Wind mediated dispersal of freshwater invertebrates in a rock pool metacommunity: differences in dispersal capacities and modes

Current evidence suggests regular overland transport of different freshwater invertebrates by wind, mainly over short distances. Yet, very little is known about the mechanism and scale of this process or about differences in wind dispersal dynamics and capacities among taxa and propagule types. We investigated wind dispersal of freshwater invertebrates in a cluster of temporary rock pools (spatial scale: 9,000 m²) in South Africa. Dispersing propagules and propagule bank fragments (i.e. aggregates of sediments and propagules) were intercepted during 1 month using a combination of windsocks (1.5 m above ground level) and sticky traps (ground level). The potential movement of propagule bank fragments (i.e. aggregates of propagules and sediments) was also simulated by tracking inter-pool movements of differently sized artificial substrate fragments similar to dry propagule bank fragments. We detected differences in the composition of dispersing communities intercepted at different altitudes (ground level and at 1.5 m). Comparison of dispersal distance distributions also revealed significant differences among taxa. Overall, larger propagule types (e.g. adult ostracods and oribatid mites) dominantly travelled near ground level while small resting eggs and cryptobiotic life stages of copepods were most frequently collected at higher altitudes (1.5 m) and dispersed over the longest distances. Finally, not only dispersal of single propagules but also ground level transport of propagule bank fragments was shown to contribute to local dispersal dynamics in temporary aquatic habitats.     

Predictions of climate change infer increased environmental harshness and altered connectivity in a cluster of temporary pools

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Relative importance of different dispersal vectors for small aquatic invertebrates in a rock pool metacommunity

The extent and frequency of passive overland dispersal of freshwater invertebrates as well as the relative importance of different dispersal vectors is not well documented. Although anecdotal evidence subscribing the feasibility of individual vectors in various aquatic systems is abundant, dispersal rates have rarely been quantified for different vectors in one study system. Earlier studies also usually investigated dispersal potential rather than actual dispersal rates. In this study we have estimated passive dispersal rates of invertebrate propagules within a cluster of temporary rock pools via water, wind and amphibians in a direct way. Overflows after heavy rains mediated dispersal of a large number of propagules through eroded channels between pools, which were collected in overflow traps. Taking into account model based predictions of overflow frequency, this corresponds with average dispersal rates of 4088 propagules/channel yr^?1. Wind dispersal rates as measured by numbers of propagules collected on sticky traps mounted between pool basins were very high (average dispersal rate: 649 propagules m^?2 in one month) and were positively related to the proximity of source populations. Finally, invertebrate propagules were also isolated from the faeces of African clawed frogs Xenopus laevis caught from the pools (on average 368 propagules/frog). The combination of short distance wind and overflow dispersal rates likely explain the dominant species sorting and mass effect patterns observed in the metacommunity in a previous study. Amphibian mediated dispersal was much less important as the Xenopus laevis population was small and migrations very rare. Based on our own results and available literature we conclude that both vector and propagule properties determine local passive dispersal dynamics of freshwater invertebrates. Accurate knowledge on rates and vectors of dispersal in natural systems are a prerequisite to increase our understanding of the impact of dispersal on ecology (colonisation, community assembly, coexistence) and evolution (gene flow, local adaptation) in fragmented environments.     

The role of metacommunity processes in shaping invertebrate rock pool communities along a dispersal gradient

Explaining the variance of local communities in a spatial‐environmental matrix is one of the core interests of ecology today. Recent progress in metacommunity theory has made a substantial contribution to this field, however good empirical data in support of available theories are still relatively scarce. In this study we sampled a cluster of 36 temporary rock pools four times during one season to assess invertebrate metacommunity structure and dynamics and to search for steering processes and variables. Both Mantel tests and redundancy models indicate that local abiotic factors were dominant over spatial factors in explaining community structure and both were acting independently. Spatial variables were only important for passive dispersers and significantly explained 11% of variation in this community component. Pools connected by temporary overflows hosted more similar communities of passive dispersers than unconnected ones while community dissimilarity significantly increased with inter‐pool distance. A negative curvilinear relation was discovered between taxon richness and isolation in passive dispersers, providing some support for existing theoretical models of Mouquet and Loreau. Of different metacommunity perspectives, a combination of species sorting and mass effects best explains the observed patterns. Additionally, priority effects and monopolization may buffer against the homogenising effects of dispersal and contribute to the distinctness of isolated communities. This is one of the first studies to present evidence for spatial patterns in aquatic communities on such a small spatial scale (a rock ledge of ±9000 m²). Bridging the gap between theory and observed patterns in natural systems is one of the main challenges for future metacommunity research. Small aquatic habitats such as pitcher plants and freshwater rock pools may well have an important role to play as model systems to study ecological processes in a natural spatially explicit environment.     

