Hierarchical spatial structure of stream fish colonization and extinction

Spatial variation in extinction and colonization is expected to influence community composition over time. In stream fish communities, local species richness (alpha diversity) and species turnover (beta diversity) are thought to be regulated by high extinction rates in headwater streams and high colonization rates in downstream areas. We evaluated the spatiotemporal structure of fish communities in streams originally surveyed by Burton and Odum 1945 (Ecology 26: 182–194) in Virginia, USA and explored the effects of species traits on extinction and colonization dynamics. We documented dramatic changes in fish community structure at both the site and stream scales. Of the 34 fish species observed, 20 (59%) were present in both time periods, but 11 (32%) colonized the study area and three (9%) were extirpated over time. Within streams, alpha diversity increased in two of three streams but beta diversity decreased dramatically in all streams due to fish community homogenization caused by colonization of common species and extirpation of rare species. Among streams, however, fish communities differentiated over time. Regression trees indicated that reproductive life‐history traits such as spawning mound construction, associations with mound‐building species, and high fecundity were important predictors of species persistence or colonization. Conversely, native fishes not associated with mound‐building exhibited the highest rates of extirpation from streams. Our results demonstrate that stream fish colonization and extinction dynamics exhibit hierarchical spatial structure and suggest that mound‐building fishes serve as keystone species for colonization of headwater streams.     

The influence of channelisation on riparian plant assemblages

1. A comparative study of species diversity and assemblage patterns of herbaceous ground‐flora communities in riparian areas was performed along natural mid‐sized lowland streams and their channelised counterparts. The areas had open vegetation and were positioned along 18 similar‐sized third to fourth order stream reaches. 2. Alpha diversity was significantly higher along natural streams both at the sample plot and reach scale. Sample plot diversity peaked at intermediate distance from the natural stream reaches, whereas it increased with increasing distance from the channelised reaches. Both gamma diversity, measured and estimated from species‐area curves, and beta diversity, which is a measure of the change in diversity between areas, was similar along the two types of streams. 3. Alpha diversity correlated with several of the measured and calibrated environmental variables. The positive correlation between bank slope and alpha diversity indicates that flooding plays a key role in maintaining high levels of diversity along natural streams. 4. Species composition varied significantly between the two stream types. A cluster analysis identified four clusters of which two clusters (one and three) primarily included species associated with sample plots in areas along natural streams. Most cluster one and three species were also identified as indicator species for this stream type. 5. Canonical correspondence analysis revealed that cluster one species were less productive species associated with high total soil carbon and nitrogen contents, whereas cluster three species were highly productive species associated with high soil moisture levels, probably partly resulting from flooding. Our results suggest that distance from the stream channel imposes a probabilistic gradient that sustains co‐occurrence of these two communities in riparian areas along natural mid‐sized lowland streams.     

Diversification rates of the “Old Endemic” murine rodents of Luzon Island,Philippines are inconsistent with incumbency effects and ecological opportunity

Diversity‐dependent cladogenesis occurs when a colonizing lineage exhibits increasing interspecific competition as it ecologically diversifies. Repeated colonization of a region by closely related taxa may cause similar effects as species within each lineage compete with one another. This may be particularly relevant for secondary colonists, which could experience limited diversification due to competition with earlier, incumbent colonists over evolutionary time. We tested the hypothesis that an incumbent lineage may diminish the diversification of secondary colonists in two speciose clades of Philippine “Old Endemic” murine rodents—Phloeomyini and Chrotomyini—on the relatively old oceanic island of Luzon. Although phylogenetic analyses confirm the independent, noncontemporaneous colonization of Luzon by the ancestors of these two clades, we found no support for arrested diversification in either. Rather, it appears that diversification of both clades resulted from constant‐rate processes that were either uniform or favored the secondary colonists (Chrotomyini), depending on the method used. Our results suggest that ecological incumbency has not played an important role in determining lineage diversification among Luzon murines, despite sympatric occurrence by constituent species within each lineage, and a substantial head start for the primary colonists.     

Assessing community functional attributes during substrate colonization: a field experiment using stream insects

Initial colonists of empty habitats tend to differ from those arriving later in terms of species identity and traits. We evaluated the dynamics of the functional attributes in aquatic insect communities during a colonization experiment under natural conditions. We tested whether the late stages of colonization show higher functional richness, diversity and specialization than early successional stages. We used 60 artificial slate samplers that were removed after 1, 3, 5, 10, 15 and 30 days of colonization. We considered five traits (with a total of 17 trait categories): feeding habits, dispersal medium, body size, body shape and locomotion. With these traits, we computed a global specialization index at the community level. Large shredders with a cylindrical body shape and fly dispersal while flying were prominent in late colonization. In contrast, early colonists tended to have flattened body and to disperse through water. Functional diversity and functional richness significantly increased in late colonization, resulting in a more specialized community. Our results suggest that any interference during the processes involved in stream insect colonization can be reflected in the community through the decrease or even lack of functional attributes.

