EFFECTS OF FIRE AND REHABILITATION SEEDING ON SAGE GROUSE HABITAT IN THE PINYON-JUNIPER ZONE

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1 EFFECTS OF FIRE AND REHABILITATION SEEDING ON SAGE GROUSE HABITAT IN THE PINYON-JUNIPER ZONE An Extension of the Gret Bsin Pinyon-Juniper Demonstrtion Are Finl Report to the Joint Fire Science Progrm Project #B-3-3- Principl Investigtors - Jenne C. Chmbers, nd Robin J. Tusch USDA Forest Service, Rocky Mountin Reserch Sttion Robert R. Blnk USDA Agriculturl Reserch Sttion Eric Fleishmn Stnford University My 5, 26

2 EXECUTIVE SUMMARY Overview Throughout much of the western United Sttes, expnsion of pinyon nd juniper species into sgebrush ecosystems is ltering the structure nd composition of sgebrush communities nd resulting in incresed fire frequency, severity, nd size. Incresed tree densities in combintion with ltered fire regimes re hving negtive effects on both sgebrush communities nd sgebrush obligtes, including sge grouse, species under considertion for listing s thretened or endngered. Mintining nd restoring hbitt of sgebrush obligte species is becoming mngement priority, nd both locl nd regionl fire mngement plns need to consider the effects of proposed ctions, including prescribed fire, on the hbitt of sgebrush obligte bird species. This study used the existing Gret Bsin Demonstrtion Project (#-2-5) to exmine the effects of prescribed fire on the hbitts nd bird species within the pinyon-juniper zone. It ddressed the following informtion needs: () The effects of fire on soil erosion nd vegettion composition nd structure of sgebrush communities over the elevtionl nd tree density grdient tht typifies the pinyon-juniper zone in the Gret Bsin. (2) The effects of rehbilittion seeding with mixtures dominted by introduced vs. ntive species on vegettion composition nd structure of sgebrush communities over the elevtionl nd tree density grdient tht typifies the pinyon-juniper zone in the Gret Bsin. (3) The effects of fire on the survivl, seedling estblishment, nd nutritionl vlue of plnt species used by sge grouse within the sgebrush communities typiclly found in the pinyon-juniper zone of the Gret Bsin. (4) The presence, bundnce, nd loction of sgebrush obligte bird species over the tree density nd elevtionl grdient tht typifies the pinyon-juniper zone within the study dringes, nd chnges tht occur in ptterns of use following fire. This effort utilized n integrted, collbortive project of the Joint Fire Sciences Progrm to ddress ech of these informtion needs. The project durtion ws from My 5, 22 to My 5, 26. A third project ws funded through the JFSP in 25 on Response of Birds, Butterflies nd their Hbitts to Mngement of Wildlnd Fuels nd Fire Regimes tht will further extend this reserch. Deliverbles The project deliverbles s outlined in the initil proposl re provided below, nd re used s frmework for orgnizing the report. Primry products re listed in text nd complete list of products is found t the end of the Report. A demonstrtion re tht cn serve s n exmple of using burn mosics to improve sge grouse hbitt. Guidelines for evluting the effects of fire on sgebrush community nd soil response to prescribed burns over the rnge of tree densities nd elevtionl grdients tht chrcterize pinyon-juniper woodlnds. Guidelines for evluting the effects of rehbilittion seeding on sgebrush community nd soil response to prescribed burns over the rnge of tree densities nd elevtionl grdients tht chrcterize pinyon-juniper woodlnds. Informtion on the effects of fire on the survivl, estblishment, nd nutrient qulity of plnt species used by sge grouse. 2

3 Informtion on the presence, bundnce, nd loction of sgebrush obligte bird species within the study dringes, nd on chnges in use ptterns following burning. A long-term demonstrtion re hs been developed to study the effects of spring-prescribed fire on pinyon-juniper woodlnds nd their ssocited sgebrush ecosystems. Informtion on the effects of spring prescribed fire on soil infiltrtion over the elevtionl nd tree density grdient within the wtershed ws included in the Finl Report for #-2-5, nd the summry publictions re included herein (Ru 25; Ru et l. 25). Chnges in vegettion composition over the sme grdients were evluted for the first three yers fter the fire (22, 23, nd 24) (Dhemers et l. 26; Dhemers nd Chmbers in preprtion) nd will be monitored gin in 26. Becuse of low initil estblishment of plnt species used in the rehbilittion seeding, there were few detectble effects of the seeding nd these dt re not included. The plots will be reevluted in 26, nd it is nticipted tht this informtion will be included in the next report. Detiled nlyses were conducted of the soil physicl nd chemicl response of mid elevtion sites within the Demonstrtion Are (Ru et l. 25). Long-term dt (5 yers) hve been collected on these sites on the responses of two soil nutrients criticl for plnt growth (nitrogen nd phosphorus) (Ru et l. in process). The informtion on the soil responses on these sites hve been relted to chnges in plnt nutritionl vlue fter fire with focus on species used by sge grouse (Ru et l. 25; Ru et l. in process). In ddition, informtion on the estblishment of plnt species used by sge grouse nd other sgebrush obligte species on both burned nd control sites hs been obtined (Bord et l. in preprtion.) Dt on the presence, bundnce, nd loction of sgebrush obligte bird species were collected throughout the wtershed nd n djcent control one yer prior to the burn nd every yer since the burn. These dt hve been used to exmine ptterns nd potentil degree of nestedness; elevtionl influences on species richness; effects of sptil scle on species composition; ptterns of sptil utocorreltion of ssemblges of birds, floristics, physiognomy, nd primry productivity; nd remote sensing vribles s predictors of species richness of birds (e.g., Fleishmn et l. 22, Seto et l. 24, Fleishmn nd Mc Nlly 26). Complimentry reserch tht hs been conducted on the Demonstrtion Are includes n ssessment of the effects of fire nd tree density on nt popultions over the elevtion grdient within the wtershed (Mont Blnc 25; Mont Blnc et l. submitted). In ddition, reserch hs been conducted on llometric methods for simulting plot biomss nd fuel lods for single lef pinyon nd techniques for quntifying litter biomss (Tusch in review). This reserch is listed in the products section, but is not reviewed in this document. The demonstrtion re hs been used for illustrting the ongoing chnges in centrl Gret Bsin pinyon-juniper woodlnds nd their ssocited sgebrush ecosystems to reserchers, mngers, NGOs, tribes nd privte individuls. It lso hs been used to show the effects of prescribed fire on these ecosystems. This hs been ccomplished through field tours from 2 through 25 nd personl tours led by the PIs nd their students. Additionlly, study results hve been presented t regionl, ntionl, nd interntionl meetings. The informtion obtined from the Demonstrtion Are is being incorported into guidelines for the mngement of these ecosystems. 3

