Donald M. Waller

Inbreeding effects in wild populations

Whether inbreeding affects the demography and persistence of natural populations has been questioned. However, new pedigree data from field populations and molecular and analytical tools for tracing patterns of relationship and inbreeding have now enhanced our ability to detect inbreeding depression within and among wild populations. This work reveals that levels of inbreeding depression vary across taxa, populations and environments, but are usually substantial enough to affect both individual and population performance. Data from bird and mammal populations suggest that inbreeding depression often significantly affects birth weight, survival, reproduction and resistance to disease, predation and environmental stress. Plant studies, based mostly on comparing populations that differ in size or levels of genetic variation, also reveal significant inbreeding effects on seed set, germination, survival and resistance to stress. Data from butterflies, birds and plants demonstrate that populations with reduced genetic diversity often experience reduced growth and increased extinction rates. Crosses between such populations often result in heterosis. Such a genetic rescue effect might reflect the masking of fixed deleterious mutations. Thus, it might be necessary to retain gene flow among increasingly fragmented habitat patches to sustain populations that are sensitive to inbreeding.

Direct and indirect effects of white-tailed deer in forest ecosystems☆

Ungulates can profoundly alter the structure and composition of forest communities via both direct and indirect mechanisms. Individual plant species often respond in a unique way to the direct effect of herbivory as a function of their sensitivity to browse damage, ungulate food preferences, and the density of ungulates present. Sustained browsing pressure can limit the regeneration of favored and susceptible woody plants and eliminate populations of favored or susceptible herbaceous plants. These losses, in turn, give rise to indirect effects via trophic cascades or physical habitat modification. These indirect effects affect many other plant and animal populations. In the mixed conifer‐hardwood forests around the Great Lakes in North America, widespread habitat modification and the extirpation of native predators and other ungulates have acted to boost populations of white-tailed deer (Odocoileus virginianus) to historically high densities. Such densities have curtailed regeneration of several important conifers (e.g. Tsuga canadensis and Thuja occidentalis) as evidenced by demographic analysis. Deer also appear to limit regeneration of Quercus and Betula in many areas. Impacts on understory herbs are harder to assess, but baseline data from 50 years ago indicate that these communities are changing in a pattern that implicates deer: grasses, sedges, and some ferns are increasing while overall herb diversity is declining. Thus, deer are playing a keystone role in these communities. We are currently assessing an additional set of questions, including: How best can we measure and represent ungulate impacts? At which densities do deer threaten forest diversity? How do impacts depend on initial plant and ungulate densities? Which species emerge as ‘winners’ or ‘losers’ in heavily-browsed landscapes? What characteristics or traits make a species susceptible to ungulate herbivory? How do ungulates affect patterns of diversity and relative abundance in ecological communities? What are the pathways by which ungulates exert indirect effects on species? and How significant are indirect effects? # 2003 Elsevier Science B.V. All rights reserved.

Microclimate moderates plant responses to macroclimate warming

Significance Around the globe, climate warming is increasing the dominance of warm-adapted species—a process described as “thermophilization.” However, thermophilization often lags behind warming of the climate itself, with some recent studies showing no response at all. Using a unique database of more than 1,400 resurveyed vegetation plots in forests across Europe and North America, we document significant thermophilization of understory vegetation. However, the response to macroclimate warming was attenuated in forests whose canopies have become denser. This microclimatic effect likely reflects cooler forest-floor temperatures via increased shading during the growing season in denser forests. Because standing stocks of trees have increased in many temperate forests in recent decades, microclimate may commonly buffer understory plant responses to macroclimate warming. Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., “thermophilization” of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that “climatic lags” may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12–67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass—e.g., for bioenergy—may open forest canopies and accelerate thermophilization of temperate forest biodiversity.

Plant strategies in relation to elevation and light in floodplain herbs

We examined the distribution of herbaceous species in southern Wisconsin floodplain forests in relation to two major environmental factors: elevation and light. Most species had definite elevation optima and could be categorized as either light specialists for high or low light, or light generalists. To test Grimes ideas regarding the existence of three primary plant strategies (competitive, stress tolerant, and ruderal), we used simple autecological indices to classify species into guilds. Higher elevation areas were dominated by tall competitive perennial forbs. As flooding frequency increased, smaller perennial forbs (stress-tolerant competitors) and tall annuals (competitive ruderals) increased in importance. In areas of greatest flooding frequency, the dichotomy in plant strategies was most extreme: slowgrowing, flood-tolerant sedges and grasses (stress tolerators) shared the ground with small, fast-maturing, annual forbs (ruderals). Tolerance to flooding in sedges and grasses may be related to the position of their shoot apex, which affords protection from mechanical damage caused by turbulent flooding. Light was important in differentiating guilds (high light specialists, light generalists, small spring-flowering forbs) only when flooding was infrequent. Grimes triangle of plant strategies maps onto axes of disturbance frequency and physiological tolerance, emphasizing the interaction of environment and adaptation in determining strategy. One environmental event, such as flood, fire, or drought, may be seen as a disturbance by short-lived ruderal species and as a stress by tolerant, long-lived perennials. Combinations of more than one important stress or disturbance further complicate Grimes model.

Differences in fitness between seedlings derived from cleistogamous and chasmogamous flowers in Impatiens capensis

It often occurred to me that it would be advisable to try whether seedlings from cross-fertilized flowers were in any way superior to those from selffertilized flowers. . .. I ought to have reflected that such elaborate provisions favouring cross-fertilization, as we see in innumerable plants, would not have been acquired for the sake of gaining a distant and slight advantage, or of avoiding a distant and slight evil.

