The Genetic Diversity of Native, Stocked, and Hybrid Populations of Brook Trout in the Southern Appalachians

Using mitochondrial DNA (mtDNA) restriction analysis and starch-gel electrophoresis of the CK-A2 locus product, we examined genetic variation in 311 brook trout (Salvelinus fontinalis) from 11 native, 5 hatchery-derived, and 8 hybrid populations in the southern Appalachian Mountains. Native southern Appalachian fish were genetically distinct from hatchery-derived fish. Southern and hatchery-derived fish were reliably distinguished based on three mtDNA restriction sites. Native southern haplotypes differed from hatchery-derived haplotypes by an average of 0.84%. Northern hatchery-derived haplotypes varied little in mtDNA compared to native southern haplotypes. Introgression of mtDNA haplotypes and the CK-A2 locus varied among populations, and introgression of allozyme and mtDNA markers was positively correlated. Continued introductions of nonnative strains of brook trout in the southern Appalachians could simplify the genetic structure of native brook trout populations and eliminate unique genotypes.     

The influence of successional processes and disturbance on the structure of Tsuga canadensis forests

Old-growth forests are valuable sources of ecological, conservation, and management information, yet these ecosystems have received little study in New England, due in large part to their regional scarcity. To increase our understanding of the structures and processes common in these rare forests, we studied the abundance of downed coarse woody debris (CWD) and snags and live-tree size-class distributions in 16 old-growth hemlock forests in western Massachusetts. Old-growth stands were compared with eight adjacent second-growth hemlock forests to gain a better understanding of the structural differences between these two classes of forests resulting from contrasting histories. In addition, we used stand-level dendroecological reconstructions to investigate the linkages between disturbance history and old-growth forest structure using an information-theoretic model selection framework. Old-growth stands exhibit a much higher degree of structural complexity than second-growth forests. In particular, old-growth stands had larger overstory trees and greater volumes of downed coarse woody debris (135.2 vs. 33.2 m3/ha) and snags (21.2 vs. 10.7 m3/ha). Second-growth stands were characterized by either skewed unimodal or reverse-J shaped diameter distributions, while old-growth forests contained bell-shaped, skewed unimodal, rotated sigmoid, and reverse J-shaped distributions. The variation in structural attributes among old-growth stands, particularly the abundance of downed CWD, was closely related to disturbance history. In particular, old-growth stands experiencing moderate levels of canopy disturbance during the last century (1930s and 1980s) had greater accumulations of CWD, highlighting the importance of gap-scale disturbances in shaping the long-term development and structural characteristics of old-growth forests. These findings are important for the development of natural disturbance-based silvicultural systems that may be used to restore important forest characteristics lacking in New England second-growth stands by integrating structural legacies of disturbance (e.g., downed CWD) and resultant tree-size distribution patterns. This silvicultural approach would emulate the often episodic nature of CWD recruitment within old-growth forests.     

Estimation of Core Terrestrial Habitat for Stream-Breeding Salamanders and Delineation of Riparian Buffers for Protection of Biodiversity

Abstract:  Many species of wildlife depend on riparian habitats for various life-history functions (e.g., breeding, foraging, overwintering). Although this unique habitat is critical for many species, delineations of riparian zones and buffers for various taxa are lacking. Typically when buffer zones are determined to mitigate edge effects, they are based on criteria that protect aquatic resources alone and do not consider impacts to wildlife and other terrestrial resources. Using two different survey methods (area-constrained daytime searches and nighttime visual encounter searches), we estimated core terrestrial habitat and buffer widths for stream-breeding salamanders in southern Appalachian streams from May to August 2004. A core terrestrial habitat of 27.0 m encompassed 95% of the salamander assemblage (four species of stream plethodontids), and an additional 50 m (to buffer edge effects) yielded a total buffer of 77.0 m. When each species of the assemblage was analyzed separately, the maximum core terrestrial habitat needed for the Blue Ridge two-lined salamander ( Eurycea wilderae ), a dominant member and the farthest-ranging species from the stream, was 42.6 m. Thus, we recommend an overall buffer width of 92.6 m in southern Appalachian streams. To protect stream amphibians and other wildlife dependent on riparian areas, land managers and policy makers must consider conserving more than aquatic resources alone. Developing core terrestrial habitat estimates and buffer zone widths for wildlife populations is a critical first step in the conservation of many semiaquatic organisms and protecting biodiversity.     

