Determinants of Tree Susceptibility to Attack by the Red Alder Bark Beetle, Alniphagus Aspericollis (Leconte) (Coleoptera: Scolytidae)
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Determinants of tree susceptibility to attack by the red alder bark beetle, Alniphagus aspericollis (LeConte) (Coleoptera: Scolytidae) Thesis by: Tristan Takaro Under the supervision of: Dr. Allan Carroll and Dr. Peter Marshall In partial fulfillment for the requirements of the degree of Bachelor of Science in Forest Sciences The University of British Columbia Faculty of Forestry Department of Forest Sciences 2013 (Submitted April 22nd, 2013) Table of contents - Introduction …………………………..............………… Page 3 - Materials and Methods …………………..............……... Page 9 o Study area ………………………..............……... Page 9 o Plot selection and description ..................……..... Page 9 o Bark beetle attack census …................................. Page 10 o Statistical analysis …………..............……..…… Page 11 - Results ……………………..............…………………… Page 12 o Study area ………………………………………. Page 12 o Bark beetle attack census ………………………. Page 13 - Discussion …………………..............………………….. Page 16 Index of tables and figures - Table 1 ….......................................................................... Page 11 - Table 2 …………………............................…………….. Page 14 - Table 3 ……………..............…..............……..……..….. Page 14 - Table 4 ………..............…………................………..….. Page 14 - Figure 1 ………..............……..…..............…………..…. Page 10 - Figure 2 ………....................……..............…………..…. Page 15 - Figure 3 ……....................……..............……………..…. Page 15 - Figure 4 …………………………………………………. Page 16 Abstract The immeasurable effect that bark beetles can have on forest ecosystems provides justification for the close study of bark beetle populations. Bark beetles that attack hardwoods have not been as extensively studied as those that attack coniferous species, and the red alder bark beetle in particular has received very little attention. Alniphagus aspericollis is a bivoltine “secondary” bark beetle that specializes on red alder. In this observational study in Vancouver, British Columbia several tree characteristics are assessed for their relationship with the likelihood of the tree being attacked by the bark beetle using logistic regression analysis. It was found that diameter at breast height (DBH) did not significantly affect the likelihood of attack, and that the level of injury of a tree is a good predictor of its likelihood of being attacked by the bark beetle. Keywords: bark beetle, secondary, Vancouver, British Columbia, injury, injuries, likelihood, diameter, DBH. Introduction Bark beetles are widely recognized as one of the most prominent and pervasive disturbance agents of the temperate forests of the world (Avtzis et al. 2012). They have been particularly destructive in North America, which is the native home to a majority of the tree-killing species of bark beetles (Stark 1995). They can devastate entire landscapes of trees during population outbreaks, causing drastic ecological and economic ramifications (Simard et al. 2012). Even when the trees are not killed, as commonly occurs with low population densities of several bark beetle species, they are commonly weakened by the attacks of the beetle which predisposes them to death by windthrow or pathogens. The immeasurable effect that bark beetles can have on forest ecosystems provides justification for the close study of bark beetle populations. The abundance of suitable host material (in terms of stand basal area and the abundance of large-diameter trees) and the stress level of the stand have been cited as key factors affecting the susceptibility of stands to bark beetle infestations (Simard et al. 2012). Among other factors, the density of attacks on an individual susceptible tree is related to the distance from that tree to a tree with emerging beetles (Cates and Alexander 1982). For the mountain pine beetle (Dendroctonus ponderosae Hopkins) tree mortality can be a result of stress associated with mechanical damage from construction, logging and vandalism among other stress-linked factors (Haverty et al. 1998). The plant stress hypothesis and the environmental constraint hypothesis both predict that changes in plant defenses of stressed plants cause decreased resistance of those plants to insect attack (Bleiker et al. 2005). During population outbreaks bark beetles in general are known to preferentially attack larger-diameter trees (Simard et al. 2012; Cates and Alexander 1982), but some species behave differently. The majority of bark beetles are known to preferentially attack weakened trees but the opposite can also occur in the case of population outbreaks of “primary” bark beetles (Avtzis et al. 2012). Bark beetles are considered