scispace - formally typeset
Search or ask a question

Showing papers in "Oecologia in 1991"


Journal ArticleDOI
TL;DR: While LSM is a useful measure of sclerophylly, its separation into leaf density and thickness may be more appropriate as they often vary independently and appear to be more responsive to environmental gradients than LSM.
Abstract: We explored the relationship between leaf specific mass (LSM) and its two components, leaf density and thickness. These were assessed on the leaves of (a) the moderately sclerophyllous tree Arbutus menziesii distributed along a natural nutrient/moisture gradient in California, (b) eight sclerophyllous shrub species on four substrates in south-western Australia, and (c) seedlings of two morphologically contrasting Hakea species grown under varying soil nutrient, moisture and light regimes in a glasshouse experiment. Leaf area, mass, LSM, density and thickness varied greatly between leaves on the same plant, different species, and with different nutrient, moisture and light regimes. In some cases, variations in LSM were due to changes in leaf density in particular or thickness or both, while in others, density and thickness varied without a net effect on LSM. At lower nutrient or moisture availabilities or at higher light irradiances, leaves tended to be smaller, with higher LSM, density and thickness. Under increased stress, the thickness (diameter) of needle leaves decreased despite an increase in LSM. We concluded that, while LSM is a useful measure of sclerophylly, its separation into leaf density and thickness may be more appropriate as they often vary independently and appear to be more responsive to environmental gradients than LSM.

651 citations


Journal ArticleDOI
TL;DR: The results indicate that species allocate resources towards a high photosynthetic assimilation rate for a brief time, or provide resistant physical structure that results in a lower rate of carbon assimilation over a longer time, but not both.
Abstract: The relationships between resource availability, plant succession, and species' life history traits are often considered key to understanding variation among species and communities. Leaf lifespan is one trait important in this regard. We observed that leaf lifespan varies 30-fold among 23 species from natural and disturbed communities within a 1-km radius in the northern Amazon basin, near San Carlos de Rio Negro, Venezuela. Moreover, leaf lifespan was highly correlated with a number of important leaf structural and functional characterisues. Stomatal conductance to water vapor (g) and both mass and area-based net photosynthesis decreased with increasing leaf lifespan (r2=0.74, 0.91 and 0.75, respectively). Specific leaf area (SLA) also decreased with increasing leaf lifespan (r2=0.78), while leaf toughness increased (r2=0.62). Correlations between leaf lifespan and leaf nitrogen and phosphorus concentrations were moderate on a weight basis and not significant on an area basis. On an absolute basis, changes in SLA, net photosynthesis and leaf chemistry were large as leaf lifespan varied from 1.5 to 12 months, but such changes were small as leaf lifespan increased from 1 to 5 years. Mass-based net photosynthesis (A/mass) was highly correlated with SLA (r2=0.90) and mass-based leaf nitrogen (N/mass) (r2=0.85), but area-based net photosynthesis (A/area) was not well correlated with any index of leaf structure or chemistry including N/area. Overall, these results indicate that species allocate resources towards a high photosynthetic assimilation rate for a brief time, or provide resistant physical structure that results in a lower rate of carbon assimilation over a longer time, but not both.

578 citations


Journal ArticleDOI
TL;DR: Analysis of plants sampled from high and low altitudes along latitudinal gradients from the equator to the polar ends of plant distribution suggests that both temperature and atmospheric pressure are responsible for the altitudinal trends in 13C discrimination.
Abstract: In an earlier paper we provided evidence that carbon isotope discrimination during photosynthesis of terrestrial C3 plants decreases with altitude, and it was found that this was associated with greater carboxylation efficiency at high altitudes. Changing partial pressures of CO2 and O2 and changing temperature are possible explanations, since influences of moisture and light were reduced to a minimum by selective sampling. Here we analyse plants sampled using the same criteria, but from high and low altitudes along latitudinal gradients from the equator to the polar ends of plant distribution. These data should permit separation of the pressure and temperature components (Fig. 1). Only leaves of fully sunlit, non-water-stressed, herbaceous C3 plants are compared. The survey covers pressure differences of 400 mbar (ca. 5000 m) and 78 degrees of latitude (ca 25 K of mean temperature of growth period). When habitats of similar low temperature (i.e. high altitude at low latitude and low altitude at polar latitude) are compared, discrimination increases towards the pole (with decreasing altitude and thus increasing atmospheric pressure). Latitudinally decreasing temperature at almost constant atmospheric pressure (samples from low altitude) is associated with a decrease in discrimination. So, polar low-altitude plants have δ13C values half way between humid tropical lowland and tropical alpine plants. It is unlikely that latitudinal changes of the light regime had an effect, since low and high altitude plants show contrasting latitudinal trends in δ13C although local altitudinal differences in overall light consumption were small. These results suggest that both temperature and atmospheric pressure are responsible for the altitudinal trends in 13C discrimination. Temperature effects may partly be related to increased leaf thickness (within the same leaf type) in cold environments. Theoretical considerations and laboratory experiments suggest that it is the oxygen partial pressure that is responsible for the pressure related change in discrimination. The study also provided results of practical significance for the use of carbon isotope data. Within a community of C3 plants, discrimination in species of similar life form, exposed to similar light, water and ambient CO2 conditions ranges over 4‰, with standard deviations for 10–30 species of ±0.6 to 1.2‰. This natural variation has to be taken into account by using a sufficient sample size and standardization of sampling in any attempt at ecological site characterization using carbon isotope data. Evidence of a pronounced genotypic component of this variation in 13C discrimination in wild C3 plant species is provided. Correlations with dry matter partitioning, mesophyll thickness and nitrogen content are also present.

