The role of periphyton abundance and quality in the microdistribution of a stream grazer, Helicopsyche borealis (Trichoptera: Helicopsychidae)

SUMMARY. 1. A series of laboratory and field experiments was performed to determine if the preference of larvae of Helicopsyche borealis (Trichoptera: Helicopsychidae) for exposed rock surfaces in streams was related to patchiness of periphyton food.
2. At low current velocities in the laboratory larvae preferred food-saturated as opposed to food-depleted portions of substrata and drifted off the substratum in response to low food levels.
3. When offered a choice of three potential foods—a diatom, a green filamentous or a blue-green filamentous alga—larvae showed no preference.
4. Larval densities on artificial subtrata in field experiments were highest in areas of high algal chlorophyll a .
5. In laboratory experiments more adults emerged from cultures containing green algae or diatoms than cultures with blue-green algae or bacteria alone.
6. Periphyton quality was not as important as abundance in controlling H. borealis microdistribution. By selecting exposed surfaces of rocks, H. borealis larvae place themselves in the microhabitat most likely to contain both abundant and high quality food patches.     

Local effects of a sedentary grazer on stream algae

1. Larvae of the sedentary aquatic caterpillar Petrophila confusalis (Walker) construct silken retreats around which they feed; outside these clearly demarcated grazed areas, stream algae are exposed to a variety of mobile grazers. Comparisons of the algal community inside and just outside grazed zones were made for third- and fifth-instar Petrophila in the South Fork of the Eel River, California. 2. Densities of both the filamentous macroalga Cladophora and diatom assemblages were significantly reduced within the grazed areas during both larval instars. Grazing of diatoms was taxonomically non-selective. 3. In spring, the grazed zones were relatively large (mean = 22.7 cm²) and visibly increased epilithic spatial patchiness. 4. Per cent composition of diatom assemblages inside and outside the grazed areas differed during the third instar but not during the fifth instar. During the third instar, the grazed zone contained more Synedra ulna (a common immigrant) and less Gomphonema olivaceum (a late-successional species).     

Dispersal of organisms in a patchy stream environment under different settlement scenarios

1.  Previous work in streams, where many organisms disperse downstream by drifting in the water column, has suggested dispersal distances could be related to the overall proportion of the stream within dead water zones (DWZ), places such as backwaters, pools or behind large obstacles, where the water is out of the main flow and is stored temporarily. However, dispersal distances might also be influenced by the spatial organization of DWZ within a reach; this would require us to distinguish between reaches containing many small DWZ compared with those having a few large ones in order to understand fully differences in dispersal patterns among reaches differing in this regard.
2.  We constructed a spatial model of flow in stream reaches, in which we varied the density and size distribution of DWZ distributed through the reach. We defined flow conditions under which stream organisms, modelled as passive particles, would settle or continue to drift downstream. We then examined the horizontal dispersal profiles of such particles through reaches differing in arrays of DWZ.
3.  Our results support those of previous studies, but suggest that the spatial arrangement of DWZ may be just as important as the overall proportion of DWZ within a reach. On a more general note, the importance of patch configuration observed in this study reflects the growing view that a thorough understanding of population and ecosystem processes will require the explicit consideration of spatial pattern in the environment.     

Spruce budworm impact, abundance and parasitism rate in a patchy landscape

The hypothesis that vegetational diversity may lessen the impact of forest insect pests by favoring natural enemies is appealing to those who seek ecologically sound solutions to pest problems. We investigated the effect of forest diversity on the impact of the spruce budworm Choristoneurafumiferana following the last outbreak, as well as the budworm's current abundance and parasitism rate, in the boreal forest of northwestern Québec. Mortality of balsam fir caused by the budworm was greater in extensive conifer stands than either in “habitat islands” of fir surrounded by deciduous forest or on true islands in the middle of a lake. Adult spruce budworm abundance, assessed by pheromone traps, did not differ significantly between the three types of sites. Larval and pupal parasitism rates were examined by transferring cohorts of laboratory-reared larvae and pupae to trees in the three site types and later collecting and rearing them. The tachinid Actiainterrupta, a parasitoid of fifth and sixth instar larvae, as well as the ichneumonid pupal parasitoids Itoplectesconquisitor, Ephialtesontario and Phaeogenesmaculicornis, caused higher mortality in the habitat islands than on true islands or in extensive stands. Exochusnigripalpistectulum, an ichneumonid that attacks the larvae and emerges from the pupae, caused greater mortality in the extensive stands of conifers. Received: 17 March 1997 / Accepted: 18 November 1997     

