Potential effects on zooplankton from species shifts in planktivorous mussels: a field experiment in the St Lawrence River

1. Suspension feeding by bivalves exceeds that by other planktivores in many North American rivers, and food webs may be altered substantially by differences in feeding patterns between native unionid mussels and invading dreissenid mussels. 2. We conducted an experiment comparing zooplanktivory by one unionid and two dreissenid species that addressed several primary questions. Is benthic planktivory important in this river? Has this linkage been altered substantially by dreissenids? Do the two dreissenid species differ in planktivory, and is this ecologically important if quagga mussels extend their geographical range? 3. Our 12‐day experiment consisted of controls (no mussels) and treatments with unionid (Elliptio complanata), quagga (Dreissena bugensis) or zebra (D. polymorpha) mussels in 3500‐L, 80‐μm mesh enclosures placed in a slackwater area of the St Lawrence River. 4. The density of the most abundant calanoid copepod Eurytemora affinis increased in the presence of dreissenids, probably as an indirect food web response. By day 12, a cumulative effect was shown by the most overwhelmingly abundant rotifer, Polyarthra, whose density declined dramatically in dreissenid enclosures compared with control and unionid enclosures. Rotifer densities in unionid enclosures were not different from controls, nor were dreissenid treatments different from each other. The effects on rotifers were probably from predation, as Chl‐a did not vary among treatments. 5. We conclude that benthic‐pelagic coupling via planktivory is important in slackwater areas. Dreissenids have strengthened this linkage, but range extension of quaggas should not appreciably alter effects produced by a similar biomass of zebra mussels.     

Invasive ecosystem engineers on soft sediment change the habitat preferences of native mayflies and their availability to predators

1. Dreissenid mussels (quagga mussels, Dreissena bugensis, and zebra mussels, D. polymorpha) are invasive species that function as ecosystem engineers in the Laurentian Great Lakes. Dreissena are increasingly abundant on silt, sand and other soft substrates; by altering benthic habitat, these mussels can alter benthic community structure. 2. We used laboratory mesocosm experiments to examine the effects of soft‐sediment Dreissena clusters on the habitat preference of Hexagenia, a native burrowing mayfly that is an important food source to fish. We conducted three experiments to test whether Hexagenia: (1) select for bare sediment, soft sediment covered with live Dreissena (added structure and food resources) or soft sediment with clusters made of empty Dreissena shells (added structure only), (2) prefer a specific density of live Dreissena on soft sediment and (3) select for or avoid sediment with an accumulation of empty Dreissena shells. 3. Contrary to initial expectations, we found that Hexagenia selected for sediment covered with live Dreissena clusters, followed by empty Dreissena shells clusters, and lastly what was previously thought to be the preferred habitat, bare sediment. Not only did Hexagenia prefer Dreissena‐covered sediment, but they also preferred high densities of Dreissena. 4. We also experimentally tested the effects of Dreissena‐covered soft sediment on the availability of Hexagenia to fish. We had three treatment levels representing three distinct habitat types: (1) bare sediment (no Dreissena) treatment in which water was turbid because of mayfly activity, (2) Dreissena‐covered sediment treatment in which water was clear because of Dreissena filtration and (3) Dreissena‐covered sediment with added turbidity. We found that in low light conditions, similar to many locations where both organisms are found to co‐occur, both yellow perch and round goby consumption of Hexagenia significantly decreased when Dreissena covered the bottom sediment. 5. These results suggest that by choosing Dreissena‐covered habitat, Hexagenia receive protection from fish predation in turbid/low light systems. However, protection from predation cannot be the only reason Hexagenia select Dreissena‐covered sediments, as Hexagenia selected for live clusters more often than empty clusters and may be a result of additional food resources.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

