Impacts of dreissenid mussel invasions on chlorophyll and total phosphorus in 25 lakes in the USA

1. Invasions of zebra and quagga mussels have had long‐term, large‐scale impacts on lake ecosystems in the USA as characterised by high abundance, broad‐scale spread and effective adaption to new environmental conditions. Due to their high filtering capacity, decreases in chlorophyll a (Chl) and total phosphorus (TP) concentrations have been reported in many affected lakes. 2. In 25 US lakes, we analysed the effects of dreissenid invasions on changes in Chl and TP concentrations, measured as the probability of a concentration decrease in the post‐invasion period and changes in Chl–TP relationships using Bayesian hierarchical regressions. We also examined whether changes in Chl and TP concentrations and in the Chl–TP relationship differed according to lake hydromorphology, such as mean depth or mixing status (mixed versus stratified lakes). 3. Our results showed that dreissenid invasions were often, but not always, associated with subsequent TP and Chl declines. Dreissenid effects on Chl and TP may be influenced by lake thermal structure. Decreases in Chl and TP were consistently found in mixed lakes where benthic–pelagic coupling is tight, while the effects were less predictable in stratified lakes. Within stratified lakes, Chl and TP reductions were more clearly discernible in deeper lakes with long water residence times. 4. Regression results demonstrated that a joint increase in slope and decrease in intercept and a tighter correlation of the Chl–TP relationship were likely to occur in dreissenid‐invaded lakes; this does not support the idea of a shift from bottom‐up to top‐down control of primary production. These results have important implications for management, suggesting that a relaxation of TP standards would be unwarranted. 5. Across lakes, the slope of the Chl–TP relationship for mixed lakes was substantially higher than that for stratified lakes before mussel invasion, indicating an important role of light in limiting primary production. The slope differences between mixed and stratified lakes decreased in the post‐invasion period, possibly because mussel filtration results in a relaxation of light limitation that is more pronounced in deeper, stratified lakes.     

Alteration of Ecosystem Function by Zebra Mussels in Oneida Lake: Impacts on Submerged Macrophytes

Dreissenid mussels (the zebra mussel Dreissena polymorpha and the quagga mussel D. bugensis) are ecosystem engineers that modify the physical environment by increasing light penetration. Such a change is likely to affect the distribution and diversity of submerged macrophytes. Filter-feeding by these mussels has been associated with increased water clarity in many North American and European lakes. In this study, we report the increase in water clarity of Oneida Lake, New York, USA, for 1975–2002 and argue that the increase was caused by zebra mussel invasion rather than declines in nutrients. Over the study period, although mean total phosphorus decreased significantly, the main increase in water clarity occurred after the zebra mussel invasion in 1991. The average depth receiving 1% surface light increased from 6.7 m to 7.8 m after the invasion of zebra mussels, representing a 23% areal expansion. The maximum depth of macrophyte colonization, as measured by diver and hydroacoustic surveys, increased from 3.0 m before the invasion of zebra mussels to 5.1 m after their establishment. In addition, macrophyte species richness increased, the frequency of occurrence increased for most species, and the composition of the macrophyte community changed from low-light–tolerant species to those tolerating a wide range of light conditions. Comparisons with observations reported in the literature indicate that increased light penetration alone could explain these changes in macrophyte distribution and diversity. Such changes will increase the importance of benthic primary production over pelagic production in the food web, thereby representing an overall alteration of ecosystem function, a process we refer to as “benthification”.     

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.     

The collapse of benthic macroalgal blooms in response to self‐shading

1. Our goal was to use physiological indicators [photosynthesis–irradiance (P–I) response, nutrient status], population level feedbacks (self‐shading) and ambient environmental conditions (dissolved nutrients, light, temperature) to improve our understanding of the seasonal and spatial population dynamics of Cladophora. 2. Cladophora grew in three distinct phases, rapid growth early in the season (May–July), a mid‐season population collapse (July–August) and autumn re‐growth. Across all sites and dates, mean net maximal photosynthesis [P_M (NET)] was 6.9 ± 3.9 mg O₂ g DM^?1 h^?1, and α was 0.055 ± 0.025 mg O₂ g DM^?1 μm photons^?1 m^?2. Mean values for critical irradiance (I_CR) and the half‐saturation light intensity (I_K), were 42.9 ± 32.1 and 189.3 ± 123.8 μm photons^?1 m^?2 s^?1 respectively. 3. At most sites growth was phosphorus‐limited. Values of α were significantly higher at a site influenced by a nutrient enriched river plume, where algal growth was phosphorus‐sufficient. 4. Photoinhibition was not apparent in any of our P–I experiments. Even if photoinhibition had been apparent during in vitro P–I experiments, population level photosynthetic rates in the field would be little affected because intense self‐shading restricts inhibiting irradiances to the upper few mm–cm of the algal canopy. 5. Our physiological (P–I response) experiments contradicted previous assertions that high ambient temperatures, or nutrient deficiency, were primary causes of mid‐summer sloughing. In our study, sloughing occurred simultaneously at nutrient enriched and nutrient deficient sites, at temperatures well below critical values found during in vitro experiments, and our indicator of physiological condition (P–I response) remained unchanged leading up to, or immediately after, the sloughing event. 6. Self‐shading can reduce the convexity of the P–I response within in vitro incubations, even when the amount of algal material is low. Our experiments used 0.08 g DM of algal material that formed clumps c. 1 cm thick. Under these conditions, we estimated negligible (<1%) effects on P_M, a 12% reduction in apparent values of α, and 14% and 17% increases in values of the α‐dependent terms I_CR and I_K, respectively. 7. Our results are consistent with the hypothesis that a population‐level negative feedback (self‐shading) is responsible for sloughing in dense macroalgal beds. Sloughing was probably inevitable once macroalgal bed density and thickness surpassed a critical threshold. Cells towards the base of the bed received insufficient light to maintain metabolic balance, began to decay and weaken, and became increasingly susceptible to physical detachment from shear stress.     

