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Infestations Within the Ozark and Ouachita National Forests

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Infestations Within the Ozark and Ouachita National Forests Forest Ecology and Management 259 (2010) 1938–1945 Contents lists available at ScienceDirect Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco Stand and individual tree characteristics associated with Enaphalodes rufulus (Haldeman) (Coleoptera: Cerambycidae) infestations within the Ozark and Ouachita National Forests L.J. Haavik *, F.M. Stephen Department of Entomology, 1 University of Arkansas, 319 AGRI, Fayetteville, AR 72701, USA ARTICLE INFO ABSTRACT Article history: Many oak decline events have been reported within the past century in the eastern U.S., and important Received 18 December 2009 causal factors often differ among them. Coincident with a recent decline event in upland oak-dominant Received in revised form 2 February 2010 forests of Arkansas, Missouri, and Oklahoma was an unexpected outbreak of a native cerambycid beetle, Accepted 4 February 2010 Enaphalodes rufulus (Haldeman), the red oak borer. A large range in estimates of oak mortality throughout affected forests was presumably due to variation in species composition, where oak-dominant areas Keywords: experienced the greatest mortality. We chose eight sites across the Ozark and Ouachita National Forests of Oak decline Arkansas, similar both topographically and by oak dominance, to determine if other stand or tree Cohort senescence characteristics were important factors in variation of E. rufulus infestations across these forests. At each site, Wood borer Physiological age we sampled 125 dead, declining or healthy host Quercus rubra L., northern red oak. We created an Classification tree estimate of the E. rufulus population level at each site during the recent outbreak using counts of dated larval gallery scars within a subset (n = 120) of all Q. rubra sampled (n = 976). We used classification tree partitioning to determine host tree characteristics that differed among dead, declining, and healthy Q. rubra. We also used classification tree partitioning, followed by logistic regression to determine stand characteristics that varied significantly among high, moderate and low infestation stands as well as between forests. Models indicated that trees which died were smallest, grew the least during the borer outbreak, and were apparently suppressed. These dying trees were likely poor competitors for resources, allowing neighboring survivors to experience a growth release during the E. rufulus outbreak. Larval survivorship was higher in trees which died, though larval densities were not greatest within these trees, which suggests that resistance in these individuals was compromised. At the stand level, differences between forests were apparently more important than those due to borer infestation. E. rufulus populations were higher at sites with lower Q. rubra basal area. This reduced basal area was likely a result of greater Q. rubra mortality at these sites during the borer outbreak in the early 2000s. ß 2010 Elsevier B.V. All rights reserved. 1. Introduction and Lachance, 1992; Thomas et al., 2002). Due to this complexity, several forest decline theories have arisen. In general these Many forest decline events have been reported throughout the theories encompass some combination of site influence or forest past three centuries on several continents (e.g. North America, history, climatic extremes and secondary invaders such as Europe, Australia, Sinclair, 1965; Kessler, 1992; Thomas et al., pathogenic fungi and/or insect attack (Manion and Lachance, 2002; Jurskis, 2005) affecting a variety of forest types either at the 1992). genus level (e.g. oak decline, Balch, 1927) or at the individual Upland oak-hickory forests in Missouri, Arkansas and Okla- species level (e.g. sugar maple decline, Minorsky, 2003). Search for homa experienced an oak decline event in the late 1990s and early primary causal mechanisms is complicated by the fact that decline 2000s affecting over 121,000 ha (300,000 acres) of public forest events arise from a complex of interacting biotic and abiotic factors land (Starkey et al., 2004). White oak species (Quercus section (Sinclair, 1965; Mueller-Dombois, 1987; Manion, 1991), and Leucobalanus) were less affected than red oak species (Quercus important factors often vary by specific decline event (Manion section Lobatae)(Starkey et al., 2004; Heitzman et al., 2007; Fan et al., 2008). A native cerambycid, Enaphalodes rufulus (Haldeman), the red oak borer, was an important secondary invader attacking * Corresponding author. Tel.: +1 479 575 3384; fax: +1 479 575 3197. members of the red oak group, primarily Q. rubra L., Quercus E-mail address: [email protected] (L.J. Haavik). coccinea Muenchh., and Quercus velutina Lam. (Stephen et al., 2001; 0378-1127/$ – see front matter ß 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2010.02.005 L.J. Haavik, F.M. Stephen / Forest Ecology and Management 259 (2010) 1938–1945 1939 Starkey et al., 2004; Fan et al., 2008). This decline event and and the USDA