Effects of hydroperiod and predation on a Mississippi River floodplain invertebrate community

 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996     

Resistance, resilience, and community dynamics in mediterranean-climate streams

Streams and rivers in mediterranean-climate regions (med-rivers) are subjected to sequential, yet contrasting hydrologic disturbances of drying and flooding. Although seasonally predictable, these disturbances can vary in intensity and duration within and among mediterranean-climate regions (med-regions). Consequently, med-rivers differ in the permanence of their aquatic habitats. To persist, species have acquired matched resistance and resilience adaptations. They gain resistance either by enduring the stress or avoiding it. Community recovery (or resilience) is achieved with cessation of hydrologic stress that permits maximization of re-colonization and reproduction. Endurance strategies are usually disturbance-specific, but avoidance enables organisms to cope with both drying and flooding, and is the prevalent resistance strategy. Correspondingly, community persistence depends to a large extent on the integrity of refuges, an aspect that has so far been little explored. Existing information suggests that seasonal community succession becomes more pronounced with increasing aridity and declining water permanence. The invertebrate community in semi-arid med-rivers can therefore undergo succession through three to four identifiable assemblages, whereas in perennial streams the difference between wet and dry period assemblages is smaller. Community turnover is influenced by the intensity of the hydrologic disturbances and varies between wet and drought years.     

Nearshore subtidal community structure compared between inner coast and outer coast sites in Southeast Alaska

Processes that structure subarctic marine communities, particularly in glaciated regions, are not well understood. This understanding is needed as a baseline and to manage these communities in the face of future climate-driven changes. This study investigates two coastal regions of Southeast Alaska with the goals to (a) identify and compare patterns of subtidal community structure for macroalgal, fish, macroinvertebrate (>5?cm), and small epibenthic invertebrate (<5?cm) communities between inner coast and outer coast sites and (b) link patterns of community structure to habitat and environmental parameters. Species assemblage and benthic habitat data were used to compare species diversity and community composition at 6?m and 12?m depths at nine inner coast and nine outer coast sites. Multivariate analysis was applied to reduce environmental variables to major gradients, to resolve community structure, and to relate community structure to habitat and environmental variables. Increased salinity and decreased temperature at outer coast sites compared with inner coast sites were associated with community structure, with greater species diversity at outer coast sites at 6?m depth. Invertebrate community composition was associated with benthic habitat, including crust and coralline algae for macroinvertebrates, and algal cover and substrate for small epibenthic invertebrates. This research suggests that marine communities in glaciated regions are strongly influenced by freshwater input and that future climate-driven changes in freshwater input will likely result in marine community composition changes.     

Persistence and stability of lotic invertebrate communities in New Zealand

1. Persistence and stability of lotic invertebrate communities were determined at an annual time scale over a 9‐year period (1990–98) at 26 river sites over the northern half of New Zealand. In addition, a number of water quality variables were measured monthly and flow information collected continuously over the same period at each site. 2. The aim of the study was to describe the levels of interannual variability in invertebrate communities, and relate community changes to variability in environmental conditions. The consequences of this temporal variability for the sensitivity of predictive models were also investigated. 3. Levels of change in environmental conditions varied significantly between years, but were relatively similar across sites. In contrast, community persistence (similarity between years in species assemblage composition), and stability (similarity between years with respect to relative abundance of species in the community) both varied significantly between sites, but changes between years were similar. Community stability was highest at sites with relatively harsh flow conditions (high coefficient of variation, high relative size of floods), and was also greater in communities dominated by Ephemeroptera. 4. Relationships between change in environmental conditions and changes in community composition and structure were relatively weak for most individual sites. However, when average levels of change for each of the 26 sites were used, communities showed greater persistence under conditions where flow conditions remained relatively constant. Water quality changes had no significant effect on community persistence when assessed for all 26 sites combined. 5. Results from this study suggest that lotic invertebrate communities fluctuate around a relatively stable state, at least over a 9‐year period. However, the extent of interannual variation in community composition and structure observed, along with the relatively low degree of cluster fidelity observed within a single region, suggests that predictive models based on reference site conditions extrapolated over several years should be applied with caution in New Zealand streams.     

