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Differential Dispersal of Hylobius Warreni Within Modified Forest

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Differential Dispersal of Hylobius Warreni Within Modified Forest BEHAVIOR Seeing the Forest Through the Trees: Differential Dispersal of Hylobius warreni Within Modified Forest Habitats 1 2 1,3,4 MATTHEW D. KLINGENBERG, NIKLAS BJO¨ RKLUND, AND BRIAN H. AUKEMA Environ. Entomol. 39(3): 898Ð906 (2010); DOI: 10.1603/EN08269 ABSTRACT Hylobius warreni Wood, also known as the Warren root collar weevil, is a ßightless insect that feeds on conifers throughout the boreal forests of Canada. Mature trees typically can withstand feeding, but larval feeding around the root collar may cause mortality to young trees. Recently, a large outbreak of mountain pine beetle (Dendroctonus ponderosae Hopkins) has killed a high proportion of mature lodgepole pine (Pinus contorta Douglas variety latifolia) across British Columbia, Canada. This raises concerns that adult weevils may migrate from mature forests with reduced host pools into adjacent young forests that had been salvaged and replanted. To study movement of these walking weevils in different habitat types, we constructed three research plots consisting of various combinations of live-, dead-, and mixed (i.e., live and dead)-tree habitats. We observed dispersal patterns of individually labeled insects using a novel insect trap attached to the base of trees. Approximately 35% of insects were recaptured over 1 mo. Weevils were least likely to be recaptured proximate to the release location when released in a habitat with dead trees. Movement rates therein were almost double the rates of insects moving through live- or mixed-tree habitats. Our Þndings support the hypothesis that H. warreni may disperse out of habitats with dead trees into areas with higher proportions of green trees. Our Þndings are discussed in the context of habitat discrim- ination and potential increases in herbivory by H. warreni in western Canada given salvage harvesting activities after outbreaks of mountain pine beetle. KEY WORDS Hylobius warreni, mark-recapture, mark-release, dispersal, matrix habitat Elucidating processes that contribute to spatial pat- other insects as well (Denno et al. 1995). Currently, terns of the dispersal of insect herbivores is a chal- for example, there is a large outbreak of mountain pine lenge, as dispersal may depend on multiple and inter- beetle, Dendroctonus ponderosae Hopkins, modifying acting factors, including the physiological status of the the forest landscape in British Columbia, Canada insects (e.g., reproductive status, energy reserves) and (Aukema et al. 2006). This outbreak has precipitated population density (e.g., migration, crowding). Hab- concern regarding how major, sudden habitat changes itat conditions, such as composition and quality of the may affect vertebrates such as woodland caribou Ran- habitat matrix or patches, may also affect insect dis- gifer tarandus caribou Gmelin (McNay et al. 2006), but persal (Nathan et al. 2003, Haynes and Cronin 2006). the effects on insect species are less well studied. The For example, within boreal forest ecosystems, habitat current study examines the effects of habitat quality alterations such as clear-cut harvesting can affect the on the spatial distribution and movement of Warren spatial distribution of insect assemblages (Welty and root collar weevil, Hylobius warreni Wood (Co- Houseweart 1985, Niemela¨ et al. 1993, Bengtsson et al. leoptera: Curculionidae). 1997, O¨ rlander et al. 1997, Helio¨la¨ et al. 2001, Lemieux Much of the natural history and ecology of H. war- and Lindgren 2004, Phillips et al. 2006). Species per- reni has been summarized by the pioneering work of turbations in forested environments are not limited to Warren (1960) and Cerezke (1994). In brief, H. war- anthropogenic impacts. For example, insect distur- reni are found throughout the boreal forests of North bances may affect the spatial or temporal dynamics of America, feeding on a variety of conifer tree species. Adults feed on coniferous foliage above ground. Fe- 1 Ecosystem Science and Management Program, University of males lay up to 25 eggs per year in the duff layer Northern British Columbia, 3333 University Way, Prince George, BC around the root collar of trees. Upon eclosion, larvae Canada V2N 4Z9. feed on the phloem tissues of the root collar, protected 2 Department of Ecology, Swedish University of Agricultural Sci- ences, P.O. Box 7044, 750 07 Uppsala, Sweden. by a hard casing consisting of a mix of frass and tree 3 Department of Entomology, University of Minnesota, 1980 Fol- resin (Cerezke 1970). After 2 yr of feeding at the root well Avenue, St. Paul, MN 55108. collar, larvae pupate and adults emerge. In the spring 4 Corresponding author: Natural Resources Canada, Canadian For- months, adults feed nocturnally on young shoots of est Service, University of Northern British Columbia, 3333 University Way, Prince George, BC Canada V2N 4Z9 (e-mail: Brian. coniferous trees in preparation for mating and ovipo- [email protected]). sition that occur throughout the summer and fall June 2010 KLINGENBERG