Biodiversity as a multidimensional construct: a review, framework and case study of herbivory's impact on plant biodiversity.

Biodiversity is inherently multidimensional, encompassing taxonomic, functional, phylogenetic, genetic, landscape and many other elements of variability of life on the Earth. However, this fundamental principle of multidimensionality is rarely applied in research aimed at understanding biodiversity's value to ecosystem functions and the services they provide. This oversight means that our current understanding of the ecological and environmental consequences of biodiversity loss is limited primarily to what unidimensional studies have revealed. To address this issue, we review the literature, develop a conceptual framework for multidimensional biodiversity research based on this review and provide a case study to explore the framework. Our case study specifically examines how herbivory by whitetail deer (Odocoileus virginianus) alters the multidimensional influence of biodiversity on understory plant cover at Black Rock Forest, New York. Using three biodiversity dimensions (taxonomic, functional and phylogenetic diversity) to explore our framework, we found that herbivory alters biodiversity's multidimensional influence on plant cover; an effect not observable through a unidimensional approach. Although our review, framework and case study illustrate the advantages of multidimensional over unidimensional approaches, they also illustrate the statistical and empirical challenges such work entails. Meeting these challenges, however, where data and resources permit, will be important if we are to better understand and manage the consequences we face as biodiversity continues to decline in the foreseeable future.

Rapid rebound of soil respiration following partial stand disturbance by tree girdling in a temperate deciduous forest.

Forests serve an essential role in climate change mitigation by removing CO2 from the atmosphere. Within a forest, disturbance events can greatly impact C cycling and subsequently influence the exchange of CO2 between forests and the atmosphere. This connection makes understanding the forest C cycle response to disturbance imperative for climate change research. The goal of this study was to examine the temporal response of soil respiration after differing levels of stand disturbance for 3 years at the Black Rock Forest (southeastern NY, USA; oaks comprise 67% of the stand). Tree girdling was used to mimic pathogen attack and create the following treatments: control, girdling all non-oaks (NOG), girdling half of the oak trees (O50), and girdling all the oaks (OG). Soil respiratory rates on OG plots declined for 2 years following girdling before attaining a full rebound of belowground activity in the third year. Soil respiration on NOG and O50 were statistically similar to the control for the duration of the study although a trend for a stronger decline in respiration on O50 relative to NOG occurred in the first 2 years. Respiratory responses among the various treatments were not proportional to the degree of disturbance and varied over time. The short-lived respiratory response on O50 and OG suggests that belowground activity is resilient to disturbance; however, sources of the recovered respiratory flux on these plots are likely different than they were pre-treatment. The differential taxon response between oaks and non-oaks suggests that after a defoliation or girdling event, the temporal response of the soil respiratory flux may be related to the C allocation pattern of the affected plant group.

Stochastic multiplicative population growth predicts and interprets Taylor's power law of fluctuation scaling.

Taylor's law (TL) asserts that the variance of the density (individuals per area or volume) of a set of comparable populations is a power-law function of the mean density of those populations. Despite the empirical confirmation of TL in hundreds of species, there is little consensus about why TL is so widely observed and how its estimated parameters should be interpreted. Here, we report that the Lewontin-Cohen (henceforth LC) model of stochastic population dynamics, which has been widely discussed and applied, leads to a spatial TL in the limit of large time and provides an explicit, exact interpretation of its parameters. The exponent of TL exceeds 2 if and only if the LC model is supercritical (growing on average), equals 2 if and only if the LC model is deterministic, and is less than 2 if and only if the LC model is subcritical (declining on average). TL and the LC model describe the spatial variability and the temporal dynamics of populations of trees on long-term plots censused over 75 years at the Black Rock Forest, Cornwall, NY, USA.

Allometric scaling of population variance with mean body size is predicted from Taylor's law and density-mass allometry.

Two widely tested empirical patterns in ecology are combined here to predict how the variation of population density relates to the average body size of organisms. Taylor's law (TL) asserts that the variance of the population density of a set of populations is a power-law function of the mean population density. Density-mass allometry (DMA) asserts that the mean population density of a set of populations is a power-law function of the mean individual body mass. Combined, DMA and TL predict that the variance of the population density is a power-law function of mean individual body mass. We call this relationship "variance-mass allometry" (VMA). We confirmed the theoretically predicted power-law form and the theoretically predicted parameters of VMA, using detailed data on individual oak trees (Quercus spp.) of Black Rock Forest, Cornwall, New York. These results connect the variability of population density to the mean body mass of individuals.

Oak loss increases foliar nitrogen, δ(15)N and growth rates of Betula lenta in a northern temperate deciduous forest.