Patterns of parasitism by insect parasitoids in patchy environments

Abstract. 1. This paper shows how the different spatial patterns of per cent parasitism in patches of different host density can be explained within a single model framework that takes into account the parasitoid's aggregative response, and the factors limiting the degree of host exploitation within patches.
2. Two contrasting laboratory examples are presented in which the distribution of searching parasitoids and the resulting levels of parasitism in different patches are both known for a range of parasitoid densities.
3. A model is described predicting the number of hosts parasitized per patch, in which the number of parasitoids searching is determined from a simple expression allowing different degrees of aggregation.
4. The model generates patterns of parasitism encompassing the two laboratory examples and a wide range of examples from the field.
5. The importance of density dependent spatial distributions of parasitism to population stability is briefly discussed.     

The dynamics of predation and competition in patchy environments

The model of N. D. Atkinson and B. Shorrocks (J. Anim. Ecol. 50, 461–471 (1981)) as two competing species distributing their progeny amongst patches according to independent negative binomial distributions. The resulting separation of the species increases the likelihood of coexistence. We have assumed a much simpler distribution of the competitors which has enabled us to explore analytically the dynamics of interactions with two competing species and a shared natural enemy in a patchy environment. Two types of natural enemy have been considered: a generalist predator whose dynamics are uncoupled from those of the two prey species, and a specialist (e.g., a parasitoid) whose dynamics are entirely coupled to those of its two prey. The following conclusions emerge. Non-aggregating generalist predators causing random predation across patches are generally destabilizing (although asymmetrical predation may in some case enhance coexistence as a result of preferential predation on the superior competitor). Predator aggregation in patches of high prey density, however, produces a switching effect which tends to promote stability. Coexistence is now even possible with high degrees of correlation in the distribution of the two prey and in situations of extreme competition where the competition coefficients exceed one. The main difference in the models with a specialist parasitoid as the natural enemy is a reduction in stability compared with the equivalent generalist-prey interaction. But stable coexistence can still readily occur if the natural enemies aggregate markedly in patches of high prey density.     

Evolutionary stability of ideal free dispersal strategies in patchy environments

A central question in the study of the evolution of dispersal is what kind of dispersal strategies are evolutionarily stable. Hastings (Theor Pop Biol 24:244-251, 1983) showed that among unconditional dispersal strategies in a spatially heterogeneous but temporally constant environment, the dispersal strategy with no movement is convergent stable. McPeek and Holt's (Am Nat 140:1010-1027, 1992) work suggested that among conditional dispersal strategies in a spatially heterogeneous but temporally constant environment, an ideal free dispersal strategy, which results in the ideal free distribution for a single species at equilibrium, is evolutionarily stable. We use continuous-time and discrete-space models to determine when the dispersal strategy with no movement is evolutionarily stable and when an ideal free dispersal strategy is evolutionarily stable, both in a spatially heterogeneous but temporally constant environment.     

Top-down control in a patchy environment: revisiting the stabilizing role of food-dependent predator dispersal

In this paper, we revisit the stabilizing role that predator dispersal and aggregation have in the top-down regulation of predator-prey systems in a heterogeneous environment. We consider an environment consisting of sites interconnected by dispersal, and propose a novel mechanism of stabilization for the case with a non-sigmoid functional response of predators. We assume that the carrying capacity of the prey is infinitely large in each site, and show that successful top-down regulation of this otherwise globally unstable system is made possible through an interplay between the unevenness of prey fitness across the sites and the rapid food-dependent migration of predators. We argue that this mechanism of stabilization is different from those previously reported in the literature: in particular, it requires a high degree of synchronicity in local oscillations of species densities across the sites. Prey outbreaks take place synchronously, but the unevenness of prey growth rates across the sites results in a pronounced difference in the species densities, and so the predator quickly disperses to the sites with the highest prey abundances. For this reason, the consumption of prey mostly takes place in the sites with high densities of prey, which assures an efficient suppression of outbreaks. Furthermore, when the total size of prey population is low, the distribution of both species among the sites becomes more even, and this prevents overconsumption of the prey by the predator. Finally, we put forward the hypothesis that this mechanism, when considered in a tri-trophic plankton community in the water column, can explain the stability of the nutrient-rich low-chlorophyll open ocean regions.     

