Showing papers in "Oecologia in 1999"

Tracing origins and migration of wildlife using stable isotopes: a review

Abstract: To understand the ecology of migratory animals it is important to link geographic regions used by individuals including breeding, wintering, and intermediate stopover sites. Previous conventional approaches used to track animal movements have relied on extrinsic markers and typically the subsequent recovery of individuals. This approach has generally been inappropriate for most small, or non-game animals. The use of intrinsic markers such as fatty acid profiles, molecular DNA analyses, and the measurement of naturally occurring stable isotopes in animal tissues offer alternative approaches. This paper reviews the use of stable isotope analyses (primarily δ13C, δ15N, δ34S, δD, δ87Sr) to trace nutritional origin and migration in animals. This approach relies on the fact that foodweb isotopic signatures are reflected in the tissues of organisms and that such signatures can vary spatially based on a variety of biogeochemical processes. Organisms moving between isotopically distinct foodwebs can carry with them information on the location of previous feeding. Such an approach has been used to track animal use of inshore versus offshore, marine versus freshwater, terrestrial C3 versus marine, terrestrial mesic versus xeric, and C3 versus C4 or Crassulacean acid metabolism foodwebs. More recently, the use of stable hydrogen isotope analyses (δD) to link organisms to broad geographic origin in North America is based on large-scale isotopic contours of growing-season average δD values in precipitation. This technique, especially when combined with the assay of other stable isotopes, will be extremely useful in helping to track migration and movement of a wide range of animals from insects to birds and mammals. Future research to refine our understanding of natural and anthropogenic-induced isotopic gradients in nature, and to explore the use of stable isotopes of other elements, is recommended.

Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies.

Abstract: The isotope enrichment ɛ* of 13C between tooth enamel of large ruminant mammals and their diet is 14.1 ± 0.5‰. This value was obtained by analyzing both the dental enamel of a variety of wild and captive mammals and the vegetation that comprised their foodstuffs. This isotope enrichment factor applies to a wide variety of ruminant mammals. Non-ruminant ungulates have a similar isotope enrichment, although our data cannot determine if it is significantly different. We also found a 13C isotope enrichment ɛ* of 3.1 ± 0.7‰ for horn relative to diet, and 11.1 ± 0.8‰ for enamel relative to horn for ruminant mammals. Tooth enamel is a faithful recorder of diet. Its isotopic composition can be used to track changes in the isotopic composition of the atmosphere, determine the fraction of C3 or C4 biomass in diets of modern or fossil mammals, distinguish between mammals using different subpathways of C4 photosynthesis,and identify those mammals whose diet is derived from closed-canopy habitats.

Effects of habitat isolation on pollinator communities and seed set

Abstract: Destruction and fragmentation of natural habitats is the major reason for the decreasing biodiversity in the agricultural landscape. Loss of populations may negatively affect biotic interactions and ecosystem stability. Here we tested the hypothesis that habitat fragmentation affects bee populations and thereby disrupts plant-pollinator interactions. We experimentally established small ”habitat islands” of two self-incompatible, annual crucifers on eight calcareous grasslands and in the intensively managed agricultural landscape at increasing distances (up to 1000 m) from these species-rich grasslands to measure effects of isolation on both pollinator guilds and seed set, independently from patch size and density, resource availability and genetic erosion of plant populations. Each habitat island consisted of four pots each with one plant of mustard (Sinapis arvensis) and radish (Raphanus sativus). Increasing isolation of the small habitat islands resulted in both decreased abundance and species richness of flower-visiting bees (Hymenoptera: Apoidea). Mean body size of flower-visiting wild bees was larger on isolated than on nonisolated habitat islands emphasizing the positive correlation of body size and foraging distance. Abundance of flower-visiting honeybees depended on the distance from the nearest apiary. Abundance of other flower visitors such as hover flies did not change with increasing isolation. Number of seeds per fruit and per plant decreased significantly with increasing distance from the nearest grassland for both mustard and radish. Mean seed set per plant was halved at a distance of approximately 1000 m for mustard and at 250 m for radish. In accordance with expectations, seed set per plant was positively correlated with the number of flower-visiting bees. We found no evidence for resource limitation in the case of mustard and only marginal effects for radish. We conclude that habitat connectivity is essential to maintain not only abundant and diverse bee communities, but also plant-pollinator interactions in economically important crops and endangered wild plants.

Leaf construction cost, nutrient concentration, and net CO 2 assimilation of native and invasive species in Hawaii

Abstract: The effects of biological invasions are most evident in isolated oceanic islands such as the Hawaiian Archipelago, where invasive plant species are rapidly changing the composition and function of plant communities. In this study, we compared the specific leaf area (SLA), leaf tissue construction cost (CC), leaf nutrient concentration, and net CO2 assimilation (A) of 83 populations of 34 native and 30 invasive species spanning elevation and substrate age gradients on Mauna Loa volcano in the island of Hawaii. In this complex environmental matrix, where annual precipitation is higher than 1500 mm, we predicted that invasive species, as a group, will have leaf traits, such as higher SLA and A and lower leaf CC, which may result in more efficient capture of limiting resources (use more resources at a lower carbon cost) than native species. Overall, invasive species had higher SLA and A, and lower CC than native species, consistent with our prediction. SLA and foliar N and P were 22.5%, 30.5%, and 37.5% higher, respectively, in invasive species compared to native ones. Light-saturated photosynthesis was higher for invasive species (9.59 μmol m−2 s−1) than for native species (7.31 μmol m−2 s−1), and the difference was larger when A was expressed on a mass basis. Leaf construction costs, on the other hand, were lower for the invasive species (1.33 equivalents of glucose g−1) than for native species (1.37). This difference was larger when CC was expressed on an area basis. The trends in the above traits were maintained when groups of ecologically equivalent native and invasive species (i.e., sharing similar life history traits and growing in the same habitat) were compared. Foliar N and P were significantly higher in invasive species across all growth forms. Higher N may partially explain the higher A of invasive species. Despite relatively high N, the photosynthetic nitrogen use efficiency of invasive species was 15% higher than that of native species. These results suggest that invasive species may not only use resources more efficiently than native species, but may potentially demonstrate higher growth rates, consistent with their rapid spread in isolated oceanic islands.