555 citations


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

442 citations


Journal ArticleDOI
TL;DR: Measurements of dry matter partitioning and growth in patchy conditions indicate a consistent positive association between the foraging characteristics of roots and shoots, supporting the hypothesis of strong interdependence of competitive abilities for light and mineral nutrients.
Abstract: There is widespread uncertainty about the nature and role of morphological plasticity in resource competition in plant communities. We have assayed the foraging characteristics of leaf canopies and root systems of eight herbaceous plants of contrasted ecology using new techniques to create controlled patchiness in light and mineral nutrient supply. The results are compared with those of a conventional competition experiment. Measurements of dry matter partitioning and growth in patchy conditions indicate a consistent positive association between the foraging characteristics of roots and shoots, supporting the hypothesis of strong interdependence of competitive abilities for light and mineral nutrients. Differences are identified in the abilities of dominant and subordinate plants to forage on coarse and fine scalcs. It is suggested that a trade-off exists in the scale (“high” in dominants) and precision (high in subordinates) with which resources are intercepted and that this trade-off contributes to diversity in communities of competing plants.

364 citations


Journal ArticleDOI
TL;DR: The result confirms earlier conclusions from gas exchange measurements that gaseous air pollutants did no long-lasting damage in an area where such damage was expected and the problem of separation of ammonium or nitrate use by roots from different soil horizons is discussed.
Abstract: Natural carbon and nitrogen isotope ratios were measured in different compartments (needles and twigs of different ages and crown positions, litter, understorey vegetation, roots and soils of different horizons) on 5 plots of a healthy and on 8 plots of a declining Norway spruce (Picea abies (L.) Karst.) forest in the Fichtelgebirge (NE Bavaria, Germany), which has recently been described in detail (Oren et al. 1988a; Schulze et al. 1989). The δ13C values of needles did not differ between sites or change consistently with needle age, but did decrease from the sun-to the shade-crown. This result confirms earlier conclusions from gas exchange measurements that gaseous air pollutants did no long-lasting damage in an area where such damage was expected. Twigs (δ13C between-25.3 and-27.8‰) were significantly less depleted in 13C than needles (δ13C between-27.3 and-29.1‰), and δ13C in twigs increased consistently with age. The δ15N values of needles ranged between-2.5 and-4.1‰ and varied according to stand and age. In young needles δ15N decreased with needle age, but remained constant or increased in needles that were 2 or 3 years old. Needles from the healthy site were more depleted in 15N than those from the declining site. The difference between sites was greater in old needles than in young ones. This differentiation presumably reflects an earlier onset of nitrogen reallocation in needles of the declining stand. δ15N values in twigs were more negative than in needles (-3.5 to-5.2‰) and showed age- and stand-dependent trends that were similar to the needles. δ15N values of roots and soil samples increased at both stands with soil depth from-3.5 in the organic layer to +4‰ in the mineral soil. The δ15N values of roots from the mineral soil were different from those of twigs and needles. Roots from the shallower organic layer had values similar to twigs and needles. Thus, the bulk of the assimilated nitrogen was presumably taken up by the roots from the organic layer. The problem of separation of ammonium or nitrate use by roots from different soil horizons is discussed.

320 citations


Journal ArticleDOI
TL;DR: This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.
Abstract: In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m−2 yr−1). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.

302 citations


Journal ArticleDOI
TL;DR: The results of the experiments show that predation risk from wading/diving animals (e.g., herons and raccoons) is much higher for larger fishes in shallow water than for these fishes in deeper water or for smaller fish in shallowWater.
Abstract: Surveys of 262 pools in 3 small streams in eastern Tennessee demonstrated a strong positive relationship between pool depth and the size of the largest fish within a pool (P 10.8 d in deep pools. In the striped shiner experiment in shallow artificial streamside troughs, no individuals 75–100 mm TL survived as long as 13 d, where-as smaller (20–25 mm) fish had 100% survival over 13 d. The results of the experiments show that predation risk from wading/diving animals (e.g., herons and raccoons) is much higher for larger fishes in shallow water than for these fishes in deeper water or for smaller fish in shallow water. We discuss the role of predation risk from two sources (piscivorous fish, which are more effective in deeper habitats, and diving/wading predators, which are more effective in shallow habitats) in contributing to the bigger fish — deeper habitat pattern in streams.

