrial DNA analyses revealed tw hapltypes, with higher frequency f hapltype 8 in the nrthern breeding herd, and significant genetic differentiatin amng nrthern and central herds (F ST = ). Micrsatellite analyses revealed allele frequencies with lw levels f subdivisin between the central and nrthern breeding herds (F ST = 0.02 in 2006 and 0.01 in 2008). These results suggest the ppulatin has tw genetically distinguishable breeding grups with strng female philpatry and male-mediated gene flw. Radi-marked adult females prvided evidence f female fidelity, but emigratin between breeding grups was substantial during Staff recmmended lng-term mnitring f micrsatellite allele and mitchndrial hapltype frequencies t track genetic diversity and ppulatin substructure. They expect F ST values t fluctuate as the ppulatin respnds t bisn density in the tw breeding herds, management actins (e.g., culling), and natural selectin. In a study partially funded and supprted by the NPS, Halbert et al. (2012) investigated the ptential fr limited gene flw acrss the Yellwstne bisn ppulatin using bld and hair samples primarily cllected frm bisn at the nrthern and western bundaries f the park during the winter migratin perid, well after the breeding seasn. Tw genetically distinct and clearly defined subppulatins were identified based n bth gentypic diversity and allelic distributins. Genetic cluster assignments were highly crrelated with sampling lcatins fr a subgrup f live capture individuals. Furthermre, a cmparisn f the cluster assignments t the tw principle winter cull sites revealed critical differences in migratin patterns acrss years. The tw Yellwstne subppulatins displayed levels f differentiatin that are nly slightly less than that between ppulatins which have been gegraphically and reprductively islated fr ver 40 years. The authrs suggested that the cntinued practice f culling bisn withut regard t pssible subppulatin structure has the ptentially negative lng-term cnsequences f reducing genetic diversity and permanently changing the genetic cnstitutin within subppulatins and acrss the Yellwstne ppulatin. 1 F ST is the prtin f ttal genetic variance cntained in a subppulatin cmpared t the ttal genetic variance. Values can range frm 0 t 1 and high F ST implies cnsiderable differentiatin amng subppulatins. 10

11 NPS staff (White and Wallen 2012) disputed sme f the assumptins and inferences made by Halbert et al. (2012) and suggested that human manipulatin had created and maintained much f the bserved ppulatin subdivisin and genetic differentiatin. Extensive mnitring f the mvements and prductivity f radi-cllared bisn since 2005, when the ppulatin reached an abundance f apprximately 5,000 bisn, suggests that emigratin and gene flw is nw much higher than suggested by Halbert et al. (2012). Allwing the bisn t migrate and disperse between breeding herds wuld be in the best interest f the bisn ppulatin fr the lng term. The NPS cntinue t allw eclgical prcesses such as natural selectin, migratin, and dispersal t prevail and influence hw ppulatin and genetic substructure is maintained in the future rather than actively managing t perpetuate an artificially created substructure. The existing ppulatin and genetic substructure may be sustained ver time thrugh natural selectin r it may nt. 4. Prmte cperative cnservatin in bisn management by partnering with states, Tribes, and thers interested in bisn health and recvery. During 2005 thrugh 2008, 213 Yellwstne bisn calves that tested negative fr brucellsis expsure were transferred frm the NPS t the Animal Plant Health Inspectin Service and Mntana Fish, Wildlife, and Parks. These bisn were mved t a research quarantine facility nrth f Yellwstne Natinal Park t evaluate if they wuld remain free f brucellsis thrugh at least their first pregnancy and calving. The quarantine feasibility study (2005 thrugh 2011) was successful and the surviving riginal bisn and their ffspring are cnsidered brucellsis-free by the state f Mntana and APHIS. The state f Mntana cmpleted envirnmental cmpliance t relcate 87 f these bisn t the Green Ranch wned by Turner Enterprises Inc. in February 2010 and the remaining 61 bisn t the Frt Peck Indian Reservatin in Mntana in March Pursuant t memranda f understanding, these bisn are underging five additinal years f assurance testing t increase public and scientific cnfidence that the bisn are truly brucellsis-free. In September 2012, there were 190 bisn at the Green Ranch in Mntana, including the surviving riginal bisn and their ffspring. All f these bisn remain the prperty f the state