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Implications to Natural Resource Management and Forest Ecosystem Health The dynamics of insect and microbial colonization also suggests some specific management tactics that can prevent the onset and spread of Red Pine Decline. For example, mixed-block rather than uniform-species planting would reduce the transmission of Leptographium fungi through root grafts. This would contain root-diseased trees within relatively small groups rather than entire stands. Where economic incentives for red pine make large-scale planting of"alternate species impractical, planting buffer strips of alternate species might serve the same purpose. Pinus strobus would probably be a better candidate than Pinus banksiana, because the former is less preferred by the root weevils in our region (Hunt et al. 1993). Judicious thinning would also reduce stress and thereby reduce stand susceptibility. However, the timing is quite important. Open light penetration can favor site suitability to root weevils by warming and drying the soil (Wilson and Millers 1983). Moreover, thinning should be accompanied by stump removal to avoid leaving a food base for the root- and lower-stem feeding species (Witcosky and Hansen 1985, Witcosky et al. 1986). Our model also suggests that severing root grafts in advance (@ 15m) of killed trees could halt the transmission of Leptographium. We have begun tests on this idea in Wisconsin. We further recommend sanitation removal of dead and living trees (including stumps) within the radius of root severing. Although the insects and fungi described here are typically viewed as forest pests, these moderately pathogenic species might serve as useful bio-indicators. Reproductive success by these organisms is contingent on their ability to detect trees that are sufficiently stressed to have impaired defensive capability, but not so severely stressed as to be available to saprophytic competitors. As such, the population densities and behavior of moderately virulent endoparasites may be indicative of reduced ecosystem vigor associated with anthropogenic disturbances or injudicious forestry practices. Our monitoring data and bioassay show this idea is feasible for at least one form of forest decline. That is, populations of some root- and lower stem-feeding insects were higher in declining than in asymptomatic stands. Moreover, different species vary in how their densities relate to the progression of decline and in how they respond to tree stress physiology. We propose that the most useful bioindicator candidates might be those that show increased numbers or behavioral alteration before any symptoms become apparent, but are not by themselves major outbreak pests. SUMMARY Interactions between feeding guilds that affect different plant parts can strongly affect the dynamics of forest declines. Environmental stresses can mediate these outcomes, especially against highly generalized plant defense systems. Such interactions among insects and fungi are primarily responsible for Red Pine Decline, a syndrome affecting mature plantation trees in the Great Lakes region of North America. Root- and lower stem-colonizing insect-fungal complexes reduce tree resistance against stem-feeding bark beetle-fungal complexes. Between-stand transmission of moderately pathogenic Ophiostomatales fungi occurs by insect vectoring, but within-stand progression also occurs through extensive root grafts in plantations. Subterranean scolytids and curculionids show a high degree of niche partitioning based on host tissue and condition, chemical ecology, microbial associates, and spatial and temporal distributions. Healthy trees can resist colonization by rapidly accumulating terpenes and phenolics to concentrations that repel or kill I. pini and inhibit O. ips spore germination and mycelial growth. However, tree constitutive and inducible defenses are compromised by root infection. Behavioral assays are generally the most sensitive to the early stages of root injury. Among these, post landing assays of I. pini acceptance behavior are most sensitive, especially those in which beetles are exposed to host reaction tissue formed in response to challenge inoculation with fungi. Host susceptibility to the root colonizing agents can be increased by environmental stresses such as reduced light, defoliation, and nitrification. An understanding of the complex