298 citations


Journal ArticleDOI
TL;DR: The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.
Abstract: The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H2O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.

267 citations


Journal ArticleDOI
TL;DR: The results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like “semi-generalist” predators, and declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.
Abstract: We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices of mustelids were derived from snow-tracking, diet composition from scat samples, and vole abundances from snap-trapping. Predation rate was estimated by the ratio of voles to mustelids and by the vole kill rate by predators (density of predator x percentage of voles in the diet). We tested the following four predictions of the hypothesis that small mustelids cause the low phase of the microtine cycle. (1) The densities of predators should lag well behind the prey abundances, as time lags tend to have destabilizing effects. The densities of stoats fluctuated in accordance with the vole abundances, whereas the spring densities of least weasels tracked the vole abundances with a half-year lag and the autumn densities with a 1-year lag. (2) Predators should not shift to alternative prey with declining vole densities. The yearly proportion of Microtus voles (the staple prey) in the diet of stoats varied widely (range 16–82%) and was positively correlated with the winter abundance of these voles. In contrast, the same proportion in the food of least weasels was independent of the vole abundance. (3) The ratio of voles to small mustelids should be smallest in poor vole years and largest in good ones. This was also observed. (4) Vole densities from autumn to spring should decrease more in those winters when vole kill rates are high than when they are low. The data on least weasels agreed with this prediction. Our results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like “semi-generalist” predators. Accordingly, declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.

261 citations


Journal ArticleDOI
TL;DR: The capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood, was examined, showing that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources.
Abstract: We examined the capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood (Populus angustifolia). A series of 14C-labeling experiments and a biomass allocation experiment showed that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources. Early gall development was dependent on aphid sinks increasing allocation from storage reserves of the stem, and later development of the progeny within the gall was dependent on resources from the galled leaf blade and from neighboring leaves. Regardless of gall position within a leaf, aphids intercepted 14C exported from the galled leaf (a non-mobilized source). However, only aphid galls at the most basal site of the leaf were strong sinks for 14C fixed in neighboring leaves (a mobilized source). Drawing resources from neighboring leaves represents active herbivore manipulation of normal host transport patterns. Neighboring leaves supplied 29% of the 14C accumulating in aphids in basal galls, while only supplying 7% to aphids in distal galls. This additional resource available to aphids in basal galls can account for the 65% increase in progeny produced in basal galls compared to galls located more distally on the leaf and limited to the galled leaf as a food resource. Developing furits also act as skins and compete with aphid-induced sinks for food supply. Aphid success in producing galls was increased 31% when surrounding female catkins were removed.

Journal ArticleDOI
N. A. O'Connor1
TL;DR: Analysis of species richness masked the complexity of species responses revealed by multivariate analyses of species abundances, and showed that different species groups selected different microhabitats on snags, particularly in response to the level of sediment deposition, which was greater on more structurally complex snags.
Abstract: Woody debris is a major structural component of south-eastern Australian lowland streams, and the decayed wood substrates provide a structurally complex habitat for macroinvertebrate colonization. I tested for the presence of a species richness-habitat complexity relationship for macroinvertebrate species inhabiting the surfaces of decayed submerged logs (snags) in a lowland stream in northern Victoria. The species-habitat complexity relationship is defined as the increase in species richness due to increased structural complexity of a habitat when area is held constant. The response of macroinvertebrates to seven treatments of artificial and natural substrates of differing levels and types of structural complexity were examined using cluster analyses and MANOVAs. These analyses revealed a significant species-habitat complexity relationship. In addition, a comparison of species evenness between simple and complex habitats supported the hypothesis that more complex habitats contained more species because they possessed more resources. Analysis of species richness, though informative, masked the complexity of species responses revealed by multivariate analyses of species abundances. These analyses showed that different species groups selected different microhabitats on snags, particularly in response to the level of sediment deposition, which was greater on more structurally complex snags. In comparison with the benthos, snags were significantly richer in species abundances, possibly related to low levels of dissolved oxygen in benthic habitats.

