Species‐Specific Barriers to Tree Regeneration in High Elevation Habitats of West Virginia

Herbaceous competition and herbivory have been identified as critical barriers to restoration of native tree species in degraded landscapes around the world; however, the combined effects of competition and herbivory are poorly understood. We experimentally manipulated levels of herbivory and herbaceous competition and analyzed the response of tree seedling performance over three growing seasons as a function of species and habitat in north‐central West Virginia. Four native tree species were planted in old field and forest experimental plots: Castanea dentata (American chestnut), Quercus rubra (red oak), Acer saccharum (sugar maple), and Picea rubens (red spruce). Red spruce demonstrated the highest growth increment and greatest survival (64%) and most consistent results among treatments and habitats. Red spruce survival was not reduced in the presence of Odocoileus virginianus (white‐tailed deer) browse and herbaceous competition; however, growth was improved by suppression of herbaceous competition. We suspect that this deciduous forest landscape would regenerate to a red spruce dominated forest if seed source was available. In contrast, the other three species tested had very low survival when exposed to deer and were more responsive to competing vegetation and habitat type. American chestnut had low survival and growth across all treatments, suggesting basic climate limitations. Vigorous natural regeneration of Prunus serotina (black cherry) occurred in forest plots where both competing herbs and deer were excluded. Our results demonstrated the importance of testing multiple potential recruitment barriers and species at once and the need for species and habitat‐specific restoration treatments.     

Growth traits of juvenile American chestnut and red oak as adaptations to disturbance

American chestnut (Castanea dentata) was a dominant species in eastern North America prior to the importation of chestnut blight. In light of recent efforts to restore viable populations of chestnut in eastern forests, an increased understanding of its association with other co‐occurring, disturbance‐adapted oak species is necessary. We evaluated crown architecture and leaf morphology in juvenile chestnut and red oak (Quercus rubra) to assess potential differences in establishment strategies of both species. We also investigated differences in nonstructural carbohydrate reserves and whole tree biomass partitioning between species. Seedlings of both species were planted in forest stands treated either with midstory removal or small patch cuts, simulating potential restoration plantings. After 5–7 years, chestnut's allocation to its root system was lower than red oak's, with chestnut saplings instead diverting resources to branches and foliage. Chestnut had lower leaf area index, greater crown projection area, and higher specific leaf area than red oak, indicating the species may have an advantage in shaded understories. There were only minor differences in nonstructural root carbohydrate reserves, between red oak and American chestnut, indicating that chestnut may respond similarly to oak by resprouting after disturbances topkill young saplings. We suggest that American chestnut has morphological and physiological attributes that allow it to function as an opportunistic and plastic species that can utilize gaps to facilitate its canopy recruitment, yet still persist after occasional surface fire. This knowledge can guide restoration strategies for this iconic species of the eastern temperate forest region.     

Increasing Red Maple Leaf Litter Alters Decomposition Rates and Nitrogen Cycling in Historically Oak-Dominated Forests of the Eastern U.S.

Without canopy-opening fire disturbances, shade-tolerant, fire-sensitive species like red maple (Acer rubrum L.) proliferate in many historically oak-dominated forests of the eastern U.S. Here, we evaluate potential implications of increased red maple dominance in upland oak forests of Kentucky on rates of leaf litter decomposition and nitrogen (N) cycling. Over 5 years, we evaluated mass loss of leaf litter and changes in total N and carbon (C) within six leaf litter treatments comprised of scarlet oak, chestnut oak, and red maple, and three mixed treatments of increasing red maple contribution to the leaf litter pool (25, 50, and 75% red maple). Over a 1.5-year period, we conducted a plot-level leaf litter manipulation study using the same treatments plus a control and assessed changes in net nitrification, ammonification, and N mineralization within leaf litter and upper (0–5 cm depth) mineral soil horizons. Red maple leaf litter contained more “fast” decomposing material and initially lost mass faster than either oak species. All litter treatments immobilized N during initial stages of decomposition, but the degree of immobilization decreased with decreasing red maple contribution. The leaf litter plot-level experiment confirmed slower N mineralization rates for red maple only plots compared to chestnut oak plots. As red maple increases, initial leaf litter decomposition rates will increase, leading to lower fuel loads and more N immobilization from the surrounding environment. These changes may reduce forest flammability and resource availability and promote red maple expansion and thereby the “mesophication” of eastern forests of the U.S.     

