Female-biased density-dependent dispersal of a tephritid fly in a fragmented habitat and its implications for population regulation

The aim of this study was to quantify the rate of dispersal as a response to density in the specialist tephritid fly Paroxyna plantaginis (the main seed predator on its patchily distributed host plant, Tripolium vulgare, Asteraceae). Marked flies were released at three different fly densities in artificial host patches. The individual histories of recaptures were recorded as well as migration between patches and invasion by unmarked flies. The loss of marked flies relative to initial density was analysed using maximum likelihood estimation. Females generally had the highest loss rate. When comparing a density-independent model with a density-dependent model of the loss rate, the density-dependent model won four times out of six for the females but not a single time for the males. A stronger immigration rate of females relative to males supported the suggested female-biased dispersal. This indicates a sit-and-wait strategy for the territorial males and a pre-emptive competition strategy for egg-laying substrates for the females. These results may be of general importance for non-frugivorous tephritid systems with unpredictable and almost ephemeral accessibility to host plants and with a dynamics characterised by a high turnover rate and high attack levels. The study presents a method for measuring the propensity of individuals to leave an area as a response to local density. It is further an example of the consequences individual behavioural responses may have on the population dynamics of a patchy population.     

The role of habitat selection behavior in population dynamics: source–sink systems and ecological traps

Ecological traps, poor-quality habitat that nonetheless attract individuals, have been observed in both natural and human-altered settings. Until recently, ecological traps were considered a kind of source–sink system, but source–sink theory does not model maladaptive habitat choice, and therefore cannot accurately represent ecological traps or predict their population-level consequences. Although recent models of ecological traps addressed this problem, they used patch-based models containing only two habitats that were very different from one another, but were internally homogeneous. These sorts of patch models may not apply to many real populations, and using them for populations in landscapes with mosaic or gradient habitat structures may be misleading. I developed models that treat source–sink dynamics and ecological traps as special cases of a single process, in which the attractiveness and quality of the habitat are separate variables that can be either positively or negatively related, and in which habitat quality varies continuously throughout the landscape. As expected, sinks are less detrimental to populations than ecological traps, in which preferential use of poor habitat elevates extinction risk. Furthermore, ecological traps may be undetected, and may even appear to be sources, when population sizes are large, but may still prevent recovery in spite of the availability of high-quality habitat when populations drop below threshold levels. Conservation biologists do not routinely consider the possibility that apparent sinks are actually traps, but since traps should be associated with the rapidly changing and novel habitat characteristics primarily produced by human activities, ecological traps should be considered an important and potentially widespread conservation concern.     

The importance of habitat and life history to extinction risk in sharks, skates, rays and chimaeras

We compared life-history traits and extinction risk of chondrichthyans (sharks, rays and chimaeras), a group of high conservation concern, from the three major marine habitats (continental shelves, open ocean and deep sea), controlling for phylogenetic correlation. Deep-water chondrichthyans had a higher age at maturity and longevity, and a lower growth completion rate than shallow-water species. The average fishing mortality needed to drive a deep-water chondrichthyan species to extinction (Fextinct) was 38-58% of that estimated for oceanic and continental shelf species, respectively. Mean values of Fextinct were 0.149, 0.250 and 0.368 for deep-water, oceanic and continental shelf species, respectively. Reproductive mode was an important determinant of extinction risk, while body size had a weak effect on extinction risk. As extinction risk was highly correlated with phylogeny, the loss of species will be accompanied by a loss of phylogenetic diversity. Conservation priority should not be restricted to large species, as is usually suggested, since many small species, like those inhabiting the deep ocean, are also highly vulnerable to extinction. Fishing mortality of deep-water chondrichthyans already exploited should be minimized, and new deep-water fisheries affecting chondrichthyans should be prevented.     

