Escaping malnutrition by shifting habitats: A driver of three-spined stickleback invasion in Lake Constance.

Fatty acids, and especially long-chain polyunsaturated fatty acids, are biologically important components in the metabolism of vertebrates, including fish. Essential fatty acids (EFA) are those that in a given animal cannot be synthesized or modified from precursors and must therefore be acquired via the diet. Because EFAs are often unevenly distributed in nature, this requirement may drive species to make behavioral or ecological adaptations to avoid malnutrition. This is especially true for fish like the three-spined stickleback (Gasterosteus aculeatus L.) of Upper Lake Constance (ULC), whose recent marine ancestors evolved with access to EFA-rich prey, but which found themselves in an EFA-deficient habitat. An unexpected and unprecedented ecological shift in the ULC stickleback population from the littoral to pelagic zones in 2012 might be linked to EFA availability, triggering ecological release and enabling them to build a hyperabundant population while displacing the former keystone species, the pelagic whitefish Coregonus wartmanni. To test this hypothesis, sticklebacks from the littoral and pelagic zones of ULC were sampled seasonally in two consecutive years, and their stomach contents and fatty acid profiles were analysed. Pelagic sticklebacks were found to possess significantly higher values of an important EFA, docosahexaenoic acid (DHA), especially during autumn. Evaluation of the DHA supply suggests that sticklebacks feeding in the littoral zone during autumn could not meet their DHA requirement, whereas DHA availability in the pelagic zone was surplus to demand. During autumn, pelagic sticklebacks consumed large amounts of DHA-rich prey, that is, copepods, whereas littoral sticklebacks relied mainly mostly on cladocerans, which provide much lower quantities of DHA. Access to pelagic zooplankton in 2012 was possibly facilitated by low densities of previously dominant zooplanktivorous whitefish. The present study offers a convincing physiological explanation for the observed expansion of invasive sticklebacks from the littoral to the pelagic zones of Lake Constance, contributing to a phase shift with severe consequences for fisheries.

Cryptic persistence and loss of local endemism in Lake Constance charr subject to anthropogenic disturbance.

In the welcome circumstance that species believed extinct are rediscovered, it is often the case that biological knowledge acquired before the presumed extinction is limited. Efforts to address these knowledge gaps, in particular to assess the taxonomic integrity and conservation status of such species, can be hampered by a lack of genetic data and scarcity of samples in museum collections. Here, we present a proof-of-concept case study based on a multidisciplinary data evaluation approach to tackle such problems. The approach was developed after the rediscovery, 40 years after its presumed extinction, of the enigmatic Lake Constance deep-water charr Salvelinus profundus. Targeted surveys led to the capture of further species and additional sympatric normal charr, Salvelinus cf. umbla. Since the lake had been subject to massive stocking in the past, an evaluation of the genetic integrity of both extant forms was called for in order to assess possible introgression. A two-step genomic approach was developed based on restriction site associated DNA (RAD). Diagnostic population genomic (single nucleotide polymorphism [SNP]) data were harvested from contemporary samples and used for RNA bait design to perform target capture in DNA libraries of archival scale material, enabling a comparison between extant and historic samples. Furthermore, life history traits and morphological data for both extant forms were gathered and compared with historical data from the past 60-120 years. While extant deep-water charr matched historical deep-water specimens in body shape, gill raker count, and growth rates, significant differences were discovered between historical and extant normal charr. These resulted were supported by genomic analyses of contemporary samples, revealing the two extant forms to be highly divergent. The results of population assignment tests suggest that the endemic deep-water charr persisted in Lake Constance during the eutrophic phase, but not one of the historical genomic samples could be assigned to the extant normal charr taxon. Stocking with non-endemic charr seems to be the most likely reason for these changes. This proof-of-concept study presents a multidisciplinary data evaluation approach that simultaneously tests population genomic integrity and addresses some of the conservation issues arising from rediscovery of a species characterized by limited data availability.

Perceived multiple stressor effects depend on sample size and stressor gradient length.

