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.     

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.     

Anthropogenic changes to leaf litter input affect the fitness of a larval amphibian

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Restoration of threatened species: a noble cause for transgenic trees

Some of the first applications of transgenic trees in North America may be for the conservation or restoration of threatened forest trees that have been devastated by fungal pathogens or insect pests. In some cases, where resistance has yet to be found in the natural population of a tree species, incorporating genes from other organisms may offer the only hope for restoration. In others, transgenics may play a role as part of an integrated approach, along with conventional breeding or biocontrol agents. American chestnut (Castanea dentata) was wiped out as a canopy species by a fungal disease accidentally introduced into the United States around 1900. Similarly, American elm (Ulmus americana) virtually disappeared as a favored street tree from Northeastern U.S. cities after the introduction of the Dutch elm disease fungus in the 1940s. In both cases, progress has been made toward restoration via conventional techniques such as selection and propagation of tolerant cultivars (American elm) or breeding with a related resistant species (American chestnut). Recently, progress has also been made with development of systems for engineering antifungal candidate genes into these “heritage trees.” An Agrobacterium-leaf disk system has been used to produce transgenic American elm trees engineered with an antimicrobial peptide gene that may enhance resistance to Dutch elm disease. Two gene transfer systems have been developed for American chestnut using Agrobacterium-mediated transformation of embryogenic cultures, setting the stage for the first tests of potential antifungal genes for their ability to confer resistance to the chestnut blight fungus. Despite the promise of transgenic approaches for restoration of these heritage trees, a number of technical, environmental, economic, and ethical questions remain to be addressed before such trees can be deployed, and the debate around these questions may be quite different from that associated with transgenic trees developed for other purposes.     

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.     

Inadequate Cold Tolerance as a Possible Limitation to American Chestnut Restoration in the Northeastern United States

The American chestnut (Castanea dentata (Marshall) Borkh.), once a major component of eastern forests from Maine to Georgia, was functionally removed from the forest ecosystem by chestnut blight (an exotic fungal disease caused by Cryphonectria parasitica (Murr.) Barr), first identified at the beginning of the twentieth century. Hybrid‐backcross breeding programs that incorporate the blight resistance of Chinese chestnut (Castenea mollissima Blume) and Japanese chestnut (Castenea crenata Sieb. & Zuc.) into American chestnut stock show promise for achieving the blight resistance needed for species restoration. However, it is uncertain if limitations in tissue cold tolerance within current breeding programs might restrict the restoration of the species at the northern limits of American chestnut's historic range. Shoots of American chestnut and hybrid‐backcross chestnut (i.e., backcross chestnut) saplings growing in two plantings in Vermont were tested during November 2006, February 2007, and April 2007 to assess their cold tolerance relative to ambient low temperatures. Shoots of two potential native competitors, northern red oak (Quercus rubra L.) and sugar maple (Acer saccharum L.), were also sampled for comparison. During the winter, American and backcross chestnuts were approximately 5°C less cold tolerant than red oak and sugar maple, with a tendency for American chestnut to be more cold tolerant than the backcross chestnut. Terminal shoots of American and backcross chestnut also showed significantly more freezing damage in the field than nearby red oak and sugar maple shoots, which showed no visible injury.     

Modelling chestnut biogeography for American chestnut restoration

Aim Chestnuts (Castanea spp.) are ecologically and economically important species. We studied the general biology, distribution and climatic limits of seven chestnut species from around the world. We provided climatic matching of Asiatic species to North America to assist the range‐wide restoration of American chestnut [C. dentata (Marsh.) Borkh.] by incorporating blight‐resistant genes from Asiatic species. Location North America, Europe and East Asia. Methods General chestnut biology was reviewed on the basis of published literature and field observations. Chestnut distributions were established using published range maps and literature. Climatic constraints were analysed for the northern and southern distribution limits and the entire range for each species using principal component analysis (PCA) of fourteen bioclimatic variables. Climatic envelope matching was performed for three Chinese species using Maxent modelling to predict corresponding suitable climate zones for those species in North America. Results Chestnuts are primarily distributed in the warm‐temperate and subtropical zones in the northern hemisphere. PCA results revealed that thermal gradient was the primary control of chestnut distribution. Climatic spaces of different species overlap with one another to different degrees, but strong similarities are shown especially between Chinese species and American species. Climatic envelope matching suggested that large areas in eastern North America have a favourable climate for Chinese species. Main conclusions The general biological traits and climatic limits of the seven chestnut species are very similar. The predictions of Chinese species climatic range corresponded with most of the historical American chestnut range. Thus, a regionally adapted, blight‐resistant, introgressed hybrid American chestnut appears feasible if a sufficiently diverse array of Chinese chestnut germplasm is used as a source of blight resistance. Our study provided a between‐continent climate matching approach to facilitate the range‐wide species restoration, which can be readily applied in planning the restoration of other threatened or endangered species.     

