Do isolation and local habitat jointly limit the structure of stream invertebrate assemblages?

1. Both local and regional processes simultaneously control species assemblages depending on spatial habitat configuration. In dendritic networks like streams, the unique spatial arrangement of habitats produces various combinations of local habitat size and isolation. Stream invertebrate assemblages could therefore be controlled by different combinations of local and regional processes, depending on their location in the network. 2. Using quantile regression, we investigated how local habitat size, local environmental conditions and spatial isolation influenced variation in assemblage composition. Adult Trichoptera and benthic macroinvertebrate assemblages were represented by non‐metric multidimensional scaling (NMDS) ordination scores, as were local environmental conditions, in four headwater stream networks in New Zealand. 3. With increasing local habitat size, there was a decrease in variation in assemblage composition (NMDS scores) of both adult Trichoptera and benthic macroinvertebrates. This relationship between habitat size and assemblage variation was related to local habitat conditions at the upper limit of assemblage variability and spatial isolation at the lower limit of assemblage variability, for both adult Trichoptera and benthic assemblages, indicating joint local and regional controls on stream invertebrate assemblages. 4. The relationships between local assemblages and their neighbours, based on community similarity scores, differed between benthic macroinvertebrates and adult Trichoptera. For benthic assemblages, the larger the stream, the more similar assemblages were to neighbouring assemblages, whereas there was no consistent relationship between assemblage similarity and stream size for adult Trichoptera. This difference in structuring could be attributed to contrasting spatial influences linked to the different dispersal modes of adults and larvae. However, because adult and benthic assemblages are not independent, the influence of life stage on spatial distribution is difficult to determine (i.e. it is essentially a ‘chicken and egg’ argument). 5. Overall, our approach using quantile regression to evaluate limit responses, rather than regressions on means, has highlighted the joint importance of local habitat and spatial processes in structuring stream invertebrate assemblages. Furthermore, we have provided evidence for the importance of the spatial network arrangement and interactions between life stages and dispersal processes, in structuring stream assemblages.     

Supply-side ecology and benthic marine assemblages

Many marine invertebrates have a planktonic stage of their life history during which widespread dispersal and much mortality occur. The numbers surviving to recruit into habitats occupied by adults are therefore very variable in time and space. Models for the structure and dynamics of benthic assemblages tend to focus on processes causing death - often assuming consistent arrivals of recruits. Supply-side ecology is a newly fashionable term to describe recent interest in the long-realized consequences of variations in recruitment. Such variations have important influences on theory and empirical research in these assemblages.     

Post-settlement dispersal: the neglected link in maintenance of soft-sediment biodiversity

Seafloor integrity is threatened by disturbances owing to human activities. The capacity of the system to recover from disturbances, as well as maintain resilience and function, depends on dispersal. In soft-sediment systems, dispersal continues after larval settlement, but there are very few measurements of how far the post-settlers disperse in nature. Spatial scales of post-settlement dispersal are, however, likely to be similar to pelagic larval dispersal because of continued, frequent, small-scale dispersal over longer periods. The consequences of this dispersal may be more important for the maintenance of biodiversity and metacommunity dynamics than is pelagic larval dispersal, because of the greater size and competency of the dispersers. We argue that an increased empirical understanding of post-settlement dispersal processes is key for predicting how benthic communities will respond to local disturbances and shrinking regional species pools, with implications for monitoring, managing and conserving biodiversity.     

Patterns of benthic mega-invertebrate habitat associations in the Pacific Northwest continental shelf waters

As human impacts and demands for ocean space increase (fisheries, aquaculture, marine reserves, renewable energy), identification of marine habitats hosting sensitive biological assemblages has become a priority. Epifaunal invertebrates, especially the structure-forming species, are an increasing conservation concern as many traditional (bottom-contact fishing) and novel (marine renewable energy) ocean uses have the potential to displace or otherwise impact these slow-growing organisms. The differences in mega-invertebrate species assemblages between high-relief rocks and low-relief sediments are well documented and likely hold for most marine environments. In anticipation of potential development of marine renewable energy faculties off Oregon and Washington (USA), a survey of the benthic invertebrate assemblages and habitats was conducted on the continental shelf of the Pacific Northwest, using video footage collected by ROV, to more finely characterize these assemblage–habitat associations. Four main associations were found: pure mud/sand dominated by sea whips and burrowing brittle stars; mixed mud–rock (which may be further divided based on size of mixed-in rocks) characterized by various taxa at small densities; consolidated rocks characterized by high diversity and density of sessile or motile mega-invertebrates; and rubble rocks showing less diversity and density than the consolidated rocks, possibly due to the disturbance generated by movement of the unconsolidated rocks. The results of this study will help classify and map the seafloor in a way that represents benthic habitats reflective of biological species assemblage distributions, rather than solely geological features, and support conservation and management planning.     