Exploitative competition between invasive herbivores benefits a native host plant

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).     

A climatic driver for abrupt mid-Holocene vegetation dynamics and the hemlock decline in New England

The mid-Holocene decline of eastern hemlock is widely viewed as the sole prehistorical example of an insect- or pathogen-mediated collapse of a North American tree species and has been extensively studied for insights into pest-host dynamics and the consequences to terrestrial and aquatic ecosystems of dominant-species removal. We report paleoecological evidence implicating climate as a major driver of this episode. Data drawn from sites across a gradient in hemlock abundance from dominant to absent demonstrate: a synchronous, dramatic decline in a contrasting taxon (oak); changes in lake sediments and aquatic taxa indicating low water levels; and one or more intervals of intense drought at regional to continental scales. These results, which accord well with emerging climate reconstructions, challenge the interpretation of a biotically driven hemlock decline and highlight the potential for climate change to generate major, abrupt dynamics in forest ecosystems.     

Processesof Tamarix invasion and floodplain development along the lower Green River, Utah.

Significant ecological, hydrologic, and geomorphic changes have occurred during the 20th century along many large floodplain rivers in the American Southwest. Native Populus forests have declined, while the exotic Eurasian shrub, Tamarix, has proliferated and now dominates most floodplain ecosystems. Photographs from late 19th and early 20th centuries illustrate wide river channels with largely bare in-channel landforms and shrubby higher channel margin floodplains. However, by the mid-20th century, floodplains supporting dense Tamarix stands had expanded, and river channels had narrowed. Along the lower Green River in eastern Utah, the causal mechanism of channel and floodplain changes remains ambiguous due to the confounding effects of climatically driven reductions in flood magnitude, river regulation by Flaming Gorge Dam, and Tamarix invasion. This study addressed whether Tamarix establishment and spread followed climate- or dam-induced reductions in annual peak flows or whether Tamarix was potentially a driver of floodplain changes. We aged 235 Tamarix and 57 Populus individuals, determined the hydrologic and geomorphic processes that controlled recruitment, identified the spatial relationships of germination sites within floodplain stratigraphic transects, and mapped woody riparian vegetation cohorts along three segments of the lower Green River. The oldest Tamarix established along several sampling reaches in 1938, and 1.50-2.25 m of alluvium has accreted above their germination surfaces. Nearly 90% of the Tamarix and Populus samples established during flood years that exceeded the 2.5-year recurrence interval. Recruitment was most common when large floods were followed by years with smaller peak flows. The majority of Tamarix establishment and Green River channel narrowing occurred long before river regulation by Flaming Gorge Dam. Tamarix initially colonized bare instream sand deposits (e.g., islands and bars), and most channel and floodplain changes followed the establishment of Tamarix. Our results suggest that Tamarix recruitment was triggered by large annual floods that were followed by years with lower peak flows, not by periods of low flow alone. Tamarix appears to have actively invaded floodplains, while Populus colonization has been limited. Thus, Tamarix invasion may have greatly influenced floodplain development and riparian vegetation composition along the lower Green River since the early 20th century.     

Quantifying Habitat Requirements of Tree-Living Species in Fragmented Boreal Forests with Bayesian Methods

Abstract:  Quantitative conservation objectives require detailed consideration of the habitat requirements of target species. Tree-living bryophytes, lichens, and fungi are a critical and declining biodiversity component of boreal forests. To understand their requirements, Bayesian methods were used to analyze the relationships between the occurrence of individual species and habitat factors at the tree and the stand scale in a naturally fragmented boreal forest landscape. The importance of unexplained between-stand variation in occurrence of species was estimated, and the ability of derived models to predict species' occurrence was tested. The occurrence of species was affected by quality of individual trees. Furthermore, the relationships between occurrence of species at the tree level and size and shape of stands indicated edge effects, implying that some species were restricted to interior habitats of large, regular stands. Yet for the habitat factors studied, requirements of many species appeared similar. Species occurrence also varied between stands; most of the seemingly suitable trees in some stands were unoccupied. The models captured most variation in species occurrence at tree level. They also successfully accounted for between-stand variation in species occurrence, thus providing realistic simulations of stand-level occupancy of species. Important unexplained between-stand variation in species occurrence warns against a simplified view that only local habitat factors influence species' occurrence. Apparently, similar stands will host populations of different sizes due to historical, spatial, and stochastic factors. Thus, habitat suitability cannot be assessed simply by population sizes, and stands lacking a species may still provide suitable habitat and merit protection.     