Scale dependency of processes structuring metacommunities of cladocerans in temporary pools of High‐Andes wetlands

Metacommunity structure can be shaped by a variety of processes operating at different spatial scales. With increasing scale, the compositional variation among local communities (beta diversity) may reflect stronger environmental heterogeneity, but may also reflect reduced exchange of organisms between habitat patches. We analyzed the spatial architecture of a metacommunity of cladoceran zooplankton in temporary pools of High Andes wetlands, with the objective of explaining the spatial dependency of its structure. The spatial distribution of the pools is hierarchical and highly discontinuous: pools are clustered within small wetlands, which lay scattered over valleys that are separated from each other by mountain ridges. We studied a total of 59 pools, belonging to six different wetlands in four different valleys. We assessed pool environmental heterogeneity and sampled active communities and dormant propagule banks of cladoceran zooplankton. Environmental heterogeneity proved very high within wetlands and showed almost no increase with increasing spatial scale. Conversely, diversity partitioning analyses indicated an increase in beta diversity with spatial scale, especially among valleys. Variation partitioning on environmental data and spatial RDA models suggested environmental heterogeneity as the most important generator of beta diversity within wetlands. At the largest spatial scale, beta diversity manifested itself mainly as a differentiation of species occurrence patterns among valleys, which could not be entirely explained by environmental variables. Our study thus presents a case where environmental control seems to be the dominant metacommunity structuring process at the smallest spatial scale, whereas neutral processes and dispersal limitation are the most likely generators of beta diversity at the largest spatial scale.     

Dispersal of aquatic organisms by waterbirds: a review of past research and priorities for future studies

1. Inland wetlands constitute ecological islands of aquatic habitat often isolated by huge areas of non-suitable terrestrial habitats. Although most aquatic organisms lack the capacity to disperse by themselves to neighbouring catchments, many species present widespread distributions consistent with frequent dispersal by migratory waterbirds.
2. A literature review indicates that bird-mediated passive transport of propagules of aquatic invertebrates and plants is a frequent process in the field, at least at a local scale. Both endozoochory (internal transport) and ectozoochory (external transport) are important processes.
3. The characteristics of the dispersed and the disperser species that facilitate such transport remain largely uninvestigated, but a small propagule size tends to favour dispersal by both internal and external transport.
4. We review the information currently available on the processes of waterbird-mediated dispersal, establishing the limits of current knowledge and highlighting problems with research methods used in previous studies. We also identify studies required in the future to further our understanding of the role of such dispersal in aquatic ecology.     

Sources of diversity in a grassland metacommunity: quantifying the contribution of dispersal to species richness

Metacommunity theory suggests a potentially important role for dispersal in diversity maintenance at local, as well as regional, scales. In addition, propagule addition experiments have shown that dispersal often limits local diversity. However, actual dispersal rates into local communities and the contribution of immigrants to observed local diversity are poorly known. We present a new approach that partitions the diversity of a target community into dispersal-maintained and dispersal-independent components. Specifically, we quantify distances through space and time to the nearest potential seed source for naturally occurring recruits in target communities by using hierarchical data on species pools (local, site, region, and seed bank). Using this "recruit tag" approach, we found that dispersal contributed 29%-57% of the seedling diversity in perennial grasslands with different successional histories. However, both dispersal and seedling mortality remained remarkably constant, in absolute terms, over succession. The considerable loss of diversity over secondary succession (66%), therefore, could be understood only by considering how these processes interact with the decreasing disturbance rate (i.e., frequency of gaps) in later-successional sites. We conclude that a metacommunity perspective is relevant and necessary to understand the diversity and community assembly of this study system.     

