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Lower Peninsula (populations 14-26) demonstrated that as the climatic differential between the origin and the test site increased, tree height decreased (p < 0.006, r2 = 0.51, n = 13 birch populations), tree survival decreased (p < 0.012, r2 = 0.49, n = 13), and tree susceptibility to borer attack increased (p < 0.006, r2= 0.51, n = 13). Similarly, in the second case, which included 12 birch populations (populations 4-7, 10, 12-18), linear regression analysis indicated that as the climatic differential between origin and test site increased, tree height decreased (p < 0.02, r2= 0.43, n = 12 birch populations), tree survival decreased (p < 0.038, r2= 0.36, n = 12), and tree susceptibility to borer attack increased (p < 0.012, r2 = 0.45, n = 12). DISCUSSION Cannell et al. (1989) state that most tree species will benefit from an increase in annual mean temperature of I°C, but genetic increases will often cause growth reductions or mortality. Similarly, Matyas (1994) states that an increase in temperature will positively affect tree growth only within each species' limits of physiological and ecological tolerance. Paper birch appears to have narrow limits of heat tolerance. In the present study, as the climatic differential increased for individual seed sources, birch trees grew slower, experienced higher mortality rates, and sustained higher rates of borer attack. Such results strongly suggest that paper birch will experience dramatic decline along its southern range in response to climatic warming, and that the bronze birch borer will likely be the principal mortality agent involved in future birch decline. Similar predictions for widespread decline of paper birch in response to global warming have been made by Pastor and Post (1988), Overpeck et al. (1991), Botkin and Nisbet (1992), Reed and Desanker (1992), Reed et al. (1992b), Solomon and Bartlein (1992), and Jones et al. (1994). Several of the general circulation models predict reduced rainfall or increased rates of evapotranspiration in association with increased air temperatures (Karl et al. 1991, IPCC 1992). Thus, if higher temperatures are combined with reduced rainfall, then many tree species will likely be even more susceptible to attack by trunk-boring insects (Mattson and Haack 1987a, 1987b). As mentioned above, differences in sensitivity to photoperiod between northern and southern seed sources could partly explain the relatively poorer performance exhibited by the more northerly birch families in the present study. However, when I restricted the birch data set to seed sources that spanned as little as 1.5 ° and 2° latitude, the same patterns of reduced paper birch growth, survival, and resistance to bronze birch borer persisted. In a Picea abies provenance study, Vaartaja (1959) reported little difference in tree growth for seed sources collected between 47°N and 52°N and then planted at 47°N. Although the general rule for photoperiod is that the further north the origin, the more photoperiod sensitive the seed source, Beuker (1994) states that the exact effect of photoperiod cannot be predicted. Although changes in photoperiod are undoubtedly important, the significant relationships found here for two relatively tight latitudinal bands, strongly suggest that the differences in seed source performance in the present study were greatly influenced by temperature. Another possible reason for the poorer growth of cold-adapted seed sources at warmer sites, is that they may have higher dark respiration rates than local sources (Kozlowski et al. 1991, Mebrahtu et al. 1991). Consequently, when growing in a warmer environment, cold-adapted sources expend more energy and have lower net assimilation rates than local sources. Reduced rates of net assimilation can help explain greater susceptibility to borer attack since compromised trees would likely produce fewer defensive compounds and have reduced rates of diameter growth (Herms and Mattson 1992). Slow stem growth has been shown to increase susceptibility of paper birch to the bronze birch borer (Herms 1991, Herms and Mattson 1991). In Michigan, as was true for much of North America, 1988 was a year of severe heat and drought (Kerr 1989, USGS 1991). In the Great Lakes region, outbreaks of several cambial-feeding forest insects occurred in 1988, including the bronze birch borer (Haack and Mattson 1989, Jones et al. 1993). Part of the rapid increase in birch mortality and borer attack between years 12 (1987) and 15 (1990) in the present study (Figs. 3B, 3C) could have been triggered by the 1988 drought. In fact, it