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Comparison of Insect, Fungal, and Mechanically Induced Defoliation of Larch: Effects on Plant Productivity and Subsequent Host Susceptibility

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Comparison of Insect, Fungal, and Mechanically Induced Defoliation of Larch: Effects on Plant Productivity and Subsequent Host Susceptibility Oecologia (1992) 90:411-416 Oecologia Springer-Verlag 1992 Comparison of insect, fungal, and mechanically induced defoliation of larch: effects on plant productivity and subsequent host susceptibility Steven C. Krause and Kenneth F. Raffa Department of Entomology, University of Wisconsin, Madison, WI 53706, USA Received October 16, 1991 / Accepted in revised form January 9, 1992 Summary. Larch sawfly, Pristiphora erichsonii Hartig, pathogens can also induce systemic antibiosis against and larch needlecast fungus, Mycosphaerella laricinia insect folivores (McIntyre et al. 1981), (R. Hartig) Neg., are early season defoliators restricted Fungal induced resistance to species of Coleoptera only to Larix host trees. Larch defoliation (100%) by (Ahmad et al. 1985), Hemiptera (Mathias et al. 1990), either the fungus or insect, but not mechanical removal, and Lepidoptera (Funk et al. 1983) has been demon- induced systemic responses that reduced sawfly con- strated for agronomic crops (Clay 1988). Trees infected sumption and digestion rates one year later. In a feeding with fungi may also be less susceptible to herbivory from behavior assay, larvae quickly abandoned seedlings gall-making species of Diptera (Bergdahl and Massola previously defoliated by M. laricinia. Adult female ovi- 1985), Homoptera (Lasota et al. 1983) and Hymenoptera position choice and egg deposition were unaffected. (Taper et al. 1986). Little is known, however, on the Seedling growth was not affected during the year of effects of fungal induced defoliation on free-feeding in- defoliation by M. laricinia, but was significantly reduced sect folivores and the productivity of host trees. one year later. Defoliation by M. laricinia reduced stem European larch, Larix decidua Miller, is a shade in- volume, radial growth, root biomass and new shoot tolerant, winter deciduous conifer (Olaczek 1986). After production. The latter tissue is the only oviposition re- spring needle growth, succulent branch shoots grow from source for larch sawfly, and, in contrast, is not influenced dormant buds for 4-6 weeks before becoming woody. by sawfly feeding. We hypothesize that M. laricinia infec- L. decidua was introduced into the United States in 1850 tion may limit larch sawfly populations where both spe- (Nyland 1965), and is now widely planted because of its cies coexist. However, this reduction is at a substantial rapid growth rate, high genetic diversity, and favorable net cost to larch productivity. wood quality (Einspahr et al. 1984). However, L. decidua is among the most susceptible species to defoliation by Key words: Larix decidua - Mycosphaerella laricinia - larch needlecast fungi (Ostry and Nichols 1989). Pristiphora erichsonii - Carbon allocation Larch needlecast, Mycosphaerella laricinia (R. Hartig) Neg., causes early season defoliation of larch in its native Europe (Hartig 1895), and was recently discovered in the United States (Patton and Spear 1983). Disease symp- Defoliation can negatively influence insect folivores by toms may appear before new- shoot growth is complete. reducing the quantity of plant tissue available for ovipo- Ascospores and conidia predominate in May-July, and sition and consumption, decreasing foliage quality August-September, respectively (Palmer and Ostry through induced defenses or nutrient depletion, and vari- 1986). The effects of M. laricinia defoliation on L. de- ous combinations thereof. Such defoliation-induced r productivity and co-occurring insect folivores are changes within host plants may partially regulate the unknown. population dynamics of some leaf-feeding insects (Bal- The larch sawfly, Pristiphora erichsonii Hartig tensweiler et al. 1977; Rhoades 1983). Although early (Hymenoptera: Tenthredinidae), is one of the major in- studies focused on the impact of plant changes on the sect defoliators on larch (Kulman 1971). Following its species causing plant injury (Whittaker and Feeny 1971 ; apparent introduction from Europe (Coppel and Leius Levin 1976), evidence now indicates that herbivory can 1955; but see Wong 1974) P. erichsonii contributed to affect interspecific associates and competitors (Suzuki widespread mortality of larch throughout the northern 1980; Karban et al. 1987). Local infections of plant United States and Canada (Nairn et al. 1962). Recent outbreaks of larch sawfly have occured on L. decidua in Offprint requests to." S. Krause Wisconsin and Canada (Drooz 1985). 