Disturbance and patch-specific responses: the interactive effects of woody debris and floods on lotic invertebrates

Disturbance may play an important role in generating patterns of abundance and distribution of biotic assemblages, particularly if its impact differs among habitat patches. Despite much speculation concerning the probable importance of spatial variation in the response of stream fauna to flooding, empirical work on patch-specific responses to spates is largely lacking. Floods typically reduce the abundance of lotic invertebrates dramatically in open-channel areas. We conducted a set of experiments to determine if faunal abundances are less affected in patches more sheltered due to the presence of woody debris dams. Specifically, we tested two hypotheses using chironomids and copepods living in a warmwater, 4th order stream: (1) the effect of flooding on the fauna varies between patches associated with debris dams versus the open channel, and (2) the absence of woody debris in a stream impedes faunal recovery throughout the channel following floods. We tested the first hypothesis by quantifying faunal abundances prior to, during, and following two floods in four patch types: mid-channel sandy patches distant from dams, coarse sediments associated with dams, fine sediments associated with dams, and leafy debris in dams. The second hypothesis was tested by removing all of the woody debris from two stretches of the stream and comparing the impact of a flood on fauna in debris-removed versus control stretches. Across all of the eight study dams, there were patchspecific faunal responses to two floods. Removal of woody debris from the stream did not prevent faunal recovery throughout the channel; however, the presence of woody debris dams did confer greater resistance of fauna to floods (as measured by no decrease in abundance during flooding) in two patch types. Abundances of chironomids and, to a lesser extent, copepods in the leafy debris of dams and in fine sediment patches associated with some dams either did not change or increased during floods, despite the fact that abundances in the dominant patch type of the stream (the sandy mid-channel) were reduced by 75–95%. All instances of faunal increase were limited to fine sediment patches associated with dams, thus entire dams cannot be labeled as flow refugia per se. Statistically, we distinguished fine patches which accumulated animals during floods from the other fine patches based on two physical attributes. Patches accumulating animals were all characterized by low water flux and nearbed flow, which likely contributed to the retention and/or passive deposition of animals. Whole dam attributes (e.g. dam size or complexity) were not useful in predicting which of the dams would accumulate animals in their fine sediments during flooding. Although structural complexity — here in the form of wood and leafy debris — is clearly important in generating biotic pattern in many ecosystems, our work underscores the need to understand what processes are responsible for the link between physical structure and biotic pattern.     

Small-sized invertebrates in a gravel stream: community structure and variability of benthic rotifers

1. The Rotifera assemblage inhabiting the streambed surface and the hyporheic zone of a gravel stream was investigated between October 1991 and October 1992. Forty-two species of Monogononta and 27 of Bdelloidea were identified. Within these two classes, dominant species differed between the surface and the hyporheic zone. At the streambed surface, the abundance of monogonont rotifers showed a seasonal pattern with significantly higher densities in pools, whereas bdelloids showed no clear temporal trend and did not differ significantly among sites. In the hyporheic zone, the depth distribution differed among the two rotifer groups, bdelloids occurred in highest densities between 0 and 30 cm sediment depth, while monogononts were most abundant at greater depths.
2. Species composition differed greatly between successive sampling dates (min. 5 to max. 26 days) at both the streambed surface and the hyporheos. At the streambed surface and in the shallow hyporheos a significantly higher percentage of species was replaced in riffles than in pools.
3. Few measured hydrophysical variables were associated with the Rotifera assemblage structure. At the streambed surface, species richness was negatively correlated with water temperature and substratum heterogeneity, and Monogononta rotifer densities declined with water depth and substratum roughness.
4. Permutation tests carried out on temporal serial correlations showed that, at riffle sites at the streambed surface, bdelloid rotifer densities, rotifer species richness and diversity did not differ significantly from a temporal, near-random pattern. The hyporheic rotifer assemblage followed similar near-random patterns.     