The role of food, weather and climate in limiting the abundance of animals

More and more studies are demonstrating that populations of animals ‐ from herbivores to top predators, vertebrates and invertebrates ‐ are limited by their food, and that the availability of this food is dictated by the weather. Satellite monitoring is revealing how cyclic and quasi‐cyclic climatic patterns, like the El Niño Southern Oscillation and the North Atlantic Oscillation, are driving and synchronising these weather‐driven changes in the supplies of food. Changes in the amount of food available operate to limit the abundance of populations largely through their influence on the survival of the very young: the Achilles heel of all populations. Any individual organism struggles to use whatever resources it can get from a mostly inhospitable environment to maximise the proliferation of its genes. Each level of a food chain is thus dependent upon, and pressing hard against the limits set by the one below. The resulting intra‐ and inter‐specific interactions produce a multitude of complex outcomes, that significantly influence the dynamics of populations, but do not determine their ultimate size. There is no density‐dependent regulation of abundance. Intra‐specific competition does not determine the size of populations, it only decides which few individuals gain access to the limited food. Nor do predators regulate their prey. They, too, are limited by their food, and the abundance and quality of food is dictated by the weather.     

The spatial population dynamics of insects exploiting a patchy food resource

The population dynamics of ten species of phytophagous insects and seven parasitoids inhabiting the flowerheads of two herbaceous plants, Centaurea nigra and Arctium minus, were studied, and three main aspects of their ecology were examined, namely, rates of population extinction, density dependence in population changes from one generation to the next, and movements between populations. The study was based on monitoring the insect populations on more than 50 patches of each plant, scattered over 5 km² of arable farmland and the results were used to test the relative importance of immigration and population regulation to the persistence of these populations. This paper describes the study of movements between patches of food plant. Experimentally planted new patches of plants were rapidly colonised by all species and this appeared to be unaffected by distance from a source population, up to the maximum distance of 800 m considered in the experiment. Large patches tended to be colonised more readily than small ones. Movements between plant patches were studied with the use of chemical markers (Rb, Sr, Dy and Cs) which were applied as chloride salts to individual patches, and which were translocated to the flowerheads and so to insects feeding on the seed, and to their parasitoids. Initial experiments in the laboratory showed that these elements could be readily detected by ICP (Inductively Coupled Plasma) mass spectrometry in the bodies of all species reared on potted plants sprayed with solutions containing them. Background levels of strontium were patchily high on the study area, but the other elements were naturally either absent or in very low concentrations. Four patches of each plant were marked with a different element in 1991. In 1992, samples of four species of tephritid fly and two parasitoids were collected from all patches, and analysed for the four elements. These analyses showed that individual of all species moved considerable distances, with movements of up to 2 km being commonly recorded. Estimates of rates of immigration to patches showed that movement plays an important role in the population dynamics of these insects. There was some evidence that immigration was density-dependent: it was highest when the resident populations (numbers per flowerhead) were low.     

Searching efficiency of the lady beetleCoccinella septempunctata larvae in uniform and patchy environments

Experiments were conducted on the searching behavior and searching efficiency of the lady beetleCoccinella septempunctata bruckii Mulsant under conditions of various prey distributions and prey densities. The larvae changed their searching behavior before and after feeding. Before feeding the larvae moved quickly and the searching paths were nearly linear. But after feeding the speed decreased and turning angle increased. The speed and turning angle reverted gradually and recovered the initial pattern 95 s after feeding. The searching efficiency differed depending on the prey distribution. At low prey density, searching was most efficient when prey were distributed uniformly. But at middle and high prey densities, searching was most efficient when prey items were highly aggregated. The observed searching behavior of 4th instarC. septempunctata larvae was likely to be optimal considering the natural distribution of colonies of their prey, aphids.     

Theoretical perspectives on the statics and dynamics of species' borders in patchy environments

Understanding range limits is a fundamental problem in ecology and evolutionary biology. In 1963, Mayr argued that "contaminating" gene flow from central populations constrained adaptation in marginal populations, preventing range expansion, while in 1984, Bradshaw suggested that absence of genetic variation prevented species from occurring everywhere. Understanding stability of range boundaries requires unraveling the interplay of demography, gene flow, and evolution of populations in concrete landscape settings. We walk through a set of interrelated spatial scenarios that illustrate interesting complexities of this interplay. To motivate our individual-based model results, we consider a hypothetical zooplankter in a landscape of discrete water bodies coupled by dispersal. We examine how patterns of dispersal influence adaptation in sink habitats where conditions are outside the species' niche. The likelihood of observing niche evolution (and thus range expansion) over any given timescale depends on (1) the degree of initial maladaptation; (2) pattern (pulsed vs. continuous, uni- vs. bidirectional), timing (juvenile vs. adult), and rate of dispersal (and hence population size); (3) mutation rate; (4) sexuality; and (5) the degree of heterogeneity in the occupied range. We also show how the genetic architecture of polygenic adaptation is influenced by the interplay of selection and dispersal in heterogeneous landscapes.     