4 The Demonstrtion Are The Demonstrtion Wtershed ws estblished s collbortive effort between the USDA Forest Service, Rocky Mountin Reserch Sttion nd Humboldt-Toiybe Ntionl Forest, nd the Nevd Bureu of Lnd Mngement. It is locted in Underdown Cnyon (39.5 N W) in the Shoshone Mountin Rnge on the Humboldt-Toiybe Ntionl Forest (Austin Rnger District) in Nye nd Lnder Counties, Nevd. Riley Cnyon, neighboring wtershed, serves s control. The geology of the wtersheds consists of welded nd non-welded silic sh flow tuff, nd the soils developed on the lluvil fns used for the reserch burns re clssified s Corse lomy mixed frigid Typic Hploxerolls. Averge nnul precipittion rnges from 23 cm t the bottom to 5 cm t the top of the dringe nd rrives mostly s winter snow nd spring rins. Both wtersheds hve strems which re ephemerl during most yers. OR ID NV UT CA UNDERDOWN CANYON The originl study ws developed to exmine both wtershed-scle nd stnd/community-scle effects of the burn tretment. Studies of the effects of the burns on strem chnnels nd wter qulity nd lso on butterfly tx were designed to be exmined t the wtershed scle. Studies of fuel lods nd of vegettion nd soil responses were designed to be evluted t the community scle. For the community-scle studies, three pirs of study sites (n=6) were locted on north fcing lluvil fns in June 2 t elevtions of 273, 23, 295, nd 2225; two sites were t 2347m. The fns rnge from two to six hectres in size. One of ech pir of fns received the burn tretment, while the other served s control. To evlute the effects of burning over the elevtionl grdient, three mcroplots (.h 2 ) with intermedite tree densities were locted on ll burn nd control sites. To exmine the effects of burning on plots with different tree cover nd fuel lods, three high tree cover nd three low tree cover mcroplots lso were locted t the mid-elevtion sites (295 nd 2225m). Tree covers verged 2% for low cover plots, 38% for intermedite cover plots, nd 78% for high tree cover plots. The community-scle burns were conducted in spring 22 (My -4) by intergency fire crews. Despite firly moist conditions t the time of the burn, every ttempt ws mde to obtin uniform burn conditions. The community-scle burns were designed to be prt of lrger wtershed- 4

5 scle burns. However, due to restrictive burn window nd concerns over potentil escpe, only the smller-scle burns were conducted in 22. Becuse of high winds nd n inbility to obtin the necessry conditions for the burns, the wtershed-scle burns were gin postponed in 23. During the third try in 24, n dditionl 9 cres were treted resulting in mosic of burned nd nonburned conditions within the wtershed. Soil Responses to the Prescribed Burn The effects of the prescribed fire on the responses of nitrogen nd phosphorus were exmined t the mid elevtion burn nd control plots immeditely before nd fter the spring burn, the fll prior to the burn, nd for 4 flls fter the smller-scle burns. This informtion hs been published in Mster s thesis (Ru 25) nd mnuscript is in process (Ru et l. in process). Specificlly, the effects of prescribed fire, vegettion type, nd time following fire on soil KCl extrctble nitrogen nd NHCO 3 extrctble phosphorus were evluted. Nitrogen nd phosphorus sttus of soils severl yers following fire cn drmticlly influence the site s recovery potentil. We found tht KCL extrctble NH 4 + incresed immeditely following prescribed fire especilly in sgebrush microsites. Extrctble NO 3 - decresed immeditely fter the burn with the lrgest losses possibly occurring under pinyon pines. However, totl extrctble inorgnic-n incresed following the fire especilly under sgebrush indicting tht NH 4 + increses out weigh NO 3 - decreses nd dominte vilble N pools immeditely following fire. Sodium bicrbonte extrctble ortho-p incresed immeditely following fire. Figure. Mens nd stndrd errors for pre nd post burn soil nutrients on the tretment site t two depths (-3 nd -8 cm) nd three microsites (interspce, under shrub, nd under tree). In the four yers following the fire microsite differences becme less obvious but lndscpe scle trends continued. In the first 8 cm of the soil profile extrctble NH 4 + remined elevted through 23 over the control with similr trend through 25. Extrctble NO 3 - nd totl extrctble inorgnic-n remined elevted over the control through 24 with the trend continuing through 5

