A species invasion mediated through habitat structure, intraguild predation and parasitism.

With field, laboratory, and modeling approaches, we examined the interplay among habitat structure, intraguild predation (IGP), and parasitism in an ongoing species invasion. Native Gammarus duebeni celticus (Crustacea: Amphipoda) are often, but not always, replaced by the invader Gammarus pulex through differential IGP. The muscle-wasting microsporidian parasite Pleistophora mulleri infects the native but not the invader. We found a highly variable prevalence of P. mulleri in uninvaded rivers, with 0–91% of hosts parasitized per sample. In addition, unparasitized natives dominated fast-flowing riffle patches of river, whereas parasitized individuals dominated slower- flowing, pooled patches. We examined the survivorship of invader and native in single and mixed-species microcosms with high, intermediate, and zero parasite prevalence. G. pulex survivorship was high in all treatments, whereas G. duebeni subsp. celticus survivorship was significantly lower in the presence of the invader. Further, parasitized G. duebeni subsp. celticus experienced near-total elimination. Models of the species replacement process implied that parasite-enhanced IGP would make invasion by G. pulex more likely, regardless of habitat and parasite spatial structure. However, where heterogeneity in parasite prevalence creates a landscape of patches with different susceptibilities to invasion, G. pulex may succeed in cases where invasion would not be possible if patches were equivalent. The different responses of parasitized and unparasitized G. duebeni subsp. celticus to environmental heterogeneity potentially link landscape patterns to the success or failure of the invasion process.

Functional responses of the intertidal amphipod Echinogammarus marinus: effects of prey supply, model selection and habitat complexity.

The influence of predation in structuring ecological communities can be informed by examining the shape and magnitude of the functional response of predators towards prey. We derived functional responses of the ubiquitous intertidal amphipod Echinogammarus marinus towards one of its preferred prey species, the isopod Jaera nordmanni. First, we examined the form of the functional response where prey were replaced following consumption, as compared to the usual experimental design where prey density in each replicate is allowed to deplete. E. marinus exhibited Type II functional responses, i.e. inversely density-dependent predation of J. nordmanni that increased linearly with prey availability at low densities, but decreased with further prey supply. In both prey replacement and non-replacement experiments, handling times and maximum feeding rates were similar. The non-replacement design underestimated attack rates compared to when prey were replaced. We then compared the use of Holling’s disc equation (assuming constant prey density) with the more appropriate Rogers’ random predator equation (accounting for prey depletion) using the prey non-replacement data. Rogers’ equation returned significantly greater attack rates but lower maximum feeding rates, indicating that model choice has significant implications for parameter estimates. We then manipulated habitat complexity and found significantly reduced predation by the amphipod in complex as opposed to simple habitat structure. Further, the functional response changed from a Type II in simple habitats to a sigmoidal, density-dependent Type III response in complex habitats, which may impart stability on the predator−prey interaction. Enhanced habitat complexity returned significantly lower attack rates, higher handling times and lower maximum feeding rates. These findings illustrate the sensitivity of the functional response to variations in prey supply, model selection and habitat complexity and, further, that E. marinus could potentially determine the local exclusion and persistence of prey through habitat-mediated changes in its predatory functional responses.

Biodiversity among mussels: separating the influence of sizes of mussels from the ages of patches

The role of habitat structure in controlling the composition of assemblages has often been studied, but is rarely manipulated so that it is distinguishable from other factors. Differences in habitat structure as determined by differences in mussel size structure may affect the diversity of assemblages associated with mussel beds. Previous studies examining the effect of the size of individual mussels in a patch on the diversity of associated macro-faunal assemblages confounded the age of the patch with the size of the mussels. We manipulated the age of mussel patches and the size of the mussels within them to test experimentally whether the size of mussels influenced the structure of associated assemblages. At one of the two locations considered, the structure of macro-faunal assemblages in patches of larger mussels differed significantly from those in patches of the same age composed of smaller mussels. At this location, the size of mussels did not affect species richness but the abundance and proportion of organisms present differed depending on the size of the mussels. Here patches of larger mussels contained greater numbers of Nematodes and Oligochaetes and a lower abundance of taxa, such as faera forsmani and Lepidonotus clava. We also found that invertebrate assemblages in general differed between the two locations. The effect of the size structure of mussels, however, varied spatially demonstrating that the effect of habitat structure on the diversity of associated assemblages is context dependent.

Phylogeographic structure of brown trout (Salmo trutta) in Britain and Ireland: glacial refugia, post-glacial colonisation, and origins of sympatric populations

The phylogeographical structure of brown trout Salmo trutta in Britain and Ireland was studied using polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) analysis of four mitochondrial DNA segments (16S/ND1, ND5/6, COXIII/ND5 and ND5/12S). Analysis of 3636 individuals from 83 sites-morphotypes revealed a total of 25 haplotypes. These haplotypes were nested in seven two-step clades. Although there was a clear geographical patterning to the occurrence of derived clades, admixture among ancestral clades was extensive throughout the studied area. A relevant feature of the data was that some populations contained mixtures of highly divergent clades. This type II phylogeographic pattern is uncommon in nature. Clade intermixing is likely to have taken place during earlier interglacials as well as since the Last Glacial Maximum. The anadromous life history of many S. trutta populations has probably also contributed to clade mixing. Based on the data presented here and published data, postglacial colonization of Britain and Ireland most likely involved S. trutta from at least five potential glacial refuges. Probable locations for such refugia were: south of England-western France, east of the Baltic Sea, western Ireland, Celtic Sea and North Sea. Ferox S. trutta, as defined by their longevity, late maturation and piscivory, exhibited a strong association with a particular clade indicating that they share a common ancestor. Current evidence indicates that the Lough Melvin gillaroo S. trutta and sonaghen S. trutta sympatric types diverged prior to colonization of Lough Melvin and, although limited gene flow has occurred since secondary contact, they have remained largely reproductively isolated due to inlet and outlet river spawning segregation. Gillaroo S. trutta may reflect descendents of a previously more widespread lineage that has declined due to habitat alterations particularly affecting outlet rivers. The mosaic-like distribution of mtDNA lineages means that conservation prioritization in Britain and Ireland should be based on the biological characteristics of local populations rather than solely on evolutionary lineages.