Oak forests dominate much of the eastern USA, but their future is uncertain due to a number of threats and widespread failure of oak regeneration. A sudden loss of oaks (Quercus spp.) could be accompanied by major changes in forest nitrogen (N) cycles with important implications for plant nutrient uptake and tree species composition. In this study, we measured the changes in N use and growth rates of black birch trees (Betula lenta L.) following oak girdling at the Black Rock Forest in southeastern New York, USA. Data were collected from nine experimental plots composed of three treatments: 100% oaks girdled (OG), 50% oaks girdled (O50) and control (C). Foliar N concentration and foliar (15)N abundance increased significantly in the oak-girdled plots relative to the control, indicating that the loss of oaks significantly altered N cycling dynamics. As mineralization and nitrification rates increase following oak loss, black birch trees increase N absorption as indicated by higher foliar N content and increased growth rates. Foliar N concentration increased by 15.5% in the O50 and 30.6% in the OG plots relative to the control, while O50 and OG plots were enriched in (15)N by 1.08‰ and 3.33‰, respectively (P < 0.0001). A 641% increase in black birch growth rates in OG plots suggests that this species is able to respond to additional N availability and/or increased light availability. The loss of oaks and subsequent increase in black birch productivity may have a lasting impact on ecosystem form and function.

Urban environment of New York City promotes growth in northern red oak seedlings.

Urbanization is accelerating across the globe, elevating the importance of studying urban ecology. Urban environments exhibit several factors affecting plant growth and function, including high temperatures (particularly at night), CO(2) concentrations and atmospheric nitrogen deposition. We investigated the effects of urban environments on growth in Quercus rubra L. seedlings. We grew seedlings from acorns for one season at four sites along an urban-rural transect from Central Park in New York City to the Catskill Mountains in upstate New York (difference in average maximum temperatures of 2.4 °C; difference in minimum temperatures of 4.6 °C). In addition, we grew Q. rubra seedlings in growth cabinets (GCs) mimicking the seasonal differential between the city and rural sites (based on a 5-year average). In the field experiment, we found an eightfold increase in biomass in urban-grown seedlings relative to those grown at rural sites. This difference was primarily related to changes in growth allocation. Urban-grown seedlings and seedlings grown at urban temperatures in the GCs exhibited a lower root: shoot ratio (urban ~0.8, rural/remote ~1.5), reducing below-ground carbon costs associated with construction and maintenance. These urban seedlings instead allocated more growth to leaves than did rural-grown seedlings, resulting in 10-fold greater photosynthetic area but no difference in photosynthetic capacity of foliage per unit area. Seedlings grown at urban temperatures in both the field and GC experiments had higher leaf nitrogen concentrations per unit area than those grown at cooler temperatures (increases of 23% in field, 32% in GC). Lastly, we measured threefold greater (13)C enrichment of respired CO(2) (relative to substrate) in urban-grown leaves than at other sites, which may suggest greater allocation of respiratory function to growth over maintenance. It also shows that lack of differences in total R flux in response to environmental conditions may mask dramatic shifts in respiratory functioning. Overall, our findings indicating greater seedling growth and establishment at a critical regeneration phase of forest development may have important implications for the ecology of urban forests as well as the predicted growth of the terrestrial biosphere in temperate regions in response to climate change.

The autotrophic contribution to soil respiration in a northern temperate deciduous forest and its response to stand disturbance.

The goal of this study was to evaluate the contribution of oak trees (Quercus spp.) and their associated mycorrhizal fungi to total community soil respiration in a deciduous forest (Black Rock Forest) and to explore the partitioning of autotrophic and heterotrophic respiration. Trees on twelve 75 × 75-m plots were girdled according to four treatments: girdling all the oaks on the plot (OG), girdling half of the oak trees on a plot (O50), girdling all non-oaks on a plot (NO), and a control (C). In addition, one circular plot (diameter 50 m) was created where all trees were girdled (ALL). Soil respiration was measured before and after tree girdling. A conservative estimate of the total autotrophic contribution is approximately 50%, as indicated by results on the ALL and OG plots. Rapid declines in carbon dioxide (CO(2)) flux from both the ALL and OG plots, 37 and 33%, respectively, were observed within 2 weeks following the treatment, demonstrating a fast turnover of recently fixed carbon. Responses from the NO and O50 treatments were statistically similar to the control. A non-proportional decline in respiration rates along the gradient of change in live aboveground biomass complicated partitioning of the overall rate of soil respiration and indicates that belowground carbon flux is not linearly related to aboveground disturbance. Our findings suggest that in this system there is a threshold disturbance level between 35 and 74% of live aboveground biomass loss, beyond which belowground dynamics change dramatically.