Environmentally-controlled,density-dependent secondary dispersal in a local estuarine crab population

The mechanisms driving the pelagic secondary dispersal of aquatic organisms following initial settlement to benthic habitats are poorly characterized. We examined the physical environmental (wind, diel cycle, tidal phase) and biological (ontogenetic, density-dependent) factors that contribute to the secondary dispersal of a benthic marine invertebrate, the blue crab (Callinectes sapidus) in Pamlico Sound, NC, USA. Field studies conducted in relatively large (0.05 km²) seagrass beds determined that secondary dispersal is primarily undertaken by the earliest juvenile blue crab instar stages (J1 crabs). These crabs emigrated pelagically from seagrass settlement habitats using nighttime flood tides during average wind conditions (speed ~5 m s^–1). Moreover, the secondary dispersal of J1 crabs was density-dependent and regulated by intra-cohort (J1) crab density in seagrass. Our results suggest that dispersal occurs rapidly following settlement, and promotes blue crab metapopulation persistence by redistributing juveniles from high-density settlement habitats to areas characterized by low postlarval supply. Collectively, these data indicate that blue crab secondary dispersal is an active process under behavioral control and can alter initial distribution patterns established during settlement. This study highlights the necessity of considering secondary dispersal in ecological studies to improve our understanding of population dynamics of benthic organisms.     

Comparison of stream benthic invertebrate assemblages among forest types in the temperate region of Japan

We compared benthic invertebrate assemblages among headwater streams in several forest types in Japan. Forests were divided into three clusters based on vegetation composition: old-growth broad-leaved forest, planted coniferous forest, and mixed forest. The numbers of individuals and families and the diversity (Shannon-Wiener) of benthic invertebrate assemblages did not differ significantly among the three forest clusters. However, principal components analysis of family abundance showed differences in the benthic invertebrate assemblages among the three forest clusters. No environmental factors were correlated with these differences. Benthic invertebrate assemblages differed depending on forest composition. The abundances of Taeniopterygidae and Athericidae in old-growth broad-leaved forest were significantly greater than in planted coniferous forest. The abundances of Heptageniidae, Baetiidae, Stenopsychidae, Uenoidae, Chironomidae, and Potamidae in planted coniferous forest were significantly greater than in old-growth broad-leaved forest. If the remaining old-growth broad-leaved forest were to be converted to coniferous plantation, species that inhabit old-growth forest may become extinct.     

Dispersal and population expansion in a direct developing marine snail (Littorina saxatilis) following a severe population bottleneck

Most marine benthic invertebrate species have planktonic larvae, and in species in which juveniles and adults have low vagility a larva is obviously an efficient way of active dispersal. A minority of benthic invertebrate species develop without any pelagic phase at all. A largely unsolved question is how and at what rate do these species disperse. We have addressed this question using the marine littoral snail Littorina saxatilis (Olivi) as an example of a species that completely lacks larval dispersal. In the Koster archipelago (north part of the Swedish west coast), L. saxatilis occupies rocky island habitats of different sizes, from large islands to small intertidal skerries (islets). In 1988 an extremely dense bloom of a toxin-producing flagellate killed more than 99% of this snail species in this area. Populations of larger islands were reduced, often to less than 1%, but were restored over 2–4 yr. In contrast, populations of small intertidal skerries were completely wiped out and thus could not increase by local recruitment. Four years later, however, four of 33 skerries (12%) were successfully recolonized with relatively dense populations, and another five had received a few founder individuals. These results indicate recruitment through founder individuals, and are rough estimates of dispersal rate in a snail species that lacks a pelagic developmental stage.     