Growth, survival and fluctuating asymmetry of Iceland scallops in a test of density-dependent growth in a natural bed

We report an experiment designed to test for overpopulation in an Iceland scallop, Chlamys islandica (O.F. Müller), slow-growing bed. Scallops were installed in pearl nets at the site of the scallop bed and in a nearby unpopulated site, near the bottom and far from the bottom. With this set-up, all groups, except that one near the bottom inside the bed, were free from potential overpopulation effects. In addition, there were two stocking densities to test for containment effects.Shell growth was higher at 2.0 m above the bottom, outside the scallop bed, than at the three other site and height combinations, which were not significantly different from each other. There was a borderline effect of site on soft tissue growth, which was probably slower at the donor site. We conclude that there was no evidence of overpopulation in the scallop bed. On both sites, soft tissue growth was slowest near the bottom, irrespective of group size, intermediate at 2.0 m height, high population density, and fastest at 2.0 m height, low population density. Survivorship was lower near the bottom, but was independent of site and of group size. Available data showed no sustained vertical patterns in phytoplankton concentration in the water column. These results suggest that food depletion occurred in the pearl nets, but that density-independent factors dominated density-dependent factors in the pearl nets near the bottom. Likely explanations involve the interaction between containment effects and current speed variations in the benthic boundary layer. Survivorship decreased with fluctuating asymmetry of the ears of the shells. Fluctuating asymmetry, therefore, may provide a means of including individual effects in growth experiments and forecasting the ability of spat to resist density-independent mortality agents.     

Opportunistic feeding by lions: non-preferred prey comprise an important part of lion diets in a habitat where preferred prey are abundant

Optimal foraging theory predicts less diverse predator diets with a greater availability of preferred prey. This narrow diet niche should then be dominated by preferred prey, with implications for predator–prey dynamics and prey population ecology. We investigated lion (Panthera leo) diets in Hluhluwe–iMfolozi Park (HiP), South Africa, to assess whether lions in a site with a high density of preferred prey (prey species weighing 92–632 kg as estimated from a published meta-analysis) have a narrow diet, consisting primarily of preferred prey. HiP is a useful study site to investigate this prediction because it is a productive landscape (with a high density of prey) where lion-preferred prey constitutes up to 33% of the prey available to lions. Furthermore, to investigate whether lions in HiP exhibit sex-specific diets as documented in other southern African populations, we estimated male and female lion diets separately. We were specifically interested in testing whether traditional approaches of estimating lion diets at the population level mask sex-specific predation patterns, with possible implications for management of lions in small to medium-sized fenced reserves. Lions in HiP preferred larger prey species (63–684 kg) and had diets with a larger proportion of preferred prey than reported in an African-wide meta-analysis. However, despite the high density of preferred prey species, 36% of lion diets still consisted of typically non-preferred species such as nyala (Tragelaphus angasii). This finding suggests that lions in HiP maintain a degree of opportunism even when preferred prey are abundant. Therefore, abundant, non-preferred prey are likely to be an important resource for lion populations. Sex-specific differences in lion diets were evident in HiP, suggesting that estimation of lion resource use and carrying capacity should consider opportunistic hunting and sex-specific differences in lion diets.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Persistence and extinction of a population in a polluted environment

Models of the persistence and extinction of a population or community in a polluted environment have been investigated in several papers. But all of those papers have a basic assumption that the capacity of the environment is so large that the change of toxicants in the environment that comes from uptake and egestion by the organisms can be neglected. This assumption is not made in this article. Some sufficient conditions on persistence or extinction of a population have been obtained, and the threshold between the two has also been obtained for most situations.     