Nutrient fluxes from water to land: seabirds affect plant nutrient status on Gulf of California islands.

Abstract: Energy and nutrient fluxes across habitat boundaries can exert profound direct and indirect effects on the dynamics of recipient systems. Transport from land to water is common and well studied; here, we document a less recognized process, substantial flows from water to land. On hyperarid, naturally nutrient poor islands in the Gulf of California, nutrient input via seabird guano directly increases N and P concentrations up to 6-fold in soils; these nutrients enrich plants. Nutrients in a long-lived cactus, a short-lived shrub, and annuals were 1.6- to 2.4-fold greater on bird versus nonbird islands. Because plant quality affects consumer growth and reproduction, we suggest that nutrient enrichment via guano ramifies to affect the entire food web on these islands.

Ant-aphid mutualisms: the impact of honeydew production and honeydew sugar composition on ant preferences.

Abstract: The honeydew composition and production of four aphid species feeding on Tanacetum vulgare, and mutualistic relationships with the ant Lasius niger were studied. In honeydew of Metopeurum fuscoviride and Brachycaudus cardui, xylose, glucose, fructose, sucrose, maltose, melezitose, and raffinose were detected. The proportion of trisaccharides (melezitose, raffinose) ranged between 20% and 35%. No trisaccharides were found in honeydew of Aphis fabae, and honeydew of Macrosiphoniella tanacetaria consisted of only xylose, glucose and sucrose. M. fuscoviride produced by far the largest amounts of honeydew per time unit (880 μg/aphid per hour), followed by B. cardui (223 μg/aphid per hour), A. fabae (133 μg/aphid per hour) and M. tanacetaria (46 μg/aphid per hour). The qualitative and quantitative honeydew production of the aphid species corresponded well with the observed attendance by L. niger. L. niger workers preferred trisaccharides over disaccharides and monosaccharides when these sugars were offered in choice tests. The results are consistent with the ants' preference for M. fuscoviride, which produced the largest amount of honeydew including a considerable proportion of the trisaccharides melezitose and raffinose. The preference of L. niger for B. cardui over A. fabae, both producing similar amounts of honeydew, may be explained by the presence of trisaccharides and the higher total sugar concentration in B. cardui honeydew. M. tanacetaria, which produced only low quantities of honeydew with no trisaccharides was not attended at all by L. niger.

Insights into nitrogen and carbon dynamics of ectomycorrhizal and saprotrophic fungi from isotopic evidence.

Abstract: The successful use of natural abundances of carbon (C) and nitrogen (N) isotopes in the study of ecosystem dynamics suggests that isotopic measurements could yield new insights into the role of fungi in nitrogen and carbon cycling. Sporocarps of mycorrhizal and saprotrophic fungi, vegetation, and soils were collected in young, deciduous-dominated sites and older, coniferous-dominated sites along a successional sequence at Glacier Bay National Park, Alaska. Mycorrhizal fungi had consistently higher δ15N and lower δ13C values than saprotrophic fungi. Foliar δ13C values were always isotopically depleted relative to both fungal types. Foliar δ15N values were usually, but not always, more depleted than those in saprotrophic fungi, and were consistently more depleted than in mycorrhizal fungi. We hypothesize that an apparent isotopic fractionation by mycorrhizal fungi during the transfer of nitrogen to plants may be attributed to enzymatic reactions within the fungi producing isotopically depleted amino acids, which are subsequently passed on to plant symbionts. An increasing difference between soil mineral nitrogen δ15N and foliar δ15N in later succession might therefore be a consequence of greater reliance on mycorrhizal symbionts for nitrogen supply under nitrogen-limited conditions. Carbon signatures of mycorrhizal fungi may be more enriched than those of foliage because the fungi use isotopically enriched photosynthate such as simple sugars, in contrast to the mixture of compounds present in leaves. In addition, some 13C fractionation may occur during transport processes from leaves to roots, and during fungal chitin biosynthesis. Stable isotopes have the potential to help clarify the role of fungi in ecosystem processes.