相似文献   

Lake morphometry determines Dreissena invasion dynamics

Predicting the ecosystem effects of invasive species and the best control strategies requires understanding population dynamics and population regulation. Invasive bivalves zebra and quagga mussels (Dreissena spp.) are considered the most aggressive invaders in freshwaters and have become major drivers of ecosystem processes in the Laurentian Great Lakes. Combining all lake-wide studies of Dreissena spp. conducted in the Great Lakes, we found that invasion dynamics are largely governed by lake morphometry. Where both species are present, quagga mussels generally become dominant in 8–13 years. Thereafter, zebra mussels remain common in shallow lakes and embayments and lake-wide Dreissena density may remain similar, while in deep lakes quagga led to a near-complete displacement of zebra mussels and an ensuing dramatic increase in overall dreissenid density. In deep lakes, overall Dreissena biomass peaked later and achieved?~?threefold higher levels than in shallow lakes. Comparison with 21 waterbodies in North America and Europe colonized by both dreissenids confirmed that patterns of invasion dynamics found in the Great Lakes are very consistent with other waterbodies, and thus can be generalized to other lakes. Our biophysical model predicted that the long-term reduction in primary producers by mussel grazing may be fourfold less in deep compared to shallow lakes due to thermal stratification and a smaller proportion of the epilimnion in contact with the bottom. While this impact remains greatest in shallow areas, we show that when lakes are vertically well-mixed, dreissenid grazing impact may be greatest offshore, revealing a potentially strong offshore carbon and phosphorus sink.

     

AN ECOLOGICAL REVIEW OF CLADOPHORA GLOMERATA (CHLOROPHYTA) IN THE LAURENTIAN GREAT LAKES1

Cladophora glomerata (L.) Kütz. is, potentially, the most widely distributed macroalga throughout the world’s freshwater ecosystems. C. glomerata has been described throughout North America, Europe, the Atlantic Islands, the Caribbean Islands, Asia, Africa, Australia and New Zealand, and the Pacific Islands. Cladophora blooms were a common feature of the lower North American Great Lakes (Erie, Michigan, Ontario) from the 1950s through the early 1980s and were largely eradicated through the implementation of a multibillion‐dollar phosphorus (P) abatement program. The return of widespread blooms in these lakes since the mid‐1990s, however, was not associated with increases in P loading. Instead, current evidence indicates that the resurgence in blooms was directly related to ecosystem level changes in substratum availability, water clarity, and P recycling associated with the establishment of dense colonies of invasive dreissenid mussels. These results support the hypothesis that dreissenid mussel invasions may induce dramatic shifts in energy and nutrient flow from pelagic zones to the benthic zone.     

Interim Responses of Benthic and Snag‐Dwelling Macroinvertebrates to Reestablished Flow and Habitat Structure in the Kissimmee River,Florida, U.S.A.

A critical component in the effort to restore the Kissimmee River ecosystem is the reestablishment of an aquatic invertebrate community typical of free‐flowing rivers of the southeastern United States. This article evaluates early responses of benthic and snag‐dwelling macroinvertebrates to restoration of flow and habitat structure following Phase I construction (interim period) of the Kissimmee River Restoration Project. Replicate benthic and snag samples were collected from remnant river channels in Pool A (Control site), and Pool C, the site of the first phase of restoration (Impact site). Samples were collected quarterly for 2 years prior to construction (baseline) and monthly or quarterly for 3 years following Phase I construction and restoration of flow. Baseline benthic data indicate a community dominated by taxa tolerant of organic pollution and low levels of dissolved oxygen, including the dipterans Chaoborus americanus (Chaoboridae) and the Chironomus/Goeldichironomus group (Chironomidae). Baseline snag data indicate a community dominated by gathering‐collectors, shredders, and scrapers. Passive filtering‐collector invertebrates were rare. Following restoration of flow, benthic invertebrate communities are numerically dominated by lotic taxa, including bivalves and sand‐dwelling chironomids (e.g. Polypedilum spp., Cryptochironomus spp., and Tanytarsini). Snags within the Phase I area support an invertebrate community dominated by passive filtering‐collectors including Rheotanytarsus spp. (Chironomidae) and Cheumatopsyche spp. (Hydropsychidae). Results indicate that restoration of flow has resulted in ecologically significant changes to the river habitat template not observed in Pool A. Observed shifts in benthic and snag macroinvertebrate community structure support previously developed hypotheses for macroinvertebrate responses to hydrologic restoration.     