Deer herbivory alters forest response to canopy decline caused by an exotic insect pest.

Hemlock woolly adelgid (HWA; Adelges tsugae) infestations have resulted in the continuing decline of eastern hemlock (Tsuga canadensis) throughout much of the eastern United States. While the initial impacts of HWA infestations have been documented, our understanding of forest response to this disturbance remains incomplete. HWA infestation is not occurring in isolation but within a complex ecological context. The role of potentially important interacting factors, such as elevated levels of white-tailed deer herbivory, is poorly understood. Despite the potential for herbivory to alter forest successional trajectories following a canopy disturbance, little is known about herbivory-disturbance interactions, and herbivory is rarely considered in assessing forest response to a co-occurring disturbance. We used repeated censuses of deer exclosures and paired controls (400 paired plots) to quantify the impact of deer herbivory on tree seedling species abundance in 10 eastern hemlock ravines that span a gradient in HWA-induced canopy decline severity. Use of a maximum likelihood estimation framework and information theoretics allowed us to quantify the strength of evidence for alternative models developed to estimate the impacts of herbivory on tree seedling abundance as a function of varying herbivore density and canopy decline severity. The exclusion of deer herbivory had marked impacts on the abundance of the studied seedling species: Acer rubrum, Acer saccharum, Betula lenta, Nyssa sylvatica, Quercus montana, and Tsuga canadensis. For all six species, the relationship between seedling abundance and deer density was either exponential or saturating. Although the functional form of the response varied among seedling species, the inclusion of both deer density and canopy decline severity measures consistently resulted in models with substantially greater support. Canopy decline resulted in higher proportional herbivory impacts and altered the ranking of herbivory impacts by seedling species. Our results suggest that, by changing species' competitive abilities, white-tailed deer herbivory alters the trajectory of forest response to this exotic insect pest and has the potential to shift future overstory composition.     

Regeneration of Native Trees in the Presence of Invasive Saltcedar in the Colorado River Delta, Mexico

Abstract:  Many riparian zones in the Sonoran Desert have been altered by elimination of the normal flood regime; such changes to the flow regime have contributed to the spread of saltcedar ( Tamarix ramosissma Ledeb.), an exotic, salt-tolerant shrub. It has been proposed that reestablishment of a natural flow regime on these rivers might permit passive restoration of native trees, without the need for aggressive saltcedar clearing programs. We tested this proposition in the Colorado River delta in Mexico, which has received a series of large-volume water releases from U.S. dams over the past 20 years. We mapped the vegetation of the delta riparian corridor through ground and aerial surveys (1999–2002) and satellite imagery (1992–2002) and related vegetation changes to river flood flows and fire events. Although saltcedar is still the dominant plant in the delta, native cottonwood (  Populus fremontii S. Wats.) and willow ( Salix gooddingii C. Ball) trees have regenerated multiple times because of frequent flood releases from U.S. dams since 1981. Tree populations are young and dynamic (ages 5–10 years). The primary cause of tree mortality between floods is fire. Biomass in the floodplain, as measured by the normalized difference vegetation index on satellite images, responds positively even to low-volume (but long-duration) flood events. Our results support the hypothesis that restoration of a pulse flood regime will regenerate native riparian vegetation despite the presence of a dominant invasive species, but fire management will be necessary to allow mature tree stands to develop.     

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.     

Projected vegetation changes for the American Southwest: combined dynamic modeling and bioclimatic-envelope approach

This study focuses on potential impacts of 21st century climate change on vegetation in the Southwest United States, based on debiased and interpolated climate projections from 17 global climate models used in the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Among these models a warming trend is universal, but projected changes in precipitation vary in sign and magnitude. Two independent methods are applied: a dynamic global vegetation model to assess changes in plant functional types and bioclimatic envelope modeling to assess changes in individual tree and shrub species and biodiversity. The former approach investigates broad responses of plant functional types to climate change, while considering competition, disturbances, and carbon fertilization, while the latter approach focuses on the response of individual plant species, and net biodiversity, to climate change. The dynamic model simulates a region-wide reduction in vegetation cover during the 21st century, with a partial replacement of evergreen trees with grasses in the mountains of Colorado and Utah, except at the highest elevations, where tree cover increases. Across southern Arizona, central New Mexico, and eastern Colorado, grass cover declines, in some cases abruptly. Due to the prevalent warming trend among all 17 climate models, vegetation cover declines in the 21st century, with the greatest vegetation losses associated with models that project a drying trend. The inclusion of the carbon fertilization effect largely ameliorates the projected vegetation loss. Based on bioclimatic envelope modeling for the 21st century, the number of tree and shrub species that are expected to experience robust declines in range likely outweighs the number of species that are expected to expand in range. Dramatic shifts in plant species richness are projected, with declines in the high-elevation evergreen forests, increases in the eastern New Mexico prairies, and a northward shift of the Sonoran Desert biodiversity maximum.     

