INTRODUCTION in the eastern UP, but there was little evidence of decline or mortality related to fungal eech bark disease (BBD) has spread across CHAPTER 11. pathogens (Kearney 2006). Inventory data roughly half of the range of American beech indicate that more than 15 million merchantable () in North America since in B beech [>22 cm diameter at breast height (d.b.h.)] it was introduced into Nova Scotia around 1890 are present on forest land in Michigan (Heyd (Erlich 1932, 1934; Gwiazdowski and others Michigan: Spread of the 2005, McCullough and others 2001). Resource 2006; Houston 1994). The nonindigenous beech managers, including foresters and wildlife scale , fagisuga Lind., mediates Advancing Front and biologists, remain concerned about potential BBD BBD by piercing the outer bark of beech trees, impacts, but their ability to plan and prioritize Stand-Level Impacts facilitating entry of the nonindigenous fungi stand-level operations (e.g., harvest, presalvage, Neonectria faginata, which colonizes only Fagus regeneration) is limited by a lack of information (Project NC–EM–09–02) spp., or (synonymous with N. about BBD distribution, spread, and effects galligena), which occurs on a variety of hardwood (Ostrofsky and McCormack 1986). Previous species in North America and Europe (Castlebury efforts to estimate BBD spread and quantify and others 2006, Houston and O’Brien 1983). The Deborah G. MccullouGh its impacts have been conducted in Eastern fungi kill small patches of phloem and cambium, States, typically years after BBD establishment. JaMes b. Wieferich and as dead tissues coalesce, large branches and Differences in topography, climate, soils, and the trunk are girdled (Burns and Houston 1987, forest attributes between the Lake States region Ehrlich 1934). Terms used to depict the three and the Northeast could potentially affect BBD stages of BBD include the “advancing front” progression and impacts. where trees are infested by beech scale, the “killing front” where trees are dying from fungal This project built on previous efforts 125 infection, followed by the “aftermath forest” undertaken following initial identi cation of BBD characterized by beech mortality, infected “cull” in Michigan. Progression of the advancing fronts trees, and, often, dense thickets of beech sprouts in Lower and Upper Michigan were monitored (Houston 1994, Houston and O’Brien 1983, from 2005 through 2009 using an adaptive Shigo 1972). sampling method to delineate distinct beech scale infestations (Schwalm 2009, Wieferich and Presence of BBD in Michigan was rst noted others 2011) and iterative modeling processes to in 2000 in Mason County in the northwestern estimate spread rates of individual infestations. Lower Peninsula and Luce County in the eastern Results through 2009 indicated that spread Upper Peninsula (UP) (McCullough and others rates varied considerably among years and 2001, O’Brien and others 2001). As of 2003, among beech scale populations (Wieferich and beech scale was present in four counties in others 2011) but were consistently lower than northwest Lower Michigan and ve counties 126 Forest Health Monitoring SECTION 3 Chapter 11 to be infested with beech scale at a previously at apreviously scale beech with infested to be one were or found more beech If sites. infested farthest the from 5to 8km circles concentric in were established 1996). points Sampling Seber and (Thompson adaptive sampling using infestations scale beech individual delineated and Advancing Front the of Progression METHODS woody material. of coarse composition and amount the and regeneration, of composition species trees, overstory other and of beech condition current to document 62 sites original the we resurveyed addition, In rates. spread to assess and Michigan Upper and (BBD) Lower in disease bark of beech fronts advancing the delineate and to monitor continue 2003. in site any to BBD in attributable mortality or decline of beech evidence There little was 21 moderate had infestations. or sites heavy while infestations, 39 light 12had sites sites, in absent was scale 2003, beech 2006). In (Kearney heavy) and (absent, light, infestation scale beech levels of (low, three and moderate, high) and area basal levels of beech three represented Sites woody material. coarse and regeneration, overstory, the of condition and composition on data baseline to collect weresites established 62 impact 2002–03, In 2007). others and Morin 2003, others and (Grif n States Northeastern in observations from derived estimates published We monitored the advancing fronts of BBD fronts We advancing the monitored were project to this in goals primary Our rates were estimated and areas of beech scale scale of beech areas and wererates estimated Spread were annually. mapped infestations and were imported into ArcGIS into were imported of plots Coordinates infested). were heavily trunk