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.     

栖木在越冬红隼（Falco tinnunculus）的觅食地与捕食方式选择中的作用

栖木是重要的生境因子,对于鸟类生境选择和利用具有重要意义。2003年9月-2004年3月,采用固定样地对照观测的方法,分析研究栖木在越冬红隼（Falco tinnunculus）的栖息地选择与捕食方式选择中的作用。研究结果表明,增加了栖木的试验区（下称试验区）,红隼的出现频率远大于未增加栖木的对照区（下称对照区）;试验区红隼个体平均停留时间显著长于对照区;在试验区红隼栖停捕食占捕食次数的77.24%,而在对照区红隼只进行飞行捕食,红隼捕食方式和样地中是否增加栖木具有极显著相关性;试验区,红隼栖停行为时间占51.8%,大于对照区（30.1%）,对照区中,红隼在空中的行为如飞行、翱翔、悬停等行为时间所占比例为34.8%,远高于试验区（12.1%）。这些结果表明,栖木在越冬红隼的栖息地选择和捕食方式选择中具有重要的作用,栖木的存在是栖停捕食的先决条件,冬季红隼偏好于具有较好栖木条件的栖息地,并且偏向于采取低消耗低收益的栖停捕食。     

Using DNA barcoding to differentiate invasive Dreissena?species (Mollusca,Bivalvia)

The zebra mussel (Dreissena polymorpha) and the quagga mussel (Dreissena rostriformis bugensis) are considered as the most competitive invaders in freshwaters of Europe and North America. Although shell characteristics exist to differentiate both species, phenotypic plasticity in the genus Dreissena does not always allow a clear identification. Therefore, the need to find an accurate identification method is essential. DNA barcoding has been proven to be an adequate procedure to discriminate species. The cytochrome c oxidase subunit I mitochondrial gene (COI) is considered as the standard barcode for animals. We tested the use of this gene as an efficient DNA barcode and found that it allow rapid and accurate identification of adult Dreissena individuals.     

Colonisation by Dreissena of Great Lakes coastal ecosystems: how suitable are wetlands?

1.  Invasive zebra ( Dreissena polymorpha ) and quagga mussels ( Dreissena bugensis ) have become widespread throughout the Great Lakes basin. However, some types of Great Lakes coastal wetlands may be unsuitable for Dreissena invasion.
2.  To test this observation, artificial substrata were placed in wetlands (with emergent vegetation) and in adjacent open water (without emergent vegetation) habitats in two types of Great Lakes coastal ecosystems: drowned river mouth (DRM) and coastal fringing systems. Wetlands in DRM systems generally have deep organic sediment and limited water movement, whereas coastal fringing wetlands generally have low to moderate amounts of organic sediment and intense wind and wave action.
3.  We did not find a significant difference in Dreissena colonisation between wetlands and adjacent open water habitat in fringing systems. However, Dreissena colonisation was significantly lower in DRM wetlands than in the adjacent open water. We also found significantly lower survival in DRM wetlands than adjacent open water habitats, whereas survival did not differ significantly in coastal fringing wetlands and the adjacent open water.
4.  Our results suggest that vulnerability to Dreissena invasion varied among wetland types with DRM wetlands being less suitable than fringing wetlands. We suggest that colonisation and survival of Dreissena is lower in wetlands with deep organic sediment and less turbulent water.     

Overwinter changes in mass and lipid content of Perca fluviatilis and Gymnocephalus cernuus

The body condition, lipid reserves and mortality of 0 and 1 year‐old perch Perca fluviatilis and ruffe Gymnocephalus cernuus , sampled during the winter in Lake Constance, Germany, were compared. Length‐frequency analyses did not provide evidence for overwinter mortality in either species. The fresh and dry mass of perch as well as their lipid contents decreased during winter, while ruffe were heavier and contained more lipid at the end of the winter. The superior performance of ruffe was mainly attributed to its sensory capabilities, which allowed it to ingest zoobenthos throughout the winter, while the zooplankton feeding of perch was constrained by low light levels. In lakes that undergo a process of re‐oligotrophication, this advantage of ruffe over perch may be even more pronounced, since lower food supply during the growth season and thus lower fish lipid content at the start of winter is probably better tolerated by ruffe than by perch.     

Crayfish predation on the common pond snail (Lymnaea stagnalis): the effect of habitat complexity and snail size on foraging efficiency

Optimal foraging theory was used to explain selective foraging by the introduced signal crayfish (Pacifastacus leniusculus) on the thin-shelled common pond snail (Lymnaea stagnalis). Crayfish predation efficiency was studied in relation to habitat complexity and snail size. In a pool experiment (area 1.3 m²) single adult crayfish were allowed to feed on four size classes of snails for one week. A pair-wise preference trial (aquarium experiment) tested if adult crayfish selectively predated on particular size classes of snail and if prey value (expressed as snail dry mass per handling time) could explain the size range of snails chosen. Crayfish preferred the smallest size classes of snails in both pool and aquaria experiments. In the pool experiment crayfish had a strong effect on snail survival. Habitat complexity did not affect overall snail survival, but resulted in reduced predation pressure on the smallest size classes of snails. Handling time and shell-thickness increased exponentially with increasing snail size, and the two smallest size classes had the highest prey values. The results suggest that crayfish can structure the abundance and size distribution of thin-shelled snails, through size-selective predation and reduction of macrophytes. The mechanisms behind the choice of snails may be based on prey value and reduced exposure time to predators and conspecifics. Crayfish effects on snail size distribution may be less pronounced in complex habitats such as macrophyte beds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Seasonal effects of zebra mussels on littoral nitrogen transformation rates in Gull Lake, Michigan, U.S.A.