Journal ArticleDOI
TL;DR: The hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery is tested is tested.
Abstract: We tested the hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery. Growth response to defoliation went from partial compensation when plants were growing at high relative growth rates (RGR) to overcompensation when plants were more stressed and growing at low RGR. Defoliation released plants from the limitation imposed by the accumulation of old and dead tissue and this release overrode the negative effect of biomass loss. Compensatory growth resulted from a higher RGR aboveground that was not associated with a reduction in RGR belowground. Time available for recovery had a major impact on the outcome of defoliation. With a short time for recovery, RGR was decreased by defoliation because an immediate increase in net assimilation rate was overridden by a reduction in the ratio of leaf area to plant weight. After defoliation, this ratio increased quickly due to a larger allocation to leaf growth and lower leaf specific weights, resulting in higher RGR. We conclude that the compensatory response to grazing depends on the type and level of stress limiting growth. Allocation and physiological responses to stress may positively or negatively affect the response to grazing and, simultaneously, grazing may alleviate or aggravate the effects of different types of stress.

Journal ArticleDOI
TL;DR: The dominant C4 grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.
Abstract: The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C4 grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C4 grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.

Journal ArticleDOI
TL;DR: Differences between the early and late successional species in the allocation of nitrogen into RuBP carboxylase and thylakoid nitrogen pools or non-photosynthetic compounds are suggested by the distinct relationships between maximum photosynthetic capacity and nitrogen content.
Abstract: We evaluated both the photosynthetic plasticity and acclimation to light of seedlings of five co-occurring tropical tree species in the Moraceae,Cecropia obtusifolia, Ficus insipida, Poulsenia armata, Brosimum alicastrum, andPseudolmedia oxyphyllaria. Distinct differences in the species' abilities to respond to increasing irradiance correlated with their known habitat breadths and successional status. The early successinalsCecropia andFicus exhibited the highest photosynthetic rates and conductance values in high light. There was a several-fold difference in assimilation across light regimes, consistent with a high physiological plasticity. When individuals grown at low light were transferred to higher irradiances, seedlings of bothCecropia andFicus produced leaves which photosynthesized at rates as high or higher than those of plants continuously grown in high light, indicating a high photosynthetic acclimation potential. In contrast, the late successionals were characterized by both a more restricted physiological plasticity and acclimation potential. Higher light levels resulted in only moderate increases in assimilation among the late successionals, and onlyBrosimum acclimated fully to increased irradiances. NeitherPoulsenia norPseudolmedia increased appreciably their photosynthetic rates when transferred to high light. This suggests that acclimation potential cannot always be inferred from plasticity responses, and calls for a reevaluation of arguments developed solely from plasticity studies. Finally, differences between the early and late successional species in the allocation of nitrogen into RuBP carboxylase and thylakoid nitrogen pools or non-photosynthetic compounds are suggested by the distinct relationships between maximum photosynthetic capacity and nitrogen content.

Journal ArticleDOI
TL;DR: Virola nobilis (Myristicaceae), a bird/mammal-dispersed tree species in Panama, may also be dispersed by a terrestrial rodent, the agouti (Dasyprocta punctata), which both appears important for tree recruitment and seed dispersal treatment.
Abstract: The data presented show thatVirola nobilis (Myristicaceae), a bird/mammal-dispersed tree species in Panama, may also be dispersed by a terrestrial rodent, the agouti (Dasyprocta punctata). Using a thread-marking method, we observed that agoutis scatterhoardedV. nobilis seeds that they found both singly or in clumps. Seed removal and seed burial rates were strongly affected by features of forest habitats, such asV. nobilis tree richness (rich vs poor) and/or forest age (old vs young), but not by seed dispersal treatment (scattered vs clumped). Predation (mostly post-dispersal) of unburied seeds by weevils was independent of habitat and dispersal treatment. Seeds artificially buried in aVirola-rich area were more likely to escape predation and become established than unburied seeds under natural conditions. The food reward for agoutis is in the germinating seedlings. The seed dispersal syndrome ofV. nobilis involves long- and short-distance dispersers which both appear important for tree recruitment.

Journal ArticleDOI
TL;DR: Comparisons suggested that species active only during the initial, less stressful months of the growing season used water less efficiently, and that physiological responses increasing the ratio of carbon fixed to water lost were common in these grassland species, and were correlated with the increase in evaporative demand and the decrease in soil moisture.
Abstract: Grassland communities of arid western North America are often characterized by a seasonal increase in ambient temperature and evaporative demand and a corresponding decline in soil moisture availability. As the environment changes, particular species could respond differently, which should be reflected in a number of physiological processes. Carbon isotope discrimination varies during photosynthetic activity as a function of both stomatal aperture and the biochemistry of the fixation process, and provides an integrated measure of plant response to seasonal changes in the environment. We measured the seasonal course of carbon isotope discrimination in 42 grassland species to evaluate changes in gas exchange processes in response to these varying environmental factors. The seasonal courses were then used to identify community-wide patterns associated with life form, with phenology and with differences between grasses and forbs. Significant differences were detected in the following comparisons: (1) Carbon isotope discrimination decreased throughout the growing season; (2) perennial species discriminated less than annual species; (3) grasses discriminated less than forbs; and (4) early flowering species discriminated more than the later flowering ones. These comparisons suggested that (1) species active only during the initial, less stressful months of the growing season used water less efficiently, and (2) that physiological responses increasing the ratio of carbon fixed to water lost were common in these grassland species, and were correlated with the increase in evaporative demand and the decrease in soil moisture.