412 Larch sawfly biology is closely tied to characteristics From 5-15 larvae (as needed) were placed on each seedling at any of its host. P. erichsonii is monophagous on larch, univol- one time, to achieve the observed fungal defoliation rates. The tine, and parthenogenetic. Adult females emerge in appropriate level of defoliation with hand shears was estimated spring from cocoons in the duff and oviposit only into visually. The entire treatment period required approximately 28 days. succulent, current year branch shoots. Larvae emerge Following the defoliations, all seedlings were transported to 7-10 days later, and feed on the foliage tufts present on Madison, W1 and held in greenhouses at ambient conditions. On older (1 + year) branches. Although needles on succulent 18 October, seedlings were moved outdoors to topless coldframes, shoots contain anti-feedant diterpenes (Wagner et al. where they were held for the winter. Seedlings infected with M. lari~ 1983), trees are not immune to complete defoliation einia were held under similar conditions nearby (6 km) to prevent (Drooz 1960). infection of the other seedlings. In spring 1988, trees within treatments were randomly assigned Current outbreaks of larch needlecast and larch saw- to larval development, larval behavior, or larch biomass productiv- fly on their native host in Wisconsin provided an oppor- ity analysis. tunity to determine whether fungal defoliation influences insect development and behavior and seedling productiv- Larval development. Recently molted fifth instar larvae were held ity. Larch seedlings were experimentally defoliated 12 h without food to allow emptying of their digestive tracts prior (100%) by sawfly larvae, fungi, and hand shears to assess to assay. Each larva was weighed (:/: 0.1 mg), and housed in a sawfly feeding performance and behavior. The root: transparent, plastic box (21.5 • 6.5 • 6.0 cm) with a known weight of foliage. After 48 h, the larva, frass and residual foliage were col- shoot biomass of seedlings was measured to assess defo- lected, oven dried for 72 h, and weighed. Larval consumption, liation effects on productivity. Two supplemental experi- digestion and feeding efficiency rates were determined gravimet- ments measured effects of artificial defoliation on sawfly rically (Waldbauer 1968). Experiments were performed in June in oviposition choice, and the susceptibility of larch trees to a growth chamber at 22-23 ~ C, 55-60% relative humidity and M. laricinia. constant light. There were 9 seedlings per treatment, with one insect per seedling. Consumption and food conversion rates (rag/day) were logl0 transformed, and digestibility arcsin (sqrt %) transformed to cor- Materials and methods rect heteroscedasticity of variances. Oneway analysis of variance (ANOVA) using the general linear models (GLM) procedure was Effect of defoliation agent on sawfly performance and conducted for each performance variable (SAS 1990). Mean separa- seedling growth tion tests were performed using least-squares means (LSM) (c~ = 0.05) (SAS 1990). A preliminary experiment demonstrated that Insects and seedlings. In October 1986, 3-year-old seedlings were these variables were not influenced by the solitary feeding con- bare-root lifted from the Department of Natural Resources state ditions used in this assay. nursery, at Boscobel, WI and placed in dark storage at 4-5 ~ C. In March 1987, trees were planted in 6 liter plastic pots containing Larval behavior. One second instar (4-5 d old) larva was placed on quartz sand : field soil: compost: sphagnum peat in a 7:1:1 : 1 ratio, each of three randomly selected branches per seedling in June. and held outdoors until use. Trees were watered regularly with a Larvae were isolated by filter paper barriers secured to the base of fertilizer solution of 17% ammoniacal nitrogen, 6% available phos- each experimental branch. Thus, larvae could either feed on one phoric acid, 6 % potash, and standard mixture of micronutrients at branch, or abandon the food source by dropping from the branch. 0.25 t per 3.8 liter water. A cardboard skirt coated with petroleum jelly extended from the Sawfiy larvae were reared from egg-laden shoots collected in stem to the drip line around each seedling to capture dispersing June, 1988 from a naturally infested L. deeidua stand in Clark larvae. Larvae were checked daily for 9 days. Because branches are County, W1 (T.24N.-R.4W., s. 18). Clipped shoots with eggs likely to have individual properties (Graham 1931 ; Gill and Halver- were placed in water until larval eclosion. Larvae were reared at son 1984), the time spent on a branch was recorded for each larva 21-23 ~ C, 40-60% relative humidity, 16 light: 8 dark, in plastic (N= 6). There were two seedlings per treatment. Insects and trees boxes (21.5 • 6.5 • 6.0 cm), and received fresh L. deeidua foliage and were kept in a walk-in chamber at 21-22 ~ C, 15 light: 9 dark, and a gentle spray of distilled water every 24-36 h (Heron and Drouin misted daily with distilled water. Data were log 10 transformed and 1969). analyzed using GLM and LSM (a=0.05) (SAS 1990). Defoliation treatments. In June 1987, 25 seedlings were randomly Larch productivity. Seedlings not used in either bioassay were har- assigned to each of four treatments. These consisted of 100 % defo- vested on 30-31 August 1988, 14 months after defoliation. Soil liation using M. larieinia, P. eriehsonii, or hand shears, or no was carefully excavated and washed from the root mass with cold defoliation (controls). Potted seedlings were transferred to the water. Roots had grown throughout the soil and were not pot appropriate field site for defofiation. A natural outbreak of M. lari- bound. Terminal leader growth was measured (ram) in 1987 and einia infection on L.
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