• 1 Zebra mussels (Dreissena polymorpha) are successful colonisers of lake littoral habitats and they interact strongly with littoral benthos. Previous research suggests that localised areas colonised by zebra mussels may be hotspots of nitrogen (N) cycling.
• 2 The effects of zebra mussels on nitrification and denitrification rates were examined approximately every other month for 1 year in Gull Lake, Michigan, U.S.A. Littoral sediment was collected from an area free of zebra mussels and distributed into shallow trays; rocks colonised with zebra mussels were placed in half of the trays, while uncolonised rocks were placed in the remaining trays. After an incubation period of 6–8 weeks in the lake, sediment and zebra mussels were collected from the trays, replaced with new sediment and zebra mussels, and placed in the lake for the next interval. In the laboratory, sediment nitrification and denitrification rates were measured for each tray.
• 3 Sediment nitrification rates did not increase in the presence of zebra mussels; instead nitrification rates were sensitive to changes in water temperature and increased with increasing exchangeable sediment ammonium. In contrast, denitrification rates increased in sediment trays with zebra mussels in the winter when nitrate (NO₃⁻) availability was high and when Chara did not grow in the trays.
• 4 Sediment denitrification was NO₃⁻‐limited in all seasons, regardless of zebra mussel treatment. However, sediment in the presence of zebra mussels responded less to NO₃⁻ addition, suggesting that NO₃⁻ limitation of denitrification can be reduced by zebra mussel activity. Zebra mussels have a seasonally variable impact on sediment denitrification rates, and this may translate into altered seasonal patterns of N cycling in localised areas of lakes where they are particularly abundant.

     

Specific amplification of the 18S rRNA gene as a method to detect zebra mussel (Dreissena polymorpha) larvae in plankton samples

An important issue in the management of zebra mussel (Dreissena polymorpha) populations is early, rapid, and accurate detection of the planktonic larvae (veliger) of the zebra mussel. The goal of this study was to explore the feasibility of developing a molecular approach for the detection of zebra mussel larvae in diverse environments. In this study a Dreissena polymorpha-specific 18S ribosomal RNA gene targeted oligonucleotide primer (ZEB-715a) and Polymerase Chain Reaction (PCR) assay was developed and compared with cross-polarized microscopy as a means to detect zebra mussel veligers in plankton samples. The design of the zebra mussel-specific primer was facilitated by sequencing nearly the complete 18S rRNA gene from the zebra mussel and three other closely related freshwater Veneroids including the quagga mussel (D. bugensis), the dark false mussel (Mytilopsis leucophaeata), and the Asian freshwater clam (Corbicula fluminea). The specificity of the primer for the zebra mussel was empirically tested by using the primer as a direct probe in a blot hybridization format. A single veliger in a plankton sample could be detected by PCR using this approach. Veliger detection sensitivity using the PCR approach was estimated to be over 300 times more sensitive than cross-polarized light microscopy based techniques. Cross-polarized light microscopy and the PCR technique were used to identify the presence of zebra mussel larvae in plankton samples that were collected from a variety of natural and industrial water sources. Detection results (presence or absence) were generally consistent between the two methods. Although additional studies will be required before routine application of molecular based veliger detection technology is available, a long-term goal of this work is the application of molecular technology to the development of a field device for the routine detection and quantification of zebra mussel veligers.     

Amplified fragment length polymorphism (AFLP) analysis of the genetic structure of the zebra mussel, Dreissena polymorpha, in the Mississippi River

1. We predicted that zebra mussel, Dreissena polymorpha (Pallas), genetic structure in the Mississippi River would follow a model of invasive species genetics, which predicts low genetic structure among populations of recently established species. This prediction was upheld in our previous genetic study using allozymes, however, one locus yielded anomalous results. 2. We employed amplified fragment length polymorphism (AFLP) analysis as a neutral marker to assess the amount of genetic structure within and among populations, and as a test of expected population structure from both invasion genetic theory, and the results from our previous study. 3. There was greater spatial differentiation, as measured by F_st, observed using AFLP's than for allozymes (P < 0.001). There was no evidence that AFLP variation conformed to an isolation by distance model, and genetic relationships of populations, as measured by AFLP markers, were not similar to those detected in our allozyme survey. 4. The lack of concordance between these two genetic marker systems probably reflects their differential responses to drift, migration, and selection occurring during this rapid invasion. Strong population structure is counter to predictions that populations of invasive species will not be differentiated, as with observations based on allozyme markers. Therefore, newly established species may require genetic surveys using multiple marker systems to evaluate population structure.