Journal ArticleDOI
TL;DR: Examination of phenotypic variation among populations, among families, within families, and the homogeneity of variances within families for morphological and life history characters of an alien, self-pollinating annual grass, Bromus tectorum, suggests that families differ in the capacity for phenotypesic response to environmental variation.
Abstract: For Bromus tectorum, an alien annual grass now widespread in western North America, we assessed the sensitivity of the phenotypic expression of populational differences in phenology and demography to variation in plant density. Plants were grown in an unheated glasshouse from seeds collected from six habitat types located along a moisture-temperature gradient. Survival to flowering was high with a mean overall survival of 91%. Survival was highest among plants from the coolest, most mesic site. Time to first flowering decreased at the higher sowing density and was dependent on seed source. Plant dry weight and average seed production was lowest within high density plots; the magnitude of this reduction was significantly dependent on seed source. Individual seed weight was also dependent on the interaction of seed source and sowing density. Seed weight did not vary as much as seed number or plant weight in response to density. Differences among sources in hierarchies of plant size and seed production were detected at low sowing densities. The degree of inequality in the distributions of plant size and seed number increased at high density.

Journal ArticleDOI
TL;DR: Evidence from controlled experiments that long straight thorns deter herbivory by browsers is reported, implying that increased thorn length is an induced defense.
Abstract: We report evidence from controlled experiments that long straight thorns deter herbivory by browsers. Cut branches of three woody species that had their thorns removed suffered significantly greater herbivory by a tethered goat than did paired intact branches. Branches on living Acacia seyal plants that had their thorns removed suffered significantly greater herbivory by a wild population of free-ranging giraffes than did intact branches on the same plants. These differences in herbivory resulted in long term losses of branch length in clipped as opposed to control branches. In addition, branches within reach of giraffes produced longer thorns and a greater density of thorns than did higher branches. These results imply that increased thorn length is an induced defense.

Journal ArticleDOI
TL;DR: N2 fixation was associated with reduced intrinsic water use efficiency and higher δ15N-values of Mimosaeae are associated with lower carbon isotope ratios (δ13C value), while the opposite trends were found in non-Mimosaceae, in which N-concentration increased with δ 15N, but δ12C was unaffected.
Abstract: Nitrogen (N2) fixation was estimated along an aridity gradient in Namibia from the natural abundance of 15N (δ15N value) in 11 woody species of the Mimosacease which were compared with the δ15N values in 11 woody non-Mimosaceae Averaging all species and habitats the calculated contribution of N2 fixation (N f ) to leaf nitrogen (N) concentration of Mimosaceae averaged about 30%, with large variation between and within species While in Acacia albida N f was only 2%, it was 49% in Acacia hereroensis and Dichrostachys cinerea, and reached 71% in Acacia melifera In the majority of species N f was 10–30% There was a marked variation in background δ15N values along the aridity gradient, with the highest δ15N values in the lowland savanna The difference between δ15N values of Mimosaceae and non-Mimosaceae, which is assumed to result mainly from N2 fixation, was also largest in the lowland savanna Variations in δ15N of Mimosaceae did not affect N concentrations, but higher δ15N-values of Mimosaeae are associated with lower carbon isotope ratios (δ13C value) N2 fixation was associated with reduced intrinsic water use efficiency The opposite trends were found in non-Mimosaceae, in which N-concentration increased with δ15N, but δ13C was unaffected The large variation among species and sites is discussed

Journal ArticleDOI
TL;DR: Investigation of photosynthetic and water relations characteristics of small juvenile and large reproductive plants were investigated during one growing season for four woody species native to Red Butte Canyon, Utah, USA, finding differences between juvenile and reproductive classes.
Abstract: Photosynthetic and water relations characteristics of small juvenile and large reproductive plants were investigated during one growing season for four woody species native to Red Butte Canyon, Utah, USA: Acer negundo, Artemisia tridentata, Chrysothamnus nauseosus, and Salix exigua. For all species, juvenile plants differed from reproductive plants in at least one of the following characters: water potential, stomatal conductance, photosynthetic rate, or water-use efficiency. Late in the growing season, mortality occurred within juvenile plants (apparently due to a lack of water), but not within reproductive plants. The observed differences between juvenile and reproductive classes are discussed in terms of environment, development, and mortality selection.

