Molecular systematics of peppermint and cleaner shrimps: phylogeny and taxonomy of the genera Lysmata and Exhippolysmata (Crustacea: Caridea: Hippolytidae)

Abstract

Shrimps from the ecologically diverse genera Lysmata and Exhippolysmata are rare among marine invertebrates because they are protandric simultaneous hermaphrodites: shrimps initially mature and reproduce solely as males, and later in life become functional simultaneous hermaphrodites. Considerable progress on the reproductive ecology of members from these two genera has been achieved during the last decade. However, several outstanding issues of systematic nature remain to be addressed. Here, a molecular phylogeny of these two genera was used to examine the overall evolutionary relationship within and between species and genera, and to answer various questions related to the systematic status of several species. The present phylogenetic analysis, including 53 sequences and 26 species of Lysmata and Exhippolysmata, indicates that semiterrestrial shrimps from the genus Merguia represent the sister group to a second natural clade composed by shrimps from the genera Lysmata and Exhippolysmata. Also, the phylogenetic analysis confirmed that the genus Lysmata is paraphyletic, and includes the genus Exhippolysmata, as noted in a preliminary study. The tree partially supports the separation of species with or without a developed accessory branch into two different genera or subgenera (i.e. Lysmata and Hippolysmata having a well-developed accessory branch, or not, respectively). The genetic distance between the cleaner shrimps Lysmata amboinensis and Lysmata grabhami was smaller than has been observed between other sister species. On the other hand, the topology of the tree indicates that these two entities are reciprocally monophyletic. Thus, this latter result, together with minor but constant differences in the colour pattern reported for these two entities, indicates that there is no reason to stop treating them as different valid species. This study enabled the long overdue resolution of standing taxonomic questions in shrimps from the genera Lysmata and Exhippolysmata. In the future, this phylogeny will help to reveal the conditions favouring the origins of several behavioural and morphological novelties in these unique shrimps. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 254–265.