Allometric scaling of resource acquisition by ruminants in dynamic and heterogeneous environments

We present a mechanistic formulation of the intake response of ruminants to vegetation biomass based solely on physiological and morphological parameters that scale allometrically with the animal's body mass. The model is applied to describe herbivore-vegetation interactions in dynamic and heterogeneous landscapes with low quality but abundant “tall grass” and high quality but sparsely available “short grass”, under two conditions: “uncoupled” (such that the effect of food intake on vegetation biomass can be neglected), or “coupled” (such that the vegetation biomass is determined by herbivore feeding). The results show that under uncoupled conditions, the minimum acceptance (proportion of vegetation consumed by the herbivore) at which the herbivore can leave its current patch without reducing its intake rate is when it has depleted the current patch by the energetic cost required to travel to another patch. The maximum acceptance at which the herbivore should leave its patch is when it has depleted the current patch by the cumulative energetic cost of traveling, handling, cropping, and digesting. Under coupled conditions, the optimal acceptance equals half the relative growth rate of the vegetation. Analytical solutions are obtained for equilibrium values for utilization of the vegetation, and for the densities of vegetation and ruminants, expressed in physiological and morphological herbivore parameters.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Modelling the effects of litter decomposition on tree diversity patterns

Current theories may not fully explain why latitudinal patterns of plant diversity differ between terrestrial and flooded ecosystems. Moreover, the co-occurrence of hyper diverse stands in lowland tierra firma (not inundated) forests and almost monospecific stands in mangroves and gallery riparian vegetation within the tropics remains enigmatic. Building on evidence from ecology and agriculture, we present a new model investigating the hypothesis that, besides the general positive feedback of plant growth by nutrients release, litter decomposition builds up an intra-specific negative feedback functionally linked with tree diversity. The model results were compared with extensive published data sets both across and within latitudinal zones. The model predicts correctly the biomass production and decomposition process, as well as the number of tree species, their relative abundance in all environmental conditions providing a novel, putative explanation also for the diversity variations observed within the tropics. The model demonstrates a possible mechanistic link between the carbon cycle and biodiversity patterns, which is interesting in the debate about advancing in the direction of a unifying ecosystem theory.     

Optimal harvesting of ecologically interdependent fish species

The optimal exploitation of a two-species predator-prey system is considered, using Lotka-Volterra-type equations. Due to the density-dependence of ecological efficiency, both species should be harvested simultaneously over a range of relative prices. Beyond the limits of this price range, either the prey species should be utilized indirectly by harvesting the predator, or the predator should be eliminated to maximize the prey yield. Neglecting harvesting costs, the simultaneous harvest of prey and predators requires that a unit of prey biomass increase in value by being “processed” by predators. Certain results from single-species fishery models are shown not to apply to multispecies models. These are as follows: (i) Optimal regulation of a free access fishery may call for subsidizing instead of taxing the harvest of predator species. (ii) Increasing the discount rate may, at “moderate” levels, imply that the optimal standing stock of biomass increases instead of decreasing. (iii) A rising price or a falling cost per unit fishing effort of a species may raise and not lower the optimal standing stock of that species.     

Ecological and social determinants of spacing behaviour in desert baboon groups

This paper investigates the determinants of individual spacing behaviour in a desert baboon population (Papio cynocephalus ursinus). Patterns of neighbour proximity and neighbour density were examined among adults in four groups under different ecological and social conditions (through instantaneous sampling during focal follows). Initial analysis of these data shows that (1) the use of vertical substrates (refuges such as tall trees and cliff faces) can confound patterns of spacing, and (2) individual differences in spacing can depend on the spatial scale over which it is measured. To minimise these substrate and scale effects, this analysis focuses on animals which are off refuges and examines spacing behaviour through its underlying statistical `dimensions' (identified through factor analysis). Analysis of these dimensions indicates that sex, group size, activity-habitat and female reproductive state can all have independent effects on spacing: (1) males are more dispersed than females in smaller groups, (2) male and female dispersion increases with time spent in foraging habitats, and (3) female dispersion is reduced during lactation. According to the hypotheses tested, these results indicate that feeding competition only affects spacing behaviour during foraging while predation risk plays little or no role in spacing. Most aspects of spacing behaviour are best explained by male reproductive strategies and their social repercussions. Received: 25 May 1998 / Accepted after revision: 18 July 1998     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Local complexity in patterns of canopy–benthos associations produces regional patterns across temperate Australasia

