Showing papers in "Oecologia in 2000"

Linking biodiversity to ecosystem function: implications for conservation ecology

Abstract: We evaluate the empirical and theoretical support for the hypothesis that a large proportion of native species richness is required to maximize ecosystem stability and sustain function. This assessment is important for conservation strategies because sustenance of ecosystem functions has been used as an argument for the conservation of species. If ecosystem functions are sustained at relatively low species richness, then arguing for the conservation of ecosystem function, no matter how important in its own right, does not strongly argue for the conservation of species. Additionally, for this to be a strong conservation argument the link between species diversity and ecosystem functions of value to the human community must be clear. We review the empirical literature to quantify the support for two hypotheses: (1) species richness is positively correlated with ecosystem function, and (2) ecosystem functions do not saturate at low species richness relative to the observed or experimental diversity. Few empirical studies demonstrate improved function at high levels of species richness. Second, we analyze recent theoretical models in order to estimate the level of species richness required to maintain ecosystem function. Again we find that, within a single trophic level, most mathematical models predict saturation of ecosystem function at a low proportion of local species richness. We also analyze a theoretical model linking species number to ecosystem stability. This model predicts that species richness beyond the first few species does not typically increase ecosystem stability. One reason that high species richness may not contribute significantly to function or stability is that most communities are characterized by strong dominance such that a few species provide the vast majority of the community biomass. Rapid turnover of species may rescue the concept that diversity leads to maximum function and stability. The role of turnover in ecosystem function and stability has not been investigated. Despite the recent rush to embrace the linkage between biodiversity and ecosystem function, we find little support for the hypothesis that there is a strong dependence of ecosystem function on the full complement of diversity within sites. Given this observation, the conservation community should take a cautious view of endorsing this linkage as a model to promote conservation goals.

Linking stable oxygen and carbon isotopes with stomatal conductance and photosynthetic capacity: a conceptual model.

Abstract: Based on measurements of δ18O and δ13C in organic matter of C3-plants, we have developed a conceptual model that gives insight into the relationship between stomatal conductance (g l) and photosynthetic capacity (A max) resulting from differing environmental constraints and plant-internal factors. This is a semi-quantitative approach to describing the long-term effects of environmental factors on CO2 and H2O gas exchange, whereby we estimate the intercellular CO2 concentration (c i) from δ13C and the air humidity from δ18O. Assuming that air humidity is an important factor influencing g l, the model allows us to distinguish whether differences in c i are caused by a response of g l or of A max. As an application of the model we evaluated the isotope data from three species in plots differing in intensity of land use (hay meadows and abandoned areas) at three sites along a south north transect in the Eastern Alps. We found three different δ18O-δ13C response patterns in native and planted grassland species (cultivated in the greenhouse). After preliminary confirmation by gas-exchange measurements we conclude that the proposed model is a promising tool for deriving carbon water relations in different functional groups from δ18O and δ13C isotope data.

The use of stable isotopes to study ecosystem gas exchange.

Abstract: Stable isotopes are a powerful research tool in environmental sciences and their use in ecosystem re- search is increasing. In this review we introduce and dis- cuss the relevant details underlying the use of carbon and oxygen isotopic compositions in ecosystem gas ex- change research. The current use and potential develop- ments of stable isotope measurements together with con- centration and flux measurements of CO2 and water va- por are emphasized. For these applications it is critical to know the isotopic identity of specific ecosystem compo- nents such as the isotopic composition of CO 2 , organic matter, liquid water, and water vapor, as well as the associated isotopic fractionations, in the soil-plant- atmosphere system. Combining stable isotopes and con- centration measurements is very effective through the use of "Keeling plots." This approach allows the identifi- cation of the isotopic composition and the contribution of ecosystem, or ecosystem components, to the exchange fluxes with the atmosphere. It also allows the estimation of net ecosystem discrimination and soil disequilibrium effects. Recent modifications of the Keeling plot ap- proach permit examination of CO2 recycling in ecosys- tems. Combining stable isotopes with dynamic flux mea- surements requires precision in isotopic sampling and analysis, which is currently at the limit of detection. Combined with the micrometeorological gradient ap- proach (applicable mostly in grasslands and crop fields), stable isotope measurements allow separation of net CO 2 exchange into photosynthetic and soil respiration compo- nents, and the evapotranspiration flux into soil evapora- tion and leaf transpiration. Similar applications in con- junction with eddy correlation techniques (applicable to forests, in addition to grasslands and crop fields) are more demanding, but can potentially be applied in com- bination with the Keeling plot relationship. The advance and potential in using stable isotope measurements should make their use a standard component in the limit- ed arsenal of ecosystem-scale research tools.

Communication between plants : induced resistance in wild tobacco plants following clipping of neighboring sagebrush

Abstract: The possibility of communication between plants was proposed nearly 20 years ago, although previous demonstrations have suffered from methodological problems and have not been widely accepted. Here we report the first rigorous, experimental evidence demonstrating that undamaged plants respond to cues released by neighbors to induce higher levels of resistance against herbivores in nature. Sagebrush plants that were clipped in the field released a pulse of an epimer of methyl jasmonate that has been shown to be a volatile signal capable of inducing resistance in wild tobacco. Wild tobacco plants with clipped sagebrush neighbors had increased levels of the putative defensive oxidative enzyme, polyphenol oxidase, relative to control tobacco plants with unclipped sagebrush neighbors. Tobacco plants near clipped sagebrush experienced greatly reduced levels of leaf damage by grasshoppers and cutworms during three field seasons compared to unclipped controls. This result was not caused by an altered light regime experienced by tobacco near clipped neighbors. Barriers to soil contact between tobacco and sagebrush did not reduce the difference in leaf damage although barriers that blocked air contact negated the effect.

Decomposition and carbon cycling of dead trees in tropical forests of the central Amazon.