Forest Service websites. Potential sites were located concurrent E. rufulus outbreak were unique in two ways. To date, it using the following criteria: oak-hickory or shortleaf pine-oak is the only oak decline event in the eastern U.S. found to be dominant forest type according to southern forest inventory and associated with E. rufulus (Millers et al., 1989). E. rufulus densities analysis (USDA Forest Service, 2001), comparable topography (ridge were 10–100 times greater than anything previously reported for tops and adjacent north and south facing slopes), and accessibility by this species within other eastern U.S. oak forests (Hay, 1974; road (within 400 m of nearest road). We selected sites which were as Donley and Rast, 1984; Fierke et al., 2005). similar as possible in topography and dominant species composition History of these second growth forests may be important in the in an attempt to hold these variables constant so that importance of recent oak decline event, as much of Ozark and Ouachita upland other variables could be observed more clearly. Generally, we forests were clear cut in the early 1900s and regeneration occurred avoided prescribed burn areas, although this was more difficult in primarily through stump sprouting (Strausberg and Hough, 1997). the Ouachita National Forest, as most oak dominant stands were Fire suppression followed throughout much of the 20th century, included in these burn areas. We visited >100 potential sites located resulting in a current simplified canopy structure of even-aged by the GIS, and finally selected eight sites somewhat evenly stands reaching maturity with little oak regeneration in the distributed throughout these national forests, that exhibited a range understory (Strausberg and Hough, 1997; Soucy et al., 2005). of observable Q. rubra mortality. Four sites were located within the Mortality estimates for this recent Ozark/Ouachita Mountains oak Ozark National Forest and four were located within the Ouachita decline event vary substantially, and range from 2% to 100% of National Forest (Fig. 1). We collected data from three stands at each basal area, apparently depending largely on species composition of study site: one on the ridge top, and one each on the adjacent south stands. Red oak dominant stands experienced 30–100% mortality, and north facing slopes. At one site in the Ouachita National Forest, whereas all stands on average, regardless of species composition, Fork Mountain, the north facing slope consisted of 100% Q. rubra experienced 2–15% mortality (Starkey et al., 2004; Guldin et al., mortality, so we collected north facing slope data from a nearby 2006; Heitzman et al., 2007). Some variation in mortality both at slope, Rattlesnake Mountain, which was 3 km north of Fork the stand and individual tree level may be due to factors other than Mountain. We collected individual tree data (described in Section species composition. Fierke et al. (2007) found that Q. rubra 2.2)from35Q. rubra at each stand, or from all live Q. rubra present, mortality and E. rufulus infestation varied by aspect as well as 105 trees per study site, and 20 dead standing or fallen Q. rubra species richness, though that study did not compare stands with per site. Overall sample sizes were: 815 live and 161 dead Q. rubra. similar aspects and different levels of E. rufulus infestation. Climate in the Ozark/Ouachita region is temperate with hot Many forest decline theories predict that advancing stand age is summers and mild winters. Mean January temperature is 4 8C, of primary importance in a decline event (Sinclair, 1965; Mueller- mean August temperature is 27 8C, and the annual mean is 16 8C. Dombois, 1987; Manion, 1991). Oak et al. (1991) found that a ratio Most precipitation occurs during spring and fall, totaling 124 cm in of site index to actual tree age as a measure of physiological age the Ozark Mountains and 150 cm in the Ouachita Mountains was a better predictor of stand maturity and susceptibility to oak (NCDC, 2009). Rock formations of limestone, sandstone and shale decline than actual tree age. If age-related changes in host comprise much of the Ozark Plateau, which is characterized by susceptibility were an important factor in E. rufulus population deep valleys, steep ledges and cliffs with elevations up to 750 m increase, then stands and/or trees which were old physiologically with slopes facing all cardinal directions (Adamski et al., 1995). The may have been the most likely to experience high borer Ouachita Mountains are oriented such that ridges run east to west populations. with maximum elevations reaching 790 m, where upper slopes are The objective of this study was to determine if particular tree or steep, gradually leveling off at lower elevations into U-shaped stand characteristics differed among areas with higher E. rufulus valleys (Guldin, in review). Soils in both forests are generally rocky, infestation density compared to those with lower populations. acidic and clay-rich, with low organic matter content (Adamski Specific characteristics measured at the stand level were mean et al., 1995).
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