Vegetation and Soil Restoration on Highly Impacted Campsites in the Eagle Cap Wilderness, Oregon

Transcription

1 Vegettion nd Soil Restortion on Highly Impcted Cmpsites in the Egle Cp Wilderness, Oregon Dvid N. Cole Dvid R. Spildie United Sttes Deprtment of Agriculture Forest Service Rocky Mountin Reserch Sttion Generl Technicl Report RMRS-GTR-185 Februry 2007

2 Cole, Dvid N.; Spildie, Dvid R Vegettion nd soil restortion on highly impcted cmpsites in the Egle Cp Wilderness, Oregon. Gen. Tech. Rep. RMRS-GTR-185. Fort Collins, CO: U.S. Deprtment of Agriculture, Forest Service, Rocky Mountin Reserch Sttion. 26 p. Abstrct We ssessed the effectiveness of plnting techniques (seeding nd trnsplnting) nd restortion tretments designed to improve the physicl, biologicl, nd chemicl properties of soils nd meliorte microclimtic conditions on six closed cmpsites in sublpine forests. Restortion tretments included scrifiction, soil mendment with orgnic mtter, compost nd soil inoculum, nd ppliction of mulch blnket. Cmpsite closure, scrifiction, plnting, nd soil mendments were successful in incresing recovery rtes. The mulch blnket hd no effect on recovery. 10 yers fter cmpsite closure, vegettion cover ws still diminished in comprison to reference conditions on nerby undisturbed sites. Prticulrly problemtic ws reestblishment of the low-growing shrub species (prticulrly Vccinium scoprium nd Phyllodoce empetriformis) tht re the most bundnt groundcover species in these forests. These species seldom estblish from seed. Moreover, survivorship nd growth rtes re unusully low for trnsplnts. Our results show the reltive ese of estblishing vrious species nd growth forms in these forests, s well s which species nd growth forms respond best to the pplied tretments. Results reinforce the importnce of voiding impcts in the first plce, the lengthy recovery periods required in these ecosystems, nd the intensive restortion efforts needed to speed recovery. Key words: compost, mulch, recretion impct, scrifiction, seeding, soil mendments, trnsplnting Authors Dvid N. Cole is Reserch Biologist with the Aldo Leopold Wilderness Reserch Institute, Missoul, MT. He received n A.B. degree in geogrphy from the University of Cliforni, Berkeley, nd Ph.D. in geogrphy from the University of Oregon. Dvid R. Spildie is Nturl Resources Specilist with Recretion Solutions, Missoul, MT. He ws biologist with the Aldo Leopold Wilderness Reserch Institute when this study ws conducted. Acknowledgments Mny individuls contributed to this project. Numerous Forest Service district personnel ssisted with implementtion of the tretments nd study, most notbly Tom Crlson, Kendll Clrk, nd Jerry Hustf. Mny others ssisted with dt collection, most notbly Jeff Comstock. Tom DeLuc coordinted nlysis of soil smples. Nel Christensen nd Rudy King ssisted with dt nlysis nd Dve Prsons nd Jeff Mrion provided useful review comments. cover photos: (top) Closed cmpsite t Horseshoe Lke 10 yers fter restortion tretments were pplied. (middle) Groundcover vegettion on smple plot, 10 yers fter scrifiction, plnting nd soil mendments. (bottom) Amending the soil of long-disturbed cmpsite with orgnic mtter, compost nd ntive soil inoculum. You my order dditionl copies of this publiction by sending your miling informtion in lbel form through one of the following medi. Plese specify the publiction title nd series number. Publishing Services Telephone: (970) FAX: (970) E-mil: rschneider@fs.fed.us Web site: Miling ddress: Publictions Distribution Rocky Mountin Reserch Sttion 240 West Prospect Rod Fort Collins, CO 80526

3 Vegettion nd Soil Restortion on Highly Impcted Cmpsites in the Egle Cp Wilderness, Oregon Dvid N. Cole, Dvid R. Spildie Contents Introduction... 1 Study Sites... 1 Methods... 1 Tretments... 1 Mesurements... 4 Dt Anlysis nd Presenttion... 5 Results... 5 Effects on Soils... 5 Effects on Totl Plnt Cover... 7 Effects on Trnsplnt Cover... 9 Effects on Seedling Cover...11 Seedling Estblishment nd Mortlity Effects on Species Richness Progress in Reltion to Reference Conditions Discussion Tretment Effects Further Reserch Tretment Needs of Individul Species Mngement Implictions A Prescription for Cmpsite Restortion Restortion nd Visitor Mngement References... 26

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5 Introduction Sublpine lkes re mgnets for recretion use throughout the mountinous wilderness of the western United Sttes. People flock to these lkes for their outstnding scenic qulities, s well s to fish, swim, picnic, nd cmp. Although use is much greter tody thn it ws before the 1960s, some populr lkes hve been regulrly visited for the better prt of century. Regulrly visited plces, prticulrly those tht serve s cmpsites, quickly become impcted. Understory vegettion disppers, s do orgnic soil horizons. Exposed minerl soil becomes highly compcted, reducing rinfll infiltrtion nd incresing erosion. Wilderness mngers re concerned bout such cmpsite impcts given their mndte to preserve nturl conditions in wilderness. While it is cler tht some level of impct is inevitble if overnight cmping is llowed, these impcts my be more numerous thn they should be or my be occurring in loctions where they re considered uncceptble. Incresingly, mngers re closing impcted sites to cmping to llow them to return to pre-disturbnce conditions. This gol hs often proved elusive, however. Recovery cnnot occur unless ll site use is curtiled. Wilderness mngers re often reluctnt to erect the signs nd ropes tht cn be effective in keeping wlkers, nglers, nd picnickers, s well s cmpers, off closed cmpsites. Even where recretionl use is effectively curtiled, recovery cn be n extremely slow process in sublpine ecosystems. Growing sesons re short nd climtic conditions during these sesons cn be hrsh. Mny of the bundnt sublpine plnts estblish infrequently nd grow slowly. Moreover, the soil on mny closed sites no longer provides good growing medium for plnts. Mny decdes without vegettion nd with miniml orgnic input hve left the plnts physiclly, biologiclly, nd chemiclly impoverished. Mngers often responded by employing vrious restortion techniques in ttempts to ccelerte successionl processes (Lester 1990). However, these efforts re often costly in terms of time nd money, nd frequently re not very successful (Moritsch nd Muir 1993). Given the lck of informtion on effective mens of restoring sublpine cmpsites, we initited long-term experiments using severl common restortion tretments. Specificlly, we ssessed the effectiveness of (1) scrifiction, (2) trnsplnting nd seeding with locl, ntive species, (3) meliorting microclimtic conditions with mulch mt, nd (4) mending soils with orgnic mtter, compost, nd soil inoculum. Tretment effects were closely followed for 10 yers. Study Sites The study ws conducted on six cmpsites in the Egle Cp Wilderness, Wllow Mountins, northestern Oregon. All sites were locted t n elevtion of 2215 to 2300 m, djcent to sublpine lkes, bout 12 to 15 km from the closest rod. They were locted in forests with n overstory of Abies lsiocrp (sublpine fir), Pinus contort (lodgepole pine), nd Pinus lbiculis (whitebrk pine). Ground cover vegettion on djcent, little disturbed sites is discontinuous (typiclly bout 50 percent cover). Ericceous dwrf shrubs, Vccinium scoprium (grouse whortleberry) nd Phyllodoce empetriformis (red mountin hether), nd the cespitose grminoids, Juncus prryi (Prry s rush) nd Crex rossii (Ross sedge), re the most bundnt species. This plnt community type occurs throughout much of the western United Sttes t high elevtions, prticulrly in loctions tht re populr destintions for wilderness recretion. The impcts of wilderness cmping re probbly more common in this community type thn ny other in the United Sttes, mking informtion bout effective restortion techniques prticulrly useful in this type. Soils re shllow, sndy, nd cidic (ph between 4.2 nd 4.8) nd re derived from grnitic substrtes (Cryochrepts nd Cryorthents). Although snow typiclly covers the ground until lte June/erly July, snowmelt is typiclly followed by hot, dry summers. The frequency of summer thunderstorms vries from yer to yer. When they re infrequent, soils cn be highly droughty for severl months (most of the growing seson). These cmpsites hve probbly exhibited high levels of impct (lck of vegettion, miniml soil orgnic horizons, nd compcted minerl soil) for t lest 50 yers. Prior to restortion, these cmpsites were typiclly bout 200 m 2 in size, with bout 100 m 2 completely devoid of vegettion. Soil orgnic horizons hd eroded wy over substntil portions of these sites nd minerl soils were so compcted tht infiltrtion rtes were reduced by lmost 50 percent (Cole nd Fichtler 1983). Potentilly minerlizble N nd microbil ctivity were lso substntilly reduced on these cmpsites (Zbinski nd others 2002). Methods Tretments Cmpsite restortion begn in 1995 with the closure of these sites to cmping, using closure signs nd USDA Forest Service RMRS-GTR