6 25 indicting tht NO 3 - domintes longer term N vilbility ner the soil surfce. Fire incresed extrctble ortho-p on the burn site ner the soil surfce through 23, nd n obvious temporl increse occurred on both sites in 23. Figure 2. Mens nd stndrd errors for pre nd post tretment ner surfce soil nutrients on the control nd tretment sites. Mens not represented by common letter re significntly different. Burn effects on NH 4 + were not noticeble over longer time periods through 52 cm of the soil profile. However, NO 3 - nd totl extrctble inorgnic-n remined elevted over the control through 22 nd the trend continued into 23 to 52 cm. There ws lso n obvious temporl pek in NO 3 - nd totl inorgnic-n in 24 for both sites to 52 cm. No fire effects were observed for extrctble ortho-p to 52 cm, but the temporl pek ppers on both sites in 23. Figure 3. Mens nd stndrd errors for pre nd post tretment soil nutrients through the profile on the control nd tretment sites. Mens not represented by common letter re significntly different. 6

7 These results indicte tht pinyon-juniper woodlnds nd their ssocited sgebrush semi-dessert communities re highly vrible in regrds to the sptil nd temporl distribution of minerl nutrients. As woodlnds expnd into semi-rid shrub communities we my see homogeniztion of soil nutrients cross the lndscpe s tree crown cover becomes more continuous, nd shift in distribution of nutrients to the soil surfce ssocited with litter fll. Incresing tree cover will lso increse the proportion of nutrients stored in bove ground biomss. These effects could prove to be deleterious if ssocited with high intensity wildfire, becuse losses from voltiliztion, convection, nd erosion could increse. Soil vilble nutrients in woodlnds chnge from one yer to the next due to climtic vribility; this complictes the nlysis of fire effects, nd emphsizes the need for representtive controls. Prescribed burning in woodlnds cn cuse immedite nd persistent effects on soil vilble nutrients. Becuse fuel nd soil moisture is high during spring prescribed burning, soil heting only occurs to shllow depths. The results re positive chnges in soil vilble nutrients. Burn effects cn be limited to surfce soils nd specific microsites, or they cn occur t soil depths of 52 cm cross the entire lndscpe. We hve documented tht these effects cn persist up to four yers following prescribed burn. This my prove beneficil to vegettion recovering on the burn site. From soil nutrient stndpoint prescribed burning in our demonstrtion woodlnd cn be successful tretment for restoring sgebrush semi-desert ecosystems. However, for extrpoltion to other sites it is importnt to consider the pre-existing understory vegettion conditions before burning, due to its influence on soil stbility nd site recovery post-burn. Plnt Community Response to the Prescribed Burn The effects of elevtion nd tree cover on the vegettion recovery of the sites following the prescribed burns were exmined in 22 nd 24. In ddition, the chnge in the cover of forb species used by sge grouse s result of the burns ws evluted. These dt re being summrized in Mster s thesis (Dhemers 26) nd in mnuscript tht is in preprtion (Dhemers et l. in process). Totl herbceous cover incresed over the elevtionl grdient within the wtershed regrdless of tretment (Figure 4). Herbceous cover incresed on ll burn sites between 23 nd 24 while decreses were seen on the low elevtion control plots. Totl herbceous eril cover on burn sites exceeded tht of their pired control plots in 24 t ll elevtions. Burning resulted in n increse in nnul forb cover. However. similr increses occurred t ll three elevtions. A significnt increse in nnul grss ws seen on burn plots, which is importnt becuse the only species in this functionl group is Bromus tectorum L. However, this species still mde up less thn one percent of the overll cover on ny site regrdless of tretment. Perennil forb cover incresed with elevtion independent of tretment. Also, burned plots hd greter cover thn their pired control plots t ll elevtions. Similrly, the nlysis for perennil grminoids showed significnt effects of burning nd elevtion. Perennil grminoid eril cover ws similr t the low nd intermedite elevtions while the high elevtion sites showed much higher cover of this functionl group for both burn nd control plots in both yers. 7

8 Percent Cover Percent Cover Percent Cover Percent Cover Percent Cover Totl Herbceous Annul Forbs Annul Grminoids Perennil Forbs Perennil Grminoids Burn Control Burn Control Burn Control 23 m 2225 m 2347 m () (b) (c) (d) (e) Figure 4. Aeril herbceous cover for burn nd control plots over three elevtions in 23 nd 24. Vlues re mens + stndrd error. There ws n overll effect of tree cover on totl herbceous cover nd burn effect (Figure 5). As tree cover incresed, totl men herbceous eril cover decresed on both burn nd control plots. However, cover ws higher on burn plots thn their pired control plots in 24. Annul grminoid cover decresed s tree cover incresed. Annul grminoids showed significnt increse on burn sites between 23 nd 24, with the highest cover occurring on the low tree cover sites in 24. Annul forb cover ws significntly higher on burn sites thn control sites, nd burn sites showed drmtic increse between 23 nd 24 in nnul forb cover. Perennil forbs showed similr ptterns to nnul grminoids, decresing s tree cover incresed. Perennil forbs recovered on burn sites to vlues equl to or greter thn tht of their pired control sites by 24, showing strong burn effect. Burning nd tree cover showed significnt effects individully nd lso showed significnt interction for perennil forbs. Control plots showed significnt decrese in perennil forbs s tree cover incresed. Perennil grminoids lso decresed s tree cover incresed. However, they seemed to decrese t lower rte thn other functionl groups. There ws significnt decrese in eril cover for control plots s tree cover incresed. Cover vlues for 24 burn sites exceeded the cover on their pired control 8