appears from the slopes of tree survival (Fig. 3B) and borer attack (Fig. 3C) for the four CD treatments, that the CD2 °, CD3 °, and CD4 ° trees were much more negatively impacted by the heat and drought than were the CD1 ° trees, suggesting that an increase in annual mean temperature of about 1°C will have limited detrimental effects on paper birch, but that increases of 2°C and more will be very damaging. In support of this contention are studies in the Upper Peninsula of Michigan by Reed et al. (1992a) and Jones et al. (1993) that report slower growth and higher mortality rates for paper birch, following the warm and dry years of 1987 and 1988 when mean summer temperatures were about 1°C above normal. In controlled studies on yellow birch, Betula alleghaniensis Britton, limited rootlet mortality was observed with a I°C increase in soil temperature, but extensive rootlet mortality occurred with increases of 6-7°C (Redmond 1955, 1957). 241
As global warming occurs, I predict that those tree species with associated cambial-feeding insects that invade the trunk will be the first tree species to exhibit widespread mortality. This guild of trunk-infesting, cambial-feeding insects poses the greatest lethal threat to their host trees (Haack and Slansky 1987, Mattson et al. 1988, Haack and Byler 1993). Trees are typically highly resistant to cambial-feeding insects, but during periods of stress, they become more susceptible to such insects (Mattson and Haack 1987a, 1987b; Millers et al. 1989). Some of the boreal tree genera that occur in Michigan that can be killed by trunk-infesting, cambial-feeding insects, and thus will likely be more at risk as climatic warming occurs, include Abies, Betula, Larix, Picea, Pinus, Populus, and Tsuga (Table 3). Although climatic warming in the northern hemisphere is predicted to bring about tree mortality along the southern edge of a species range, the northern range of these same tree species may expand (Cannell et al. 1989, Woodward 1992, Sykes and Prentice 1995). In fact, paper birch has been documented to be expanding its range northward into the tundra, possibly in response to climatic warming (Woodward 1992). Table 3.--Boreal tree genera of the Great Lakes region that will likely experience widespread insect-induced mortality as a result of climatic warming and the corresponding trunk-infesting, cambial-feeding insect that will likely be the major mortality agent. Host tree Insect species Genus Common name Species Common name Family Abies Fir Pityokteins sparsus (LeConte) Balsam fir bark beetle Scolytidae Betula Birch Agrilus anxius Gory Bronze birch borer Buprestidae Larix Larch Dendroctonus simplex LeConte Eastern larch beetle Scolytidae Picea Spruce Dendroctonus rufipennis (Kirby) Spruce beetle Scolytidae Pinus Pine Ips pini (Say) Pine engraver beetle Scolytidae Populus Aspen Agrilus liragus Barter and Brown Bronze poplar borer Buprestidae Tsuga Hemlock Melanophilafulvoguttata (Harris) Hemlock borer Buprestidae The present study shows that data from genetic test plantings of forest tree species can provide insight into how treeinsect interactions may change as air temperatures increase in the future. Similar studies should be conducted by others. This would be particularly useful in areas where the local geography allows seed to be collected from several temperature regimes within a narrow latitudinal band and thereby allow the effects of temperature and photoperiod to be partly disentangled. ACKNOWLEDGMENTS I wish to thank the staff of Michigan State University, Fred Russ Experimental Forest, for field assistance in this study; Paul Bloese, Michigan State University, Department of Forestry and Michigan Cooperative Tree Improvement Program, for making available early tree performance data from this paper birch plantation; and Paul Bloese, Daniel Herms, Dow Gardens, and Robert Lawrence, USDA Forest Service, for reviewing an earlier draft of this manuscript. LITERATURE CITED AKERS, R.C. and NIELSEN, D.G. 1990. Reproductive biology of the bronze birch borer (Coleoptera: Buprestidae) on selected trees. J. Entomol. Sci: 25: 196-203. ANDERSON, R.F. 1944. The relation between host condition and attacks by the bronzed birch borer. J. Econ. Entomol. 37: 588-596. 242
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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
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 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 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
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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