A food web analysis of the juvenile blue crab, Callinectes sapidus, using stable isotopes in whole animals and individual amino acids

Abstract: The stable isotope compositions (C and N) of plants and animals of a marsh dominated by Spartina alterniflora in the Delaware Estuary were determined. The study focused on the juvenile stage of the Atlantic blue crab, Callinectes sapidus, and the importance of marsh-derived diets in supporting growth during this stage. Laboratory growth experiments and field data indicated that early juvenile blue crabs living in the Delaware Bay habitat fed primarily on zooplankton, while marsh-dwelling crabs, which were enriched in 13C relative to bay juveniles, utilized marsh-derived carbon for growth. In laboratory experiments, the degree to which juvenile blue crabs isotopically fractionated dietary nitrogen, as well as the growth rate, depended on the protein quality of the diet. The range of δ13C of amino acids in laboratory-reared crabs and their diets was almost 20‰, similar to the isotopic range of amino acids of other organisms. In laboratory studies, the δ13C of nonessential and essential amino acids in the diet were compared to those in juvenile crabs. Isotopic fractionation at the molecular level depended on diet quality and the crabs' physiological requirements. Comparison of whole-animal isotope data with individual amino acid C isotope measurements of wild juvenile blue crabs from the bay and marsh suggested a different source of total dietary carbon, yet a shared protein component, such as zooplankton.

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves.

Abstract: We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species -Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina- with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots.

Partitioning of soil water among canopy trees in a seasonally dry tropical forest

Abstract: Little is known about partitioning of soil water resources in species-rich, seasonally dry tropical forests. We assessed spatial and temporal patterns of soil water utilization in several canopy tree species on Barro Colorado Island, Panama, during the 1997 dry season. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θv), and sap flow were measured concurrently. Evaporative fractionation near the soil surface caused soil water δD to decrease from about –15‰ at 0.1 m to –50 to –55‰ at 1.2 m depth. Groundwater sampled at the sources of nearby springs during this period yielded an average δD value of –60‰. θv increased sharply and nearly linearly with depth to 0.7 m, then increased more slowly between 0.7 and 1.05 m. Based on xylem δD values, water uptake in some individual plants appeared to be restricted largely to the upper 20 cm of the soil profile where θv dropped below 20% during the dry season. In contrast, other individuals appeared to have access to water at depths greater than 1 m where θv remained above 45% throughout the dry season. The depths of water sources for trees with intermediate xylem δD values were less certain because variation in soil water δD between 20 and 70 cm was relatively small. Xylem water δD was also strongly dependent on tree size (diameter at breast height), with smaller trees appearing to preferentially tap deeper sources of soil water than larger trees. This relationship appeared to be species independent. Trees able to exploit progressively deeper sources of soil water during the dry season, as indicated by increasingly negative xylem δD values, were also able to maintain constant or even increase rates of water use. Seasonal courses of water use and soil water partitioning were associated with leaf phenology. Species with the smallest seasonal variability in leaf fall were also able to tap increasingly deep sources of soil water as the dry season progressed. Comparison of xylem, soil, and groundwater δD values thus pointed to spatial and temporal partitioning of water resources among several tropical forest canopy tree species during the dry season.

Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans

Abstract: The diet of extant elephants (Loxodonta in Africa, Elephas in Asia) is dominated by C3 browse although some elephants have a significant C4 grass component in their diet. This is particularly noteworthy because high-crowned elephantid cheek teeth represent adaptation to an abrasive grazing diet and because isotopic analysis demonstrates that C4 vegetation was the dominant diet for Elephas in Asia from 5 to 1 Ma and for both Loxodonta and Elephas in Africa between 5–1 Ma. Other proboscideans in Africa and southern Asia, except deinotheres, also had a C4-dominated diet from about 7 Ma (when the C4 biomass radiated in tropical and subtropical regions) until their subsequent extinction.

Role of brown bears (Ursus arctos) in the flow of marine nitrogen into a terrestrial ecosystem.

Abstract: We quantified the amount, spatial distribution, and importance of salmon (Oncorhynchus spp.)-derived nitrogen (N) by brown bears (Ursus arctos) on the Kenai Peninsula, Alaska. We tested and confirmed the hypothesis that the stable isotope signature (δ15N) of N in foliage of white spruce (Picea glauca) was inversely proportional to the distance from salmon-spawning streams (r=–0.99 and P<0.05 in two separate watersheds). Locations of radio-collared brown bears, relative to their distance from a stream, were highly correlated with δ15N depletion of foliage across the same gradient (r=–0.98 and –0.96 and P<0.05 in the same two separate watersheds). Mean rates of redistribution of salmon-derived N by adult female brown bears were 37.2±2.9 kg/year per bear (range 23.1–56.3), of which 96% (35.7±2.7 kg/year per bear) was excreted in urine, 3% (1.1±0.1 kg/year per bear) was excreted in feces, and <1% (0.3± 0.1 kg/year per bear) was retained in the body. On an area basis, salmon-N redistribution rates were as high as 5.1±0.7 mg/m2 per year per bear within 500 m of the stream but dropped off greatly with increasing distance. We estimated that 15.5–17.8% of the total N in spruce foliage within 500 m of the stream was derived from salmon. Of that, bears had distributed 83–84%. Thus, brown bears can be an important vector of salmon-derived N into riparian ecosystems, but their effects are highly variable spatially and a function of bear density.

Ecosystem properties and microbial community changes in primary succession on a glacier forefront.