Long-term population and community patterns of benthic macroinvertebrates and fishes in Northern California Mediterranean-climate streams

Long-term studies can document temporal patterns in freshwater ecosystems, and this is particularly important in mediterranean-climate (med-climate) regions because of strong interannual variation in precipitation amounts and consequently stream flow. We review long-term studies of populations and communities of benthic macroinvertebrate and fishes from sites throughout the med-climate region of California and develop generalities that may apply broadly to med-climate streams worldwide. Severe drought may result in community shifts, and alter age-structure in both macroinvertebrates and fishes. Within-year seasonal patterns in macroinvertebrate communities can be influenced by annual variability in flow regimes. Macroinvertebrate biological-monitoring metrics with consistently low intra-annual variability may be especially applicable in med-climate streams, as is the use of different temporal windows to describe reference periods to reduce influence of interannual variability on impact detection. Long-term data can be used to develop macroinvertebrate-based metrics that can either show or be independent of climate-change effects. Most macroinvertebrate species are temporally rare in their annual occurrence. Multiple components of natural flow regimes can favor native over invasive fishes. Long-term, quantitative information from med-climate streams is generally lacking, which is a hindrance to both management practices and development of appropriate ecological constructs.     

Effects of fish predation and habitat type on stream benthic communities

The influence of habitat on interactions between a fish predator (brown trout Salmo trutta) and a benthic invertebrate community was studied in nine field enclosures (8 ×3 m) in a creek in southern Sweden. Three habitat treatments were tested, a shallow sandy habitat, a deep habitat containing a mixture of large and small cobbles and a moderately deep habitat with large cobbles. The one month-long experiment showed that there were no major differences in the abundance and biomass of the benthic macroinvertebrate fauna among these habitats as no functional groups of invertebrates and only a few taxa differed between treatments. Invertebrate drift rates decreased over time, which was probably related to seasonal changes in invertebrate life cycles or to effects of predation independent of habitat type, as there was no difference between treatments.     

The influence of habitat structure and flow permanence on invertebrate communities in karst spring systems

The macroinvertebrate fauna of five karst (limestone) springbrook systems with contrasting physical habitat and discharge patterns were investigated to examine the role of flow permanence and habitat structure on macroinvertebrate community composition. Clear physical differences were identified between perennial and intermittent springs and individual sampling stations. However, flow permanence, water temperature and the input of leaf litter exerted a greater influence on the aquatic invertebrate community than habitat structure. Perennial sites were characterised by a greater abundance of macroinvertebrates and greater Ephemeroptera, Plecoptera and Trichoptera (EPT) richness than intermittent sites. The fauna of all of the springbrook systems examined were dominated by relatively common and ubiquitous taxa (e.g. Gammarus pulex) although a number of taxa displaying life cycle adaptations to ephemeral aquatic habitats (e.g. Limnephilus auricula and Stenophylax permistus) were recorded at intermittent sites.     

Food selection of Coregonus lavaretus in a brackish water ecosystem

This study examined how variability in the abundance and biomass structure of benthic invertebrates affected the feeding choice of the whitefish Coregonus lavaretus on a hard bottom habitat of the brackish Baltic Sea. In general, crustaceans such as Idotea balthica and Gammarus spp. were preferred over molluscs. Although being the most numerous taxon in the invertebrate samples, Mytilus trossulus was the lowest ranking in C. lavaretus food preference. The availability of benthic invertebrate prey set the dietary range of fish but the selectivity largely described fish feeding within this range. There was no clear link between fish predation and the dominance structure of benthic invertebrate communities, suggesting that species composition, abundance and biomass of invertebrate species had no impact on the feeding selectivity of the fish. Thus, while fish predation may not affect the dominant species within a benthic community, due to strong selectivity fish may impose strong pressure on some rarer but highly preferred invertebrate prey species.     

