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Morphotypes revision, population genetics, and demographic history of Sinotaia quadrata in East Asia

YE BIN 東北大学

2020.09.25

概要

The systematics of species in the genus Sinotaia (Gastropod: Viviparidae: Bellamyinae) has been a complex and controversial issue since the nineteenth century and continues to the present day. The type species Sinotaia quadrata exists in many transitional forms, and the validity of some species similar to S. quadrata requires confirmation. Thus, phylogenetic relationships for species related to S. quadrata were reconstructed based on 16S rRNA and mitochondrial cytochrome oxidase subunit I (COI) genes to reveal the invalid classifications of these species. The shell outline morphology was compared based on the type materials of most species in the genus Sinotaia from the initial publications. The admixture of phylogeny and uniform shape variance suggested that some species of Sinotaia should be revised to morphotypes instead of over-classified species. Here, microsatellite markers and landmark-based geometric morphometric analysis were used to examine the genetic differentiation and shell shape variations of S. quadrata in both China and Japan. An inconsistency between genetic variations and shape variations was detected, thus suggesting that using minor shell shape variations is an invalid way to classify each described instance of S. quadrata as an independent species. The shell shape variation has no clear separations but shows plasticity under various local environments. The molecular phylogeny, genetic diversity, and shell morphology provided evidence for revising some species related to S. quadrata, especially for the eight species widely distributed in China and East Asia.

Anthropogenic factors potentially affect observed biogeographical patterns in population genetics, but the effects of ancient human activities on the original patterns created by natural processes are unknown. A widely distributed freshwater snail species in East Asia, Sinotaia quadrata, was used to investigate this issue. It is unclear if S. quadrata in Japan was introduced from China, and how different human uses and varying geographic patterns affect the contemporary population genetics between the two regions. Thus, the demography of S. quadrata and its genetic structure in Japan and continental East Asia were detected. Results showed that S. quadrata populations first naturally migrated from continental East Asia to Japan, which is associated with the ancient period in Japanese geohistory (about 70,000 years ago). They were then artificially introduced in association with human movements in two recent periods (about 8000 and 1200 years ago). Populations in different parts of Japan have their own sources. Natural migration in the ancient period and artificial introduction in the recent period suggest that the population distribution is affected by both the geohistory of East Asia and the history of human expansion. In the background of the historical migration and introduction, contemporary populations in the two regions show different genetic patterns. Population divergence levels were significantly correlated with geographical patterns in Japan, and significantly correlated with human interventions variables in continental East Asia, suggesting that long-term geographical isolation is likely the major factor that shaped the contemporary population genetics in Japan, while modern human uses are likely the major factor in continental East Asia. These preliminary results show a complex demography and unusual genetic patterns in the contemporary populations for a common freshwater snail and are of significance to determine the historical formation and contemporary patterns of biogeography in Japan and continental East Asia.

The ancient geological patterns and climate change, such as the alternation of land and sea, the formation of lakes and rivers have important influence on the species diversification and biogeographic patterns. In East Asia, there were close paleogeographical relationships among the continent, the Korean peninsula, and the Japanese archipelago. The frequently changes of paleographical patterns since Pleistocene, reflected by the repeated shelf exposure in the regions connecting with continental East Asia and the Japanese archipelago, potentially played an important role in shaping the demographic history of this species in East Asia. To investigate the demographic history of S. quadrata influenced by the natural process of paleogeographical changes in the ancient times, the double digest restriction site associated DNA sequencing (ddRAD-seq) was used for higher precise analyses. Results showed that the population size changed associated with the sea level changes in the early Pleistocene, particularly, was highly consistent with the largest sea level changes in the Last Glacial Maximum (LGM). The non-continental populations increased during the early Pleistocene (~ 2.3−0.8 Ma) and shrank to the small sizes during the mid-Pleistocene (~ 0.8−0.13 Ma) probably because of the larger variations of climate and the possible paleogeographical changes that totally made the Japanese archipelago separated during the mid-Pleistocene. Then, during the LGM, populations encountered size outbreaks probably because they received migrants from continental East Asia through the paleo-river systems developed on the exposed ECS land bridge and through the narrow strait channel that had low-salinity surface water caused by injection of large continental paleo-rivers. The non-continental populations of S. quadrata were then replenished and increased rapidly in the warming up climate after the LGM. The divergence between lineages was also consistent with the period of sea level changes. The ancient river systems on the exposed East China Sea land bridge in the Last Glacial Maximum (LGM) would be corridors for the dispersal of S. quadrata. The migration route of S. quadrata was likely form the east of continent to the southwest of Japan (Kyushu) passing the southern part of Korean peninsula, and then arrived to the other parts of Japan. The alternation of sea and land and the paleographic changes had important influence on the evolutionary process of freshwater species in East Asia.

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