Multiple stressors are continuously deteriorating surface waters worldwide, posing many challenges for their conservation and restoration. Combined effect types of multiple stressors range from single-stressor dominance to complex interactions. Identifying prevalent combined effect types is critical for environmental management, as it helps to prioritise key stressors for mitigation. However, it remains unclear whether observed single and combined stressor effects reflect true ecological processes unbiased by sample size and length of stressor gradients. Therefore, we examined the role of sample size and stressor gradient lengths in 158 paired-stressor response cases with over 120,000 samples from rivers, lakes, transitional and marine ecosystems around the world. For each case, we split the overall stressor gradient into two partial gradients (lower and upper) and investigated associated changes in single and combined stressor effects. Sample size influenced the identified combined effect types, and stressor interactions were less likely for cases with fewer samples. After splitting gradients, 40 % of cases showed a change in combined effect type, 30 % no change, and 31 % showed a loss in stressor effects. These findings suggest that identified combined effect types may often be statistical artefacts rather than representing ecological processes. In 58 % of cases, we observed changes in stressor effect directions after the gradient split, suggesting unimodal stressor effects. In general, such non-linear responses were more pronounced for organisms at higher trophic levels. We conclude that observed multiple stressor effects are not solely determined by ecological processes, but also strongly depend on sampling design. Observed effects are likely to change when sample size and/or gradient length are modified. Our study highlights the need for improved monitoring programmes with sufficient sample size and stressor gradient coverage. Our findings emphasize the importance of adaptive management, as stress reduction measures or further ecosystem degradation may change multiple stressor-effect relationships, which will then require associated changes in management strategies.

Stickleback mass occurrence driven by spatially uneven parasite pressure? Insights into infection dynamics, host mortality, and epizootic variability.

Since 2012, a massive invasion of the three-spined stickleback (Gasterosteus aculeatus) has taken place into the pelagic area of Lake Constance. This species, which had previously been restricted to the littoral zone, is now the dominant pelagic fish and the previously dominant whitefish (Coregonus wartmanni) has suffered severe reductions in growth and recruitment. In this study, in total, 2871 sticklebacks were collected via monthly sessions over a 4-year period in pelagic and benthic areas of Lake Constance and examined for signs of infection with Schistocephalus solidus, a parasite known to be potentially fatal. The infection risk to sticklebacks increases throughout the course of the year and is size- and sex-dependent. Habitat has only a marginal impact. All parasite-induced harm is imparted after stickleback spawning and parental care is over. The results did not support the hypothesis that the invasion of the pelagic area might be driven by parasite-avoiding behaviour. Furthermore, the impact of the parasite is likely to be limited to post-reproductive adults, thereby ensuring stable reproduction of the hosts despite high rates of transmission and mortality. In consequence, stickleback stock development is independent of S. solidus infection, leading to secure coexistence of host and parasite even at extraordinary high host levels.

Size matters? Species- and size-specific fish predation on recently established invasive quagga mussels Dreissena rostriformis bugensis Andrusov 1897 in a large, deep oligotrophic lake.

Since its first appearance in Lake Constance in 2016 the invasive quagga mussel Dreissena rostriformis bugensis has come to dominate the mussel community and now occurs in hyperabundant densities over the whole lake bottom. A lake-wide field study was conducted between 2019 and 2020 to obtain a systematic insight into the importance of this novel source of potential prey for the native fish community. In total 664 fish of 20 different species were caught and their digestive tracts were analysed. Meanwhile, quagga mussels were sampled to assess their calorific energy value by size and season. Regressions of septum length on maximum valve length were used to evaluate size-specific mussel consumption by the three dominant quagga-eating fish species. The study shows that nearly all benthivorous fish are able to forage efficiently on quaggas. However, in the case of one keystone species with very high commercial interest, the benthic whitefish Coregonus spp., quagga consumption is more limited as only individuals larger than 35 cm consumed quaggas in relevant amounts. The energy content of quagga mussels is positively size dependent with seasonal effect and elevated values during summer for medium- and large-sized mussels. Even at its peak, the calorific value of quaggas by weight is much lower than that of endemic aquatic invertebrates, while single mass is high. Future implications for fish stocks, food web integrity and possible fishery management options are discussed.

All day-long: Sticklebacks effectively forage on whitefish eggs during all light conditions.