Across subtidal coasts of temperate Australasia, canopy–benthos associations are mostly understood from broadly defined studies of kelp forests within eastern Australia and north-eastern New Zealand. We tested the hypotheses that (1) benthic assemblages differ between monospecific stands of Ecklonia radiata and stands that comprise E. radiata mixed with other canopy-forming species, (2) patterns observed locally (i.e. within sites 1–10 km apart) match those observed among regions (>1,000 km apart) for which (3) eastern Australia and northern New Zealand are representative of other regions of temperate Australasia. Benthic assemblages almost always differed among monospecific, mixed, and open stands indicating that failure to distinguish between superficially similar habitats can lead to over-generalised conclusions about the ecology of kelp forests. Patterns of differences among stands did not change between western and southern Australia but differed from eastern Australia, and patterns from all regions were distinct from New Zealand (WA=SAEANZ). Whilst local patterns were complex, the major morphological groups that often characterise benthos (i.e. encrusting coralline algae and turf-forming algae) revealed patterns that could be related across space from local to regional scales. These findings demonstrate that knowledge about the configuration of canopy-forming species will improve confidence about the representativeness of results and that any local complexity need not impede searches for generality when the spatial limits of patterns are also understood.Communicated by M.S. Johnson, Crawley     

Tidal regime dictates the cascading consumptive and nonconsumptive effects of multiple predators on a marsh plant

Prey perception of predators can dictate how prey behaviorally balance the need to avoid being eaten with the need to consume resources, and this perception and consequent behavior can be strongly influenced by physical processes. Physical factors, however, can also alter the density and diversity of predators that pursue prey. Thus, it remains uncertain to what extent variable risk perception and antipredator behavior vs. variation in predator consumption of prey underlie prey-resource dynamics and give rise to large-scale patterns in natural systems. In an experimental food web where tidal inundation of marsh controls which predators access prey, crab and conch (predators) influenced the survivorship and antipredator behavior of snails (prey) irrespective of whether tidal inundation occurred on a diurnal or mixed semidiurnal schedule. Specifically, cues of either predator caused snails to ascend marsh leaves; snail survivorship was reduced more by unrestrained crabs than by unrestrained conchs; and snail survivorship was lowest with multiple predators than with any single predator despite interference. In contrast to these tidally consistent direct consumptive and nonconsumptive effects, indirect predator effects differed with tidal regime: snail grazing of marsh leaves in the presence of predators increased in the diurnal tide but decreased in the mixed semidiurnal tidal schedule, overwhelming the differences in snail density that resulted from direct predation. In addition, results suggest that snails may increase their foraging to compensate for stress-induced metabolic demand in the presence of predator cues. Patterns from natural marshes spanning a tidal inundation gradient (from diurnal to mixed semidiurnal tides) across 400 km of coastline were consistent with experimental results: despite minimal spatial variation in densities of predators, snails, abiotic stressors, and marsh productivity, snail grazing on marsh plants increased and plant biomass decreased on shorelines exposed to a diurnal tide. Because both the field and experimental results can be explained by tidal-induced variation in risk perception and snail behavior rather than by changes in snail density, this study reinforces the importance of nonconsumptive predator effects in complex natural systems and at large spatial scales.     

Amount or pattern? Grassland responses to the heterogeneity and availability of two key resources

Patterns of resource availability and heterogeneity shape the composition, productivity, and dynamics of plant assemblages in a wide variety of terrestrial ecosystems. Despite this, the responses of plant assemblages to simultaneous changes in the availability and heterogeneity of more than a single resource are virtually unknown. To fill this gap, microcosms consisting of assemblages formed by Lolium perenne, Plantago lanceolata, Anthoxantum odoratum, Holcus lanatus, and Trifolium repens were grown in a factorial experiment with the following treatments: nutrient availability (NA), water availability (WA), spatial nutrient heterogeneity (NH), and temporal water heterogeneity (WH). Assemblages exhibited precise root foraging patterns in response to nutrient heterogeneity, which were modified by NA and WA. A series of two- and three-way interactions involving the four factors evaluated determined biomass production, the belowground: aboveground biomass ratio, the patterns of root biomass allocation with depth, and the relative contribution to aboveground biomass of Lolium and Anthoxanthum. In all cases, these interactions explained significant amounts of the variation found in the data. Our study demonstrates that considering the interactions between resource availability and heterogeneity allows for a refinement of predictions that can detectably reduce the error associated with extrapolating from single factor analyses.     