ET AL.: WEEVIL DISPERSAL IN MODIFIED FOREST HABITATS 899 (Cerezke 1994). Adults may live for up to 5 yr, and both interior hybrid spruce (Picea glauca ϫ engelman- remain ßightless throughout their entire life cycles nii) and Douglas-Þr (Pseudotsuga menziesii). Small (Cerezke 1994). gaps had formed where weevils had previously killed Forest health issues with H. warreni are predomi- young trees, although only live trees were used for the nately related to larval feeding. Adult feeding is not study. A trap was placed on each tree, and the contents detrimental to the tree host, but larval feeding may were emptied each morning. Weevils were returned inhibit growth, or even result in tree death, by girdling to the plantation near the bases of the trees where they of the stem and disrupting translocation (Cerezke were captured. After settling on a working prototype 1974). Although insects do not feed on dead trees, and Þnding no evidence of avoidance behavior larvae and pupae may mature on residual stumps after (Bjo¨rklund 2009), we collected 300 weevils from 8 harvesting, thus supplying weevils to the regenerat- May 2007 to 5 June 2007. Insects were stored in growth ing forest. Populations are frequently maintained chambers at 7ЊC to reduce the insectsÕ metabolism throughout the life of the stand (Cerezke 1973). Mor- until experiment initiation (Toivonen and Viiri 2006). tality to young trees by larval feeding may be exac- Labeling of Insects. Each weevil was labeled in the erbated by root deformation resulting from planting laboratory by etching the elytra, a technique used practices (Robert and Lindgren 2006), as well as by previously for other ground-dwelling arthropods root rots introduced at feeding scars (Whitney 1961, (Winder 2004). We had previously experimented with 1962). latex paint, but the paint rubbed off over time as the H. warreni appears to be an emerging concern in insects burrowed through leaf litter. Insects were se- forests in western Canada, particularly in areas with cured in petri dishes using plasticine (Flair Leisure extensive ongoing reforestation or regeneration after Products, Surrey, United Kingdom) before etching the current outbreak of mountain pine beetle. labels into the elytra with a high speed Dremel rotary Whereas a vast amount of research exists on the ecol- drill (engraving cutter bit number 106). To accentuate ogy and management of related Hylobius spp. such as markings, elytral etchings were traced using nontoxic Hylobius pales Herbst and Hylobius radicis Buchanan latex-based model paint (Humbrol Paints, Hornby in the northeastern United States (Corneil and Wilson Hobbies, Kent, United Kingdom). Each weevil was 1984, Hunt et al. 1993) and Hylobius abietis L. in labeled with a unique dot-and-number code repre- northern Europe (Nordlander et al. 1997, Leather et senting 0Ð99 in either blue, green, or yellow for tracing al. 1999, Bylund et al. 2004, Nordlander et al. 2005, to speciÞc release points. To investigate whether la- Wallertz et al. 2006), including studies on dispersal beling affected insect survival, 12 labeled and 12 un- patterns (Rieske and Raffa 1990, Eidmann 1997), little labeled weevils were placed in petri dishes (10-cm information exists on the movement or dispersal pat- diameter) containing clippings of lodgepole pine and terns of H. warreni (Schroff et al. 2006). In this study, moistened Þlter paper. New lodgepole pine branches we examine the dispersal patterns and movement rates were added every week, and Þlter paper was moist- of H. warreni through different environments contain- ened as necessary. Insect survival in the groups of ing live and dead trees using a label, release, and labeled and unlabeled insects was monitored over a recapture experiment using a novel trap. We hypoth- 2-wk period. esize that weevils may exhibit the highest movement The sexes of weevils released for the experiment rates through habitats with dead trees, which, in nat- were not identiÞed, because at the time of these ex- ural settings could complicate reforestation efforts as periments, a noninvasive technique for sexing had not a result of emigration from stands with high propor- yet been developed for this species (Cerezke 1994, but tions of pine killed by D. ponderosae. now see O¨ hrn et al. 2008). Dissections of 12 randomly selected weevils captured in the same forest used to obtain weevils for this experiment revealed approxi- Materials and Methods mately equal proportions of males and females (7:5), Trap Development and Collection of Insects. H. consistent with previous Þeld observations (Stark warreni were collected daily from a Ϸ15-yr-old pine 1959, Warren 1960, Cerezke 1994). forest near Prince George, British Columbia, Canada Plot Design and Insect Release. We established (53Њ55Ј14“N, 122Њ49Ј10”W), using a Bjo¨rklund funnel three experimental plots at the Prince George Tree trap attached to the base of each tree (Bjo¨rklund Improvement Station, British Columbia, Canada 2009). These traps exploit the insectsÕ nocturnal feed- (53Њ46Ј18“N, 122Њ43Ј4”W), with a mix of different hab- ing behavior and capture them alive, without chem- itat types.
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