Concordance among stream assemblages and spatial autocorrelation along a fragmented gradient

Patterns of spatial autocorrelation of biota and distributional similarity (concordance) between assemblages of different organism groups have important implications in both theoretical ecology and biodiversity conservation. Here we report environmental gradients and spatial distribution patterns of taxonomic composition among stream fish, benthic macroinvertebrate, and diatom assemblages along a fragmented stream in south‐western France. We quantified spatial patterns of lotic assemblage structure along this stream, and we tested for concordance in distribution patterns among the three taxonomic groups. Our results showed that both environmental characteristics and stream assemblages were spatially autocorrelated. For stream fish and diatom assemblages, these patterns reflected assemblage changes along the longitudinal stream gradient, whereas environmental variables and benthic macroinvertebrates exhibited a more patchy spatial pattern. Cross‐taxa concordance was significant between stream fish and diatoms, and between stream fish and benthic macroinvertebrates. The assemblage concordance between stream fish and diatoms could be attributed to similar responses along the longitudinal gradient, whereas those between stream fish and benthic macroinvertebrates may result from biotic interactions. Based on potential dispersal capacities of taxa, our results validated the hypotheses that weakly dispersing taxa exhibit greater concordance than highly dispersing ones and that dispersal capacities affect how taxonomic groups respond to their local environment. Both diatoms and highly dispersing stream fish were affected by stream fragmentation (i.e. the number of dams between sites), while the effect of fragmentation was not significant for invertebrates that fly well in their adult stage, thus emphasizing the importance of the way of dispersal. These results suggest that addressing the effects of dispersal capacity on stream assemblage patterns is crucial to identifying mechanisms behind patterns and to better understanding the determinants of stream biodiversity.     

Fish foraging effects on benthic assemblages along a warm-temperate stream: differences among drift feeders, benthic predators and grazers

In streams, physical and biotic conditions change from the headwaters to the mouth, shaping longitudinal patterns in community structure. We examined how fish foraging effects on periphyton and benthic invertebrates changed along a longitudinal gradient of a warm-temperate stream in southwestern Japan. We established three study sites according to changes in the composition of fish feeding guilds (upper site characterized by drifting-invertebrate feeders, Oncorhynchus masou ; middle site by benthic invertebrate feeders, Rhinogobius spp.; lower site by the presence of periphyton grazers, Sicyopterus japonicus ), and performed two manipulative experiments to examine effects of different fish assemblages on periphyton and benthic invertebrate abundances. Results of an exclosure experiment suggested that fishes had no effect on the benthic assemblages at the upper and middle sites whereas fishes reduced the abundances of both periphyton and invertebrates on stone surfaces at the lower site, where both benthic invertebrate feeders and grazers inhabited. A subsequent enclosure experiment showed that the reduction of invertebrate densities at the lower site was caused by the grazers rather than benthic invertebrate feeders. These experimental results suggested that effects of fishes on benthic assemblages are intensified downstream, owing to the occurrence of the grazing fish. Furthermore, observational data based on field sampling suggested that such grazing effects were reflected in longitudinal patterns in periphyton and invertebrate abundances. Overall results emphasize an important role of the grazing fish ( S. japonicus ) in shaping longitudinal patterns in benthic assemblage structure.     

Spatial autocorrelation of assemblages of benthic invertebrates and its relationship to environmental factors in two upland rivers in southeastern Australia

The nature of spatial autocorrelation of biota may reveal much about underlying ecological and biological factors responsible for producing those patterns, especially dispersal processes (drift, adult flight, etc.). We report here on assemblage‐level autocorrelation in the benthic‐invertebrate assemblages (retained in sieves of 300 µm mesh) of riffles in two adjacent, relatively pristine rivers in southeastern Victoria, Australia (40‐km reaches of the Wellington and Wonnangatta Rivers). These are related to patterns of autocorrelation in physical and catchment conditions (‘environmental variables’) in the vicinity of the sampling points. Both the invertebrate assemblages and environmental variables were autocorrelated at small scales (= 8 km) in the Wellington River in one of the sampling years (1996). Dissimilarities of invertebrate assemblages were correlated with dissimilarities of environmental variables in both sampling years (1996 and 1997) in that river. Environmental variables were autocorrelated in the Wonnangatta River, but this was not expressed as autocorrelation in the assemblages of invertebrates, which were not autocorrelated at any scale studied. Individual environmental variables showed different spatial patterns between the two rivers. These results suggest that individual rivers have their own idiosyncratic patterns and one cannot assume that even similar, geographically adjacent rivers will have the same patterns, which is a difficulty for ecological assessment and restoration.     