Exposure to herbivores increases seedling growth and survival of American chestnut (Castanea dentata) through decreased interspecific competition in canopy gaps

The successful development of early stages of blight‐resistant hybrid stock has increased hopes for restoration of American chestnut (Castanea dentata) to eastern North American forests. However, these forests have undergone substantial ecological change in the century since the functional extirpation of American chestnut, and it remains unknown to what extent American chestnut will be able to recolonize contemporary forests. In particular, high densities of white‐tailed deer (Odocoileus virginianus) and competition with mesophytic tree species such as maple (Acer) may impede chestnut regeneration, much as they affect oak (Quercus). We used a split‐plot analysis of variance (ANOVA) design to examine the effects of canopy gaps and herbivory on survival and growth of third generation backcrossed (BC₃) hybrid chestnut seedlings over two growing seasons in central Indiana, U.S.A. Only 4 of 588 (0.7%) seedlings in closed‐canopy plots survived to the end of the study, as opposed to 264 of 589 (45%) seedlings in gap plots. Within the gap treatment, fencing was associated with reduced chestnut survival as well as reduced herbivory and increased cover of non‐chestnut vegetation. Our results indicate that herbivory may indirectly benefit chestnut regeneration by suppressing competition. However, this beneficial effect is likely context‐dependent and additional work is needed to establish the conditions under which it occurs.     

Interactions between a fungal pathogen, foliar properties, and generalist herbivores

American [Castanea dentata (Marsh) Borkh.] × Chinese [Castanea mollissima Blume] chestnut (Fagac, ae) hybrids are a novel system in which to study influences of phytopathogenic fungi and woody plant hybridization on herbivore susceptibility, as the hybrids are well characterized with regard to resistance to the chestnut blight fungus [Cryphonectria parasita (Murr) Barr (Endothia) Diaporthales: Valsaceae] and variability is present. We chose two groups of resistance‐rated backcross chestnut that shared an F1 parent and had different American parents. Foliage from both backcross groups and the parent trees was sampled on three dates for use in feeding assays with gypsy moth larvae [Lymantria dispar (L.) [Lepidoptera: Lymantriidae], adult Japanese beetles [Popillia japonica Newman (Coleoptera: Scarabaeidae)], and fall webworm larvae [Hyphantria cunea Drury (Lepidoptera: Arctiidae)], respectively. Foliar analyses were performed concurrently and included carbohydrate, tannin, and nitrogen content, toughness, and density. Blight resistance had almost no effect on herbivore performance or foliar chemistry. When the parent trees and backcross groups were compared, however, significant differences in gypsy moth performance and Japanese beetle consumption were evident. There were no differences in fall webworm consumption. Most foliar characteristics measured differed among chestnut genotypes at some point in the season, and all varied seasonally. No clear pattern emerges with respect to the relationship among blight resistance, herbivore susceptibility, foliar properties, and plant genotype, and more research is needed to separate these effects.     

Shredder abundance and leaf breakdown in an Appalachian Mountain stream

SUMMARY.

• 1 Breakdown rates of dogwood (Cornus florida L.), red maple (Acer rubrum L.) and white oak (Quercus alba L.) leaves were investigated at two first-order and two second-order sites in an Appalachian Mountain stream.
• 2 Leaves exposed in mesh bags were sampled on eight occasions over a 207 day period and breakdown rates were compared using an exponential decay model.
• 3 There was a consistent ranking in leaf breakdown rate within each site, i.e. dogwood > red maple > white oak, and all species broke down faster at second-than at first-order sites.
• 4 Our data suggest that differences in species-specific leaf breakdown rates were largely a function of shredder abundance on the leaves.

     

Influence of Tree Species on Forest Nitrogen Retention in the Catskill Mountains, New York, USA

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day ¹⁵N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot ¹⁵N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for ¹⁵N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. ¹⁵Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in ¹⁵N recovery as a result of N fertilization. Aboveground biomass became a significantly greater ¹⁵N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.     