Summer habitat associations of bats between riparian landscapes and within riparian areas

The present study examines those features which promote bat feeding in agricultural riparian areas and the riparian habitat associations of individual species. Activity of Nathusius' pipistrelle (Pipistrellus nathusii), common pipistrelle (Pipistrellus pipistrellus), soprano pipistrelle (Pipistrellus pygmaeus), Leisler's bat (Nyctalus leisleri), and Myotis species (Myotis sp.) were recorded, and their habitat associations both "between" and "within" riparian areas were analyzed. General feeding activity was associated with reduced agricultural intensity, riparian hedgerow provision, and habitat diversity. Significant habitat associations for P. pipistrellus were observed only within riparian areas. Myotis species and P. pygmaeus were significantly related to indices of landscape structure and riparian hedgerow across spatial scales. Myotis species were also related to lower levels of riffle flow at both scales of analysis. The importance of these variables changed significantly, however, between analysis scales. The multi-scale investigation of species-habitat associations demonstrated the necessity to consider habitat and landscape characteristics across spatial scales to derive appropriate conservation plans.

Habitat Isolation Reduces the Temporal Stability of Island Ecosystems in the Face of Flood Disturbance

Natural ecosystems are increasingly exposed to multiple anthropogenic stressors, including land-use change, deforestation, agricultural intensification, and urbanisation, all of which have led to widespread habitat fragmentation, which is also likely to be amplified further by predicted climate change. The potential interactive effects of these different stressors cannot be determined by studying each in isolation, although such synergies have been largely ignored in ecological field studies to date. Here, we use a model system of naturally fragmented islands in a braided river network, which is exposed to periodic inundation, to investigate the interactive effects of habitat isolation and flood disturbance. Food web structure was similar across the islands during periods of hydrological stability, but several key properties were altered in the aftermath of flood disturbance, based on distance of the islands from the regional source pool of species: taxon richness and mean food chain length declined with habitat isolation after flooding, while the proportion of basal species increased. Greater species turnover through time reflected the slower process of re-colonisation on the more distant islands following disturbance. Increased variability of several food web properties over a 1-year period highlighted the reduced temporal stability of isolated habitat fragments. Many of these effects reflected the differential successes of predator and prey species at re-colonising the islands: even though larger, more mobile consumers may reach the more distant islands first, they cannot establish populations until the lower trophic levels have successfully reassembled. These results highlight the susceptibility of fragmented ecosystems to environmental perturbations. © 2013 Elsevier Ltd.

Community structure, diversity and spatial organization of enchytraeids in Mediterranean urban holm oak stands

There is a strong northern bias in Europe as regards enchytraeid community ecology, particularly in urban settings. We approached the enchytraeid assemblages of urban holm oak stands in Naples and Siena adopting a high intensity sampling that, for the first time in the Mediterranean climate zone, would ensure that the data collected be representative of the target populations. Structural parameters (diversity and evenness, biomass, size classes, aggregation) were compared across different spatial (regional, urban district, within habitat) and temporal scales (season and year). Species richness was found to change significantly only at regional scale; background data suggest that this may depend on the higher environmental heterogeneity occurring at Naples. Differences in size class structure were significant only on a seasonal scale and within either city separately. With one exception (Fridericia bulbosa s.s.), the patterns of spatial aggregation of the common species were fairly robust and the total range of patchiness was consistent with previous studies, despite the different sampling methodologies. The size of the sampling unit, the number of replicates per plot and the number of plots proposed in this study appear suitable to obviate the difficulties of evaluating Mediterranean enchytraeid communities.

Ecological dynamics of extinct species in empty habitat networks. 2. The role of host plant dynamics

This paper explores the relative effects of host plant dynamics and butterfly-related parameters on butterfly persistence. It considers an empty habitat network where a rare butterfly (Cupido minimus) became extinct in 1939 in part of its historical range in north Wales, UK. Surviving populations of the butterfly in southern Britain were visited to assess use of its host plant (Anthyllis vulneraria) in order to calibrate habitat suitability and carrying capacity in the empty network in north Wales. These data were used to deduce that only a portion ( similar to 19%) of the host plant network from north Wales was likely to be highly suitable for oviposition. Nonetheless, roughly 65,460 eggs (3273 adult equivalents) could be expected to be laid in north Wales, were the empty network to be populated at the same levels as observed on comparable plants in surviving populations elsewhere. Simulated metapopulations of C. minimus in the empty network revealed that time to extinction and patch occupancy were significantly influenced by carrying capacity, butterfly mean dispersal distance and environmental stochasticity, although for most reasonable parameter values, the model system persisted. Simulation outputs differed greatly when host plant dynamics was incorporated into the modelled butterfly dynamics. Cupido minimus usually went extinct when host plant were at low densities. In these simulations host plant dynamics appeared to be the most important determinant of the butterfly's regional extirpation. Modelling the outcome of a reintroduction programme to C. minimus variation at high quality locations, revealed that 65% of systems survived at least 100 years. Given the current amount of resources of the north Wales landscape, the persistence of C. minimus under a realistic reintroduction programme has a good chance of being successful, if carried out in conjunction with a host plant management programme.