Habitat structure,resources and diversity: the separate effects of surface roughness and macroalgae on stream invertebrates

Habitat structure has pervasive effects on community composition and diversity, with physically complex habitats often containing more species than physically simple ones. What factors or mechanism drive this pattern is little understood, but a complicating problem is that different sources of habitat structure can be confounded in both surveys and experiments. In this study, we carried out an experiment in which two sources of habitat structure, attached macroalgae and substrate surface texture, were separately manipulated to discern their joint and separate effects upon the diversity and composition of colonizing macroinvertebrates in a stony, upland stream. Because stream algae vary markedly in abundance in both space and time, we also sampled the epilithon of stream stones at two spatial scales on eight dates over 2 years to gain some preliminary data on how stream algae vary between individual substrata over time. Experimental substrata had either a smooth (siltstones, sandstones, crystal-poor felsic volcanics, plain paving bricks) or rough (granodiorites, crystal-rich felsic volcanics, sand-blasted paving bricks) surface. We allowed these substrata to be colonized naturally by macroalgae, mostly the filamentous red alga Audouinella hermannii. Half of each of the rough and smooth substrata were selected at random and the macroalgae gently sheared off. All substrata were defaunated with a household insecticide with little field persistence, set out randomly through the study riffle, and invertebrates allowed to colonize them for 14 days. Some substrata were sampled immediately to check the efficacy of faunal and algal removals, which proved to be successful. Experimental results showed that both surface texture and macroalgae increase species richness independently of each other. Surface texture had no effect on densities, while macroalgae increased colonization densities, but rarefaction showed that both sources of habitat structure increased species richness above values expected simply on the basis of the numbers of colonists. However, reference stones with high macroalgal cover had the same species richness as those with low cover, suggesting that the effects of macroalgae on species richness are transient relative to those associated with surface texture. Epilithon samples taken at different times suggest that the magnitude of spatial variation in plant growth alters with time. If plants generally recolonize rough surfaces more quickly than smooth, then the effects of habitat structure on macroinvertebrates ought to be strongest after major disturbances during growing seasons of plants. Received: 1 September 1999 / Accepted: 10 January 2000     

The role of fungi in the nutrition of stream invertebrates

Dead leaves falling into streams are an important food source for many invertebrates. They are generally made more palatable and more nutritious if they are first colonized by aquatic hyphomycetes and other micro-organisms. At least two mechanisms appear to be responsible for this conditioning effect: microbial production (addition of easily digested microbial compounds to the nutritionally poor leaf substrate), and microbial catalysis (conversion of indigestible leaf substances into digestible subunits by microbial enzymes). Different invertebrate species vary in their ability to take advantage of microbial conditioning. This appears to be influenced by their mobility, the range of their food resources and their ability to overcome defense mechanisms of leaf-colonizing microorganisms.     

Disturbance history influences the distribution of stream invertebrates by altering microhabitat parameters: a field experiment

1. We investigated the effects of local disturbance history and several biotic and abiotic habitat parameters on the microdistribution of benthic invertebrates after an experimental disturbance in a flood‐prone German stream. 2. Bed movement patterns during a moderate flood were simulated by scouring and filling stream bed patches (area 0.49 m²) to a depth of 15–20 cm. Invertebrates were investigated using ceramic tiles as standardized substrata. After 1, 8, 22, 29, 36 and 50 days, we sampled one tile from each of 16 replicates of three bed stability treatments (scour, fill and stable controls). For each tile, we also determined water depth, near‐bed current velocity, the grain size of the substratum beneath the tile, epilithic algal biomass and standing stock of particulate organic matter (POM). 3. Shortly after disturbance, total invertebrate density, taxon richness and density of the common taxa Baetis spp. and Chironomidae were highest in stable patches. Several weeks after disturbance, by contrast, Baetis spp. and Hydropsychidae were most common in fill and Leuctra spp. in scour patches. The black fly Simulium spp. was most abundant in fill patches from the first day onwards. Community evenness was highest in scour patches during the entire study. 4. Local disturbance history also influenced algal biomass and POM standing stock at the beginning of the experiment, and water depth, current velocity and substratum grain size throughout the experiment. Scouring mainly exposed finer substrata and caused local depressions in the stream bed characterized by slower near‐bed current velocity. Algal biomass was higher in stable and scour patches and POM was highest in scour patches. In turn, all five common invertebrate taxa were frequently correlated with one or two of these habitat parameters. 5. Our results suggest that several ‘direct’ initial effects of local disturbance history on the invertebrates were subsequently replaced by ‘indirect’ effects of disturbance history (via disturbance‐induced changes in habitat parameters such as current velocity or food).     