Predation by the salt marsh killifish Fundulus heteroclitus (L.) in relation to prey size and habitat structure: Consequences for prey distribution and abundance

Laboratory feeding preference experiments show that the maximum size of prey eaten (the snail Melampus bidentatus (Say) and the amphipod Orchestia grillus (Bosc)) increases with increasing size of the predator, Fundulus heteroclitus (L.). Melampus > 7 mm in shell height escape predation by even the largest killifish. In the laboratory, consumption of prey is reduced in high marsh habitat relative to low marsh, particularly in the case of larger fish. Low marsh has few grass stems per unit area, while high marsh is considerably more complex, with dense small stems providing cover for prey and reducing successful fish hunting. The population of Melampus in low marsh within Great Sippewissett salt marsh consists mainly of large snails but this size is rare in high marsh. The size-distribution is inverse for Orchestia, with large amphipods more abundant in high marsh. The construction of fences excluding Fundulus from the marsh surface led to low marsh size-distributions of Melampus and Orchestia resembling those of high marsh, in agreement with the laboratory results. Killifish predation seems to be an important factor regulating the abundance and size-distribution of the two prey species in the two marsh habitats.     

Predation by xanthid crabs on early post-settlement gastropods: the role of prey size, prey density, and habitat complexity

Small predators in marine benthic communities create a hazardous environment for newly settled invertebrates, especially for the smallest individuals. To explore the effects of predation on a newly settled gastropod, queen conch (Strombus gigas Linnaeus), by a xanthid crab (Micropanope sp.), prey size, prey density, and habitat complexity were manipulated in five laboratory experiments. All crabs >3.1 mm CW killed all conch <2 mm SL when individual crabs (<14 mm carapace width (CW)) were offered individual conch that were 2–35 days old after metamorphosis (1.2–8.8 mm shell length (SL)). Only 10% of the crabs >5.0 mm CW, however, killed conch that were >5.0 mm SL, suggesting that conch may reach a size refuge from xanthid crabs at 5 mm SL. Furthermore, when given a choice, crabs (4.8 mm CW) preferred smaller conch (2.0 mm SL) to larger (3.7 mm SL), suggesting that 1 week of additional growth in shell length is advantageous to survivorship. Proportional mortality decreased as conch density increased when crabs were offered conch at seven different densities (two to 96 individuals). Crabs proved to be effective predators regardless of the amount of seagrass structure provided in a microcosm experiment, and could consume two conch in 10 s. The high densities of xanthid crabs that occur in the wild, their effectiveness as predators, and their large appetites point to the important role that small predators may potentially play in structuring the population dynamics of their small prey immediately after settlement.     

Dispersal capability in a habitat specialist bush cricket: the role of population density and habitat moisture

1. In fragmented landscapes many insect species depend on a regular exchange of individuals between subpopulations to ensure the persistence of the population. Thus, the ability to disperse is of particular relevance. 2. However, in some insect species mobility is not a fixed trait. Hence, knowing the causes of phenotypic plasticity is of great importance when evaluating whether a species is able to survive in fragmented landscapes or not. 3. A multi‐year field study was conducted to identify possible causes of macroptery in the wing‐dimorphic habitat specialist Metrioptera brachyptera L. and to quantify its dispersal capability (% macropters). Therefore, 746 individuals of the species were caught on 135 plots. Additionally, environmental variables that possibly induce the development of macropters (population density and habitat moisture) were recorded. 4. Dispersal capability of M. brachyptera was very low. Less than 3% were long‐winged. The statistical analysis revealed that the proportion of long‐winged M. brachyptera was strongly correlated with high bush‐cricket densities and not with habitat moisture. 5. The low dispersal capability of M. brachyptera leads to the conclusion that individual exchange between isolated populations is limited or even impossible. Habitat specialists, like M. brachyptera, may thus be unable to respond to rapid changes in the availability of suitable habitats by dispersing, and hence may be especially dependent on habitat management activities that promote the long‐term stability of existing habitat patches.     