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.     

Ectomycorrhizal characterization of an American chestnut <Emphasis Type="SmallCaps">(</Emphasis><Emphasis Type="Italic">Castanea dentata</Emphasis><Emphasis Type="SmallCaps">)</Emphasis>-dominated community in Western Wisconsin

Circa 1900, a farmer from the eastern US planted 11 American chestnut (Castanea dentata) seeds on a newly established farm near West Salem in western Wisconsin. These trees were very successful, producing a large stand of over 6,000 trees. Since this area is well outside the natural range of chestnut, these trees remained free from chestnut blight until 1987. In the West Salem stand, chestnuts are the dominant species of a mixed forest community, reminiscent of the chestnut–oak ecosystems of pre-1900 Appalachia. To identify putative mycorrhizal associates of chestnut in this unique forest, our approach was twofold: (1) an extensive fruiting body survey was conducted for four seasons that yielded approximately 100 putative mycorrhizal species and (2) a belowground molecular approach was used to generate DNA sequences of the internal transcribed spacer region from ectomycorrhizae. Unexpectedly, chestnut did not appear to be the dominant underground ectomycorrhizal-forming plant species. This study highlights the need to identify the plant host species when conducting belowground molecular-based surveys and provides preliminary identification of ectomycorrhizal fungi associated with a disjunct stand of American chestnut. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Community composition and diversity of riparian forests regulate decomposition of leaf litter in stream ecosystems

Riparian forest plantings are a well‐established restoration technique commonly used to stabilize banks and intercept nutrient flow from adjacent agricultural fields. Tree species planted for these efforts may not reflect mature forest communities within the same region. Given contemporary research on links between biodiversity and ecosystem functioning, we conducted a leaf‐litter decomposition study to investigate how mixing of detrital resources that reflect forest community composition would regulate in‐stream leaf litter. Leaf litter bags containing material from a mature forest (Liriodendron tulipifera, Acer rubrum, Quercus rubra, full factorial treatments = 7) and a restored riparian forest (Cornus sericea, Fraxinus pennsylvanica, Platanus occidentalis, full factorial treatments = 7) were deployed in a stream reach that experienced riparian reforestation in 2004. Litter from the restored riparian community had less mass remaining (45.28 ± 2.27%) than that from the mature riparian community (54.95 ± 2.19%) after 5 weeks. In addition, mixed litter treatments in the restored riparian community had less mass remaining (40.54 ± 2.37%) than single‐species treatments (51.80 ± 4.05%), a pattern not observed in the mature forest community. Results highlight the importance of planting mixed‐species assemblages as this structure may regulate processes such as decomposition and food‐web structure, processes often not targeted in the restoration plans.     

Leaf‐Litter Mixtures Affect Breakdown and Macroinvertebrate Colonization Rates in a Stream Ecosystem

Previous work in terrestrial and aquatic ecosystems has suggested that the relationship between breakdown rates of leaf litter and plant species richness may change unpredictability due to non‐additive effects mediated by the presence of key‐species. By using single‐ and mixed‐species leaf bags (7 possible combinations of three litter species differing in toughness; common alder [Alnus glutinosa ], sweet chestnut [Castanea sativa ], and Spanish oak [Quercus ilex ilex ]), I tested whether leaf species diversity, measured as richness and composition, affects breakdown dynamics and macroinvertebrate colonization (abundance, richness and composition) during 90 days incubation in a stream. Decomposition rates were additive, i.e., observed decomposition rates were not different from expected ones. However, decomposition rates of individual leaf species were affected by the mixture, i.e., there were species‐specific responses to mixing litter. The invertebrate communities colonizing the mixtures were not richer and more diverse in mixtures than in single‐species leaf bags. On the opposite, mixing leaf species had a negative, non‐additive effect on rates of shredder and taxa colonization and on macroinvertebrate diversity. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of plant litter diversity,species, origin and traits on larval toad performance

