Macroinvertebrate community response to natural and forest harvest gradients in western Oregon headwater streams

1. To examine the effects of forest harvest practices on headwater stream macroinvertebrates, we compiled a 167 site database with macroinvertebrate, fish, physical habitat and catchment land cover data from the three forested ecoregions in western Oregon. For our analysis, headwater streams were defined by catchment areas <10 km² and perennial water during summer low flows. Almost all sites in the database were selected using a randomised survey design, constituting a representative sample of headwater streams in these ecoregions. 2. Macroinvertebrate taxonomic and functional feeding group composition were very similar among the three ecoregions in the study area (Coast Range, Cascades and Klamath Mountains). On average, 55% of the individuals at each site were in the orders Ephemeroptera, Plecoptera or Trichoptera. Dipteran taxa (mostly chironomids) accounted for another 34%. At almost all sites, non‐insects made up <10% of the macroinvertebrate assemblage. Almost half (49%) of the assemblages were collectors; remaining individuals were about evenly divided among scrapers, shredders and predators. 3. There were 189 different macroinvertebrate taxa at the 167 sites with richness at individual sites ranging from 7 to 71 taxa. Ordination by non‐metric multidimensional scaling revealed a strong association between % Ephemeroptera, especially Baetis, and site scores along the first axis. This axis was also strongly related to % coarse substratum and fast water habitat. The second axis was strongly related to % intolerant individuals, site slope and altitude. No strong relationships were evident between any ordination axis and either logging activity, presence/absence of fish, catchment size or ecoregion. 4. Based on macroinvertebrate index of biotic integrity (IBI) scores, 62% of the sites had no impairment, 31% of the sites had slight impairment and only 6% of the sites had moderate or severe impairment. IBI scores were not strongly related to forest harvest history. All four severely impaired sites and five of the seven sites with moderate impairment were lower altitude, shallower slope stream reaches located in the Coast Range with evidence of agricultural activity in their catchment or riparian zone. % sand + fine substratum was the environmental variable most strongly related to macroinvertebrate IBI.     

Self-organisation in streams: the relationship between movement behaviour and body condition in a headwater salamander

1. When movement behaviour is correlated with traits affecting fitness, it may affect population performance directly, independent of extrinsic habitat conditions. 2. In a previous 3‐year, capture–recapture study, upstream movement by the salamander Gyrinophilus porphyriticus compensated for low reproduction in the upper 500 m of Merrill Brook, a first‐order stream in northern New Hampshire (U.S.A.). This immigrant subsidy resulted from excess reproduction in the downstream section (i.e. the lower 500 m of stream length) and from consistently upstream‐biased movement by salamanders. Reproduction in the two stream sections was positively correlated with mean body condition. Using 6 years of capture–recapture data, this study examines whether the movement behaviour of G. porphyriticus is related to body condition and thereby directly influences mean body condition and reproduction in the two sections of Merrill Brook. 3. Upstream‐biased movement and greater mean body condition in the downstream section were consistent across 6 years of data collection. In Merrill Brook and four other streams, however, individuals with high body condition were more likely to move upstream and low‐condition individuals were more likely to move downstream. Movement direction was unrelated to the size, sex and initial location of individuals. Body condition was positively related to growth rate, further supporting its link to reproductive potential, and positively autocorrelated through time in individuals that moved. 4. Results of this 6‐year study suggest that the movement behaviour of G. porphyriticus partially compensated for environmental factors differentiating mean body condition and reproduction along Merrill Brook and illustrate the potential for this form of self‐organisation to occur in linear habitats such as streams and rivers.     