Geographic relatedness and predictability of Escherichia coli along a peninsular beach complex of Lake Michigan.

To determine more accurately the real-time concentration of fecal indicator bacteria (FIB) in beach water, predictive modeling has been applied in several locations around the Great Lakes to individual or small groups of similar beaches. Using 24 beaches in Door County, Wisconsin, we attempted to expand predictive models to multiple beaches of complex geography. We examined the importance of geographic location and independent variables and the consequential limitations for potential beach or beach group models. An analysis of Escherichia coli populations over 4 yr revealed a geographic gradient to the beaches, with mean E. coli concentrations decreasing with increasing distance from the city of Sturgeon Bay. Beaches grouped strongly by water type (lake, bay, Sturgeon Bay) and proximity to one another, followed by presence of a storm or creek outfall or amount of shoreline enclosure. Predictive models developed for beach groups commonly included wave height and cumulative 48-h rainfall but generally explained little E. coli variation (adj. R2=0.19-0.36). Generally low concentrations of E. coli at the beaches influenced the effectiveness of model results presumably because of low signal-to-noise ratios and the rarity of elevated concentrations. Our results highlight the importance of the sensitivity of regressors and the need for careful methods evaluation. Despite the attractiveness of predictive models as an alternative beach monitoring approach, it is likely that FIB fluctuations at some beaches defy simple prediction approaches. Regional, multi-beach, and individual beach predictive models should be explored alongside other techniques for improving monitoring reliability at Great Lakes beaches.

Dichloro(norbornadiene)platinum(II): a comparison with dichloro(cyclooctadiene)platinum(II).

The crystal structure of the title compound, (bicyclo[2.2.1]hepta-2,5-diene)dichloroplatinum(II), [PtCl2(C7H8)], has been determined from single-crystal X-ray analysis. The coordination sphere about the Pt atom is pseudo-square planar, with shorter Pt-C distances than in the corresponding dichloro(cyclooctadiene)platinum(II) complex.

Energy investment in leaves of red maple and co-occurring oaks within a forested watershed.

Despite its recent expansion in eastern US forests, red maple (Acer rubrum L.) generally exhibits a low leaf photosynthetic rate, leaf mass per unit area (LMA) and leaf nitrogen concentration ([N]) relative to co-occurring oaks (Quercus spp.). To evaluate these differences from the perspective of leaf energy investment, we compared leaf construction cost (CC) and leaf maintenance cost (MC) with leaf photosynthetic rate at saturating photon flux density and ambient CO2 partial pressure (Amax) in red maple and co-occurring red oak (Quercus rubra L.) and chestnut oak (Quercus prinus L.). We also examined relationships among leaf physiological, biochemical and structural characteristics of upper-canopy leaves of these three species at lower (wetter) and upper (drier) elevation sites of a watershed in the Black Rock Forest, Cornwall, NY, USA. Although A(max), leaf [N], leaf carbon concentration ([C]) and LMA were significantly less in red maple than in either oak species at both sites, CC per unit leaf area of red maple was 28.2 and 35.4% less than that of red oak at the lower and upper site, respectively, and 38.8 and 32% less than that of chestnut oak at the lower and upper site, respectively. Leaf MC per unit leaf area, which was positively associated with leaf CC (r2 = 0.95), was also significantly lower in red maple than in either oak species at both sites. When expressed per unit leaf area, A(max) was positively correlated with both CC (r2 = 0.65) and MC (r2 = 0.59). The cost/benefit ratio of CC/Amax of red maple was significantly less than that of chestnut oak at the lower site, however, CC/A(max) did not exhibit any significant interspecific differences at the upper site. Expressed per unit leaf area, CC was correlated positively with LMA (r2 = 0.90), leaf [N] (r2 = 0.97), and leaf [C] (r2 = 0.89), and negatively correlated with leaf molar carbon to nitrogen ratio (r2 = 0.92). Combined with red maple's general success in many oak-dominated forests, our findings suggest that reduced leaf-level photosynthetic capacity and related leaf characteristics in red maple are partially balanced by lower energy and resource requirements for leaf biomass construction and maintenance, which could enhance the competitive success of this species.

Analysis of the growth of rimu (Dacrydium cupressinum) in South Westland, New Zealand, using process-based simulation models.