9 sites in ll cses for perennil grminoids, but no significnt effect of tree cover or significnt differences between burn plots were seen. Percent Cover Percent Cover Percent Cover Percent Cover Percent Cover Totl Herbceous Annul Forbs Annul Grminoids Perennil Forbs Perennil Grminoids Burn Control Burn Burn Low Intermedite Control High Control () (b) (c) (d) (e) Figure 5. Aeril herbceous cover for burn nd control plots over tree cover grdient in 23 nd 24. Vlues re mens + stndrd error. The potentil sge grouse diet on our sites ws clculted s the sum of the cover vlues for the forbs tht sge grouse utilize for their spring/ summer diet. These species include Eriogonum spp., Astrglus spp., Crepis ccumint, Agoseris gluc, Collinsi prviflor, Phlox grcilis, Antennri dimorph, Antennri rose, nd Phlox longifoli. Elevtion significntly ffected the cover of these species (Figure 6). Aeril cover ws highest t the intermedite elevtion regrdless of tretment. Burning hd n overll effect with low nd intermedite elevtions showing greter cover of these importnt forbs t burn sites thn their pired control sites. Tree cover hd significnt effect on grouse food sources, with percent eril cover decresing s tree cover incresed regrdless of tretment. Cover for burn plots in 24 exceeded tht of their pired control plots t ll tree cover clss, suggesting positive effect of burning on these sites. 9

10 4 Elevtion Comprison Men Percent Aeril Cover Burn Control Burn Control Burn Control 23 m 2225 m 2347 m Tree Cover Comprison Men Percent Aeril Cover Burn Control Burn Control Burn Control Low Intermedite High Figure 6. Aeril cover of forbs utilized by sge grouse s food. The top grph shows the elevtion grdient nlysis while the bottom grph shows the nlysis for tree cover clsses. Vlues re men + stndrd error. Results of this study hve fcilitted the development of stte-nd-trnsition model tht cn be used for the mngement of fuels nd fire in these ecosystems (Figure 7). Where tree overstory is t reltively low to intermedite cover, nturl or prescribed fire typiclly results in mortlity of both the trees nd non-root sprouting shrubs nd cretes suitble conditions for increses in understory herbceous biomss. Ares with ntive herbceous perennils in the understory usully return to ntive grss nd forb communities. In the bsence of fire, sgebrush increses over time, especilly where overgrzing decreses competition from grsses nd forbs. Repeted fire nd proper grzing will mintin successionl mosic dominted by vrying proportions of shrubs nd herbceous species. Pinyon nd juniper estblishment is fcilitted by shrubs, nd given tree seed source nd the bsence of fire the trees expnd into shrub dominted communities. In the bsence of fire, tree cover increses nd there is progressive depletion of understory shrubs nd herbceous species. If biotic threshold hs been crossed, fire cn shift community dominnce to ntive nnul grsses nd forbs, which my or my not recover to sgebrush community depending on the presence of residul perennil ntives nd seed source. The presence of invsive nnul grsses cn result in lterntive vegettion communities, grss-fire cycle, nd progressive degrdtion of the site. More recently, exotic perennil forbs re invding these communities with unknown ecologicl consequences. Mintining

11 sustinble sgebrush ecosystems requires n understnding of community recovery potentil t different stges of tree expnsion. Low-Intermedite Pinyon-Juniper Overstory/Sge/Grss Understory Tree Seed Source Fire Fire Proper Grzing Sgebrush/Grss Community No Fire Overgrzing Grss/Forb Community No Disturbnce Intermedite-High Pinyon-Juniper Overstory nd/or Depleted Understory Fire Fire/ Seeds Ntive Annuls/Residul Grss/Forb Community Fire/ Seeds Invsive Annul Grss/ Perennil Forb Community Seeds Sgebrush/ Grss/Forb Community? Fire Figure 7. A stte nd trnsition digrm for pinyon nd juniper ecosystems. Fire Effects on Plnt Nutritionl Vlue Pinyon nd juniper expnsion into sgebrush-grsslnds hs been shown to be detrimentl to understory vegettion nd sgebrush ecosystems. We exmined the effects of spring prescribed fire on understory boveground biomss nd tissue nutrient concentrtions of grss nd forb species tht occur in sgebrush ecosystems. This study ws done in conjunction with the reserch on soil nutritionl response to fire described bove. The dt re reported in Mster s thesis (Ru 25) nd mnuscript is in process (Ru et l. submitted). Six ntive understory plnt species (Crepis cumint, Eriogonum umbelltum, Eriogonum eltum, Po secund secund, Festuc idhoensis, nd Lupinus rgenteus) importnt for ntive sgebrush obligte forgers were chosen to represent the understory plnt community. L. rgenteus is lso importnt for system nutrient cycling nd nitrogen fixtion. Plnts were collected from three microsites (under tree cnopy, under shrub cnopy, nd interspce) common in trnsitionl woodlnds the summer before spring prescribed burn nd ech of two summers following the burn. Soils were lso collected from corresponding loctions t two depths (-8 nd -52 cm) to determine burning x soil x plnt x microsite interctions. Microsite hd n impct on soil nutrients but ws typiclly not reflected by plnt tissue concentrtions with the exception of F. idhoensis. Burning cused increses in soil surfce nd profile ortho-p nd inorgnic-n in both yers following fire (Figure 8). All species studied hd distinct nutritionl chrcteristics nd independent responses to burning. Burning did not ffect boveground plnt biomss or nutrient concentrtions in the first yer with the exception of F. idhoensis which hd incresed tissue P. By the second yer ll species except C. cumint hd sttisticlly significnt responses to