to be “primary” if they preferentially attack and kill vigorous trees (Marchetti et al. 2011). A bark beetle is considered “secondary” if it is most often observed to reproduce in trees that are weakened or recently killed or in slash (Teen 2012). High stand and individual tree susceptibility to bark beetle attack has been found to be closely linked with defoliation of the stand or tree, and this pattern has been observed in several bark beetle- conifer systems (Bowers et al. 1996). In a 2009 study on a secondary bark beetle known as Pseudips mexicanus, every attacked tree was observed to have some amount of scarring, a broken top, or a dwarf mistletoe infection (Smith et al. 2009). Bark beetle species that infest coniferous trees have been more heavily studied than those that affect hardwood species, possibly due to the fact that the majority of phloemophagous bark beetle species are exclusive to conifers, as are the majority of tree-killing species (Ohmart 1989). This could be attributed to the qualitative complexity of induced responses of angiosperms to inoculation by fungal associates of bark beetles and the effectiveness of this response, or to the greater difference between costs and nutritive benefits of attacking hardwoods than conifers (Ohmart 1989). One hardwood-attacking bark beetle, the banded elm bark beetle (Scolytus schevyrewi Semenov), was found to have similar attack success as measured by the density of exit holes in 10-cm and 24-cm (diameter at breast height, DBH) bolts of Siberian elm, Ulmus pumila L. (Lee et al. 2011). S. schevyrewi is native to Asia, was first observed in North America in 2003, and has since expanded to occur in 28 US states and four Canadian provinces (Lee et al. 2011). A native elm bark beetle, Hylurgopinus rufipes Eichhoff, was found to occur in very low densities on trees with < 10 cm DBH (Anderson and Holliday 2003). H. rufipes is one of the most destructive hardwood bark beetles because it is a vector for Dutch elm disease (Anderson and Holliday 2003), and S. schevyrewi may be a vector for it as well (Lee et al. 2011). Interestingly Anderson and Holliday also found that the densities of entrance holes was very low in trees with DBH < 10 cm but densities plateau at approximately 3 holes per 100 cm2 (300 per m2) once DBH reaches about 20 cm (Anderson and Holliday 2003). They also found that overwintering beetles were almost always found in the lower 25 cm of the bole, likely due to the thicker bark there providing more protection from the weather (Anderson and Holliday 2003). Yet another elm bark beetle, Scolytus kashmirensis, has been observed to prefer trees that are older and taller for feeding (Buhroo 2012). In a study on the dieback of ash in Slovakia bark beetles of the Hylesinus genus (fraxini and crenatus species) were found to preferentially attack ash trees that were weakened by a pathogen that causes patches of bark necrosis (Kunca et al. 2011). Red alder is a common, short-lived hardwood tree in the Pacific Northwest. While individual trees can live for over 100 years, stands of red alder typically “break up” at around 50-70 years of age depending on site productivity (Deal 2006). In the Pacific Northwest mortality of red alder is said to increase rapidly in stands over 90 years old, and very few red alder trees remain once the stand has reached 130 years (Deal 2006). Red alder plays an important role as a both a nitrogen-fixer and a pioneer species that is good at colonizing new sites with exposed mineral soil. The species has traditionally been viewed as having low economic value relative to the species which eventually replace it with succession (Douglas-fir, Western hemlock, Sitka spruce, etc.) in the Pacific Northwest and it was widely considered to be a “weedy” species (Deal 2006). In a general technical report by the United States Department of Agriculture (USDA) published in 2006 prices of red alder sawlogs were said to equal or exceed those of Douglas-fir sawlogs (Deal 2006). While red alder lumber hasn’t been valued very highly in the past, demand for it has continually increased in the recent past (Deal 2006), providing economic incentive to understand more about the insects that infest it. Much of the study of the insects that infest red alder (Alnus rubra Bong.) has focused on defoliators while bark beetles have not received as much attention (Deal 2006; Niemiec et al. 1995; Markham and Chanway 1998; Myers and Williams 1987). A species of ambrosia beetle described as new in 1984 and named Gnathotrichus alniphagus was found on Alnus firmifolia in southwest Morelos, Mexico (Wood 1984). The alder bark beetle (Alniphagus aspericollis LeConte) is a specialist on red alder and has been found to occur throughout the range of its host (Borden 1969). The species was originally described under the genus Hylesinus by LeConte in 1876 (Chamberlin