Abstract: Decomposition rate constants were measured for boles of 155 large dead trees (>10 cm diameter) in central Amazon forests. Mortality data from 21 ha of permanent inventory plots, monitored for 10–15 years, were used to select dead trees for sampling. Measured rate constants varied by over 1.5 orders of magnitude (0.015–0.67 year–1), averaging 0.19 year–1 with predicted error of 0.026 year. Wood density and bole diameter were significantly and inversely correlated with rate constants. A tree of average biomass was predicted to decompose at 0.17 year–1. Based on mortality data, an average of 7.0 trees ha–1 year–1 died producing 3.6 Mg ha–1 year–1 of coarse litter (>10 cm diameter). Mean coarse litter standing-stocks were predicted to be 21 Mg ha–1, with a mean residence time of 5.9 years, and a maximum mean carbon flux to the atmosphere of 1.8 Mg C ha–1 year–1. Total litter is estimated to be partitioned into 16% fine wood, 30% coarse wood, and 54% non-woody litter (e.g., leaves, fruits, flowers). Decomposition rate constants for coarse litter were compiled from 20 globally distributed studies. Rates were highly correlated with mean annual temperature, giving a respiration quotient (Q 10) of 2.4 (10°C–1).

Competition and intraguild predation among three sympatric carnivores

Abstract: We examined the relative roles of dominance in agonistic interactions and energetic constraints related to body size in determining local abundances of coyotes (Canis latrans, 8–20 kg), gray foxes (Urocyon cinereoargenteus, 3–5 kg) and bobcats (Felis rufus, 5–15 kg) at three study sites (hereafter referred to as NP, CP, and SP) in the Santa Monica Mountains of California. We hypothesized that the largest and behaviorally dominant species, the coyote, would exploit a wider range of resources (i.e., a higher number of habitat and/or food types) and, consequently, would occur in higher density than the other two carnivores. We evaluated our hypotheses by quantifying their diets, food overlap, habitat-specific abundances, as well as their overall relative abundance at the three study sites. We identified behavioral dominance of coyotes over foxes and bobcats in Santa Monica because 7 of 12 recorded gray fox deaths and 2 of 5 recorded bobcat deaths were due to coyote predation, and no coyotes died as a result of their interactions with bobcats or foxes. Coyotes and bobcats were present in a variety of habitats types (8 out of 9), including both open and brushy habitats, whereas gray foxes were chiefly restricted to brushy habitats. There was a negative relationship between the abundances of coyotes and gray foxes (P=0.020) across habitats, suggesting that foxes avoided habitats of high coyote predation risk. Coyote abundance was low in NP, high in CP, and intermediate in SP. Bobcat abundance changed little across study sites, and gray foxes were very abundant in NP, absent in CP, and scarce in SP; this suggests a negative relationship between coyote and fox abundances across study sites, as well. Bobcats were solely carnivorous, relying on small mammals (lagomorphs and rodents) throughout the year and at all three sites. Coyotes and gray foxes also relied on small mammals year-round at all sites, though they also ate significant amounts of fruit. Though there were strong overall interspecific differences in food habits of carnivores (P<0.0001), average seasonal food overlaps were high due to the importance of small mammals in all carnivore diets [bobcat-gray fox: 0.79±0.09 (SD), n=4; bobcat-coyote: 0.69±0.16, n=6; coyote-gray fox: 0.52±0.05, n=4]. As hypothesized, coyotes used more food types and more habitat types than did bobcats and gray foxes and, overall, coyotes were the most abundant of the three species and ranged more widely than did gray foxes. We propose that coyotes limit the number and distribution of gray foxes in Santa Monica Mountains, and that those two carnivores exemplified a case in which the relationship between their body size and local abundance is governed by competitive dominance of the largest species rather than by energetic equivalences. However, in the case of the intermediate-sized bobcat no such a pattern emerged, likely due to rarity or inconsistency of agonistic interactions and/or behavioral avoidance of encounters by subordinate species.

Long distance seed dispersal by wind: measuring and modelling the tail of the curve.

Abstract: The size and shape of the tail of the seed dispersal curve is important in determining the spatial dynamics of plants, but is difficult to quantify. We devised an experimental protocol to measure long-distance dispersal which involved measuring dispersal by wind from isolated individuals at a range of distances from the source, but maintaining a large and constant sampling intensity at each distance. Seeds were trapped up to 80 m from the plants, the furthest a dispersal curve for an individual plant has been measured for a non-tree species. Standard empirical negative exponential and inverse power models were fitted using likelihood methods. The latter always had a better fit than the former, but in most cases neither described the data well, and strongly under-estimated the tail of the dispersal curve. An alternative model formulation with two kernel components had a much better fit in most cases and described the tail data more accurately. Mechanistic models provide an alternative to direct measurement of dispersal. However, while a previous mechanistic model accurately predicted the modal dispersal distance, it always under-predicted the measured tail. Long-distance dispersal may be caused by rare extremes in horizontal wind speed or turbulence. Therefore, under-estimation of the tail by standard empirical models and mechanistic models may indicate a lack of flexibility to take account of such extremes. Future studies should examine carefully whether the widely used exponential and power models are, in fact, valid, and investigate alternative models.

Host plant species effects on arbuscular mycorrhizal fungal communities in tallgrass prairie

Abstract: Symbiotic associations between plants and arbuscular mycorrhizal (AM) fungi are ubiquitous in many herbaceous plant communities and can have large effects on these communities and ecosystem processes. The extent of species-specificity between these plant and fungal symbionts in nature is poorly known, yet reciprocal effects of the composition of plant and soil microbe communities is an important assumption of recent theoretical models of plant community structure. In grassland ecosystems, host plant species may have an important role in determining development and sporulation of AM fungi and patterns of fungal species composition and diversity. In this study, the effects of five different host plant species [Poa pratensis L., Sporobolus heterolepis (A. Gray) A. Gray, Panicum virgatum L., Baptisia bracteata Muhl. ex Ell., Solidago missouriensis Nutt.] on spore communities of AM fungi in tallgrass prairie were examined. Spore abundances and species composition of fungal communities of soil samples collected from patches within tallgrass prairie were significantly influenced by the host plant species that dominated the patch. The AM fungal spore community associated with B. bracteata showed the highest species diversity and the fungi associated with Pa. virgatum showed the lowest diversity. Results from sorghum trap cultures using soil collected from under different host plant species showed differential sporulations of AM fungal species. In addition, a greenhouse study was conducted in which different host plant species were grown in similar tallgrass prairie soil. After 4 months of growth, AM fungal species composition was significantly different beneath each host species. These results strongly suggest that AM fungi show some degree of host-specificity and are not randomly distributed in tallgrass prairie. The demonstration that host plant species composition influences AM fungal species composition provides support for current feedback models predicting strong regulatory effects of soil communities on plant community structure. Differential responses of AM fungi to host plant species may also play an important role in the regulation of species composition and diversity in AM fungal communities.