Abstract: We studied microbial community composition in a primary successional chronosequence on the forefront of Lyman Glacier, Washington, United States. We sampled microbial communities in soil from nonvegetated areas and under the canopies of mycorrhizal and nonmycorrhizal plants from 20- to 80-year-old zones along the successional gradient. Three independent measures of microbial biomass were used: substrate-induced respiration (SIR), phospholipid fatty acid (PLFA) analysis, and direct microscopic counts. All methods indicated that biomass increased over successional time in the nonvegetated soil. PLFA analysis indicated that the microbial biomass was greater under the plant canopies than in the nonvegetated soils; the microbial community composition was clearly different between these two types of soils. Over the successional gradient, the microbial community shifted from bacterial-dominated to fungal-dominated. Microbial respiration increased while specific activity (respiration per unit biomass) decreased in nonvegetated soils over the successional gradient. We proposed and evaluated new parameters for estimating the C use efficiency of the soil microbial community: “Max” indicates the maximal respiration rate and “Acc” the total C released from the sample after a standard amount of substrate is added. These, as well as the corresponding specific activities (calculated as Max and Acc per unit biomass), decreased sharply over the successional gradient. Our study suggests that during the early stages of succession the microbial community cannot incorporate all the added substrate into its biomass, but rapidly increases its respiration. The later-stage microbial community cannot reach as high a rate of respiration per unit biomass but remains in an “energy-saving state,” accumulating C to its biomass.

Ecological and evolutionary aspects of isoprene emission from plants

Abstract: Isoprene (2-methyl-1,3,-butadiene), produced by many woody and a few herbaceous plant species, is the dominant volatile organic compound released from vegetation. It represents a non-trivial carbon loss to the plant (typically 0.5–2%, but much higher as temperatures exceed 30°C), and plays a major role in tropospheric chemistry of forested regions, contributing to ozone formation. This review summarizes current knowledge concerning the occurrence of isoprene production within the plant kingdom, and discusses other aspects of isoprene biology which may be of interest to the ecological community. The ability to produce significant amounts of isoprene may or may not be shared by members of the same plant family or genus, but emitting species have been found among bryophytes, ferns, conifers and Ephedra and in approximately one-third of the 122 angiosperm families examined. No phylogenetic pattern is obvious among the angiosperms, with the trait widely scattered and present (and absent) in both primitive and derived taxa, although confined largely to woody species. Isoprene is not stored within the leaf, and plays no known ecological role as, for example, an anti-herbivore or allelopathic agent. The primary short-term controls over isoprene production are light and temperature. Growth in high light stimulates isoprene production, and growth in cool conditions apparently inhibits isoprene, production of which may be induced upon transfer to warmer temperatures. The stimulation of isoprene production by high irradiance and warm temperatures suggests a possible role in ameliorating stresses associated with warm, high-light environments, a role consistent with physiological evidence indicating a role in thermal protection.

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

Abstract: We used data from a 15-year experiment in a C4-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C3 plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C4 grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C4-dominated grassland can also be directly related to community structure independent of disturbance.

Seed size and seedling emergence: an allometric relationship and some ecological implications.

Abstract: We develop a geometric model predicting that maximum seedling emergence depth should scale as the cube root of seed weight. We tested the prediction by planting seeds from 17 species ranging in weight from 0.1 to 100 mg at a variety of depths in a sand medium. The species were spread across 16 genera and 13 families, all occurring in fire-prone fynbos shrublands of South Africa. Maximum emergence depth was found to scale allometrically with seed weight with an exponent of 0.334, close to the predicted value. We used the allometry to predict recruitment response to experimentally simulated variation in fire intensity. Five species with small ( 10 mg) seeds were planted at ≤20-mm and 40-mm depths and exposed to low and high heat treatments and a control. The allometric equation predicted that species with large seeds would be able to emerge from a depth of 40 mm but those with small seeds would not. Only 1% of 481 seedlings from small-seeded species emerged from the 40-mm planting compared with 40% of 626 seedlings from the large-seeded group. The simulated fire treatments killed seeds in shallow, but not deeper, soil layers. At simulated high fire intensities, seedling emergence was poor in small-seeded species but good in large-seeded species, with most seedlings emerging from the 40-mm planting depth. Seed size could be a useful general predictor of recruitment success under different fire intensities in this system. We suggest that allometric relationships in plants deserve wider attention as predictive tools.

Evolution of flight morphology in a butterfly that has recently expanded its geographic range

Abstract: Individuals colonizing unoccupied habitats typically possess characters associated with increased dispersal and, in insects, colonization success has been related to flight morphology. The speckled wood butterfly, Pararge aegeria, has undergone recent major expansions in its distribution: in the north of its range, P. aegeria has colonized many areas in north and east England, and in the south, it was first recorded on Madeira in 1976. We examined morphological traits associated with flight and reproduction in the northern subspecies tircis, and in the southern subspecies aegeria, from sites colonized about 20 years ago in northern England and on Madeira, respectively. Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were from individuals reared in a common environment and there were significant family effects in most of the variables measured. Compared with individuals from sites continuously occupied in recent history, colonizing individuals were larger (adult live mass). In the subspecies tircis, colonizing individuals also had relatively larger thoraxes and lower wing aspect ratios indicating that evolutionary changes in flight morphology may be related to colonization. However, sex by site interactions in analyses of thorax mass and abdomen mass suggest different selection pressures on flight morphology between the sexes in relation to colonization. Overall, the subspecies aegeria was smaller (adult live mass) and had a relatively larger thorax and wings, and smaller abdomen than subspecies tircis. Evolutionary changes in flight morphology and dispersal rate may be important determinants of range expansion, and may affect responses to future climate change.

