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WARGO, RM. and HAACK, R.A. 1991. Understanding the physiology of dieback and decline diseases and its management implications for oak, p. 147-158. In The Oak Resource in the Upper Midwest: Implications for Management Pro- ceedings, 3-6 June 1991, Winona, MN. Minnesota Extension Service, Univ. Minn. NR-BU-5663-S WILLIAMS, D.W. and LIEBHOLD, A.M. 1995. Forest defoliators and climate change: Potential changes in spatial distribu- tion of outbreaks of western spruce budworm (Lepidoptera: Tortricidae) and gypsy moth (Lepidoptera: Lyrnantriidae). Environ. Entomol. 24: 1-9. WILSON, L. F. and HAACK, R.A. 1990. The bronze birch borer. U.S. Dept. Agric. For. Serv., North Central For. Exp. Sta. Northern Hardwood Note 7.09. 3 p. WOODWARD, EI. 1992. A review of the effects of climate on vegetation: Ranges, competition, and composition, p. 105- 123. In Peters, R.L. and Lovejoy, T.E., eds. Global Warming and Biological Diversity. Yale Univ. Press, New Haven, CT. WRIGHT, J.W. 1976. Introduction to Forest Genetics. Academic Press, New York. 247
VARIATIONS IN SPRUCE NEEDLE CHEMISTRY AND IMPLICATIONS FOR THE LITTLE SPRUCE SAWFLY, PRISTIPHORA ABIETINA C. SCHAFELLNER, R. BERGER, J. MATTANOVICH, and E. FUHRER Institute of Forest Entomology, Forest Pathology, and Forest Protection Universit_it ftir Bodenkultur, Hasenauerstrasse 38, A 1190 Vienna, Austria INTRODUCTION The plant-herbivore system of Norway spruce, Picea abies Karst., and the little spruce Sawfly, Pristiphora abietina Christ (Hymenoptera: Tenthredinidae), provides a good opportunity to study the synchronization between the host's bud burst and the beginning of larval feeding. Adults of the sawfly emerge in spring at about the same time as spruce bud break and oviposit in the soft, partially-expanded needles. Every single spruce bud is suitable for oviposition for only a few days. The emerging larvae almost always remain on the expanding bud where they hatch and only under starving conditions do they switch buds. After 2 or 3 weeks, depending on weather conditions, the larvae are fully grown and drop to the ground to form cocoons. They overwinter in the litter until the next spring. The larvae of the little spruce sawfly are early-spring feeders and restricted to feeding on expanding spruce needles. Their success and survival depend on the exact phenological coincidence of bud break and oviposition, because larvae can initiate feeding only in flushing buds. Seasonal variations in the timing of bud break can therefore be of defensive value for the trees: trees that are genetically predisposed to flush needles earlier or later than the rest of the stand, thereby preceding or following the sawfly swarming, may escape larval feeding. During the last 3 decades the sawfly has become an important spruce pest in several parts of Austria. The most severe attacks occur in lowland areas where the natural deciduous forests have been replaced by monocultures of Norway spruce. As the larvae feed exclusively on current-year needles, the affected trees do not die, but repeated infestations lead to deformed, bushy crowns and to reduced height and volume increments. In recent years, an increase in frequency and duration of epidemics has been observed, particularly in areas exposed to air pollutants like sulfur and nitrogen (Sierpinski 1985, Berger 1992). There is good reason to believe that some air pollutants may favor the sawfly's success, either by improving food quality due to changes in needle chemistry and/or by weakening the tree's defense system (Schafellner et al. 1993, Schafellner et al. 1994, Berger und Katzensteiner 1994). This paper reports a study on the most important food quality parameters of newly emerging needles during the short period of larval feeding, and the changes that occur during the weeks of rapid growth. Additionally, the effect of excess nitrogen input (via fertilization) on needle chemistry and the nutritional value of the spruce needles for the feeding larvae is demonstrated. METHODS Three even-aged stands of trees 16 years old, were selected for study to see if food quality parameters show significant site-specific variation. Stand A is located at ca. 550 m elevation in Hochstrass, Lower Austria, and stands B and C are located at ca. 720 m elevation in the Hausruck, Upper Austria. Moderate sawfly attack was observed at stand A during the mid- 1980's. Throughout the 3 years of attack, tree flushing and sawfly attack for every individual tree was recorded (Holzer t988). The stand was attacked irregularly so that affected and unaffected individuals often stood close together, although the trees did not vary in the timing of bud break and should have been suitable for oviposition. From spring 1988 onwards, the sawflies vanished from the area so that during the 1989 sampling year, no frass was collected. Based on the data collected during the years of attack, 15 trees that had been affected once and 15 trees that remained unaffected, were selected for Mattson, W.J., Niemel_i, P., 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. 248
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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
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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
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 236 and 237: FURNISS, M.M., LIVINGSTON, R.L., an
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 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
Page 286 and 287: The Connectivity Index (CI). The CI
Page 288 and 289: FUNCTIONAL HETEROGENEITYANALYSIS :
Page 290 and 291: The Modified Hazard Map. The next l
Page 292 and 293: systems facilitate mapping of fores
Page 294 and 295: KOLASA, J. and ROLLO, C.D. 1991. In
Page 296: ) i,i_i)i_)i_i_ U.S. Departme_]t of
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