Aluminium in aquatic environments: abundance and ecotoxicological impacts

Aquatic Ecology - Aluminium (Al) is a common chemical element released into the aquatic environment from the Earth’s crust and many anthropogenic activities. It may be present in various...     

The effect of branching angle on adaptive growth in patchy environments

In clonal plants, the genetic individual (genet) develops via the production of multiple physiological individuals (ramets). The branching angle between the ramets can significantly influence the shape of the developing clone. We investigated the adaptive significance of this phenomenon by means of a spatially explicit dynamic model of clonal growth. We studied the effect of the branching angle on the efficiency of filling habitat patches, varying the sizes and shapes of the patches. Two growth forms were compared: the Narrow Range (NR) versus Wide Range (WR) form. In the NR plant, the branching angle was always acute, while in the WR plant, both acute and wide angles could occur. We hypothesized that the NR plant would be less successful, as narrower branching constrained the plant’s ability to turn. The simulations revealed an opposite trend: the NR plant occupied more space in most of the simulated habitats. However, the effect was weak in general. We conclude that the branching angle between ramets is likely to be a neutral trait in terms of natural selection.     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

Richness and abundance of stream fish communities in a fragmented neotropical landscape

Environmental Biology of Fishes - Environmental conditions influence ecological processes that shape stream community diversity and abundance. Deforestation has the potential to limit available...     

Cellular automaton models for competition in patchy environments: Facilitation, inhibition, and tolerance

We have developed cellular automaton models for two species competing in a patchy environment. We have modeled three common types of competition: facilitation (in which the winning species can colonize only after the losing species has arrived) inhibition (in which either species is able to prevent the other from colonizing) and tolerance (in which the species most tolerant of reduced resource levels wins). The state of a patch is defined by the presence or absence of each species. State transition probabilities are determined by rates of disturbance, competitive exclusion, and colonization. Colonization is restricted to neighboring patches. In all three models, disturbance permits regional persistence of species that are excluded by competition locally. Persistence, and hence diversity, is maximized at intermediate disturbance frequencies. If disturbance and dispersal rates are sufficiently high, the inferior competitor need not have a dispersal advantage to persist. Using a new method for measuring the spatial patterns of nominal data, we show that none of these competition models generates patchiness at equilibrium. In the inhibition model, however, transient patchiness decays very slowly. We compare the cellular automaton models to the corresponding mean-field patch-occupancy models, in which colonization is not restricted to neighboring patches and depends on spatially averaged species frequencies. The patch-occupancy model does an excellent job of predicting the equilibrium frequencies of the species and the conditions required for coexistence, but not of predicting transient behavior.     

The role of colonization in the dynamics of patchy populations of a cyclic vole species

The crash phase of vole populations with cyclic dynamics regularly leads to vast areas of uninhabited habitats. Yet although the capacity for cyclic voles to re-colonize such empty space is likely to be large and predicted to have become evolved as a distinct life history trait, the processes of colonization and its effect on the spatio-temporal dynamics have been little studied. Here we report from an experiment with root voles (Microtus oeconomus) specifically targeted at quantifying the process of colonization of empty patches from distant source patches and its resultant effect on local vole deme size variation in a patchy landscape. Three experimental factors: habitat quality, predation risk and inter-patch distance were employed among 24 habitat patches in a 100 × 300-m experimental area. The first-born cohort in the spring efficiently colonized almost all empty patches irrespective of the degree of patch isolation and predation risk, but this was dependent on habitat quality. Just after the initial colonization wave the deme sizes in patches of the same quality were underdispersed relative to Poisson variance, indicating regulated (density-dependent) settlement. Towards the end of the breeding season local demographic processes acted to smooth out the initial post-colonization differences among source and colonization patches, and among patches of initially different quality. However, at this time demographic stochasticity had also given rise to a large (overdispersed) variation in deme sizes that may have contributed to an overshadowing of the effect of other factors. The results of this experiment confirmed our expectation that the space-filling capacity of voles is large. The costs associated with transience appeared to be so low, at least at the spatial scale considered in this experiment, that such costs are not likely to substantially constrain habitat selection and colonization in the increase phase of cyclic patchy populations.