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Abstract
Marine symbiotic Palaemonidae, comprising over 600 species, live in association with marine invertebrates of different phyla, like Cnidaria, Echinodermata, Mollusca, Porifera, and Tunicata. A phylogenetic study is performed on a clade of bivalve- and ascidian-associated endosymbiotic shrimp species (Caridea: Palaemonidae), using morphological and molecular data. A Total Evidence approach is used in order to include all currently known ingroup species in an evolutionary framework. Ancestral state reconstruction analyses are performed to identify host-switching events and ancestral ranges. The clade, including Ascidonia, Conchodytes, Dactylonia, Odontonia, and Pontonia, and various smaller genera, is recovered as monophyletic, with an ascidian-associated ancestral host state. At least six interphylum host switches are tentatively identified, with members of Odontonia and Notopontonia switching back to an ascidian host affiliation after the ancestral host switch of the clade including Conchodytes, Odontonia and related genera, from an ascidian- to a bivalve host. The clade including Ascidonia and Pontonia was recovered to have an ancestor with an East Pacific/Atlantic distribution. The other studied genera remained in the original ancestral Indo-West Pacific range. We hypothesize that similar internal environments of shrimp hosts from different phyla will function as hot spots for interphylum host switching in various lineages of symbionts.
Abstract
Most marine shrimp species of the family Palaemonidae are characterized by symbiotic associations with hosts that belong to a wide range of invertebrate phyla. One clade of related endosymbiotic species has evolved to live inside the branchial chambers of ascidians and the mantle cavities of bivalve molluscs. The phylogeny of this clade (comprising 15 bivalve-associated species in the genera Anchistus, Neoanchistus, and Paranchistus, and three species of ascidian-associated species in the genus Dasella) is the topic of the present study, which is based on both morphological and molecular data. A concatenated phylogeny reconstruction was built by using the markers coi and 16S. With the help of a total evidence approach (with a scored morphological datamatrix), species could be added for which no molecular data were available. An ancestral character state analysis was performed to detect host switches. In contrast to another endosymbiotic clade, the ancestral host state was found to be slightly in favour of bivalves. The phylogenetic relevance of morphological features is discussed with a focus on a tympanal organ located in the major chelipeds of some bivalve-associated shrimp species. This little-known anatomical structure is illustrated by sem and µCT scans. Its possible function is discussed. In the phylogeny reconstructions, both Anchistus and Paranchistus were found to be polyphyletic. They were reclassified to obtain a more natural classification: Anchistus custoides and Anchistus custos were transferred to the resurrected genus Ensiger. Paranchistus liui, P. nobilii, P. pycnodontae, and P. spondylis were transferred to Polkamenes gen. nov. Anchistus pectinis and P. ornatus were transferred to Tympanicheles gen. nov. Per genus, a key to the species is provided.