12 burning. The most common response ws for incresed boveground plnt weight nd tissue N concentrtions, e.g., the responses of Eriogonum umbelltum in Figure 9. Plnt response to burning ppered to be best linked to the burn tretment nd the soil vrible profile ortho-p. Figure 8. Men extrctble soil nutrient concentrtions (ug g - ) nd stndrd errors in 2, 22, nd 23 for the soil surfce (-8cm) nd soil profile (-52cm) t the control (295 m) nd burned (2225 m) loctions. P-vlues of mens comprisons for control vs. burn sites in yer vs. 2, vs. 3, nd overll re indicted on ech grph. 2

13 The percent cover of the species studied either ws not ffected or incresed following fire. Incresed cover coupled with incresed understory plnt biomss nd tissue nutrient concentrtions should increse bove ground pools of minerl nutrients, nd produce sites more fvorble to ntive forgers including sge grouse. Fll burning in these woodlnds my not hve the sme beneficil effects due to decresed fuel moisture nd incresed fire severity. Figure 9. Men bove-ground plnt weight (g), tissue nutrient concentrtions (ug g - ), nd stndrd errors for Eriogonum umbelltum in 2, 22, nd 23 on the control (295 m) nd burned (2225 m) loctions. P-vlues of mens comprisons for control vs. burn sites in yer vs. 2, vs. 3, nd overll re indicted on ech grph. 3

14 Fire Effects on Plnt Estblishment We exmined the effects of spring prescribed fire on seedling emergence nd survivl of ntive plnt species tht occur in sgebrush ecosystems nd tht re importnt for ntive sgebrush obligte forgers. This study ws complimentry to the reserch described bove on soil nd plnt nutrient reltions following spring prescribed fire. A mnuscript is in process (Bord et l. in preprtion). One shrub (Artemisi tridentt vseyn), two forbs (Eriogonum umbelltum nd Lupinus rgenteus) nd two grsses (Po secund secund) nd Festuc idhoensis) were selected from the understory plnt community. Plnts were seeded into three microsites (under tree cnopy, under shrub cnopy, nd interspce) on the mid-elevtion control nd burn plots. Seeding ws conducted on the control plot the fll before the spring prescribed burn (2) nd the fll fter the burn (22). Becuse of the dely in the smll-scle burns, seeding ws conducted on the burn plot only the fll fter the burn (22). Seedling emergence nd survivl ws monitored on the control plot in summer (Figures & b). Emergence Survivl P. secund F. idhoensis Ar. tridentt ssp. vseyn Proportion Proportion Proportion b b b 5 b b b bc bc b b b bc bc 7 c 6 bc b 8 9 c 4 2 b 29 c b b b 9 6 b b b bc c c 5 8 b 9 3 c 7 bc c IS US UT IS US UT IS US UT IS US UT IS US UT IS US UT Control Control Burn Control Control Burn c c 5 Figure. Proportionl emergence nd survivl of A. tridentt, F. idhoensis, nd P. secund on control plots in 22 nd 23 nd burn plots in 23 in interspce, undershrub nd undertree microsites. The numbers of individuls tht emerged nd tht were live t the end of the study re indicted bove the brs. Unlike letters indicte significnt differences mong yers, tretments nd microsites. 4

15 Emergence Survivl L. rgenteus E. umbelltum Proportion Proportion b b b 33 bc 5 bc bc bc bc 22 b c b b 2 3 b 2 b IS US UT IS US UT IS US UT IS US UT IS US UT IS US UT Control 22 Control 23 Burn 23 Control 22 Control 23 Burn 23 Figure b. Proportionl emergence nd survivl of E. umbelltum nd L. rtgenteus on control plots in 22 nd 23 nd burn plots in 23 in interspce, undershrub nd undertree microsites. The numbers of individuls tht emerged nd tht were live t the end of the study re indicted bove the brs. Unlike letters indicte significnt differences mong yers, tretments nd microsites. Emergence of the shrub, A. tridentt vseyn, ws generlly low (Figure.). In 22, none of the Artemisi seedlings tht emerged survived. In 23, A. tridentt vseyn seedlings tht emerged on the unburned sites under shrubs nd on the burned site under trees did not survive. There were no other tretment differences in seedling survivl. The two grsses hd similr emergence nd survivl ptterns (Figure.). In 22 there were no differences in emergence mong microsites. In 23, reltively dry yer, overll emergence ws lower nd the under tree microsite hd significntly higher seedling emergence thn the other microsites. Both grss species hd the highest survivl under trees on the unburned site in both yers. Survivl ws higher in 23 thn 22 for both grsses. On the burned site, neither grss survived in undershrub loctions nd survivl in under shrub loctions ws nil. The forbs exhibited different emergence nd survivl ptterns thn the grsses (Figure b.). Eriogonum umbelltum hd higher emergence thn L. rgenteus, but exhibited no difference in emergence mong yers, tretments or microsites. Lupinus rgenteus hd higher emergence in 23 in interspces on the unburned site, nd interspces hd higher emergence thn undertree res on the burned site. Eriogonum umbelltum hd higher survivl thn L. rgenteus, but neither forb showed ny differences mong yers, tretments or microsites. Seedling estblishment in semi-rid ecosystems is typiclly low nd vries mong species. Estblishment of Artemisi species is episodic nd, similr to other studies, ws very low on the Demonstrtion re. Seedling estblishment of the grss species ws higher thn tht of Artemisi, but survivl ws still reltively low on the burn re. Overll emergence ws high for both forb species nd survivl, especilly of E. umbelltum, ws reltively high on the burn plots. Forb species re n importnt component of the diets of sgebrush obligte species, nd 5

