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FURNISS, M.M., LIVINGSTON, R.L., and MCGREGOR, M.D. 1981. Development of a stand susceptibility classification for Douglas-fir beetle, p. 115-128. In Hedden, R.L., Barras, S.J., and Coster, J.E., tech. coords. Hazard-Rating Systems in Forest Insect Pest Management: Symposium Proceedings. Gen. Tech. Rep. WO-27. Washington, DC: U.S. Department of Agriculture, Forest Service. FURNISS, R.L. and CAROLIN, V.M. 1977. Western forest insects. Misc. Publ. 1339. Washington, DC: U.S. Department of Agriculture, Forest Service. HARRINGTON, T.C. 1988. Leptograhium species, their distributions, hosts and insect vectors, p. 1-40. In Harrington, T.C. and F.W. Cobb, Jr., eds. Leptographium Root Diseases on Conifers. American Phytopathological Society Press, St. Paul, MN. JOHNSON, N.E. and BELLUSCHI, RG. 1969. ttost-finding behavior of the Douglas-fir beetle. J. For. 67: 290-295. LIEUTIER, E 1993. Induced defence reaction of conifers to bark beetles and their associated Ophiostoma species, p. 225- 234. In Wingfield, M.J., Seifert, K.A., and Webber, J.E, eds. Ceratocystis and Ophiostoma Taxonomy, Ecology, and Pathogenicity. American Phytopathological Society Press, St. Paul, MN. REED, A.N., HANOVER, J.W., and FURNISS, M.M. 1986. Douglas-fir and western larch: chemical and physical properties in relation to Douglas-fir bark beetle attack. Tree Physiol. 1: 277-287. REID, R.W., WHITNEY, H.S., and WATSON, J.A. 1967. Reactions of lodgepole pine to attack by Dendroctonus ponderosae Hopkins and blue stain fungi. Can. J. Bot. 45:1115-1126. ROSS, D.A. 1967. Wood- and bark-feeding Coleoptera of felled western larch in British Columbia. J. Entomol. Soc. Brit .... Columbia 64: 23-24. ROSS, D.W., FENN, R, and STEPHEN, F.M. 1992. Growth of southern pine beetle associated fungi in relation to the induced wound response in loblolly pine. Can. J. For. Res. 22:1851-1859. RUDINSKY, J.A. 1966. Host selection and invasion by the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, in coastal Douglas-fir forests. Can. Entomol. 98: 98-111. RUMBOLD, C.T. 1936. Three blue-staining fungi, including two new species, associated with bark beetles. J. Agric. Res. 52: 419-437. SOLHEIM, H. 1988. Pathogenicity of some Ips typographus-associated blue-stain fungi to Norway spruce. Medd. Nor. inst. skogforsk. 40(14): 1-11. SOLHEIM, H. 1993. Ecological aspects of fungi associated with the spruce beetle Ips typographus in Norway, p. 235-242. In Wingfield, M.J., Seifert, K.A., and Webber, J.F., eds. Ceratocystis and Ophiostoma Taxonomy, Ecology, and Pathogenicity. American Phytopathological Society Press, St. Paul, MN. WHITNEY, H.S. 1982. Relationships between bark beetles and symbiotic organisms, p. 183-211. In Mitton, J.B. and Sturgeon, K.B., eds. Bark Beetles in North American Conifers. University of Texas Press, Austin, TX. 227
INTERRUPTION OF BARK BEETLE AGGREGATION BY A VIGOR-DEPENDENT PINUS HOST COMPOUND KENNETH R. HOBSON Utah State University, Department of Forest Resources and USDA Forest Service, Forestry Sciences Laboratroy, Logan, Utah 84322, USA INTRODUCTION The attraction of bark beetles to pheromones has been abundantly demonstrated in the last quarter century (Wood 1982, Borden 1985). Much less is known about the response of bark beetles to host volatiles. This paper reports preliminary evidence that one common host compound of North American conifers is a strong repellent or interruptant of aggregation for several species of bark beetles. It also reviews work that suggests that the level of this compound may be an indicator of host tree stress typically associated with bark beetle infestation in several species of pines. This compound, methyl chavicol, fulfills the two essential requirements for a kairomonal cue that can explain the selective infestation of susceptible trees by bark beetles, lit is correlated to tree susceptibility, and bark beetles respond to it. The search for a biochemical indicator of host tree stress that might serve as an olfactory cue for bark beetles is not new (Rudinsky 1962 and references therein, Cobb et al. 1968). Miller et al. (1968) examined the resin from smog-damaged ponderosa pines, Pinus ponderosa, in the San Bernardino Mountains east of Los Angeles. These trees were being selectively infested by mountain pine beetle, Dendroctonus ponderosae, and western pine beetle, Dendroctonus brevicomis. Of the six monoterpenes examined, none were shown to be significantly different between damaged and healthy trees. However, Cobb et al. (1972) found a strong, significant difference in the level of one compound, methyl chavicol, in the foliage of smogdamaged ponderosa pines and healthy trees. Smog-damaged trees had 71% less methyl chavicol in their foliage - by far the sharpest phytochemical difference found. More recently Nebeker et al. (1995) measured 18 host volatiles from