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From the point of view of trees, the evolutionary response in terms of defense seems to be consistent with a concept k_own from deciduous species like oak or beech, as masting or mast seeding (Silvertown 1981). tn other words, in some years there will be high beech-mast or acorn production, but no seeds in the years in between. In the latter years, pest populations will decrease markedly. The strategy has probably evolved against seed predators but it could also work against _omedefoliators. The second line of defense is simply the variability of bud burst (Mitcherlich and Wellenstein 1942). Large variation in bud burst between provenances exist, but also large variation occurs within provenances. However, variability as a defense will mean that some larvae always will survive. The third line of defense against flush-feeders is a chemical one. Apparently, rather few spruce herbivores are deterred from feeding on newly burst needles to such an extent that they prefer old needles (Schopf 1986). In contrast, in pine it seems essential that flush feeders avoid plant defenses and here male flowers are of vitae importance. In pine, male flowers are a more predictable resource than in spruce. Larvae that hatch after the flowering period will die and hence the risk of early hatch is less because male flowers are most often present. However, the strategy might fail; they might hatch on a tree without flowers, or worse in a year without flowers in the whole stand. In the context of the phenological window it is actually necessary to consider the choices made by the female moth. Where does she deposit her eggs? Does she in fact spread the risk so that at least some of her larvae will survive? But at the same _ime,does she abandon an optimal strategy of maximizing short term fitness? Essential to all these considerations are _hedevelopment time of buds and eggs, both factors being related to spring temperatures, although probably not in the same fashioa. I hypothesize that the position of the eggs is a key factor in this synchronization simply because the temperature varies so much between microsites. 1#mantria monacha females deposit their egg clusters mainly on the tree trunk under bark scales. Egg numbers and egg hatch differ consideraNy between north- and south-facing sides of the trunk (Raae 1979), and it might be that this is the mechanism the females employ when they spread the risk. This could be the way whereby females ensure that some of their progeny in this variable environment will hatch when the phenological window is open. Another bet-hedging mechanism couid be that after laying their first batch and becoming lighter, females fly to another tree and deposit the rest of their eggs. If the first tree is in the middle of a dense stand and the next is at the southern edge, the temperature regime is different and accordingly the bets change. litis interesting to observe that many flush feeders have wingless females or have reduced flight capability due to their heavy loads of eggs. In L monacha, females are winged but poor fliers and it is likely that the females lay their eggs close to if not directly on the tree where they pupated, thereby increasing the likelihood of synchronization between the egg i_atchand bud burst. In conclusion, avoiding defenses of new needles is absolutely necessary for pine defoliators but of minor importance _o spruce de_bliators. Bet hedging might prove to be the evolutionary response by the defoliators in order to find the host's phenological window. LITERATURE CITED DEN BOER, RJ. 1968. Spreading of risk and stabilization of animal numbers. Acta Biotheoretica 18:165-194. FEEENY, P 1970. Seasonal changes in oak leaf tannins and nutrients as a cause of spring feeding by winter moth caterpillars. Ecology 51' 565-582. KEDA, T., MATSUMURA, E, and BENJAMIN, D.M. 1977. Chemical basis for feeding adaptation of pine sawflies, ,_e_:dlpr_on rug(frorls and Neodiprion swainei. Science 197: 497-498. JENSEN, T.S. 1988. Variability of Norway spruce needles; performance of spruce sawflies (Gilpinia hercyniae). Oecologia 77:313-32(t.