16 estblishment of these species indictes tht they could be seeded following prescribed fire to increse their composition on burned sites. Fire Effects on Birds From 2 to 25, dt were collected on the distribution nd bundnce of breeding birds from totl of 298 points in the Shoshone, Toiybe, nd Toquim rnges (Humboldt Toiybe Ntionl Forest, Lnder nd Nye Counties, Nevd). At those points, we lso collected dt on vegettion structure, plnt composition, nd other ttributes of bird hbitt. Most of the points were locted in pinyon-juniper or riprin woodlnd. Fourteen points were locted in the Underdown Cnyon Demonstrtion Wtershed nd ten were locted in Riley Cnyon, n djcent control wtershed. Eleven points (six treted, five control) were estblished in Wll Cnyon (Toiybe Rnge), in which wildfire occurred in high-density pinyon-juniper woodlnd in 2. We recorded totl of 88 species of birds from the project tsk re from 2 to 25. One species, Northern Goshwk, is identified s Sensitive by the U.S. Forest Service. An dditionl 23 species re priorities for Nevd Prtners in Flight. The totl re treted with prescribed fire in the Underdown Cnyon Demonstrtion Wtershed prior to 24 (~2 h) ws insufficient to ssess relibly the response of birds nd their hbitts to wtershed-scle burns. Multiple-yer surveys in the Demonstrtion Wtershed nd beyond, however, fforded us vluble opportunity to exmine deterministic nd stochstic influences on species richness nd composition of birds. In ddition, surveys provided insights to monitoring pproches likely to provide relible informtion bout biologicl responses to n rry of environmentl chnges, including but not limited to fuels mngement. The following is summry of some of the mjor inferences nd conclusions bsed on this work. Ptterns nd potentil mechnisms of nestedness (Fleishmn et l. 22). Nestedness nlyses test the degree to which species present in reltively species-poor loctions re proper subsets of species present in reltively species-rich loctions. One cn predict the sequence in which species will be extirpted from or colonize set of terrestril or qutic islnds occupied by nested ssemblge; the ccurcy of these predictions is positively correlted with the degree of nestedness, nd does not depend on knowledge of underlying mechnisms. Assemblges cn be nested by multiple phenomen, including but not limited to species-specific probbilities of extinction, species-specific probbilities of coloniztion, nd nestedness of resources or hbitt types. We used nested subsets nlysis to exmine distribution ptterns of birds in 83 loctions in the centrl Gret Bsin. Site re nd topogrphy did not pper to differ in their influence on nestedness of birds. In ddition, riprin dependence hd little discernible effect on nested distribution ptterns of birds. Elevtionl influences on species richness of birds (Fleishmn nd Dobkin in review). From 2 25, we conducted point counts in 28 loctions in three mountin rnges. The men elevtion of riprin points (n = 3) versus non-riprin points (n = 5) ws not significntly different cross the three mountin rnges or within ny mountin rnge. The proportion of riprin versus non-riprin points, however, differed considerbly mong the three mountin rnges. In the Toiybe Rnge, 75% of points were riprin, s compred with 34% in the Shoshone Mountins nd just 5% in the Toquim Rnge. The men elevtion of points dominted by different types of vegettion (spen, mixed tree, pinyon-juniper, shrub, nd 6

17 willow) ws not significntly different cross the three mountin rnges or within ny mountin rnge. The proportion of points dominted by trees versus shrubs differed somewht mong the three rnges (4% in the Shoshone Mountins, 69% in the Toquim Rnge, nd 52% in the Toiybe Rnge). When dt for the three mountin rnges were pooled, severl of the tested reltionships between species richness nd elevtion were sttisticlly significnt, but only smll proportion of the vrince in species richness ws explined by elevtion. All sttisticlly significnt correltions in the Shoshone Mountins, Toiybe Rnge, nd cross the three mountin rnges were negtive (species richness decresed unimodlly s elevtion incresed), wheres ll sttisticlly significnt correltions in the Toquim Rnge were positive (species richness incresed unimodlly s elevtion incresed). The response of species richness to elevtion in the Toiybe Rnge generlly ws not sttisticlly significnt. Drmtic differences mong rnges in proportion of lnd-cover types nd vilbility of surfce wter my drive the estblishment of distinct vin ssemblges with different numbers nd proportions of riprin obligtes nd different distributionl ptterns. Effects of sptil scle on species composition of birds (Mc Nlly et l. 24). We exmined whether vrition in species composition of breeding birds in the centrl Gret Bsin depended on smpling grin (the smllest resolvble unit of study) nd on the reltive proximity of smpling units cross the lndscpe. Dt were compiled t three smpling grins, sites (verge 2 h), cnyons (verge 74 h), nd mountin rnges. First, we investigted whether men similrity of species composition mong the three rnges differed s function of the grin size t which dt were compiled. Second, we explored whether men similrity of species composition ws greter for cnyons within the sme mountin rnge thn for cnyons within different mountin rnges. Third, we exmined whether men similrity of species composition t the site level ws different for sites within the sme cnyon, sites within different cnyons in the sme mountin rnge, nd sites within cnyons in different mountin rnges. Men similrity of species composition incresed s the smpling grin incresed. Men similrity of species composition t the cnyon level nd site level ws ffected by reltive proximity of smpling loctions; turnover incresed s the reltive isoltion of smpling loctions incresed. The sensitivity of turnover to smpling grin likely reflects the effect of locl environmentl heterogeneity: s smpling grin increses, biotic ssemblges pper more homogeneous. The degree of vrition in species composition suggested tht sptilly extensive smpling will be more effective for drwing inferences bout regionl ptterns of species richness nd composition thn intensive smpling t reltively few, smller sites. Ptterns of sptil utocorreltion of ssemblges of birds, floristics, physiognomy, nd primry productivity. We exmined reltionships between sptil ptterns of species richness of birds nd three ttributes of birds hbitts, txonomic composition of vegettion (floristics), physicl structure of vegettion (physiognomy), nd NDVI, mesure of photosyneticlly ctive rdition estimted from stellite imges, t 279 loctions in the centrl Gret Bsin (Mc Nlly nd Fleishmn in review). We fitted sptil utocorreltion functions (semivriogrms) to distnce-bsed dt for these four vribles. The semivriogrm for bird species richness ws liner, wheres semivriogrms for the three hbitt vribles were either sphericl (vegettion composition) or power-lw (vegettion structure, NDVI). The lck of reltionships between sptil ptterns of the three hbitt vribles nd species richness suggests tht species richness ws not controlled by these vribles. The sptil pttern for species composition of birds, in which the best-fit semivriogrm ws sphericl with 3-km rnge, ws more concordnt with sptil ptterns of hbitt vribles such s vegettion composition (Fleishmn nd Mc Nlly 7