Macroinvertebrate community structure and biological traits related to flow permanence in a Mediterranean river network

In Mediterranean climate areas, the great seasonal variability in temperature and rainfall is considered to be an evolutionary pressure that constrains plant and animal communities and their biological traits. Droughts alter habitat availability (changes of flow alter riffle-pool sequences), although habitat characteristics may also exacerbate drought to some extent. Using a simple quantitative index based on the proportion of conglomerate bedrock versus gravel and cobbles, pools versus riffles and winter versus summer flow, we show how habitat characteristics (in terms of substratum and flow) may influence the permanency of a stream site and how flow permanence constrains macroinvertebrate community structure and biological traits. Annual and seasonal macroinvertebrate richness, and the EPT (Ephemeroptera, Plecoptera and Trichoptera) and OCH (Odonata, Coleoptera and Hemiptera) metrics differed between permanent, intermittent and ephemeral sites, but not between permanent and intermittent sites. In contrast, distinct biological traits were observed in the three flow categories, although permanent sites presented few significant traits which was attributed to the stability of the habitat. Intermittent sites were dominated by taxa with pool-like strategies, while ephemeral sites were characterized by fauna with life-history adaptations to floods and droughts. In contrast to most traits (e.g., dissemination, reproduction, substrate relation), which were more constrained by local flow and substrate characteristics, life-cycle characteristics did not differ significantly among flow categories. This pattern can be explained by the features of the Mediterranean climate and particularly its high seasonal predictability, which serves as a large-scale filter of life-cycle traits, independently of local hydromorphological characteristics. Our findings indicate that drought is related to habitat characteristics and that local habitat variability favours organisms with certain traits, while other traits are independent of habitat variability on this scale and are probably affected by other large-scale habitat characteristics. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: S. Declerck     

Non-Random Variability in Functional Composition of Coral Reef Fish Communities along an Environmental Gradient

Changes in the coral reef complex can affect predator-prey relationships, resource availability and niche utilisation in the associated fish community, which may be reflected in decreased stability of the functional traits present in a community. This is because particular traits may be favoured by a changing environment, or by habitat degradation. Furthermore, other traits can be selected against because degradation can relax the association between fishes and benthic habitat. We characterised six important ecological traits for fish species occurring at seven sites across a disturbed coral reef archipelago in Indonesia, where reefs have been exposed to eutrophication and destructive fishing practices for decades. Functional diversity was assessed using two complementary indices (FRic and RaoQ) and correlated to important environmental factors (live coral cover and rugosity, representing local reef health, and distance from shore, representing a cross-shelf environmental gradient). Indices were examined for both a change in their mean, as well as temporal (short-term; hours) and spatial (cross-shelf) variability, to assess whether fish-habitat association became relaxed along with habitat degradation. Furthermore, variability in individual traits was examined to identify the traits that are most affected by habitat change. Increases in the general reef health indicators, live coral cover and rugosity (correlated with distance from the mainland), were associated with decreases in the variability of functional diversity and with community-level changes in the abundance of several traits (notably home range size, maximum length, microalgae, detritus and small invertebrate feeding and reproductive turnover). A decrease in coral cover increased variability of RaoQ while rugosity and distance both inversely affected variability of FRic; however, averages for these indices did not reveal patterns associated with the environment. These results suggest that increased degradation of coral reefs is associated with increased variability in fish community functional composition resulting from selective impacts on specific traits, thereby affecting the functional response of these communities to increasing perturbations.     

Characterization of tropical near-shore fish communities by coastal habitat status on spatially complex island systems

Synopsis We present a protocol for characterizing near-shore fish habitat as well as fish communities for Andros Island, Bahamas, a complex coastal-reef island system. Benthic assessments and beach seine surveys were carried out at sites varying in coastal and benthic characteristics. Temporal variability affected fish community composition, indicating that attempts to characterize a fish community should include sampling evenly across tides, times of day, and seasons. Univariate and multivariate analyses revealed that each site harbored a unique fish community, with the greatest variability within each site attributed to seasonal changes. Measures of diversity (Shannon–Weiner Index and number of species) were markedly different at sites with varying coverage of seagrass and macro-algae and extents of disturbance. Total abundance of fishes was positively related to the percent of bare sand. We suggest that thorough sampling of coastal fish communities can be applied to comparative and long-term studies. This protocol for the characterization of complex island habitats can be applied to ecological studies aimed at understanding the responses of fishes to small-scale changes in coastal areas and habitat structure due to land use and shoreline alterations.     