healthy lodgepole, Pinus contorta, and those diseased by dwarf mistletoe, Arceuthobium americanum; armillaria root disease, Armillaria mellea; or comandra blister rust, Cronartium comandrae. The latter two diseases are reported to be the most common predisposing agents of lodgepole to mountain pine beetle attack in the Intermountain West (Tkacz and Schmitz 1986). In the case of both predisposing diseases of lodgepole, there was a large and significant difference in the level of 4-allylanisole (a synonym for methyl chavicol) between healthy and diseased trees. This was not true for dwarf mistletoe, which was not shown to be associated with bark beetle infestation (Tkacz and Schmitz 1986). For both comandra and armillaria, the difference in the level of methyl chavicol was one of the greatest phytochemical differences found between diseased and healthy trees. Trees infected with comandra had 43.6% less methyl chavicol than healthy trees; trees infected with armillaria had 63% less methyl chavicol than healthy trees. Cobb et al. and Nebeker et al. both found the strongest biochemical differences between diseased and healthy trees in one compound. These results begged the question, "What is the response of bark beetles to methyl chavicol?" Electroantennograms (EAG) show a strong response to methyl chavicol. In 1989, Peter White found that estragole (methyl chavicol) produced the third highest response of all 11 host compounds tested on the red turpentine beetle, Dendroctonus valens (White and Hobson 1993). In 1993 Armand Whitehead showed a strong EAG response of mountain pine beetle to methyl chavicol (Hobson et al. in prep). A large EAG response does not indicate that a compound will be a strong attractant or repellent. However, ecologically relevant compounds that produce a large EAG are likely to have behavioral significance (Masson and Mustaparta 1990). These data encouraged us to conduct behavioral field studies. Mattson, W.J., Niemel_i, R, and Rousi, M., eds. 1996. Dynamics of forest herbivory: quest for pattern and principle. USDA For. Serv. Gen. Tech. Rep. NC-183, N.C. For. Exp. Sta., St. Paul, MN 55108. 228
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DYNAMICS OF FOREST HERBIVORY: QUEST
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TABLE OF CONTENTS Seeking The Rules
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32. Companion planting of the nitro
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GRIMALSKY, ¥.I., Research Institut
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POUTTU, ANTTI, Finnish Forest Resea
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persist in mature leaves and affect
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THE SINK-SOURCE HYPOTHESIS: TYPE AN
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late season defoliation decreases t
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BLANCHE, C.A., LORIO, EL., Jr., SOM
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NIEMELA, R, TUOMI, J,, and LOJANDER
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induced reactions have been studied
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summer, but the other sawfly specie
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LARSSON, S., BJ(_RKMAN, C., and GRE
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STAND AND LANDSCAPE DIVERSITY AS A
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not related to resistance to the se
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In contrast to the situation in eas
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ROOT, R.B. 1973. Organization of a
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We tlhus neglect two potential sour
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c__ c__ 1T1 Ill a) a) 0 0 0 e 1 o e
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c_ >l.s S S S m N < DN DN < D c DN
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:hen k > 0 suggest that the cue or
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oth as protective weapons and as po
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DEFENSE THEORIES AND BIRCH RESISTAN
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P 45 A L 40 A T 35 A B 30 I L 25 I
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However, hares showed strong prefer
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DUGLE, J.A. 1966. A taxonomic study
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accommodate findings related to mic
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Table l.--Mean ±S.E. area eaten by
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Sulfhydryl-Disulfide Mechanisms In
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Iraa prior experimental analysis of
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FRAZIER, J.L. and HEITZ, J.R. 1975.