of the (onewhitewashed or more aspects (3) scale); or of (4) beech (several patchy clumps scale); beech (2) low-density (scattered, trace (1) as: absent; ranked scale and beech examined were visually trees beech plots, all In measured. were encountered 10 trees trees), beech rst the (e.g., sites nonforested roadside in and access (>10trees limited with d.b.h.). sites cm In for overstory by recorded was species d.b.h. and present) (if were trees concentrated, infested and of where 10) beech factor established was area abasal with aprism using (determined infestation. asingle as were considered subsequently and coalesced some infestations spread, scale beech as Over time, or satellites. other infestation scale beech primary the and infestation of the edge the between habitat of uninfested km ≥20 was there if distinct to be were considered beech). Infestations (with sites of uninfested by abuffer surrounded was infestation the until repeated was process the and were revisited points to infested closest points uninfested the moved year, surveyor the Each on. and revisited, would not be area the torecorded ensure were points site, “no beech” infested of an km 8 were present within no beech If recorded. was point uninfested an out until farther established was point another point, uninfested In forested sites, a variable radius plot radius avariable forested sites, In ® 10.1 (ESRI 2012), 2012), (ESRI 10.1 infestations were calculated using minimum three transects (25.5 m by 1 m wide) in each site. convex polygons for each distinct infestation Decay classes for coarse woody material (CWM) and statewide. were de ned as: (0) fresh material with intact bark and no obvious decay; (1) bark sloughing Impacts of BBD to Date off or absent, but with solid inner sapwood; (2) The 62 sites with BBD impact plots, including some decay; small wood chunks break off under 34 sites in 7 counties in Upper Michigan and impact, but rm center; (3) decaying; loses form 28 sites in 14 counties in Lower Michigan, under impact; and (4) decayed and form lost were revisited in 2011–13 to assess impacts of (Kearney and others 2005). Regeneration plots BBD. Similar methods were used in 2002–03 were established equidistantly between the and 2011–13 to assess beech scale and beech center plot and each of the four subplots in the condition. The center plot and four subplots, cardinal directions in the sites. Seedlings (<30.5 18.3 m from the center in each cardinal direction cm tall), saplings (>30.5 cm tall; <2.5 cm d.b.h.) (all plots 7.3-m radius), were relocated, and GPS and recruits (>2.5 cm d.b.h.) were tallied by coordinates were recorded at the center plot. species within a 2.4-, 3.5-, and 7.3-m radius of Species, d.b.h., and a visual estimate of beech the plot center, respectively. scale abundance were recorded on trees in all ve subplots. Twelve additional beech trees RESULTS growing at equally spaced azimuths and within Progression of the Advancing Front 60 m of the perimeter of the four subplots were tagged in 2002 and were also reexamined. These From 2011 through 2013, we established a trees were originally selected using prioritized total of 544 sites (with beech) in 28 counties criteria: (1) largest tree with dead tissue or at in Lower Michigan and 9 counties in Upper 127 least one canker, (2) any tree (pole-sized or Michigan to monitor the advancing front. We larger) with a canker, (3) largest tree with C. examined 1,854 live beech trees in these sites; fagisuga present, (4) any tree (pole-sized or larger) d.b.h. of these beech ranged from 4.2 to 119.5 with C. fagisuga, or (5) the largest tree near the cm and averaged 30.0 ± 0.4 cm. There were 187 azimuth. We measured d.b.h., number of cavities sites with infested beech. On average, 86.0 ± and estimated crown dieback, transparency, and 1.6 percent of the beech trees within infested beech scale densities on each tree. Overstory plots had at least some beech scale. Across all species composition and tree and beech scale sites, beech made up 53 ± 1.1 percent of the total abundance were also recorded along three basal area (all species), which averaged 16.8 2 transects (each 25.5 m by 10 m) in the sites. ± 0.4 m /ha. Along with beech, sugar maple (Acer saccharum), red maple (Acer rubrum), and Frequency, species, size, and decay class of northern red oak (Quercus rubra) dominated coarse woody material were recorded along the overstory. 