interactions underlying forest declines can improve our ability to relate various syndromes to management practices, anthropogenic stress, and natural succession. These systems can also serve as useful models to improve our overall understanding of plant-insect and insect-fungal interactions. The underlying mechanisms also suggest some specific tactics for preventing or ameliorating Red Pine Decline. Marginally pathogenic agents such as the organisms described here may also serve as useful indicators of forest ecosystem health. Because their existence is contingent on detecting trees under sufficient stress to fully resist attack but not yet available to saprophytic competitors, the population densities, behavior, and developmental success of these insects and fungi n-fight be good indicators of incipient stress at the stand or tree level. 219
ACKNOWLEDGMENTS This work was supported by USDA CSRS Competitive Grant 90-3725P-5581, the University of Wisconsin Graduate School, the University of Wisconsin College of Agricultural and Life Sciences, Mclntire Stennis, and Hatch Regional Project W-187. We would also like to thank our colleagues L. Rieske, S. Krause, and K. Kleiner )'or input on the ideas discussed in this paper. REFERENCES CITED BERRYMAN, A.A. 1972. Resistance of conifers to invasion by bark beetle-fungus associations. Bio Science 22: 598-602. CATES, R.G. and ALEXANDER, H. 1982. Host resistance and susceptibility, p. 212-263. In Mitton, J.B. and Sturgeon, K.B., eds. Bark Beetles in North American Conifers: A System for the Study of Evolutionary Biology. University Texas Press, Austin. CHRISTENSEN, E. and ERICSSON, A. 1986. Starch reserves in Picea abies in relation to defense reaction against a bark beetle transmitted blue-stain fungus, Ceratocuptispolonica. Can. J. For. Res. 16: 78-83. CHRISTENSEN, E. and HORNTVEDT, R. 1983. Combined lps Ceratocystis attack on Norway spruce and defensive mechanisms of the trees. Zeit. angew. Entomol. 96:110-118. DUNN, J.E, POTTER, D.A., and KIMMERER, T.W. 1990. Carbohydrate reserves, radial growth, and mechanisms of resistance of oak trees to phloem-boring insects. Oecologia 83: 458-468. ELKINTON, J.W. and WOOD, D.L. 1980. Feeding and boring behavior of the bark beetle lps paraconfusus (Coleoptera Scolytidae) on the bark of a host and non-host tree species. Can. Entomol. 112: 797-809. ERICSSON, A., HELLQUIST, C., LANGSTROM, B., LARSSON, S., and TENNOW, O. 1985. Effects on growth of simulated and induced shoot pruning by Tomicus piniperda as related to carbohydrate and nitrogen dynamics in Scots pine. J. Applied Ecol. 22: 105-124. HODGES, J.D., ELAM, W.W., WATSON, W.E, and NEBEKER, T.E. 1979. Oleoresin characteristics and susceptibility for four southern pines to southern pine beetle attacks. Can. Entomol. 11: 889-896. HOUSTON, D.R. 1981. Stress triggered tree diseases_the diebacks and declines. NE-INF-41-81. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 4-7. HUETTL, R.E, FINK, S., LUTZ, H-J., POTH, M., and WISMEWSKI, J. 1990. Forest decline, nutrient supply and diagnostic fertilization in Southwestern Germany and in Southern California. For. Ecol. and Manage. 30: 341-350. HUNT, D.W.A., LINTEREUR, G., SALOM, S., and RAFFA, K.E 1993. Performance and preference of Hylobius radicis Buchanan, and H. pales (Herbst) (Coleoptera:Curculionidae)on various Pinus species. Can. Entomol. 125:1003- 1010. KIMMERER, T.W. and KOZLOWSKI, T.T. 1982. Ethylene, ethane, acetyl aldehyde, and ethanol production by plants under stress. Plant Physiol. 69: 840-847. KLEIN, R.M. and PERKINS, T.D. 1988. Primary and secondary causes and consequences of contemporary forest decline. The Bot. Rev. 54: 1-43. KLEPZIG, K.D., RAFFA, K.E, and SMALLEY, E.B. 1991. Association of insect-fungal complexes with Red Pine Decline in Wisconsin. For. Sci. 37: 1119-1139. 220
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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
Page 178 and 179: Laboratory Procedures The week afte
Page 180 and 181: E 120 Nogent-sur-Vernisson Remnings
Page 182 and 183: Interdependence Between Reaction Zo
Page 184 and 185: 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 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 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
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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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