Monitoring adaptive genetic responses to environmental change

Widespread environmental changes including climate change, selective harvesting and landscape alterations now greatly affect selection regimes for most organisms. How animals and plants can adapt to these altered environments via contemporary evolution is thus of strong interest. We discuss how to use genetic monitoring to study adaptive responses via repeated analysis of the same populations over time, distinguishing between phenotypic and molecular genetics approaches. After describing monitoring designs, we develop explicit criteria for demonstrating adaptive responses, which include testing for selection and establishing clear links between genetic and environmental change. We then review a few exemplary studies that explore adaptive responses to climate change in Drosophila, selective responses to hunting and fishing, and contemporary evolution in Daphnia using resurrected resting eggs. We further review a broader set of 44 studies to assess how well they meet the proposed criteria, and conclude that only 23% fulfill all criteria. Approximately half (43%) of these studies failed to rule out the alternative hypothesis of replacement by a different, better‐adapted population. Likewise, 34% of the studies based on phenotypic variation did not test for selection as opposed to drift. These shortcomings can be addressed via improved experimental designs and statistical testing. We foresee monitoring of adaptive responses as a future valuable tool in conservation biology, for identifying populations unable to evolve at sufficiently high rates and for identifying possible donor populations for genetic rescue. Technological advances will further augment the realization of this potential, especially next‐generation sequencing technologies that allow for monitoring at the level of whole genomes.

Relative performance of selfed and outcrossed progeny in impatiens-capensis

This study compares survival and growth of progeny derived from chasmogamous (CH) and cleistogamous (CL) flowers in Impatiens capensis, a forest annual. When progeny were grown in the field, CH seeds had significantly higher survival rates over winter (64% versus 56%), and the survival advantage of outcrossed progeny was not attributable to seed weight differences. No differences in seedling growth were detected. Greenhouse comparisons revealed no difference in seed survival but a 30% growth advantage to CH seedlings. We found no changes in developmental homeostasis of three leaf shape characters between inbred and outbred progeny, nor was there any difference in variability within CH and CL families. The outcrossing advantage observed in these experiments could not have been caused by avoidance of sib competition. Theory predicts that self‐pollinated progenies may be more variable than outcrossed progenies if rare, recessive alleles are important contributors to genetic variances. Electrophoretic markers indicate that progeny derived from CH flowers are predominantly outcrossed (at least 54‐97%).

SHIFTS IN SOUTHERN WISCONSIN FOREST CANOPY AND UNDERSTORY RICHNESS, COMPOSITION, AND HETEROGENEITY

We resurveyed the under- and overstory species composition of 94 upland forest stands in southern Wisconsin in 2002-2004 to assess shifts in canopy and understory richness, composition, and heterogeneity relative to the original surveys in 1949-1950. The canopy has shifted from mostly oaks (Quercus spp.) toward more mesic and shade-tolerant trees (primarily Acer spp.). Oak-dominated early-successional stands and those on coarse, nutrient-poor soils changed the most in canopy composition. Understories at most sites (80%) lost native species, with mean species density declining 25% at the 1-m2 scale and 23.1% at the 20-m2 scale. Woody species have increased 15% relative to herbaceous species in the understory despite declining in absolute abundance. Initial canopy composition, particularly the abundance of red oaks (Quercus rubra and Q. velutina), predicted understory changes better than the changes observed in the overstory. Overall rates of native species loss were greater in later-successional stands, a pattern driven by differential immigration rather than differential extirpation. However, understory species initially found in early-successional habitats declined the most, particularly remnant savanna taxa with narrow or thick leaves. These losses have yet to be offset by compensating increases in native shade-adapted species. Exotic species have proliferated in prevalence (from 13 to 76 stands) and relative abundance (from 1.2% to 8.4%), but these increases appear unrelated to the declines in native species richness and heterogeneity observed. Although canopy succession has clearly influenced shifts in understory composition and diversity, the magnitude of native species declines and failure to recruit more shade-adapted species suggest that other factors now act to limit the richness, heterogeneity, and composition of these communities.

Improving and integrating data on invasive species collected by citizen scientists

Limited resources make it difficult to effectively document, monitor, and control invasive species across large areas, resulting in large gaps in our knowledge of current and future invasion patterns. We surveyed 128 citizen science program coordinators and interviewed 15 of them to evaluate their potential role in filling these gaps. Many programs collect data on invasive species and are willing to contribute these data to public databases. Although resources for education and monitoring are readily available, groups generally lack tools to manage and analyze data. Potential users of these data also retain concerns over data quality. We discuss how to address these concerns about citizen scientist data and programs while preserving the advantages they afford. A unified yet flexible national citizen science program aimed at tracking invasive species location, abundance, and control efforts could be designed using centralized data sharing and management tools. Such a system could meet the needs of multiple stakeholders while allowing efficiencies of scale, greater standardization of methods, and improved data quality testing and sharing. Finally, we present a prototype for such a system (see www.citsci.org).

Neglect of Genetic Diversity in Implementation of the Convention on Biological Diversity

Genetic diversity is the foundation for all biological diversity; the persistence and evolutionary potential of species depend on it. World leaders have agreed on the conservation of genetic diversity as an explicit goal of the Convention on Biological Diversity (CBD). Nevertheless, actions to protect genetic diversity are largely lacking. With only months left to the 2010-biodiversity target, when the 191 parties to the CBD have agreed on achieving a significant reduction of the rate of biodiversity loss, gene-level diversity is still not being monitored, and indicators and thresholds that can be used to devise strategies to conserve this important component of biodiversity are missing. Immediate action is needed to ensure that genetic diversity is not neglected in conservation targets beyond 2010.