Herbs and grasses as an allochthonous resource in open-canopy headwater streams

1. The organic matter dynamics of streams dominated by herbs and grass on their banks are poorly understood, despite the fact that such streams are common worldwide. Further, herbs and grasses can provide large quantities of detritus to stream food webs, and particularly small streams can be heavily shaded by overhanging vegetation, perhaps limiting in‐stream primary production. 2. We quantified the standing crop of edge vegetation and associated macroinvertebrate communities along three headwater streams with herbaceous and grass riparian vegetation on agricultural land in the Piedmont of Maryland, U.S.A., measured the decomposition of four common species of herbs and grasses using experimental leaf packs, and removed edge vegetation experimentally to determine the effect of shading on benthic algal production. 3. Large standing crops of plant material (average range: 68–276 g ash‐free dry mass per m⁻²), composed largely of monocotyledons, were found at all three study streams. These values are similar to those for coarse particulate organic matter in deciduous forested streams in the eastern U.S.A. In addition, diverse assemblages of shredding macroinvertebrates were observed at all three study sites. 4. Decomposition of the herbs was faster than that of the grasses, and both decomposed faster than most deciduous tree leaf litter. The decomposition rates of the herbs and grasses were significantly related to leaf quality as measured by leaf nitrogen content. Macroinvertebrate shredders colonized all experimental leaf packs, and the colonization of the herbs was faster than that of the grasses. 5. The accrual of chlorophyll‐a after the removal of shading vegetation was faster than that measured prior to removal as well as that in an unmanipulated control reach. 6. Given that the standing crop of organic matter in streams with herbs and grass along their banks was similar to that in forested streams, that the organic matter was rich in nitrogen and used by detritivores, and riparian shading limited algal growth, we suggest that herbaceous and grass plant material may be an important allochthonous food resource in such systems.     

Macroinvertebrate communities of non-glacial high altitude intermittent streams

1. Macroinvertebrate assemblages of five non‐glacial intermittent high altitude headwater streams (above 1400 m – Serra da Estrela, Portugal), with dry periods of different lengths (0–3 months), were investigated in nearly undisturbed conditions to (i) examine spatial differences and identify environmental variables responsible for the observed invertebrate patterns, (ii) assess the association of dry period length with invertebrate community structure and (iii) determine the influence of using different taxonomic identification levels (order, family and genus) to assess invertebrate community patterns. 2. More than 100 macroinvertebrate genera were identified. Insects clearly dominated these communities with more than 95% of total captures and around 95% of the total richness. Diptera were the most rich and abundant group with chironomid occurrences comprising over 70% of macroinvertebrate captures. 3. The highest taxon richness, diversity, EPT (Ephemeroptera + Plecoptera + Trichoptera) and OCH (Odonata + Coleoptera + Heteroptera) genus richness, the greatest number of exclusive and characteristic taxa identified by the Indicator Value (IndVal), and a distinct community structure shown by Canonical Correspondence Analyses (CCA), were found in the only stream that was never totally dry, with pools lasting over summer. Environmental gradients that spatially structured the macroinvertebrate communities were always related to flow variations. 4. Over time, the highest abundances found in these systems were also related to flow variations and maximum genus richness occurred in the connected pools or in isolated pools. Streams with longer dry periods presented a distinct recolonization phase, with higher abundance of the stonefly larvae Nemoura sp. and the presence of the chironomid larvae Krenosmittia sp., possibly arriving from the hyporheos. 5. Taxonomic level of invertebrate identification was vital for recognizing the characteristic taxa (IndVal) of streams yet was not critical for identifying streams with the highest macroinvertebrate richness/diversity or structuring environmental gradients. 6. Overall, this study emphasizes the variability of high altitude intermittent streams macroinvertebrate communities, despite spatial proximity. This variability was probably related to flow intermittency and hydrologic permanence, different vegetation covers and riverbed substrata. Consequently, the establishment of reference conditions should involve long‐term data collections and more detailed physical characterization. Also, these findings have significant implications for accurately predicting the ecological consequences of future climate change in high altitude scenarios.     