6 rope. The closures were highly effective. Over the 10 yers of the study, we found only few instnces where it ppered someone hd wlked on the plots. Nobody cmped on ny of the sites. A three-fctor experiment, using split plot design ws employed. Twelve tretment plots (1.5 m by 1.5 m) were estblished on ech cmpsite. The soil ws scrified on these plots. Scrifiction involved the use of shovels, picks, pitchforks, hoes, nd hnd kneding to brek up compction nd clods to depth of bout 15 cm. Substntil mixing of soil horizons ws unvoidble in our resolve to develop crumb texture. On severl sites, numerous tree roots were cut nd removed during scrifiction. This intensity of scrifiction exceeds tht commonly undertken on wilderness cmpsites. A control plot (not treted in ny wy) ws estblished within the closed re on prt of the cmpsite tht ws not scrified. This plot ws used in n nlysis, seprte from the fctoril experiment, of the effect of scrifiction in the bsence of mulch, soil mendments, nd plnting. Of the three fctors in the split plot experiment, the mulch tretment ws the fctor used to estblish whole plot units becuse it ws most fesible to pply mulch blnkets over lrge res. Six contiguous plots on ech site were covered with biodegrdble mulch mde of strw interwoven with cotton string nd jute (Bionet )(fig. 1). The other six contiguous plots were not mulched. Within ech of the two mulch whole plots, three levels of soil mendment nd two levels of plnting were ssigned to split-plot units in completely rndomized design. Ech combintion of soil mendment nd plnting occurred in ech whole plot. Figure 2 illustrtes the lyout for one of the cmpsites. Ech cmpsite hd unique ordering of tretments within the mulch whole plots. Ech cmpsite provided one of six replictes. Figure 1. The ppernce of the mulch blnket immeditely fter plnting. There were three levels of the soil mendment fctor. Within ech whole plot, two tretment plots (split-plot units) received no mendments. Another two plots were mended with orgnic mtter nd inoculted with ntive soil. The orgnic mtter ws mix of pet moss (20 percent) nd well decomposed, loclly collected orgnic mtter. The dry pet moss ws mixed with wter before ppliction. A 2.5 cm lyer of this orgnic mteril ws mixed with minerl soil to depth of 7.5 cm (fig. 3). Soil from the rooting zone of locl trnsplnts ws the source for the inoculum. About 1.2 liters of soil were mixed with bout 20 liters of wter to mke slurry. Three liters of this slurry were sprinkled over ech plot nd rked into the soil. The finl two plots were mended with compost in ddition to the orgnic mtter nd inoculum tretment. We dded 2.5 cm of commercilly vilble compost (sewge sludge/log yrd wste compost with C:N of pproximtely 20:1; Ekocompost, Missoul, Montn), lightly wtered nd rked into the top 10 cm of orgnic nd minerl soil. Figure 2. Distribution of tretments for one cmpsite, illustrting completely rndom ssignment of tretments within mulch whole-plot units, s well s the seprte non-scrified control. Tretments re: mulch or no mulch; seed or no seed; no soil mendment, orgnics mendment or orgnics nd compost mendment; nd scrified or not scrified. 2 USDA Forest Service RMRS-GTR

7 Figure 3. Appliction of soil mendments. The two levels of plnting were plnted nd unplnted. Within ech whole plot, three plots were plnted (seeded nd trnsplnted) nd three were not. Seeding involved (1) collecting seed loclly from severl species with mture seed, (2) dividing vilble seed into equl quntities for ech seeded plot, (3) pinch-brodcsting seed over the plot, nd (4) rking seed into the upper 2.5 cm of soil. Seeded species vried between cmpsites depending on loclly vilble plnts with mture seed. Juncus prryi nd Phleum lpinum (lpine timothy) were seeded on three of the cmpsites. Antennri lpin (lpine pussytoes), Antennri lnt (woolly pussytoes), nd Sibbldi procumbens (creeping sibbldi) were seeded on two cmpsites. Aster lpigenus (lpine ster), Dnthoni intermedi (timber otgrss), Penstemon globosus (globe penstemon), nd Sitnion hystrix (bottlebrush squirreltil) were ech seeded on one cmpsite. Loclly vilble seed ws unusully limited due to the unusully short growing seson in One of the cmpsites (t Crescent Lke) ws not seeded due to lck of mture seed in the vicinity. Trnsplnting involved (1) digging up enough trnsplnts in the vicinity to plnt equl numbers of ech species in ech plot, (2) digging hole nd plcing trnsplnts in the hole, long with Vit-strt (vitmin B-1) to reduce trnsplnt shock (fig. 4), nd (3) giving ech trnsplnt 0.6 liters of wter. Plots not plnted were given n equivlent mount of wter. Most trnsplnt plugs were between 5 nd 25 cm in dimeter, nd most plots received five to six plugs (fig. 5). Most plugs contined only one species, but some contined more thn one. Trnsplnted species vried between cmpsites. Vccinium scoprium nd Juncus prryi were intentionlly trnsplnted on five of the six cmpsites. Phyllodoce empetriformis, Crex rossii, Luzul hitchcockii (smooth woodrush), nd Sibbldi procumbens were intentionlly trnsplnted on two cmpsites. Species tht were intentionlly trnsplnted on only one cmpsite Figure 4. Plnting trnsplnt. Figure 5. Typicl trnsplnt cover nd density immeditely fter plnting. were Abies lsiocrp, Achille millefolium (yrrow), Antennri lpin, Antennri lnt, Aster lpigenus, Clmgrosts cndensis (bluejoint reedgrss), Dnthoni intermedi, Hypericum formosum (western St. John s-wort), Oryzopsis exigu (little ricegrss), Pinus contort, Polemonium pulcherrimum (showy polemonium), nd Spire betulifoli (shiny-lef spire). Thirteen other species were unintentionlly included in plugs. Nomenclture follows Hitchcock nd Cronquist (1973). Seeding nd trnsplnting occurred only in the centrl 1 m 2 of ech plot. Mesurements were lso confined to this centrl re, leving 0.5 m buffer between the mesured portion of ech treted plot. In 1996, when it ppered tht soils were extremely dry, plots were wtered severl times. We did this becuse reports from erlier cmpsite restortion projects in the Pcific Northwest indicted tht it is common for most seedlings to die during prolonged periods of summer drought USDA Forest Service RMRS-GTR

8 (Lester 1990). All plots were given n equl mount of wter (bout 2 liters per plot). No supplementl wtering occurred in lter yers. Climtic vribility clerly influenced temporl ptterns of plnt response, primrily by determining the vilbility of soil moisture. Although there ws yerly vrition, growing seson conditions generlly becme incresingly droughty over the 10-yer study period. For the first 4 yers of the study, the lte snowpck ws unusully deep, suggesting tht erly seson conditions were much less droughty thn norml. This ws not the cse for the finl 6 yers of the study. In 1996, the first growing seson fter restortion, when plots were occsionlly irrigted, the summer ws dry but cool. In 1997, the summer ws cool nd wet. In 1998, the summer ws hot nd dry, nd plnts were not given supplementl wter. Long-term, regionl drought set in with the hot, dry summer of 2000 nd the generlly low precipittion tht fell throughout Drought persisted through Mesurements We mesured trnsplnt survivl nd growth, seedling density nd plnt cover t lest once every yer for 7 yers (fig. 6). In ddition, cover ws ssessed fter 10 yers. Trnsplnts For ech trnsplnt, we mesured rel extent of cnopy cover (using 1 m squre PVC frme with 5 cm by 5 cm grid) nd mximum height. Mesurements were tken immeditely fter trnsplnting (September 1995) nd in ech September therefter, for 7 yers (through 2002). Seedlings Seedling estblishment ws ssessed beginning in erly July Every two weeks from erly Figure 6. Qudrts used to ssess plnt cover, seedling density, nd trnsplnt growth. July to erly September (four times), ll estblished seedlings were mpped. Ech seedling ws identified by species, nd colored toothpick ws plced next to it to denote dte of estblishment. This mde it possible to ssess period of estblishment nd deth if mortlity occurred. In 1997, seedlings tht germinted in 1996 were identified on the bsis of their size, loction, nd species. New seedlings (the 1997 cohort) were identified in the surveys conducted every two weeks. In some plots, seedlings were so numerous tht they were ssessed in subplots. In 1998, seedling ssessment occurred twice, in mid-july nd erly September. In subsequent yers (through 2002), we ssessed the density of seedlings (plnts tht hd estblished by seed since the beginning of the study) only in September. Ech yer, 10 individuls of seeded species were rndomly selected on ech plot, nd their height ws mesured in September. In 1996 nd 1997, we excvted four individuls of the sme species tht were growing within the treted plot, but outside the re where mesurements were tken. Their root nd shoot biomss ws mesured following clening nd drying. In 1998, we mesured the height of the tllest individul of the seeded species, which we found to be highly correlted with biomss. This voided the need for further destructive smpling. In 1997 nd 1998, height nd biomss mesurements were tken only on seedlings tht germinted nd estblished in In 1996 nd 1997, trnsplnt rel extent nd seedling loctions were digitized to llow sptil nlysis. Plnt cover Every September between 1996 nd 2002, totl plnt cover ws oculrly estimted to the closest percent if cover ws 10 percent or less nd in 10 percent increments therefter. Totl vegettion cover ws ssessed, s ws the cover of ech trnsplnted species nd ech species tht hd estblished from seed. Finl cover estimtes were tken in September 2005, 10 yers fter seeding nd trnsplnting. Reference conditions Totl vegettion cover nd the cover of individul species were estimted in undisturbed plots ner to ech restored cmpsite. The mens from these six plots re used s trgets for successful restortion. Soil chrcteristics Soils nlyses were conducted twice during the study. Three yers fter the restortion tretments, soil smples were collected on ech cmpsite from (1) plots tht received the tretment of orgnics, inoculum, nd compost, (2) untreted prts of the closed cmpsite, nd (3) undisturbed sites djcent to the cmpsites. Prmeters ssessed were: microbil biomss C, bsl respirtion rtes, totl orgnic crbon, totl nitrogen, mmonium, potentilly minerlizble nitrogen, nd 4 USDA Forest Service RMRS-GTR