Macrozoobenthos of the Small Aral Sea

Benthic assemblages of the Small Aral Sea were studied in 1999. The species composition, density, and biomass of benthic invertebrates were determined. The vertical distribution pattern and its dynamics in the course of desiccation of the sea were analyzed. It is shown that the vertical distribution of the benthos has changed substantially over the last 30 years. The biomass of the benthos in the inshore zone increased, while in the central part of the sea, benthic invertebrate populations declined and gradually disappeared. These changes are likely to be due to the accumulation of organic matter in the bottom sediments and the weakening of hydrodynamic processes, which could lead to oxygen depletion in deep water. In the 1990s, the benthic assemblages of the Small Aral Sea were relatively stable and highly productive. It is concluded that the Aral Sea is a promising area for fishery restoration.     

A Hierarchical Classification of Benthic Biodiversity and Assessment of Protected Areas in the Southern Ocean

An international effort is underway to establish a representative system of marine protected areas (MPAs) in the Southern Ocean to help provide for the long-term conservation of marine biodiversity in the region. Important to this undertaking is knowledge of the distribution of benthic assemblages. Here, our aim is to identify the areas where benthic marine assemblages are likely to differ from each other in the Southern Ocean including near-shore Antarctica. We achieve this by using a hierarchical spatial classification of ecoregions, bathomes and environmental types. Ecoregions are defined according to available data on biogeographic patterns and environmental drivers on dispersal. Bathomes are identified according to depth strata defined by species distributions. Environmental types are uniquely classified according to the geomorphic features found within the bathomes in each ecoregion. We identified 23 ecoregions and nine bathomes. From a set of 28 types of geomorphic features of the seabed, 562 unique environmental types were classified for the Southern Ocean. We applied the environmental types as surrogates of different assemblages of biodiversity to assess the representativeness of existing MPAs. We found that 12 ecoregions are not represented in MPAs and that no ecoregion has their full range of environmental types represented in MPAs. Current MPA planning processes, if implemented, will substantially increase the representation of environmental types particularly within 8 ecoregions. To meet internationally agreed conservation goals, additional MPAs will be needed. To assist with this process, we identified 107 spatially restricted environmental types, which should be considered for inclusion in future MPAs. Detailed supplementary data including a spatial dataset are provided.     

Ecological and evolutionary significance of dispersal by freshwater invertebrates

Traditional expectations for how widely and how often freshwater invertebrates disperse differ from empirical data. Freshwater invertebrates have been characterized as frequent, widespread dispersers, particularly those that are transported passively. Our review finds that this characterization may describe the potential for dispersal in some taxa, but it is not an accurate generalization for actual dispersal rates. High variance among habitats and taxonomic groups is a consistent theme. Advances in population genetics may help resolve these issues, but underlying assumptions should be carefully tested. Further, even unbiased estimates of gene flow may not equate with individual movement, because not all dispersers survive and reproduce. Some freshwater invertebrates may exist in classic Levins metapopulations. However, other species fit into a broader metapopulation definition, where temporal dispersal via diapause is functionally equivalent to spatial dispersal. In the latter case, local extinctions and rescue effects may be rare or absent. Finally, limited dispersal rates in many taxa suggest that theories of freshwater community assembly and structure can be made more robust by integrating dispersal and local processes as joint, contingent regulators. Recent research on freshwater invertebrate dispersal has substantially advanced our basic and applied understanding of freshwaters, as well as evolutionary ecology in general.     

