Abstract
The upside-down jellyfish Cassiopea is a globally distributed, semi-sessile, planktonically dispersed scyphomedusa. Cassiopea occurs in shallow, tropical inshore marine waters on sandy mudflats and is generally associated with mangrove-dominated habitats. Controversy over the taxonomy of upside-down jellyfishes precedes their introduction to the Hawaiian Islands during the Second World War, and persists today. Here we address the global phylogeography and molecular systematics of the three currently recognized species: Cassiopea andromeda, C. frondosa, and C. xamachana. Mitochondrial cytochrome c oxidase I (COI) sequences from Australia, Bermuda, Fiji, the Florida Keys, the Hawaiian Islands, Indonesia, Palau, Panama, Papua New Guinea, and the Red Sea were analyzed. Highly divergent COI haplotypes within the putative species C. andromeda (23.4% Kimura 2-parameter molecular divergence), and shared haplotypes among populations of two separate putative species, C. andromeda and C. xamachana from different ocean basins, suggest multiple anthropogenic introductions and systematic confusion. Two deeply divergent O’ahu haplotypes (20.3%) from morphologically similar, geographically separate invasive populations indicate long-term (14–40 million years ago) reproductive isolation of phylogenetically distinct source populations and cryptic species. Data support at least two independent introductions to the Hawaiian Islands, one from the Indo-Pacific, another from the western Atlantic/Red Sea. Molecular phylogenetic results support six species: (1) C. frondosa, western Atlantic (2) C. andromeda, Red Sea/western Atlantic/Hawaiian Islands (3) C. ornata, Indonesia/Palau/Fiji (4) Cassiopea sp. 1, eastern Australia (5) Cassiopea sp. 2, Papua New Guinea and (6) Cassiopea sp. 3, Papua New Guinea/Hawaiian Islands.



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Acknowledgements
We are grateful to Professor Mike Hadfield for providing support and research facilities at the Kewalo Marine Laboratory, University of Hawai’i, and to Professor John Benzie for facilities at the Centre for Marine and Coastal Studies, University of New South Wales (UNSW). We thank William Puleloa, Tom Iwai, Lori Colin, Bert Hoeksema, Harilaos Lessios, Don de Maria, Laura Martin, David Miller, Kylie Pitt, Alan Nelson, Kirk Murakami, and Dan Lindstrom for providing medusae specimens. Thanks to Kualoa Ranch and Mid-Pacific Golf Course for providing access. François Seneca assisted in laboratory DNA extractions. Thanks to Dr. Jonathan Gardner, Victoria University, New Zealand for valuable comments on an earlier draft of this manuscript. We also thank Aquarium Solutions International, Ruhr-Universität Bochum, and the Vice-Chancellor’s Post-doctoral Fellowship scheme (UNSW) for financial support. The experiments performed during the course of this study are in full compliance with the current laws of the United States.
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Communicated by P.W. Sammarco, Chauvin
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Holland, B.S., Dawson, M.N., Crow, G.L. et al. Global phylogeography of Cassiopea (Scyphozoa: Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands. Marine Biology 145, 1119–1128 (2004). https://doi.org/10.1007/s00227-004-1409-4
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DOI: https://doi.org/10.1007/s00227-004-1409-4