Acetylene reduction (nitrogen fixation) in a Delaware,USA salt marsh

Acetylene reduction (nitrogen fixation) was measured in several vegetational areas in a Delaware, USA salt marsh. Samples were collected for 1 yr and the results showed a seasonally variable pattern of acetylene reduction at all stations. Peak rates were generally recorded during the later summer and early fall (September–October). The seasonality was influenced mainly, although not exclusively, by the soil temperature. In addition, samples collected in short Spartina alterniflora stands exhibited rates which were up to 20-fold higher than those found in samples from tall S. alterniflora stands. Over 50% of the total yearly ethylene production occurred from mid-August until the beginning of December at the tall and short S. alterniflora stations. Maximum activity occurred at 5 cm depth for all stations. Surface activity accounted for only 3–4% of the total measured in the top 20 cm. Addition of glucose or mannitol resulted in considerable increases in activity, thus suggesting that heterotrophic acetylene reduction is carbon and/or energy limited. The results obtained in this study indicate that the measured rates are only potential rates and that considerable caution must be used in extrapolating from acetylene reduction rates to nitrogen fixation rates in situ.     

Combined effect of fire and water scarcity on vegetation patterns in arid lands

In many arid zones around the word, the vegetation spontaneously forms regular patterns to optimize the use of the scarce water resources. The patterns act as early warning signal that fragile ecosystems may suddenly undergo irreversible shifts, thus, interpreting the structural shape of vegetation patterns is crucial to deciphering the ecosystem history and its expected further development. The sudden and irreversible shift of delicate ecosystems as a consequence of minor variation of the climatic forcing has been studied extensively in the past. The attitude of the ecosystem to recover after a catastrophic event, such as fire, did not receive as much attention so far. Here we modelled fire, as a sudden shift of the ecosystem state variables and functionality and evaluated post-fire scenarios under the hypothesis that two major feedbacks shaped the vegetation patterns: a positive feedback between preferential infiltration and plant growth, and a second feedback between infiltration and vegetation burning. A simple model solving a set of partial differential equations for soil moisture, plant biomass, surface water and dead biomass balance predicted significantly diverse post-fire vegetation patterns depending on the fire severity and on the degree of soil water repellency induced by the vegetation burning.     

Spatial and temporal variability of phytoplankton biomass and taxonomic composition around Elephant Island,Antarctica, during the summers of 1990–1993

During the austral summers of 1990–1993, phytoplankton studies were conducted in the vicinity of Elephant Island, Antarctica, to investigate the spatial and temporal variability of phytoplankton biomass and taxonomic composition. There was much intraannual variability, with a trend for increasing biomass from January–February (Leg I) to February–March (Leg II), except in the 1993 studies. There was also a change in phytoplankton composition between the two legs. During 1990–1991 the increase was due mostly to diatoms, during 1992 mostly to an increase of flagellates; during 1993 there was a decrease in total biomass between the two legs, with diatoms decreasing, so that dinoflagellates, which increased slightly in numbers, dominated the biomass during the second leg. There was also much inter-annual variability, with the summers of 1990–1991 having greater biomass and higher proportions of microplanktonic diatoms than that of 1992–1993, which had a higher proportion of flagellates. Cluster analyses revealed the presence of four major phytoplankton assemblages, with varying geographical distributions. The northwestern portion of the grid (Drake Passage waters), was characterized by nanoplanktonic diatoms during 1990–1991 and 1993, but by nanoplanktonic flagellates during 1992. The central area (Drake-Bransfield confluence) was characterized by microplanktonic diatoms in 1990–1991, but by cryptophytes or flagellates in 1992–1993. The south and southeastern portion of the area (Bransfield Strait waters) was characterized mainly by either cryptophytes or other flagellates during all 4 yr. The spatial and temporal variability of phytoplankton could not be ascribed specifically to the geographical extent of the different water masses found in the study area, but appears to be due to changing growth conditions in the upper water column as influenced by physical mixing and meteorological conditions, as well as to effects of differential grazing.     