Niche relations among three sympatric Mediterranean carnivores

Abstract: Previous studies carried out in the Donana National Park reported that red foxes (Vulpes vulpes) were killed by Iberian lynxes (Lynx pardinus), whereas similar-sized Eurasian badgers (Meles meles) were not. Therefore, we predicted that fox would avoid lynx predation risk by niche segregation whereas we did not expect such a segregation between badger and lynx. As an approach for evaluating our predictions, we compared their diet, activity patterns, and habitat use in an area of Donana where the three carnivores are sympatric. Lynxes preyed almost uniquely on European rabbits (Oryctolagus cuniculus), and though badgers and foxes were omnivorous, rabbits also were a major prey, resulting in high overlaps throughout the year. However, badgers preyed largely on small rabbits, whereas lynxes and foxes preyed mainly on medium-sized rabbits. There were also interspecific differences in activity patterns. Maximum levels of activity among lynxes were during sunrise and dusk (49–67%). Foxes were most active during dusk and night (34–67%), and badgers were mainly nocturnal (53–87%). Though there were seasonal differences in the amount of activity of each species, specific activity patterns changed little throughout the year. There was a strong difference in annual habitat use by the three species (P < 0.0001). Lynxes used mainly the Mediterranean scrubland during both the active (PMAX) and the resting (PMIN) periods. During PMIN, foxes used the Mediterranean scrubland intensively (40% of locations on average), but during PMAX, they used the pastureland much more intensively despite this habitat being poorer in their main prey (rabbits). As a consequence, foxes and lynxes exhibited segregation in their habitat use during the active period. Badgers also used the Mediterranean scrubland intensively, especially during PMIN. There were no seasonal differences in habitat use for lynx and fox, but there was for badgers (P < 0.015). Within the study area, the three species selected habitat suggesting they were sensitive to factors such as vegetation and prey abundance. However, in general, carnivore habitat use did not correlate with rabbit abundance. We propose that foxes avoided lynxes by using, during activity, habitats not frequented by lynxes, and that a low predation risk associated with the distinctive foraging mode of badgers may facilitate its coexistence with other carnivores.

Growth, development, and survival of a generalist predator fed single- and mixed-species diets of different quality.

Abstract: It is often assumed that prey species consumed by generalist predators are largely, though not entirely, equivalent in terms of their value to the predators. In contrast to this expectation, laboratory feeding experiments uncovered distinctly varied developmental responses of a generalist predator, the wolf spider Schizocosa, to different experimental diets. Naive Schizocosa attacked and fed upon all the prey species offered; however, highly divergent patterns of survival, development, and growth of Schizocosa spiderlings reared on different single-prey diets revealed a wide spectrum of prey qualities. Spiderlings fed the collembolan Tomocerus bidentatus sustained the highest overall rates of survival, growth, and development. Fruit flies (Drosophila melanogaster) were intermediate-quality prey: spiders fed Drosophila initially exhibited rates of survival, growth, and development equal to those of spiders on a diet of T. bidentatus, but after about 3 months, rates declined markedly. Fungus gnats (Sciaridae; Bradysia sp.) and conspecfic spiderlings are low-quality prey for Schizocosa: a sole diet of either of these prey types resulted in positive but markedly submaximal rates of growth, retarded rates of development, and survival rates much lower than that supported by a diet of Drosophila. Worst were the collembolans Folsomia candida and Isotoma trispinata, and the aphid Aphis nerii: spiderlings fed solely one of these species did not grow and died without molting. A. nerii is classified as poor quality because survival was no better than that of starved controls. F. candida and I. trispinata were toxic: survival of Schizocosa hatchlings fed these collembolans was lower than that of starved controls. A mixed diet of T. bidentatus and fruit flies yielded positive synergistic effects with respect to growth, but development and rate of survival were similar to those of spiders on a sole diet of T. bidentatus. Including toxic prey did not produce a better diet, while inclusion of toxic prey with prey of higher quality created diets that were no better than the toxic prey alone. The results of these experiments suggest that prey species that are similar in morphology and behavior, and that are initially killed and consumed, may differ dramatically in their suitability as food for generalist arthropod predators.

Foliage physiology and biochemistry in response to light gradients in conifers with varying shade tolerance.

Abstract: To examine the predictability of leaf physiology and biochemistry from light gradients within canopies, we measured photosynthetic light-response curves, leaf mass per area (LMA) and concentrations of nitrogen, phosphorus and chlorophyll at 15-20 positions within canopies of three conifer species with increasing shade tolerance, ponderosa pine [Pinus ponderosa (Laws.)], Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. Adjacent to each sampling position, we continuously monitored photosynthetically active photon flux density (PPFD) over a 5-week period using quantum sensors. From these measurements we calculated FPAR: integrated PPFD at each sampling point as a fraction of full sun. From the shadiest to the brightest canopy positions, LMA increased by about 50% in ponderosa pine and 100% in western hemlock; Douglas fir was intermediate. Canopy-average LMA increased with decreasing shade tolerance. Most foliage properties showed more variability within and between canopies when expressed on a leaf area basis than on a leaf mass basis, although the reverse was true for chlorophyll. Where foliage biochemistry or physiology was correlated with FPAR, the relationships were non-linear, tending to reach a plateau at about 50% of full sunlight. Slopes of response functions relating physiology and biochemistry to ln(FPAR) were not significantly different among species except for the light compensation point, which did not vary in response to light in ponderosa pine, but did in the other two species. We used the physiological measurements for Douglas fir in a model to simulate canopy photosynthetic potential (daily net carbon gain limited only by PPFD) and tested the hypothesis that allocation of carbon and nitrogen is optimized relative to PPFD gradients. Simulated photosynthetic potential for the whole canopy was slightly higher (<10%) using the measured allocation of C and N within the canopy compared with no stratification (i.e., all foliage identical). However, there was no evidence that the actual allocation pattern was optimized on the basis of PPFD gradients alone; simulated net carbon assimilation increased still further when even more N and C were allocated to high-light environments at the canopy top.

