Multispecies fish tracking across newly created shallow and deep habitats in a forward-restored lake

Background: Freshwater fish communities typically thrive in heterogenous ecosystems that offer various abiotic conditions. However, human impact increasingly leads to loss of this natural heterogeneity and its associated rich fish communities. To reverse this trend, we need guidelines on how to effectively restore or recreate habitats for multiple fish species. Lake Markermeer in the Netherlands is a human-created 70,000-ha lake with a uniform 4 m-water depth, steep shorelines, high wind-induced turbidity, and a declining fish community. In 2016, a forward-looking restoration project newly created a 1000-ha five-island archipelago in this degrading lake, which offered new sheltered shallow waters and deep sand excavations to the fish community. Methods: In 2020, we assessed how omnivorous and piscivorous fish species used these new habitats by tracking 78 adult fish of five key species across local and lake-scales. We monitored spring arrival of adult fish and assessed local macro-invertebrate and young-of-the-year fish densities. Results: Adult omnivorous Cyprinidae and piscivorous Percidae arrived at the archipelago in early spring, corresponding with expected spawning movements. During the productive summer season, 12 species of young-of-the-year fish appeared along the sheltered shorelines, with particularly high densities of common roach (Rutilus rutilus) and European perch (Perca fluviatilis). This suggests the sheltered, shallow, vegetated waters formed new suitable spawning and recruitment habitat for the fish community. Despite highest food densities for adult fish in the shallowest habitats (&lt; 2-m), adult fish preferred minimally 2-m deep water. After spawning most Cyprinidae left the archipelago and moved long distances through the lake system, while most Percidae remained resident. This may be related to (1) high densities of young-of-the-year fish as food for piscivores, (2) medium food densities for omnivores compared to elsewhere in the lake-system, or (3) the attractiveness of 30-m deep sand excavations that were newly created and frequently used by one-third of all tracked fish. Conclusions: New littoral zones and a deep sand excavation constructed in a uniform shallow lake that lacked these habitat types attracted omnivorous and piscivorous fish species within four years. Both feeding guilds used the littoral zones for reproduction and nursery, and notably piscivorous fish became residents year-round.</p

Seed dispersal by waterbirds: a mechanistic understanding by simulating avian digestion

Waterbirds disperse plant species via ingestion and egestion of seeds (endozoochory). However, our understanding about the regulating effects of seed traits, underlying mechanisms and possible (co)evolutionary processes is limited by our traditional reliance on data from feeding experiments with living waterbirds. Here, we overcome these limitations by developing and applying a new bioassay that realistically simulates digestive processes for Anseriformes waterbirds. We test three hypotheses: 1) seed survival and germination are most affected by mechanical digestion in the waterbird gizzard; 2) seed size, hardness, imbibition and shape regulate seed survival; and 3) plants growing in aquatic habitats benefit most from endozoochory by waterbirds. Experiments with 28 200 seeds of 48 plant species demonstrated species-specific seed survival that was entirely determined by digestion in the avian gizzard. Intestinal digestion did not affect seed survival but affected seed establishment (germinability and germination time) for 21% of the species. Large, hard seeds survived the simulations the best, in contrast to generally higher seed survival for smaller seeds during in vivo experiments. This mechanistically explains that small seeds escape digestive processes rather than being inherently more resistant (the ‘escape mechanism'), while large seeds are retained until fully digested or regurgitated (the ‘resistance and regurgitation mechanism'). Plants growing in wetter habitats had similar seed survival, but digestive processes stimulated their germinability and accelerated their germination more than for terrestrial plants. This indicates a relative advantage of endozoochory for plant species growing in wet habitats, possibly reflecting a co-evolutionary response related to dormancy breaking by gut passage. Simulating seed gut passage using a bioassay allowed establishing mechanisms and identifying relevant seed traits involved in seed dispersal by waterbirds. This information enhances our understanding of how animal species shape plant species distributions, which is extremely relevant now that current anthropogenic pressures already severely impact plant dispersal capacities

Animal-Mediated Dispersal in Understudied Systems

Peer reviewe

Direct and indirect effects of native plants and herbivores on biotic resistance to alien aquatic plant invasions