Disturbance, biological legacies and community development in stream mesocosms

Disturbances reduce the biota in stream ecosystems, and leave biological legacies, including remnant species, which potentially influence post-disturbance community development but are poorly understood. We investigated whether three remnant species, the snail Radix peregra, the mayfly Serratella ignita and the freshwater shrimp Gammarus pulex, affected community development in mesocosms that mimicked disturbed habitat patches in streams. Following 21 days of colonisation, we found that the occurrence of legacy effects depended on the identity of the remnant species. Radix had the strongest effect. By bulldozing epilithon, the snails acted as ecological engineers that promoted settlement of filter feeders (Simuliidae) and invertebrate predators (especially Pentaneura and Aphelocheirus) and strongly deterred settlement of non-predatory chironomids (e.g. Heterotrissocladius and Microtendipes). Gammarus increased in density (by 665%) where remnant, probably through rapid reproduction. Baetis and Pentaneura were scarce, and Asellus absent, in remnant Gammarus treatments, as a consequence of interference and/or predation by the amphipods. In contrast, Serratella tolerated the colonisation of immigrant species and did not affect the structure of the developing benthic community. Despite the observed effects on the presence and abundance of benthos, remnant fauna had no significant effect on assemblage taxon richness, or that of any specific trophic group. The contrasting effects of remnant species on immigrant colonisation echoed differences in their life-history traits and foraging behaviours. Our results indicate that biota can generate spatial patchiness of epilithon and benthic invertebrates in stream ecosystems.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Surviving out in the cold: Antarctic endemic invertebrates and their refugia

Aim To identify Antarctic palaeoendemic taxa and their probable glacial refugia from regional groups of endemic species records. Location Antarctica. Methods We compiled a list of Antarctic non‐marine invertebrates from published literature, and then deleted all records relating to non‐endemic, zoochoric (phoretic and parasitic), marine and partially identified species to leave only the elements endemic to Antarctica. We then used cluster analysis and principal components analysis to identify regional groupings within this endemic fauna. Results Some 170+ of the reported 520+ Antarctic invertebrates are free‐living and endemic, but only nine of these are pan‐Antarctic, with the majority having either ‘continental’/eastern or ‘maritime’/western distributions. Main conclusions All invertebrates endemic to continental Antarctica are confined to, or found adjacent to, ice‐free palaeorefugial mountains, nunataks and coastal exposures. By contrast, only one maritime Antarctic palaeorefugium has been identified, and most endemic taxa are currently associated with coastal lowland neorefugia. We suggest which regions of Antarctica (1) are likely to be refugial, and (2) simply require more data in order that the nature and origin of their fauna can be elucidated.     

Covariation of stream community structure and biomass of algae, invertebrates and fish with forest cover at multiple spatial scales

1. To evaluate the spatial extent of the effects of forest cover on stream ecosystems, we measured algae, invertebrate, and fish biomass and invertebrate and fish community structure in 38 small first- to third-order streams in the National Capital Region of Canada along with forest cover at different spatial scales.
2. We considered 55 spatial scales of forest cover including several buffer widths (doubling 10–320 m) and lengths (doubling 10–1280 m, entire riparian distance upstream from sampling area) and entire catchments to determine which spatial scale maximized the correlation with biomass and metrics of community structure.
3. The proportion of variability in biomass and structural metrics explained by forest cover generally increased with increasing scale, suggesting that catchment-wide disturbances are the most influential determinants of benthic and fish communities.
4. Catchment forest cover explained more variation in algal (adjusted r ²   =   0.54), invertebrate (adjusted r ²   =   0.51) and fish (adjusted r ²   =   0.33) biomass than structural metrics of invertebrates and fish (adjusted r ²   =   0.08–0.27).
5. Analyses of the partial effects of forest cover at three scales (reach, riparian and the entire catchment) on biomass and community structure metrics identified catchment and reach scales as being most influential and never detected a significant partial effect of forest cover at the riparian scale.
6. These results suggest that maintenance or protection of reach and riparian buffers alone will not sufficiently protect stream function and structure from catchment-wide impacts.     