f Mntana until February 2015, at which time Turner Enterprises Inc. will return t Mntana Fish, Wildlife, and Parks all the surviving riginal bisn frm the quarantine feasibility study and 25% f their ffspring. At that time, Turner Enterprises Inc. will gain wnership f the remaining ffspring. In September 2012, there were 72 bisn at the Frt Peck Indian Reservatin. Thirty-ne f these bisn were suppsed t be transferred this summer t the Frt Belknap Indian Reservatin in Mntana. Hwever, a Mntana judge granted an injunctin blcking further relcatin f the bisn. Per agreement, up t 25% f the prgeny f these bisn will be made available t the state f Mntana. In September 2012, the Superintendent f Yellwstne Natinal Park signed an agreement with the InterTribal Buffal Cuncil fr ccasinally transferring sme Yellwstne bisn t them fr transprt t slaughter and subsequent distributin f bisn meat and ther parts t American Indian tribes. During Octber 2012, staff at Yellwstne Natinal Park cnsulted with members f American Indian tribes assciated with Yellwstne Natinal Park during tw cnference phne calls regarding the management f Yellwstne bisn and pssible transfers f bisn t the tribes. 11

12 Staff at Yellwstne Natinal Park wrked with the federal, state, and tribal agencies invlved with the management f Yellwstne bisn t develp a prtcl in Nvember 2012 that describes prcedures and respnsibilities fr future transfers f Yellwstne bisn t (1) apprved slaughter facilities and subsequent cnsumptin, (2) terminal pastures frm which bisn wuld be harvested within 120 days, and (3) peratinal quarantine facilities. In Nvember 2012, staff at Yellwstne Natinal Park develped a prpsal fr the Directr f the NPS t cnsider establishing an peratinal quarantine facility that can eventually hld up t 1,000 bisn and transferring apprximately 250 Yellwstne bisn testing negative fr brucellsis expsure t the facility fr several years. Bisn that successfully cmplete the quarantine requirements wuld be cnsidered brucellsis-free and culd be used fr cnservatin and/r t supprt the culture and nutritin f American Indian tribes. Risk Management (Prevent Brucellsis Transmissin frm Bisn t Livestck) 5. Estimate the prbabilities (i.e., risks) f brucellsis transmissin amng bisn, cattle, and elk, and the elk feed grunds in Wyming and nrthern Yellwstne. NPS staff cllabrated with clleagues at the Agricultural Research Service and University f Mntana t gentype 10 variable number f tandem repeat DNA lci in 58 B. abrtus islates frm bisn, elk, and cattle and test which wildlife species was the likely rigin f recent utbreaks f brucellsis in cattle in the greater Yellwstne area (Beja-Pereira et al. 2009). Findings suggested that islates frm cattle and elk were nearly identical, but highly divergent frm bisn islates. Thus, elk, nt bisn, were the reservir species f rigin fr these cattle infectins. NPS staff cllabrated with clleagues at the U.S. Gelgical Survey and ther agencies and universities t assess several plausible hyptheses fr bserved increases in the serprevalence f brucellsis in several free-ranging elk ppulatins f Wyming (Crss et al. 2009). Free-ranging elk appear t be a maintenance hst fr B. abrtus in sme areas. Brucellsis serprevalence in free-ranging elk increased frm 0-7% in t 8-20% in in fur herd units nt assciated with feed grunds. These serprevalence levels, which are cmparable t units where elk are aggregated n feed grunds, are unlikely t be sustained by dispersal f elk frm feeding areas with high serprevalence r an lder age structure. The rate f serprevalence increase was related t the ppulatin size and density f each herd unit. Enhanced elk-t-elk transmissin in free-ranging ppulatins may be ccurring due t larger winter elk aggregatins. Elk ppulatins inside and utside f the greater Yellwstne area that traditinally did nt maintain brucellsis may nw be at-risk due t recent ppulatin increases. In particular, sme neighbring ppulatins f Mntana elk were 5-9 times larger in 2007 than in the 1970 s with sme aggregatins cmparable t the Wyming feed grund ppulatins. NPS staff cllabrated with clleagues at Clrad State University t develp a Bayesian state space mdel t guide adaptive management f the Yellwstne bisn ppulatin by assimilating frm nging ppulatin mnitring and designed studies f ppulatin prcesses (Hbbs et al. 2009). 12

