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Galls, leaf miners, or leaf folds were counted on 50 shoots per plant for hybrids and S. eriocephala and on 300 shoots for S. sericea in 1991 only. Data are expressed as number of herbivores per 300 shoots. Because leaf folds of Phyllocolpa r_igritaand Phyllocolpa sp. nov. were not distinguished on S. sericea in 1991, we combined them for analyses in all years. All other species densities are considered separately. Species densities were considered statistically independent among plants within taxa for all years (Fritz et al. 1994), and species-wide significance tests rather than table-wide significance tests were applied. ANOVAs were performed for each year separately. We first performed a multivariate ANOVA, using loge transformed densities of alt herbivore species, and then we performed univariate ANOVAs for each species separately. Orthoganol contrasts were performed to test for the fit of each herbivore species to the hypotheses. We performed only two orthoganol contrasts to test the significance of herbivore densities on willow taxa (species) to avoid overparameterization of the analysis, since only three taxonomic groups were present (SAS Institute 1985). To test the Additive hypothesis, we tested the significance of the hybrid-midparent contrast (Contrast A) using sequential Bonferroni analysis with specieswide significance at P < 0.05/2 = 0.025 (Rice 1989). If this test was significant, we rejected the Additive hypothesis and then tested the contrast between herbivore density on hybrids and density on the numerically closest parent (Contrast B) (P < 0.05/ 1 = 0.05). This contrast tested if there was a significant departure from the Dominance hypothesis. If the Dominance hypothesis was rejected, we inspected the means to determine if the Hybrid Susceptibility hypothesis (density on hybrids exceeded that of the highest parent) or the Hybrid Resistance hypothesis (density on hybrids was lower than the lowest parent) was supported, or if the density of hybrids was intermediate between that of one parent and the midparent value. This last possibility would support a partial Dominance hypothesis. RESULTS Multivariate ANOVAs showed highly significant contrasts between herbivore densities on hybrids and the mean herbivore densities of parent species for all three years (Table 1). This is a rejection of the Additive hypothesis overall. Further multivariate contrasts could not be made since the test of the Dominance hypothesis requires comparison to the numerically nearest parent, which would not be the same for all herbivores. In 1991, five herbivore species had significant hybrid vs. parents contrasts (Contrast A) at P < 0.025, and two herbivore species had marginally significant hybrid vs. parents contrasts at P < 0.05 (Table 1). In 1992, four species had significant hybrid vs. parents contrasts (Table 1). Three species that had significant contrasts in 1991 did not have significant contrasts in 1992. In 1993, five species had significant hybrid vs. parent contrasts (Table 1). Overall, then, there appear to be differences among the years in the species of herbivores that had significant departures from the Additive hypothesis. To illustrate the year-to-year differences in more detail and to draw conclusions about which hypotheses mentioned above are supported by each species, we present the separate results for each year for each species, which included all censused plants. Both contrasts for Phyllonoo, cter salicifoliella were significant in 1991 and 1993 (Table 1), and the means support for the Hybrid Susceptibility hypothesis (Table 2). In I992, neither contrast was significant, which supported the Additive hypothesis. The other leaf miner Phyllocnistis sp. also showed differences between years. In 1991 and 1993, the first contrast was not significant, supporting the Additive hypothesis (Tables 1 & 2). Fritz et al. (1994) concluded that this species supported the Hybrid Susceptibility hypothesis in 1991, because of the marginally significant contrast. In 1992 the Dominance hypothesis was supported with densities on S. eriocephala and hybrids being equal and lower than on S. sericea (Table 2). Among species of the leaf galling guild, Phyllocolpa termMatis showed consistent results in each year (Tables I & 2). In all cases the Hybrid Susceptibility hypothesis was supported. Phyllocolpa spp. supported