Population genetic history of the dreissenid mussel invasions: expansion patterns across North America

This study tests population genetic patterns across the Eurasian dreissenid mussel invasions of North America—encompassing the zebra mussel Dreissena polymorpha (1986 detection) and the quagga mussel D. rostriformis bugensis (detected in 1990, which now has largely displaced the former in the Great Lakes). We evaluate their source-spread relationships and invasion genetics using 9–11 nuclear microsatellite loci for 583 zebra mussels (21 sites) and 269 quagga mussels (12 sites) from Eurasian and North American range locations, with the latter including the Great Lakes, Mississippi River basin, Atlantic coastal waterways, Colorado River system, and California reservoirs. Additionally, mtDNA cytochrome b gene sequences are used to verify species identity. Our results indicate that North American zebra mussels originate from multiple non-native northern European populations, whereas North American quagga mussels trace to native estuaries in the Southern Bug and Dnieper Rivers. Invasive populations of both species show considerable genetic diversity and structure (zebra F _ST = 0.006–0.263, quagga F _ST = 0.008–0.267), without founder effects. Most newer zebra mussel populations have appreciable genetic diversity, whereas quagga mussel populations from the Colorado River and California show some founder effects. The population genetic composition of both species changed over time at given sites; with some adding alleles from adjacent populations, some losing them, and all retaining closest similarity to their original composition. Zebra mussels from Kansas and California appear genetically similar and assign to a possible origin from the St. Lawrence River, whereas quagga mussels from Nevada and California assign to a possible origin from Lake Ontario. These assignments suggest that overland colonization pathways via recreational boats do not necessarily reflect the most proximate connections. In conclusion, our microsatellite results comprise a valuable baseline for resolving present and future dreissenid mussel invasion pathways.     

Multi‐scale habitat modification by coexisting ecosystem engineers drives spatial separation of macrobenthic functional groups

By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large‐scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co‐occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co‐occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. This understanding can profitably be used to improve ecosystem‐based management and conservation actions.     

Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river

1. Algal growth in lotic systems is controlled either from the bottom‐up (e.g. nutrients and light, which determine growth rates) or from the top‐down (e.g. grazing pressure, which reduces accumulated biomass). Nutrient‐enriched streams that support large and diverse grazing macroinvertebrate populations and those with shaded riparian corridors rarely suffer from excessive algal growth. 2. In this study, the density of benthic algivorous macroinvertebrates was experimentally manipulated in shaded and open nutrient‐enriched stream habitats of the Owennagearagh River, south‐west Ireland. The ability of macroinvertebrate grazers and riparian shade to control benthic algal growth [particularly the nuisance alga, Cladophora glomerata (L. Kütz)] was investigated. Three sites with markedly different concentrations of plant nutrients (one site upstream and two sites downstream of the sewage outfall) were selected. The density of grazing invertebrates colonising ceramic tiles was reduced using high‐voltage localised electric pulses. Replicates of treatment (grazer‐excluded) and control (grazed) tiles were deployed in open and shaded (<25 and >80% canopy cover, respectively) patches of stream bed, in each site. 3. After 2‐week Cladophora cover, periphytic chlorophyll a and biofilm ash‐free dry mass (AFDM) were quantified for all experimental tiles. Values for all three parameters were highest on grazer‐excluded tiles from open patches. Grazed tiles from open patches accrued little Cladophora and had significantly lower levels of chlorophyll a and AFDM. Nutrient inputs were found to have an impact on the density of grazing invertebrates, with higher densities of Baetis nymphs at the most nutrient‐enriched site. 4. Our results demonstrate that in eutrophic, high‐light streams, filamentous algae can quickly accumulate to nuisance levels in the absence of invertebrate grazers. In future, greater attention should be paid to the role of grazing invertebrates in controlling nuisance algae in streams, in addition to algal–nutrient relationships.     

Effects of structural habitat on drift distance and benthic settlement of the caddisfly,Ceratopsyche sparna