Long-term effects of local disturbance history on mobile stream invertebrates

The patchy distribution of benthic invertebrates in streams and rivers is an important and widely researched phenomenon. Previous studies on reasons for this patchiness have neglected the potential role of local disturbance history, probably because most lotic invertebrates are mobile and any effect of disturbance history was thought to be short-lived. Here we demonstrate for a New Zealand gravel-bed stream that local disturbance history can have long-term effects on the distribution of highly mobile stream invertebrates. Buried scour chains (100 at each of three 20-m sites within a 350-m reach) indicated that a spate with a return period of 5 months caused a mosaic of bed patches with different stabilities. More than 2 months after the spate, we took random, quantitative samples at each site from five patches that had experienced 4 cm or more of scour during the spate, from five patches with 4 cm or more of fill, and from five stable patches. Density of the dominant invertebrate taxon, the highly mobile mayfly Deleatidium spp., and densities of another three of the seven most common taxa differed significantly between patch stability categories. Larvae of Deleatidium, the black fly Austrosimulium spp. and the dipteran Eriopterini were most abundant in fill patches, whereas Isopoda were most abundant in scour patches. Total invertebrate densities and densities of six common taxa also differed between sites, although these were only 95–120 m apart. These results show that local disturbance history can have long-term effects on lotic invertebrates and be an important cause of invertebrate patchiness. The observed effects might have been even stronger had we sampled sooner after the spate or after a large flood. Disturbance history may influence invertebrates both directly (through dislodgement or mortality) and indirectly, through effects on the spatial distribution of their resources. Our results suggest that the role of disturbance in structuring animal communities dominated by mobile species may be more important than previously thought. Received: 25 January 2000 / Accepted: 14 April 2000     

Idiosyncratic relationships between pond invertebrates and environmental, temporal and patch-specific predictors of incidence

The incidence of eight species of microcrustacea and seven species of Dytiscidae (adults) was recorded from 30 small, temporary pools, in January and early summer, over six years. Incidence of each species, both in January and in summer, was modelled using logistic regression. Three categories of predictor variables were included in the models. Firstly systematic environmental variables; length of previous summer's dry phase, inter-pond links caused by flooding, distance between ponds and macrophyte structural density. Secondly temporal factors; year and previous incidence. Thirdly individual ponds, representing patch-specific effects. Logistic regression provided effective models of incidence for most species. However individual species varied in the factors that provided significant prediction of presence and absence. Systematic, temporal and patch-specific predictors were all significant predictors for different species of both microcrustacea and Dytiscidae. No single factor dominated most models. Individual factors showed positive or negative relationships with incidence with different species. Length of previous summer dry-phase was the most common predictor for microcrustacea and beetles in January. Summer models were more varied; length of previous dry phase and flood linkage between ponds and macrophyte density were significant predictors in many models, year and distance less important. Patch-specific predictors were significant for five of the microcrustacea and four of the beetles. The results suggest that individual species show idiosyncratic responses to systematic, temporal and patch-specific factors, rather than most species responding in a similar way to one or two dominant influences.     

The effects of electrofishing on the invertebrates of a Lake District stream

Summary The experiments were performed in Dale Park Beck, a stony stream in the English Lake District. Two operators electrofished the sampling area (length 20 m in April and July 1970, and 40 m in May 1971) three times (runs 1, 2, 3) in each experiment.Electrofishing caused a marked increase in the number of invertebrates drifting out of the sampling area, and nearly all taxa taken in the bottom samples were also found in the drift samples. The fish shocker was chiefly responsible for the increased drifting of Plecoptera, Ephemeroptera and Gammarus pulex, and these taxa were dislodged from the substratum more easily than Trichoptera, Coleoptera, Diptera and Polycelis felina. The increased drifting of the latter taxa was chiefly due to the disturbance of the substratum by the two operators.Most of the invertebrates drifting from the upstream end of the experimental section returned to the bottom within the sampling area. The invertebrate drift out of the sampling area came chiefly from the downstream end of the section, and was equivalent to a loss of only 5% from the total benthos in the sampling area (losses varied between <1 and 13% for individual taxa).     