     

Community diversity outweighs effect of warming on plant colonization

Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m²). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.     

Evaluating effects of weed cutting on water level and ecological status in Danish lowland streams

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Community monopolization: local adaptation enhances priority effects in an evolving metacommunity

The diversity and composition of biological communities might often depend on colonization history because early colonists can exclude future colonists through a priority effect. These priority effects, which have been observed across a wide variety of ecosystems, often arise because early colonists have sufficient time to use available resources efficiently and subsequently withhold them from invaders. Here, we explore the extent to which rapid local adaptive evolution contributes to the pervasiveness of these priority effects. Using an individual-based simulation, we show that early colonization allows the descendants of colonists to adapt to novel conditions and reduce the establishment success of an initially ecologically equivalent competing species. Our model predicts that slight differences in colonization timing and adaptive capacity between species can substantially alter the dynamics and diversity of communities. We also show that priority effects and gene flow can generate a novel mechanism for the expansion and retraction of species distributions in a metacommunity. Our results suggest that local adaptation combined with stochastic colonization events can obscure direct relationships between species distributions and environmental gradients. Given the increasing recognition of rapid, microgeographic evolution in natural populations, we expect that evolutionary priority effects could affect the structure and dynamics of many natural metacommunities.     

Leaf breakdown in streams differing in catchment land use

1. The impact of changes in land use on stream ecosystem function is poorly understood. We studied leaf breakdown, a fundamental process of stream ecosystems, in streams that represent a range of catchment land use in the Piedmont physiographic province of the south‐eastern United States. 2. We placed bags of chalk maple (Acer barbatum) leaves in similar‐sized streams in 12 catchments of differing dominant land use: four forested, three agricultural, two suburban and three urban catchments. We measured leaf mass, invertebrate abundance and fungal biomass in leaf bags over time. 3. Leaves decayed significantly faster in agricultural (0.0465 day^?1) and urban (0.0474 day^?1) streams than in suburban (0.0173 day^?1) and forested (0.0100 day^?1) streams. Additionally, breakdown rates in the agricultural and urban streams were among the fastest reported for deciduous leaves in any stream. Nutrient concentrations in agricultural streams were significantly higher than in any other land‐use type. Fungal biomass associated with leaves was significantly lower in urban streams; while shredder abundance in leaf bags was significantly higher in forested and agricultural streams than in suburban and urban streams. Storm runoff was significantly higher in urban and suburban catchments that had higher impervious surface cover than forested or agricultural catchments. 4. We propose that processes accelerating leaf breakdown in agricultural and urban streams were not the same: faster breakdown in agricultural streams was due to increased biological activity as a result of nutrient enrichment, whereas faster breakdown in urban streams was a result of physical fragmentation resulting from higher storm runoff.     

Detrital processing in streams exposed to acidic precipitation in the Central Appalachian Mountains

Continuing high rates of acidic deposition in the eastern United States may lead to long-term effects on stream communities, because sensitive catchments are continuing to lose anions and cations. We conducted a two-year study of the effects of pH and associated water chemistry variables on detrital processing in three streams with different bedrock geology in the Monongahela National Forest, West Virginia. We compared leaf pack processing rates and macroinvertebrate colonization and microbial biomass (ATP concentration) on the packs in the three streams. Breakdown rates of red maple and white oak leaf packs were significantly lower in the most acidic stream. The acidic stream also had significantly lower microbial and shredder biomass than two more circumneutral streams. Shredder composition differed among streams; large-particle detritivores dominated the shredder assemblages of the two circumneutral streams, and smaller shredders dominated in the acidic stream. Within streams, processing rates for three leaf species were not significantly different between the two years of the study even though invertebrate and microbial communities were different in the two years. Thus, macroinvertebrate and microbial communities differed both among streams that differed in their capacity to buffer the effects of acidic precipitation and among years in the same stream; these differences in biotic communities were not large enough to affect rates of leaf processing between the two years of the study, but they did significantly affect processing rates between acidic and circumneutral streams.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.     