A potential role for overland dispersal in shaping aquatic invertebrate communities in arid regions

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Effect of light stress from phytoplankton on the relationship between aquatic vegetation and the propagule bank in shallow lakes

1. The way light stress controls the recruitment of aquatic plants (phanerogams and charophytes) is a key process controlling plant biodiversity, although still poorly understood. Our aim was to investigate how light stress induced by phytoplankton, that is, independent from the aquatic plants themselves, determines the recruitment and establishment of plant species from the propagule bank. The hypotheses were that an increase in light stress (i) decreases abundance and species richness both of established aquatic plants and of propagules in the bank and (ii) decreases the recruitment success of plants from this bank. 2. These hypotheses were tested in 25 shallow lakes representing a light stress gradient, by sampling propagule banks before the recruitment phase and when the lakes are devoid of actively growing plants (i.e. at the end of winter), established vegetation at the beginning of the summer and phytoplankton biomass (chlorophyll a) during the recruitment and establishment phase. 3. The phytoplankton biomass was negatively correlated with the richness and abundance of established vegetation but was not correlated with the propagule bank (neither species richness nor propagule abundance). The similarity between the propagule bank and established vegetation decreased significantly with increasing phytoplankton biomass. 4. The contrast in species composition between the vegetation and the propagule bank at the highest light stress suggests poor recruitment from the propagule bank but prompts questions about its origin. It could result from dispersal of propagules from neighbouring systems. Propagules could also originate from a persistent propagule bank formerly produced in the lake, suggesting strong year‐to‐year variation in light stress and, as a consequence, in recruitment and reproductive success of plants.     

Successional colonization of temporary streams: An experimental approach using aquatic insects

The metacommunity concept studies the processes that structure communities on local and regional scales. This concept is useful to assess spatial variability. However, temporal patterns (e.g., ecological succession and colonization) are neglected in metacommunity studies, since such patterns require temporally extensive, and hard to execute studies. We used experimental habitats in temporary streams located within the Brazilian Cerrado to evaluate the importance of succession for the aquatic insect metacommunity. Five artificial habitats consisting of wrapped crushed rock were set transversally to the water flow in five streams. The habitats were sampled weekly to assess community composition, and replaced after sampling to identify new potential colonizers. We analyzed the accumulation of new colonizers after each week using a logistic model. We selected pairs of experimental habitats and estimated the Bray-Curtis dissimilarity index to assess the community composition trajectory during the experiment. We used the dissimilarity values in ANOVA tests, identifying the importance of time and space for the community. The number of new taxa stabilized in the third week, and we estimated a weekly increase of 1.61 new taxa in the community after stabilization. The overall pattern was a small change on community composition, but one stream had a higher weekly turnover. Our results showed a relevant influence of time in the initial communities of aquatic insects of temporary streams. However, we must observe the temporal pattern in a spatial context, once different streams have different successional history regarding number of taxa and community turnover. We highlight the importance of aerial dispersal and movement to seek oviposition sites as an important factor in determining colonization patterns.     

Dormant propagule banks integrate spatio-temporal heterogeneity in cladoceran communities

The dynamics of populations of short-lived organisms are very patchy, both in space and time. The production of dormant propagules, however, results in an effective increase in generation time. We hypothesize that prolonged dormancy, together with variable regeneration niches, result in integration of temporal variability in community structure. In addition, in aquatic habitats, mechanisms such as sediment focussing can contribute to the integration of spatial variability. We tested the hypothesis that dormant propagule banks integrate spatial and temporal variation in active zooplankton communities. This was done by comparing cladoceran species richness and the community structure of hatchling assemblages retrieved from propagule bank samples collected on a single occasion with assemblages encountered in active community samples covering spatial variation (littoral and pelagic zone), diel (day and night), intra-year (May–October) and inter-year variation (1996–2000). The egg bank community structure differed significantly from the active community structure, but the dissimilarity decreased as spatial and temporal variation was better covered by the active community samples. Furthermore, the identification of all fully grown hatchlings (n=214) yielded an equally high number of species (n=22) to that occurring in all active community samples together (a total of 1,730 individuals were analysed). We conclude that the analysis of dormant propagules may form a cost-efficient alternative tool to the analysis of active community samples for an integrated assessment of cladoceran communities.     