Loss of genetic diversity in the North American mayfly Ephemerella invaria associated with deforestation of headwater streams

1. Terrestrial dispersal by aquatic insects increases population connectivity in some stream species by allowing individuals to move outside the structure of the stream network. In addition, individual survival and reproductive success (as well as dispersal) are tightly linked to the quality of the terrestrial habitat. 2. In historically forested catchments, deforestation and altered land use have the potential to interfere with mayfly dispersal or mating behaviours by degrading the quality of the terrestrial matrix among headwater streams. We hypothesised that loss of tree cover in first‐order catchments would be associated with an increase in population substructure and a decrease in genetic diversity of mayfly populations. 3. To test this hypothesis, we investigated spatial patterns of genetic variation in the common mayfly Ephemerella invaria across a gradient of deforestation in the central piedmont region of eastern United States. Intraspecific genetic diversity and population substructure were estimated from data obtained using fluorescent amplified fragment length polymorphism (AFLP) markers. 4. We found that mayfly populations had low population substructure within headwater stream networks and that genetic diversity was strongly negatively correlated with mean deforestation of the first‐order catchments. The large‐scale pattern of population substructure followed a pattern of isolation by distance (IBD) in which genetic differentiation increases with geographical distance, but assignment tests placed a few individuals into populations 300 km away from the collection site. 5. Our results show that loss of genetic diversity in this widespread aquatic insect species is co‐occurring with deforestation of headwater streams. 6. Most arguments supporting protection of headwater streams in the United States have centred on the role of these streams as hydrological and biogeochemical conduits to downstream waters. Our work suggests that headwater stream land use, and specifically tree cover, may have a role in the maintenance of regional genetic diversity in some common aquatic insect species.     

Export of invertebrates and detritus from fishless headwater streams in southeastern Alaska: implications for downstream salmonid production

1. We examined the export of invertebrates (aquatic and terrestrial) and coarse organic detritus from forested headwaters to aquatic habitats downstream in the coastal mountains of southeast Alaska, U.S.A. Fifty‐two small streams (mean discharge range: 1.2–3.6 L s^?1), representing a geographic range throughout southeast Alaska, were sampled with 250‐μm nets either seasonally (April, July, September) or every 2 weeks throughout the year. Samples were used to assess the potential subsidy of energy from fishless headwaters to downstream systems containing fish. 2. Invertebrates of aquatic and terrestrial origin were both captured, with aquatic taxa making up 65–92% of the total. Baetidae, Chironomidae and Ostracoda were most numerous of the aquatic taxa (34, 16 and 8%, respectively), although Coleoptera (mostly Amphizoidae) contributed the greatest biomass (30%). Mites (Acarina) were the most numerous terrestrial taxon, while terrestrial Coleoptera accounted for most of the terrestrial invertebrate biomass. 3. Invertebrates and detritus were exported from headwaters throughout the year, averaging 163 mg invertebrate dry mass stream^?1 day^?1 and 10.4 g detritus stream^?1 day^?1, respectively. The amount of export was highly variable among streams and seasons (5–6000 individuals stream^?1 day^?1 and <1–22 individuals m^?3 water; <1–286 g detritus stream^?1 day^?1 and <0.1–1.7 g detritus m^?3 water). Delivery of invertebrates from headwaters to habitats with fish was estimated at 0.44 g dry mass m^?2 year^?1. We estimate that every kilometre of salmonid‐bearing stream could receive enough energy (prey and detritus) from fishless headwaters to support 100–2000 young‐of‐the‐year (YOY) salmonids. These results illustrate that headwaters are source areas of aquatic and terrestrial invertebrates and detritus, linking upland ecosystems with habitats lower in the catchment.     