Journal ArticleDOI
TL;DR: Field experiments withMercenaria mercenaria in a relatively high-energy environment demonstrated that clams on unvegetated sand flats failed to grow during autumn while those within seagrass beds grew substantially, suggesting that sand flats, where predation rates are higher, may be sites of lower clam growth than seagRass beds because of greater consumer interference with clam feeding.
Abstract: Field experiments withMercenaria mercenaria in a relatively high-energy environment demonstrated that clams on unvegetated sand flats failed to grow during autumn while those within seagrass beds grew substantially. Clam growth rates at the seagrass margin that first receives the faster-flowing, flood-tidal currents were about 25% less than at the opposite edge. In a second experiment, pruning, which reduced average blade length by 50–75%, was shown to enhance near-bottom current velocities and to reduce shell growth ofMercenaria during summer by about 50%. As in the first experiment, clams in the unvegetated sand flats exhibited no net growth. Clam mortality, caused mostly by predatory crabs and whelks, was much higher on sand flats than in seagrass beds and intermediate in clipped seagrass. Although consistent with some previous reports, these growth results are still surprising given that they contradict the generalization that suspension feeders grow faster under more rapid current regimes. Three types of indirect interactions might explain the observed effect of seagrass on growth of buried clams: (1) altering food supply; (2) changing the intensity of biological disturbance on feeding clams; and/or (3) affecting the physical stability of the sediments. Previous research on this question has focused almost exclusively on processes that alter food supply rates. In this study, food concentrations, as indicated by suspended chla, were 30% higher inside than outside one seagrass bed, whereas chla concentrations in two other beds were not different from those on adjacent sand flats. This result is sufficient to show that more intense food depletion was not induced by the reduction in flow velocities under the seagrass canopy. Nevertheless, the possible small difference in food concentrations between vegetated and unvegetated bottom seems insufficient to explain the absence of growth of sand-flat clams, especially given the virtual lack of food limitation among suspension feeders in this system. Two data sets demonstrated that the effects of biological disturbance agents cannot be ignored. An outdoor laboratory experiment showed that even in the absence of physical contact between predator and prey the presence of a whelk reduces the amount of time spent feeding byMercenaria. This result suggests that sand flats, where predation rates are higher, may be sites of lower clam growth than seagrass beds because of greater consumer interference with clam feeding. Furthermore, clam siphons are proportionately larger inside seagrass than on sand flats, implying that siphon nipping may not be as intense inside seagrass. This process, too, would reduce net growth of sand-flat clams. Finally, no explicit test was conducted of the hypothesis that enhanced sediment transport in the absence of flow baffling and root binding by seagrass inhibits net growth of clams on high-energy sand flats. Nevertheless, this is a reasonable explanation for the pattern of enhanced growth of seagrass clams, and could serve to explain the otherwise unexplained pattern of lower clam growth at the edge of the seagrass bed that experiences the faster flood-tidal current velocities. Each broad process, changing fluid dynamics, altering consumer access, and varying sediment stability, represents a mechanism whereby habitat structure, provided by the dominant plant, has an important indirect influence on the functional value of the habitat for resident animals.

Journal ArticleDOI
TL;DR: In this paper, a positive correlation was observed between the amount of phenotypic variation and population size in two perennial, outbreeding plant species, Salvia pratensis and Scabiosa columbaria.
Abstract: The amount of genetic variation within a population is, among other things, related to population size. In small populations loss of genetic variation due to high levels of genetic drift and inbreeding may result in decline of individual fitness and increase the chance of population extinction. This chain of processes is known as genetic erosion. In this study we tested the genetic erosion hypothesis by investigating the relation between morphological variation and population size in two perennial, outbreeding plant species, Salvia pratensis and Scabiosa columbaria. To relate phenotypic variation to genetic variation the experiments were performed under common environmental conditions. For both species a positive correlation was observed between the amount of phenotypic variation and population size (Salvia r=0.915; Scabiosa r=0.703). Part of this variation is likely to have a genetic base, although maternal effects were present in the seedling and juvenile life stages. Differences between populations could in both species be attributed to parameters related to fitness, i.e. growth rate in Salvia and reproductive effort in Scabiosa. Discriminant functions reflecting these parameters did not however discriminate between large and small populations.Results are discussed in relation to the common environment approach and to electrophoretic results obtained earlier (Van Treuren et al. 1991).