The soil food web of two beech forests (Fagus sylvatica) of contrasting humus type: stable isotope analysis of a macro- and a mesofauna-dominated community.

Abstract: The structure of the soil food web in two beech (Fagus sylvatica) forests, the Gottinger Wald and the Solling forest (Northern Germany), was investigated using variations in tissue 15N concentrations of animal species or taxa. The Gottinger Wald is located on a limestone plateau and characterized by mull humus with high macrofauna activity, particularly of Lumbricidae, Diplopoda and Isopoda. In contrast, the Solling forest is located on a sandstone mountain range and characterized by moder humus. The soil fauna of this forest is dominated by mesofauna, particularly by Collembola, Enchytraeidae and Oribatida. In June 1995 soil fauna was sampled using heat extraction. Three soil layers were analysed at each of the sites. 15N/14N ratios of bulk material increased strongly with soil depth in both forests. This also applied to the water-soluble fraction at the Gottinger Wald, but not at the Solling. Generally, the water-soluble fraction was more enriched in 15N than the bulk materials. For most animals studied 15N/14N ratios varied little with soil depth. In both forests soil animals could be classified either as saprophages, including microphytophages, or predators. On average, the δ15N of predatory taxa (Chilopoda, Araneida, Gamasina, Staphylinidae) exceeded that of saprophagous or microphytophagous taxa (Lumbricidae, Isopoda, Diplopoda, Collembola, Oribatida, Enchytraeidae) by 4.4 and 3.9‰ for the Gottinger Wald and the Solling, respectively. We assume that most of the saprophagous or microphytophagous taxa studied consist of primary and secondary decomposers and hypothesize that predators prey more on secondary than primary decomposers. Generally, average δ15N values differed little between saprophagous (Lumbricidae, Diplopoda, Isopoda) and microphytophagous taxa (Collembola, Oribatida). The variations in δ15N values of species within these taxa consistently exceeded the variation between them, indicating that the species of each of these taxa form a continuum from primary to secondary decomposers. Also, variations in δ15N values within predatory taxa in most cases exceeded that between taxa excluding top predators like Sorex. We conclude that using higher taxonomic units in soil food web analysis is problematic and in general not consistent with nature. Higher taxonomic units may only be useful for depicting very general trophic groupings such as predators or microbi-detritivores.

Altitude trends in conifer leaf morphology and stable carbon isotope composition

Abstract: The natural ratio of stable carbon isotopes (δ 13 C) was compared to leaf structural and chemical characteristics in evergreen conifers in the north-central Rockies, United States. We sought a general model that would explain variation in δ 13 C across altitudinal gradi- ents. Because variation in δ13C is attributed to the shifts between supply and demand for carbon dioxide within the leaf, we measured structural and chemical variables related to supply and demand. We measured stomatal density, which is related to CO 2 supply to the chlorop- lasts, and leaf nitrogen content, which is related to CO 2 demand. Leaf mass per area was measured as an inter- mediate between supply and demand. Models were test- ed on four evergreen conifers: Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii, and Pinus contor- ta, which were sampled across 1800 m of altitude. We found significant variation among species in the rate of δ13C increase with altitude, ranging from 0.91‰ km -1 for A. lasiocarpa to 2.68‰ km -1 for Pinus contorta. Leaf structure and chemistry also varied with altitude: stomatal density decreased, leaf mass per area increased, but leaf nitrogen content (per unit area) was constant. The regressions on altitude were particularly robust in Pinus contorta. Variables were derived to describe the balance between supply and demand; these variables were stomata per gram of nitrogen and stomata per gram of leaf mass. Both derived variables should be positively related to internal CO 2 supply and thus negatively relat- ed to δ 13 C. As expected, both derived variables were negatively correlated with δ13C. In fact, the regression on stomatal density per gram was the best fit in the study (r 2 =0.72, P<0.0001); however, the relationships were species specific. The only general relationship observed was between δ 13 C and LMA: δ 13 C (‰)=-32.972+ 0.0173〈LMA (r2=0.45, P<0.0001). We conclude that species specificity of the isotopic shift indicates that ev- ergreen conifers demonstrate varying degrees of func- tional plasticity across environmental gradients, while the observed convergence of δ 13 C with LMA suggests that internal resistance may be the key to understanding inter-specific isotopic variation across altitude.

Influence of soil porosity on water use in Pinus taeda

Abstract: We analyzed the hydraulic constraints im- posed on water uptake from soils of different porosities in loblolly pine (Pinus taeda L.) by comparing genetical- ly related and even-aged plantations growing in loam versus sand soil. Water use was evaluated relative to the maximum transpiration rate (E crit ) allowed by the soil- leaf continuum. We expected that trees on both soils would approach Ecrit during drought. Trees in sand, how- ever, should face greater drought limitation because of steeply declining hydraulic conductivity in sand at high soil water potential ( Ψ S ). Transport considerations sug- gest that trees in sand should have higher root to leaf ar- ea ratios (AR:AL), less negative leaf xylem pressure ( Ψ L), and be more vulnerable to xylem cavitation than trees in loam. The A R :A L was greater in sand versus loam (9.8 vs 1.7, respectively). This adjustment maintained about 86% of the water extraction potential for both soils. Trees in sand were more deeply rooted (>1.9 m) than in loam (95% of roots <0.2 m), allowing them to shift water uptake to deeper layers during drought and avoid hydraulic failure. Midday Ψ L was constant for days of high evaporative demand, but was less negative in sand (-1.6 MPa) versus loam (-2.1 MPa). Xylem was more vulnerable to cavitation in sand versus loam trees. Roots in both soils were more vulnerable than stems, and expe- rienced the greatest predicted loss of conductivity during drought. Trees on both soils approached Ecrit during drought, but at much higher Ψ S in sand (<-0.4 MPa) than in loam (<-1.0 MPa). Results suggest considerable phenotypic plasticity in water use traits for P. taeda which are adaptive to differences in soil porosity.