Patterns and causes of demographic variation in a harvested moose population: evidence for the effects of climate and density-dependent drivers

1.?Better understanding of the mechanisms affecting demographic variation in ungulate populations is needed to support sustainable management of harvested populations. While studies of moose Alces alces L. populations have previously explored temporal variation in demographic processes, managers responsible for populations that span large heterogeneous landscapes would benefit from an understanding of how demography varies across biogeographical gradients in climate and other population drivers. Evidence of thresholds in population response to manageable and un-manageable drivers could aid resource managers in identifying limits to the magnitude of sustainable change. 2.?Generalized additive models (GAMs) were used to evaluate the relative importance of population density, habitat abundance, summer and winter climatic conditions, primary production, and harvest intensity in explaining spatial variation in moose vital rates in Ontario, Canada. Tree regression was used to test for thresholds in the magnitudes of environmental predictor variables that significantly affected population vital rates. 3.?Moose population growth rate was negatively related to moose density and positively related to the abundance of mixed deciduous habitat abundant in forage. Calf recruitment was negatively related to a later start of the growing season and calf harvest. The ratio of bulls to cows was related to male harvest and hunter access, and thresholds were evident in predictor variables for all vital rate models. 4.?Findings indicate that the contributions of density-dependent and independent factors can vary depending on the scale of population process. The importance of density dependence and habitat supply to low-density ungulate populations was evident, and management strategies for ungulates may be improved by explicitly linking forest management and harvest. Findings emphasize the importance of considering summer climatic influences to ungulate populations, as recruitment in moose was more sensitive to the timing of vegetation green-up than winter severity. The efficacy of management decisions for harvested ungulates may require regional shifts in targets where populations span bioclimatic gradients. The use of GAMs in combination with recursive partitioning was demonstrated to be an informative analytical framework that captured nonlinear relationships common in natural processes and thresholds that are relevant to population management in diverse systems.     

Evolution of rotifer population dynamics in a heterogeneous habitat: mathematical modeling

We present the results of mathematical modeling of a rotifer species inhabiting two coupled habitats with different environmental conditions. We use the modified Consensus model and show that the exchange between the habitats can lead to chaotization of originally regular plankton dynamics and synchronization of plankton biomass oscillations. As a result, the invasion of a chaotic regime takes place.     

Genetic differences in fruit-set patterns are determined by differences in fruit sink strength and a source : sink threshold for fruit set

Background and Aims

Fruit set in indeterminate plant species largely depends on the balance between source and sink strength. Plants of these species show fluctuations in fruit set during the growing season. It was tested whether differences in fruit sink strength among the cultivars explained the differences in fruit-set patterns.

Methods

Capsicum was chosen as a model plant. Six cultivars with differences in fruit set, fruit size and plant growth were evaluated in a greenhouse experiment. Fruit-set patterns, generative and vegetative sink strength, source strength and the source : sink ratio at fruit set were determined. Sink strength was quantified as potential growth rate. Fruit set was related to total fruit sink strength and the source : sink ratio. The effect of differences observed in above-mentioned parameters on fruit-set patterns was examined using a simple simulation model.

Key Results

Sink strengths of individual fruits differed greatly among cultivars. Week-to-week fruit set in large-fruited cultivars fluctuated due to large fluctuations in total fruit sink strength, but in small-fruited cultivars, total fruit sink strength and fruit set were relatively constant. Large variations in week-to-week fruit set were correlated with a low fruit-set percentage. The source : sink threshold for fruit set was higher in large-fruited cultivars. Simulations showed that within the range of parameter values found in the experiment, fruit sink strength and source : sink threshold for fruit set had the largest impact on fruit set: an increase in these parameters decreased the average percentage fruit set and increased variation in weekly fruit set. Both were needed to explain the fruit-set patterns observed. The differences observed in the other parameters (e.g. source strength) had a lower effect on fruit set.

Conclusions

Both individual fruit sink strength and the source : sink threshold for fruit set were needed to explain the differences observed between fruit-set patterns of the six cultivars.     