128 Forest Health Monitoring SECTION 3 Chapter 11 expanded to 7088 km 7088 to expanded in 2009, had expanded to 9200 km to 9200 2009, expanded had in encompassed <3010 km <3010 encompassed infestation Charlevoix-Crawford-Emmet The occupied <2 km <2 occupied County Midland in relatively new infestation population encompassing 255 km encompassing population satellite Adistinct 2005. since monitored been has infestation scale beech continuous to 14.8 <2 km/year. from ranged infestations 2013,through but rate of of the spread individual 2013 2005 from 3.1through and ±0.3 km/year average 2011 from an rate of 3.3 ±0.4 km/year at spread Lower Michigan in fronts Advancing Annual spread rates of the advancing front in in front advancing of rates spread the Annual UP. of the portion northwestern the in occur beech with areas a few scattered although points, by no-beech now is ringed Peninsula Upper central the in edge infestation of the leading encompassed at least 13 at least encompassed 530 km area in the UP, estimated at 6214 UP, the estimated in km area 2011. in scale-infested infestation The beech primary the with coalesced infestation This front. edge main of the leading west of the 45 km 2009, more than in County Menominee in 16 575 km More than (Kent Co.) found was infested. to be Rapids Grand center near anature in area small 2014, in early when a occurred detection recent infestation, which encompassed 7727 km 7727 encompassed which infestation, 2011). others and (Wieferich The Mason-Wexford 2009 through 2005 were from monitored that infestations scale beech four distinct survey had increased to km 287 increased had In the UP of Michigan, a single and nearly nearly and asingle of Michigan, UP the In In Lower Michigan, we continued to we to continued Lower Michigan, In 2 in Lower Michigan are now infested. now infested. are Lower Michigan in 2 when detected in 2011 in when detected but 2 in 2013. In addition, a a 2013. in addition, In 2 in 2011 in but had 2 by 2013. The most 2 in 2013. in The 2 was identi ed identi ed was 2 in 2013. 2013. in 2 in 2005, 2005, in 2

through 2013. 2013. through at spread 3.9 2005 from ±0.7infestation km/year 2011 between year 2013. and average, On the km/ ±0.8 to 2.0 2007 and 2005 between year km/ ±2.2 6.2 from have ranged Michigan Upper (e.g., snap). 11 beech Only 314 of the in beech broken had trunk the along 46 and but standing, were 80 dead fallen, and died 5had were alive, (79 percent) trees Most 2002. in were tagged that subplots of the perimeter the around trees (averagesites 13 of respectively). 7and percent, Lower Michigan in than 16 respectively) percent, (average sites of 12 and percent Peninsula Upper in higher levels were generally transparency and dieback Canopy died. had trees ofpercent the 8 only 2002, in scale beech had that sites in dead was by 2013. area basal Even beech the of percent <7 and trees beech of9 percent the only Lower Michigan, in 2013.in contrast, In were dead 49 trees of percent the 2002, in scale beech with were infested that sites Peninsula Upper the In dead. plots are our in area basal beech of 26 percent the and trees beech of the 23 percent Michigan; Upper in progressing is front (average d.b.h.). ±0.4 cm 30 The killing (average beech live d.b.h.) and cm ±0.8 34 of dead beech size the in difference little was by 2013. died had 62 sites the in There examined trees beech 18 1,440 of percent the Overall, infested. were heavily that sites 44 including but 62 sites, bythe 2013, 55 were sites infested, Date BBD to of Impacts We were able to nd 724 of the 744 beech We 744 of beech were the 724 able to nd 23 of present was in scale beech 2002–03, In Lower Michigan had died, while roughly 30 DISCUSSION percent of the 368 beech in the Upper Peninsula Spread rates of the advancing front of BBD were dead. In 2002, 500 of the tagged trees were in Michigan, monitored since 2005, have varied not yet infested by beech scale, but by 2013, only among years and among distinct beech scale 189 trees had not been colonized. Average d.b.h. infestations. Between 2005 and 2007, Schwalm of live and dead beech trees was 33.5 ± 1.0 cm (2009) reported populations spread at rates of 4.0 and 23.5 ± 4.7 cm in Lower Michigan and 34.3 ± and 1.5 km/year in Upper and Lower Michigan, 0.8 cm and 39.5 ± 1.1 cm in the UP, respectively. respectively. Wieferich and others (2011) Radial growth was higher for trees that were monitored 12 distinct infestations and reported uninfested in 2003 than for trees that were spread rates of individual infestations varying already colonized by beech scale in 2003 (2.5 ± from 1.0 to 8.0 km/year in Lower Michigan. 