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REICHARD, R 1993. From RNA to DNA,
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; trees, and between and within ind
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Encapsulated objects are data struc
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_re3.--The Lignum model can be cont
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SEVERE DEFOLIATION Tree Fine Root +
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RUMBAUGH, J., BLAHA, M., PREMERLANI
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oo0 a b ol ¢' _achiman_i ()® 1,eA
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1000 Conspicuous Defo i at ion 100
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2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
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Qualitative Changes in [,eaves and
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t00 2 Yr 1 Yr o_ 80 Treatment Treat
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Table 2.--Body size of Q. purzctate
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06 b AddedBSA < 04 _ 2rag -'=' i:3
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Severe Defol iat ion Site-A 1993 _
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KAMATA, N. and IGARASHI, M. 1995b.
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enclosure %r 20 post-diapausing sec
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2,500 F 000 F NF NF b _7 a _;_ a _7
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8O D 09 i 4,'.) F AS4L a Z___7 2o !
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2,500 _ st2-st4 El st5 0 st6 [_ pup
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96 SLANSKY, E, Jr. 1990. Insect nut
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later, we selected one more floweri
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Table 2.---Mean spruce budworm perf
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Table 3..--Mean spruce budworm perf
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FOLIVORE FEEDING ON MALE CONIFER FL
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Table 3.--Lymantria monacha prefere
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500- 400- i 300 v ¢ m a 200- I00 L
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From the point of view of trees, th
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THE BLACKMARGINED APHID AS A KEYSTO
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Wood et al. (1987) report that M. c
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MIZELL, R.E 1991. Pesticides and be
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METHODS Study Site The study took p
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the samples. Fertilization had subs
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Table 2.--Percent nutrient content
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Some tree responses to fertilizatio
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PHYTOCHEMICAL PROTECTION AND NATURA
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Indirect effects of abiotic factors
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HUNTER, M.D. and PRICE, P.W. 1992.