Utilisation of a coastal grassland by geese after managed re-alignment

In this study we evaluate the effect of coastal re-alignment on the utilisation of coastal grasslands by staging geese. We assessed vegetation change and utilisation by geese using repeated mapping and regular dropping counts in both the restored marsh and adjacent reference sites. All measurements were started well before the actual re-alignment. In addition, we studied the effects of livestock grazing on vegetation and geese, using exclosures. The vegetation transformed from fresh grassland into salt-marsh vegetation. A relatively large proportion of the de-embanked area became covered with secondary pioneer vegetation, and the overall cover of potential food plants for geese declined. Goose utilisation had initially dropped to low levels, both in autumn and in spring, but it recovered to a level comparable to the reference marsh after ten years. Exclosure experiments revealed that livestock grazing prevented the establishment of closed swards of grass in the poorly drained lower area of the restored marsh, and thereby negatively affected goose utilisation of these areas during spring staging. Goose grazing in the restored marsh during spring showed a positive numerical response to grass cover found during the preceding growing season. (1) The value of restored salt marsh as foraging habitat for geese initially decreased after managed re-alignment but recovered after ten years. (2) Our findings support the idea that the value of foraging habitats depends largely on the cover of forage plants and that this can be manipulated by adjusting both grazing and drainage.     

Evolutionary diversity and spatiotemporal dynamics of a spatial game

In this paper, the evolution of cooperation is studied by a spatially structured evolutionary game model in which the players are located on a two-dimensional square lattice. Each player can choose one of the following strategies: “always defect” (ALLD), “tit-for-tat” (TFT), and “always cooperate” (ALLC). Players merely interact with four immediate neighbors at first and adjust strategies according to their rewards. First, the evolutionary dynamics of the three strategies in non-spatial population is investigated, and the results indicate that cooperation is not favored in most settings without spatial structure. Next, an analytical method, which is based on comparing the local payoff structures, is introduced for the spatial game model. Using the conditions derived from the method as criteria, the parameter plane for two major parameters of the spatial game model is divided and nine representative regions are identified. In each parameter region, a distinct spatiotemporal dynamics is characterized. The spatiotemporal dynamics not only verify that the spatial structure promote the evolution of cooperation but also reveal how cooperation is favored. Our results show that spatial structure is the keystone of the evolution of intraspecific diversity.     

Movement of foraging Tundra Swans explained by spatial pattern in cryptic food densities

We tested whether Tundra Swans use information on the spatial distribution of cryptic food items (below ground Sago pondweed tubers) to shape their movement paths. In a continuous environment, swans create their own food patches by digging craters, which they exploit in several feeding bouts. Series of short (<1 m) intra-patch movements alternate with longer inter-patch movements (>1 m). Tuber biomass densities showed a positive spatial auto-correlation at a short distance (<3 m), but not at a larger distance (3-8 m). Based on the spatial pattern of the food distribution (which is assumed to be pre-harvest information for the swan) and the energy costs and benefits for different food densities at various distances, we calculated the optimal length of an inter-patch movement. A swan that moves to the patch with the highest gain rate was predicted to move to the adjacent patch (at 1 m) if the food density in the current patch had been high (>25 g/m2) and to a more distant patch (at 7-8 m) if the food density in the current patch had been low (<25 g/m2). This prediction was tested by measuring the response of swans to manipulated tuber densities. In accordance with our predictions, swans moved a long distance (>3 m) from a low-density patch and a short distance (<3 m) from a high-density patch. The quantitative agreement between prediction and observation was greater for swans feeding in pairs than for solitary swans. The result of this movement strategy is that swans visit high-density patches at a higher frequency than on offer and, consequently, achieve a 38% higher long-term gain rate. Swans also take advantage of spatial variance in food abundance by regulating the time in patches, staying longer and consuming more food from rich than from poor patches. We can conclude that the shape of the foraging path is a reflection of the spatial pattern in the distribution of tuber densities and can be understood from an optimal foraging perspective.     

