Variability in macrozoobenthic assemblages along a gradient of environmental conditions in the stream water of karst caves (Lower Shakuranskaya Cave, western Caucasus)

The fauna of the stream water in the Lower Shakuranskaya Cave in central Abkhazia, western Caucasus, was studied. This cave has a large inlet and an extended entrance ecotone area of approximately 60 m, which makes it a convenient area for studying macrozoobenthic assemblages across a gradient of environmental factors. The cave has 13 species of stygobionts, 10 species of stygophiles and 18 species of stygoxenes. The number of species and the abundance and biomass of stygobionts per station were the highest near the boundary of the photic zone, at a distance of 50-60 m from the entrance to the cave, and gradually decreased toward both the remote parts of the cavity and the cave exit. The most abundant stygobionts were gastropod mollusks of the Hydrobiidae family, and Xiphocaridinella shrimp comprised the main part of the biomass. It has been shown that the main environmental factors determining the distribution of macrozoobenthos are luminosity and distance from the entrance to a cave. According to the differences in their reactions to these environmental factors, several groups of species were identified. In addition, three main assemblages of macrozoobenthic species were described: (1) an assemblage of epigean species near the cave entrance area; (2) stygobionts in remote parts of the cave outside the photic zone; and (3) a mixed assemblage in the cave ecotone, where a faint light penetrates. The specific details related to the faunal structure in the ecotone of the cave are discussed, as well as active and passive methods by which stygoxenes invade underground cavities.