Demographic,mechanistic and density-dependent determinants of population growth rate: a case study in an avian predator

Identifying the determinants of population growth rate is a central topic in population ecology. Three approaches (demographic, mechanistic and density-dependent) used historically to describe the determinants of population growth rate are here compared and combined for an avian predator, the barn owl (Tyto alba). The owl population remained approximately stable (r approximately 0) throughout the period from 1979 to 1991. There was no evidence of density dependence as assessed by goodness of fit to logistic population growth. The finite (lambda) and instantaneous (r) population growth rates were significantly positively related to food (field vole) availability. The demographic rates, annual adult mortality, juvenile mortality and annual fecundity were reported to be correlated with vole abundance. The best fit (R(2) = 0.82) numerical response of the owl population described a positive effect of food (field voles) and a negative additive effect of owl abundance on r. The numerical response of the barn owl population to food availability was estimated from both census and demographic data, with very similar results. Our analysis shows how the demographic and mechanistic determinants of population growth rate are linked; food availability determines demographic rates, and demographic rates determine population growth rate. The effects of food availability on population growth rate are modified by predator abundance.     

Predation by juvenile Platichthys flesus (L.) on shelled prey species in a bare sand and a drift algae habitat

Due to increasing eutrophication of the coastal Baltic waters, drifting algae are a common phenomenon. Drifting algal mats accumulate on shallow sandy bottoms in late summer and autumn, and affect the ambient fauna. Juvenile flounder, Platichthys flesus, utilize these habitats during their first few years. They feed on benthic meio- and macrofauna; part of their diet consists of shelled species, such as Ostracods, and juvenile Hydrobia spp. and Macoma balthica. Earlier studies have shown that up to 75% of ostracods and 92% of hydrobiids survive the gut passage of juvenile flounder, while all M. balthica are digested by the fish. We conducted laboratory experiments to study how the shelled prey responded to a drift algal mat, and the predation efficiency of juvenile P. flesus on these prey species on bare sand and with drifting algae (50% coverage). Hydrobia spp. utilized the drift algae as a habitat and, after 1 h, 50% had moved into the algae; ostracods and M. balthica were more stationary and, after 96 h, only 23 and 12%, respectively, were found in the algae. For the predation efficiency of P. flesus, a two-way ANOVA with habitat (algae, bare sand) and predation (fish, no fish) as factors revealed that both algae and predation affected negatively the survival of all three prey species. The algae, thus, affected the predation efficiency of juvenile P. flesus and the consumption of prey was much reduced in the algal treatments compared to the bare sand. This was due probably to increased habitat complexity and the ability of prey, especially hydrobiids, to use the algal mat as a refuge. Altered habitat structure due to drift algae, together with the resultant changes in habitat (refuge) value for different prey species, may profoundly change the structure of benthic communities.     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Intraspecific variation in prey quality: a comparison of nutrient presence in prey and nutrient extraction by predators

Prey quality can have large impacts on the survival, growth and behavior of predators. A number of studies have examined how different species of prey vary in quality. However, far less is known about intraspecific variation in the quality of prey for predators and even less about what nutrients are extracted from prey by predators. We examined how the sex, feeding level and developmental status of prey affected the quantities of nutrients present in prey bodies and the quantities of nutrients that could be extracted from prey by spiders. Female and well‐fed prey were larger and had more nutrients than male and food‐limited prey, respectively. After taking into account differences in prey size, spiders extracted relatively more lipid and less protein from female and well‐fed prey than from male and food‐limited prey, respectively. Mealworms were of higher quality than adult mealworm beetles; spiders were able to extract more lipid, protein and other nutrients from larvae than adults. While lipid present in prey was a good predictor of lipid consumed, protein present in prey was not a reliable predictor of protein consumed. The variation in prey quality that we observed within a single species of prey (i.e. well‐fed vs food‐limited crickets) was as large as variation in quality among the three species of prey used in these experiments. Intraspecific variation in prey quality may be an important factor affecting predatory arthropods, especially in habitats or at times of year when one species of prey is abundant. Further studies are needed to examine the consequences of intraspecific variation in prey quality on the life history and behavior of predators.