Quaternary Vicariance of Lotic Coeliccia in the Ryukyu-Taiwan Islands Contrasted with Lentic Copera.

Abstract

Lotic dragonflies and damselflies are expected to be more affected by vicariance than lentic sister species. We demonstrated that severe vicariant speciation acted on lotic Coeliccia in contrast to lentic Copera damselflies, which are both included in the family Platycnemididae. We constructed maximum likelihood and Bayesian inference trees of these Platycnemididae species from the continental islands of Ryukyu (Amami, Okinawa, and Yaeyama islands), Taiwan, and Japan relative to Chinese species using raxmlGUI and BEAST, based on the mitochondrial COI gene (682 bp), COII gene (494 bp), 16SrRNA (478 bp), and the nuclear 28SrRNA gene (807 bp). In BEAUti, we calibrated the splitting age of the MRCA of all the Coeliccia species as 1.55-0.15 million years ago (Ma), a date that corresponds to a geologic constraint: the Okinawa trough and associated straits, including the Yilan basin in Taiwan, began to rift at 1.55 Ma, isolating the Ryukyu-Taiwan islands from the Chinese continent. The vicariance split Coeliccia into the Ryukyu-side clade of Coeliccia ryukyuensis (Coe. r. ryukyuensis in Okinawa and Coe. r. amamii in Amami) and Coeliccia flavicauda (Coe. f. masakii in Yaeyama and Coe. f. flavicauda in southern Taiwan), and the Chinese-side clade of Coeliccia cyanomelas (northern Taiwan and China), separated by the Okinawa trough. These Coeliccia species were further deeply differentiated to form local populations on the major islands and some of the minor islands. The Copera clade constituted a sister of the lotic Coeliccia clade, but genetic differentiation was not recognizable in lentic Copera between China, Taiwan, and Japan. Base substitution rates applying a strict clock model were estimated for COI: 0.0783, COII: 0.0803, 18SrRNA: 0.0186, 28SrRNA: 0.00577, and combined: 0.0408 substitutions/site/myr, and these rates are relatively high.