Species Diversity and Driving Factors of Benthic and Zooplanktonic Assemblages at Different Stages of Thermokarst Lake Development: A Case Study in the Lena River Delta (Middle Siberia)

Global climate change might result in permafrost thaw and the formation of thermokarst landscapes that release long-term carbon stocks as greenhouse into the atmosphere, thereby initiating a positive climate feedback. These processes are mediated by biological activity, including by microbes, vascular plants and animals, whereas the role of invertebrates in thermokarst ecosystems remains poorly understood. We investigated the diversity and assemblage structures of zooplankton (mainly Copepoda, Cladocera), microbenthos (testate amoebae) and meio- (Copepoda and Cladocera) and macrozoobenthos (mollusks, crustaceans, insects and annelids) from a range of water bodies representing different stages of thermokarst lake formation in the southern part of the Lena River Delta (Central Siberia). Altogether, 206 species of testate amoeba, mollusk, crustacean, insect and annelid taxa were identified. A total of 60 species of macrozoobenthos (mainly insects) and 62 species of testate amoebae were detected in the water bodies of the Lena River Delta for the first time. The species richness of zooplankton and meio- and macrozoobenthos was greater in the large thermokarst lakes than in the polygonal ponds due to the freezing of the latter in the winter. In contrast, the species richness of protists was higher in the polygonal ponds, which was related to the habitat preferences of testate amoebae. Fish grazing strongly affected the macrobenthos assemblages but not the smaller-sized organisms. Water acidity and temperature were the main environmental drivers of the assemblage structure of testate amoeba and microcrustacean. The species structure of the macroinvertebrate assemblages was significantly explained by water acidity, permafrost depth and size of the water area. It means that small size organisms with their short generation times are sensitive to more dynamic factors such as temperature and may serve as indicators of ecosystem changes due to global climate warming. In contrast, large size organisms are affected by driven factors that appear during thermokarst lakes formation and permafrost degradation.