The influence of initial colonization by hydropsychid caddisfly larvae on the development of stream invertebrate assemblages

Many aspects of the colonization history of a disturbed site can influence the development of a biological community. Initial colonization is known to play a significant role in community development because of the facilitative or inhibitory effects that `pioneer' species can have on subsequently arriving taxa. We performed an experiment to assess how initial colonization by two species of benthic invertebrates (Trichoptera: Hydropsychidae) might influence the development of stream faunal assemblages. Substrate baskets initially colonized by either Hydropsyche depravata or Ceratopsyche bronta were placed alongside control baskets in a recently flooded stream. After baskets had colonized for 30 days, we found that species composition in treatment baskets was identical to that in control baskets, indicating that the caddisfly taxa had no selective effects on colonization of other macroinvertebrate species. We did, however, find that C. bronta facilitated the recruitment of all species in the colonist pool leading to greater overall abundance and biomass of macroinvertebrates in the final assemblages. In contrast, H. depravata had no effect on the abundance or biomass of colonizing invertebrates. The differential effects of these two taxa on abundance and biomass may have been related to differences in microhabitat complexity created by the construction of their retreats and catchnets. The results of this study support the growing recognition that colonization history does influence the structure of lotic communities, but they also suggest that even closely related taxa can play different roles as initial colonists in community development.     

Dispersal of plant fragments in small streams

1. Streams are subject to frequent natural and anthropogenic disturbances that cause sediment erosion and loss of submerged vegetation. This loss makes downstream transport and retention of vegetative propagules on the streambed very important for re‐establishing vegetation cover. We measured dispersal and retention of macrophyte stem fragments (15–20 cm long) along 300 m long reaches of four small to medium sized Danish lowland streams. 2. The number of drifting stem fragments declined exponentially with distance below the point of release. This finding makes the retention coefficient (k, m⁻¹) in the exponential equation a suitable measure for comparisons among different macrophyte species, and between stream reaches of different hydrology and vegetation cover. 3. Buoyancy of macrophyte tissue influenced retention. Elodea canadensis stems drifted below the water surface, and were more inclined to be retained in deeper water associated with submerged plants and obstacles in the streambed. Ranunculus peltatus stems were more buoyant, drifted at the water surface, and were more inclined to be trapped in shallow water and in riparian vegetation. 4. The retention coefficient of drifting stems increased with the relative contact between the flowing water and streambed, bank and vegetation. Thus, the retention coefficients were highest (0.02–0.12 m⁻¹) in shallow reaches with a narrow, vegetation‐free flow channel. Here there were no significant differences between E. canadensis and R. peltatus. Retention coefficients were lowest (0.0005–0.0135 m⁻¹) in deeper reaches with wider vegetation‐free flow channels. Retention of E. canadensis was up to 16 times more likely than retention of R. peltatus. 5. Overall, the longitudinal position in the stream system of source populations of species capable of producing numerous stems, the species‐specific retention coefficients of stems, and the retention capacity of stream reaches should be important for species distribution in perturbed stream systems. Retention of stems is probably constrained in headwaters by the small downstream flux of stem fragments because of the restricted source area, and constrained in downstream reaches by small retention coefficients. Macrophyte retention may, consequently, peak in medium‐sized streams.     

Distinct colonization waves underlie the diversification of the freshwater sculpin (Cottus gobio) in the Central European Alpine region

Ecological speciation and adaptive radiation are key processes shaping northern temperate freshwater fish diversity. Both often involve parapatric differentiation between stream and lake populations and less often, sympatric intralacustrine diversification into habitat‐ and resource‐associated ecotypes. However, few taxa have been studied, calling for studies of others to investigate the generality of these processes. Here, we test for diversification within catchments in freshwater sculpins in a network of peri‐Alpine lakes and streams. Using 8047 and 13 182 restriction site‐associated (RADseq) SNPs, respectively, we identify three deeply divergent phylogeographic lineages associated with different major European drainages. Within the Aare catchment, we observe populations from geographically distant lakes to be genetically more similar to each other than to populations from nearby streams. This pattern is consistent with two distinct colonization waves, rather than by parapatric ecological speciation after a single colonization wave. We further find two distinct depth distribution modes in three lakes of the Aare catchment, one in very shallow and one in very deep water, and significant genomewide differentiation between these in one lake. Sculpins in the Aare catchment appear to represent an early‐stage adaptive radiation involving the evolution of a lacustrine lineage distinct from parapatric stream sculpins and the repeated onset of depth‐related intralacustrine differentiation.     