Biotic resistance to alien plant invasions is mainly determined by ecological interactions in two layers of the food web: competition with native plant species and herbivory by native herbivores. While the direct effect of native plants on alien plant performance via competition has been well documented across ecosystems, less is known about the direct and indirect effects of herbivores in providing biotic resistance. Our main aims were to determine whether temperate native aquatic plants and herbivores can provide biotic resistance to plant invasions, understand the underlying mechanisms and search for potential interactive effects of competition and herbivory on invader performance (i.e. growth). We mimicked natural temperate mesotrophic and eutrophic freshwater lakes in mesoscosms, by growing three native submerged plant species in monocultures (Ceratophyllum demersum, Myriophyllum spicatum and Potamogeton perfoliatus) at three competition levels (no, low and high) without and with the native aquatic generalist snail Lymnaea stagnalis. We subsequently simulated an early stage of establishment of the South American highly invasive alien plant species Egeria densa. We found that competition by native plant biomass significantly reduced invader performance but depended on native species identity. Herbivory had no direct negative effect on invader performance as the snails fed mainly on the available filamentous algae, which are commonly found in meso- and eutrophic systems, instead of on the plants. However, the consumption of filamentous algae by herbivores indirectly had positive effects on the invader total biomass, thus facilitating the invasion by E. densa. Nonetheless, these indirect effects worked through different pathways depending on the native plant identity. Synthesis. We found evidence for biotic resistance through competition by native plant species. However, we show that herbivores can indirectly facilitate South American plant E. densa invasion promoting its growth through selective feeding on filamentous algae, but this effect depends on the native plant species involved. Our experiment illustrates the important role of indirect interactions to understand the potential of biotic resistance in natural ecosystems

Impacts of shelter on the relative dominance of primary producers and trophic transfer efficiency in aquatic food webs: Implications for shallow lake restoration

Wind-induced turbulence can strongly impact ecological processes in shallow lake ecosystems. The creation of shelter against wind can be expected to affect both primary producers and herbivores in aquatic food webs. Shelter may benefit particular primary producers more than others by changing relative resource availabilities for different primary producers. Herbivore community compositions may be affected either directly or indirectly as a consequence of changes in their food quantity and quality that, in turn, may affect the transfer efficiency between primary producers and herbivores. A reduction in trophic transfer efficiency resulting from wind-induced turbulence potentially can lead to declines of higher trophic levels, but is generally understudied. Here, we focus on the impact of wind on aquatic primary producers and trophic transfer efficiency. We hypothesised that reducing wind-induced turbulence will stimulate higher trophic production in shallow lakes. However, the multitude of impacts of wind-induced turbulence on aquatic food webs make it challenging to predict the direction of change when creating sheltered conditions. We tested our hypothesis in the shallow waters of a newly constructed archipelago named Marker Wadden in lake Markermeer in the Netherlands. Lake Markermeer has experienced declining numbers of birds and fish. These declines have been related to wind-induced sediment resuspension that potentially limits primary production and trophic transfer efficiency. Marker Wadden is a large-scale restoration project that aims to add sheltered and heterogeneous habitat to the otherwise mostly homogeneous lake, thus targeting the potential problems associated with wind-induced turbulence. We executed a 2-month manipulative field mesocosm experiment in the shallow waters of Marker Wadden to study the effect of reduced wind-induced turbulence (i.e., shelter) on aquatic food webs. Specifically, we studied the effects on primary producers, trophic transfer efficiency between phytoplankton and zooplankton (using zooplankton biomass divided by phytoplankton Chl a as a proxy), and benthic fauna. The experiment consisted of three treatments: no shelter, shelter without macrophytes and shelter with submerged macrophytes (Myriophyllum spicatum) present at the start of the experiment. Our results clearly showed that under unsheltered conditions phytoplankton was the dominant primary producer, whereas in sheltered conditions submerged macrophytes became dominant. Interestingly, submerged macrophytes appeared rapidly in the sheltered treatment where first no macrophytes were visibly present; hence, at the end of the experiment, there was little difference among the sheltered treatments with and without initial presence of submerged macrophytes. Despite that phytoplankton concentrations were 23-fold higher under the unsheltered conditions, this did not result in higher zooplankton biomass. This can be explained by a five-fold greater trophic transfer efficiency between phytoplankton and zooplankton under the sheltered conditions. Furthermore, under the sheltered conditions the Gastropoda density reached 746 individuals m−2, whereas no Gastropoda were found under the no shelter treatment. These findings indicate that for shallow lakes that are negatively affected by wind-induced turbulence, measures aimed at ameliorating this stressor can be effective in facilitating submerged macrophyte recovery, increasing Gastropoda densities and restoring trophic transfer efficiency between phytoplankton and zooplankton. Ultimately, this may support higher trophic levels such as fish and water birds by increasing their food availability in shallow lake ecosystems

Impacts of shelter on the relative dominance of primary producers and trophic transfer efficiency in aquatic food webs : Implications for shallow lake restoration