9 severl indictors of the crbon utiliztion cpbilities of the microbil community. Detils of the lbortory nlyses nd results cn be found in Zbinski nd others (2002). In 2003, 8 yers fter the restortion tretments, soil smples were gin tken on ech cmpsite. As before, smples were tken from untreted prts of the closed cmpsite nd from undisturbed sites djcent to the cmpsites. This time, however, the effects of six different tretments were ssessed: plnted plots tht received ech of the three soil mendment tretments nd unplnted plots tht received ech of the three soil mendment tretments. In ech cse, only plots tht were not covered with the mulch mt were smpled. Soil nlysis procedures were identicl to those used 5 yers erlier, except tht crbon utiliztion cpbilities were not investigted. Dt Anlysis nd Presenttion Some of the results of this study re presented in more detil elsewhere. Zbinski nd others (2002) describe results of the erly soils study. Cole nd Spildie (2006) nd Cole (in press) present detiled results for trnsplnts nd seedlings, respectively. In this report, results presented elsewhere re only briefly reviewed. Most ttention is given to the effect of restortion on soil 8 yers fter tretment nd to how plnt cover responded to tretment over the 10-yer study period. The soils dt re nlyzed using pired t-tests nd univrite nlyses of vrince in SPSS 9.0. First we tested differences between untreted prts of the cmpsite nd undisturbed control sites using pired t-tests. Then we tested the effects of the min fctors, soil mendment, nd plnting on soil chrcteristics. Most dt were squre-root trnsformed to better comply with ssumptions bout normlity. Where there were significnt min effects, we ssessed the significnce of differences between mens using Tukey s honestly significnt differences test, which djusts for multiple comprisons. Then we ssessed which tretments differed from the untreted cmpsite using Dunnett s tests. Finlly, we ssessed which tretments differed from undisturbed controls, lso using Dunnett s tests. For the cover dt, we performed repeted mesures nlyses of vrince, pproprite for split-plot designs (using n utoregressive covrince structure, PROC MIXED in SAS 9.1). Most dt were squreroot trnsformed to better comply with ssumptions bout normlity. In mny cses, tretment effects vried significntly with time since tretment (in other words, interctions with time were significnt). In these cses, we describe tretment effects for ech of the 10 yers of the experiment, but the significnce of effects is only ssessed t the end of the experiment, in In those cses where tretment interctions with time were not significnt, we report results of the repeted mesures nlyses. To ssess the hypothesis tht scrifiction hs positive effects, the control (the plot tht ws not even scrified) ws compred to the one plot on ech cmpsite tht ws scrified but not mulched, mended, or plnted. For the three fctors included in the split plot design (mulch, soil mendment, nd plnting), min effects of ech fctor nd interctions mong fctors were ssessed. Interctions mong the tretments were never sttisticlly significnt. Therefore, we simply report the min effects of tretments. We report ANOVA tretment effects when there re two tretments being compred (effects of scrifiction, plnting, nd mulch). For the soil mendments, we report Dunnett s tests, djusted for multiple comprisons, of differences between ech of the two mendments nd the non-mended tretment, s well s Tukey-Krmer tests, djusted for multiple comprisons, of differences between the two mendments. Results Effects on Soils Compred to undisturbed controls, unrestored cmpsite soils hd significntly lower orgnic crbon (C), totl nitrogen (N), mmonium (NH4), potentilly minerlizble nitrogen (PMN), nd microbil biomss (fig. 7). Bsl respirtion rtes (n indictor of the mgnitude of microbil popultions) did not differ significntly. These results, bsed on soils collected in 2003, 8 yers fter the beginning of the experiment, re similr to those reported for soils collected in 1998 (Zbinski nd others 2002). In 1998, bsl respirtion rtes did differ between cmpsites nd controls if the soil on controls ws collected immeditely underneth vegettion, rther thn in open res between vegettion clumps. Zbinski nd others (2002) lso report tht control sites hd higher mesures for n indictor of the functionl diversity of microbil popultions, the number of substrtes metbolized in crbon utiliztion profiles. Since cmping reduced levels of most of these prmeters, we hypothesized tht soil mendments nd plnting might ccelerte soil recovery nd increse levels of these prmeters. The soil mendments hd much more pronounced effects on soil chrcteristics thn whether or not plots were trnsplnted nd seeded USDA Forest Service RMRS-GTR

10 Figure 7. Effect of restortion tretments on soil chrcteristics. (tble 1). Compred to soil in plots tht were only scrified, the soil in plots mended with orgnics nd compost hd significntly higher levels of orgnic C, totl N, potentilly minerlizble N, bsl respirtion rtes, nd microbil biomss (fig. 7). In the plots mended with orgnics only (no compost), soils hd significntly higher levels of orgnic C nd totl N thn on scrified plots nd significntly lower levels of orgnic N, potentilly minerlizble N, bsl respirtion rtes, nd microbil biomss thn on orgnics nd compost plots. Plots mended with orgnic mtter nd compost differed significntly from untreted cmpsites for ll prmeters other thn bsl respirtion. Moreover, plots mended with orgnics nd compost were not significntly different from controls for ny of these prmeters. This suggests mendment with orgnics nd compost 6 USDA Forest Service RMRS-GTR

11 Tble 1. Anlysis of vrince results (F vlues) for the effect of soil mendments nd plnting on soil chrcteristics. Source of vrition df Orgnic C Totl N NH 4 PMN Respirtion Biomss C Soil mendment ** 18.5** * 5.1* 3.6* Plnting 1 5.9* 7.1* Interction Significnce: *<0.05; **<0.01 F ws sufficient to meliorte mny of the effects of longterm cmping on these soils, without exceeding levels found in undisturbed soils. Plots mended with orgnics only hd soil chrcteristics tht were consistently intermedite between those for untreted cmpsites nd control sites. However, orgnics only plots differed significntly from untreted cmpsites only in orgnic C. They differed from controls only in potentilly minerlizble N. Plots tht were only scrified did not differ significntly from untreted cmpsites for ny of these prmeters. They differed from controls for ll prmeters other thn NH 4 nd bsl respirtion. For most of these prmeters, the soil in plots tht were trnsplnted nd seeded ws not significntly different from the soil in plots tht were not plnted. Contrry to our hypotheses, orgnic C ws significntly lower on plnted plots (6.5 percent) thn it ws on plots tht were not plnted (8 percent). Similrly, totl N ws significntly lower on plnted plots (0.25 percent) thn it ws on plots tht were not plnted (0.28 percent). Neither of these differences seems substntil, but clerly plnting lone does not led to rpid recovery of soil properties. Effects on Totl Plnt Cover Following closure nd restortion of the cmpsites, totl plnt cover incresed from zero to 12 percent 10 yers lter (tble 2). Totl cover reched mximum of lmost 15 percent in 1999 nd declined somewht therefter. This response pttern ws prticulrly pronounced on plots tht were mended with orgnics nd compost, suggesting tht the effect of mendments ws most drmtic in the first few yers following tretment. The pttern ws lso more pronounced on plnted plots, prticulrly for seeded species. Totl vegettion cover is still substntilly below the 50 percent cover tht is typicl of djcent undisturbed vegettion. Initilly, trnsplnts ccounted for most of the cover. After 10 yers, however, the totl cover of plnts tht germinted from seed both intentionlly seeded nd volunteer (11 percent) pproximted tht of trnsplnts (12 percent) on plots tht were plnted. Similrly, seeded plnts were initilly much more bundnt thn plnts tht volunteered (plnts tht germinted from seed in the soil seedbnk or tht dispersed nturlly onto the site). After 10 yers, cover of volunteers ws only slightly lower (5 percent) thn cover of seeded plnts (6 percent) on seeded plots. On plots tht were not seeded nd trnsplnted, volunteers ccounted for ll the vegettion cover. Non-ntive species were bsent. Tretment effects Restortion tretments vried in effectiveness (fig. 8). Mere closure ws not successful. On plots tht were not scrified, men cover never exceeded 1 percent nd ws only 0.4 percent 10 yers fter closure. Scrifiction lone ws slightly more successful (fig. 9). Plnt cover ws significntly greter on plots tht were scrified (but not seeded, mended with orgnics, or mulched) thn on control plots tht were closed Tble 2. Vrition in plnt cover (percent) over the 10-yer study All plnts (totl cover) Trnsplnts Seeded plnts Volunteer seedlings b For trnsplnts nd seeded plnts, cover is the men of those plots tht were plnted. b Volunteer seedlings cover is derived from ll plnt cover on unplnted plots, nd from the cover of seedlings of species tht were not seeded on plnted plots. USDA Forest Service RMRS-GTR

12 Figure 8. Vrition in effectiveness of tretments 2 yers fter tretment. Tble 3. Effect of scrifiction on plnt cover, repeted mesures nlysis of vrince results. Effect df F p Scrifiction (S) * Yer (Y) Interction (S*Y) * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypothesis tht cover would increse with scrifiction. Figure 9. Effect of scrifiction (but no other tretments) on plnt cover. to use but received no restortion tretments (tble 3). However, differences were smll. 10 yers fter closure, men plnt cover on scrified-only plots ws less thn 3 percent. The plnting (both trnsplnting nd seeding) tretment hd the most pronounced effect on plnt cover (tble 4). Mgnitude of effect vried significntly with yer since plnting (tht is, the interction between plnting nd yer ws significnt). Consequently, the significnce of tretment effects ws ssessed t the end of the study, in 2005 (tble 5). 10 yers fter plnting, cover ws more thn three times greter on plnted plots Tble 4. Effects of plnting, soil mendments, nd mulching on plnt cover, repeted mesures nlysis of vrince results. Effect df F p Plnting (P) <0.01* Soil mendment (S) <0.01* Mulch (M) P*S P*M S*M Yer (Y) <0.01* Y*P <0.01* Y*S <0.01* Y*M * Denotes sttisticlly significnt difference (α = 0.05). Higher order interctions were not significnt nd re not shown. 8 USDA Forest Service RMRS-GTR