Zooplankton biomass and indices of grazing and metabolism during a late winter bloom in subtropical waters

The development of the so-called late winter bloom in subtropical water was studied in an oceanic area north of the Canary Islands from January to May 2000. Zooplankton was sampled at short-term intervals (1–4 days) during the bloom (January–March), and biomass, indices of grazing (gut fluorescence) and metabolism (electron transfer system activity, ETS) were measured in four different size fractions (100–200, 200–500, 500–1000 and >1000 µm). During the bloom, ETS activity and gut fluorescence increased before the development of zooplankton biomass. At the end of February, the presence of an impressive cloud of dust formed in the Sahara desert was related to an increase in chlorophyll and small zooplankton a week later. The increments in biomass were the consequence of consumption by zooplankton as inferred from the indices of grazing and metabolism. Estimated grazing from gut fluorescence and gut evacuation rates during the period of study accounted for 55% of the assessed total ingestion from respiration and normal values of assimilation, showing the importance of the non-pigmented food in the diet of zooplankton in these waters. In contrast, the sharp decreases in zooplankton biomass observed during the bloom appeared during the dark period of the moon, the days in which the diel vertical migrants reach the shallower layers, in agreement with previous works in the area. Thus, the development of the late winter bloom in this region is suggested to be driven by the interplay between resource and consumer controls.Communicated by S.A. Poulet, Roscoff     

Foraging patterns of yellow-bellied marmosts: role of kinship and individual variability

Summary Two colonies of yellow-bellied marmots (Marmota flaviventris) at an elevation of 2900 m in Colorado were studied to elucidate the role of various behavioral and ecological factors as determinants of spatial foraging patterns. The locations of known individuals were periodically recorded. These locality data were plotted as three-dimensional block diagrams, the peak heights representing the frequency of observation. Predation risk and vegetation distribution influenced the location of foraging areas; kinship was an important factor in the determination of the amount of foraging area shared between individual marmots. Spatial overlap tended to be greater among close kin, but this was modified by individual behavioral characteristics, reproductive state, the existence of separate burrow systems within a colony, and the age of the animal. Mothers and juveniles, and littermates as young and resident yearlings, had nearly identical foraging areas.     

Prey-capture benefits in a mixed-species group of Amazonian tamarins,Saguinus fuscicollis and S. mystax

Summary Tamarins of the genus Saguinus feed on a wide range of arthropods and small vertebrates, which compose a critical component of their diet. This paper examines the foraging patterns and capture success of the Avila-Peres saddle-back (S. fuscicollis avilapiresi) and the red-capped moustached tamarin (S. mystax pileatus) in very stable mixed-species groups, and whether and how any foraging benefits for either species resulted from their association. Moustached tamarins actively searched for prey items which were mainly well exposed on the midstorey foliage. Saddle-back tamarins, on the other hand, foraged at lower heights, largely by manipulating a variety of microhabitats potentially concealing embedded prey. The foraging activity of the numerically dominant and larger-bodied moustached tamarins often resulted in prey items escaping to lower substrates, usually the forest leaf-litter. The beating effect of this species substantially facilitated captures of large, mobile prey items by saddle-backs, which were highly adept at locating and retrieving flushed prey. It is estimated that, while saddle-backs obtained 66–73% of their prey biomass from flushed items, this proportion was substantially lower (2–9%) for moustached tamarins. Commensal insectivory appears to involve a highly asymmetric benefit to saddle-backs, and a low cost to moustached tamarins, which partly explains the stability of mixed-species groups. Correspondence to the present address     

Influences on dusky dolphin (<Emphasis Type="Italic">Lagenorhynchus obscurus</Emphasis>) fission-fusion dynamics in Admiralty Bay,New Zealand

In societies characterized by a high degree of fission-fusion dynamics, individuals adjust their grouping patterns according to the shifting balance of costs and benefits associated with grouping. This study examines influences on fission-fusion dynamics for dusky dolphins (Lagenorhynchus obscurus) in Admiralty Bay, New Zealand. This area is an important foraging habitat for dusky dolphins during the winter and spring. Admiralty Bay has little predation risk, but nearshore mussel farms may infringe on available habitat. I used generalized estimating equations to determine the influences of coordinated foraging, predation risk, and presence of mussel farms on party size, rate of fission-fusion, and behavioral state. I conducted 168 boat-based group focal follows totaling 168 h. The proportion of individuals observed foraging was positively related to party size and rate of party fusion. Resting had no effect on party size and did not vary according to location. Near mussel farms, traveling decreased, and rate of party fission decreased. I conclude that (1) coordinated foraging strategies are a primary influence on fission-fusion dynamics within this population, (2) dolphins may respond to decreased predation risk by not adjusting party size or location during resting, and (3) areas near mussel farms are not used for traveling.