The Impact of Hydrological Restoration on Benthic Aquatic Invertebrate Communities in a New Zealand Wetland

Drainage is a major disturbance affecting wetlands, as drains lower water tables and convert lentic habitats to lotic ones. Consequently, invertebrate communities in drained wetlands are likely to differ from those in unimpacted wetlands. This study investigated the effect of hydrological restoration on invertebrate communities in small drains in a New Zealand fen. Invertebrates were collected over 4 summers from 10 drains within the wetland, one of which was blocked as part of a restoration program. The sampling protocol thus represented a Before‐After Control‐Impact experiment. Invertebrate community composition varied over the 4 years, but variability was greatest in the manipulated drain before and after it was blocked. Relative abundance of the amphipod Paraleptamphopus decreased after blockage, whereas those of the midges Chironomus zelandicus and Tanypodinae increased. Relative abundances of these taxa in control sites were unchanged. Hydraulic restoration thus had a demonstrable impact on the invertebrate communities. The invertebrate community of the blocked drain was compared to that of natural wetlands in undisturbed catchments. Similarity was very low prior to drain blockage, but increased following drain blockage. Invertebrate communities in the restored drain were more similar to those of low pH wetlands than high pH wetlands. Given the goal of restoring the communities to those similar to natural conditions, this was a beneficial result. These results, coupled with studies that showed a decline in the cover of alien pasture grasses around the blocked drain, suggest that drain blockage represents a cost‐effective way of restoring wetland plant and aquatic invertebrate communities, especially where connectivity allows for the natural recruitment of these organisms into restored areas.     

Drought ecohydrology alters the structure and function of benthic invertebrate communities in mountain streams

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Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.     

Mechanisms of maintenance of tropical freshwater fish communities in the face of disturbance

Community resistance to, and resilience from, perturbation will determine the trajectory of recovery from disturbance. Although selective timber extraction is considered a severe disturbance, fish communities from headwater streams around Danum Valley Field Centre, Sabah, Malaysia, showed few long-term changes in species composition or abundance. However, some species showed short-term (< 18 months) absence or decrease in abundance. These observations suggested that both resistance and resilience were important in maintaining long-term fish community structure. Resistance to perturbation was tested by monitoring fish communities before and after the creation of log-debris dams, while resilience was investigated by following the time-course of recolonization following complete removal of all fish. High community resistance was generally shown although the response was site-specific, dependent on the composition of the starting community, the size of the stream and physical habitat changes. High resilience was demonstrated in all recolonization experiments with strong correlations between pre- and post-defaunation communities, although there was a significant difference between pool and riffle habitats in the time-course of recovery. These differences can be explained by the movement characteristics of the species found in the different habitats. Resilience appeared to be a more predictable characteristic of the community than resistance and the implications of this for ensuring the long-term persistence of fish in the area are discussed.     

Effects of floods versus low flows on invertebrates in a New Zealand gravel-bed river