Changes in drought response strategies with ontogeny in Quercus rubra: implications for scaling from seedlings to mature trees.

Abstract: We investigated scaling of physiological parameters between age classes of Quercus rubra by combining in situ field measurements with an experimental approach. In the in situ field study, we investigated changes in drought response with age in seedlings, juveniles, and mature trees of Q. rubra. Throughout the particularly dry summer of 1995 and the unusually wet summer of 1996 in New England, we measured water potential of leaves (ΨLeaf) and gas exchange of plants at three sites at the Harvard Forest in Petersham, Massachusetts. In order to determine what fraction of the measured differences in gas exchange between seedlings and mature trees was due to environment versus ontogeny, an experiment was conducted in which seedlings were grown under light and soil moisture regimes simulating the environment of mature trees. The photosynthetic capacity of mature trees was three-fold greater than that of seedlings during the wet year, and six-fold greater during the drought year. The seedling experiment demonstrated that the difference in photosynthetic capacity between seedlings and mature trees is comprised equally of an environmental component (50%) and an ontogenetic component (50%) in the absence of water limitation. Photosynthesis was depressed more severely in seedlings than in mature trees in the drought year relative to the wet year, while juveniles showed an intermediate response. Throughout the drought, the predawn leaf water potential (ΨPD) of seedlings became increasingly negative (-0.4 to -1.6 MPa), while that of mature trees became only slightly more negative (-0.2 to -0.5 MPa). Again, juveniles showed an intermediate response (-0.25 to -0.8 MPa). During the wet summer of 1996, however, there was no difference in ΨPD between seedlings, juveniles and mature trees. During the dry summer of 1995, seedlings were more responsive to a major rain event than mature trees in terms of ΨLeaf , suggesting that the two age classes depend on different water sources. In all age classes, instantaneous measurements of intrinsic water use efficiency (WUEi), defined as C assimilation rate divided by stomatal conductance, increased as the drought progressed, and all age classes had higher WUEi during the drought year than in the wet year. Mature trees, however, showed a greater ability to increase their WUEi in response to drought. Integrated measurements of WUE from C isotope discrimination (Δ) of leaves indicated higher WUE in mature trees than juveniles and seedlings. Differences between years, however, could not be distinguished, probably due to the strong bias in C isotope fractionation at the time of leaf production, which occurred prior to the onset of drought conditions in 1995. From this study, we arrive at two main conclusions.

Contributions of stable-isotope data to elucidating food webs of Mediterranean rocky littoral fishes.

Abstract: The food webs of rocky infra-littoral ecosystems in the Mediterranean have been little studied. In this investigation stable isotopes and dietary data were compared in an attempt to describe features of the food webs concerned. δ13C and δ15N were determined for plants, invertebrates and fishes from the Bay of Calvi, Corsica. Dietary data were derived from the literature. δ13C of plants ranged from –8.59‰ to –33.74‰, of benthic invertebrates from –17.0‰ to –20.52‰, of planktonic invertebrates from –20.08‰ to –22.34‰ and of fishes from –16.27‰ to –19.59‰. δ15N was generally greater at higher trophic levels. δ15N of plants was 0.95–2.92‰, of benthic invertebrates 1.69–6.54‰, of planktonic invertebrates 3.51–6.82‰ and of fishes 4.63–9.77‰. 13C enrichment tended to be associated with benthic food chains and 13C depletion with planktonic chains. Stable-isotope data suggested more varied diets for many species than implied by gut-contents data. Omnivory and trophic plasticity were widespread, and many consumers fed lower down the food chain than previous studies had suggested. Both stable-isotope and gut-contents analysis resolved differences between fishes feeding on planktonic and benthic prey and indicated that the herbivorous fish Sarpa salpa fed on a diet substantially different from that of other fishes. Zooplankton were important in the diets of several consumers (both primary and secondary), as was plankton derived detritus. One species of fish previously identified as planktivorous was shown to feed largely on benthic organisms, whilst several species of benthic invertebrates may feed on plankton-derived detritus. Although herbivores seemed to obtain most of their C from macroalgae, δ15N data suggested that many of these animals supplemented their intake of N, although gut-contents analysis did not provide evidence for such uptake. The isotopic data have elucidated several features of the food web which we would not otherwise have detected.

Effects of disturbance frequency, intensity, and area on assemblages of stream macroinvertebrates

Abstract: Disturbance frequency, intensity, and areal extent may influence the effects of disturbance on biological communities. Furthermore, these three factors may have interacting effects on biological diversity. We manipulated the frequency, intensity, and area of disturbance in a full-factorial design on artificial substrates and measured responses of benthic macroinvertebrates in a northern Vermont stream. Macroinvertebrate abundance was lower in all disturbance treatments than in the undisturbed control. As in most other studies in streams, species density (number of species/sample) was lower in disturbed treatments than in undisturbed controls. However, species density is very sensitive to total abundance of a sample, which is usually reduced by disturbance. We used a rarefaction method to compare species richness based on an equivalent number of individuals. In rarefied samples, species richness was higher in all eight disturbed treatments than in the undisturbed control, with significant increases in species richness for larger areas and greater intensities of disturbance. Increases in species richness in response to disturbance were consistent within patches, among patches with similar disturbance histories, and among patches with differing disturbance histories. These results provide some support for Huston’s dynamic-equilibrium model but do not support the intermediate-disturbance hypothesis. Our analyses demonstrate that species richness and species density can generate opposite patterns of community response to disturbance. The interplay of abundance, species richness, and species density has been neglected in previous tests of disturbance models.

Predictable risk to native plants in weed biological control.