13 Tble 5. Effects of plnting, soil mendments, nd mulching on plnt cover in Effect df F p Plnting (P) <0.01* Soil mendment (S) * Mulch (M) P*S P*M S*M * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypotheses tht cover would increse with tretment. Figure 11. Effect of soil mendments on plnt cover. Figure 10. Effect of plnting on plnt cover. (19 percent) thn on plots tht were not plnted (6 percent) (fig. 10). Soil mendments were lso effective, but to lesser degree (tble 4). Agin, mgnitude of effect vried with yer since tretment, being lest pronounced in the first yers following closure (fig. 11). 10 yers fter tretment, plnt cover on plots tht received orgnics nd compost mendments ws significntly greter thn on plots tht received no mendments (djusted Dunnett s multiple comprison, t = 3.34, p = 0.01). The plnt cover on plots tht were mended with orgnics only ppered greter thn on plots tht received no mendments, but differences were not sttisticlly significnt (djusted Dunnett s multiple comprison, t = 1.41, p = 0.16). Differences between the orgnics nd compost nd orgnics only plots were lso not sttisticlly significnt (djusted Tukey-Krmer multiple comprison, t = 1.93, p = 0.09). Mulching with biodegrdble mt did not hve significnt effect on plnt cover, over the entire length of the study (tble 4), for 2005 (tble 5), or for ny other yer (fig. 12). Figure 12. Effect of mulching on plnt cover. Effects on Trnsplnt Cover Trnsplnts differed from plnts tht grew from seed, both in temporl ptterns of response nd in the effectiveness of the restortion tretments. Although there ws nnul vrition, the totl cover of trnsplnts fter 10 yers ws comprble to their cover immeditely fter trnsplnting (fig. 13). As reported in detil in Cole nd Spildie (2006), this temporl pttern resulted from growth in trnsplnt size offsetting the mortlity of individul trnsplnts. In the first 7 yers of the study, 32 percent of the trnsplnts died; however, the men re of survivors incresed 39 percent. Growth forms vried in response. The cover of smll trees incresed most, where they were plnted, while the cover of shrubs decresed most (fig. 13). As Cole nd Spildie (2006) report, most plnted trees (79 percent) survived, nd the size of the trees incresed gretly (men increse in re of 243 cm 2 ). Grminoids survived USDA Forest Service RMRS-GTR

14 Figure 13. Chnge in cover of trnsplnts since trnsplnting: ll plnts nd growth forms. most frequently (87 percent); they lso grew substntilly (men of 56 cm 2 ). Forbs survived less frequently (72 percent) nd grew more modestly (men of 56 cm 2 ). Shrubs, in contrst, hd poor survivorship (45 percent) nd experienced little growth (men increse in re of 0 cm 2 ). Tretment effects Beyond closure nd trnsplnting, other restortion tretments hd reltively little effect on plnt cover. Mulch hd no effect on trnsplnt cover (tble 6). Trnsplnt cover did vry significntly mong the soil mendments nd there ws no interction between yer nd tretment (tble 6). Overll, trnsplnt cover on plots mended with orgnics nd compost ws significntly greter thn on plots tht were only scrified (djusted Dunnett s multiple comprison, t = 2.46, p = 0.03). As Cole nd Spildie (2006) report, mendments incresed trnsplnt growth rtes but not survivl. During the period of time between bout two nd 6 yers fter trnsplnting, it ws possible to visully identify the composted plots by the lrge stture of mny of the trnsplnts. However, the mgnitude of effect declined Tble 6. Effects of soil mendments nd mulching on trnsplnt cover, repeted mesures nlysis of vrince results. Effect df F p Soil mendment (S) * Mulch (M) S*M Yer (Y) <0.01* Y*S Y*M * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypothesis tht cover would increse with tretment. Figure 14. Effect of soil mendments on trnsplnt cover. towrd the end of the 10-yer study (fig. 14). In 2005, plots mended with orgnics nd compost did not hve significntly more trnsplnt cover thn plots tht were only scrified (djusted Dunnett s multiple comprison, t = 0.89, p = 0.31) nd these plots could no longer be esily identified visully, on the bsis of trnsplnt stture. Grminoids were the only growth form for which trnsplnt cover ws significntly higher on orgnics nd compost plots thn on unmended plots (djusted Dunnett s multiple comprison, t = 2.49, p = 0.04). Cole nd Spildie (2006) report tht grminoid trnsplnt growth ws lso gretest on orgnics nd compost plots. Responses of individul species Despite smll smple size, some insight into the vrible response of individul species cn be gined (tble 7). All seven of the Achille millefolium trnsplnts died. Otherwise, survivorship vried between 43 percent nd 100 percent. Note tht becuse only trnsplnt cover ws ssessed in 2005, survivorship nd chnge in size of individul trnsplnt cn only be estimted through Of the most bundnt species, survivorship ws notbly high (100 percent) for Juncus prryi nd Crex rossii. It ws notbly low for Vccinium scoprium (45 percent) nd Phyllodoce empetriformis (50 percent). Of the eighteen species tht survived to 2002, eight species grew substntilly (increse in re of more thn 20 percent), while six species declined substntilly in size. The net effect of survivorship nd growth is reflected in cover vlues in tble 7. Cover incresed over the 10-yer period for nine species nd decresed for 10 species. The species tht incresed most were trees nd grminoids: Pinus contort, Abies lsiocrp, Clmgrostis cndensis, Juncus prryi, nd Crex rossii. The species tht decresed most were Vccinium 10 USDA Forest Service RMRS-GTR

15 Tble 7. Responses of individul trnsplnted species. Survivl b Survivor re (cm 2 ) Cover d (%) Species Number (%) Abies lsiocrp Achille millefolium Antennri lpin Antennri lnt Aster lpigenus Clmgrostis cndensis Crex rossii Dnthoni intermedi Festuc viridul Hypericum formosum Juncus prryi Luzul hitchcockii Oyzopsis exigu Phyllodoce empetriformis Pinus contort Polymonium pulcherrimum Sibbldi procumbens Spire betulifoli Vccinium scoprium Number of individuls of ech species tht were originlly trnsplnted. b Percentge of originl trnsplnts tht survived until c Men re of those originl trnsplnts tht survived, in 1995 nd d Men cover of ll plots trnsplnted with the given species. scoprium, Phyllodoce empetriformis, Dnthoni intermedi, Aster lpigenus, nd Achille millefolium. For the species tht trnsplnted successfully, plnts survived nd grew well whether soils were mended or not. Nevertheless, in the cse of both Juncus prryi nd Crex rossii, growth rtes were significntly greter on plots mended with either orgnics or orgnics nd compost, in comprison to unmended plots (Cole nd Spildie 2006). Of the species tht trnsplnted lest successfully, it ws possible to ssess the effect of soil mendments on Vccinium scoprium nd Phyllodoce empetriformis. Soil mendments hd little effect on the trnsplnting success of P. empetriformis. For V. scoprium trnsplnts, soil mendments ffected trnsplnt survivl but not growth (fig. 15). Survivl ws substntilly greter on plots tht received either of the soil mendment tretments, strting the fourth yer following trnsplnting. Although differences between tretments were not significnt when the entire length of the experiment is exmined (Wilcoxon chi-squre = 2.37, p = 0.15), they were significnt in 2002 (Person chi-squre = 4.62, p = 0.05), the lst yer trnsplnts were counted. Despite incresed survivorship, men cover of V. scoprium trnsplnts in 2005 ws not significntly greter on orgnics nd compost plots (2 percent) thn on unmended plots (1.3 percent) (djusted Dunnett s multiple comprison, t = 0.56, p = 0.17) Figure 15. Effect of soil mendments on survivorship of Vccinium scoprium. Effects on Seedling Cover In contrst to the totl cover of trnsplnts, the cover of plnts tht grew from seed (seedlings) doubled over 10 yers (fig. 16). However, this increse ws not consistent. Seedling cover incresed lmost threefold in the first 4 yers following closure, declined substntilly for the next 3 yers nd then incresed slightly between 2002 nd Both seeded species nd volunteers exhibit this temporl pttern of response, lthough the increse in cover of volunteers hs been more slow nd stedy. Cover of seeded species fter 10 yers ws only USDA Forest Service RMRS-GTR

16 Figure 16. Chnge in seedling cover since cmpsite closure nd seeding. slightly greter thn it ws the first summer fter seeding. Cover of volunteers ws three times greter in 2005 thn it ws in In 1996, the cover of seeded species on plots tht were seeded ws 3.5 times the cover of volunteer species. In 2005, cover of seeded species ws only 1.3 times the cover of volunteers. As reported in detil in Cole (in press), the density of seeded species ws gretest in 1996, while the density of volunteers ws gretest in Consequently, increses in cover reflect increses in the size of seedlings tht more thn offset seedling mortlity. Most seedling cover is provided by grminoids, while shrub nd tree seedling cover is miniml (fig. 17). This prtilly reflects the fct tht shrub nd tree species were not seeded nd tht only grminoids were seeded on two of the sites. But grminoids nd forbs were lso the most bundnt volunteer seedlings. Both grminoids nd forbs incresed in cover over the 10 yers, but cover of tree seedlings declined. Although the cover of wht we clled shrub seedlings incresed, mny of these seedlings my hve sprouted from roots rther thn reproduced from seed. Consequently, we re hesitnt to try to describe the response of shrub seedlings beyond noting tht they remin negligible, even 10 yers fter closure. The vrible temporl pttern of incresing cover is lrgely explined by the response of grminoids. Only one non-ntive individul estblished on these plots, nd it only survived for two sesons. Non-ntive species re lrgely bsent t the elevtion where these cmpsites re locted; they re bundnt long trils nd on cmpsites t lower elevtions (below 2000 m). Tretment effects Scrifiction hd smll but positive effect on seedling cover. As noted erlier, plnt cover on plots tht were only scrified (ll seedlings) ws significntly greter thn on plots tht were not scrified. Cole (in press) reports tht 7 yers fter closure, plnt density on scrified plots ws more thn three times the density on plots tht were not scrified. In terms of cover, however, the mgnitude of difference ws never more thn couple percent. The more elborte tretments vried substntilly in effect. Appliction of the biodegrdble mulch mt did not hve cler effect on the cover of seedlings. In contrst, seeding hd pronounced effect (tble 8). Although mgnitude of effect declined somewht over the 10-yer study, seeded plots hd two times more seedling cover in 2005 thn plots tht were not seeded (fig. 18). Results re less conclusive regrding the effect of plnting tretments on the cover of volunteer seedlings. One might expect either incresed success for volunteers on plots with trnsplnts due to fcilittion Tble 8. Effects of seeding, soil mendments, nd mulching on seedling cover, repeted mesures nlysis of vrince results. Effect df F p Seeding (SD) <0.01* b Soil mendment (S) * b Mulch (M) SD*S SD*M S*M Yer (Y) <0.01* b Y*SD Y*S <0.01* Y*M Figure 17. Chnge in seedling cover of different growth forms since cmpsite closure. * Denotes sttisticlly significnt difference (α = 0.05). Higher order interctions were not significnt nd re not shown. b One-tiled test of hypothesis tht cover would increse with tretment. 12 USDA Forest Service RMRS-GTR