Decreased plant diversity is expected to reduce ecosystem function. Although many studies have examined effects of plant species on trophic interactions, information regarding effects of native or non‐native plant diversity on performance of individuals of higher trophic levels is limited. We reared larval American toad Anaxyrus americanus tadpoles in outdoor mesocosms containing litter of 1, 3, 6 or 12 plant species drawn randomly from a pool of 24 (15 native, 9 nonnative) species. Tadpole performance varied significantly among litter types in single litter treatments and pH and litter C:N were significant predictors of tadpole performance. Metamorphs were larger in mixtures than expected based on performance in single species treatments, suggesting a non‐additive effect of diversity. Litter diversity did not affect probability of survival or probability of metamorphosis. Plant origin (native or non‐native) had no significant effect on amphibian performance. Our study suggests some benefits to tadpole development at low levels of plant diversity, but questions assumed benefits of increased plant diversity and assumed detrimental effects of nonnative plant species for a common larval amphibian. Presence of specific plant species with strong negative effects on tadpole performance may outweigh diversity benefits in brown food webs.     

Impacts of cattle on amphibian larvae and the aquatic environment

• 1 Agricultural practices such as cattle farming may have direct or indirect negative effects on larval amphibians by decreasing water quality through deposition of nitrogenous waste, causing eutrophication, and grazing shoreline vegetation that contributes to detrital cover and food.
• 2 We sampled amphibian larvae on the Cumberland Plateau, Tennessee, U.S.A., twice per week, water quality twice per month and algal and detrital biomass once per month at seven wetlands (three cattle‐access and four non‐access) from March to August 2005 and 2006.
• 3 In general, species richness and diversity of amphibian larvae were greater in wetlands without cattle. Mean relative abundance of green frog (Rana clamitans) and American bullfrog (Rana catesbeiana) tadpoles was greater in non‐access wetlands. Body size of some ranid larvae was larger in cattle‐access wetlands but this trend did not exist for juveniles or adults. Dissolved oxygen was lower, while specific conductivity and turbidity were higher in cattle‐access wetlands. Mean biomass of detritus was lower in cattle‐access wetlands compared to non‐access wetlands; no differences were detected in algal biomass.
• 4 Given the negative impacts of cattle on water quality, detrital biomass, larval amphibian species richness and relative abundance of some amphibian species, we recommend that farmers consider excluding these livestock from aquatic environments.

     

Living in the litter: the influence of tree leaf litter on wetland communities

Empirical research in streams has demonstrated that terrestrial subsidies of tree leaf litter influence multiple community factors including composition, diversity and growth of individuals. However, little research has examined the importance of tree litter species on wetlands, which are ubiquitous across the landscape and serve as important habitats for a unique and diverse community of organisms. Using outdoor mesocosms, we assessed the impact of 12 litter monocultures and three litter mixtures (from both broadleaf and conifer trees) on pond communities containing gray tree frog tadpoles Hyla versicolor, periphyton, phytoplankton and zooplankton. We found that leaf litter species had substantial and differential impacts on all trophic groups in the community including effects on algal abundance, zooplankton density and amphibian growth. In many instances, patterns of responses were specific to individual litter species yet some responses, including both pH values and periphyton biomass, were generalizable to broad taxonomic groups. In addition, while most responses of litter mixtures were additive, we found evidence for antagonistic effects of litter mixing among responses of periphyton and amphibian body mass. Our results highlight the potential impact of human and naturally driven changes in forest composition on wetland communities through associated changes in leaf litter.     

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.     

Impact of site‐induced mouse caching and transport behaviour on regeneration in Castanea crenata

Abstract. Dispersal and retrieval site selection by mice, transport distance, cache depth, and emergence and survival of seedlings of Castanea crenata (Japanese chestnut) were investigated by a magnet‐locating experiment in two habitat conditions (gap vs. forest understorey). Magnets were inserted into nuts (n= 450) and the nuts placed in the edge of forest gaps. Although wood mice (Apodemus speciosus and A. argenteus) initially buried nuts singly in shallow surface caches near the nut source, by the following spring these cached nuts were retrieved and re‐cached in larger, deeper caches farther from the source, particularly in forest understories, probably to reduce the threat of pilferage. All the nuts cached in the forest understories were consumed, but 4 seedlings emerged in gaps, apparently because of lower foraging activity in the gaps by the mice. Seed size was not correlated with cache depth or cache site selection. With increasing seed size, transport distance increased, particularly in gaps, possibly due to a greater potential energy gain (relative to handling cost to the cacher), or to attempts to prevent density‐ or mass‐dependent loss of caches by other foragers. Variable seed dispersal behaviour based on variation in seed size may influence the chances of colonization and distribution of the light‐demanding Castanea trees in mosaic landscapes and may play an important role in community organization and dynamics.     