0.2 and 1.6 ± 0.2 cm, respectively). The highest spread rate was recorded when two Coarse woody material (CWM) was originally distinct infestations coalesced and encountered in every site in 2011–13, and subsequently expanded at a rate of 14.3 km/ 732 pieces were recorded. Beech made up 68 year. In Upper Michigan, Wieferich and others percent of the 310 fresh, identi able pieces, and (2011) reported the maximum spread rate of the beech CWM was most abundant in heavily single advancing front was 11.0 km/year. Our infested sites. Total CWM volume ranged from estimates indicate spread in the Upper Peninsula 7 to 312 m3/ha and averaged 74 ± 10.5 and has slowed, probably because nearly the entire 50 ± 8.4 m3/ha in sites in the Upper Peninsula range of beech has been colonized. Spread rates and Lower Michigan, respectively. Beech CWM in Lower Michigan remain highly variable, volume averaged 14.7 ± 3.2 m3/ha across all sites. however, which may reect the fragmented 129 distribution of beech and forested land in general A total of 4,978 overstory trees, representing in this part of the State. We expect spread rates 19 species, were measured in subplots and will slow as the advancing front reaches areas transects in the 62 sites in 2011–13. Beech and in the central and eastern Lower Peninsula sugar maple dominated the overstory. Other where beech volume is relatively low. Recent common species (from most to least dominant) identi cation of localized infestations in Midland included red maple, eastern hemlock (Tsuga and Kent Counties indicates long-range dispersal canadensis), paper birch (Betula papyrifera), and of beech scale continues, probably as a result of red oak. As in 2002, regeneration was dominated birds transporting eggs or crawlers. Uninfested by beech and maple. Beech accounted for only beech within 15 km of beech scale infestations 11 percent of seedlings but made up 63 percent should be treated as high-risk sites, and BBD of all saplings, followed by sugar maple, which impacts should be considered as management accounted for 18 percent of the saplings. plans are developed. 130 Forest Health Monitoring SECTION 3 Chapter 11 (DiGregorio and others 1999). others and (DiGregorio overstory the into maples of young sugar recruitment and growth radial increase can BBD from resulting gaps canopy indicate studies previous and sites, our in abundant consistently maple Sugar was die. beech overstory as change likely will composition overstory Michigan, Upper In none Lower were Michigan. present in and Peninsula, Upper the in sites three only in thicket beech asingle we observed common, were saplings 1984). beech Patterson Although 1995, and others and 2003, Kobe Twery maple (Hane red maple and sugar including saplings, and seedlings ofrates nonbeech survival and availability reduce light Thickets 2003, Houston 1994). 2003, Hane others and (Grif n stems beech of susceptible abundance the by BBD, caused trees increasing beech healthy relatively of rapidthe death previously Northeastern forests, may proliferate following 2003. in were infested that sites our to in what we observed comparable is rate 1992). Whitemore and mortality Krasny This 2012, wave Livingston and of BBD rst (Kasson the during died reportedly beech overstory of the or percent 50 more States, Northeastern In killed. been had we examined trees beech ofpercent the 2001), others and 25 butMcCullough almost 2005, others and (Heyd Kearny 2005, to 2002–03 BBD present was in attributable mortality beech Little advance. to continues front where killing the Peninsula, Upper the Beech thickets, which are common in many many in common are which thickets, Beech pronounced of most in BBD are Effects needed to understand why beech mortality rates rates mortality why beech needed to understand be 2001). will others study and (O’Brien Further Lower Michigan in to nd dif cult remains of Neonectria evidence while Lower Michigan, in than Peninsula Upper the in more frequently have observed been bodies Fruiting virulence. likely involves Neonectria The difference scale. by beech colonized been have beech of the or all nearly all sites, of our many in and Peninsula, Upper the present in been for long it has as as at least Lower Michigan present in been has scale Beech Lower Michigan. in than sites Peninsula Upper the in greater 1979). Lea and Mize 2005, others and 1934, Mencuccini (Erlich studies other in noted similarly been has scale by beech infested of trees 2003. Reduced growth in were infested that of trees those than 65 higher percent almost were 2003 in were not that infested of trees rates growth Radial growth. reduced radial has infestation scale beech indicate also data 2011). nuts (Hamelin of beech abundance Our with associated is reproduction bear indicate for States, example, Northeastern in Studies 2011, Houston 1990). and Tubbs (Hamelin succumb trees overstory as of mast production by reduced offset may be bene ts such although 1986, 2010), others Strojny others and and Morrison 2006, 1997, Anderson and Kahler others 1983, and (Davis Gilbert salamanders and mammals, for habitat numerousprovide birds, may changes These woody material. of coarse apulse and snags in increase an in resulted It is not clear why beech mortality is so much much so is mortality why beech not is clear It has Peninsula Upper the in mortality Beech sp. infection rates or or rates infection sp. remain relatively low in Lower Michigan despite Grif n, J.M.; Lovett, G.M.; Arthur, M.A.; Weathers, K.C. the presence of beech scale for more than 15 2003. 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