1. Floods and low flows are hydrological events that influence river ecosystems, but few studies have compared their relative importance in structuring invertebrate communities. Invertebrates were sampled in riffles and runs at eight sites along 40 km of a New Zealand gravel‐bed river every 1–3 months over 2.5 years, during which time a number of large flood and low flow events occurred. Flows were high in winter and spring, and low in summer and autumn. Four flow‐related variables were calculated from hydrological data: flow on the day of sampling (Q_sample), maximum and minimum flow between successive samples (Q_max and Q_min, respectively), and the number of days since the last bed‐moving flood (N_days). 2. The invertebrate community was summarised by relative densities of the 19 most abundant taxa and four biotic metrics [total abundance, taxon richness, the number of Ephemeroptera, Plecoptera and Trichoptera taxa (i.e. EPT richness), and per cent EPT]. Invertebrate density fluctuated greatly, and was high in summer and autumn, and low during winter and spring. Stepwise multiple regression (SMR) analysis was used to investigate relationships between the invertebrate community and season, flow, habitat and water temperature. 3. Seasonal variables were included in almost 50% of the SMR models, while flow‐related variables were included in >75% of models. Densities of many taxa were negatively correlated to Q_min and Q_max, and positively correlated to N_days, suggesting that while high flows reduced invertebrate densities, densities recovered with increasing time following a flood. Although season and flow were confounded in this study, many of the taxa analysed display little seasonal variation in abundance, suggesting that flow‐related variables were more important in structuring communities than seasonal changes in density associated with life‐cycles. 4. Five discrete flood and low flow events were identified and changes to invertebrate communities before and after these events examined. Invertebrate densities decreased more commonly after floods than after low flows, and there was a significant positive relationship between the number of taxa showing reductions in density and flood magnitude. Densities of most invertebrates either remained unchanged, or increased after low flow events, except for four taxa whose densities declined after a very long period (up to 9 months) of low flow. This decline was attributed to autogenic sloughing of thick periphyton communities and subsequent loss of habitat for these taxa. 5. Invertebrate communities changed more after floods and the degree of change was proportional to flood magnitude. Community similarity increased with increasing time since the last disturbance, suggesting that the longer stable flows lasted, the less the community changed. These results suggest that invertebrate communities in the Waipara River were controlled by both floods and low flows, but that the relative effects of floods were greater than even extended periods of extreme low flow. 6. Hydraulic conditions in riffles and runs were measured throughout the study. Riffles had consistently faster velocities, but were shallower and narrower than runs at all measured flows. Invertebrate density in riffles was expressed as a percentage of total density and regressed against the flow‐related variables to see whether invertebrate locations changed according to flow. Significant negative relationships were observed between the per cent density of common taxa in riffles and Q_sample, Q_max and Q_min. This result suggests either that these animals actively drifted into areas of faster velocity during low flows, or that their densities within riffles increased as the width of these habitats declined.     

Effects of natural hydrological variability on fish assemblages in small Mediterranean streams: Implications for ecological assessment

Small Mediterranean streams are shaped by predictable seasonal events of flooding and drying over an annual cycle, and present a strong inter and intra-annual variation in flow regime. Native fish assemblages in these streams are adapted to this natural environmental variability. The distinction of human-induced disturbances from the natural ones is thus a crucial step before assessing the ecological status of these streams. In this aim, the present study evaluates the effects of natural hydrological variability on fish assemblages from disturbed and least disturbed sites in small intermittent streams of south Portugal. Data were collected over the last two decades (1996–2011) in 14 sites located in the Guadiana and Sado river basins. High variability of fish assemblages was strongly dependent on human-induced disturbances, particularly nutrient/organic load and sediment load, and on natural hydrological variability. Natural hydrological variability can act jointly with anthropogenic disturbances, producing changes on fish assemblages structure of small intermittent streams. In least disturbed sites, despite the natural disturbances caused by inter-annual rainfall variations (including drought and flood events), fish assemblages maintained a long-term stability and revealed a high resilience. On the contrary, disturbed sites presented significantly higher variability on fish assemblages and a short and long-term instability, reflecting a decrease on the resistance and resilience of fish assemblages. Under these conditions, fish fauna integrity is particularly vulnerable and the ecological assessment may be influenced by natural hydrological variations. High hydrological variability (especially if it entails high frequency of dryer years and meaningful cumulative water deficit) may affect the impact of the human pressures with significant and consistent consequences on fish assemblage composition and integrity. In this study, fish metrics that maximize the detection of human degradation and minimize the response to natural variability were based on the relative abundance of native species (insectivorous species, eurytopic species, water column species, native lithophilic species), relative abundance of species with intermediate tolerance and relative number of exotic species. Results highlight the importance of assessing temporal variability on stream biomonitoring programs and emphasize the need to improve the assessment tools, accounting for long-term changes in fish assemblages, namely by selecting the most appropriate fish metrics that respond to anthropogenic disturbances but exhibit low natural temporal variability, essential both in the characterization of the biological reference conditions and in the development of fish indexes in intermittent streams.     