The effects of predation and detritus on the structure of a stream insect community: a field test

Summary The role of predator density and plant detritus in determining patterns of insect abundance was tested in an Appalachian stream community (Reeds Creek, Pendleton Co, West Virginia). Insect colonization was followed over a 21-day period in field enclosures containing different densities of sculpins (Cottus bairdi and Cottus girardi). Sculpins caused no significant reductions in prey abundance; however, the combined effect of a guild of vertebrate predators caused a significant depression in the abundance of Chironomidae and the stonefly Leuctra. Since Chironomidae comprised approximately 85% of the total benthic fauna, vertebrate predation had an important role in the overall structure of the invertebrate community.Total insect abundance and diversity were correlated to the presence of plant detritus. However, this relationship was taxon specific, as abundance was highly correlated to detritus for only selected insect taxa. In general, abundance was best correlated with coarse particulate organic matter (CPOM), compared to fine particulate organic matter (FPOM) or total detritus (FPOM+CPOM+Whole leaf organic matter (LVOM)).This experiment illustrates that the influence of various potential structuring mechanisms is highly taxon dependent.     

Disturbance and trajectory of change in a stream fish community over four decades

Communities can change gradually or abruptly, and directionally (to an alternate state) or non-directionally. We briefly review the history of theoretical and empirical perspectives on community change, and propose a new framework for viewing temporal trajectories of communities in multivariate space. We used a stream fish dataset spanning 40 years (1969–2008) in southern Oklahoma, USA, emphasizing our own 1981–2008 collections which included well-documented, extreme drought and flood events, to assess dynamics of and environmental factors affecting the fish community. We evaluated the trajectory of the Brier Creek community in multivariate space relative to trajectories in 27 published studies, and for Brier Creek fish, tested hypotheses about gradual versus event-driven changes and persistence of shifts to alternate states. Most species were persistent, qualitatively, across the four decades, but varied widely in abundance, with some having unusually strong reproduction after extreme droughts. The community had an early period of relatively gradual and directional change, but greater displacement than predicted at random after two consecutive extreme droughts midway through the study (1998 and 2000). But, the community subsequently returned toward its former state in the last decade. This fish community is characterized by species that are tolerant of environmental extremes, and have life history traits that facilitate population recovery. The community appears “loosely stable” about a long-term average condition, but the impacts of the two consecutive droughts were substantial, and may foretell future dynamics of this or other communities in a changed global climate if disturbance events become more frequent or severe.     

Salamander diversity alters stream macroinvertebrate community structure

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Bioturbation by a dominant detritivore in a headwater stream: litter excavation and effects on community structure

Bioturbation can affect community structure by influencing resource distribution and habitat heterogeneity. Bioturbation by detritivores in small headwater streams could affect community structure by reintroducing buried detrital resources into the food web and could also affect the distribution of various taxa on detritus. We evaluated the ability of the caddisfly Pycnopsyche gentilis to uncover experimentally buried leaves in a headwater stream. Packs of leaves were placed in enclosures and covered with a known volume of sediment. We added 0, 3 or 6 large Pycnopsyche to the enclosures which were permeable to most other invertebrate taxa. Leaf packs were sampled after 23 days and leaf pack mass, the amount of sediment covering the leaf packs, and macro‐ and microinvertebrate densities on leaf packs were quantified. There was a significant negative relationship between Pycnopsyche density and leaf pack mass. Pycnopsyche also reduced the volume of sediment covering leaf packs. Pycnopsyche had complex effects on the abundance of invertebrate taxa associated with the leaves. Some taxa exhibited their highest abundance in the 3 Pycnopsyche treatment while others exhibited non‐significant increases as Pycnopsyche density increased. These results suggest that the beneficial effects of Pycnopsyche (e.g. uncovering leaves which increases the availability of habitat and food) outweigh any negative effects (e.g. disturbance, encounter competition) of the caddisfly when it is present at lower densities. However, the negative impacts of Pycnopsyche appear to outweigh the positive effects via sediment removal at higher caddisfly densities for some taxa. Our results suggest that bioturbating organisms in streams have the potential to reintroduce organic matter to detrital food webs and affect the distribution and abundance of benthic taxa associated with organic matter.     