Growth and population size of crayfish in headwater streams: individual- and higher-level consequences of acidification

1. Environmental stress may have indirect positive effects on population size through modification of food‐web interactions, despite having negative effects on individuals. Here we evaluate the individual‐ and population‐level effects of acidification on crayfish (Cambarus bartonii) in headwater streams of the Allegheny Plateau (PA, U.S.A.) with field experiments and survey data. Median baseflow pH of 24 study reaches in nine streams varied from 4.4 to 7.4, with substantial variation found both among and within streams. 2. Two bioassays were conducted to evaluate the relationship between stream pH and crayfish growth rates. Growth rates were always higher in circumneutral reaches than in acidic reaches. Crayfish originating in acidic water grew less when transplanted into neutral water than did crayfish originating in neutral water, providing some evidence for a cost of acclimation to acidity. 3. Stream surveys showed that fish were less abundant and crayfish more abundant in acidified streams than in circumneutral streams. Crayfish density was sixfold higher in reaches with the lowest pH relative to circumneutral reaches. Large crayfish made up a higher proportion of crayfish populations at sites with high fish biomass, consistent with the hypothesis that fish predation on small individuals may be limiting crayfish population size at these sites. 4. Although individual crayfish suffered lower growth in acidified streams, increased acidity appeared to cause an increase in crayfish population size and shifts in size structure, possibly by relieving predation pressure by fish.     

BIODIVERSITY RESEARCH: Conserving macroinvertebrate diversity in headwater streams: the importance of knowing the relative contributions of α and β diversity

Aim We investigated partitioning of aquatic macroinvertebrate diversity in eight headwater streams to determine the relative contributions of α and β diversity to γ diversity, and the scale dependence of α and β components. Location Great Dividing Range, Victoria, Australia. Methods We used the method of Jost (Ecology, 2007, 88, 2427–2439) to partition γ diversity into its α and β components. We undertook the analyses at both reach and catchment scales to explore whether inferences depended on scale of observation. Results We hypothesized that β diversity would make a large contribution to the γ diversity of macroinvertebrates in our dendritic riverine landscape, particularly at the larger spatial scale (among catchments) because of limited dispersal among sites and especially among catchments. However, reaches each had relatively high taxon richness and high α diversity, while β diversity made only a small contribution to γ diversity at both the reach and catchment scales. Main conclusions Dendritic riverine landscapes have been thought to generate high β diversity as a consequence of limited dispersal and high heterogeneity among individual streams, but this may not hold for all headwater stream systems. Here, α diversity was high and β diversity low, with individual headwater stream reaches each containing a large portion of γ diversity. Thus, each stream could be considered to have low irreplaceability since losing the option to use one of these sites in a representative reserve network does not greatly diminish the options available for completing the reserve network. Where limited information on individual taxonomic distributions is available, or time and money for modelling approaches are limited, diversity partitioning may provide a useful ‘first‐cut’ for obtaining information about the irreplaceability of individual streams or subcatchments when establishing representative freshwater reserves.     

Predicting fine‐scale distributions of peripheral aquatic species in headwater streams

Headwater species and peripheral populations that occupy habitat at the edge of a species range may hold an increased conservation value to managers due to their potential to maximize intraspecies diversity and species' adaptive capabilities in the context of rapid environmental change. The southern Appalachian Mountains are the southern extent of the geographic range of native Salvelinus fontinalis and naturalized Oncorhynchus mykiss and Salmo trutta in eastern North America. We predicted distributions of these peripheral, headwater wild trout populations at a fine scale to serve as a planning and management tool for resource managers to maximize resistance and resilience of these populations in the face of anthropogenic stressors. We developed correlative logistic regression models to predict occurrence of brook trout, rainbow trout, and brown trout for every interconfluence stream reach in the study area. A stream network was generated to capture a more consistent representation of headwater streams. Each of the final models had four significant metrics in common: stream order, fragmentation, precipitation, and land cover. Strahler stream order was found to be the most influential variable in two of the three final models and the second most influential variable in the other model. Greater than 70% presence accuracy was achieved for all three models. The underrepresentation of headwater streams in commonly used hydrography datasets is an important consideration that warrants close examination when forecasting headwater species distributions and range estimates. Additionally, it appears that a relative watershed position metric (e.g., stream order) is an important surrogate variable (even when elevation is included) for biotic interactions across the landscape in areas where headwater species distributions are influenced by topographical gradients.     