Riparian forest composition affects stream litter decomposition despite similar microbial and invertebrate communities

Cross-boundary flows of energy and nutrients link biodiversity and functioning in adjacent ecosystems. The composition of forest tree species can affect the structure and functioning of stream ecosystems due to physical and chemical attributes, as well as changes in terrestrial resource subsidies. We examined how variation in riparian canopy composition (coniferous, deciduous, mixed) affects adjacent trophic levels (invertebrate and microbial consumers) and decomposition of organic matter in small, coastal rainforest streams in southwestern British Columbia. Breakdown rates of higher-quality red alder (Alnus rubra) litter were faster in streams with a greater percentage of deciduous than coniferous riparian canopy, whereas breakdown rates of lower-quality western hemlock (Tsuga heterophylla) litter were independent of riparian forest composition. When invertebrates were excluded using fine mesh, breakdown rates of both litter species were an order of magnitude less and were not significantly affected by riparian forest composition. Stream invertebrate and microbial communities were similar among riparian forest composition, with most variation attributed to leaf litter species. Invertebrate taxa richness and shredder biomass were higher in A. rubra litter; however, taxa evenness was greatest for T. heterophylla litter and both litter species in coniferous streams. Microbial community diversity (determined from terminal restriction fragment length polymorphisms) was unaffected by riparian forest or litter species. Fungal allele richness was higher than bacterial allele richness, and microbial communities associated with lower-quality T. heterophylla litter had higher diversity (allele uniqueness and richness) than those associated with higher-quality A. rubra litter. Percent variation in breakdown rates was mostly attributed to riparian forest composition in the presence of invertebrates and microbes; however, stream consumer biodiversity at adjacent trophic levels did not explain these patterns. Riparian and stream ecosystems and their biotic communities are linked through exchange and decomposition of detrital resources, and we provide evidence that riparian forest composition affects stream ecosystem catabolism despite similarities in microbial and invertebrate communities.     

Invasibility and Effects of Amur Honeysuckle in Southwestern Ohio Forests

The Asian exotic Amur honeysuckle ( Lonicera maackii [Rupr.] Herder) has become the dominant shrub in many forests in southwestern Ohio and in some other locations in the eastern United States. Our research focused on the invasibility of forest communities and relationships of L. maackii to the abundance of tree seedlings and herbs. We surveyed 93 forest stands near Oxford, Ohio (USA) to determine L. maackii cover, time since invasion, tree canopy cover, tree basal area, and a shade tolerance index. Stepwise multiple regression indicated that greater than one-half of the variation (r ² = 0.550) in Lonicera cover was correlated to five variables (in descending order of importance): tree canopy cover, distance from Oxford, shade tolerance index, tree basal area, and time since invasion. The results suggest that high light levels and proximity to an abundant seed source increase forest invasibility. Tree seedling density, species richness of seedlings, and herb cover were all inversely related to L. maackii cover. When Lonicera becomes abundant, future structure and composition of forests could be affected and local populations of herbs threatened.     

Germination and Establishment of the Native Plains Cottonwood (Populus deltoides Marshall subsp. monilifera) and the Exotic Russian-olive (Elaeagnus angustifolia L.)