We conducted two experiments in flow-through, artificial streams to examine how habitat structure affected drift and benthic resettlement of larval hydropsychid caddisflies (Ceratopsyche sparna). In the first experiment, we quantified drift distance and the number of times larvae re-entered the drift in 9.0 × 0.51-m channels with contiguous patches (ea. 2.5-m long) of biofilm-covered gravel, biofilm-covered cobbles (– Cladophora), and Cladophora-covered cobbles (+ Cladophora). In the second experiment, we tracked nocturnal movements of larvae after benthic settlement in 2.8 × 0.1-m channels, each containing one of the three habitat types. In experiment 1, drift distance was (1) greatest in gravel and lowest in cobbles + Cladophora, (2) inversely related to hydraulic roughness of habitats, (3) independent of body size, and (4) similar for live and dead larvae. Average drift distance was relatively short (<2.5 m), regardless of habitat type. Number of drift re-entries also varied among habitats, being greatest in gravel and lowest in cobbles + Cladophora. No larvae re-entered the drift after settling in Cladophora patches. Results from experiment 2 revealed that drift propensities were higher for larvae in biofilm-covered gravel and cobbles than in cobbles + Cladophora. Larvae remaining in substrate patches (i.e. not drifting) laid fewer draglines in biofilm-covered stones than in Cladophora patches. Extent of benthic movement (i.e., crawling) by non-drifting larvae did not differ significantly among habitats. However, distance moved did differ with flow direction, being 4× greater downstream than upstream. These results highlight how local substrate and hydraulic conditions interact to affect small-scale movements of caddisfly larvae.     

Ecosystem engineers maintain a rare species of butterfly and increase plant diversity

We evaluated whether ecosystem engineers can accomplish two conservation goals simultaneously: (1) indirectly maintain populations of an endangered animal through habitat modification and (2) increase riparian plant diversity. We tested for effects of a prominent ecosystem engineer, the beaver Castor canadensis, on populations of St. Francis' satyr butterfly Neonympha mitchellii francisci and plant species richness and composition. We performed our test by surveying riparian vegetation communities in all stages of beaver‐influenced wetland succession. We found that beavers created wetland habitats that supported plant species not found elsewhere in riparian zones and increased plant species diversity across the landscape by creating a novel combination of patch types. Our results confirmed what others have found about engineering effects on plant diversity, but these results further demonstrated a case where ecosystem engineers indirectly maintain populations of rare animals by modifying the composition and diversity of plant communities within wetlands. Our research demonstrates how an ecosystem engineer can influence habitat availability and composition of plant communities important for an endangered insect, and maintain overall plant species diversity by increasing habitat heterogeneity.     

Microhabitat selection by the invasive amphipod Echinogammarus ischnus and native Gammarus fasciatus in laboratory experiments and in Lake Erie

1. The amphipod Echinogammarus ischnus was first reported in the Laurentian Great Lakes during 1995. However, analysis of archived samples revealed the presence of the species from western Lake Erie in 1994 and possibly as early as 1993. 2. Surveys conducted in Lake Erie between 1995 and 1998 revealed that Echinogammarus was the dominant amphipod on rocks covered by Dreissena molluscs compared with those fouled by the filamentous alga Cladophora, while Gammarus fasciatus used both Dreissena and Cladophora substrata extensively. 3. In laboratory habitat selection studies, Echinogammarus chose Dreissena‐ over Cladophora‐encrusted rocks and bare rocks, while Gammarus occupied the more complex substrata equally. 4. Field colonisation experiments demonstrated that the densities of Echinogammarus and Gammarus were positively correlated when the total density of the species was low, in contrast to the large‐scale natural distribution of the species that revealed a strong inverse relationship. 5. The on‐going replacement of Gammarus by Echinogammarus in the Laurentian Great Lakes may be related to the stronger affinity of the latter for substrata fouled by Dreissena, a genus with which it co‐evolved.     

Change in diel catchability of young-of-year yellow perch associated with establishment of dreissenid mussels