Patterns of benthic mega-invertebrate habitat associations in the Pacific Northwest continental shelf waters: a reassessment

As human impacts and demands for ocean space increase (fisheries, aquaculture, marine reserves, renewable energy), identification of marine habitats hosting sensitive biological assemblages has become a priority. Epifaunal invertebrates, especially the structure-forming species, are an increasing conservation concern as many traditional (bottom-contact fishing) and novel (marine renewable energy) ocean uses have the potential to displace or otherwise impact these slow-growing organisms. The differences in mega-invertebrate species assemblages between high-relief rocks and low-relief sediments are well documented and likely hold for most marine environments. In anticipation of potential development of marine renewable energy faculties off Oregon and Washington (USA), a survey of the benthic invertebrate assemblages and habitats was conducted on three rocky reefs on the continental shelf of the Pacific Northwest, using video footage collected by remotely operated vehicle, to more finely characterize these assemblage–habitat associations. Benthic assemblages appeared to first group by depth (50–80 vs. 100–120 m), then by relief (consolidated rocks vs. unconsolidated rocks and soft sediments). Consolidated rocks were characterized at each site by a combination of various sponges, gorgonians, sea anemones and echinoderms; unconsolidated rocks were characterized at Grays Bank by sea anemones and burrowing brittle stars, and at Bandon-Arago by sponges and echinoderms; soft sediments were characterized at Grays Bank and Siltcoos Reef by sea whips and burrowing brittle stars, as well as pink shrimps and sea stars at Siltcoos Reef, and at Bandon-Arago by sponges, gorgonians and echinoderms. The results of this study will help classify and map the seafloor in a way that represents benthic habitats reflective of biological species assemblage distributions, rather than solely geological features, and support conservation and management planning.     

Dispersal in drift-prone macroinvertebrates: a case for density-independence

1. Studies of dispersal of macroinvertebrates in streams and rivers tend to be focused on drift, whilst benthic movements are usually considered to be less important.
2. Field-enclosure experiments with the mayfly Baetis rhodani indicate that net dispersal in this species is simply a proportional loss of individuals from the benthos.
3. Neither net upstream or downstream movements exhibited evidence of density-dependence in the form of curvilinear relationships between initial and final densities.
4. The net number of animals moving upstream did not differ significantly from the net number moving downstream.
5. The probable mechanisms behind density-independent dispersal are discussed, as are the implications for our understanding of population dynamics in relation to invertebrate drift.     

Development of a microplate-based fluorescence immunoassay using quantum dot streptavidin conjugates for enumeration of putative marine bacteria, Alteromonas sp., associated with a benthic harpacticoid copepod

Attached bacteria inhabit the surfaces of many marine animals--a process that may play important roles in the survival and transport through aquatic systems. However, efficient detection of these bacteria has been problematic, especially small aquatic animals such as benthic harpacticoid copepod. Quantum dots (QD) have recently emerged as a significant tool in immunofluorescence detection because of their unique properties compared to other fluorescent probes. In the present study, a polyclonal antibody was raised against the Gram-negative marine bacterium, Alteromonas sp. A microplate-based immunofluorescence bioassay using QD strepavidin conjugates was developed for quantifying putative Alteromonas sp. cells located on the surfaces of a marine harpacticoid copepod, Microarthridion littorale. The number of attached Alteromonas sp. was estimated to be 10(2)+/-8 CFU using this method. The QD approach, coupled to a microplate assay can potentially provide an efficient and accurate method for rapidly detecting multiple bacteria species attached to small invertebrate animals because of their unique excitation and emission characteristics.     

The influence of scale and geography on relationships between stream community composition and landscape variables: description and prediction

SUMMARY 1. We analysed an existing database of macroinvertebrates and fish in the context of a newly established geographical information system (GIS) of physical features to determine the relationships between stream community composition and physical factors measured at three landscape scales – catchment, reach and bedform. Both an exploratory (concordance analysis) and a predictive ( ausrivas ) approach were used.
2. The environmental variables that most successfully accounted for variation in macroinvertebrate assemblages were mainly 'natural' and at the catchment-scale (relief ratio, basin diameter, etc.) but the human-induced physical setting of percentage of pasture in the riparian zone was also influential. For fish, 'natural' variables were also dominant, but these were mostly at the bedform or reach scales and land use featured strongly.
3. Geographic location accounted for some of the variation in invertebrate assemblages, partly because geography and influential conditions/resources are correlated but also because different species may have evolved in different places and have not colonised every 'ecologically appropriate' location. Geographic location was not influential in accounting for variation in assemblages of strongly flying invertebrates, supporting the hypothesis that organisms having high dispersal potential can be expected to break down geographic barriers more readily than those with poor dispersal powers. In accord with what is known about the local evolution and restricted distributions of native and exotic species, history (reflected in geography) appeared to account for some variation in fish assemblages.
4. Given their different mathematical bases, the fact that exploratory and predictive analyses yielded similar results provides added confidence to our conclusions.     