The expression of dehydrin proteins in desiccation-sensitive (recalcitrant) seeds of temperate trees

Proteins that have homology with dehydrins have been identified immunologically in the desiccationsensitive (recalcitrant) seeds of English oak (Quercus robur L.), European chestnut (Castanea sativa L.), horse chestnut (Aesculus hippocastanum L.), sycamore (Acer psuedoplatanus L.) and silver maple (Acer saccharinum L.), and in the desiccation-tolerant seeds of Norway maple (Acer platanoides L.). The mRNA for a late embryogenesis abundant (LEA) protein (dehydrin) was also detected by Northern blotting, using a cDNA clone (D11) from cotton embryos, in the recalcitrant and orthodox seeds. Medium-stringency washing was required to detect this hybridization. InQ. robur the amount of dehydrin protein increased during seed development, andLEA mRNA was induced by limited desiccation and by abscisic acid. Confirmation of the presence of dehydrin mRNA in matureQ. robur andC. sativa seeds was obtained by in-vitro translation of the extracted polyadenylated RNA followed by analysis of the immunoprecipitation products. Thus the presence of dehydrin proteins is not sufficient to confer desiccation tolerance on truly recalcitrant seeds, nor can their presence or absence be used as clear criteria for identification of recalcitrant seeds.     

The phosphorus status of northern hardwoods differs by species but is unaffected by nitrogen fertilization

Northern hardwood forests in the eastern US exhibit species-specific influences on nitrogen (N) cycling, suggesting that their phosphorus (P) cycling characteristics may also vary by species. These characteristics are increasingly important to understand in light of evidence suggesting that atmospheric N deposition has increased N availability in the region, potentially leading to phosphorus limitation. We examined how P characteristics differ among tree species and whether these characteristics respond to simulated N deposition (fertilization). We added NH₄NO₃ fertilizer (50 kg ha^?1 year^?1) to single-species plots of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.), eastern hemlock (Tsuga canadensis (L.) Carr.), American beech (Fagus grandifolia Ehrh.), and yellow birch (Betula alleghaniensis Britt.), in the Catskill Mountains, New York from 1997 to 2007. Species differences were observed in foliar, litter and root P concentrations, but all were unaffected by a cumulative N fertilization of 550 kg/ha. Similarly, measures of soil P availability and biotic P sufficiency differed by species but were unaffected by N fertilization. Results suggest species exhibit unique relationships to P as well as N cycles. We found little evidence that N fertilization leads to increased P limitation in these northern hardwood forests. However, species such as sugar maple and red oak may be sufficient in P, whereas beech and hemlock may be less sufficient and therefore potentially more sensitive to future N-stimulated P limitation.     

Branch growth and leaf numbers of red maple (Acer rubrum L.) and red oak (Quercus rubra L.): response to defoliation

Summary Branch growth and leaf formation from terminal and from lateral buds of red maple (Acer rubrum L.) and red oak (Quercus rubra L.) were measured in response to simulated insect defoliation. A single large branch representative of the crown of each tree was used for enumeration of growth and of bud numbers throughout three successive years of 0, 50, 75, and 100% leaf removal for the entire tree. Leaf number per tree for both species after the last year of defoliation was reduced in direct proportion to the severity of defoliation, in comparison to the predefoliation status of the trees. Bud number per tree for red maple, but not for red oak, was also reduced in proportion to severity of defoliation.Averaged over all defoliation treatments, defoliation reduced branch growth more than leaf production. Furthermore, the reduction in branch growth and leaf production was greater in red oak than in red maple. Three years of successive defoliation reduced the mean lateral plus terminal branch growth by 40% in red oak and by 23% in red maple, while leaf number was reduced 22% in red oak and remained unchanged in red maple. In red maple, 100% defoliation caused greater branch death than the 50 or 75% defoliation treatments, and the amount of death was greater after each successive year of defoliation. In contrast to red maple, undefoliated red oak incurred a substantial amount of branch death throughout the study which was little affected by defoliation treatment.     

Multiple interacting ecosystem drivers: toward an encompassing hypothesis of oak forest dynamics across eastern North America

Many forests of eastern North American are undergoing a species composition shift in which maples (Acer spp.) are increasingly important while oak (Quercus spp.) regeneration and recruitment has become increasingly scarce. This dynamic in species composition occurs across a large and geographically complex region. The elimination of fire has been postulated as the driver of this dynamic; however, some assumptions underlying this postulate have not been completely examined, and alternative hypotheses remain underexplored. Through literature review, and a series of new analyses, we examined underlying assumptions of the “oak and fire” hypothesis and explored a series of alternative hypotheses based on well‐known ecosystem drivers: climate change, land‐use change, the loss of foundation and keystone species, and dynamics in herbivore populations. We found that the oak–maple dynamic began during a shift in climate regime‐from a time of frequent, severe, multi‐year droughts to a period of increased moisture availability. Anthropogenic disturbance on the landscape changed markedly during this same time, from an era of Native American utilization, to a time characterized by low population densities, to Euro‐American settlement and subsequent land transmogrification. During the initiation of the oak‐maple dynamic, a foundation species, the American chestnut, was lost as a canopy tree across a broad range. Several important browsers and acorn predators had substantial population dynamics during this period, e.g. white‐tailed deer populations grew substantially concurrent with increasing oak recruitment failure. In conclusion, our analyses suggest that oak forests are reacting to marked changes in a suite of interlocking factors. We propose a “multiple interacting ecosystem drivers hypothesis”, which provides a more encompassing framework for understanding oak forest dynamics.     