Russian-olive ( Elaeagnus angustifolia ) is a small Eurasian tree that has escaped from cultivation and become naturalized, primarily along watercourses throughout the western United States. We examined germination and establishment of Russian-olive and plains cottonwood ( Populus deltoides ), the principal native riparian tree of the Great Plains, under a range of experimental moisture and light conditions. The fewest seedlings established under the driest conditions; seedling biomass was predictably lower in the shade; root-to-shoot ratios were higher for cottonwood, higher in the sun, and higher under drier conditions. Several interactions were also significant. The timing of germination and mortality varied between plains cottonwood and Russian-olive: cottonwood germinated in mid-June in all treatments in a single pulse with subsequent mortality; the timing and amount of Russian-olive germination differed substantially across treatments with little net mortality. Differences in life-history traits of these species, including seed size, viability, and dispersal, help explain treatment differences. Russian-olive will likely remain an important component of riparian communities along both unregulated and regulated western rivers because it succeeds under conditions optimal for cottonwood establishment and under many conditions unfavorable for cottonwood. Furthermore, many western states still encourage planting of Russian-olive, and control techniques tend to be labor-intensive and expensive.     

Effects of Timber Harvesting on Southern Appalachian Salamanders

We compared the species richness and abundance of salamanders on six recent clearcuts (< 10 years old) with that of salamanders on 34 mature forest stands (>50 years old) in southern Appalachian forests in western North Carolina, U.S.A. Catches of salamanders from plots in mature forest stands were about five times higher than those on recent clearcuts. Almost all species and major taxonomic groups of salamanders were adversely affected by timber removal. Mean number of species collected per plot was about twice as great in mature forest stands as in clearcuts. Analyses of stand age versus salamander catch for 47 plots indicate that 50–70 years are required for populations to return to predisturbance levels following cutting. We conservatively estimate that clearcutting in U.S. national forests in western North Carolina results in a loss of nearly 14 millton salamanders annually. It also is chronically reducing regional populations by more than a quarter of a billion salamanders (9%) below that which could be sustained if mature forests were not cut.     

Tree-Dependent Lichens and Beetles as Indicators in Conservation Forests

We tested whether the conspicuous lichen Lobaria pulmonaria indicates the number of tree-dependent, red-listed species in a hemiboreal forest in southern Sweden. In 18 naturally regenerated, mainly old deciduous forest plots considered to be of high or very high conservation value, the number of red-listed tree lichens or wood beetles was not positively correlated with the area of the forest stands studied (8–56 ha). The 8 stands with L. pulmonaria had about nine (median) red-listed lichens, but 10 stands without L. pulmonaria had only about one such species, a highly significant difference. Similarly, the variation between stands in the number of red-listed wood beetles was considerable, but it was very weakly correlated with the number of red-listed lichens. The number of red-listed wood beetles dependent on dead trees was not different in areas with or without Lobaria pulmonaria , but the number of red-listed wood beetles dependent on hollow trees was higher in stands with Lobaria pulmonaria (median of seven species) than in those without (three species) the lichen. Stands with this lichen species also contained significantly more of other types of lichens that have been proposed as indicators of forest continuity according to three lists. We stress the need for identification of species that could serve as indicators of different types of forest continuity and identify some organism groups that may indicate the different types.     

Defoliation levels of oriental spruce by Ips typographus (L.) in relation to elevation and exposure

A study concerning the effects of elevation and exposure of the spruce forests on defoliation levels of oriental spruce (Picea orientalis (L.) Link.) by Ips typographus L. was carried out during 2005 and 2006 in Artvin-Hatila National Park, Turkey Nine spruce stands were selected at 3 zones of elevations (1000-1350 m, 1350-1700 m and 1700-2000 m) and at different aspects to assess the role of elevation and exposure in the crown defoliation level and body length of beetles. Influence of bark thickness and trunk diameter at 1.3 m on the damage caused by the pest was investigated as well. The results of the study were as follows: (1) The mean defoliation level was highest at 1700-2000 m following by 1350-1700 m and 1000-1350 m. (2) The highest defoliation levels occurred on southern slopes following by eastern and northern slopes at 1700-2000 m. (3) No statistical differences were found in the mean bark thickness between tree defoliation levels 1, 2, 3 and 4. (4) Mean trunk diameters of dead trees (level 4) were significantly greater than those with defoliation levels 0, 1 and 2. (5) Mean body length of I. typographus at upper zones was significantly higher than those at middle and lower zones.     

Spectral analysis of forest model time series

The use of spectral analysis to elucidate the cyclic behavior in time series generated by a forest stand growth simulation model is discussed. A stand-level simulator, FORET, for an Appalachian deciduous forest is described. An estimate of the power spectral density of the total biomass time series is calculated. The power spectral density estimate indicates a dominant cyclic behavior with a period of about 200 years. In addition the spectral density is approximately bandlimited. This characteristic makes possible the application of the sampling theorem for analysis of sampling rates.