亚热带森林转换对土壤无脊椎动物群落结构的影响

受人类活动干扰的增加,亚热带森林频繁转换为次生林和人工林,可能显著影响土壤无脊椎动物群落结构及其生态功能,但当前的认识并不一致。因此,于2022年7月调查了亚热带天然常绿阔叶林转换为次生林、米槠人工林、杉木人工林后土壤无脊椎动物群落结构特征。共捕获土壤无脊椎动物659只,丰度为26540只/m²,隶属1门6纲13目59科,其中蚁科和球角 虫 兆 科为优势类群。森林转换改变了土壤无脊椎动物群落组成和多样性。天然林向米槠人工林和杉木人工林转换后,土壤无脊椎动物丰度和类群均明显降低,其中大型土壤无脊椎动物丰度的响应更为敏感,在2种林型中分别显著降低了33.58%和36.53%。尽管林型转换对土壤无脊椎动物群落多样性指数无显著影响,但改变了土壤无脊椎动物群落组成,其中天然林与杉木人工林群落组成极不相似(J ＜ 0.25),等节 虫 兆 科为杉木人工林优势类群,占比达到59.84%。冗余分析显示,土壤湿度、凋落物现存量和凋落物磷含量是影响土壤无脊椎动物群落的主要因子,对土壤无脊椎动物群落的解释率为69.30%。可见,林型转换可能通过改变土壤理化性质和凋落物质量,调控土壤无脊椎动物群落结构。     

Low‐level effects of inert sediments on temperate stream invertebrates

1. The delivery, entrainment and deposition of inert fine sediments are among the most significant contributors to stream and river impairment worldwide. Associated ecological effects have been observed frequently, but specific experiments to identify sensitivity and avoidance behaviour in stream organisms are few, particularly in headwaters. 2. In a field‐experiment, we added fine sand at low levels (c. 4–5 kg m^?2) to 10 m reaches of two replicate headwater streams in the Usk catchment (Wales, U.K.) over two periods (autumn and summer). Upstream reaches were used as control in a classic before‐after‐control‐impact design. Invertebrate drift and benthic composition were measured for 2 days before and 1 day after sediment impact. 3. Sediment addition significantly increased overall drift density (by 45%) and propensity (by 200%), with effects largest on the night following addition rather than immediately (i.e. within 9 h). The mayflies Baetis rhodani, B. muticus and Ecdyonurus spp., simuliid and chironomid dipterans, and helodid beetles were the strongest contributors. 4. There were no marked effects on benthic composition, but density declined in treated reaches by 30–60%, particularly in B. rhodani, Ecdyonurus spp. and Leuctra hippopus + L. moselyi. 5. All effects were consistent between both seasons and streams. 6. These data show how even low‐level, short‐term, increases in fine sediment loading to upland, stony streams can reduce overall benthic density through increased drift. We suggest that the likely cause of the delayed drift response was a change in habitat quality which prompted avoidance behaviour. Longer‐term experiments are required to assess whether these effects reduce fitness or explain the losses of some types of organisms observed recently in sediment‐impaired reaches of this and other catchments.     

Size structure of the metazoan community in a Piedmont stream

We characterized the size structure of virtually the entire metazoan community in a fourth order, sandybottomed Piedmont stream during late summer. Our study, the first to sample across all habitat types and sizes of metazoans in an aquatic ecosystem, indicates that at the community level, stream size spectra may be bimodal for the benthos or trimodal when fish are included. Animals spanning 10 orders of magnitude in dry mass (from gastrotrichs to fish) were quantitatively collected from nine habitat types. The bimodal benthic size spectrum was characterized by a meiofaunal component (mostly oligochaetes and micro-crustacea) and a macrobenthic component (mostly the introduced asiatic clam, Corbicula fluminea). Insects contributed little to overall standing crop. Size-specific contribution to whole-community metabolism was assessed using allometric equations for respiration, and we found a distinctly bimodal distribution across the entire metazoan size range, with peaks in the meiofaunal and benthic macrofaunal size ranges. Our bimodal benthic size spectrum is similar to that observed for marine benthos but not to other freshwater benthic systems, possibly because the entire range of habitat types and/or animal sizes were not sampled in the latter. Numerous factors may influence size spectra in stream ecosystems, including local geomorphic (habitat) conditions, water level fluctuations, species introductions, and predation processes.     

Determinants of stream bryophyte community structure: bringing ecology into conservation

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