Individual specialization and trophic adaptability of northern pike ( Esox lucius ): an isotope and dietary analysis

Abstract: Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike (“pike-only”) and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) (“pike-other”). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization.

Growth and reproduction of Antarctic vascular plants in response to warming and UV radiation reductions in the field

Abstract: Along the west coast of the Antarctic Peninsula springtime ozone depletion events can lead to a two-fold increase in biologically effective UV-B radiation (UV-BBE) and summer air temperatures have risen ≈1.5°C during the past 50 years. We manipulated levels of UV radiation and temperature around Colobanthus quitensis (a cushion-forming plant, Caryophyllaceae) and Deschampsia antarctica (a tussock grass) along the Peninsula near Palmer Station for two field seasons. Ambient levels of UV were manipulated by placing filters that either transmitted UV (filter control), absorbed UV-B (reducing diurnal levels of UV-BBE by about 82%), or absorbed both UV-B and UV-A (reducing UV-BBE and UV-ABE by about 88 and 78%, respectively) on frames over naturally growing plants from November to March. Half the filters of each material completely surrounded the frames and raised diurnal and diel air temperatures around plants by an average of 2.3°C and 1.3°C, respectively. Reducing UV or warming had no effect on leaf concentrations of soluble UV-B absorbing compounds, UV-B absorbing surface waxes or chlorophylls. Warming had few effects on growth of either species over the first season. However, over the second field season warming improved growth of C. quitensis, leading to a 50% increase in leaf production (P < 0.10), a 26% increase in shoot production, and a 6% increase in foliar cover. In contrast, warming reduced growth of D. antarctica, leading to a 20% decline in leaf length, a 17% decline in leaf production (P < 0.10), and a 5% decline in foliar cover. Warming improved sexual reproduction in both species, primarily through faster development of reproductive structures and greater production of heavier seeds. Over the second field season, the percentage of reproductive structures that had reached the most developed (seed) stage in C. quitensis and D. antarctica was 20% and 15% higher, respectively, under warming. Capsules of C. quitensis produced 45% more seeds under warming and these seeds were 11% heavier. Growth of D. antarctica was improved when UV was reduced and these effects appeared to be cumulative over field seasons. Over the second season, tillers produced 55% more leaves and these leaves were 32% longer when UV-B was reduced. Tillers produced 137% more leaves that were 67% longer when both UV-B and UV-A were reduced. The effects of UV reduction were not as pronounced on C. quitensis, although over the second season cushions tended to be 17% larger and produce 21% more branches when UV-B was reduced, and tended to be 27% larger and produce 38% more branches when both UV-B and UV-A were reduced (P < 0.10). Few interactions were found between UV reduction and warming, although in the absence of warming, reducing UV led to slower development of reproductive structures in both species. The effects of warming and UV reduction were species specific and were often cumulative over the two field seasons, emphasizing the importance of long-term field manipulations in predicting the impacts of climate change.

Stable isotopes (δD and δ 13 C) are geographic indicators of natal origins of monarch butterflies in eastern North America

Abstract: Wing membranes of laboratory and field-reared monarch butterflies (Danaus plexippus) were analyzed for their stable-hydrogen (δD) and carbon (δ13C) isotope ratios to determine whether this technique could be used to identify their natal origins. We hypothesized that the hydrogen isotopic composition of monarch butterfly wing keratin would reflect the hydrogen isotope patterns of rainfall in areas of natal origin where wings were formed. Monarchs were reared in the laboratory on milkweed plants (Asclepias sp.) grown with water of known deuterium content, and, with the assistance of volunteers, on native milkweeds throughout eastern North America. The results show that the stable hydrogen isotopic composition of monarch butterflies is highly correlated with the isotopic composition of the milkweed host plants, which in turn corresponds closely with the long-term geographic patterns of deuterium in rainfall. Stable-carbon isotope values in milkweed host plants were similarly correlated with those values in monarch butterflies and showed a general pattern of enrichment along a southwest to northeast gradient bisecting the Great Lakes. These findings indicate that natal origins of migratory and wintering monarchs in Mexico can be inferred from the combined δD and δ13C isotopic signatures in their wings. This relationship establishes that analysis of hydrogen and carbon isotopes can be used to answer questions concerning the biology of migratory monarch butterflies and provides a new approach to tracking similar migratory movements of other organisms.

Restored cut-away peatland as a sink for atmospheric CO2.