13 There was 6.8 times mre supprt in the data fr frequency-dependent (i.e., ppulatin expands as bisn numbers increase) than density-dependent brucellsis transmissin (i.e., area used by a ppulatin is fixed). The average cntinuus rate f brucellsis transmissin was 3.9 per year (95% CI = 2.6, 5.3), and the basic reprductive rati (i.e., number f new infectius individuals created by a single infectius individual) averaged 2.5 (95% CI = 1.9, 3.7). Frty percent (95% CI = 27, 55) f yearling and adult females wuld need t be susceptible fr the disease t establish in the ppulatin. Estimatin f ppulatin grwth in the presence f brucellsis λ = 1.06 (95% CI = 1.01, 1.10) reflected the depressing effect f the disease n recruitment. In the absence f brucellsis, the ppulatin was predicted t grw at a rate f λ = 1.11 (95% CI = 1.06, 1.15). Serprevalence f adult females varied between 40 and 60% during the last tw decades, but nly 10% f adult females were infectius. The prbability that a susceptible bisn wuld becme infected via hrizntal transmissin varied annually between 10 and 20%. The prbability f vertical transmissin was 0.22 (95% CI = 0.03, 0.50), but accunted fr fewer than 12% f transmissin events. The annual remval f ser-psitive females was likely t maintain the ppulatin belw 3,500 animals five years int the future and had a high prbability ( ) f reducing ser-prevalence belw 10%. In cntrast, the traditinal practice f episdic remvals f large numbers f untested animals was unlikely t maintain abundance <3,500 bisn five years int the future and wuld fail t reduce serprevalence. Hwever, annual remvals f females created substantial risks f reducing the ppulatin t <2,500 bisn and increasing the adult and yearling male prprtin f the ppulatin t >40%. The NPS reviewed and prvided cmments n a draft f the Kilpatrick et al. (2009) article that used a mdel t integrate epidemilgical and eclgical data t quantify and assess the spatitempral relative risk f transmissin f Brucella frm bisn t cattle utside Yellwstne under different scenaris. The risk f transmissin f brucellsis frm bisn t cattle is likely t be a relatively rare event, even under a n plan (n management f bisn) strategy. The risk f transmissin f brucellsis frm bisn t cattle will increase with increasing bisn numbers and severe snw fall r thawing and freezing events. As the area bisn ccupy utside Yellwstne in the winter is enlarged and verlaps cattle grazing lcatins, the risk f transmissin will increase. Thus, adaptive management measures t minimize risk f transmissin will be mst effective. Risk f transmissin culd be effectively managed with lwer csts, but land use issues and the larger questin f bisn ppulatin management and mvement utside the park might hinder the prspect f slutins that will please all stakehlders. NPS staff estimated the timing and lcatin f parturitin events that may have shed tissues infected by B. abrtus during April t mid-june, (Jnes et al. 2010). Observed abrtins ccurred frm January thrugh 19 May, while peak calving (80% f births) ccurred frm 25 April t 26 May, and calving was finished by 5 June. Observed parturitin events ccurred in the park and n the Hrse Butte peninsula in Mntana, where cattle were nt present at any time f the year. Allwing bisn t ccupy public lands utside the park where cattle are never present (e.g. Hrse Butte peninsula) until mst bisn calving is cmpleted (late May r early June) is nt expected t significantly increase the risk f brucellsis transmissin frm bisn t cattle because: 1) bisn parturitin is essentially cmpleted weeks befre cattle ccupy nearby ranges, 2) female bisn meticulusly cnsume birthing tissues, 3) ultravilet light and heat degrade B. abrtus n tissues, vegetatin and sil, 4) scavengers remve fetuses and 13

14 remaining birth tissues and 5) management maintains separatin between bisn and cattle n nearby ranges. Allwing bisn t ccupy public lands utside the park thrugh their calving seasn will help cnserve bisn migratry behavir and reduce stress n pregnant females and their newbrn calves. The risk f brucellsis transmissin t cattle can still be minimized thugh effective management f bisn distributin. NPS staff cllabrated with clleagues at Mntana State University t analyze cnditins facilitating cntact between bisn and elk n a shared winter range in the Madisn headwaters area f Yellwstne during 1991 thrugh 2006 (Prffitt et al. 2010). Spatial verlap between bisn and elk increased thrugh winter as snw pack increased and peaked when late-term abrtin events and parturitin ccurred fr bisn. Wlves cntributed t immediate, shrt-term respnses by elk that increased spatial verlap with bisn, but lnger-term respnses t wlves resulted in elk distributins that reduced spatial verlap with bisn. Despite this relatively high risk f transmissin, levels f elk expsure t B. abrtus (2-4%) were similar