18 26). Resolving the disprity between ptterns for species richness nd ssemblge composition of birds could provide vluble informtion, especilly for conservtion plnning. Distinguishing between species on the bsis of ecologicl speciliztion (i.e., functionl groups or guilds) my be required to revel reltionships between sptil ptterns of birds nd their resources. Remote sensing vribles s predictors of species richness of birds (Seto et l. 24). The bility to predict sptil ptterns of species richness using few esily mesured environmentl vribles would fcilitte timely evlution of potentil impcts of nthropogenic nd nturl disturbnces on biologicl diversity nd ecosystem function. Two common hypotheses mintin tht funl species richness cn be explined in prt by either locl vegettion heterogeneity or primry productivity. We exmined the potentil of single-dte Normlized Difference Vegettion Index (NDVI) dt, which is ssocited with vegettion productivity nd derived from Lndst Themtic Mpper (TM) imges, to predict species richness of birds. Results indicted positive correltion, but with no definitive functionl form, between species richness nd productivity. The strongest reltionships between species richness of birds nd NDVI were observed t lrger smpling grins nd extent, t which severl mesures of NDVI explined more thn 5% of the vrition in species richness. Potentil of bird conservtion to protect co-occurring species (Betrus et l. 25). Vlidtion tht conservtion of certin species effectively protects high proportion of co-occurring species is rre. Our previous work hd suggested tht n umbrell index bsed on geogrphic distribution nd life history chrcteristics would mximize the proportion of conspecifics protected per unit re conserved. Using bird nd butterfly dt from three mountin rnges in the Gret Bsin, we exmined whether umbrell species lso would confer protection to species in different txonomic groups. Further, we ddressed the sptil trnsferbility of umbrell species by considering whether species identified s umbrells in one mountin rnge would be effective umbrells in other mountin rnges. Overll, equl proportions of species would be protected using either cross-txonomic umbrell species or sme-txon umbrell species. Our dt suggested tht in given mountin rnge, umbrell species identified using dt from the sme mountin rnge versus different mountin rnge would be eqully effective. The bility of one set of umbrell species to confer protection to co-occurring species, however, my vry mong txonomic groups nd geogrphic regions. Potentil of birds s indictors of species richness (Fleishmn et l. 25, Thomson et l. 25). Indictor species models my be cost-effective pproch to estimting species richness cross lrge res. Obtining relible distributionl dt for indictor species (nd therefore relible estimtes of species richness) often requires longitudinl dt, tht is, surveys for indictor species repeted for severl yers or time steps. Mximum informtion must be extrcted from such dt. We compred the influence of presence/bsence dt nd reporting rte dt (the proportion of survey yers in which species ws present) on models of species richness bsed on indictor species. Using dt on birds nd butterflies, we evluted models of species richness for one txonomic group bsed on indictor species drwn from the sme txonomic group nd from different group. We lso evluted models of combined species richness of both txonomic groups bsed on indictor species drwn from either group. We identified suites of species whose occurrence ptterns explined s much s 7% of devince in species richness of different txonomic group. Vlidtion tests reveled strong correltions between observed nd predicted species richness, with 83 % of the observed vlues flling within the 95% credible intervls of the predictions. Whether reporting rte dt improved the 8