1. Non‐native mussels have increased water clarity in many lakes and streams in North America and Europe. Diel variation in catchability of some fish species has been linked to visibility during survey trawls (used to measure escapement). 2. Water clarity increased in nearshore areas of western Lake Erie by the early 1990s, following passage of legislation in 1972 to improve water quality (e.g. reduce phosphorus loading) and the invasion of dreissenid mussels (Dreissena spp.) beginning in 1987. 3. We hypothesised that increased water clarity in Lake Erie resulted in decreased catchability of young‐of‐year (age‐0) yellow perch (Perca flavescens Mitchill) during daylight compared to during night. We used a two‐tiered modelling approach to test this hypothesis on the ratio (R) of catch per hour (CPH) during night to CPH during daylight in bottom trawl surveys conducted during 1961–2005. 4. First, we examined seven a priori models. The first model, the ‘null’ model, represented no change in R over time. Three more models tested whether the timing of the change in R was associated with passage of water quality legislation only, dreissenids only (two‐period models) and both legislation and dreissenids (three‐period models). Three additional models included a 3‐year lag before the effects of legislation, dreissenids or both occurred. Secondly, all possible two‐ and three‐period models with a minimum of 2 years per time period were explored a posteriori. The a posteriori procedure determined the temporal transitions to higher R that were best supported by the data, without regard to a priori hypotheses. 5. Night CPH was greater than daylight CPH in 3 of 11 years during 1961–72, in 10 of 15 years during 1973–87, and in 14 of 18 years during 1988–2005. During 1991–2005 night CPH exceeded daylight CPH in all years except one, and night CPH was more than twice daylight CPH in 10 years during this period. 6. The best a priori model had two periods, with a break between 1990 and 1991, corresponding to 3 years after the dreissenid invasion. Similarly, the best two‐ and three‐period a posteriori models both had breaks between 1990 and 1991. The results supported our hypothesis that age‐0 yellow perch exhibited a transition to lower catchability during daylight compared to night, and the timing of the transition coincided with the establishment of dreissenid mussels. 7. The most plausible mechanism for our results was increased visibility of the trawl during daylight, resulting in increased avoidance of the trawl. These results have potential applications wherever non‐native mussels have increased water clarity.     

Mussels as ecosystem engineers: Their contribution to species richness in a rocky littoral community

Mussels are important ecosystem engineers in marine benthic systems because they aggregate into beds, thus modifying the nature and complexity of the substrate. In this study, we evaluated the contribution of mussels (Brachidontes rodriguezii, Mytilus edulis platensis, and Perna perna) to the benthic species richness of intertidal and shallow subtidal communities at Cerro Verde (Uruguay). We compared the richness of macro-benthic species between mussel-engineered patches and patches without mussels but dominated by algae or barnacles at a landscape scale (all samples), between tidal levels, and between sites distributed along a wave exposition gradient. Overall, we found a net increase in species richness in samples with mussels (35 species), in contrast to samples where mussels were naturally absent or scarce (27 species). The positive trend of the effect did not depend upon tidal level or wave exposition, but its magnitude varied between sites. Within sites, a significant positive effect was detected only at the protected site. Within the mussel-engineered patches, the richness of all macro-faunal groups (total, sessile and mobile) was positively correlated with mussel abundance. This evidence indicates that the mussel beds studied here were important in maintaining species richness at the landscape-level, and highlights that beds of shelled bivalves should not be neglected as conservation targets in marine benthic environments.     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.     

Thirty-Year Time Series of PCB Concentrations in a Small Invertivorous Fish (<Emphasis Type="Italic">Notropis Hudsonius</Emphasis>): An Examination of Post-1990 Trajectory Shifts in the Lower Great Lakes

The production and use of polychlorinated biphenyls (PCBs) have been restricted in North America since the 1970s; yet, PCBs are still detected in all components of the Great Lakes ecosystems. Our objective was to determine how total PCB (PCB_T) concentrations in spottail shiner (Notropis hudsonius) changed over the period 1975–2007 in the lower Great Lakes. Trends were best described by three basic models: (1) piecewise models where concentrations followed a decreasing trend before the break point (T) and an increasing trend post-T (Lake St. Clair, eastern Lake Erie, and upper Niagara River); (2) piecewise models where concentrations decreased both pre- and post-T but where the rate of decline post-T was less than that pre-T (western Lake Erie and Niagara River’s Tonawanda Channel); and (3) linear models where concentrations declined at a constant rate across the entire temporal range (lower Niagara River and western Lake Ontario). Piecewise models best described the trends in shallow areas that are susceptible to full water-column mixing whereas constant-slope models best described trends in deeper areas. For piecewise models, T typically occurred during the years 1988–1992. Two events coincided with this timing: (1) a sustained shift towards warming summer temperatures and (2) the proliferation of dreissenid mussels (Dreissena spp.). The weight-of-evidence suggests that the dreissenid invasions were a more likely driving factor behind the observed trends.     