t thse in free-ranging elk ppulatins that d nt cmmingle with bisn (1-3%), suggesting that B. abrtus transmissin frm bisn-t-elk under natural cnditins is rare. Management f brucellsis in elk ppulatins culd fcus n reducing elk-t-elk transmissin risk and, t the extent feasible, curtailing practices that increase elk density and grup sizes during the ptential abrtin perid. NPS staff cllabrated with clleagues at Clrad State University t develp Bayesian mdels t estimate rates f incidence and rutes f transmissin f B. abrtus bacteria amng Yellwstne bisn during and assessed the reprductive csts (C. Geremia, Natinal Park Service, unpublished data). The median prbabilities f hrizntal (frm unrelated bisn) and vertical (frm mther) expsure t calves were 0.10 (95% credible interval = 0.03, 0.22) and 0.10 (0.00, 0.28), respectively; thugh the distributin fr vertical transmissin was skewed left with mst f the prbability clser t zer. Prbabilities that adult bisn were expsed t brucellsis since the preceding parturitin seasn varied frm and snw pack severity exacerbated incidence. We detected a measureable prbability (0.01, 0.12) f bisn recrudescing frm a latent t an infectius state. There was a reprductive cst f diminished birth rates fllwing brucellsis infectin, with nly 59% f serpsitive and recently sercnverting females with calves cmpared t 79% f sernegative females with calves. These results suggest brucellsis is maintained thrugh mixed transmissin mdes and the duratin f infectin may extend beynd the acute phase. NPS and Animal Plant and Health Inspectin Service staff cllabrated with clleagues at the University f Califrnia-Davis n a spatially-explicit assessment f brucellsis transmissin risk amng bisn, elk, and cattle in the nrthern prtin f the greater Yellwstne area (Schumaker et al. 2010). Ppulatin size and winter severity were majr determinants influencing bisn mvements t lwer elevatin winter grazing areas, verlapping with federally-regulated dmestic cattle grazing alltments. Increasing ppulatin size resulted in higher herd densities and increased bacterial shedding. Median ttal risk t cattle frm elk and bisn was 3.6 cattle-expsure event-days (95% P.I ). The estimated percentage f cattle expsure risk frm the Yellwstne bisn herd 14

15 was small ( % f ttal risk) cmpared with elk which cntributed % f the ttal risk. Natural herd migratin and bundary management peratins were imprtant in minimizing the cntributin f bisn t cattle expsure risk, which supprts cntinued bundary management peratins fr separatin between bisn and cattle. Transmissin risks t elk frm elk in ther ppulatins r frm bisn were very small. Minimal pprtunity exists fr B. abrtus transmissin frm bisn t elk under current natural cnditins in the nrthern greater Yellwstne area. Management alternatives that reduce bisn serprevalence are unlikely t substantially reduce transmissin risk frm elk t cattle. Strategies that decrease elk herd densities and grup sizes and reduce elk-t-elk transmissin culd reduce the verall risk t cattle grazing in the nrthern prtin f the greater Yellwstne area. Effrts shuld be taken t reduce the cmingling f cattle and elk, especially during the late gestatin perid fr elk, when spntaneus elk abrtins pse a risk fr interspecies disease transmissin. Bisn vaccinatin did nt meaningfully reduce B. abrtus transmissin risk t cattle. Effective strategies included delaying the turn-n date t cattle grazing alltments, reducing elk serprevalence, reducing the number f cattle at-risk, r prhibiting the cmingling f elk and cattle n individual premises. Staff frm the Mntana Department f Fish, Wildlife, and Parks estimated the persistence f bacteria n fetal tissue, sil, and vegetatin, and scavenging n infectius materials frm birth and abrtin sites near the nrthern and western bundaries f Yellwstne Natinal Park during (Aune et al. 2012). Brucella bacteria can persist n fetal tissues and sil r vegetatin fr days depending n mnth, temperature, and expsure t sunlight. Bacteria purpsely applied t fetal tissues persisted lnger in February than May and did nt survive n tissues beynd 10 June regardless f when they were set ut. Brucella abrtus field strain persisted up t 43 days n sil and vegetatin at naturally cntaminated bisn birth r abrtin sites. Fetuses were scavenged by a variety f birds and mammals in areas near Yellwstne and mre rapidly inside than utside the park bundary. Mdels derived frm the data determined a 0.05% chance f bacterial survival beynd 26 days (95% Credible Interval f days) fr a cntaminatin event in May. The University f Mntana and cllabratrs (including the NPS) examined transmissin f B. abrtus between bisn, elk, and cattle using nine variable-number tandem repeat (VNTR) markers n DNA frm bacterial islates frm 98 tissue samples frm gegraphically-distinct ppulatins f all three hst species in Idah, Mntana, and Wyming (M. O Brien, University f Mntana, unpublished data). Hapltype netwrk assessments f genetic relatedness amng Brucella islates suggested substantial interspecific transmissin between elk and bisn ppulatins in bth Wyming and Mntana. Brucella gentypes frm the 2008 cattle utbreak in Wyming matched elk Brucella gentypes cnfirming elk as the likely surce. Hwever, Brucella frm the tw recent utbreaks (2008, 2010) in Mntana cattle had gentypes similar t bth bisn and elk. Because wild bisn have been excluded frm these cattle areas, this finding suggests transmissin ccurred between bisn and elk in Yellwstne in the past, befre eventually being transmitted amng elk and by elk in the Paradise Valley t cattle. Identical Brucella gentypes amng many elk ppulatins in Mntana suggests that brucellsis may have becme established in Mntana thrugh intraspecific transmissin 15

16 amng ppulatins, withut all infected elk riginating as immigrants frm Wyming r by transmissin frm Yellwstne bisn. 6. Estimate age-specific rates f bisn testing serpsitive and sernegative fr brucellsis that are als culture psitive and the prtin f serpsitive bisn that react psitively n serlgic tests due t expsure t crss-reactive agents ther than B. abrtus (e.g., Yersinia). NPS staff cllabrated with clleagues at the University f Mntana t investigate if Y. enterclitica sertype O:9 caused false-psitive reactins in brucellsis serlgical tests fr bisn using culturing techniques and multiplex PCR (See et al. 2012). Y. enterclitica was nt detected in samples f feces cllected frm 53 Yellwstne bisn culled frm the ppulatin and 113 free-raming bisn frm thrughut the greater Yellwstne ecsystem. These findings suggest Y. enterclitica O:9 crss-reactivity with B. abrtus antigens is unlikely t cause false psitive serlgy tests in bisn, and that Y. enterclitica prevalence is lw in these bisn. NPS and Animal and Plant Health Inspectin Service staff sampled mre than 400 bisn that were cnsigned t slaughter during winter and cllected bld and tissues t estimate the prprtin f serpsitive and sernegative bisn that were actively infected with B. abrtus (i.e., culture psitive; Treanr et al. 2011). Remving brucellsis-infected bisn is expected t reduce the level f ppulatin infectin, but test and slaughter practices may instead be remving mainly recvered bisn. Recvered animals culd prvide prtectin t the verall ppulatin thrugh the effect f herd immunity, thereby reducing the spread f disease. Identifying recvered bisn is difficult because serlgic tests (i.e., bld tests) detect the presence f antibdies, indicating expsure, but cannt distinguish active frm inactive infectin. Age-specific serlgy and B. abrtus culture results frm slaughtered bisn were integrated t estimate prbabilities f active brucellsis infectin using a Bayesian framewrk. Infectin prbabilities were assciated with age in yung bisn (0-5 years ld) and with elevated antibdy levels in lder bisn (>5 years ld). Results indicate that Yellwstne bisn acquire B. abrtus infectin early in life but typically they recver as they grw lder. A tl was develped t allw bisn management t better reflect the prbability that particular animals are infective, with the aim f cnserving Yellwstne bisn while reducing the risk f brucellsis transmissin t cattle. Flurescent plarizatin assay (FPA) values were higher in serpsitive bisn that were culture psitive cmpared t serpsitive bisn that were culture negative, supprting that active infectin is assciated with increased antibdy prductin. The tw cvariates (age and FPA) have management applicatin t identify the prbability f active infectin within specified credible intervals. This wuld allw fr remving bisn that mst likely cntribute t brucellsis maintenance in the ppulatin, while keeping bisn that cntribute t herd immunity which reduces brucellsis transmissin. Estimatin f true infectin prbabilities can replace culling practices (such as the slaughter f all serpsitive individuals) that cnflict with bisn cnservatin. Cmbining selective remval f infectius bisn with additinal management practices, such as vaccinatin, has the ptential t advance an effective brucellsis reductin prgram. 7. Estimate the timing and prtin f remvals frm the central and nrthern herds each winter, including the prtin f remvals frm each age and sex class and calf-cw pairs. 16