Meta‐community theory and stream restoration: evidence that spatial position constrains stream invertebrate communities in a mine impacted landscape

Mining activities, particularly acid mine drainage, often result in adverse effects on stream diversity and ecosystem functioning, and increased concern about these effects has generated a focus on restoration of mine‐impacted waterways. However, many stream restoration projects have not led to increased stream diversity and ecological recovery. One reason for this failure may be that restoration practitioners focus on local environmental conditions and fail to consider the importance of dispersal as a driver of stream invertebrate composition. To test this hypothesis, we used a meta‐community analysis to compare the influence of the local stream conditions with the regional supply of colonists. Invertebrate communities and physico‐chemical conditions were sampled in 37 streams across a mine‐impact gradient on the Stockton Plateau, West Coast of New Zealand's South Island. We found that pH, temperature, dissolved metals, and sediment significantly influenced invertebrate community composition. Furthermore, the spatial location of streams was a good predictor of stream diversity and invertebrate communities, independent of local environmental conditions. This result indicates an important role for regional dispersal barriers in determining stream invertebrate communities. Consequently, consideration of both the locations and strategic preservation of future colonist source streams and potential dispersal barriers during mine planning would enhance post‐mining restoration.     

Species richness and assemblage structure of Trichoptera in Danish streams

• 1 Species richness and assemblages of Trichoptera from 157 ‘pristine’ Danish lowland stream sites were analyzed, for dependence on geographical position of the sites and simple physical variables, using two way indicator species analysis and detrended correspondence analysis.
• 2 A total of 106 species were recorded, representing ≈ 90% of the species pool known from Danish streams. Only seven species occurred at more than half the sites, whereas an additional 11 species were found at more than a quarter of the sites.
• 3 Although sites showed significant regional differences in environmental variables (stream order, width, slope and presence/absence of riparian forest), species richness and assemblages were primarily correlated with stream order, width and slope. Maximum richness was found at the largest (5th order) stream sites.
• 4 Regional differences in species assemblages were found, with several species being absent from the islands Funen and Bornholm. Species assemblages also differed between forested and non‐forested stream sites.
• 5 We found evidence that stream temperature may be of only minor importance in determining Trichoptera species richness and assemblage composition in Danish streams compared to other size‐related physical factors.

     

PHYLOGEOGRAPHY OF BATRACHOSPERMUM MACROSPORUM (BATRACHOSPERMALES,RHODOPHYTA) FROM NORTH AND SOUTH AMERICA1

Phylogeographic trends in Batrachospermum macrosporum Mont. were investigated using the mitochondrial intergenic spacer between the cytochrome oxidase subunit 2 and 3 genes (cox2‐3). A total of 11 stream segments were sampled with seven in the coastal plain of North America and four in tropical areas of South America. Fifteen thalli were sampled from seven streams, 14 thalli from two streams, and eight thalli from two streams. There were 16 haplotypes detected using 149 individuals. Of the eight haplotypes from locations in North America, all were 334 base pairs (bp) in length, and of those from South America, five were 344 bp, and three were 348 bp. Two individual networks were produced: one for the haplotypes from North America and another for those from South America, and these could not be joined due to the large number of base pair differences. This split between haplotypes from North and South America was confirmed with sequence data of the rbcL gene. There was very little genetic variation among the haplotypes from the North American locations, leading us to hypothesize that these are fairly recent colonization events along the coastal plain. In contrast, there was high variation among haplotypes from South America, and it would appear that the Amazon serves as a center of diversity. We detected considerable variation in haplotypes among streams, but frequently, a single haplotype in an individual stream segment, which is consistent with data from previous studies of other batrachospermalean taxa, may suggest a single colonization event per stream.     

Land use change affects macroinvertebrate community size spectrum in streams: the case of Pinus radiata plantations

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Genetic assessment of the effects of streamscape succession on coho salmon Oncorhynchus kisutch colonization in recently deglaciated streams

Measures of genetic diversity within and among populations and historical geomorphological data on stream landscapes were used in model simulations based on approximate Bayesian computation (ABC) to examine hypotheses of the relative importance of stream features (geomorphology and age) associated with colonization events and gene flow for coho salmon Oncorhynchus kisutch breeding in recently deglaciated streams (50–240 years b.p .) in Glacier Bay National Park (GBNP), Alaska. Population estimates of genetic diversity including heterozygosity and allelic richness declined significantly and monotonically from the oldest and largest to youngest and smallest GBNP streams. Interpopulation variance in allele frequency increased with increasing distance between streams (r = 0·435, P < 0·01) and was inversely related to stream age (r = –0·281, P < 0·01). The most supported model of colonization involved ongoing or recent (<10 generations before sampling) colonization originating from large populations outside Glacier Bay proper into all other GBNP streams sampled. Results here show that sustained gene flow from large source populations is important to recently established O. kisutch metapopulations. Studies that document how genetic and demographic characteristics of newly founded populations vary associated with successional changes in stream habitat are of particular importance to and have significant implications for, restoration of declining or repatriation of extirpated populations in other regions of the species' native range.     