Two process-based models were used to identify the environmental variables limiting productivity in a pristine, mature forest dominated by rimu (Dacrydium cupressinum Sol. ex Lamb.) trees in South Westland, New Zealand. A model of canopy net carbon uptake, incorporating routines for radiation interception, photosynthesis and water balance was used to determine a value for quantum efficiency when climate variables were not limiting. The annual net carbon uptake by the canopy was estimated to be 1.1 kg C m(-2) and the quantum efficiency 22.6 mmol mol quanta(-1). This value of quantum efficiency, combined with other parameters obtainable from the literature, was then used in a model of forest productivity (3-PG), to simulate changes in net productivity and the allocation of carbon to tree components. The model was adjusted to match a measured stem increment of 10.6 Mg ha(-1) over a period of 13 years. To achieve this while maintaining a low, but stable value for leaf area index, it was necessary to set the site fertility rating very low and select high values for the parameters describing the proportional allocation of total carbon to roots. This approach highlighted nutrient availability as the principal constraint on productivity for the ecosystem and identified critical measurements that will be necessary for using the model to predict the effects of climate change on carbon sequestration. The low rates of carbon uptake and productivity are consistent with the low nutrient supply available from the highly leached, acid soils, most likely attributable to frequent saturation and a very shallow aerobic zone.

Photosynthetic characteristics in canopies of Quercus rubra, Quercus prinus and Acer rubrum differ in response to soil water availability.

Photosynthesis and related leaf characteristics were measured in canopies of co-occurring Quercus rubra L. (red oak), Quercus prinus L. (chestnut oak) and Acer rubrum L. (red maple) trees. Mature (20+ m tall) trees were investigated at sites of differing soil water availability within a catchment (a drier upper site and a wetter lower site). Leaf photosynthetic characteristics differed significantly between species and in response to site and position in the canopy. Photosynthetic capacity (A max) was significantly greater at the wetter site in all canopy strata in A. rubrum but not in Q. rubra or Q. prinus. Our findings for A. rubrum are generally consistent with those predicting that species with higher specific leaf area (SLA) will have higher A max per unit leaf nitrogen (N) and that species with leaves with lower SLA (e.g. Q. rubra and Q. prinus) will have shallower slopes of the A max-N relationship. Importantly, the relationships between A max and N area (and by implication photosynthetic nitrogen-use efficiency, PNUE) differed in A. rubrum between the sites, with PNUE significantly lower at the drier site. The lower photosynthetic capacity and PNUE must substantially reduce carbon acquisition capacity in A. rubrum under these field conditions. Maximum stomatal conductance (g smax) differed significantly between species, with g smax greatest in Q. rubra and Q. prinus. In Q. rubra and Q. prinus, g smax was significantly lower at the upper site than the lower site. There was no significant response of g smax to site in A. rubrum. These stomatal responses were consistent with the C i/C a ratio, which was significantly lower in leaves of Q. rubra and Q. prinus at the upper site, but did not differ between sites in A. rubrum. Leaf δ13C was significantly lower in A. rubrum than in either Q. rubra or Q. prinus at both sites. These findings indicate differences in stomatal behaviour in A. rubrum which are likely to contribute to lower water use efficiency at both sites. Our results support the hypothesis that the two Quercus species, in contrast to A. rubrum, maintain photosynthetic capacity at the drier site whilst minimising transpirational water loss. They also suggest, based primarily on physiological evidence, that the ability of A. rubrum to compete with other species of these deciduous forests may be limited, particularly in sites of low moisture availability and during low rainfall years.

Comparisons of carbon isotope discrimination in populations of aridland plant species differing in lifespan.

Carbon isotope discrimination (Δ) was compared between populations of dominant perennial plant species, differing in life expectancy, in two deserts with contrasting vegetation types. In both deserts, plants of the shorter-lived species showed significantly higher Δ and greater intrapopulation variance in this character compared to the long-lived species. These results indicate underlying differences in gas-exchange physiology, and suggest a positive correlation between water-use efficiency and lifespan in desert plants. Differences in variance for this character may reflect greater microenvironmental variation experienced by shorter-lived plants and/or different forms of selection acting on water-use traits. Spatial distributions were significantly clustered for the shorter-lived species and significantly uniform for the long-lived species, indicating that competition has been important in the development of the long-lived populations. The long-lived Larrea tridentata showed a significant, negative correlation between Δ and Thiessen polygon area, suggesting a positive relationship between water-use efficiency and longevity within this species. This relationship was weakly supported in the other warm desert species, Encelia farinosa, but was not observed within populations of the cold desert species, Gutierrezia microcephala and Coleogyne ramosissima. These results suggest that Δ reflects key aspects of plant metabolism related to lifespan; these differences may ultimately influence interactions among desert plants and the structure of desert plant communities.

Differential utilization of summer rains by desert plants.

Seasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.