17 Figure 18. Effect of plnting on seedling cover. Figure 20. Effect of soil mendments on seedling cover. (remember tht seeded plots were lso trnsplnted) or decresed success due to competition. Volunteer cover ws consistently higher on plots tht were not seeded, strting the third yer fter closure (fig. 19). However, this difference ws not sttisticlly significnt over the entire length of the study (repeted mesures nlysis of vrince, F = 1.95, p = 0.24) or in 2005 (univrite nlysis of vrince, F = 0.37, p = 0.57). Therefore, we cnnot confidently conclude tht plnting decresed the cover of volunteer species. However, we lso cnnot be confident tht plnting does not hve n dverse effect, given the results illustrted in figure 19. More detiled exmintion of the dt indictes tht volunteer cover ws much higher on unplnted plots on only two of the cmpsites. Cole (in press) reported tht the density of volunteers did not differ between plnted nd unplnted plots, suggesting tht if there is n dverse effect of plnting on volunteers, it is reduction in size rther thn density of plnts. Amending soils lso hd positive effect on seedling cover. The effectiveness of soil mendments vried over the course of the study (fig. 20), being lest effective in the erly yers nd most effective 4 or 5 yers fter tretment. Becuse the interction with yer since tretment ws sttisticlly significnt (tble 8), soil mendment effects re only evluted in 2005, the finl yer of the study (tble 9). In 2005, men seedling cover on plots with orgnics nd compost mendments ws lmost three times the seedling cover of unmended plots (djusted Dunnett s multiple comprison, t = 2.37, p = 0.03). Cover on plots with the orgnics only tretment ws not significntly different from either unmended plots (djusted Dunnett s multiple comprison, t = 1.06, p = 0.13) or the orgnics nd compost plots (djusted Tukey- Krmer multiple comprison, t = 1.31, p = 0.20). Responses of different species Both seeded species nd volunteers responded positively to the orgnics nd compost soil mendment (tble 10). Generlly, grminoids nd forbs responded positively to soil mendments, while tree seedlings did not (tble 10). Of the seeded species, only Sitnion hystrix did not estblish well initilly. It estblished t low densities Tble 9. Effects of seeding, soil mendments, nd mulching on seedling cover in Effect df F p Seeding (SD) <0.01* Soil mendment (S) * Mulch (M) SD*S SD*M S*M Figure 19. Effect of plnting on cover of volunteers. * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypotheses tht cover would increse with tretment. USDA Forest Service RMRS-GTR

18 Tble 10. Men (s.e.) seedling cover in 2005, 10 yers fter tretment, on plots with vrious soil mendments. Soil mendment Orgnics None Orgnics nd compost Seeded species 2.7 (1.3) 5.1 (2.3) 11.2 (4.5) Volunteers 3.0 (0.7) 4.4 (0.7) 6.7 (1.4) Grminoids 2.9 (0.7) 4.2 (0.8) 7.9 (1.6) Forbs 0.4 (0.2) 1.6 (0.9) 3.8 (1.4) Tree seedlings 0.5 (0.1) 0.4 (0.1) 0.3 (0.1) Tretments tht differ significntly from untreted plots. on only two of six seeded plots nd ws bsent on ll plots 10 yers lter (tble 11). The two dditionl seeded grsses, Dnthoni intermedi nd Phleum lpinum, estblished in bundnce initilly, but provided miniml cover 10 yers lter. D. intermedi survived on only one of six seeded plots, while P. lpinum survived on 10 of 18 seeded plots. Neither of these species is well represented in the undisturbed djcent vegettion, however. In contrst, the two most common grminoids in the undisturbed vegettion, Juncus prryi nd Crex rossii, incresed in cover over the 10-yer period (tble 11). J. prryi ws seeded on three of the cmpsites. It volunteered on ll cmpsites nd on 38 of the 54 plots where it ws not seeded. Although the density of seeded J. prryi declined from mximum of 251 seedlings in 1996 to only 36 seedlings in 2002 (s reported in Cole in press), mortlity ws more thn offset by growth of those plnts tht survived. 10 yers fter tretment, J. prryi seedling cover on seeded plots ws 4 percent. Along with the 4 percent cover of J. prryi trnsplnts, totl cover of this species exceeded its men cover of 6 percent in undisturbed vegettion within severl yers. On sites tht were not seeded or trnsplnted, men J. prryi cover ws 1.1 percent. Although J. prryi trnsplnts responded positively to soil mendment nd Cole (in press) reports tht the density of J. prryi seedlings ws higher on mended soils, the cover of J. prryi seedlings did not vry significntly mong soil mendments (repeted mesures nlysis of vrince, F = 0.22, p = 0.81). In 2005, the men cover of J. prryi seedlings ws 2.6 percent on unmended plots, 1.9 percent on orgnics plots, nd 3.6 percent on orgnics nd compost plots. Crex rossii volunteered on 69 of the 72 plots tht were t lest scrified. Within 3 yers, C. rossii cover exceeded the 1 percent cover typicl of undisturbed sites. 10 yers fter closure, men C. rossii cover ws 3.1 percent. C. rossii cover responded positively to both soil mendments. In 2005, men cover of C. rossii ws 1.2 percent on non-mended plots, 3.2 percent on orgnics plots, nd 4.7 percent on orgnics nd compost plots (djusted Dunnett s multiple comprison, t = 2.18, p = 0.05 for the difference between non-mended nd orgnics plots nd t = 3.37, p < 0.01 for the difference between non-mended nd orgnics nd compost plots). Differences between orgnics nd orgnics nd compost plots were not sttisticlly significnt (djusted Tukey s multiple comprison, t = 1.17, p = 0.25). Forbs responded less vribly. Aster lpigenus nd Sibbldi procumbens both estblished in modest numbers on virtully ll the plots on which they were seeded. Density of these species hs declined precipitously (Cole in press), but cover in 2005 is still bout 50 percent of their mximum cover (reched the first yer fter seeding) nd not different from typicl cover on undisturbed sites. Antennri lpin nd Penstemon globosus estblished in much greter bundnce nd, despite declining density, hve incresed in cover over the 10 yers. Although smple size is too smll for sttisticl nlysis, both of these species pper to hve responded Tble 11. Men cover (percent) t the end of ech growing seson for seeded species nd the most bundnt volunteers Antennri lpin seeded (n=12) Antennri lpin volunteer (n=5) Antennri lnt seeded (n=6) Aster lpigenus seeded (n-6) Dnthoni intermedi seeded (n=6) Juncus prryi seeded (n=17) Juncus prryi volunteer (n=37) Penstemon globosus seeded (n=6) Phleum lpinum seeded (n=18) Sibbldi procumbens seeded (n=11) Sitnion hystrix seeded (n=2) Crex rossii volunteer (n=69) Men cover on plots where the species estblished (n). 14 USDA Forest Service RMRS-GTR

19 Figure 21. Abundunt lrge Penstemon globosus seedlings on plot mended with orgnics nd compost, 6 yers fter tretment. fvorbly to the soil mendments. In 2005, men cover of A. lpin ws 0.6 percent on non-mended plots, 2 percent on orgnics plots, nd 5.5 percent on orgnics nd compost plots. In 2005, men cover of P. globosus ws 0.5 percent on non-mended plots, 6.2 percent on orgnics plots, nd 15.0 percent on orgnics nd compost plots (fig. 21). Antennri lnt estblished t more modest densities but lso incresed in cover. It lso ppers to hve responded fvorbly to soil mendments. In 2005, men cover of A. lnt ws 0.5 percent on non-mended plots, 1.2 percent on orgnics plots, nd 1.4 percent on orgnics nd compost plots. Seedling Estblishment nd Mortlity Additionl insights into seedling response to restortion cn be glened from the more frequent nd detiled ssessments of seedlings tht were conducted the first 3 yers following restortion. Seedlings becme estblished throughout the two-month ssessment period (erly July to erly September), but two-thirds of the seedlings becme estblished (cotyledons were welldeveloped) in erly August bout one month fter snowmelt. On seeded plots, the number of newly estblished seeded species declined gretly ech yer, from 521/m 2 the first yer, to 159/m 2 the second yer nd 44/m 2 the third yer (fig. 22). The number of newly estblished volunteers declined from 30/m 2 the first yer, to 24/m 2 the second yer, nd 18/m 2 the third yer. Mortlity rtes during the growing seson, once seedlings were estblished (hd well-developed cotyledons), were very low for the first two sesons (less thn 1 percent for seeded species nd 5 to 12 percent for Figure 22. Seedling density (#/m 2, in squres) nd survivl rtes (in tringles) of seeded species nd volunteers from cohorts of seedlings tht estblished in 1996, 1997, nd Densities re for erly nd lte summer of ech yer nd survivl for intervening periods. volunteers) (fig. 22). Overwinter mortlity rtes were higher (22 to 33 percent for seeded species nd 15 to 42 percent for volunteers). Mortlity of seeded species ws much higher during the third growing seson (45 percent), the first hot, dry summer without supplementl wtering (supplementl wtering occurred the first summer; the second summer ws cool, with frequent precipittion). Mortlity of volunteers during the third growing seson ws lso higher thn in erly yers but much less thn for seeded species (22 percent). Only 33 percent of the estblished seedlings of seeded species survived three growing sesons, while 53 percent of the estblished seedlings of volunteers survived. Consequently, the proportion of seedlings tht were volunteers incresed from 5 percent fter the first growing seson to 14 percent t the end of the third growing seson. This proportion hd incresed to 20 percent fter 7 yers. For the first three sesons it ws possible to distinguish tretment effects on seedling estblishment rtes from effects on mortlity rtes. In ech of the first three sesons, the number of seedlings tht estblished ws much greter on seeded plots thn on plots tht were not seeded (tble 12 nd fig. 22), but neither the mulch nor the soil mendments ffected ntlity rtes. USDA Forest Service RMRS-GTR