Colonisation of Introduced Timber by Algae and Invertebrates, and its Potential Role in Aquatic Ecosystem Restoration

As part of a habitat restoration experiment wood substrates (red gum) were introduced to two lowland streams of SE Australia in which habitat has been severely degraded by deposition of sand eroded from higher in the catchment. We monitored net primary production (NPP) and community respiration (CR), nutrient concentrations and the succession of algae and invertebrates (abundance and species richness), sampling at 2, 4, 8, 12, 16 and 20 weeks. Colonisation by algae was rapid, and there were distinct changes in the assemblages over the first 4 weeks. Thereafter, changes were much less marked. There were also differences in nutrient concentrations and some measures of algal abundance between the two creeks. As with the algae, invertebrates colonised these substrates extremely rapidly, peaking in abundance and richness in week 8. Invertebrate abundances closely tracked changes in the abundance of algae. By the end of the study both algal and invertebrate communities were in apparent decline, with sharp decreases in invertebrate and algal abundance and invertebrate species richness. Rates of GPP also declined toward the end of the experiment, and this coincided with the detachment of large mats of filamentous algae and the recession of flows over the summer months. However, in both streams the added timber quickly created habitat with high levels of primary production in an otherwise strongly heterotrophic stream system. These hotspots of autotrophic production were quickly colonised by high numbers of macroinvertebrates indicating timber addition may provide an effective means of augmenting habitat for algae and invertebrates in sanded streams.     

The response of perennial and temporary headwater stream invertebrate communities to hydrological extremes

The headwaters of karst rivers experience considerable hydrological variability, including spates and streambed drying. Extreme summer flooding on the River Lathkill (Derbyshire, UK) provided the opportunity to examine the invertebrate community response to unseasonal spate flows, flow recession and, at temporary sites, streambed drying. Invertebrates were sampled at sites with differing flow permanence regimes during and after the spates. Following streambed drying at temporary sites, dewatered surface sediments were investigated as a refugium for aquatic invertebrates. Experimental rehydration of these dewatered sediments was conducted to promote development of desiccation-tolerant life stages. At perennial sites, spate flows reduced invertebrate abundance and diversity, whilst at temporary sites, flow reactivation facilitated rapid colonisation of the surface channel by a limited number of invertebrate taxa. Following streambed drying, 38 taxa were recorded from the dewatered and rehydrated sediments, with Oligochaeta being the most abundant taxon and Chironomidae (Diptera) the most diverse. Experimental rehydration of dewatered sediments revealed the presence of additional taxa, including Stenophylax sp. (Trichoptera: Limnephilidae) and Nemoura sp. (Plecoptera: Nemouridae). The influence of flow permanence on invertebrate community composition was apparent despite the aseasonal high-magnitude flood events. Flow permanence was also critical in determining the community response to the spate flows. Following streambed drying at temporary sites, the surficial sediments overlying the karstic bedrock functioned as an effective refugium for several taxa. The development of aquatic insects following experimental rehydration indicated that these taxa survived in dewatered sediments as desiccation-resistant eggs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Sonja Stendera     

Landscape geometry determines community response to disturbance

Ecological communities are impacted by anthropogenic changes in both habitat geometry (i.e. amount, shape, fragmentation and connectivity of habitat) and disturbance regime. Although the effect of each of these drivers on diversity is well-documented, few studies have considered how habitat geometry and disturbance interact to affect diversity. We used a miniature landscape of moss patches to experimentally manipulate both habitat geometry and disturbance frequency on microarthropod communities. Species richness and abundance in local patches declined linearly with disturbance rate in all experimental landscapes, but the speed of this decline (a measure of ecological resilience) depended on the size and connectivity of the surrounding region. Reductions in region size had little effect on community resilience to disturbance until habitat loss resulted in complete loss of connectivity between patches, suggesting a threshold in community response to habitat loss. Beyond this threshold, repeated disturbance resulted in rapid declines in patch species richness and abundance and substantial changes in community composition. These effects of habitat geometry and disturbance on diversity were scale-dependent. Gamma (regional-scale) diversity was unaffected by habitat geometry, suggesting experimental reductions in alpha (local-scale) diversity were offset by increases in beta diversity. There was no effect of body size, abundance, or trophic position in determining species response to disturbance. Taxonomic grouping had a weak effect, with oribatids less affected by drought. We conclude that, in this system, dispersal from the surrounding metacommunity is vital in allowing recovery of local communities from disturbance. When habitat loss and fragmentation disrupt this process, extinctions result. Studies that examine separately the effects of habitat alterations and disturbance on diversity may therefore underestimate their combined effects.