Journal ArticleDOI
TL;DR: Increases in the apparent quantum efficiency (AQE) for all C3 species suggest that elevated CO2 may increase photosynthetic rate relative to ambient CO2 over a wide range of light conditions.
Abstract: Seedlings of nine tropical species varying in growth and carbon metabolism were exposed to twice the current atmospheric level of CO2 for a 3 month period on Barro Colorado Island, Panama. A doubling of the CO2 concentration resulted in increases in photosynthesis and greater water use efficiency (WUE) for all species possessing C3 metabolism, when compared to the ambient condition. No desensitization of photosynthesis to increased CO2 was observed during the 3 month period. Significant increases in total plant dry weight were also noted for 4 out of the 5 C3 species tested and in one CAM species, Aechmea magdalenae at high CO2. In contrast, no significant increases in either photosynthesis or total plant dry weight were noted for the C4 grass, Paspallum conjugatum. Increases in the apparent quantum efficiency (AQE) for all C3 species suggest that elevated CO2 may increase photosynthetic rate relative to ambient CO2 over a wide range of light conditions. The response of CO2 assimilation to internal Ci suggested a reduction in either the RuBP and/or Pi regeneration limitation with long term exposure to elevated CO2. This experiment suggests that: (1) a global rise in CO2 may have significant effects on photosynthesis and productivity in a wide variety of tropical species, and (2) increases in productivity and photosynthesis may be related to physiological adaptation(s) to increased CO2.

Journal ArticleDOI
TL;DR: Environmentally-based maternal effects can significantly influence the expression of offspring dispersal potential, growth rate, and offspring fecundity and contribute to natality and survival in natural populations and, hence, to population growth potential.
Abstract: The nutritional environment of the parental generation of the polyphagous gypsy moth, Lymantria dispar, can significantly influence the growth and reproductive potential of the next generation through environmentally-based maternal effects. In the first experiment, members of the parental generation were reared on red oak trees (Quercus rubra L.) with known defoliation and phenolic levels. Diet in the offspring generation was homogeneous (synthetic diet). With genetic effects accounted for 1) offspring attained greater pupal weights when their mothers fed on trees with higher leaf damage levels, 2) daughters had a shorter prefeeding stage, a trait associated with dispersal tendency, when their mothers experienced higher condensed tannin levels, and 3) sons had lower pupal weights when their mothers experienced greater condensed tannin levels. In the second experiment, members of the parental generation were reared on either red or black oak (Q. velutina) trees. Offspring of each mother were divided among four diets: red oak, chestnut oak (Q. prinus L.), a standard synthetic diet, and a low-protein synthetic diet. The parental host species accounted for 24% of the variation in daughters' development time; offspring diet accounted for 52%. When mothers were reared on black oak rather than red oak, their offspring developed significantly faster when the F1 diet was chestnut oak. Environmentally-based maternal effects can significantly influence the expression of offspring dispersal potential, growth rate, and offspring fecundity. These traits contribute to natality and survival in natural populations and, hence, to population growth potential. Theoretical models of insect population dynamics demonstrate that the presence of a time delay in a density dependent response can induce destabilization. Maternal effects act on a time delay and may participate in the destabilization characteristic of outbreak species.

Journal ArticleDOI
TL;DR: The high quality of Puccinellia allowed geese to gain mass in spring, but the metabolizable energy of this plant species declined during the staging period, and Plantago and Triglochin increased in importance in supplying the geese with components with which to build their body reserves.
Abstract: The diet composition of Brent Geese Branta bernicla on a salt-marsh was quantified. Puccinellia maritima was the principal food species, while Plantago maritima and Triglochin maritima were less commonly taken. Festuca rubra only acted as a substitute for Puccinellia when production of the latter species dropped. The metabolizable energy of the food plants ranged from 5 to 11 kJ·g-1. By assessing the ingestion rates of geese feeding on different food species, the net intake rate could be derived. Plantago and Triglochin appeared to be the most profitable plants to eat. The proportion of these species in the diet was restricted by (1) the capacity of the alimentary tract, since high intake rates combined with high water contents of the food plants easily led to overfill; and (2) the limited distribution of these plants, in combination with their rapid depletion by grazing geese. These latter factors led to an unequal allocation among individual geese. Most Plantago and Triglochin was obtained by dominant pairs within the flocks. The high quality of Puccinellia allowed geese to gain mass in spring, but the metabolizable energy of this plant species declined during the staging period, and Plantago and Triglochin increased in importance in supplying the geese with components with which to build their body reserves. The timing of the onset of spring growth of the various food species differed between years, and plant phenology was shown to have a profound effect on the final body reserves of the geese.

