New insights into the phylogeny of Spyridia (Ceramiales, Rhodophyta) species with uncinate spines, focusing on the structure of determinate branches

Abstract

ABSTRACT Twenty-one samples belonging to species of Spyridia Harvey with uncinate (= hook-shaped) spines on determinate lateral branches were investigated for their morphology with a focus on the structure of determinate branches, and for comparative chloroplast-encoded rbcL and nuclear LSU rDNA sequence analysis to elucidate their taxonomy and phylogeny. Currently, four Spyridia species with uncinate spines are recognized worldwide: S. alternans, S. cupressina, S. horridula and S. hypnoides. Of them, S. hypnoides has been recognized as the most common species with uncinate spines. In this study, we show that S. aculeata needs to be resurrected from S. hypnoides, and recognized as a distinct species based on samples from Israel, Red Sea, near the type locality. Spyridia aculeata is characterized by the spiral arrangement of determinate branches, incompletely corticated determinate branches, naked basal segment of determinate branches, by the presence of only acropetal cortication in the nodes of determinate branches, and by uncinate spines at the terminal node and on the first and second nodes of determinate branches. In our molecular analyses based on rbcL and LSU rDNA, although four Spyridia species with uncinate spines are supported on phylogenetic trees, they are not a monophyletic group. The feature of uncinate spines in Spyridia is recognized as a polyphyletic character. Our phylogenetic analysis using rbcL and LSU rDNA sequences reveals high gene sequence divergence (6.8–7.0% for rbcL and 1.2–1.3% for LSU rDNA) between samples of S. aculeata and S. hypnoides. Therefore, the distribution of S. hypnoides may be restricted to the Indian Ocean, whereas S. aculeata is widely distributed in the Atlantic Ocean including the Mediterranean Sea and the Red Sea. Another Spyridia species with uncinate spines, S. alternans, is recognized as a synonym of S. horridula by our detailed morphological observations of its type specimen.