JENSEN, T.S. 199 la. Integrated pest management of the nun moth, Lymatltria monacha (Lepidoptera: Lymantriidae) in Denmark. For. Ecol. Mgmt. 39: 29-34. JENSEN, T.S. t99 lb. Patterns of nutrient utilization in the needle feeding guild. In Baranchikov et at., eds. Forest insect guilds: Patterns of interaction with host trees. GTR-NE-153. Radnor. PA: U.S. Department of Agriculture, Forest Service. JENSEN, T.S. 1992. The role of plant stress in the population dynamics of nun moths. IUFRO Conference Zakopane, Poland 1991. LARSSON, S. and TENOW, O. 1980. Needle-eating insects and grazing dynamics in a mature Scots pine forest in Central Sweden. In Persson, T., ed. Structure and function of northern coniferous forests: an ecosystem study. Ecol. Bull. 32: 269-306. MITCHERLICH, H. and WELLENSTEIN, G. 1942. Die Nonne an Frtih- und Sp_ittreiberformen der Fichte. Mon. Z. ang. Ent. 15: 94-125. NIEMEL,_, R and HAUKtOJA, E. 11982. Seasonal patterns in species richness of herbivores: Macrolepidopteran larvae on Finnish deciduous trees. Ecol. Entomol. 7: 169-175. PHILIPPI, T. and SEGER, J. 1989. Hedging one's evolutionary bets, revisited. TREE 4:41:44. RAAE, K. 1979. The distribution of nun moth eggs in old spruce stands. Unpubl. rep. Agricultural Univ., Copenhagen. SCHONHERR, J. 1989. Outbreak characteristics of Lymantriids. In Wallner, W.E. and McManus, K.A., ed. Lymantriidae: A comparison of features of new and old world tussock moths. GTR-NE-123. Radnor, PA: U.S. Department of Agriculture, Forest Service. SCHOPE R. 1986. The effect of secondary needle compounds on the development of phytophagous insects. For. Ecol. Man. 15: 54-64. SEGER, J. and BROCKMANN, H.J. 1987. What is bet-hedging? Oxford Surv. Evol. Biol. 4: 182-211. SILWERTOWN, J.W. 1981. The evolutionary ecology of mast seeding in trees. Biol. J. Linn. Soc. 14: 235-240. VARLEY, G.C. and GRADWELL, G.R. 1970. Recent advances in insect population dynamics. Ann. Rev. Ent. 15: 1-24. 111
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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,
Page 70 and 71: ; trees, and between and within ind
Page 72 and 73: Encapsulated objects are data struc
Page 74 and 75: _re3.--The Lignum model can be cont
Page 76 and 77: SEVERE DEFOLIATION Tree Fine Root +
Page 78 and 79: RUMBAUGH, J., BLAHA, M., PREMERLANI
Page 80 and 81: oo0 a b ol ¢' _achiman_i ()® 1,eA
Page 82 and 83: 1000 Conspicuous Defo i at ion 100
Page 84 and 85: 2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
Page 86 and 87: Qualitative Changes in [,eaves and
Page 88 and 89: t00 2 Yr 1 Yr o_ 80 Treatment Treat
Page 90 and 91: Table 2.--Body size of Q. purzctate
Page 92 and 93: 06 b AddedBSA < 04 _ 2rag -'=' i:3
Page 94 and 95: Severe Defol iat ion Site-A 1993 _
Page 96 and 97: KAMATA, N. and IGARASHI, M. 1995b.
Page 98 and 99: enclosure %r 20 post-diapausing sec
Page 100 and 101: 2,500 F 000 F NF NF b _7 a _;_ a _7
Page 102 and 103: 8O D 09 i 4,'.) F AS4L a Z___7 2o !
Page 104 and 105: 2,500 _ st2-st4 El st5 0 st6 [_ pup
Page 106 and 107: 96 SLANSKY, E, Jr. 1990. Insect nut
Page 108 and 109: later, we selected one more floweri
Page 110 and 111: Table 2.---Mean spruce budworm perf
Page 112 and 113: Table 3..--Mean spruce budworm perf
Page 114 and 115: FOLIVORE FEEDING ON MALE CONIFER FL
Page 116 and 117: Table 3.--Lymantria monacha prefere
Page 118 and 119: 500- 400- i 300 v ¢ m a 200- I00 L
Page 122 and 123: THE BLACKMARGINED APHID AS A KEYSTO
Page 124 and 125: Wood et al. (1987) report that M. c
Page 126 and 127: MIZELL, R.E 1991. Pesticides and be
Page 128 and 129: METHODS Study Site The study took p
Page 130 and 131: the samples. Fertilization had subs
Page 132 and 133: Table 2.--Percent nutrient content
Page 134 and 135: Some tree responses to fertilizatio
Page 136 and 137: PHYTOCHEMICAL PROTECTION AND NATURA
Page 138 and 139: Indirect effects of abiotic factors
Page 140 and 141: HUNTER, M.D. and PRICE, P.W. 1992.