Abstract: Data on field host use of 112 insects, 3 fungi, 1 mite, and 1 nematode established for biological control of weeds in Hawaii, the continental United States, and the Caribbean indicate that the risk to native flora can be judged reliably before introduction. Virtually all risk is borne by native plant species that are closely related to target weeds. Fifteen species of insects introduced for biological control use 41 native plant species; 36 of which are congeneric with target weeds, while 4 others belong to two closely allied genera. Only 1 of 117 established biological organisms uses a native plant unrelated to the target weed. Thus the elements of protection for the native flora are the selection of weed targets that have few or no native congeners and the introduction of biological control organisms with suitably narrow diets.

Numerical responses of different trophic groups of invertebrates to manipulations of plant diversity in grasslands.

Abstract: We studied the effects of plant diversity on abundance of invertebrate herbivores, parasitoids and predators in two grassland communities (one in Switzerland and one in Sweden) in which plant species richness and functional diversity have been experimentally manipulated. Among herbivores, the abundance of only the most sessile and specialised groups (leafhoppers and wingless aphids) was affected by plant diversity. At both sites, numbers of leafhoppers in sweep net samples showed a linear, negative relationship with plant species number whereas numbers of wingless aphids in suction samples increased with the number of plant functional groups (grasses, legumes, and non-legume forbs) present in the plot. Activity of carabid beetles and spiders (as revealed by pitfall catches) and the total number of predators in pitfalls at the Swiss site decreased linearly with increases in the number of plant species and plant functional groups. Abundance of more specialised enemies, hymenopteran parasitoids, was not affected by the manipulations of plant diversity. Path analysis and analysis of covariance indicated that plant diversity effects on invertebrate abundance were mostly indirect and mediated by changes in plant biomass and cover. At both sites, plant species composition (i.e. the identity of plant species in a mixture) affected numbers of most of the examined groups of invertebrates and was, therefore, a more important determinant of invertebrate abundance in grasslands than plant species richness per se or the number of plant functional groups. The presence of legumes in a mixture was especially important and led to higher numbers of most invertebrate groups. The similarity of invertebrate responses to plant diversity at the two study sites indicates that general patterns in abundance of different trophic groups can be detected across plant diversity gradients under different environmental conditions.

Species richness and parasitism in a fragmented landscape: experiments and field studies with insects on Vicia sepium.

Abstract: Effects of habitat fragmentation on species diversity and herbivore-parasitoid interactions were analyzed using the insect community of seed feeders and their parasitoids in the pods of the bush vetch (Vicia sepium L.). Field studies were carried out on 18 old meadows differing in area and isolation. The area of these meadows was found to be the major determinant of species diversity and population abundance of endophagous insects. Effects of isolation were further analyzed experimentally using 16 small plots with potted vetch plants isolated by 100–500 m from vetch populations on large old meadows. The results showed that colonization success greatly decreased with increasing isolation. In both cases, insect species were not equally affected. Parasitoids suffered more from habitat loss and isolation than their phytophagous hosts. Minimum area requirements, calculated from logistic regressions, were higher for parasitoids than for herbivores. In addition, percent parasitism of the herbivores significantly decreased with area loss and increasing isolation of Vicia sepium plots, supporting the trophic-level hypothesis of island biogeography. Species with high rates of absence on meadows and isolated plant plots were not only characterized by their high trophic level, but also by low abundance and high spatial population variability. Thus conservation of large and less isolated habitat remnants enhances species diversity and parasitism of potential pest insects, i.e., the stability of ecosystem functions.

Density compensation in neotropical primate communities: evidence from 56 hunted and nonhunted Amazonian forests of varying productivity

Abstract: Density compensation is a community-level phenomenon in which increases in the abundance of some species may offset the population decline, extirpation, or absence of other potentially interacting competitors. In this paper we examine the evidence for density compensation in neotropical primate assemblages using data from 56 hunted and nonhunted, but otherwise undisturbed, forest sites of Amazonia and the Guianan shields from which population density estimates are available for all diurnal primate species. We found good evidence of density compensation of the residual assemblage of nonhunted mid-sized species where the large-bodied (ateline) species had been severely reduced in numbers or driven to local extinction by subsistence hunters. Only weak evidence for density compensation, however, was detected in small-bodied species. These conclusions are based on the effects of ordinal measures of hunting pressure on the aggregate primate biomass across different size classes after controlling for the effects of forest type and productivity. These results are interpreted primarily in relation to patterns of niche partitioning between different primate functional groups or ecospecies. This study suggests that while overhunting drastically reduces the average body size in multi-species assemblages of forest vertebrates, depletion of large-bodied species is only partially offset (i.e. undercompensated) by smaller taxa.

Relationships between fine root dynamics and nitrogen availability in Michigan northern hardwood forests.

Abstract: Minirhizotrons were used to observe fine root (≤1 mm) production, mortality, and longevity over 2 years in four sugar-maple-dominated northern hardwood forests located along a latitudinal temperature gradient. The sites also differed in N availability, allowing us to assess the relative influences of soil temperature and N availability in controlling fine root lifespans. Root production and mortality occurred throughout the year, with most production occurring in the early portion of the growing season (by mid-July). Mortality was distributed much more evenly throughout the year. For surface fine roots (0–10 cm deep), significant differences in root longevity existed among the sites, with median root lifespans for root cohorts produced in 1994 ranging from 405 to 540 days. Estimates of fine root turnover, based on the average of annual root production and mortality as a proportion of standing crop, ranged from 0.50 to 0.68 year–1 for roots in the upper 30 cm of soil. The patterns across sites in root longevity and turnover did not follow the north to south temperature gradient, but rather corresponded to site differences in N availability, with longer average root lifespans and lower root turnover occurring where N availability was greater. This suggests the possibility that roots are maintained as long as the benefit (nutrients) they provide outweighs the C cost of keeping them alive. Root N concentrations and respiration rates (at a given temperature) were also higher at sites where N availability was greater. It is proposed that greater metabolic activity for roots in nitrogen-rich zones leads to greater carbohydrate allocation to those roots, and that a reduction in root C sink strength when local nutrients are depleted provides a mechanism through which root lifespan is regulated in these forests.