Species divergence and the measurement of microbial diversity

Diversity measurement is important for understanding community structure and dynamics, but has been particularly challenging for microorganisms. Microbial community characterization using small subunit rRNA (SSU rRNA) gene sequences has revealed an extensive, previously unsuspected diversity that we are only now beginning to understand, especially now that advanced sequencing technologies are producing datasets containing hundreds of thousands of sequences from hundreds of samples. Efforts to quantify microbial diversity often use taxon-based methods that ignore the fact that not all species are equally related, which can therefore obscure important patterns in the data. For example, alpha-diversity (diversity within communities) is often estimated as the number of species in a community (species richness), and beta-diversity (partitioning of diversity among communities) is often based on the number of shared species. Methods for measuring alpha- and beta-diversity that account for different levels of divergence between individuals have recently been more widely applied. These methods are more powerful than taxon-based methods because microorganisms in a community differ dramatically in sequence similarity, which also often correlates with phenotypic similarity in key features such as metabolic capabilities. Consequently, divergence-based methods are providing new insights into microbial community structure and function.     

Critical landscape attributes that influence fish population dynamics in headwater streams

Three landscape attributes are likely to have strong effects on the rate-dependent processes determining fish population dynamics in headwater streams: (1) functional interactions at terrestrial-aquatic ecotones and their influence on temporal and spatial variation in resource supply and predator-prey interactions, (2) large-scale spatial habitat relationships and their effect on resource use and fish movement, and (3) presence of refugia from harsh environmental conditions and their influence on fish survival and emigration/immigration rates. Elucidating how these factors interact over a range of temporal and spatial scales should be a major goal of lotic fish ecologists.     

Sediment deposition from eroding peatlands alters headwater invertebrate biodiversity

Land use and climate change are driving widespread modifications to the biodiverse and functionally unique headwaters of rivers. In temperate and boreal regions, many headwaters drain peatlands where land management and climate change can cause significant soil erosion and peat deposition in rivers. However, effects of peat deposition in river ecosystems remain poorly understood. We provide two lines of evidence—derived from sediment deposition gradients in experimental mesocosms (0–7.5 g/m²) and headwaters (0.82–9.67 g/m²)—for the adverse impact of peat deposition on invertebrate community biodiversity. We found a consistent negative effect of sediment deposition across both the experiment and survey; at the community level, decreases in density (1956 to 56 individuals per m² in headwaters; mean 823 ± 129 (SE) to 288 ± 115 individuals per m² in mesocosms) and richness (mean 12 ± 1 to 6 ± 2 taxa in mesocosms) were observed. Sedimentation increased beta diversity amongst experimental replicates and headwaters, reflecting increasing stochasticity amongst tolerant groups in sedimented habitats. With increasing sedimentation, the density of the most common species, Leuctra inermis, declined from 290 ± 60 to 70 ± 30 individuals/m² on average in mesocosms and >800 individuals/m² to 0 in the field survey. Traits analysis of mesocosm assemblages suggested biodiversity loss was driven by decreasing abundance of invertebrates with trait combinations sensitive to sedimentation (longer life cycles, active aquatic dispersal of larvae, fixed aquatic eggs, shredding feeding habit). Functional diversity metrics reinforced the idea of more stochastic community assembly under higher sedimentation rates. While mesocosm assemblages showed some compositional differences to surveyed headwaters, ecological responses were consistent across these spatial scales. Our results suggest short‐term, small‐scale stressor experiments can inform understanding of “real‐world” peatland river ecosystems. As climate change and land‐use change are expected to enhance peatland erosion, significant alterations to invertebrate biodiversity can be expected where these eroded soils are deposited in rivers.     