Susceptibility of quagga mussels (Dreissena rostriformis bugensis) to hot-water sprays as a means of watercraft decontamination

The recent spread of dreissenid mussels to various bodies of water in the western US has sparked interest by many state and federal agencies to develop protocols to stop further expansion. Quagga mussels (Dreissena rostriformis bugensis) are of particular importance as they are currently the most widespread dreissenid species in the region. This project examined the susceptibility of quagga mussels to hot-water sprays at different temperatures and durations of spray contact at Lake Mead (Nevada-Arizona, USA). Emersed adult quagga mussels were exposed to hot-water sprays at 20, 40, 50, 54, 60, 70, and 80°C for 1, 2, 5, 10, 20, 40, 80, and 160 s. Sprays at ≥60°C for 5 s were shown to be 100% lethal. Sprays of 54°C for 10 s, 50°C for 20 s, and 40°C for 40 s also resulted in 100% mortality. A spray temperature of 60°C for 5 s is recommended for mitigating fouling by quagga mussels.     

Invasive mussels modify the cycling,storage and distribution of nutrients and carbon in a large lake

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Nutrient management and structural shifts in fish assemblages: Lessons learned from an Area of Concern in Lake Ontario

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Relative Distributions of Dreissena bugensis and Dreissena polymorpha in the Lower Don River System,Russia

A survey was conducted in the lower Don River system in Russia to confirm the presence of Dreissena bugensis, and to compare its distribution relative to that of Dreissena polymorpha. In 1999 and 2001–2002, dreissenid mussels were collected at 15 sites in the main river, in connecting reservoirs, and in a major tributary, the Manych River. Collections were made near stations where long‐term monitoring data on total mineral (sum of principal ions) and calcium content were available. Both dreissenid species were found at all sites, with D. bugensis comprising 4–75% of all dreissenids at individual sites. D. bugensis was relatively more abundant than D. polymorpha in the Manych River where total mineral and calcium content was significantly higher than in the Don River, suggesting the two species may have different calcium requirements. Examination of archived samples indicated that D. bugensis was present in the Don River system as early as the 1980s, presenting the unresolved enigma of why D. bugensis has not displaced D. polymorpha as the dominant species as typically found over shorter time periods in other water bodies. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Spatially heterogeneous trends in nearshore and offshore chlorophyll a concentrations in lakes Michigan and Huron (1998–2013)

1. Nutrient abatement programmes have been successfully implemented around the globe to reduce nutrient loading into aquatic ecosystems. Concurrently, the worldwide spread of invasive filter feeders, such as dreissenid mussels, may alter nutrient dynamics in invaded systems by sequestering nutrients away from pelagic zones and reducing primary production in offshore areas. Such is the case in the Laurentian Great Lakes of North America, where decades of nutrient abatement and the establishment of dreissenid mussels have seemingly resulted in more oligotrophic conditions and altered spatial patterns of nutrient availability and primary production.
2. Recent studies have focused on whole lake trends in primary production despite spatial differences in tributary inputs, bathymetry, and other environmental conditions that can affect primary production in nearshore areas. Thus, we hypothesised that trends in nearshore chlorophyll concentrations in different areas may diverge in a manner consistent with spatial differences in nutrient input. To evaluate these differences in surface chlorophyll responses, we assessed temporal trends in four different areas of Lake Michigan and three different areas of Lake Huron.
3. We hypothesised that in lakes Huron and Michigan, nearshore zones have experienced slower declines of chlorophyll concentrations relative to offshore zones. To assess this hypothesis, we estimated temporal trends of surface water chlorophyll concentrations (a proxy for primary production) from satellite imagery from 1998 to 2013. We calculated average surface chlorophyll concentrations for 10-m depth intervals ranging from the shore (0–10 m) to offshore (>90 m) during representative months of May, July, and September. We then analysed these data to determine if long-term trends in surface chlorophyll varied by season, proximity to the shoreline, and water depth.
4. The rates of annual change in chlorophyll concentrations in nearshore areas were markedly different to offshore trends in both lakes. Chlorophyll concentrations declined overtime in offshore areas, but nearshore chlorophyll concentrations were either stable (in May) or increased (in July and September) throughout the time series.
5. Differences in chlorophyll concentrations among areas were prominent in Lake Michigan. While differences between the northern and southern basin have been previously documented, trends in chlorophyll concentrations also differed between the eastern and western sides of Lake Michigan. Despite similar bathymetry and geographic features to Lake Michigan, regional trends were not observed in Lake Huron.
6. The results of this study are generally consistent with the nearshore shunt hypothesis, which predicts that dreissenid filtering, nutrient re-suspension, and continued nutrient loading from tributaries can cause an increase in primary production in nearshore areas during periods of offshore oligotrophication. Thus, the localised effects of nutrient abatement programmes in a given lake will be influenced by complex interactions between lake bathymetry and the presence of non-native filter feeding organisms.