Correlations between foliar δ15N and nitrogen concentrations may indicate plant-mycorrhizal interactions.

Abstract: Nitrogen isotope measurements may provide insights into changing interactions among plants, mycor- rhizal fungi, and soil processes across environmental gradients. Here, we report changes in δ 15 N signatures due to shifts in species composition and nitrogen (N) dy- namics. These changes were assessed by measuring fine root biomass, net N mineralization, and N concentrations and δ 15 N of foliage, fine roots, soil, and mineral N across six sites representing different post-deglaciation ages at Glacier Bay, Alaska. Foliar δ 15 N varied widely, between 0 and -2‰ for nitrogen-fixing species, between 0 and -7‰ for deciduous non-fixing species, and be- tween 0 and -11‰ for coniferous species. Relatively constant δ 15 N values for ammonium and generally low levels of soil nitrate suggested that differences in ammo- nium or nitrate use were not important influences on plant δ 15 N differences among species at individual sites. In fact, the largest variation among plant δ15N values were observed at the youngest and oldest sites, where soil nitrate concentrations were low. Low mineral N con- centrations and low N mineralization at these sites indi- cated low N availability. The most plausible mechanism to explain low δ 15 N values in plant foliage was a large isotopic fractionation during transfer of nitrogen from mycorrhizal fungi to plants. Except for N-fixing plants, the foliar δ 15 N signatures of individual species were gen- erally lower at sites of low N availability, suggesting ei- ther an increased fraction of N obtained from mycorrhi- zal uptake (f), or a reduced proportion of mycorrhizal N transferred to vegetation ( T r ). Foliar and fine root nitro- gen concentrations were also lower at these sites. Foliar N concentrations were significantly correlated with δ 15 N in foliage of Populus, Salix, Picea, and Tsuga hetero- phylla, and also in fine roots. The correlation between δ 15 N and N concentration may reflect strong underlying relationships among N availability, the relative allocation of carbon to mycorrhizal fungi, and shifts in either f or Tr.

How plants shape the ant community in the Amazonian rainforest canopy: the key role of extrafloral nectaries and homopteran honeydew.

Abstract: Ant-plant interactions in the canopy of a lowland Amazonian rainforest of the upper Orinoco, Venezuela, were studied using a modified commercial crane on rails (Surumoni project). Our observations show a strong correlation between plant sap exudates and both abundance of ants and co-occurrence of ant species in tree canopies. Two types of plant sap sources were compared: extrafloral nectaries (EFNs) and honeydew secretions by homopterans. EFNs were a frequent food source for ants on epiphytes (Philodendron spp., Araceae) and lianas (Dioclea, Fabaceae), but rare on canopy trees in the study area, whereas the majority of trees were host to aggregations of homopterans tended by honeydew-seeking ants (on 62% of the trees examined). These aggregations rarely occurred on epiphytes. Baited ant traps were installed on plants with EFNs and in the crowns of trees from three common genera, including trees with and without ant-tended homopterans: Goupia glabra (Celastraceae), Vochysia spp. (Vochysiaceae), and Xylopia spp. (Annonaceae). The number of ant workers per trap was significantly higher on plants offering one of the two plant sap sources than on trees without such resources. Extrafloral nectaries were used by a much broader spectrum of ant species and genera than honeydew, and co-occurrence of ant species (in traps) was significantly higher on plants bearing EFNs than on trees. Homopteran honeydew (Coccidae and Membracidae), on the other hand, was mostly monopolised by a single ant colony per tree. Homopteran-tending ants were generally among the most dominant ants in the canopy. The most prominent genera were Azteca, Dolichoderus (both Dolichoderinae), Cephalotes, Pheidole, Crematogaster (all Myrmicinae), and Ectatomma (Ponerinae). Potential preferences were recorded between ant and homopteran species, and also between ant-homopteran associations and tree genera. We hypothesize that the high availability of homopteran honeydew provides a key resource for ant mosaics, where dominant ant colonies and species maintain mutually exclusive territories on trees. In turn, we propose that for nourishment of numerous ants of lower competitive capacity, Philodendron and other sources of EFNs might be particularly important.

Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types.

Abstract: The structural causes of variation in leaf mass per area, and of variations in leaf structure accounted for by leaf habit and life form, were explored in a set of laboratory-grown seedlings of 52 European woody species. The leaf traits analysed included density, thickness, saturated mass/dry mass, and leaf nitrogen per mass and per area. Other traits described the anatomy of leaves, most of them relating to the lamina (proportions of palisade and spongy parenchymata, epidermis, air space and sclerified tissues, expressed as volume per leaf area, and per-cell transversal areas of epidermis and parenchymata), and another referring to the mid rib (transversal section of sclerified tissues). Across the whole set of species leaf mass per area was correlated with leaf density but not with thickness, and this was confirmed by taxonomic relatedness tests. Denser leaves corresponded with greater proportion of sclerified tissues in the lamina, smaller cells and lower water and N contents, but no relation was found with the proportion of air space in the lamina. Taxonomic relatedness analysis statistically supported the negative association of leaf density with saturated to dry leaf mass ratio. Thicker leaves also exhibited greater volume per leaf area and greater individual cell area in each of the tissues, particularly parenchyma. Mean leaf mass per area and leaf thickness were lower in deciduous than in evergreen species, but no significant differences in leaf density, proportion of sclerified tissues in the lamina or cell area were found between the two groups. Leaf mass per area was higher in trees and subshrubs than in shrubs and climbers-plus-scramblers, this rank being equal for leaf density and proportion of sclerified tissues in the lamina, and reversed for cell area. Given the standardised environment and ontogenetic phase of the seedlings, we conclude that variation in leaf structure and anatomy among species and species groups has a strong genetic basis, and is already expressed early in the development of woody plants. From an ecological viewpoint, we can interpret greater leaf mass per area across this species set as greater allocation to support and defence functions, as shown predominantly by species from resource-poor environments.