The effect of light and seed mass on seed germination of common herbaceous species from the eastern Qinghai‐Tibet Plateau

Smaller seeds might encounter more severe selective pressure than larger ones because they have fewer food reserves and are more easily buried; thus, seed mass can be considered to be directly related to the effect of light on germination. To investigate the effect of light on seed germination and associated seed mass variation within a whole plant community, we compiled germination data for common herbaceous species from an alpine meadow on the eastern Qinghai‐Tibet plateau. The results showed the following. (i) Light had a general positive effect on seed germination. Under light, the proportion of species with lower germinability was decreased, mean germination percentage was increased by 20% and the speed of germination was doubled. (ii) Irrespective of light environment, species with medium‐sized seeds (seed mass ranging from 0.11 to 0.5 mg) had higher germination percentage and speed when compared with species within the largest seed mass group. (iii) The germination of smaller‐seeded species was more dependent on light stimulation than larger‐seeded ones. In darkness, the species within the smallest seed mass group had the lowest percentage and speed of germination; however, under light, the species within the largest seed mass group had the lowest percentage and speed of germination. Our results suggested that the germination characteristics and especially seeds’ response to light among species in the alpine meadow might be an adaptation to natural selective pressure.     

Physical and biological changes to a lengthening stream gradient following a decade of rapid glacial recession

In mountains, environmental gradients are steep in both terrestrial and aquatic systems, and climate change is causing upward shifts of physical and biological features of these gradients. Glacial streams are an interesting system to evaluate such shifts both because streams have a linear nature (for simplicity of analysis), and because the stream habitat will at least temporarily lengthen as it follows receding glaciers upward. The Tschierva Glacier, Swiss Alps, receded 482 m upstream from 1997 to 2008. We tested the null hypothesis that the physical and biological stream gradient below this glacier maintained the same structure between these time periods, but simply shifted upward following the glacial source. We compared longitudinal patterns of water temperature and zoobenthic community structure in 1997 and 2007–2008 during three seasons (spring, summer, fall) along the uppermost ca. 5 stream km. Upward shifts were evident, including colonization of the newly exposed stream reaches by cold‐adapted taxa, and the appearance in 2007/2008 of four lower‐altitude species that were previously absent. Overall, however, results rejected the null hypothesis, instead revealing significant changes in gradient structures. These included a more steeply increasing temperature profile downstream of the glacier and increased amplitude of seasonal community turnover in 2007/2008 vs. 1997. Long‐term (1955–2007) flow records revealed increasing short‐term and seasonal hydrologic variability, which might have influenced the increased intra‐annual community variability. The steepening of the temperature gradient was likely caused by a warming lake‐outlet tributary upon which glacial influence was diminished between 1997 and 2007/2008. These results suggest that upward‐shifting gradients in glacial streams can involve complex interactions with other landscape elements and that local‐scale climate response can progress even more rapidly than the rate of glacial recession.     

Convergence of fish communities from the littoral zone of reservoirs

1. Understanding factors that regulate the assembly of communities is a main focus of ecology. Human‐engineered habitats, such as reservoirs, may provide insight into these assembly processes because they represent novel habitats that are subjected to colonization by fishes from the surrounding river basin or transported by humans. By contrasting community similarity within and among reservoirs from different drainage basins to nearby stream communities, we can test the relative constraints of reservoir habitats and regional species pools in determining species composition of reservoirs. 2. We used a large spatial database that included intensive collections from 143 stream and 28 reservoir sites within three major river basins in the Great Plains, U.S.A., to compare patterns of species diversity and community structure between streams and reservoirs and to characterize variation in fish community structure within and among major drainage basins. We expected reservoir fish faunas to reflect the regional species pool, but would be more homogeneous that stream communities because similar species are stocked and thrive in reservoirs (e.g. planktivores and piscivores), and they lack obligate stream organisms that are not shared among regional species pools. 3. We found that fish communities from reservoirs were a subset of fishes collected from streams and dominant taxa had ecological traits that would be favoured in lentic environments. Although there were regional differences in reservoir fish communities, species richness, patterns of rank abundance and community structure in reservoir communities were more homogonous across three major drainage basins than for stream communities. 4. The general pattern of convergence of reservoir fish community structure suggests their assembly is constrained by local factors such as habitat and biotic interactions, and facilitated by the introduction of species among basins. Because there is a reciprocal transfer of biota between reservoirs and streams, understanding factors structuring both habitats is necessary to evaluate the long‐term dynamics of impounded river networks.