20 As noted before, mortlity rtes were generlly low the first two sesons. Mortlity rtes were significntly higher on plots tht were not plnted for the first 2 yers (tble 13), reflecting the fct tht mortlity rtes were higher for volunteers thn for seeded species those yers (fig. 22). In 1998, however, the summer ws hot nd dry nd plnts were not wtered. Mortlity rtes were much higher tht summer (fig. 22). Under these conditions, seeded plots experienced much greter mortlity (33 percent) thn plots tht were not seeded (17 percent) (tbles 13 nd 14). Higher mortlity on seeded plots did little to offset the greter estblishment rte on seeded plots. Seedling density on seeded plots ws more thn five times the density on non-seeded plots fter three growing sesons (Cole in press). Plots mended with orgnics nd compost experienced much lower mortlity rtes (13 percent) thn plots mended with orgnics only (23 percent) nd plots without mendments (33 percent) (tble 13 nd 14). As reported by Cole (in press), seedlings growing in plots mended with orgnics nd compost grew much more rpidly thn seedlings tht did not receive this tretment. Both root nd shoot biomss of seedlings ws significntly greter in plots mended with orgnics nd compost, both in 1996 nd in 1997 (tble 15). Tble 12. Effects of seeding, soil mendments, nd mulching on seedling ntlity (density of newly estblished seedlings) in ech of the first three yers fter tretment Effect df F p F p F p Seeding (SD) <0.01* * 8.9 <0.01* Soil mendment (S) Mulch (M) SD*S SD*M S*M * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypothesis tht ntility would increse with tretment. Tble 13. Effects of seeding, soil mendments, nd mulching on seedling mortlity in ech of the first 3 yers fter tretment Effect df F p F p F p Seeding (SD) * * * Soil mendment (S) * Mulch (M) SD*S SD*M S*M * Denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypothesis tht mortlity would decrese with tretment. Tble 14. Men (s.e.) percent mortlity of seedlings in ech of the first 3 yers fter tretment. Soil mendment Seeding tretment Orgnics nd None Orgnics compost Not seeded Seeded (3) 14(4) 12(4) 16(3) 8(2) (5) 3(3) 12(6) 14(4) 3(1) (7) 23(4) 13(3) 17(3) 32(6) Tretments tht differ significntly from untreted plots. 16 USDA Forest Service RMRS-GTR

21 Tble 15. Root nd shoot biomss, men (s.e.) on plots with different soil mendment tretments. Becuse we mpped ll seedlings tht estblished for the first 2 yers, it ws lso possible to ssess the effect of trnsplnts on seedling estblishment nd survivl. As noted erlier, the density of volunteers on trnsplnted plots ws not significntly different from density on plots tht were not plnted. This suggests tht trnsplnts hve little effect on seedling estblishment nd survivl, either positive or negtive. To explore reltionships further, we compred seedling density within the cnopy of trnsplnts to density outside the cnopy. Two yers fter seeding, seedling densities were significntly lower within the cnopy (pired t-test, t = 2.35, p < 0.01). We then ssessed whether seedlings tended to estblish close to, or fr from, trnsplnts. This ws ccomplished by clculting the distnce from seedlings to the nerest trnsplnt, s well s the men distnce from 100 rndom points to the nerest trnsplnt. The differences between these distnces ws clculted nd verged. On verge, seedlings were locted 1.1 cm closer to trnsplnts thn expected, but this difference ws not sttisticlly significnt (one-smple t-test, t = 1.72, p = 0.10). Effects on Species Richness Soil mendment Orgnics nd None Orgnics compost 1996 root biomss (mg) 1.7 (0.3) 1.9 (0.4) 3.4 (0.7) 1996 shoot biomss (mg) 3.7 (1.1) 6.1 (1.6) 8.6 (2.0) 1997 root biomss (mg) 30 (14) 38 (3) 112 (29) 1997 shoot biomss (mg) 71 (42) 62 (18) 231 (66) Amended plots tht differed significntly from non-mended plots. 10 yers fter tretment, the men 1 m 2 plot hd four species. At the scle of the entire cmpsite, men species richness for the six closed cmpsites ws 13 species. This is comprble to the men of 15 species found on undisturbed reference sites. Tretments vried gretly in their effectiveness in restoring species richness. Plots tht were scrified (but not treted in ny other wy) hd significntly more species (men of 1.8 species, s.e. = 0.5) thn plots tht were not scrified (men of 0.7 species, s.e. = 0.3) (one-tiled t-test, t = 1.83, p = 0.05). Of the more elborte tretments, only plnting ws effective in incresing species richness (tble 16). At the plot level of nlysis, men (s.e.) species richness ws 2.2 (0.5) species on unplnted plots nd 6.3 (0.4) species on plnted plots. At the cmpsite level of nlysis, species richness ws 5.0 (0.7) species on unplnted plots nd 11.2 (1.4) species on plnted plots. Progress in Reltion to Reference Conditions Clerly, progress hs been mde in restoring both the soil nd vegettion on these cmpsites. However, degree of progress vried mong tretments nd mong sites, s well s mong the mesures of progress tht we employed. To ssess progress, it is helpful to compre conditions on the closed cmpsites to cmpsite conditions prior to cmpsite closure nd to reference conditions on undisturbed control sites. As illustrted in figure 7, mendment of cmpsite soil with orgnics nd compost resulted in ner-complete restortion of the soil chrcteristics we exmined. Soils on these plots did not differ significntly from undisturbed controls, while ll chrcteristics were higher thn on unrestored portions of the cmpsite. In contrst, plots tht were scrified only showed little recovery. Soils on these plots generlly differed significntly from undisturbed controls nd were not different from soils on unrestored portions of the cmpsite. Plots mended with orgnics only were intermedite, but more similr to those tht were scrified only. It is importnt to remember, however, tht if we hd ssessed physicl chrcteristics of the soil (such s porosity or infiltrtion rtes), both degree of progress nd the reltive success of the different tretments might be different. The 12 percent men vegettion cover on cmpsites, 10 yers fter closure, represents substntil progress in vegettion recovery compred to the 0 percent cover tht typified these cmpsites prior to closure. However, vegettion cover is still fr from the 50 percent cover tht is Tble 16. Effects of plnting, soil mendments, nd mulching on species richness in Effect df F p Plnting (P) <0.01* Soil mendment (S) Mulch (M) P*S P*M S*M * denotes sttisticlly significnt difference (α = 0.05). One-tiled test of hypotheses tht richness would increse with tretment. USDA Forest Service RMRS-GTR

22 Figure 23. Plot tht ws plnted nd mended with orgnics nd compost, with 50 percent cover fter 7 yers. typicl of undisturbed sites. Progress vried both mong cmpsites nd mong tretments. On plots tht were given the most effective tretments (scrified, plnted, nd mended with orgnics nd compost), men cover fter 10 yers ws 27 percent, bout one-hlf of the cover of reference sites. On one of the cmpsites, plots tht were given this tretment were t 50 percent cover fter 10 yers (fig. 23). In ddition to interest in incresing vegettion cover, we were lso interested in restoring vegettion similr in composition to undisturbed conditions. As illustrted in tble 17, most of the vegettion on the closed cmpsites ws herbceous. This contrsts shrply with composition of the groundcover on undisturbed sites. Shrubs ccounted for 57 percent of the cover on undisturbed sites but only bout 11 percent of the cover on the cmpsites in 2005 (fig. 24). In the undisturbed vegettion, grminoids ccounted for 26 percent of the cover; on the cmpsites they ccounted for 65 percent of the cover. Finlly forbs mde up 17 percent of the cover of undisturbed vegettion nd 24 percent of the cover on cmpsites. Cmpsite composition ws most similr to reference conditions immeditely fter the restortion tretments (tble 17 nd fig. 24). Dissimilrity incresed Figure 24. Reltive cover of growth forms on closed cmpsites nd undisturbed control sites. to mximum in 1999, when totl cover lso peked, nd hs declined since. Most tretments were not beneficil in restoring the ntive species composition. Trnsplnting ws the only effective wy of estblishing shrubs on the sites. However, even on plots tht were plnted nd received the orgnics nd compost tretment, the vegettion ws 12 percent shrubs, 53 percent grminoids, nd 35 percent forbs. Shrubs comprised bout 40 percent of the vegettion cover on plnted plots on two of the cmpsites (fig. 25), but were virtully bsent on two of the other sites. Of the 13 species with men cover of more thn 0.1 percent on undisturbed sites, only Crex rossii, Antennri lpin, nd Sibbldi procumbens hd similr cover levels when ll tretments re included (tble 18). On plots tht were plnted nd mended with orgnics nd compost, Juncus prryi, Pinus contort, nd Luzul hitchcockii lso were surviving t levels t lest equl to those on reference sites. In contrst, even on trnsplnted nd composted sites, Vccinium Tble 17. Vrition in cover of different growth forms on cmpsites over the 10-yer study Grminoids Forbs Shrubs Trees USDA Forest Service RMRS-GTR