Migratory strategies of waterbirds shape the continental‐scale dispersal of aquatic organisms

Long distance dispersal (LDD) of propagules is an important determinant of population dynamics, community structuring and biodiversity distribution at landscape, and sometimes continental, scale. Although migratory animals are potential LDD vectors, migratory movement data have never been integrated in estimates of propagule dispersal distances and LDD probability. Here we integrated migratory movement data of two waterbird species (mallard and teal) over two continents (Europe and North America) and gut retention time of different propagules to build a simple mechanistic model of passive dispersal of aquatic plants and zooplankton. Distance and frequency of migratory movements differed both between waterbird species and continents, which in turn resulted in changes in the shapes of propagule dispersal curves. Dispersal distances and the frequency of LDD events (generated by migratory movements) were mainly determined by the disperser species and, to a lesser extent, by the continent. The gut retention time of propagules also exerted a significant effect, which was mediated by the propagule characteristics (e.g. seeds were dispersed farther than Artemia cysts). All estimated dispersal curves were skewed towards local‐scale dispersal and, although dispersal distances were lower than previous estimates based only on the vector flight speed, had fat tails produced by LDD events that ranged from 230 to 1209 km. Our results suggest that propagule dispersal curves are determined by the migratory strategy of the disperser species, the region (or flyway) through which the disperser population moves, and the propagule characteristics. Waterbirds in particular may frequently link wetlands separated by hundreds of kilometres, contributing to the maintenance of biodiversity and, given the large geographic scale of the dispersal events, to the readjustment of species distributions in the face of climate change.     

Connectivity and propagule sources composition drive ditch plant metacommunity structure

The fragmentation of agricultural landscapes has a major impact on biodiversity. In addition to habitat loss, dispersal limitation increasingly appears as a significant driver of biodiversity decline. Landscape linear elements, like ditches, may reduce the negative impacts of fragmentation by enhancing connectivity for many organisms, in addition to providing refuge habitats. To characterize these effects, we investigated the respective roles of propagule source composition and connectivity at the landscape scale on hydrochorous and non-hydrochorous ditch bank plant metacommunities. Twenty-seven square sites (0.5 km² each) were selected in an agricultural lowland of northern France. At each site, plant communities were sampled on nine ditch banks (totaling 243 ditches). Variables characterizing propagule sources composition and connectivity were calculated for landscape mosaic and ditch network models. The landscape mosaic influenced only non-hydrochorous species, while the ditch network impacted both hydrochorous and non-hydrochorous species. Non-hydrochorous metacommunities were dependent on a large set of land-use elements, either within the landscape mosaic or adjacent to the ditch network, whereas hydrochorous plant metacommunities were only impacted by the presence of ditches adjacent to crops and roads. Ditch network connectivity also influenced both hydrochorous and non-hydrochorous ditch bank plant metacommunity structure, suggesting that beyond favoring hydrochory, ditches may also enhance plant dispersal by acting on other dispersal vectors. Increasing propagule sources heterogeneity and connectivity appeared to decrease within-metacommunity similarity within landscapes. Altogether, our results suggest that the ditch network's composition and configuration impacts plant metacommunity structure by affecting propagule dispersal possibilities, with contrasted consequences depending on species' dispersal vectors.     