The three-spined stickleback Gasterosteus aculeatus invaded Lake Contance in the 1940s and expanded in large numbers from an exclusively shoreline habitat into the pelagic zone in 2012. Stickleback abundance is very high in the pelagic zone in winter near the spawning time of pelagic whitefish Coregonus wartmanni, and it is hypothesized that this is triggered by the opportunity to consume whitefish eggs. Field sampling has qualitatively confirmed predation of whitefish eggs by stickleback, but quantification has proven difficult due to stormy conditions that limit sampling. One fundamental unknown is if freshwater stickleback, known as visual feeders, can successfully find and eat whitefish eggs during twilight and night when whitefish spawn. It is also unknown how long eggs can be identified in stomachs following ingestion, which could limit efforts to quantify egg predation through stomach content analysis. To answer these questions, 144 individuals were given the opportunity to feed on whitefish roe under daylight, twilight, and darkness in controlled conditions. The results showed that stickleback can ingest as many as 100 whitefish eggs under any light conditions, and some individuals even consumed maximum numbers in complete darkness. Furthermore, eggs could be unambiguously identified in the stomach 24 hours after consumption. Whitefish eggs have 28% more energy content than the main diet of sticklebacks (zooplankton) based on bomb-calorimetric measurements, underlining the potential benefits of consuming eggs. Based on experimental results and estimates of stickleback abundance and total egg production, stickleback could potentially consume substantial proportions of the total eggs produced even if relatively few sticklebacks consume eggs. Given the evidence that stickleback can feed on eggs during nighttime spawning and may thereby hamper recruitment, future studies aimed at quantifying actual egg predation and resulting effects on the whitefish population are urgently needed.

Differentiation of two swim bladdered fish species using next generation wideband hydroacoustics.

Monitoring fish populations in large, deep water bodies by conventional capture methodologies requires intensive fishing effort and often causes mass mortality of fish. Thus, it can be difficult to collect sufficient data using capture methods for understanding fine scale community dynamics associated with issues such as climate change or species invasion. Hydroacoustic monitoring is an alternative, less invasive technology that can collect higher resolution data over large temporal and spatial scales. Monitoring multiple species with hydroacoustics, however, usually requires conventional sampling to provide species level information. The ability to identify the species identity of similar-sized individuals using only hydroacoustic data would greatly expand monitoring capabilities and further reduce the need for conventional sampling. In this study, wideband hydroacoustic technology was used in a mesocosm experiment to differentiate between free swimming, similar-sized individuals of two swim-bladdered species: whitefish (Coregonus wartmanni) and stickleback (Gasterosteus aculeatus). Individual targets were identified in echograms and variation in wideband acoustic responses among individuals, across different orientations, and between species was quantified and visually examined. Random forest classification was then used to classify individual targets of known species identity, and had an accuracy of 73.4% for the testing dataset. The results show that species can be identified with reasonable accuracy using wideband hydroacoustics. It is expected that further mesocosm and field studies will help determine capabilities and limitations for classifying additional species and monitoring fish communities. Hydroacoustic species differentiation may offer novel possibilities for fisheries managers and scientists, marking the next crucial step in non-invasive fish monitoring.

Quantification of alizarin red S uptake in coregonid eggs after mass-marking.

An easy method to measure the uptake rate of the persistent dye alizarin red S (ARS) during marking of whitefish eggs was established and used to measure the ARS content in three different whitefish species during and at the end of the marking procedure. Those values show that only 6-10% of the ARS in the marking solution will be absorbed by the eggs (0.0061-0.0119 mg per egg). Additional analyses 6, 15 and 36 months after marking showed ARS levels below the response level (<6.9 µg kg-1 ).

Invasion strategy and abiotic activity triggers for non-native gobiids of the River Rhine.

The 24 hour activity patterns of three non-native gobiids (round goby Neogobius melanostomus, Western tubenose goby Proterorhinus semilunaris and bighead goby Ponticola kessleri) were assessed over 46 consecutive months between 2011 and 2014 from their occurrence in the cooling water intake of a nuclear power plant on the River Rhine, Germany. In total, 117717 gobiids were identified and classified. The occurrence of all three species varied strongly between sampling years, and species-specific activity triggers were identified. The activity of juveniles of all three gobiids species was positively temperature dependent while adult tubenose goby activity appeared to be negatively temperature dependent. Increasing fluvial discharge in the adjoining main river stimulated the activity of juvenile round goby but inhibited activity of adult tubenose goby. Except for adult bighead goby, activity was also structured by time of day, but with no uniform mean. Meteorological factors such as precipitation, air pressure and duration of sunshine hours had little or no influence on gobiid activity. On selected rare occasions, mainly at night, all three species exhibited pulsed swarming behaviour, with thousands of individuals recorded in the intake water. Round goby swarms exhibited both the highest intensity and the largest swarming individuals, suggesting a potential competitive advantage over tubenose and bighead goby. Electric fishing surveys in natural river stretches corroborated this observation. Negative effects on the native fish fauna were apparent only for the bullhead, Cottus gobio. The activity triggers identified offer a unique insight into the invasion mechanisms of these ecosystem-changing non-native gobiids.