Turbidity-mediated interactions between invasive filter-feeding mussels and native bioturbating mayflies

1. Invasive dreissenid mussels are known to cause large ecosystem changes because of their high filter‐feeding capacity, while native bioturbators may interfere with the mussels filter feeding. In this experiment, we investigated indirect environmental interactions between invasive filter‐feeding dreissenid mussels (zebra and quagga mussels) and native recolonizing bioturbating hexagenid mayflies (Hexagenia) at two mussel densities and two Hexagenia densities in a 2‐month long laboratory experiment. 2. Mean turbidity increased with increasing density of Hexagenia and decreased with increasing density of mussels. Turbidity showed the fastest decline at the highest mussel density, and no decline or a lower rate of decline at the low mussel density, dependent on Hexagenia density. 3. Mussel growth decreased with increasing Hexagenia density at low but not at high mussel density. Moreover, growth of mussels decreased as a function of increased mean turbidity at low mussel density but not at high mussel density. Filtering activity at the highest mussel density increased after introduction of food at the lower two densities of Hexagenia, but was constantly high at the highest Hexagenia density. 4. There was no difference in emergence of Hexagenia among the treatments, but mortality of Hexagenia was higher in the presence of mussels than in their absence. 5. Our results indicate that interactions between dreissenids and hexagenids are mediated through the sediment, and depend on density of both dreissenids and hexagenids. As the natural densities of these animals vary considerably within lakes, their growth and survival because of indirect environmental interactions is expected to vary spatially.     

Fouling mussels (Dreissena spp.) colonize soft sediments in Lake Erie and facilitate benthic invertebrates

• 1 We conducted survey and transplant studies to determine whether colonization and residency on soft sediments by introduced, fouling mussels (Dreissena polymorpha and D. bugensis) were affected by physical disturbance, and whether Dreissena presence in turn influenced the diversity and population densities of other benthic invertebrates. Surveys revealed that colony density was typically higher at moderate depths than at shallower and greater ones. However, the largest, midsummer colonies and greatest coverage of sediments by mussels occurred at deeper sites.
• 2 Disturbance of transplanted colonies varied by depth and colony size, with deeper and larger colonies experiencing the lowest destruction rates. Colony destruction rate was positively correlated with current velocity adjacent to the lakebed.
• 3 Absence of mussel colonies at shallow sites was not determined by recruitment or substrate limitation, as recruit density was higher and sediment characteristics more suitable for postveliger settlement at shallow than at deeper sites. Rather, seasonal storms have much stronger effects in shallow than in deep water.
• 4 Mussel residency on soft sediment has profound effects on invertebrate biodiversity. Invertebrate species (taxon) richness and total abundance were positively correlated with mussel colony area. Mussel‐sediment habitat supported between 462 and 703% more taxa, and between 202 and 335% more individuals (exclusive of Dreissena) than adjacent soft‐sediment lacking mussels.
• 5 Results from this study illustrate that physical disturbance directly limits the distribution of mussels on soft sediments, and the diversity and abundance of other benthic invertebrates in consequence.

     

Ecosystem engineering by invasive exotic beavers reduces in-stream diversity and enhances ecosystem function in Cape Horn,Chile

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers (Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A Review of the Biology and Ecology of the Quagga Mussel (Dreissena bugensis), a Second Species of Freshwater Dreissenid Introduced to North America

SYNOPSIS. North America's Great Lakes have recently been invadedby two genetically and morphologically distinct species of Dreissena.The zebra mussel (Dreissena polymorpha) became established inLake St. Clair of the Laurentian Great Lakes in 1986 and spreadthroughout eastern North America. The second dreissenid, termedthe quagga mussel, has been identified as Dreissena bugensisAndrusov, 1897. The quagga occurs in the Dnieper River drainageof Ukraine and now in the lower Great Lakes of North America.In the Dnieper River, populations of D. polymorpha have beenlargely replaced by D. bugensis; anecdotal evidence indicatesthat similar trends may be occurring in the lower LaurentianGreat Lakes. Dreissena bugensis occurs as deep as 130 m in theGreat Lakes, but in Ukraine is known from only 0–28 m.Dreissena bugensis is more abundant than D. polymorpha in deeperwaters in Dneiper River reservoirs. The conclusion that NorthAmerican quagga mussels have a lower thermal maximum than zebramussels is not supported by observations made of populationsin Ukraine. In the Dnieper River drainage, quagga mussels areless tolerant of salinity than zebra mussels, yet both dreissenidshave acclimated to salinities higher than North American populations;eventual colonization into estuarine and coastal areas of NorthAmerica cannot be ignored.