Natural and experimentally altered hydraulic architecture of branch junctions in Acer saccharum Marsh. and Quercus velutina Lam. trees

 The functional xylem anatomy and the hydraulic conductivity of intact and treated branch junctions of the diffuse-porous sugar maple (Acer saccharum Marsh.) were compared to those of the ring-porous black oak (Quercus velutina Lam.). Maple shoots possessed greater growth intensity than those of oak. The extensive growth of the maple trees resulted in about a two-fold increase in xylem production in the maple branches. Branches were altered by removing a patch of bark from the base of a branch (near a junction) leaving a bridge of bark on the upper or lower side of the branch. The experimentally treated branch junctions revealed that, in oak, most (up to 92%) of the water flows in the lower side of a branch, where most of the large vessels occurred. In maple, most of the conductive tissue was observed to form in the upper side of the branches, which was equally or more conductive than the lower side. A treatment of longitudinal, parallel scratches in the bark-bridge, which reduced earlywood vessel width, substantially decreased conductivity (to only 15%) in oak, but had no effect on conductivity in maple. In maple, such wounding stimulated more wood formation and increased conductivity. In both trees, a narrow bridge at the junction induced more wood formation and higher conductivity in the branch. The mechanisms controlling wood formation and water flow in branch junctions of ring- and diffuse-porous trees are discussed. Received: 14 February 1996 / Accepted: 27 May 1996     

TEMPORAL AND SPATIAL PATTERNS OF SPECIFIC LEAF WEIGHT IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Temporal and spatial patterns of specific leaf weight (SLW, g/m²) were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan to evaluate the utility of SLW as an index of leaf photosynthetic capacity. No significant diurnal changes in SLW were found. Specific leaf weight decreased and then increased during leaf expansion in the spring. Most species, especially those located in the understory, then had relatively constant SLW for most of the growing season, followed by a decline in SLW during autumn. Specific leaf weight decreased exponentially down through the canopy with increasing cumulative leaf area index. Red oak (Quercus rubra), paper birch (Betula papyrifera), bigtooth aspen (Populus grandidentata), red maple (Acer rubrum), sugar maple (A. saccharum), and beech (Fagus grandifolia) generally had successively lower SLW, for leaves at any one level in the canopy. On a given site, comparisons between years and comparisons of leaves growing within 35 cm of each other showed that differences in SLW among species were not due solely to microenvironmental effects on SLW. Bigtooth aspen, red oak, and red maple on lower-fertility sites had lower SLW than the same species on higher-fertility sites. Maximum CO₂ exchange rate, measured at light-saturation in ambient CO₂ and leaf temperatures of 20 to 25 C, increased with SLW. Photosynthetic capacities of species ranked by SLW in a shaded habitat suggest that red oak, red maple, sugar maple, and beech are successively better adapted to shady conditions.     

Population dynamics and growth patterns for a cohort of northern red oak (Quercus rubra) seedlings

Summary I studied the survival and development of a 1986 cohort of northern red oak (Quercus rubra L.) seedlings growing under a variety of overstory and microsite conditions in a northern hardwood forest dominated by northern red oak, red maple (Acer rubrum L.) paper birch (Betula papyrifera Marsh.), and scattered white pine (Pinus strobus L.). Fifty naturally regenerating seedlings of oak were randomly selected in each of three canopy classes: no overstory, partial overstory, and complete overstory. Growth and mortality were measured for six years. Seedling height growth decreased with overstory density, with less growth evident with even a partial overstory. Seedling survival also declined with overstory density and depended on microtopography to a lesser extent. After six years, 92% of the seedlings survived in the open, compared to 54% under the partial overstory, and 36% under the complete overstory. The open environment, in which woody and herbaceous regrowth formed a low canopy reducing light intensities to about 50% of full sunlight, provided a favorable site for the growth and survival of northern red oak.     

