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242<br />

14.3 Medusae (Cnidaria)<br />

W.M. Graham and K.M. Bayha<br />

14.3.1 Phyllorhiza punctata (Scyphozoa)<br />

The scyphomedusa Phyllorhiza punctata appeared suddenly and in spectacular<br />

numbers in the northern Gulf of Mexico during the summer of 2000 (Graham<br />

et al. 2003). Phyllorhiza punctata was first described from Pt. Jackson,<br />

Australia (von Lendenfeld 1884), and, presumably due to its conspicuous<br />

medusa stage, has a relatively well-documented history of invading tropical<br />

and subtropical environments around the globe over the past 200 years. The<br />

first recorded invasion of P. punctata was into the Swan-Canning estuary,<br />

Perth (Western Australia) between 1837–1838, presumably by early Australian<br />

shipping (Rippingale and Kelly 1995). P. punctata subsequently<br />

appeared in Pearl Harbor, Hawaii, USA (1941; Devaney and Eldridge 1977),<br />

Laguna Joyunda, Puerto Rico (1945 at the latest; Garcia-Sais and Durbin<br />

1993), San Diego, California, USA (1990 at the latest; Colin and Arneson 1995),<br />

Danajon Bank, Bohol Island, The Philippines (Heeger et al. 1992), and Bahia<br />

de Todos os Santos, Brazil (between 1991–1999; da Silveira and Cornelius<br />

2000). P. punctata also appeared in the eastern Mediterranean (in 1965, as referenced<br />

by Galil et al. 1990), but apparently did not persist there. This welldocumented<br />

history of invasion over an extended period may make P. punctata<br />

a particularly instructive model toward understanding invasions of<br />

jellyfish in particular, and of invasive marine species in general.<br />

The occurrence of the very large population estimated at 10 ¥ 10 6 medusae<br />

across the north-central Gulf of Mexico (Alabama, Mississippi, and Louisiana)<br />

in the summer of 2000 was unexpected, as the species had never been<br />

documented north of the Caribbean Sea. The timing of the occurrence was<br />

coincident with the incursion of tropical water from the Caribbean into the<br />

Gulf of Mexico. Using a modeling approach to assess transport mechanisms<br />

via currents measured during the 2000 bloom, Johnson et al. (2005) were able<br />

to suggest a plausible advection of medusae out of the Caribbean Sea by the<br />

northward-flowing Loop Current in the Gulf of Mexico. However, in the<br />

absence of direct evidence of transport of medusae from the Caribbean to the<br />

northern Gulf via this current (Johnson et al. 2005), we must also consider<br />

that cryptic populations have existed for some time in the Gulf of Mexico.<br />

This is supported by anecdotal reports by fishermen of this species occurring<br />

as small populations around coastal Louisiana for at least several years prior<br />

to 2000. P. punctata has been documented in the coastal waters of south-central<br />

Louisiana in each summer since 2000 (unpublished data).<br />

The potential ecological and economic impacts of P. punctata were judged<br />

as high, and it was feared that the ecology of the northern Gulf of Mexico<br />

would be altered permanently, along with the valuable fishing industry that<br />

depends on it (Graham et al. 2003). These fears were fueled by the costs of P.

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