Passive external transport of freshwater invertebrates by elephant and other mud‐wallowing mammals in an African savannah habitat

1. Recent findings hint at the potential importance of mammals affecting the spatial dynamics of aquatic organisms in areas where mammals live in close association with water. Perhaps the most iconic example of such an environment is the African savannah. 2. We investigated dispersal patterns of freshwater organisms among a set of temporary ponds in SE Zimbabwe to test the hypothesis that large mammals, and particularly African elephants (Loxodonta africana), can be important vectors of aquatic organisms. Dispersal kernels were reconstructed by hatching mud collected from ‘rubbing’ trees located at increasing distances from a set of isolated ponds. To assess the relative importance of other mammalian vectors, the vertical distribution of mud on rubbing trees was mapped and related to the body size of candidate vector species. 3. Laboratory hatching of mud samples revealed large numbers of propagules of 22 invertebrate taxa as well as some aquatic macrophytes. Dispersing communities reflected source communities and diverged with increasing distance from the source. Both dispersal rates and richness of transported taxa decreased significantly with dispersal distance. No indications for differences in dispersal capacity among propagule types were detected. Instead, common propagules were more likely to travel greater distances. Most mud was attached to trees at heights >1.5 m, implicating elephants as the dominant vector. Vertical distributions of tree mud, however, also revealed clustering at heights up to 50 cm and 90–120 cm corresponding to the height of warthog, rhinoceros and buffalo, respectively. Finally, variation in the vertical distribution of mud on trees in combination with differences in vector vagility suggests that local differences in vector species composition may affect passive dispersal dynamics of aquatic organisms. 4. Based on vagility and vector load, mud‐wallowing mammals emerge as highly effective vectors that, in some areas, may be more important in transporting aquatic organisms than traditionally recognised vectors such as waterbirds. Since most large‐ and medium‐sized mammals currently have restricted geographic distributions, it is likely that mammal‐mediated dispersal was more important in the past.     

Dormancy and dispersal as mediators of zooplankton population and community dynamics along a hydrological disturbance gradient in inland temporary pools

At some stage in their life cycle, most zooplankton in temporary waters produce dormant eggs that assemble in a persistent egg bank to cope with unfavourable conditions. As part of a risk-spreading strategy, only a fraction of the egg bank hatches during a single inundation. Besides this dispersal in time, resistant dormant eggs also disperse in space via vectors including wind, water and animals. The structure and functioning of the dormant egg bank has important consequences for (meta) population and (meta) community structure and dynamics. Here, we merge empirical and theoretical data into a conceptual framework for the study of population and community responses in temporary ponds along a gradient in hydrological disturbance. Overall, we conclude that changes in hydrological disturbance may compromise both the abiotic (i.e. water quality) and biotic (i.e. population and community processes) integrity of temporary pools which is especially relevant in light of ongoing anthropogenic alterations in the hydrology of inland waters.     

Strong spatial influence on colonization rates in a pioneer zooplankton metacommunity

The magnitude of community-wide dispersal is central to metacommunity models, yet dispersal is notoriously difficult to quantify in passive and cryptic dispersers such as many freshwater invertebrates. By overcoming the problem of quantifying dispersal rates, colonization rates into new habitats can provide a useful estimate of the magnitude of effective dispersal. Here we study the influence of spatial and local processes on colonization rates into new ponds that indicate differential dispersal limitation of major zooplankton taxa, with important implications for metacommunity dynamics. We identify regional and local factors that affect zooplankton colonization rates and spatial patterns in a large-scale experimental system. Our study differs from others in the unique setup of the experimental pond area by which we were able to test spatial and environmental variables at a large spatial scale. We quantified colonization rates separately for the Copepoda, Cladocera and Rotifera from samples collected over a period of 21 months in 48 newly constructed temporary ponds of 0.18–2.95 ha distributed in a restored wetland area of 2,700 ha in Doñana National Park, Southern Spain. Species richness upon initial sampling of new ponds was about one third of that in reference ponds, although the rate of detection of new species from thereon were not significantly different, probably owing to high turnover in the dynamic, temporary reference ponds. Environmental heterogeneity had no detectable effect on colonization rates in new ponds. In contrast, connectivity, space (based on latitude and longitude) and surface area were key determinants of colonization rates for copepods and cladocerans. This suggests dispersal limitation in cladocerans and copepods, but not in rotifers, possibly due to differences in propagule size and abundance.     