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A° ADDITIVE C. HYBRID SUSCEPTIBILI
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Galls, leaf miners, or leaf folds w
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Table 2.--Summary of the hypotheses
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the inheritance patterns of seconda
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FLOATE, K.D. and WHITHAM, T.G. 1993
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Elements of the Suitability/Defense
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Table l.--Comparing the mean number
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ACKNOWLEDGEMENTS The authors sincer
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SINGH, D.R 1986_ Breeding for resis
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The objectives of this presentation
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Table 1.--Average weevil attack on
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1977, Kline and Mitchell 1979, Wood
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Further research is underway to cla
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A SUGI CLONE HIGHLY PALATABLE TO HA
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RESULTS Seasonal Stability of Palat
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-
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OGAWA, A., NAGATA, Y., and SUKENO,
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MATERIALS AND METHODS Field Work In
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Laboratory Procedures The week afte
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E 120 Nogent-sur-Vernisson Remnings
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Interdependence Between Reaction Zo
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alance hypothesis (Lorio 1986) cann
Page 186 and 187: SOLHEIM, H., LANGSTROM, B., and HEL
Page 188 and 189: In a second experiment, three 30-ye
Page 190 and 191: Considering biochemical pathways, i
Page 192 and 193: - = 120 N /'_" =40 1/ _ j Days 0 _
Page 194 and 195: BIGGS, A.R. 1985. Suberized boundar
Page 196 and 197: DIFFERENTIAL SUSCEPTIBILITY OF WHIT
Page 198 and 199: Table 2.--Mortality of white fir pr
Page 200 and 201: Results from the geographic range p
Page 202 and 203: esulting seedlings were then rooted
Page 204 and 205: -0.2 0 5 82. _j -0,4 e a a a D4 E I
Page 206 and 207: 1960, Kalkstein 1976). Increased su
Page 208 and 209: RYAN, B.F., JOtNER, B.L., and RYAN,
Page 210 and 211: It is seldom feasible to control wa
Page 212 and 213: 15 15 u. 10 |O O N tal 5 5 g ao o 4
Page 214 and 215: Only recently have we conducted stu
Page 216 and 217: esistance to beetle attack. Another
Page 218 and 219: LORIO, EL., Jr'. and HODGES, J.D. 1
Page 220 and 221: EFFECTS OF ROOT INHABITING INSECT-F
Page 222 and 223: OCCURRENCE OF ROOT AND STEM SUBCORT
Page 224 and 225: ]Predisposition of Root-infected Tr
Page 226 and 227: chemistry associated with root colo
Page 228 and 229: Implications to Natural Resource Ma
Page 230 and 231: KLEPZIG, K.D., RAFFA, K.F., and SMA
Page 232 and 233: RAFFA, K.F., PHILLIPS, T., and SALO
Page 234 and 235: METHODS The study was installed in
Page 238 and 239: METHODS In the spring and summer of
Page 240 and 241: The importance of a compound as a r
Page 242 and 243: WERNER, R.A. 1995. Toxicity and rep
Page 244 and 245: 41 40 42 \ 4t 42 41 42 ",,,\ 42 40
Page 246 and 247: Table 1.--Summary data for the 26 p
Page 248 and 249: One weakness of using data from gen
Page 250 and 251: Lower Peninsula (populations 14-26)
Page 252 and 253: AUCLAIR, A.ND., WORREST, R.C., LACH
Page 254 and 255: KELLOMAKI, S., HANNINEN, H., and KO
Page 256 and 257: WARGO, RM. and HAACK, R.A. 1991. Un
Page 258 and 259: investigation. Sampling was done on
Page 260 and 261: Figure 2.--Changes in protein preci
Page 262 and 263: q) J 400 T J ! i i i i o control tr
Page 264 and 265: attacked trees than in control tree
Page 266 and 267: ACIDIC DEPOSITION, DROUGHT, AND INS
Page 268 and 269: Table 1.--Impact of acidic fog upon
Page 270 and 271: MENGEL, K., BREININGER, T., and LUT
Page 272 and 273: THE RESISTANCE OF SCOTCH PINE TO DE
Page 274 and 275: EXPERIMENTAL METHODS Experiments we
Page 276 and 277: C w Q. 1,200 1,000 o 800 C_ --- 600
Page 278 and 279: 0 .................................
Page 280 and 281: FONTAtNE, R.G. 1985. Forty years of
Page 282 and 283: Host Defenses An important componen
Page 284 and 285: Functional Heterogeneity of Forest
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The Connectivity Index (CI). The CI
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FUNCTIONAL HETEROGENEITYANALYSIS :
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The Modified Hazard Map. The next l
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systems facilitate mapping of fores
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KOLASA, J. and ROLLO, C.D. 1991. In
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) i,i_i)i_)i_i_ U.S. Departme_]t of
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