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The sands of time: rediscovery of the genus Neozoanthus (Cnidaria: Hexacorallia) and evolutionary aspects of sand incrustation in brachycnemic zoanthids


The zoanthid family Neozoanthidae (Anthozoa: Hexacorallia: Zoantharia) was described in 1973 from Madagascar as a monogeneric and monotypic taxon, and never reported again in literature. In 2008–2010, numerous zoanthid specimens fitting the morphological description of Neozoanthus were collected in the Ryukyu Islands, Okinawa, Japan, and the Great Barrier Reef (GBR), Australia. Utilizing these specimens, this study re-examines the phylogenetic position of Neozoanthidae and analyzes the evolutionary history of sand incrustation in zoanthids through phylogenetic and ancestral state reconstruction analyses. Specimens were colonial, partially incrusted with large, irregular sand and debris, zooxanthellate, and found from the intertidal zone to depths of approximately 30 m. Phylogenetic results utilizing mitochondrial 16S ribosomal DNA and cytochrome oxidase subunit I sequences show the presence of two Neozoanthus species groups, one each from Japan and the GBR. Unexpectedly, the molecular results also show Neozoanthus to be very closely related to the genus Isaurus, which as a member of the family Zoanthidae, is not sand incrusted. These results suggest that during evolution zoanthids can acquire and lose the ability to incrust sand with relative rapidity.

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The Census of Coral Reef Ecosystems (CReefs) Australia Project is sponsored by BHP Billiton in partnership with the Great Barrier Reef Foundation and the Australian Institute of Marine Science (AIMS). CReefs is a field program of the Census of Marine Life. In Japan, all MISE laboratory members (2008–2010) are thanked for field help, as are Dr. Ryuji Asami (Rising Star Program) and Ryuji Suzuki (Marine Service Kamui, Tokunoshima). CReefs Australia project leader Dr. Julian Caley, field logistics manager Shawn Smith (both AIMS) and all Heron Island 2009 CReefs survey participants are thanked for their help in specimen collection. JDR was partially supported by the Rising Star Program at the University of the Ryukyus, and a grant from the Japan Society for the Promotion of Science (“Wakate B” # 2170088). Specimens from Australia were collected under Great Barrier Reef Marine Park Authority permit # G32313.1 and Queensland Fisheries permit # 95152. Two anonymous reviewers’ comments greatly improved the manuscript.

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Correspondence to James Davis Reimer.

Additional information

Communicated by C. Riginos.

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Fig. S1

Scanning electron microscope image of a cross-section of specimen MISE 539 showing incrusted sponge spicules (ss) with relatively intact mesoglea (m) and remains of locations of other embedded materials (rough areas surrounding mesoglea). Scale = 200 μm. (TIFF 758 kb)

Fig. S2

Maximum likelihood (ML) tree of cytochrome oxidase subunit 1 (COI) sequences for zoanthid specimens. Values at branches represent ML probabilities (>50%) and Bayesian posterior probabilites (>0.50), respectively. Sequences newly obtained in this study in bold. Sequences/species names from previous studies in regular font with GenBank Accession Number. For specimen information see Table S1. (EPS 365 kb)

Supplementary material 3 (DOC 58 kb)

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Reimer, J.D., Hirose, M., Irei, Y. et al. The sands of time: rediscovery of the genus Neozoanthus (Cnidaria: Hexacorallia) and evolutionary aspects of sand incrustation in brachycnemic zoanthids. Mar Biol 158, 983–993 (2011).

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  • Coral Reef
  • Markov Chain Monte Carlo
  • Great Barrier Reef
  • Reef Flat
  • Sphincter Muscle