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H.F. CEREZKE Abstract Resume WARREN ROOTCOLLAR WEEVIL, HYLOBZUS WARREN1 WOOD (COLEOPTERA: CURCULIONIDAE), IN CANADA: ECOLOGY, BEHAVIOR, DAMAGE RELATIONSHIPS, AND MANAGEMENT H.F. CEREZKE Canada Department of Natural Resources, Canadian Forest Service, 5320 - 122 Street, Edmonton, Alberta, Canada T6H 3S5 Abstract The Canadian Entomologist 126: 1383- 1442 (1994) Warren rootcollar weevil, Hylobius warreni Wood, is a pest of several conifer species and is distributed widely throughout the boreal forest in Canada. Literature on this weevil is reviewed and interpreted to provide comprehensive coverage of its ecology, behavior, and impacts, and the implications for its management. New information is provided to fill gaps in our knowledge of H. warreni. These include distribution and hosts, immature stages, sex ratios, and daily and seasonal activity patterns of adults. Sex ratios, abundance, longevity, and pedestrian dispersal behavior of adults within the forest and on trees are described from mark, release, and recapture studies conducted over several years using a newly designed interception trap. Daily trap captures are correlated with previous night temperatures and this relationship is a tool for predicting adult captures. Fecundity and oviposition behavior on the host tree are described from laboratory- and field-reared adults. Population levels of H. warreni are compared across a variety of lodgepole pine stands in Alberta. Weevil numbers within forests are described in relation to tree size and age, depth of duff material around tree bases, and stand density. Analysis of weevil feeding scars distributed at the root collar base of mature pine stems were used to describe the likely temporal pattern of population abundance and change during stand develop- ment. Information supportive of this pattern is presented from surveys in young and mature stands. A new method of assessing stands of lodgepole pine for suitability of habitat for H. warreni takes into account the cumulative nature of this weevil's injury and reflects its temporal pattern of success in relation to stand conditions. The effects of a pre- commercial thinning treatment on H. warreni populations and its girdling injury were investigated in a 25-year-old lodgepole pine stand over an 8-year period. Compared with control plots, thinning treatment caused a 5-fold increase in weevil numbers per tree and increased the tree attack incidence by 2-fold, but resulted in an average 6% reduction in partial stem girdling of attacked trees. Information about the effects of various forest management practices on H. warreni abundance and survival is reviewed and areas requiring research are identified. Cerezke, H.F. 1994.Le Charanqon de Warren, Hylobius warreni Wood (Coleoptera:Curculionidae) au Canada. The Canadian Entomologist 126: 1383- 1442. Resume Le Charan~conde Warren, Hylobius warreni Wood, parasite plusieurs espbces de conifbres et est bien rCpandu dans la for& borCale canadienne. Les travaux sur cette espbce ont CtC rCvisCs et interprCtCs dans le but d'obtenir le plus d'informations possible sur l'Ccologie, le comportement et les effets de ce parasite avant d7Ctablirdes programmes de contrble. De nouvelles donnCes viennent combler les lacunes dans nos connaissances de l'espbce: rkpartition des hbtes, stades immatures, rapports m2les:femelles, activitCs quotidiennes et saisonnibres des adultes. Les donnCes sur les rapports miiles:femelles, l'abondance, la longCvitC et le comportement de dispersion pCdestre des adultes dans la forEt et sur les arbres ont CtC obtenues par marquage-capture-recapture durant plusieurs anntes au cours desquelles un nouveau type de pibge a CtC utilisC. Les captures quoti- diennes dans les pibges se sont avCrtes en corrClation avec la tempCrature de la nuit preckdente et celle relation permet de prCdire les captures des adultes. La fCconditC et le 1384 THE CANADTAN~MoLOGIST November/December 1994, comportement de ponte sur les arbres hates sont dCcrites B partir de donnCes sur des adultes ClevCs en laboratoire et en nature. Les densitks de population d'H. warreni ont ttt comparkes au sein de plusieurs for& de pins de Murray en Alberta. Le nombre de charan~onstrouvts a CtC mis en relation avec la taille et l'2ge des arbres, avec la profondeur de la couche d'humus autour de la base des arbres et avec la densit6 de la forCt. L'analyse des cicatrices laissts par les charan~onsse nounissant B la base du tronc d'arbres B maturitt, B la naissance des racines, a permis de dCcrire la courbe probable d'abondance de la population en fonction du temps au cows du dCveloppement de la forCt. Le pattern obtenu est confirm6 par des donntes obtenues dans des for& jeunes et des for& B maturitC. Une nouvelle mCthode d'kaluation des forCts de pins de Murray comme hates de populations d'H. warreni tient compte de la nature cumulative des blessures causCes par le charan~onet reflkte son succ&sen fonction du temps en relation avec les conditions de la forst. Les effets d'un Cclaircissement prkliminaire de la forCt pour des fins commerciales sur les populations d'H. warreni