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Take Me to Your Leader: Does Early Successional Nonhost Vegetation Spatially Inhibit Pissodes Strobi (Coleoptera: Curculionidae)?

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Take Me to Your Leader: Does Early Successional Nonhost Vegetation Spatially Inhibit Pissodes Strobi (Coleoptera: Curculionidae)? PLANTÐINSECT INTERACTIONS Take Me to Your Leader: Does Early Successional Nonhost Vegetation Spatially Inhibit Pissodes strobi (Coleoptera: Curculionidae)? JORDAN M. KOOPMANS,1 HONEY-MARIE C. DE LA GIRODAY,1,2 B. STAFFAN LINDGREN,1 1,2,3 AND BRIAN H. AUKEMA Environ. Entomol. 38(4): 1189Ð1196 (2009) ABSTRACT The spatial inßuences of host and nonhost trees and shrubs on the colonization patterns of white pine weevil Pissodes strobi (Peck) were studied within a stand of planted interior hybrid spruce [Picea glauca (Moench) Voss ϫ Picea engelmannii (Parry) ex Engelm.]. Planted spruce accounted for one third of all trees within the stand, whereas the remaining two thirds were comprised of early-successional nonhost vegetation, such as alder (Alnus spp.), paper birch (Betula papyrifera Marsh.), black cottonwood [Populus balsamifera ssp. trichocarpa (T. Ng.) Brayshaw], lodgepole pine [Pinus contorta (Dougl.) ex Loud.], trembling aspen (Populus tremuloides Michx), willow (Salix spp.), and Canadian buffaloberry [Shepherdia canadensis (L.) Nutt.]. Unlike the spruce trees, nonhost vegetation in the stand was not uniformly distributed. Spatial point process models showed that Canadian buffaloberry, paper birch, black cottonwoood, and trembling aspen had negative associa- tions with damage caused by the weevil, even though the density of the insectsÕ hosts in these areas did not change. Moreover, knowing the locations of these nonhost trees provided as much, or more, inference about the locations of weevil-attacked trees as knowing the locations of suitable or preferred host trees (i.e., those larger in size). Nonhost volatiles, the alteration of soil composition, and overstory shade are discussed as potential explanatory factors for the patterns observed. New research avenues are suggested to determine whether nonhost vegetation in early successional stands might be an additional tool in the management of these insects in commercially important forests. KEY WORDS nonhost volatiles, angiosperm volatiles, plantÐinsect interactions, Engelmann spruce weevil, spatial point processes The white pine weevil Pissodes strobi (Peck) (Co- chemical responses and genetic variation therein leoptera: Curculionidae), native to North America, is (King et al. 1997, 2004; Alfaro et al. 2000, 2008; Tomlin a herbivorous insect that reproduces in the terminal et al. 2000). Physical characteristics, such as thickness leaders of young spruce and pine. In the spring, fe- of the primary cortex and the size of trees and their males oviposit in leaders of host trees generally Ͼ5yr leaders, also inßuence host selection (VanderSar and of age (He and Alfaro 2000). Developing larvae girdle Borden 1977, Kiss and Yanchuk 1991, Turnquist and the shoots, often leading to creases, crooks, and forks Alfaro 1996, King et al. 1997, van den Driessche 1997, in the tree (Silver 1968). Although mortality of trees Manville et al. 2002). Spatial factors that potentially is uncommon, stem defects and associated growth inßuence insect distribution have received less atten- reductions may reduce lumber yield by as much as tion, however (He and Alfaro 1997). Some of these 40% in commercially important forests (Alfaro et al. factors affect the vigor of young spruce, including 1997). After a brief pupation within the leader, adults overstory composition, sun exposure, soil drainage, emerge between July and September to overwinter in and edge vegetation (Lavallee et al. 1996, Taylor et al. the duff layer or on the tree itself in coastal areas 1996, Simard and Hannam 2000). Certain properties of (Silver 1968). White pine weevils complete their life these factors may be exploited in silvicultural recom- cycle in 1 yr (Silver 1968). Infestations may continue mendations to reduce impacts of white pine weevil. in stands of spruce for up to 50 yr (He and Alfaro Inßuences of spatial distribution and diversity of 2000). nonhost species, and their impacts on the host-selec- Past research on the distribution and impact of tion behavior, distribution, and abundance of white white pine weevils on host trees has focused primarily pine weevil, has not yet been studied to our knowl- on defensive mechanisms of the trees, including edge. New spruce forests are typically established as the result of natural or anthropogenic disturbances 1 University of Northern British Columbia, 3333 University Way, and therefore may contain diverse early successional Prince George, British Columbia V2N 429, Canada. species (Denslow 1980), such as alder (Alnus spp.), 2 Natural Resources Canada, Canadian Forest Service, 3333 Uni- versity Way, Prince George, British Columbia V2N 4Z9, Canada. trembling aspen (Populus tremuloides Michx), willow 3 Corresponding author, e-mail: [email protected]. (Salix spp.), and paper birch (Betula papyrifera 1190 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 4 Marsh.), as well as various shrub species. Recent