19 explntory nd predictive bility of cross-txonomic models depended on the txonomic group of the indictor species. The discrepncy in predictive bility ws smller for sme-txon models. Our methods provide mnger with the mens to mximize the informtion obtined from longitudinl survey dt. PRODUCTS SUPPORTED BY JFSP # B-3-3- Peer-reviewed Publictions Fleishmn, E., C.J. Betrus, R.B. Blir, R. Mc Nlly, nd D.D. Murphy. 22. Nestedness nlysis nd conservtion plnning: the importnce of plce, environment, nd life history cross txonomic groups. Oecologi 33: Fleishmn, E., C.J. Betrus, nd R.B. Blir. 23. Effects of sptil scle nd txonomic group on prtitioning of butterfly nd bird diversity in the Gret Bsin. Lndscpe Ecology 8: Fleishmn, E., C.J. Betrus, nd L.P. Bulluck. 23. Annul vribility of species richness nd composition of breeding birds in the centrl Gret Bsin. Gret Bsin Birds 6(): Biley, S-A., S. Anderson, K. Crney, E. Clelnd, M.C. Horner-Devine, G. Luck, L.A. Moore, C. Betrus, nd E. Fleishmn. 24. Primry productivity nd species richness: reltionships mong functionl guilds, residency groups nd vgility clsses t multiple sptil scles. Ecogrphy 27: Fleishmn, E. nd R. Mc Nlly (24). Linking models of species occurrence nd lndscpe reconstruction. Trnsctions of the Western Section of the Wildlife Society 38/39: 4. Mc Nlly, R., E. Fleishmn, L. Bulluck, nd C. Betrus. 24. Comprtive influence of sptil scle on bet diversity within regionl ssemblges of birds nd butterflies. Journl of Biogeogrphy 3: Seto, K.C., E. Fleishmn, J.P. Fy, nd C.J. Betrus. 24. Linking sptil ptterns of butterfly nd bird species richness with Lndst TM derived NDVI. Interntionl Journl of Remote Sensing 25: Betrus, C.J., E. Fleishmn, nd R.B. Blir. 25. Cross-txonomic potentil nd sptil trnsferbility of n umbrell species index. Journl of Environmentl Mngement 74: Fleishmn, E. 25. Identifiction nd conservtion ppliction of signl, noise, nd txonomic effects in diversity ptterns. Invited pper. Animl Biodiversity nd Conservtion 28: Fleishmn, E., R. Mc Nlly, nd J.R. Thomson. 24 (25). Chllenges nd opportunities for conserving funl biodiversity in rid ecosystems. Invited pper. Annls of Arid Zone 43: Fleishmn, E., J.R. Thomson, R. Mc Nlly, D.D. Murphy, nd J.P. Fy. 25. Predicting species richness of multiple txonomic groups using indictor species nd genetic lgorithms. Conservtion Biology 9: Thomson, J.R., E. Fleishmn, R. Mc Nlly, nd D.S. Dobkin. 25. Influence of the temporl resolution of dt on the success of indictor species models of species richness cross multiple txonomic groups. Biologicl Conservtion 24:

20 Bulluck, L.P., E. Fleishmn, C.J. Betrus, nd R.B. Blir. 26. Sptil nd temporl vrition in species occurrence rte ffects the ccurcy of occurrence models. Globl Ecology nd Biogeogrphy 5: Fleishmn, E. nd R. Mc Nlly. 26. Ptterns of sptil utocorreltion of ssemblges of birds, floristics, physiognomy, nd primry productivity in the centrl Gret Bsin, USA. Diversity nd Distributions 2: Chmbers, J. C. 25. Fire relted restortion issues in woodlnd nd rngelnd ecosystems. Pges In L. Tylor, J. Zelnik, S. Cdwllder, nd B. Hughes (compilers). Mixed Fire Regimes: Ecology nd Mngement. Symposium Proceedings. November 7-9, 24, Spokne, WA. AFE MIXC3. Ru, B. M., J. C. Chmbers, R. R. Blnk, nd W. W. Miller. 25. Hydrologic response to prescribed fire in centrl Nevd pinyon-juniper (Pinus monophyll-juniperus osteosperm) woodlnd. Journl of Rnge Mngement. 58: Suring, Lowell, Michel J. Wisdom, Robin J. Tusch, Richrd F. Miller, Mry M. Rowlnd, Lind Schueck, nd Cr W. Meinke. 25. Chpter 4: Modeling threts to sgebrush nd other shrublnd communities. Pges 4-49 in Wisdom, M.J., M.M. Rowlnd, nd L.H. Suring, editors. Hbitt threts in the sgebrush ecosystem: methods of regionl ssessment nd pplictions in the Gret Bsin. Allince Communictions Group, Lwrence, Knss, USA. D Antonio, C. nd J. C. Chmbers. 26. Using ecologicl theory to mnge or restore ecosystems ffected by invsive plnt species. In D. Flk, M. Plmer, nd J. Zedler (eds). Foundtions of Restortion Ecology. Islnd Press, Covelo, CA. in press. Wisdom, Michel J., Robin J. Tusch nd Mry M. Rowlnd. 26. Effective mngement strtegies for sge-grouse nd sgebrush: A question of trige? Proceedings Ntionl Acdemy of Sciences (in press) Mnuscripts Submitted or In-Preprtion Fleishmn, E. nd D.S. Dobkin. Response of vin species richness to elevtion in the centrl Gret Bsin. Gret Bsin Birds. Submitted. Fleishmn, E., R. Donnelly, nd J.P. Fy. Applictions of nestedness nlyses to biodiversity conservtion in developing lndscpes. Lndscpe nd Urbn Plnning. Submitted. Mc Nlly, R. nd E. Fleishmn. Reltionships between sptil vrition in ecosystem mesures nd species richness of birds in desert montne system. Ecogrphy. Submitted. Mont Blnc, E., J. C. Chmbers, nd P. Brussrd. The effects of fire nd tree density on nt popultions in pinyon-juniper dominted wtersheds. Western North Americn Nturlist. Submitted. Thomson, J.R., E. Fleishmn, R. Mc Nlly, nd D.S. Dobkin. Comprison of predictor sets for species richness nd the number of rre species of butterflies nd birds. Journl of Biogeogrphy. Submitted. Ru, B. M., J. C. Chmbers, R. R. Blnk, nd D. W. Johnson. Effects of pinyon (Pinus monophyll) nd juniper (Juniperus osteosperm) expnsion nd prescribed fire on understory biomss nd nutrient concentrtion in Gret Bsin sgebrush ecosystems. Plnt Ecology. Submitted. Ru, B. M., R. R. Blnk, J. C. Chmbers, nd D. W. Johnson. Pinyon-juniper expnsion nd prescribed fire effects on soil in Gret Bsin sgebrush ecosystems. Journl of Environmentl Qulity. Submitted. 2

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3.

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3. The nlysis of vrince (ANOVA) Although the t-test is one of the most commonly used sttisticl hypothesis tests, it hs limittions. The mjor limittion is tht the t-test cn be used to compre the mens of only

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