Journal ArticleDOI
TL;DR: The strong negative relationship between the occurrence of fish and the egg predators suggests the surprising conclusion that the presence of fish actually protects aquatic anuran eggs from predation in this tropical system, and allows aquatic oviposition to dominate only in those waterbodies with moderate to high densities of fish.
Abstract: Terrestrial oviposition with free-living aquatic larvae is a common reproductive mode used by amphibians within the central Amazonian rainforest. We investigated the factors presently associated with diversity of microhabitats (waterbodies) that may be maintaining the diversity of reproductive modes. In particular, desiccation, predation by fish, competition with other anurans and water quality were examined in 11 waterbodies as possible forces leading to the evolution of terrestrial oviposition. Predation experiments demonstrated that fish generally do not eat anuran eggs, and that predacious tadpoles and dytiscid beetle larvae are voracious predators of anuran eggs. The percentage of species with terrestrial oviposition was only weakly correlated with the occurrence of pond drying, pH and oxygen concentration, suggesting that anurans in this tropical community are able to use the range of water quality available for egg development. There was a tendency for terrestrial oviposition to be associated with the number of species of tadpoles using the waterbody, but we consider this to be spurious as there was no obvious competitive mechanism that could result in this relationship. The percentage of species with terrestrial oviposition was significantly positively related to our index of egg predation pressure, and negatively related to our index of fish biomass. Egg predation pressure was also negatively related to the index of fish biomass. These results allow us to discount as improbable the hypothesis that predation by fish on anuran eggs was an important selective pressure leading to terrestrial oviposition in this community. The strong positive relationship between terrestrial oviposition and our index of egg predation pressure indicates that these predators have exerted, and are exerting, a significant selective pressure for terrestrial oviposition. The strong negative relationship between the occurrence of fish and the egg predators suggests the surprising conclusion that the presence of fish actually protects aquatic anuran eggs from predation in this tropical system, and allows aquatic oviposition to dominate only in those waterbodies with moderate to high densities of fish. Our results suggest that terrestrial oviposition is a “fixed predator avoidance” trait.

Journal ArticleDOI
TL;DR: In this paper, the authors investigated the differences in feeding ecology of the blue tit between the mainland and the island of Corsica and found that the food items brought to the nestlings were much more diverse on Corsica than on the mainland, including many fewer caterpillars and a wider range of taxa.
Abstract: High variation in laying date and clutch size of the blue tit between a Mediterranean mixed habitat on the mainland, southern France, and a sclerophyllous habitat on the island of Corsica is hypothesized to be related to differences in the food supply. The diet of the nestlings and feeding frequencies were studied using camera nestboxes and electronic chronographs. Food items brought to the nestlings were much more diverse on Corsica than on the mainland, including many fewer caterpillars and a wider range of taxa. However, when expressed as a volume index, prey items were on average larger on Corsica than on the mainland. Feeding frequencies were significantly lower on Corsica. A good correlation was found in both habitats between laying date and the caterpillar peak date, although both the leafing development of oaks and the peak of abundance of caterpillars occurred 3 weeks later in the Corsican sclerophyllous trees than in the mainland deciduous ones. Differences in the feeding ecology of tits between the two habitats are discussed in the light of the evergreen habit, which means that only 30% of leaves are available for phyllophagous insects instead of 100% in deciduous trees. the combination of a late and low food supply in evergreen trees is the best explanation for the differences in breeding traits betwen the two populations.

Journal ArticleDOI
TL;DR: Home areas of adults of red-backed salamanders were significantly more segregated intrasexually and more aggregated intersexually than would be expected from a random distribution and intersexual overlap of home areas was significantly greater than intrasexual home area overlap.
Abstract: I used a mark-recapture study to estimate home areas for 107 red-backed salamanders (Plethodon cinereus) in a natural forest habitat. Both males and females of this species defend feeding territories, but I presume that some individuals in this relatively highdensity population (approximately 2.8 salamanders per m2) are nonterritorial floaters. Although territorial salamanders exhibited greater numbers of tail autotomies, they had significantly longer relative tail lengths. This difference suggests that territorial individuals gain benefits from territorial ownership. From the observation that home area size was inversely correlated with body size, I infer that larger animals gained higher quality foraging areas. Home areas of adults were significantly more segregated intrasexually and more aggregated intersexually than would be expected from a random distribution. Furthermore, intersexual overlap of home areas was significantly greater than intrasexual home area overlap. Territorial defense of feeding areas by male and female red-backed salamanders therefore also may play a role in mating behavior.

Journal ArticleDOI
TL;DR: The amount of food that polar bears consume from terrestrial food webs appears negligible, even though some bears spend 1/3 or more of each year on land during the seasons of greatest primary productivity.
Abstract: The mean stable-carbon isotope ratios (δ13C) for polar bear (Ursus maritimus) tissues (bone collagen −15.7‰, muscle −17.7‰, fat −24.7‰) were close to those of the same tissues from ringed seals (Phoca hispida) (−16.2‰, −18.1‰, and −26.1‰, respectively), which feed exclusively from the marine food chain. The δ13C values for 4 species of fruits to which polar bears have access when on land in summer ranged from −27.8 to −26.2‰, typical of terrestrial plants in the Arctic. An animal's δ13C signature reflects closely the δ13C signature of it's food. Accordingly, the amount of food that polar bears consume from terrestrial food webs appears negligible, even though some bears spend 1/3 or more of each year on land during the seasons of greatest primary productivity.