SPECIES RESPONSE TO A MOISTURE GRADIENT IN CENTRAL ILLINOIS FORESTS

Polar and Gaussian ordination applied to data collected from 37 forest sites in central Illinois resulted in a continuous and gradual change in species composition along a moisture gradient. A series of overlapping species success curves formed by plotting Importance Values over stands ordered along the gradient varied continuously in modal location and habitat width. Blackjack oak and black oak dominated upland sandy sites. Black oak, white oak, and shagbark hickory were the most important species on exposed, upper slope positions or ridge tops with silt-loam soils. Red oak, sugar maple, American elm, and bur oak dominated sheltered locations on lower slope positions and stream terraces. Sycamore, silver maple, and cottonwood were leading tree species in floodplain forests. Conversion of black, white, and red oak forests on silt-loam sites to sugar maple, white ash, and red elm dominance is evident by high densities of these shade tolerant species in the understory. Composition of forests at the extreme ends of the moisture gradient is more stable than the mesic sites. Maximum tree diversity occurred on mesic sites and decreased toward the extreme ends of the moisture gradient. However, competitive exclusion of shade intolerant species by sugar maple and other species has caused a decrease in understory diversity on mesic sites. Diversity decreased from canopy to understory strata in lowland forests and increased on xeric sites.     

Comparative efficacy of gypsy moth (Lepidoptera: Erebidae) entomopathogens on transgenic blight‐tolerant and wild‐type American,Chinese, and hybrid chestnuts (Fagales: Fagaceae)

American chestnut (Castanea dentata [Marsh.] Borkh.) was once the dominant hardwood species in Eastern North America before an exotic fungal pathogen, Cryphonectria parasitica (Murrill) Barr, functionally eliminated it across its range. One promising approach toward restoring American chestnut to natural forests is development of blight‐tolerant trees using genetic transformation. However, transformation and related processes can result in unexpected and unintended phenotypic changes, potentially altering ecological interactions. To assess unintended tritrophic impacts of transgenic American chestnut on plant–herbivore interactions, gypsy moth (Lymantria dispar L.) caterpillars were fed leaf disks excised from two transgenic events, Darling 54 and Darling 58, and four control American chestnut lines. Leaf disks were previously treated with an LD₅₀ dose of either the species‐specific Lymantria dispar multiple nucleopolyhedrovirus (LdMNPV) or the generalist pathogen Bacillus thuringiensis subsp. kurstaki (Btk). Mortality was quantified and compared to water blank controls. Tree genotype had a strong effect on the efficacies of both pathogens. Larval mortality from Btk‐treated foliage from only one transgenic event, Darling 54, differed from its isogenic progenitor, Ellis 1, but was similar to an unrelated wild‐type American chestnut control. LdMNPV efficacy was unaffected by genetic transformation. Results suggest that although genetic modification of trees may affect interactions with other nontarget organisms, this may be due to insertion effects, and variation among different genotypes (whether transgenic or wild‐type) imparts a greater change in response than transgene presence.     

Selection,caching, and consumption of hardwood seeds by forest rodents: implications for restoration of American chestnut

Recent field trials on blight‐resistant hybrids (BC₃F₃) of American chestnut (Castanea dentata) and Chinese chestnut (C. mollissima) have intensified planning for widespread restoration of Castanea to eastern U.S. forests. Restoration will likely rely on natural seed dispersal from sites planted with chestnut; however, we do not know how dispersal agents such as granivorous rodents will respond to hybrid chestnuts. At one extreme, excessive seed consumption may impede restoration. Alternatively, scatter‐hoarding rodents might facilitate the spread of chestnut by dispersal of seeds from restoration plantings. We conducted trials with five rodent species to quantify foraging preferences and to evaluate the potential role of granivores in chestnut restoration. Specifically, we presented seeds from American and hybrid chestnuts (BC₃F₂) with other common mast species and recorded the proportion of seeds removed and the fates of tagged seeds. Mice, chipmunks, and flying squirrels harvested both chestnut types preferentially over larger, tougher black walnut, hickory, and red oak seeds, but fox squirrels and eastern gray squirrels preferred larger seeds to chestnuts. All rodents consumed a greater proportion of the chestnuts than other seed types. American and hybrid chestnut also differed in important ways: except for fox squirrels, rodents preferentially removed American chestnuts over hybrid chestnuts, but we estimated that fox squirrels carried a greater proportion of hybrid chestnuts beyond our tag search area, suggesting that hybrids may be dispersed farther and cached more often than American chestnut. These differences indicate that hybrid chestnut may not be functionally equivalent to American chestnut with regard to seed–granivore interactions.     