work Measurements for spruce and pine [lodgepole pine has suggested that the presence of nonhost vegetation Pinus contorta variety latifolia (Engelm.) ex S. Wats.] such as red alder (Alnus rubra Bong.) may affect the trees included age, root collar diameter, height, evi- levels of white pine weevil abundance (McLean 1989, dence of white pine weevil activity, and the presence Almond 2006). Increased plant diversity may decrease of other insects. The ages of spruce trees were esti- impacts by herbivores that exhibit narrow host mated by counting the number of whorls and con- breadths (Bianchi et al. 2006) but also may increase Þrmed by counting growth rings on a randomly se- impacts by more generalist herbivores (Koricheva et lected subset of trees destructively sampled poststudy. al. 2006). Root collar diameter was measured with a 150-mm In this study, we explore spatial factors that may caliper, and tree height was measured with a standard inßuence host selection and distribution of P. strobi. 8-m tape measure. Weevil activity was identiÞed by We Þt spatial point process models to a data set from the presence of oviposition punctures, fecal plugs, a population census of Ͼ1,000 trees in a stand con- emergence holes, larval cavities on broken leaders, taining moderate levels of white pine weevil. We seek and wilted (i.e., shepherdÕs crooks) or dead leaders to answer two questions regarding the pattern of wee- (Lavallee et al. 1996). In this paper, we refer only to vil distribution. First, does nonhost vegetation have those trees possessing dead or dying leaders as suc- either a positive or negative association with distribu- cessfully colonized. Four categories of weevil coloni- tion of white pine weevil attack? Second, if such an zation were identiÞed: 1, 2, and 3 yr of colonization or association exists, how does it compare with known never colonized. Spruce trees were judged to have predictors of weevil attack distribution, including multiple years of weevil colonization if they showed physical attributes of colonized spruce trees such as damage on the initial leader with subsequent attack of size? Expanding our understanding of insect interac- new shoots vying for apical dominance. An elevation tions with host and nonhost vegetation, in combina- grid was developed by recording elevation values at tion with existing knowledge of spruce defense mech- 10-m intervals along the transect and at two to four anisms, may yield new silvicultural tools for locations on the horizontal axis using a GPS (Meridian contending with this herbivore in commercially im- Gold Magellan; Thales Navigation, San Dima, CA). portant forests. Data were collected from 26 May to 17 June 2008. Data Analysis. Maps for the locations of all trees, including weevil-colonized trees, were created using the spatstat package v.1.13Ð3 in R v.2.6.2 (Ihaka and Materials and Methods “ ” Gentleman 1996, Baddeley and Turner 2005, R De- Site Selection and Data Collection. We censused a velopment Core Team 2008). Spatial trends in the x 0.154-ha stand of interior hybrid spruce with relatively and y directions of height and root collar diameter of even-aged trees displaying a moderate level of P. strobi colonized versus uncolonized spruce trees were stud- abundance near Prince George, British Columbia, ied using linear regression. Further effects of weevil Canada (53Њ52Ј N, 122Њ47Ј W). The spruce had been colonization status on height and diameter of spruce hand-planted and were 12 yr of age at the time of trees were studied by examining the signiÞcance of study. The stand also contained a variety of other such a term added to the prior regression models that nonhost vegetation from natural regeneration. With accounted for spatial trends. the exception of willow (Salix spp.), trees and shrubs Spatial point process models were used to examine were identiÞed to the species or subspecies level. For the effects of spatial trends (i.e., x and y, as well as the purposes of analysis, alder, which consisted of elevation), spatial attributes of host plants (i.e., height green alder [Alnus viridis ssp. crispa (Ait.) Turrill], and diameter at root collar), and locations of nonhost sitka alder [Alnus viridis ssp. sinuata (Regel) A. and D. plants on the prevalence of white pine weevil attack. Lo¨ve], and mountain alder [Alnus incana ssp. tenuifo- The response variable for each model, intensity (␭), a lia (Nutt.) Breit.] were grouped together at the genus spatially explicit estimated density of weevil attacks, level. Every live tree with Ͼ1-cm root collar diameter was measured as the number of trees with damaged was censused. (i.e., dead or dying) leaders per square meter. Co- Spatial location data for each tree was determined variates were converted to either density surfaces (for using a compass, vertex, and corresponding distance point processes such as locations of trees or shrubs) or transponder (Haglo¨f Sweden Vertex III v.1.4; Haglo¨f, a smoothed interpolation (for the marks of a point Madison, MS), with slope values and horizontal dis- process) before Þtting. Both types of surfaces incor- tance measurements corrected based on the internal porated a Gaussian kernel density smoother as a rep- clinometer.
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