Caribou movement as a correlated random walk

Abstract: Movement is a primary mechanism coupling animals to their environment, yet there exists little empirical analysis to test our theoretical knowledge of this basic process. We used correlated random walk (CRW) models and satellite telemetry to investigate long-distance movements of caribou, the most vagile, non-volant terrestrial vertebrate in the world. Individual paths of migratory and sedentary female caribou were quantified using measures of mean move length and angle, and net squared displacements at each successive move were compared to predictions from the models. Movements were modelled at two temporal scales. For paths recorded through one annual cycle, the CRW model overpredicted net displacement of caribou through time. For paths recorded over shorter intervals delineated by seasonal behavioural changes of caribou, there was excellent correspondence between model predictions and observations for most periods for both migratory and sedentary caribou. On the smallest temporal scale, a CRW model significantly overpredicted displacements of migratory caribou during 3 months following calving; this was also the case for sedentary caribou in late summer, and in late winter. In all cases of overprediction there was significant positive autocorrelation in turn direction, indicating that movements were more tortuous than expected. In one case of underprediction, significant negative autocorrelation of sequential turn direction was evident, indicating that migratory caribou moved in straightened paths during spring migration to calving grounds. Results are discussed in light of known migration patterns and possible limiting factors for caribou, and indicate the applicability of CRW models to animal movement at vast spatial and temporal scales, thus assisting in future development of more sophisticated models of population spread and redistribution for vertebrates.

Plant species diversity as a driver of early succession in abandoned fields: a multi-site approach.

Abstract: Succession is one of the most studied processes in ecology and succession theory provides strong predictability. However, few attempts have been made to influence the course of succession thereby testing the hypothesis that passing through one stage is essential before entering the next one. At each stage of succession ecosystem processes may be affected by the diversity of species present, but there is little empirical evidence showing that plant species diversity may affect succession. On ex-arable land, a major constraint of vegetation succession is the dominance of perennial early-successional (arable weed) species. Our aim was to change the initial vegetation succession by the direct sowing of later-successional plant species. The hypothesis was tested that a diverse plant species mixture would be more successful in weed suppression than species-poor mixtures. In order to provide a robust test including a wide range of environmental conditions and plant species, experiments were carried out at five sites across Europe. At each site, an identical experiment was set up, albeit that the plant species composition of the sown mixtures differed from site to site. Results of the 2-year study showed that diverse plant species mixtures were more effective at reducing the number of natural colonisers (mainly weeds from the seed bank) than the average low-diversity treatment. However, the effect of the low-diversity treatment depended on the composition of the species mixture. Thus, the effect of enhanced species diversity strongly depended on the species composition of the low-diversity treatments used for comparison. The effects of high-diversity plant species mixtures on weed suppression differed between sites. Low-productivity sites gave the weakest response to the diversity treatments. These differences among sites did not change the general pattern. The present results have implications for understanding biological invasions. It has been hypothesised that alien species are more likely to invade species-poor communities than communities with high diversity. However, our results show that the identity of the local species matters. This may explain, at least partly, controversial results of studies on the relation between local diversity and the probability of being invaded by aliens.

Field deposition of Bt transgenic corn pollen: lethal effects on the monarch butterfly.

Abstract: We present the first evidence that transgenic Bacillus thuringiensis (Bt) corn pollen naturally deposited on Asclepias syriaca; common milkweed, in a corn field causes significant mortality of Danaus plexippus L (Lepidoptera: Danaidae) larvae Larvae feeding for 48 h on A syriaca plants naturally dusted with pollen from Bt corn plants suffered significantly higher rates of mortality at 48 h (20±3%) compared to larvae feeding on leaves with no pollen (3±3%), or feeding on leaves with non-Bt pollen (0%) Mortality at 120 h of D plexippus larvae exposed to 135 pollen grains/cm2 of transgenic pollen for 48 h ranged from 37 to 70% We found no sub-lethal effects on D plexippus adults reared from larvae that survived a 48-h exposure to three concentrations of Bt pollen Based on our quantification of the wind dispersal of this pollen beyond the edges of agricultural fields, we predict that the effects of transgenic pollen on D plexippus may be observed at least 10 m from transgenic field borders However, the highest larval mortality will likely occur on A syriaca plants in corn fields or within 3 m of the edge of a transgenic corn field We conclude that the ecological effects of transgenic insecticidal crops need to be evaluated more fully before they are planted over extensive areas

Fire effects on ecosystem nitrogen cycling in a Californian bishop pine forest

Abstract: Fire can cause severe nitrogen (N) losses from grassland, chaparral, and temperate and boreal forest ecosystems. Paradoxically, soil ammonium levels are markedly increased by fire, resulting in high rates of primary production in re-establishing plant communities. In a manipulative experiment, we examined the influence of wild-fire ash residues on soil, microbial and plant N pools in a recently burned Californian bishop pine (Pinus muricata D. Don) forest. Ash stimulated post-fire primary production and ecosystem N retention through direct N inputs from ash to soils, as well as indirect ash effects on soil N availability to plants. These results suggest that redistribution of surface ash after fire by wind or water may cause substantial heterogeneity in soil N availability to plants, and could be an important mechanism contributing to vegetation patchiness in fire-prone ecosystems. In addition, we investigated the impact of fire on ecosystem N cycling by comparing 15N natural abundance values from recently burned and nearby unburned P. muricata forest communities. At the burned site, 15N natural abundance in recolonising species was similar to that in bulk soil organic matter. By contrast, there was a marked 15N depletion in the same species relative to the total soil N pool at the unburned site. These results suggest that plant uptake of nitrate (which tends to be strongly depleted in 15N because of fractionation during nitrification) is low in recently burned forest communities but could be an important component of eco- system N cycling in mature conifer stands.