描述Meta-种群动态的耦合映象格子模型

Ｍｅｔａ－种群观点为在局部种群之上的空间尺度上描述种群的生态学过程提供了一种途径。但是由于植物所拥有的若干特殊性质（如种子休眠、有限扩散和局部适应）使得对某些植物种使用原有的Ｍｅｔａ－种群概念和模型存在许多困难。因此,为了能更精确地反映植物Ｍｅｔａ－种群动态,本文将Ｍｅｔａ－种群动态分解为斑块内局部种群动态和斑块间的扩散过程两个分量。用Ｌｏｇｉｓｔｉｃ方程表述每个斑块内的局部种群动态,用扩散系数ε表示斑块间的扩散,建立了描述Ｍｅｔａ－种群动态的耦合映象格子模型。对单峰映象ｘｎ＋１＝１－ａｘ２ｎ给出了耦合映象格子的时空行为,这对Ｍｅｔａ－种群动态的研究可能是非常重要的。     

Latitudinal gradient of nestedness and its potential drivers in stream detritivores

Understanding what mechanisms shape the diversity and composition of biological assemblages across broad‐scale gradients is central to ecology. Litter‐consuming detritivorous invertebrates in streams show an unusual diversity gradient, with α‐diversity increasing towards high latitudes but no trend in γ‐diversity. We hypothesized this pattern to be related to shifts in nestedness and several ecological processes shaping their assemblages (dispersal, environmental filtering and competition). We tested this hypothesis, using a global dataset, by examining latitudinal trends in nestedness and several indicators of the above processes along the latitudinal gradient. Our results suggest that strong environmental filtering and low dispersal in the tropics lead to often species‐poor local detritivore assemblages, nested in richer regional assemblages. At higher latitudes, dispersal becomes stronger, disrupting the nested assemblage structure and resulting in local assemblages that are generally more species‐rich and non‐nested subsets of the regional species pools. Our results provide evidence that mechanisms underlying assemblage composition and diversity of stream litter‐consuming detritivores shift across latitudes, and provide an explanation for their unusual pattern of increasing α‐diversity with latitude. When we repeated these analyses for whole invertebrate assemblages of leaf litter and for abundant taxa showing reverse or no diversity gradients we found no latitudinal patterns, suggesting that function‐based rather than taxon‐based analyses of assemblages may help elucidate the mechanisms behind diversity gradients.     

Are native cyprinids or introduced salmonids stronger regulators of benthic invertebrates in South African headwater streams?

The introduction of nonnative salmonids in the Southern Hemisphere generally leads to a reduction in invertebrate abundance and changes in assemblage composition. In the Cape Floristic Region of South Africa, introduced rainbow trout Oncorhynchus mykiss is the dominant predator in many headwater streams, where they have replaced small‐bodied native fishes such as Breede River redfin Pseudobarbus burchelli. To examine the consequences of this species replacement on food web structure, we used a month‐long field experiment to compare the top‐down effects of Breede River redfin and rainbow trout on benthic invertebrate assemblages (abundance and composition) and basal resources (periphyton and particulate organic matter) in 1 × 1.5 m of plastic cages. Benthic invertebrate abundance was more strongly depleted in the cages with redfin than in the cages with trout, and redfin and trout had distinct effects on invertebrate assemblage composition. On the other hand, neither redfin nor trout had a significant influence over standing stocks of periphyton or organic matter, implying that their differential effects on benthic invertebrates did not cascade down to the base of the stream food web in our experiment. Gut content analysis showed that aquatic invertebrates contributed more to the diet of redfin, while terrestrial invertebrates contributed more to the diet of trout, which may be responsible for the relatively weak effect of trout on aquatic invertebrates. This pattern contrasts with nonnative salmonid impacts elsewhere in the Southern Hemisphere. That trout can strongly alter the structure of benthic invertebrate assemblages, in addition to severely depleting native fish abundance, in Cape Floristic Region headwater streams should be weighed into management decisions, and our findings highlight the need for a detailed understanding of species‐specific top‐down effects where native predators are replaced by invasive predators.