Seed dispersal by waterbirds in southern Africa: comparing the roles of ectozoochory and endozoochory

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Contrasting metacommunity structure and beta diversity in an aquatic‐floodplain system

Habitat connectivity and dispersal interact to structure metacommunities, but few studies have examined these patterns jointly for organisms across the aquatic–terrestrial ecotone. We assessed metacommunity structure and beta diversity patterns of instream benthic invertebrates, riparian carabid beetles (Order: Coleoptera; Family: Carabidae) and riparian spiders (Order: Araneae) at fifteen sites in a river‐floodplain system. Sampling took place over a three‐year period (2010–2012) in the Rhine‐Main‐Observatory LTER site on the Kinzig River, central Germany. This allowed disentangling the combined influence, and temporal variability, of habitat connectivity (i.e. between aquatic and terrestrial) and dispersal ability (i.e. between spiders and beetles, and aerial and aquatic dispersing invertebrates) on the dominant paradigms structuring these metacommunities. We found mostly consistent differences in the manner that metacommunities were structured between groups, with lower levels of variability explained for beetles compared to the other groups. Beetles were consistently structured more by turnover than nestedness components, with greater beta diversity than expected by chance and a minor spatial compared to environmental signal emerging with variance partitioning. Conversely, spiders and benthic invertebrates had lower beta diversity and greater nestedness than null expectation, and a clearer spatial signal controlling metacommunity structure. Our results suggest varying levels of mass effects and species sorting shape river‐floodplain metacommunities, depending on habitat connectivity and dispersal ability. That is, greater connectivity and lower fragmentation along the river compared to the terrestrial zone promoted mass effects, and differences in overall dispersal ability and mode (i.e. active and passive) for instream and riparian communities shifted paradigms between mass effects and species sorting.     

Hatching phenology,life history and egg bank size of fairy shrimp Branchipodopsis spp. (Branchiopoda,Crustacea) in relation to the ephemerality of their rock pool habitat

In temporary aquatic habitats, permanence and the severe disturbance associated with desiccation are strong selective agents expected to lead to differentiation in life history strategies in populations experiencing different disturbance regimes. Besides optimal timing of hatching of dormant life stages, maturation and reproduction, pool inhabitants also benefit from the acquisition of reliable cues for the quality of the ambient environment. We investigated whether hatching patterns, life history characteristics and egg bank size of Branchipodopsis fairy shrimp (Branchiopoda, Anostraca) inhabiting a cluster of temporary rock pools in South Africa reflect variation in habitat stability and hatching cues. Long-term hydrological variation was used to select pools along a gradient of habitat stability. Initial conductivity was a good indicator for the length of inundations. No hatching occurred under elevated conductivities, which may present a mechanism to avoid abortive hatching. Egg bank size was unaffected by habitat size or habitat stability but instead was related to cover by a protective sheet of dry aquatic vegetation, which presumably counteracts egg bank erosion by wind when pools are dry. Life history but not hatching phenology reflected some aspects of habitat stability. Fairy shrimp populations in ephemeral pools started reproduction earlier than populations in more stable habitats. Additional common garden or transplant experiments, however, will be required to assess the relative importance of environmental and genetic components in explaining the observed variation and acquire more insight into the trade-offs that lie at the base of the evolution of life history strategies along the pond permanence gradient.     

Ecological,Dynamic and Taxonomic Problems Due to <Emphasis Type="Italic">Ludwigia</Emphasis> (Onagraceae) in France

We analyzed meta-community structure of bdelloid rotifers colonizing mosses along an 80 meter section of Rio Valnava in NW Italy. Bdelloid rotifers are small animals living associated with a substratum; colonization in bdelloids can be produced by active animals moving along the riverbed, or by passive dormant propagules, moved by wind. To detect which kind of colonization might be stronger at different spatial scales, we designed a spatially nested sampling experiment at three hierarchical levels: (1) single sample, (2) 10 communities inside each pool, (3) complete section of 10 pools. Assessing species richness and species similarity of communities, and coherence and nestedness of bdelloid meta-communities, we found that different forces may drive species composition at different spatial scales: at the largest scale, colonization of propagules may over-ride direct dispersal between pools, while at the scale of the single pool, differential movements of species give a nested structure to the meta-communities. The number of species increased as the level of analysis increased, even though this study was carried out along only a small stream section.