Wind-induced turbulence can strongly impact ecological processes in shallow lake ecosystems. The creation of shelter against wind can be expected to affect both primary producers and herbivores in aquatic food webs. Shelter may benefit particular primary producers more than others by changing relative resource availabilities for different primary producers. Herbivore community compositions may be affected either directly or indirectly as a consequence of changes in their food quantity and quality that, in turn, may affect the transfer efficiency between primary producers and herbivores. A reduction in trophic transfer efficiency resulting from wind-induced turbulence potentially can lead to declines of higher trophic levels, but is generally understudied. Here, we focus on the impact of wind on aquatic primary producers and trophic transfer efficiency. We hypothesised that reducing wind-induced turbulence will stimulate higher trophic production in shallow lakes. However, the multitude of impacts of wind-induced turbulence on aquatic food webs make it challenging to predict the direction of change when creating sheltered conditions. We tested our hypothesis in the shallow waters of a newly constructed archipelago named Marker Wadden in lake Markermeer in the Netherlands. Lake Markermeer has experienced declining numbers of birds and fish. These declines have been related to wind-induced sediment resuspension that potentially limits primary production and trophic transfer efficiency. Marker Wadden is a large-scale restoration project that aims to add sheltered and heterogeneous habitat to the otherwise mostly homogeneous lake, thus targeting the potential problems associated with wind-induced turbulence. We executed a 2-month manipulative field mesocosm experiment in the shallow waters of Marker Wadden to study the effect of reduced wind-induced turbulence (i.e., shelter) on aquatic food webs. Specifically, we studied the effects on primary producers, trophic transfer efficiency between phytoplankton and zooplankton (using zooplankton biomass divided by phytoplankton Chl a as a proxy), and benthic fauna. The experiment consisted of three treatments: no shelter, shelter without macrophytes and shelter with submerged macrophytes (Myriophyllum spicatum) present at the start of the experiment. Our results clearly showed that under unsheltered conditions phytoplankton was the dominant primary producer, whereas in sheltered conditions submerged macrophytes became dominant. Interestingly, submerged macrophytes appeared rapidly in the sheltered treatment where first no macrophytes were visibly present; hence, at the end of the experiment, there was little difference among the sheltered treatments with and without initial presence of submerged macrophytes. Despite that phytoplankton concentrations were 23-fold higher under the unsheltered conditions, this did not result in higher zooplankton biomass. This can be explained by a five-fold greater trophic transfer efficiency between phytoplankton and zooplankton under the sheltered conditions. Furthermore, under the sheltered conditions the Gastropoda density reached 746 individuals m−2, whereas no Gastropoda were found under the no shelter treatment. These findings indicate that for shallow lakes that are negatively affected by wind-induced turbulence, measures aimed at ameliorating this stressor can be effective in facilitating submerged macrophyte recovery, increasing Gastropoda densities and restoring trophic transfer efficiency between phytoplankton and zooplankton. Ultimately, this may support higher trophic levels such as fish and water birds by increasing their food availability in shallow lake ecosystems

Impacts of sediment resuspension on phytoplankton biomass production and trophic transfer: Implications for shallow lake restoration

Wind-induced sediment resuspension in shallow lakes may enhance nutrient availability while reducing light availability for phytoplankton growth, thereby affecting the entire food-web. Lake restoration projects that reduce wind-induced resuspension are expected to enhance trophic transfer efficiencies, thereby improving food-web structure and functioning. Yet, reduced resuspension may also lead to lower nutrient concentrations in the water column, promote benthic algae development, reduce phytoplankton biomass production and thereby reduce secondary production by zooplankton. Lake Markermeer is a shallow delta lake in The Netherlands subject to wind-induced sediment resuspension. Restoration project Marker Wadden consists of newly built islands aiming to reduce sediment resuspension and promote higher trophic levels. Here, we tested the effects of reduced sediment resuspension on phytoplankton biomass build-up, benthic algae development, and zooplankton abundances at different temperatures in a 14-day indoor microcosm experiment. We used Marker Wadden sediment with three resuspension intensities combined with three temperatures, to also test effects of higher temperatures in shallow sheltered waters. Reduced sediment resuspension decreased nutrient concentrations and phytoplankton biomass build-up, while increasing light availability and enhancing benthic algae biomass development. Reduced sediment resuspension furthermore increased zooplankton biomass. Enhanced sediment resuspension and higher temperatures synergistically interacted, maintaining a high level of inorganic suspended solids. Our experimental results are in line with long-term seasonal observations from Lake Markermeer. Our findings demonstrate that for shallow lakes suffering from wind effects, measures such as Marker Wadden aimed at reducing sediment resuspension can be effective in restoring secondary production and supporting higher trophic levels

