Description of Calliactis tigris sp. nov.: reconciling taxonomy and phylogeny in hermit-crab symbiotic anemones (Cnidari

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ORIGINAL ARTICLE

Description of Calliactis tigris sp. nov.: reconciling taxonomy and phylogeny in hermit-crab symbiotic anemones (Cnidaria: Actiniaria: Hormathiidae) Luciana C. Gusmão 1

&

E. Rodríguez 1 & Marymegan Daly 2

Received: 22 April 2019 / Accepted: 11 August 2019 # Gesellschaft für Biologische Systematik 2019

Abstract The symbiosis between sea anemones and hermit crabs is ubiquitous in the marine environment (except in the poles), occurring from shallow to deep waters; it involves one or more anemones living on a shell inhabited by a hermit crab. The anemone-crab partnership is a mutualism in which hermit crabs provide a hard substrate, increased access to oxygenated waters and food supply, in exchange for defense by the anemone. The vast majority of the sea anemone partners belong to three genera in family Hormathiidae: Adamsia, Calliactis, Paracalliactis. Given the remarkable nature of the symbiosis, hormathiid partners have been hypothesized to represent a monophyletic group. This has been rejected by Gusmão and Daly et al. (2010) and confirmed by our phylogenetic analysis using molecular markers (12S, 16S, 18S, 28S, COIII). We expand the results of Gusmão and Daly et al. (2010) by finding a monophyletic Paracalliactis, which was left untested in their analyses. Thus, characters of taxonomic significance associated to the symbiotic habit are interpreted as functional rather than phylogenetic. We reconcile taxonomy and the present evolutionary framework to avoid defining taxonomic groups based on characters prone to convergence. We formalize the synonymy of Adamsia and Calliactis and provide updated diagnoses for the valid genera Calliactis and Paracalliactis to bring more stability to the group. Under this new framework, we describe Calliactis tigris sp. nov. from Australia based on 21 specimens collected off the coast of New South Wales and Queensland and differentiate it from congeners and other hermit crab symbionts recorded in the Pacific Ocean. Keywords Symbiosis . Mutualism . Convergence . Adamsia . Paracalliactis . Australia

Introduction The symbiosis between sea anemones and hermit crabs is ubiquitous in the marine environment (except in the poles), occurring from shallow to deep waters; it involves one or more anemones living on a gastropod shell inhabited by a

Electronic supplementary material The online version of this article (https://doi.org/10.1007/s13127-019-00414-2) contains supplementary material, which is available to authorized users. * Luciana C. Gusmão [email protected] 1

Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA

2

Museum of Biological Diversity, Ohio State University, 1315 Kinnear Road, Columbus, OH 43215, USA

hermit crab (Gusmão and Daly 2010). The anemone-crab partnership is a mutualism (Brooks and Gwaltney 1993; Mainardi and Rossi 1969) in which hermit crabs provide a hard substrate that prevents the anemone’s burial and increase access to positive conditions, including oxygenated wat

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