Ecological and phylogenetic aspects of the spring diet of three palaearctic species of swans.

The quality of swans' nutrition at spring migration stopovers is important for their successful breeding. It is of great interest to study the differences in nutrition of different swan species when sharing the same habitat. Microscopic analysis of Cygnus olor, C. cygnus, and C. columbianus bewickii feces collected in the eastern part of the Gulf of Finland in February-April 2014-2019 was performed. We measured food preferences of the three swan species using non-metric multidimensional scaling (NMDS). The width and overlap of dietary niches were also calculated. The diet of C. olor consists almost entirely of soft submerged aquatic vegetation, mainly macroalgae. Samples of the other two species except macroalgae contained large amounts of young shoots and roots of rigid semi-submerged and coastal vegetation. The dietary niche of C. cygnus is the most isolated because it is dominated by thick rhizomes of Phragmites australis, which are hardly used by other swan species. The diet of Bewick's swans was similar in many respects to that of the Mute swan, but Bewick's swans much more often preferred vegetative parts of submerged and semi-submerged plants, such as Stuckenia pectinata, Potamogeton perfoliatus, Sparganium sp., Nuphar lutea, and others. Notably, the dietary niches of Mute swan and Whooper swan overlapped as much as possible in February March during a period of severe food shortage, in contrast to later periods in spring when food was more abundant and varied. In general, differences in diets are well explained by differences in the morphology of birds. Comparison of tarsometatarsus indices shows that C. olor is the most water-related species. C. olor has the longest neck and its beak has the strongest filter features, whereas beaks of the other two species shows noticeable "goose-like grazing" features. Moreover, C. Cygnus has the most powerful beak. These features are due to the history of species. The formation of C. olor occurred during the Miocene-Pliocene of the Palaearctic in the warm eutrophic marine lagoons of the Paratethys with abundant soft submerged vegetation. The evolution of C. cygnus and C. c. bewickii took place in Pleistocene. At that time, periglacial and thermokarst water bodies on permafrost became widespread in the Palearctic, as well as dystrophic peat lakes with much poorer submerged aquatic vegetation, but well-developed coastal and semi-submerged vegetation.