Paai- en opgroeigebieden voor vis in en rond Marker Wadden

Wageningen Marine Research rapport C058/21 | 5 van 66 Samenvatting Marker Wadden is een eilandengroep die sinds 2016 aangelegd wordt in het Markermeer. In het kader van het Kennis- en Innovatieprogramma Marker Wadden (KIMA) is onderzocht hoe de nieuw gevormde habitats op en rond Marker Wadden worden benut als paaigebieden en opgroeigebieden voor jonge vis en op die manier bijdragen aan de visproductie en voedselbasis voor visetende watervogels. Dit rapport vormt een achtergronddocument voor het KIMA-syntheserapport 2022 (KIMA, concept 2021) dat het onderzoek in de eerste 5 jaar van het ontstaan van de archipel samenvat. Voor het onderzoek naar paai- en opgroeigebieden voor vis zijn intensieve visstandbemonsteringen uitgevoerd naar het voorkomen en de abundantie van vislarven en jonge vis in het voorjaar en de zomer (april/mei-september) van 2019 (eerste eiland,) en 2020 en 2021 (alle eilanden). In juni 2020 is tevens eenmaal een bemonstering uitgevoerd in de nieuwe habitats van Trintelzand. Waar mogelijk werd wadend in ondiepe oeverzones bemonsterd met respectievelijk een larvennet, RAVON-net en een broedzegen. Bij instabiele bodems (met name in de eerste periode na aanleg) werd ook vanuit Canadese kano’s gevist. In 2020 en 2021 zijn de bemonsteringen in samenwerking met onderzoeksprogramma Natuur in Productie (Nederlands Instituut voor Ecologie, Rijksuniversiteit Groningen en Radboud Universiteit) uitgevoerd. Determinatie van jonge vis gebeurde grotendeels in het veld. Kleine larven werden verzameld op formol voor nadere identificatie in het lab (microscoop). In aanvulling op het onderzoek naar paai- en opgroeigebieden rond de eilanden van Marker Wadden zijn in 2018 in het westelijk Markermeer verschillende bemonsteringsmethoden getest voor het vaststellen van soortsamenstelling van vis in en tussen fonteinkruidvelden, een type habitat dat zich op grote schaal ontwikkelt tussen de eilanden van Marker Wadden en Trintelzand. In augustus 2019 is daarnaast nog het voorkomen van vis in zandwinputten rond Marker Wadden onderzocht. Belangrijke habitats voor productie van jonge vis zijn ondiepe oeverzones, met name zones met een combinatie van oevervegetatie (riet en moerasandijvie) en onderwatervegetatie (fonteinkruid, aarvederkruid, kranswier), terwijl kale zand- en sliboevers vooralsnog relatief weinig te bieden hebben voor de meeste soorten jonge vis. Trintelzand en Marker Wadden hebben vergelijkbare functies doordat het beschutte gebieden zijn met geleidelijke land-water overgangen zoals die elders in het Markermeer weinig te vinden zijn. Dit heeft meerwaarde als paai- en opgroeigebied voor vele soorten vis. Binnen deze gebieden is er een grote variatie in habitats en ecologische waarde, ook tussen seizoenen en jaren, mede als gevolg van variatie in ontwikkeling van de watertemperatuur in het voorjaar en van wind die fluctuaties in waterpeil veroorzaakt (Marker Wadden zijn “windwadden”). De zandwinputten (met een diepte van 10-30 m) bieden een additioneel type habitat wat door zowel grote vis (met name snoekbaars) als kleine vis (met name spiering en jonge baars) gebruikt wordt. Het gebied bevindt zich nog in een pionierssituatie maar zal zich waarschijnlijk verder ontwikkelen richting een door waterplanten gedomineerde toestand, zowel oevers als de geleidelijke land- waterovergangen als de diepere delen van het meer in de luwte van de eilanden. In bemonsteringen met stortkuil en zegen werd daar een variatie aan jonge vis, inclusief (kleinere) roofvis als jonge snoekbaars en baars aangetroffen. De belangrijkste conclusie tot dusver is dat geleidelijke land-waterovergangen met flauwe taluds in de luwte van eilanden zorgen voor een mozaïek van vegetatierijke oeverzones voor paaien en ondiepe baaien met ondergedoken vegetatie die van belang zijn voor de productie van jonge vis. Op de schaal van het Markermeer werd de afgelopen jaren echter een zeer lage visstand waargenomen waarbij de aanleg van kale zandoevers op voorheen relatief visrijke stenen oevers, alsmede baggerwerkzaamheden, vermoedelijk een rol van betekenis hebben gespeeld. Het is belangrijk te noteren dat het tot dusver gaat om pionierssituaties die in ontwikkeling zijn. Dit zijn langjarige processen waarbij verschillende vormen van beheer mogelijk zijn die uiteindelijk zullen bepalen welke habitatontwikkeling en welke visproductie gerealiseerd kan worden