Abstract: In a field study, we examined the relationship between vegetation, abiotic factors and the CO2 exchange dynamics of a cut-away peatland 20 years after production had ended. The main objective was to determine the effect of rewetting on the CO2 exchange dynamics, measured separately in Eriophorum vaginatum tussocks and intertussocks (almost non-vegetated surfaces) using closed-chamber techniques, one growing season before and three growing seasons after the rewetting treatment. Rewetting lowered total respiration (RTOT) and increased gross photosynthesis (PG), which resulted in a higher incorporation of CO2 into the system. The seasonal CO2 balance for the almost continuously submerged section of the rewetted site became positive 2 years after rewetting (9.1 g CO2-C m−2), and it was still higher in the 3rd year (64.5 g CO2-C m−2), i.e. the system accumulated carbon. In the driest section of the rewetted site the seasonal balance increased strongly, but the balance was still negative during the 3 years following rewetting with losses from the system of 44.1, 26.1, 38.3 g CO2-C m−2 in 1995, 1996 and 1997 respectively. At the control site seasonal balance during 1995–1997 varied between ecosystem C losses of 41.8 and 95.3 in an area with high Eriophorum cover and between 52.1 and 109.9 g CO2-C m−2 with lower cover. Simulation of a cut-away peatland with dense Eriophorum vegetation (Eriophorum cover 70%) showed that if the water level (WT) is low, the seasonal CO2 balance of the ecosystem can reach the compensation point of no net C change (PG = RTOT) only if weather conditions are favourable, but with a high WT the seasonal CO2 balance would be positive even under varying weather conditions. It can be concluded that with dense Eriophorum vegetation a restored cut-away peatland acts as a functional mire and becomes a sink for atmospheric CO2.

Isotopic tracking of foraging and long-distance migration in northeastern Pacific pinnipeds.

Abstract: We investigated the impact of foraging location (nearshore vs offshore) and foraging latitude (high vs middle) on the carbon (δ13C) and nitrogen (δ15N) isotope compositions of bone collagen of northern fur seals (Callorhinus ursinus), harbor seals (Phoca vitulina), California sea lions (Zalophus californianus), and northern elephant seals (Mirounga angustirostris). Nearshore-foraging harbor seals from California had δ13C values 2.0‰ higher than female northern elephant seals foraging offshore at similar latitudes. Likewise, nearshore-foraging harbor seals from Alaska had values 1.7‰ higher than male northern fur seals, which forage offshore at high latitudes. Middle-latitude pinnipeds foraging in either the nearshore or offshore were 13C enriched by ∼1.0‰ over similar populations from high latitudes. Male northern elephant seals migrate between middle and high latitudes, but they had δ13C values similar to high-latitude, nearshore foragers. Female northern fur seal δ13C values were intermediate between those of high- and middle-latitude offshore foragers, reflecting their migration between high- and middle-latitude waters. The δ13C values of California sea lions were intermediate between nearshore- and offshore-foraging pinnipeds at middle latitudes, yet there was no observational support for the suggestion that they use offshore food webs. We suggest that their “intermediate” values reflect migration between highly productive and less-productive, nearshore ecosystems on the Pacific coasts of California and Mexico. The relative uniformity among all of these pinnipeds in δ15N values, which are strongly sensitive to trophic level, reveals that the carbon isotope patterns result from differences in the δ13C of organic carbon at the base of the food web, rather than differences in trophic structure, among these regions. Finally, the magnitude and direction of the observed nearshore-offshore and high-to middle-latitude differences in δ13C values suggest that these gradients may chiefly reflect differences in rates and magnitudes of phytoplankton production as well as the δ13C value of inorganic carbon available for photosynthesis, rather than the input of 13C-enriched macroalgal carbon to nearshore food webs.

Resource partitioning between sympatric wild and domestic herbivores in the Tarangire region of Tanzania

Abstract: The effect of the introduction of an exotic species (cattle) into a native African herbivore assemblage was investigated by studying resource partitioning between zebu cattle, wildebeest and zebra. Resource partitioning was investigated by analysing grass sward characteristics (such as sward height and percentage nitrogen in leaves) of feeding sites selected by the different herbivore species. Linear discriminant analysis was used to determine whether a distinction could be made between feeding sites selected by the different animal species or whether the animal species showed overlap in resource use by selecting similar feeding sites. Wildebeest and zebra did not show overlap in resource use except in the wet season when resources were ample. Cattle showed overlap in resource use with zebra in the early wet season and with wildebeest in the early dry season, seasons when food limitation is likely. In the wet season, cattle showed overlap in resource use with both zebra and wildebeest. Implications of these results for competitive relationships between livestock and wildlife are discussed. We suggest that interpretation of overlap in resource use may be different for an assemblage of long-term coexisting native species as compared to an assemblage of native and exotic species. Among native herbivores, overlap in resource use is not expected based on evolutionary segregation. In a native assemblage to which an exotic species has been introduced, however, overlap in resource use can occur under food-limited conditions and consequently implies competition.

The importance of scale of patchiness for selectivity in grazing herbivores.

Abstract: The notion that spatial scale is an important determinant of foraging selectivity and habitat utilization has only recently been recognized. We predicted and tested the effects of scale of patchiness on movements and selectivity of a large grazer in a controlled field experiment. We created random mosaics of short/high-quality and tall/low-quality grass patches in equal proportion at grid sizes of 2×2 m and 5×5 m. Subsequently, we monitored the foraging behaviour of four steers in 16 20×40 m plots over 30-min periods. As predicted on the basis of nutrient intake maximization, the animals selected the short patches, both by walking in a non-random manner and by additional selectivity for feeding stations. The tortuosity of foraging paths was similar at both scales of patchiness but selectivity was more pronounced in large patches than in small ones. In contrast, the number of bites per feeding station was not affected by patch size, suggesting that selection between and within feeding stations are essentially different processes. Mean residence time at individual feeding stations could not be successfully predicted on the basis of the marginal-value theorem: the animals stayed longer than expected, especially in the less profitable patch type. The distribution of the number of bites per feeding station suggests a constant probability to stay to feed or to move on to the next feeding station. This implies that the animals do not treat larger patches as discrete feeding stations but rather as a continuous resource. Our results have important implications for the application of optimal foraging theory in patchy environments. We conclude that selectivity in grazers is facilitated by large-scale heterogeneity, particularly by enhancing discrimination between feeding stations and larger selection units.