Units of freezing of deep supercooled water in woody xylem

The low temperature exotherms (LTE) of 1-year-old twigs of Haralson apple (Malus pumila Mill.), shagbark hickory (Carya ovata [Mill.] K. Koch), green ash (Fraxinus pennsylvanica Marsh), honey locust (Gleditsia triacanthos L.), American chestnut (Castanea dentata [Marsh] Borkh.), and red oak (Quercus rubra L.) were determined by differential thermal analysis (DTA). In one type of experiment freezing during a DTA experiment was halted for up to 2.5 hours after part of the supercooled water had frozen at temperatures between −25 and −42 C. Upon resumption of cooling the freezing started within 2 C of the stopping temperature. In a second type of experiment living and dead cells were microscopically observed in the same ray after partial freezing in the DTA apparatus. In another experiment, the LTE persisted even after tangential and radial sectioning of the twig to 0.13 millimeters. In a final experiment the LTE of a single multiseriate ray of red oak had the same shape as the LTE of wood with many uniseriate rays.     

Tree resistance to Lymantria dispar caterpillars: importance and limitations of foliar tannin composition

The ability of foliar tannins to increase plant resistance to herbivores is potentially determined by the composition of the tannins; hydrolyzable tannins are much more active as prooxidants in the guts of caterpillars than are condensed tannins. By manipulating the tannin compositions of two contrasting tree species, this work examined: (1) whether increased levels of hydrolyzable tannins increase the resistance of red oak (Quercus rubra L.), a tree with low resistance that produces mainly condensed tannins, and (2) whether increased levels of condensed tannins decrease the resistance of sugar maple (Acer saccharum Marsh.), a tree with relatively high resistance that produces high levels of hydrolyzable tannins. As expected, when Lymantria dispar L. caterpillars ingested oak leaves coated with hydrolyzable tannins, levels of hydrolyzable tannin oxidation increased in their midgut contents. However, increased tannin oxidation had no significant impact on oxidative stress in the surrounding midgut tissues. Although growth efficiencies were decreased by hydrolyzable tannins, growth rates remained unchanged, suggesting that additional hydrolyzable tannins are not sufficient to increase the resistance of oak. In larvae on condensed tannin-coated maple, no antioxidant effects were observed in the midgut, and levels of tannin oxidation remained high. Consequently, neither oxidative stress in midgut tissues nor larval performance were significantly affected by high levels of condensed tannins. Post hoc comparisons of physiological mechanisms related to tree resistance revealed that maple produced not only higher levels of oxidative stress in the midgut lumen and midgut tissues of L. dispar, but also decreased protein utilization efficiency compared with oak. Our results suggest that high levels of hydrolyzable tannins are important for producing oxidative stress, but increased tree resistance to caterpillars may require additional factors, such as those that produce nutritional stress.     

Environmentally-based maternal effects: a hidden force in insect population dynamics?

Summary The nutritional environment of the parental generation of the polyphagous gypsy moth, Lymantria dispar, can significantly influence the growth and reproductive potential of the next generation through environmentally-based maternal effects. In the first experiment, members of the parental generation were reared on red oak trees (Quercus rubra L.) with known defoliation and phenolic levels. Diet in the offspring generation was homogeneous (synthetic diet). With genetic effects accounted for 1) offspring attained greater pupal weights when their mothers fed on trees with higher leaf damage levels, 2) daughters had a shorter prefeeding stage, a trait associated with dispersal tendency, when their mothers experienced higher condensed tannin levels, and 3) sons had lower pupal weights when their mothers experienced greater condensed tannin levels. In the second experiment, members of the parental generation were reared on either red or black oak (Q. velutina) trees. Offspring of each mother were divided among four diets: red oak, chestnut oak (Q. prinus L.), a standard synthetic diet, and a low-protein synthetic diet. The parental host species accounted for 24% of the variation in daughters' development time; offspring diet accounted for 52%. When mothers were reared on black oak rather than red oak, their offspring developed significantly faster when the F1 diet was chestnut oak. Environmentally-based maternal effects can significantly influence the expression of offspring dispersal potential, growth rate, and offspring fecundity. These traits contribute to natality and survival in natural populations and, hence, to population growth potential. Theoretical models of insect population dynamics demonstrate that the presence of a time delay in a density dependent response can induce destabilization. Maternal effects act on a time delay and may participate in the destabilization characteristic of outbreak species.