23 exceeded those on reference sites for ll species other thn Gultheri humifus, Achille millefolium, nd Sitnion hystrix (tble 18). Forty-one different species pper to hve volunteered on closed cmpsites (tble 19). However, s mny s 10 of these species my be incorrectly clssified s volunteers, either becuse they were overlooked in trnsplnted plugs or becuse they sprouted from roots of estblished plnts. About hlf of the species tht volunteered were species tht we plnted on the sites. Twenty species volunteered only on closed cmpsites. On verge, we trnsplnted nd/or seeded six or seven different species on ech cmpsite. After 10 yers, the men number of species growing on cmpsites ws 13. This level of species richness is close to the men of 15 species growing on undisturbed control sites. Figure 25. Unusully luxurint growth of Vccinium scoprium nd Phyllodoce empetriformis trnsplnts fter 7 yers. scoprium nd Phyllodoce empetriformis hd men cover of 2.5 percent nd 0.4 percent, respectively. On reference sites, these species hve men cover of 27 percent nd 7.1 percent, respectively. Of the less common species, plnting resulted in coverges tht Discussion The tretments we pplied to these highly disturbed cmpsites were prtilly successful in restoring these sites. There hs been significnt progress in soil nd vegettion recovery, prticulrly on those plots tht were plnted nd mended with orgnics nd compost. Some soil impcts hve lso been mitigted. Vegettion Tble 18. Men percent cover of plnted species on ll cmpsite plots, plots tht were plnted nd mended with orgnics nd compost, nd undisturbed reference sites. Number Plnted nd Species of cmps All tretments composted Undisturbed Vccinium scoprium Phyllodoce empetriformis Juncus prryi Abies lsiocrp Pinus contort Crex rossii Antennri lnt Antennri lpin Festuc viridul Veronic cusickii Sibbldi procumbens Gultheri humifus Luzul hitchcockii Achille millefolium <0.1 Aster lpigenus <0.1 Clmgrostis cndensis <0.1 Dnthoni intermedi <0.1 Hypericum formosum 1 <0.1 <0.1 <0.1 Oryzopsis exigu <0.1 Polemonium pulcherrimum <0.1 Spire betulifoli <0.1 Penstemon globosus <0.1 Phleum lpinum <0.1 Sitnion hystrix <0.1 Species tht were unintentionlly nd non-systemticlly trnsplnted. USDA Forest Service RMRS-GTR

24 Tble 19. Species tht volunteered the number of cmps nd percent of plots where they estblished, s well s men percent cover, in Species Number of cmps Plot frequency Men cover Crex rossii Juncus prryi Muhlenbergi filiformis Vccinium scoprium Pinus contort Pice engelmnnii Antennri lpin Polygonum kellogii Crex pheocephl Crex micropter Abies lsiocrp 3 26 <0.1 Lewisi pygme 3 15 <0.1 Erigeron peregrinus 3 6 <0.1 Gyophytum rmosissimum 2 14 <0.1 Polemonium pulcherrimum 2 10 <0.1 Sibbldi procumbens 2 10 <0.1 Hiercium grcile 2 6 <0.1 Luzul hitchcockii 2 3 <0.1 Sprgue umbellt 1 10 <0.1 Clmgrostis cndensis 1 6 <0.1 Linnthstrum nuttllii 1 3 <0.1 Penstemon fruticosus 1 3 <0.1 Veronic cusickii 1 3 <0.1 Epilobium ngustifolium 1 1 <0.1 Festuc viridul 1 1 <0.1 Hypericum formosum 1 1 <0.1 Juncus mertensinus 1 1 <0.1 Ligusticum tenuifolium 1 1 <0.1 Oryzopsis exigu 1 1 <0.1 Penstemon globosus 1 1 <0.1 Phleum lpinum 1 1 <0.1 Polygonum phytolccefolium 1 1 <0.1 Spire betulifoli 1 1 <0.1 Aster lpigenus Antennri lnt Sgin sginoides Cstillej chrysnth Cerstium vulgtum Luzul cmpestris Roripp islndic Sitnion hystrix Plnts of these species my hve been trnsplnts or sprouts rther thn volunteers. cover hs incresed nd mny species hve been reestblished on the sites. However, totl vegettion cover remined diminished fter 10 yers (fig. 26). Even more problemtic is the miniml cover provided by shrubs, prticulrly Vccinium scoprium nd Phyllodoce empetriformis. Neither of these species responded well to the tretments we pplied. Trnsplnting helped, but more thn one-hlf of the shrub trnsplnts died nd the soil mendments did not result in incresed growth. Tretment Effects Most of the tretments contributed to recovery. Little recovery would hve occurred if cmpsites hd not been effectively closed to ll humn entry. Conversely, closure lone ws not very effective. Men cover on plots tht were closed to use but not scrified ws 0.5 percent fter 10 yers. Scrifiction ws effective in promoting recovery, but s with closure, it is not sufficient. Men cover on plots tht were only scrified ws 2.5 percent fter 10 yers. Mulch blnkets Surprisingly, the mulch blnkets hd no pprecible effect, despite the fct tht they re frequently employed nd recommended (Urbnsk nd Chmbers 2002) nd hve been shown to be effective in reserch conducted in other high-elevtion ecosystem types (Petersen nd others 2004). Fttorini (2001) reported tht mulch blnkets on ski runs in the Alps hd 20 USDA Forest Service RMRS-GTR

25 Figure 26. After 10 yers of recovery, cover hs incresed, but is still sprse. no effect on trnsplnt survivl. Their primry effect ws reduction in flowering. In our study, lck of effect my reflect the unusully good growing conditions tht existed for the first two summers. Survivl of both trnsplnts nd seedlings ws extremely high in these sesons. By the third summer, when hotter nd drier conditions previled, the mts hd lredy disintegrted. While they were not worthwhile under our working conditions, we re reluctnt to conclude tht they would not be worthwhile under other circumstnces. More reserch on mulch mts is needed. They cn lso hve other beneficil effects, such s deterring recretionl use of the site nd reducing erosion on steep slopes. Soil mendments The orgnics nd compost soil mendment ws clerly effective in ccelerting recovery. It ws effective in rising levels of orgnic crbon nd potentilly minerlizble nitrogen, nd incresing microbil ctivity in soils where these chrcteristics hd been dversely ffected by decdes of disturbnce nd lck of orgnic inputs (fig. 7). These chnges, perhps long with other unmesured chnges (such s bulk density, porosity, nd wter holding cpcity), likely explin the higher levels of vegettion cover on plots given this mendment. Numerous other studies, conducted in diverse ecosystems (such s Crvc nd others 2002), hve reported positive effects of compost on restortion success. This mendment ws more effective in promoting the cover of seedlings thn of trnsplnts. Our creful monitoring of seedling estblishment nd mortlity, long with our root nd shoot biomss mesurements, show tht while this mendment hd no effect on germintion nd estblishment rtes, it benefited seedlings by mking them less prone to mortlity fter they hd become estblished. For the first 2 yers, t lest, seedlings growing in soil mended with orgnics nd compost hd much lrger shoots nd root systems thn seedlings growing in soil tht did not receive this mendment (fig. 27). This suggests tht much of the beneficil effect of this tretment cme from the nutritionl input of the compost. The effect of the dditionl nutrients in the compost on trnsplnts ws visully obvious, prticulrly 2 to 5 yers fter trnsplnting. As reported in Cole nd Spildie (2006), trnsplnts growing in plots mended with orgnics nd compost were unusully lrge. By 2005, presumbly becuse the dditionl nutrients supplied by the compost hd been depleted, trnsplnts on composted plots were no longer visully lrger. Perhps the most importnt long-term beneficil effect of this tretment ws incresed survivorship for Vccinium scoprium trnsplnts. Reestblishment of V. scoprium is both the most importnt nd the most chllenging spect of restoring these sites. Any tretment tht hs positive effect is importnt. Results regrding the effectiveness of the mendment of soils with locl orgnic mtter, pet moss, nd Figure 27. Unusully lrge Phleum lpinum seedlings on plots mended with compost, 2 yers fter tretment. USDA Forest Service RMRS-GTR