Recent successional processes investigated by pollen analysis of closedcanopy forest sites

Forest succession was investigated by pollen analysis of two mor-humus sections and of peat from a 3 m-diameter hollow under mixed conifer-hardwood forest in north-central Massachusetts, USA. The humus profiles recorded a major forest perturbation caused by the removal of Castanea dentata by the chestnut blight (1910–1912), and the peat from the hollow extended the record beyond the time of colonial settlement (1733). Fagus grandifolia was a forest dominant before 1733 but declined abruptly upon settlement. Castanea, a late Holocene immigrant to the area, rapidly increased its pollen representation after settlement until the epidemic of the chestnut blight. Forest succession following the loss of Castanea involved the successive rise to dominance of Betula, Quercus, Acer rubrum, and Tsuga canadensis. These vegetational changes conform to observations made during studies of forest-stand composition by other workers. Allogenic factors such as logging, disease, and wind have initiated major compositional change, which has been modified by autogenic successional processes such as the gradual rise to dominance of Tsuga canadensis around one of the humus sections. The two humus sites resolve fine-scale pattern in former vegetation such as differences in the distribution of Pinus strobus and Castanea over 200 m, the distance between the mor-humus sites. These within-forest sites permit investigations of fine-scale vegetational patterns and processes that are of interest to forest ecologists.     

Comparative chloroplast and nuclear DNA analysis of Castanea species in the southern region of the USA

Boundaries between American Castanea species (Castanea dentata, the American chestnut and C. pumila var. pumila, the Allegheny chinkapin, and var. ozarkensis, the Ozark chinkapin) have been difficult to establish because of intraspecific variation, interspecific similarities and the incidence of chestnut blight, which has prevented trees from maturing. In this study, informative chloroplast (cp) DNA and nuclear sequences from Castanea taxa were analyzed to gain a better understanding of their phylogeography in North America. Our emphasis has been on the most southern Castanea population in the Appalachian region, known for its morphological diversity. This Ruffner Mountain (Alabama) population shows a high number of unique haplotypes, which can be divided into two main groups. One group shares homology with the widespread and evolutionarily recent C. dentata haplotype. The other group shares homology with American chestnuts and Allegheny chinkapin taxa from southern states. This group has been the result of recent and more ancient cp capture and hybridization, indicative of hybrid zone clustering and glacial refugial origin. The range of C. pumila must have been more extensive along the Coastal Plains region, since only a few mutations separate the Ozark chinkapin from the main Allegheny chinkapin haplotype. The geographic origin of the American Castanea species complex appears to be in the Gulf Coast region.     

Synergistic effects of the invasive Chinese tallow (Triadica sebifera) and climate change on aquatic amphibian survival

Changes in climate and the introduction of invasive species are two major stressors to amphibians, although little is known about the interaction between these two factors with regard to impacts on amphibians. We focused our study on an invasive tree species, the Chinese tallow (Triadica sebifera), that annually sheds its leaves and produces leaf litter that is known to negatively impact aquatic amphibian survival. The purpose of our research was to determine whether the timing of leaf fall from Chinese tallow and the timing of amphibian breeding (determined by weather) influence survival of amphibian larvae. We simulated a range of winter weather scenarios, ranging from cold to warm, by altering the relative timing of when leaf litter and amphibian larvae were introduced into aquatic mesocosms. Our results indicate that amphibian larvae survival was greatly affected by the length of time Chinese tallow leaf litter decomposes in water prior to the introduction of the larvae. Larvae in treatments simulating warm winters (early amphibian breeding) were introduced to the mesocosms early in the aquatic decomposition process of the leaf litter and had significantly lower survival compared with cold winters (late amphibian breeding), likely due to significantly lower dissolved oxygen levels. Shifts to earlier breeding phenology, linked to warming climate, have already been observed in many amphibian taxa, and with most climate models predicting a significant warming trend over the next century, the trend toward earlier breeding should continue if not increase. Our results strongly suggest that a warming climate can interact with the effects of invasive plant species, in ways we have not previously considered, to reduce the survival of an already declining group of organisms.