the Additive hypothesis in 1991 and 1993 (Tables 1 & 2). In each year, including 1992, densities on hybrids were intermediate between the two parental species. In 1992, the first contrast was significant and the second contrast was not significant, supporting the Dominance hypothesis (Tables t & 2). Pontania sp. supported the Additive hypothesis in 1991 and 1992, but the Dominance hypothesis was supported in 1993 (Tables 1 & 2). Iteomyia salicifolia supported the Hybrid Susceptibility hypothesis in 1991 and 1993 (Tables 1 & 2). In each year, including 1992, densities on hybrids were greater than on either parental species. In 1992, the first contrast was not significant at P < 0.025 (although it was marginally significant at P < 0.05), thereby supporting the Additive hypothesis, even though the mean density on hybrids was more than twice as high as on S. eriocephala (Table 2). The last leaf galling species, A. tetanothrix, supported the Dominance hypothesis in 1991 and 1993 (Tables 1 & 2). In both of 135
Table 1. --Summary of multivariate and univariate ANOVAs of herbivore densities on S. sericea, S. eriocephala, and hybrid plants for 1991, 1992, and 1993. F-values and degrees of freedom are shown for the univariate tests. Contrast A was considered significant if its P value was less than the Bonferroni criterion of P < 0.025. F-values for the multivariate ANOVAs are based on Wilks' lambda (other tests gave similar results). Herbivore species are grouped by feeding guild. YEAR 1991 1992 1993 CONTRAST A B A B A B F(1,61) F(1,61) F(1,92) F(1,92) F(1,184) F(1,184) Multivariate 11.22? -- 11.28t -- 7.04? -- Leaf Miners Phyllonorycter salicifoliella 10.31"** 5.94* 2.06 0.68 7.27** 5.82*** Phyllocnistis sp. 4.24* 4.67* 13.02? 0.22 0.45 5.35* Leaf Gallers Phyllocolpa terminalis 18.35t 10.13"** 16.09? 10.26'** 36.91t 27.93? Phylocolpa spp. 2.18 0.31 6.13"* 0.03 1.66 11.75? Pontania sp. 0.87 8.60*** 4.28* 0.16 6.86** 1.82 lteomyia salicifolia 13.04? 13.50t 4.32* 0.69 16.95? 11.93? Aculops tetanothrix 13.97t 2.27 30.23t 7.62*** 11.88t 2.34 Leaf Folders Leaf folder-LF 4.49* 1.74 0.73 0.40 0.92 0.52 Leaf folder -V 16.99? 9.30*** 0.44 2.36 0.40 0.47 Stem Gallers Rabdophaga rigidae 1.99 0.21 0.50 0.47 2.13 0.22 salicisbrassicoides 3.24 0.06 4.77* 0.11 0.13 0.44 Total Herbivores 0.42 9.47*** 1.30 0.32 11.987 4.62* * - P < .05, ** P < .025, *** P < .01, # - P < .001 these years, densities on S. sericea and hybrids were equal and lower than on S. eriocephala. In 1992, however, the Hybrid Resistance hypothesis was supported. Mite gall density on hybrid plants was significantly lower than on S. sericea. It should be noted, however, that in all three years the numbers of mite galls were lower on the hybrid plants than on the nearest parent. For the leaf folding guild, LF supported the Additive hypothesis in all years (Tables 1 & 2), but the first contrast was marginally significant in 1991 (Table 1). Species V supported the Hybrid Susceptibility hypothesis in 1991, but the Additive hypothesis was supported in 1992 and 1993 (Tables 1 & 2). In the stem galling guild, both R. rigidae and R. salicisbrassicoides supported the Additive hypothesis consistently in each year (Tables 1 & 2), but densities of R. salicisbrassicoides in 1992 were marginally significant for the first contrast, nearly supporting the Dominance hypothesis. Seven of the 11 herbivore species showed significant changes in the hypotheses that were supported between years. In five of the seven cases, densities of the herbivores had declined between the years of the change; in two cases densities had increased or there was little change. Lower herbivore densities, therefore, might have contributed to the changes in which hypotheses were supported. 136
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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
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 120 and 121: From the point of view of trees, th
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 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
Page 170 and 171: RESULTS Seasonal Stability of Palat
Page 172 and 173: -
Page 174 and 175: OGAWA, A., NAGATA, Y., and SUKENO,
Page 176 and 177: 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 _
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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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