Page 142 and 143: A° ADDITIVE C. HYBRID SUSCEPTIBILI
Page 144 and 145: Galls, leaf miners, or leaf folds w
Page 146 and 147: Table 2.--Summary of the hypotheses
Page 148 and 149: the inheritance patterns of seconda
Page 150 and 151: FLOATE, K.D. and WHITHAM, T.G. 1993
Page 152 and 153: Elements of the Suitability/Defense
Page 154 and 155: Table l.--Comparing the mean number
Page 156 and 157: ACKNOWLEDGEMENTS The authors sincer
Page 158 and 159: SINGH, D.R 1986_ Breeding for resis
Page 160 and 161: The objectives of this presentation
Page 162 and 163: Table 1.--Average weevil attack on
Page 164 and 165: 1977, Kline and Mitchell 1979, Wood
Page 166 and 167: Further research is underway to cla
Page 168 and 169: 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
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SOLHEIM, H., LANGSTROM, B., and HEL
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In a second experiment, three 30-ye
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Considering biochemical pathways, i
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- = 120 N /'_" =40 1/ _ j Days 0 _
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BIGGS, A.R. 1985. Suberized boundar
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DIFFERENTIAL SUSCEPTIBILITY OF WHIT
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Table 2.--Mortality of white fir pr
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Results from the geographic range p
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esulting seedlings were then rooted
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-0.2 0 5 82. _j -0,4 e a a a D4 E I
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1960, Kalkstein 1976). Increased su
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RYAN, B.F., JOtNER, B.L., and RYAN,
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It is seldom feasible to control wa
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15 15 u. 10 |O O N tal 5 5 g ao o 4
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Only recently have we conducted stu
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esistance to beetle attack. Another
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LORIO, EL., Jr'. and HODGES, J.D. 1
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EFFECTS OF ROOT INHABITING INSECT-F
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OCCURRENCE OF ROOT AND STEM SUBCORT
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]Predisposition of Root-infected Tr
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chemistry associated with root colo
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Implications to Natural Resource Ma
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KLEPZIG, K.D., RAFFA, K.F., and SMA
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RAFFA, K.F., PHILLIPS, T., and SALO
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METHODS The study was installed in
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FURNISS, M.M., LIVINGSTON, R.L., an
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METHODS In the spring and summer of
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The importance of a compound as a r
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WERNER, R.A. 1995. Toxicity and rep
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41 40 42 \ 4t 42 41 42 ",,,\ 42 40
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Table 1.--Summary data for the 26 p
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One weakness of using data from gen
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Lower Peninsula (populations 14-26)
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AUCLAIR, A.ND., WORREST, R.C., LACH
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KELLOMAKI, S., HANNINEN, H., and KO
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WARGO, RM. and HAACK, R.A. 1991. Un
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investigation. Sampling was done on
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Figure 2.--Changes in protein preci
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q) J 400 T J ! i i i i o control tr
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attacked trees than in control tree
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ACIDIC DEPOSITION, DROUGHT, AND INS
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Table 1.--Impact of acidic fog upon
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MENGEL, K., BREININGER, T., and LUT
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THE RESISTANCE OF SCOTCH PINE TO DE
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EXPERIMENTAL METHODS Experiments we
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C w Q. 1,200 1,000 o 800 C_ --- 600
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0 .................................
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FONTAtNE, R.G. 1985. Forty years of
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Host Defenses An important componen
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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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