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Jackson2013-Status and Trendsof Caribbean Coral Reefs

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Pacific lionfish Pterois volitans throughout the<br />

entire wider <strong>Caribbean</strong> over the past decade<br />

(De Leon et al. 2011; Hackerott et al. 2013) or<br />

the uncontrolled spread of the alga Caulerpa<br />

taxifolia in the northern Mediterranean<br />

(Meinesz et al. 1993, 2001). The effects<br />

of lionfish on <strong>Caribbean</strong> invertebrates <strong>and</strong><br />

fishes could be very considerable, especially<br />

in exacerbating the consequences of<br />

overfishing by depleting juvenile parrotfishes<br />

<strong>and</strong> surgeonfishes (Albins <strong>and</strong> Hixon 2011).<br />

However, it is too soon to tell whether native<br />

predators might eventually have an impact<br />

of lionfish, especially in marine reserves<br />

where predators could regain their former<br />

abundance (Mumby et al. 2013).<br />

Far too little attention has been paid,<br />

however, to the introduction of the myriad<br />

marine organisms we cannot see, including<br />

virtually all microorganisms <strong>and</strong> pathogens.<br />

The case of the unidentified pathogen that<br />

caused the mass mortality of Diadema<br />

antillarum in 1983-1984 is a case in point.<br />

Diadema mortality began next door to the<br />

<strong>Caribbean</strong> entrance to the Panama Canal,<br />

whence it spread like wildfire on ocean<br />

currents eastward to Trinidad <strong>and</strong> Tobago<br />

<strong>and</strong> northward throughout the western<br />

<strong>Caribbean</strong>, Greater Antilles, <strong>and</strong> Florida all<br />

the way to Bermuda, with mortality in the<br />

eastern <strong>Caribbean</strong> arriving from both the<br />

north <strong>and</strong> the south in 1984 (Lessios et al.<br />

1984; Lessios 1988). Introduction via ballast<br />

water from the Pacific is seemingly the only<br />

explanation.<br />

This begs the question of why so many<br />

<strong>Caribbean</strong> coral diseases first appeared in<br />

the 1970s <strong>and</strong> early 1980s for which there<br />

is no compelling environmental explanation.<br />

These first coral diseases include black b<strong>and</strong><br />

disease that first appeared in 1973 <strong>and</strong><br />

white b<strong>and</strong> disease that first appeared in<br />

1977 (possibly earlier) that is by far the most<br />

devastating of the coral diseases through<br />

its virtual elimination of formerly dominant<br />

Acropora palmata <strong>and</strong> A. cervicornis<br />

throughout most of the <strong>Caribbean</strong> by<br />

the early 1980s (Weil <strong>and</strong> Rogers 2011).<br />

Temperatures were not excessively warm in<br />

the 1970s <strong>and</strong> heating in relation to El Nino<br />

in 1983 was small compared to the episodes<br />

in 1995, 1998, 2005, <strong>and</strong> 2010. There is<br />

also no evidence of a pervasive decline in<br />

<strong>Caribbean</strong> water quality before the 1980s or<br />

later.<br />

In contrast, the volume of international<br />

shipping exploded in the late 1960s with<br />

the advent of bulk carriers <strong>and</strong> enormous<br />

cruise ships that discharged untold volumes<br />

of ballast water into coastal waters before<br />

stricter regulations may have begun to take<br />

effect (Carlton 1996). Greater speed of<br />

transport among distant ports may also be a<br />

contributing factor. Many introduced species<br />

have been transported by ballast water, <strong>and</strong><br />

this is especially true for microbes that have<br />

been calculated to be transported in numbers<br />

on the order of 10 20 /year into the lower<br />

Chesapeake Bay alone (Ruiz et al. 2000;<br />

Drake et al. 2007). None of this proves that<br />

Diadema disease or WBD were introduced<br />

into the <strong>Caribbean</strong> from another ocean. But<br />

given the numbers, it is remarkable that all<br />

marine diseases have not been introduced<br />

throughout the global ocean.<br />

Once introduced, different environmental<br />

factors may retard or promote the growth<br />

of introduced species including species that<br />

cause disease. But it is important not to<br />

confuse the causes of an initial outbreak from<br />

factors that may subsequently promote or<br />

inhibit its spread <strong>and</strong> increase.<br />

It is therefore of considerable interest that the<br />

<strong>Caribbean</strong> is by far the most geographically<br />

isolated of all the tropical oceans with<br />

extensive coral reefs. The <strong>Caribbean</strong> has<br />

been separated from the vast Pacific to the<br />

west by the Isthmus of Panama for about<br />

3½ million years (Coates <strong>and</strong> Stallard 2013;<br />

Jackson <strong>and</strong> O’Dea 2013) <strong>and</strong> from the<br />

tropical Indian Ocean to the east by the<br />

continents of Africa <strong>and</strong> Asia, the subtropical<br />

Mediterranean, <strong>and</strong> the inhospitable eastern<br />

Atlantic. It is therefore of considerable interest<br />

that the effects of diseases on <strong>Caribbean</strong><br />

reef associated species have been so much<br />

more extreme than in other tropical seas.<br />

Nothing like the mass mortality of Diadema<br />

has affected any echinoderm throughout the<br />

entire Indian Ocean or Pacific, nor has any<br />

genus of Indo-Pacific coral suffered such<br />

broad extirpation as <strong>Caribbean</strong> Acropora.<br />

All of this suggests that isolation has been a<br />

64 STATUS AND TRENDS OF CARIBBEAN CORAL REEFS: 1969-2012

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