Experimental reductions in stream flow alter litter processing and consumer subsidies in headwater streams

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Do body size and diet breadth affect partitioning of species diversity? A test with forest Lepidoptera

Two of the major themes resulting from recent macroecological research are the central roles that body size and niche breadth may play as determinants of species geographical distribution. Unanswered questions, however, linger regarding how similarities in body size or niche breadth affect the allocation of α‐ and β‐diversity across spatial scales. Using data on moth diversity in the eastern deciduous forest of North America, we tested the predictions that smaller‐bodied and diet‐restricted species would have lower levels of α‐diversity within forest stands and greater β‐diversity at higher sampling scales compared to larger or more generalist species. Moths were sampled using a nested sampling design consisting of three hierarchical levels: 20 forest stands, 5 sites and 3 ecoregions. Body size for 492 species was estimated as mean forewing length, and diet breadth was assessed from the published literature. Moth species were then classified according to body size (small or large) or diet breadth (generalist or restricted), and partitioning was conducted on each group. Diversity partitions for large‐ and small‐bodied species yielded similar patterns. When observed diversity components differed from those derived from our null model, a consistent pattern was observed: α‐diversity was greater than expected, β‐diversity among forest stands was less than expected, and β‐diversity among sites and ecoregions was higher than expected. In contrast, diet‐restricted moths contributed significantly less to stand‐level α‐diversity than generalist feeders. Furthermore, specialists contributed to a greater proportion of β‐diversity across scales compared to generalist moths. Because absolute measures of β‐diversity among stands were greater for generalists than for restricted feeders, we suggest that regional β‐diversity of forest moths may be influenced by several possible factors: intraspecific aggregation of diet‐restricted species, local fluctuations in population size of eruptive generalists and small geographical distributions of generalist moths than predicted by the geographical extent of putative host plants     

Computing β-diversity with Rao's quadratic entropy: a change of perspective

Ecologists have traditionally viewed β-diversity as the ratio between γ-diversity and average α-diversity. More recently, an alternative way of partitioning diversity has been proposed for which β-diversity is obtained as the difference between γ-diversity and average α-diversity. Although this additive model of diversity decomposition is generally considered superior to its multiplicative counterpart, in both models β-diversity is a formally derived quantity without any self-contained ecological meaning; it simply quantifies the diversity excess of γ-diversity with respect to average α-diversity. Taking this excess as an index of β-diversity is a questionable operation. In this paper, we show that a particular family of α-diversity measures, the most celebrated of which is Rao's quadratic entropy, can be adequately used for summarizing β-diversity. Our proposal naturally leads to a new additive model of diversity for which, given two or more sets of plots, overall plot-to-plot species variability can be additively partitioned into two non-negative components: average variability in species composition within each set of plots and the species variability between the set of plots. For conservation purposes, the suggested change of perspective in the summarization of β-diversity allows for a flexible analysis of spatial heterogeneity in ecological diversity so that different hierarchical levels of biotic relevance (i.e. from the genetic to the landscape level) can be expressed in a significant and consistent way.     

Species diversity in neutral metacommunities: a network approach

Biologists seek an understanding of the processes underlying spatial biodiversity patterns. Neutral theory links those patterns to dispersal, speciation and community drift. Here, we advance the spatially explicit neutral model by representing the metacommunity as a network of smaller communities. Analytic theory is presented for a set of equilibrium diversity patterns in networks of communities, facilitating the exploration of parameter space not accessible by simulation. We use this theory to evaluate how the basic properties of a metacommunity – connectivity, size, and speciation rate – determine overall metacommunity γ -diversity, and how that is partitioned into α - and β -components. We find spatial structure can increase γ -diversity relative to a well-mixed model, even when θ is held constant. The magnitude of deviations from the well-mixed model and the partitioning into α - and β -diversity is related to the ratio of migration and speciation rates. γ -diversity scales linearly with metacommunity size even as α - and β -diversity scale nonlinearly with size.     

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