The effects of long-term nitrogen loading on grassland insect communities

Abstract: Just as long-term nitrogen loading of grasslands decreases plant species richness and increases plant biomass, we have found that nitrogen loading decreases insect species richness and increases insect abundances. We sampled 54 plots that had been maintained at various rates of nitrogen addition for 14 years. Total insect species richness and effective insect diversity, as well as herbivore and predator species richness, were significantly, negatively related to the rate of nitrogen addition. However, there was variation in trophic responses to nitrogen. Detritivore species richness increased as nitrogen addition increased, and parasitoids showed no response. Insect abundances, measured as the number of insects and insect biovolume (an estimate of biomass), were significantly, positively related to the rate of nitrogen addition, as were the abundances of herbivores and detritivores. Parasitoid abundance was negatively related to the rate of nitrogen addition. Changes in the insect community were correlated with changes in the plant community. As rates of nitrogen addition increased, plant species richness decreased, plant productivity and plant tissue nitrogen increased, and plant composition shifted from C4 to C3 grass species. Along this gradient, total insect species richness and effective insect diversity were most strongly, positively correlated with plant species richness. Insect biovolume was negatively correlated with plant species richness. Responses of individual herbivores varied along the nitrogen gradient, but numbers of 13 of the 18 most abundant herbivores were positively correlated with their host plant biomass. Although insect communities did not respond as strongly as plant communities, insect species richness, abundance, and composition were impacted by nitrogen addition. This study demonstrates that long-term nitrogen loading affects the entire food chain, simplifying both plant and insect communities.

Ecophysiological comparison of direct and indirect defenses in Nicotiana attenuata

Abstract: After herbivore attack, plants launch a suite of direct and indirect defense responses that must be coordinated if plants are to realize a fitness benefit from these responses. Here we characterize the volatile emissions in the native tobacco plant, Nicotiana attenuata Torr. ex Wats., that are elicited by tobacco hornworm (Manduca sexta L.) attack and are known to function as attractants for parasitoids. To provide the first ecophysiological comparison of examples of both types of defense in the same species, we characterize the elicitation and signaling mechanisms, the resources required, and the potential costs and benefits of the volatile release and compare these traits with those of the well-described induced direct defense in this species, nicotine production. The release of (E)-β-ocimene, cis-α-bergamotene and linalool is dramatically induced within 24 h by application of methyl jasmonate (MeJA), caterpillar feeding, and the treatment of mechanical wounds with larval oral secretions (OS), but not by mechanical damage alone. Plants from different geographic locations produce volatile blends that differ in composition. The most consistently released component from all genotypes, cis-α-berga-motene, is positively related to the amount of MeJA and the level of wounding if OS are applied to the wounds. The volatile release is strongly light dependent, dropping to undetectable quantities during dark periods, even when temperatures are elevated to match those of the light period. Inhibitors of wound-induced jasmonate accumulation (salicylates and auxins), which are known to inhibit wound-induced nicotine production, do not inhibit the release of volatiles. By individually inducing different leaf positions with OS and, on other plants, excising them after induction, we demonstrate that the emission is largely a systemic, whole-plant response, which is maximally triggered when the second fully expanded leaf is induced. We conclude that while both are whole-plant, systemic responses that utilize recently acquired resources for their production and are activated by the jasmonate cascade, the elicitation of the volatile release exhibits greater tissue sensitivity and utilizes additional signaling components than does nicotine production. In contrast to the large investment of fitness-limiting resources required for induced nicotine production or the resources used in benzyl acetone release from flowers for pollinator attraction, the resource requirements for the volatile release are minor. Hence the argument that the volatile release incurs comparatively large physiological costs cannot be supported in this system.

Spatial patterns in abundance of a damselfish reflect availability of suitable habitat.

Abstract: For species with metapopulation structures, variation in abundance among patches can arise from variation in the input rate of colonists. For reef fishes, variability in larval supply frequently is invoked as a major determinant of spatial patterns. We examined the extent to which spatial variation in the amount of suitable habitat predicted variation in the abundance of the damselfish Dascyllus aruanus, an abundant planktivore that occupies live, branched coral throughout the Indo-Pacific. Reef surveys established that size, branching structure and location (proximity to sand) of the coral colonies together determined the "suitability" of microhabitats for different ontogenetic stages of D. aruanus. Once these criteria were known, patterns of habitat use were quantified within lagoons of five Pacific islands. Availability of suitable habitat generally was an excellent predictor of density, and patterns were qualitatively consistent at several spatial scales, including among different lagoons on the same island, among different islands and between the central (French Polynesia and Rarotonga) and western (Great Barrier Reef, Australia) South Pacific. A field experiment that varied the amount of suitable coral among local plots indicated that habitat for settlers accounted for almost all of the spatial variation in the number of D. aruanus that settled at that location, suggesting that spatial patterns of abundance can be established at settlement without spatial variation in larval supply. Surveys of four other species of reef-associated fish revealed that a substantial fraction of their spatial variation in density also was explained by availability of suitable reef habitat, suggesting that habitat may be a prevalent determinant of spatial patterns. The results underscore the critical need to identify accurately the resource requirements of different species and life stages when evaluating causes of spatial variation in abundance of reef fishes.

Meteorological forcing of plankton dynamics in a large and deep continental European lake.

Abstract: The timing of various plankton successional events in Lake Constance was tightly coupled to a large- scale meteorological phenomenon, the North Atlantic Oscillation (NAO). A causal chain of meteorological, hydrological, and ecological processes connected the NAO as well as winter and early spring meteorological conditions to planktonic events in summer leading to a remarkable memory of climatic effects lasting over al- most half a year. The response of Daphnia to meteoro- logical forcing was most probably a direct effect of al- tered water temperatures on daphnid growth and was not mediated by changes in phytoplankton concentrations. High spring water temperatures during "high-NAO years" enabled high population growth rates, resulting in a high daphnid biomass as early as May. Hence, a critical Daphnia biomass to suppress phytoplankton was reached earlier in high-NAO years yielding an early and longer- lasting clear-water phase. Finally, an earlier summer de- cline of Daphnia produced in a negative relationship be- tween Daphnia biomass in July and the NAO. Meteoro- logical forcing of the seasonal plankton dynamics in Lake Constance included simple temporal shifts of pro- cesses and successional events, but also complex chang- es in the relative importance of different mechanisms. Since Daphnia plays an important role in plankton suc- cession, a thorough understanding of the regulation of its population dynamics provides the key for predictions of the response of freshwater planktonic food webs to glob- al climate change.