Links between the detritivore and the herbivore system: effects of earthworms and Collembola on plant growth and aphid development.

Abstract: Effects of Collembola (Heteromurus nitidus and Onychiurus scotarius) and earthworms (Aporrectodea caliginosa and Octolasion tyrtaeum) on the growth of two plant species from different functional groups (Poa annua and Trifolium repens), and on the development of aphids (Myzus persicae) were investigated in a laboratory experiment lasting 20 weeks. Using soil from a fallow site which had been set aside for about 15 years, we expected that nitrogen would be of limited supply to plants and hypothesized that the soil animals studied, particularly earthworms, would increase nutrient availability to plants and thereby also modify aphid reproduction and development. Plant growth was modified strongly by the presence of soil animals. Earthworms caused a more than twofold increase in shoot and root mass of P. annua but increased that of T. repens by only 18% and 6%, respectively. However, earthworms neither affected plant shoot/root ratio nor the nitrogen concentration in plant tissue. In contrast, the presence of Collembola caused a reduction in plant biomass particularly that of P. annua roots, but plant tissue nitrogen concentration was increased, although only slightly. Aphid reproduction on T. repens was lowered in the presence of Collembola on average by 45% but on P. annua increased by a factor of about 3. It is concluded that Collembola decrease aphid reproduction on more palatable host plants like T. repens but increase that on less palatable ones like P. annua. Earthworm presence also affected aphid reproduction but the effect was less consistent than that of Collembola. In the presence of earthworms, aphid reproduction was in one experimental period increased by some 70%. Earthworms also modified the numbers of Collembola and their vertical distribution in experimental chambers. Exploitation of deeper soil layers by H. nitidus was increased but, generally, O. scotarius numbers were reduced whereas those of H. nitidus increased in earthworm treatments. The presence of Collembola also influenced earthworm body mass during the experiment. In general it declined, but in the presence of Collembola loss of body mass of A. caliginosa was more pronounced. We conclude that inhibiting effects between Collembola and earthworms resulted from the use of a common resource, litter material rich in nitrogen. This is supported by the higher C/N ratio of the litter material in the presence of earthworms and Collembola by the end of the experiment. Effects of soil invertebrates like Collembola and earthworms on plant performance and aphid development are assumed to be modified by complex direct and indirect interactions among soil animal groups.

Experimental test of the effect of maternal hormones on larval quality of a coral reef fish.

Abstract: Maternal hormones can play an important role in the development of fish larvae. Levels of the stress hormone, cortisol, in females are elevated by social interactions and transferred directly to the yolk of eggs, where they may influence developmental rates. In some vertebrates, prenatal exposure to high levels of testosterone determine early growth rates, social status and reproductive success. The present study examined whether post-fertilization exposure of eggs of the tropical damselfish, Pomacentrus amboinensis (Pomacentridae), to natural levels of cortisol or testosterone directly affects larval morphology at hatching. Maternal and egg levels of cortisol and testosterone varied widely among clutches of eggs from local populations around Lizard Island on the Great Barrier Reef. The morphology of larvae produced by these local fish populations also varied widely and differed significantly among sites (e.g., standard length: 2.6–3.4 mm; yolk sac area: 0.01–0.13 × 10−2 mm2). Laboratory experiments showed that elevated cortisol levels in the egg reduced larval length at hatching, while slight elevations in testosterone increased yolk sac size. The influence of testosterone, and to a smaller extent cortisol, on larval morphology differed among egg clutches. These differences were partly explained by differences in initial egg hormone levels. Morphological changes induced by experimental hormonal regimes encompassed the entire range of variability in body attributes found in field populations. It is unclear whether cortisol influences growth alone or development rate or both. Testosterone appears to influence yolk utilization rates, and has no significant effect on growth, in contrast to its role in later developmental stages. Maternally derived cortisol and testosterone are important in regulating growth, development, and nutritive reserves of the embryo and larvae of this fish species. Factors that influence the maternal levels of cortisol and testosterone may have a major impact on larval mortality schedules and, therefore, on which breeding individuals contribute to the next generation.

Estimating the uptake of traffic-derived NO2 from 15N abundance in Norway spruce needles

Abstract: The 15N ratio of nitrogen oxides (NOx) emitted from vehicles, measured in the air adjacent to a highway in the Swiss Middle Land, was very high [δ15N(NO2) = +5.7‰]. This high 15N abundance was used to estimate long-term NO2 dry deposition into a forest ecosystem by measuring δ15N in the needles and the soil of potted and autochthonous spruce trees [Picea abies (L.) Karst] exposed to NO2 in a transect orthogonal to the highway. δ15N in the current-year needles of potted trees was 2.0‰ higher than that of the control after 4 months of exposure close to the highway, suggesting a 25% contribution to the N-nutrition of these needles. Needle fall into the pots was prevented by grids placed above the soil, while the continuous decomposition of needle litter below the autochthonous trees over previous years has increased δ15N values in the soil, resulting in parallel gradients of δ15N in soil and needles with distance from the highway. Estimates of NO2 uptake into needles obtained from the δ15N data were significantly correlated with the inputs calculated with a shoot gas exchange model based on a parameterisation widely used in deposition modelling. Therefore, we provide an indication of estimated N inputs to forest ecosystems via dry deposition of NO2 at the receptor level under field conditions.