26 ntive soil inoculum (but no compost) re less conclusive. This mendment ws n ttempt to reintroduce ntive soil biot nd meliorte some of the dverse physicl properties of cmpsite soils, by incresing wter holding cpcity nd reducing susceptibility to compction but to not meliorte the reduced levels of plnt-vilble nitrogen on cmpsite soils. For virtully every mesure of soil chrcteristics nd vegettion response, vlues for plots with this mendment were intermedite between those for plots tht were not mended nd plots tht were mended with orgnics nd compost. In 2005, totl cover on orgnics plots ws 1.3 times the cover on unmended plots. Seedling cover ws 1.6 times s high, while trnsplnt cover ws not different. These results re in line with our expecttions. However, for most of our mesures of response, differences between this tretment nd the other two tretments were not sttisticlly significnt t n lph level of For effects on totl cover nd seedling cover, p-vlues were 0.16 nd 0.13, respectively. Therefore, we cnnot conclude with high degree of confidence tht ddition of locl orgnic mtter ws beneficil. Conversely, the power of sttisticl tests ws low, given our smll smple size nd substntil vrition mong the six cmpsites. Consequently, we lso cnnot conclude with high degree of confidence tht the incresed cover observed on plots tht received the orgnics mendment is not the result of the tretment. The beneficil effects we cn be most confident bout (differences were sttisticlly significnt t n lph of 0.05) re incresed orgnic crbon nd totl nitrogen in the soil, survivl of Vccinium scoprium trnsplnts, growth of Crex rossii trnsplnts nd density of Crex rossii seedlings. Seeding nd trnsplnting Both seeding nd trnsplnting were effective. On plots tht were both seeded nd trnsplnted, men cover of volunteers fter 10 yers ws 3.9 percent. Seeding dded nother 6.3 percent, while trnsplnting dded nother 12 percent for totl of 20.2 percent cover on trnsplnted nd seeded plots. On plots tht were not seeded nd trnsplnted, men cover fter 10 yers (ll volunteers) ws 5.8 percent. Trnsplnts hd little effect, either positive or negtive, on seedlings when plnted t the densities t which we plnted them. The fct tht seedling density ws lower within trnsplnt cnopies suggests tht trnsplnting might inhibit seedling estblishment if plnted more densely. Tretments nd recovery rtes Tble 20 provides mens for compring the effectiveness of different tretments in vrious combintions. It ssumes liner extension into the future of the recovery rtes tht Tble 20. Estimted time (in yers) to recovery of 50 percent vegettion cover, ssuming liner extrpoltion of the pst 10 yers. Yers to recovery Closure scrifiction mulch mt orgnics compost 56 + seeding 31 + trnsplnting 19 Trnsplnting, but not seeding 36 On plots with orgnics nd compost mendments. occurred over the pst 10 yers, reporting the number of yers it would tke to restore the trget condition of 50 percent vegettion cover. Although future recovery rtes re unlikely to be equivlent to rtes during the first 10 yers, the tble emphsizes the lengthy recovery periods required regrdless of tretment, s well s the need to pply intensive tretments if we intend to restore these sites within our lifetime. By working the soil, recovery time cn be reduced to bout 50 yers, while with the ddition of plnting it cn be reduced to bout 20 yers (tble 20). Of the plnting tretments, seeding is more effective thn trnsplnting in reducing recovery time where soils re mended. In the bsence of soil mendments (tble 21), trnsplnting is more effective thn seeding in reducing recovery time; however, in the bsence of soil mendments, recovery time exceeds 30 yers even with both seeding nd trnsplnting. Our evlution of tretment effectiveness chnges, however, if our criterion for success is the reestblishment of ntive species composition. Tretments were much more effective in restoring species richness thn in restoring cover. The men number of species on cmpsites ws 13, close to the 15 species tht is typicl of undisturbed sites. Seeding nd trnsplnting were by fr the most effective tretment. Scrifiction hd smll but sttisticlly significnt positive effect, but mulching nd mending soils hd no effect. Tble 21. Estimted time (in yers) to recovery of 50 percent vegettion cover, without soil mendments, ssuming liner extrpoltion of the pst 10 yers. Yers to recovery Closure + scrifiction seeding 84 + trnsplnting 34 Trnsplnting, but not seeding 4 22 USDA Forest Service RMRS-GTR

27 Tretments were less effective in restoring the ntive species composition of these sites thn in restoring cover. After 10 yers, shrub cover ws only 11 percent, compred to 57 percent on reference sites. The most bundnt species on reference sites, Vccinium scoprium nd Phyllodoce empetriformis, hd men cover on cmpsites of only 2.5 percent nd 0.4 percent, respectively. Of the tretments, trnsplnting ws by fr the most effective for estblishing these species. The only P. empetriformis plnts on the cmpsites fter 10 yers were trnsplnts. We observed few smll V. scoprium plnts tht were not trnsplnts (probbly stem sprouts rther thn seedlings), but virtully ll V. scoprium cover is provided by trnsplnts. In combintion with trnsplnting, the soil mendments my lso be effective becuse more V. scoprium trnsplnts survived on mended soils. Further Reserch It is importnt to remember tht these generl ctegories of tretment (such s plnting nd soil mendment) refer to very specific tretments. If we hd used different plnting tretments (different species, lrger or smller trnsplnts, more trnsplnts, more or less seed) results would hve been t lest quntittively different. Further experimenttion with plnting techniques would be worthwhile. Prticulrly useful would be experimenttion with plnting of estblished plnts grown from seed in nurseries. Theoreticlly it should be possible to reduce recovery time gretly by plnting trnsplnts t densities severl times greter thn the five to seven trnsplnt plugs per squre meter used in this study. Experimenttion with dense plntings of Vccinium scoprium nd Phyllodoce empetriformis, both of which survived no more thn hlf the time, would be prticulrly useful. Opportunities for further experimenttion with soil mendments re even more diverse. As noted erlier, mendments cn serve to meliorte the dverse effects of cmping both on soil physicl properties nd on nutritionl properties. Applying loclly-derived orgnic mtter of differing types nd quntities could be tried. Highly-decomposed orgnics might be prticulrly effective in incresing wter-holding cpcity nd reducing the tendency for scrified soils to become compcted by rinfll nd snow loding. Mterils such s long pine needles, twigs nd smll brnches might be effective in promoting wter infiltrtion into soils. Other sources of long-lsting supplementl nutrition, relesed in smll quntities, could be employed. For exmple, Pschke nd others (2000) report tht the orgnic fertilizer Biosol ws effective in promoting plnt growth on rodcuts. The depth of scrifiction nd depth to which mendments re mixed could lso be vried. Finlly, it would be worthwhile to replicte the tretments pplied in this study, in different plces, to ssess if the results re site-specific. Tretment Needs of Individul Species Individul species vried gretly in how esy they were to restore to reference conditions. This study provided substntil insight into the tretments necessry to effectively restore the most bundnt species. Grminoids Juncus prryi nd Crex rossii, the most bundnt grminoids, trnsplnted well. Every trnsplnt of these species survived nd grew substntilly (Cole nd Spildie 2006). J. prryi lso estblished well when seeded nd modest number of seedlings volunteered. These seedlings survived in sufficient quntities to pproximte trget conditions fter 10 yers. Although C. rossii ws not seeded, it volunteered profusely, exceeding trget conditions fter 10 yers. Results suggest tht trnsplnting my be unnecessry for these species nd even seeding my be unnecessry for C. rossii. However, C. rossii estblished nd survived much more frequently on plots with either of the soil mendments, suggesting tht mendment with orgnics is importnt if this species is left to reestblish on its own (fig. 28). Forbs The most bundnt forbs plnted were Antennri lnt, Antennri lpin, nd Sibbldi procumbens. Both A. lnt nd S. procumbens becme well-estblished only where they were plnted. Both Figure 28. Abundnt Crex rossii seedlings tht volunteered on plots mended with orgnic mtter. USDA Forest Service RMRS-GTR

28 Figure 29. Percent cover of the Pinus contort trnsplnt on this plot incresed 15-fold in 10 yers. species estblished well from trnsplnts nd seed, lthough A. lnt estblished prticulrly well from seed. Trnsplnting ws more effective for S. procumbens. A. lpin trnsplnts often did not survive or grow well. However, this species estblished prolificlly from seed. In fct, it volunteered often enough to suggest tht seeding my be unnecessry. For ech of these species, cover ws greter on mended plots thn on plots tht were not mended. Where they were plnted, S. procumbens reched its trget coverge even on unmended plots, A. lpin reched its trget only on mended plots (either mendment), nd A. lnt did not rech its trget on ny tretment. Veronic cusickii, the most bundnt forb not intentionlly plnted, estblished only in the plugs we trnsplnted on the cmpsites. Trees Most of the smll trees tht we trnsplnted survived nd grew well, prticulrly Pinus contort (fig. 29). Seedlings of P. contort, Abies lsiocrp, nd Pice engelmnnii volunteered in substntil numbers but growth rtes were so low tht cover fter 10 yers ws negligible. Soil mendments hd no effect on either trnsplnts or tree seedlings. Shrubs The most difficult species to estblish were the shrubs. It is quite likely tht none of the shrubs estblished from seed. We found six new shoots of Vccinium scoprium on plots tht hd not been trnsplnted, but it is likely tht these re root sprouts rther thn seedlings. Therefore, trnsplnting is criticl to restortion of shrub species. However, trnsplnt survivorship ws 50 percent or less for ll three shrub species, V. scoprium, Phyllodoce empetriformis, nd Spire betulifoli. Soil mendments incresed survivorship for V. scoprium, but not the other shrubs. Amendments hd no effect on shrub trnsplnt growth or cover. Clerly, identifiction of tretments tht cn increse survivorship nd growth of V. scoprium nd P. empetriformis trnsplnts is the most importnt reserch need. Trnsplnting t much greter densities thn we used might lso prove successful. Uncommon species None of the uncommon species plnted would hve estblished often on these cmpsites without plnting. Of these, Penstemon globosus ws by fr the most successful. It estblished profusely when seeded. It survived t higher densities nd grew more rpidly on plots tht hd been mended, lthough it survived t dequte densities on non mended plots. We seeded Phleum lpinum on three of the cmpsites. Lrge numbers of seedlings germinted nd estblished, but most of them hve eventully died. Still, in 2005, P. lpinum cover exceeded its trget cover. Another grss, Dnthoni intermedi, responded in similr mnner. Lrge numbers of seedlings estblished, but fter 10 yers, few were live. Smll quntities of the Aster lpigenus tht estblished from seed were still live in 2005, but none of the few Sitnion hystrix seedlings survived. By trnsplnting t covers tht exceeded their trget covers, Aster lpigenus, Clmgrostis cndensis, Dnthoni intermedi, Luzul hitchcockii, Oryzopsis exigu, nd Polemonium pulcherrimum ll exceeded trget covers fter 10 yers, despite low survivorship for L. hitchcockii nd poor growth for A. lpigenus. Hypericum formosum trnsplnts did very poor, but few survived fter 10 yers. None of the trnsplnts of Achille millefolium survived. This ws surprising, since A. millefolium is often weedy plnt. Mngement Implictions A Prescription for Cmpsite Restortion Our results suggest the following prescription should provide vegettive recovery of cmpsites in this vegettion type on time scle of decdes rther thn century or longer Effectively close the cmpsite to ll use. Rope off the perimeter nd post signs tht instruct people to sty off the site nd explin why. Scrify soils to depth of t lest 15 cm. Brek up ll clods to produce crumb texture. Amend soils with t lest 2.5 cm lyer of loclly collected, well decomposed orgnic mtter. Add n equivlent mount of compost. Alterntively, mix in smller mount of bioorgnic fertilizer. For exmple, 24 USDA Forest Service RMRS-GTR