et ceux des blessures d'annklation ont CtC Ctudits pendant 8 ans au sein d'une forCt de pins de Murray de 25 ans. La comparaison avec des for& tCmoins a rtvtlt que 1'Cclaircissement a entrain6 une augmentation considerable (X5) du nombre de LharanCons par arbre, a augment6 l'incidence des attaques aux arbres par un facteur de 2, mais a rksultt en une rkduction moyenne de 6% des anntlations partielles des tiges des arbres attaques. L'information sur les effets des pratiques d'am~na~ementdes for& sur l'abondance et la survie d'H. warreni est rtvisCe et les zones de recherche qui doivent Ctre approfondies sont indentifkes. [Traduit par la RCdaction] INTRODUCTION In North America, there are seven known species of weevils in the genus Hylobius. The following species occur in Canada: H. congener Dalla Torre, Shenkling, and Marshall, H. pales (Herbst), H. pinicola (Couper), H. radicis Buchanan, and H. warreni Wood. All attack and feed on coniferous hosts (Wood 1957; Warren 1958; Warner 1966). Warren rootcollar weevil, Hylobius warreni Wood, first described in 1957, is indigenous to coniferous forests in North America (Wood 1957; Warner 1966). Its larvae feed on the root and root collar region of trees, causing partial to complete girdling of roots and stem, large open wounds, and resinosis (Warren 1956b; Cerezke 1970~).Live healthy trees are susceptible to attack at a young age and these trees are subject to repeated attacks at irregular intervals through to stand maturity or harvest (Cerezke 1969, 1970~).Highest populations of the weevil and its incidence of attacks tend to occur on trees of vigorous crown classes and on high-productivity sites (Cerezke 1969, 1970b, 1970~;Ives and Rentz 1993). Trees may be killed directly from the girdling injury, and their wounds may provide entry courts for root and stem diseases (Warren 1956b; Whitney 1961; Cerezke 1974). Partially girdled trees may also accumulate losses in growth from repeated attacks and be made less windfinn because of weakened roots and stems (Cerezke 1974). Different habitat conditions are created during the process of forest removal and the early establishment of the new forest, and most of these conditions influence the population dynamics of H. warreni. The cut stumps left behind after harvesting allow established resident populations of H. warreni to complete their development, but the newly planted or naturally grown seedlings provide continuity of habitat as breeding sites for subsequent generations of the weevil (Cerezke 1973~).Thus early establishment of H. warreni into young stands is often assured at an age when the risk of tree mortality from larval girdling is maximal (Cerezke 1970c, 1974; Ives and Rentz 1993). Various silvicultural treatments are often applied in young stands to help achieve optimal growth. Studies in stands of white spruce [Picea glauca (Moench) Voss] and lodgepole pine (Pinus contorta Dougl. var. latifolia Engel.) in western Canada have shown that this weevil is distributed in relation to various site and stand parameters. Its abundance and survival can therefore be influenced Volume 126 THE CANADIAN ENTOMOLOGIST 1385 by harvesting practices and probably also by post-harvest treatments including prescribed bums, scarification, and stump removal (Warren 1956b, 1956d; Stark 1959~;Cerezke 1969, 1970a, 1970b, 1973~;Byford and McLean 1991). Such treatments as precommercial thinnings and spacings, stand fertilization, and alternate species selection may all influence H. warreni abundance and its injury (Goyer and Benjamin 1972; Wilson and Millers 1983; Selander and lmmonen 1992; Hunt et al. 1993). The earliest collections in Canada of adult H. warreni were probably made during 1930-1935, but concern for its injury to trees did not develop until the late 1940s and early 1950s (Daviault 1949). Early investigations in Saskatchewan and Manitoba were prompted by the concern that its larval wounds on white spruce provided courts of entry for root and stem diseases (Warren and Whitney 1951). Since 1949, a considerable body of information has accumulated on the biology, ecology, behavior, and tree damage relationships of H. warreni, especially in western Canada. Much of this information is contained in published reports, a number of unpublished file and internal reports, theses, and as unpublished data, much of which is not readily available. There is a need to compile and synthesize this information into a single document and thereby make it available to land managers and researchers. The objectives of this paper are to review all sources of information on H. warreni in Canada, present a synthesis and interpretation of the knowledge about this weevil, and formulate guidelines for coping with this insect in managed forests and plantations. The content of this document is arranged under four main sections. The first three sections (geographical distribution and hosts; life history and behavior; and population dynamics and impact) present the main body of scientific information; the last section focuses on forest management guidelines and research needs.
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