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EPA's Vessel General Permit and Small Vessel General

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planktonic larvae form before settling <strong>and</strong> metamorphosizing on appropriate substrates,<br />

preferably coralline algae (Bak 1977, Sammarco 1980, Rylaarsdam 1983). Initial calcification<br />

ensues <strong>and</strong> develop into daughter corallites.<br />

Studies indicate that larger colonies (as measured by surface area of the live colony) have higher<br />

fertility <strong>and</strong> fecundity rates; colonies with a branch length longer than 3.5 inches were fertile <strong>and</strong><br />

over 80% of colonies with branches longer than 6.7 inches were fertile (Soong <strong>and</strong> Lang 1992).<br />

Estimated size at sexual maturity is 6.7 inches in branch length <strong>and</strong> the smallest known<br />

reproductive colony was 3.5 inches in branch length (Soong <strong>and</strong> Lang 1992).<br />

Biological <strong>and</strong> physical factors affect spatial <strong>and</strong> temporal patterns of recruitment. These include<br />

substrate availability <strong>and</strong> community structure, grazing pressure, fecundity, mode <strong>and</strong> timing of<br />

reproduction, behavior of larvae, hurricane disturbance, physical oceanography, the structure of<br />

established coral assemblages, <strong>and</strong> chemical cues (Lewis 1974, Birkel<strong>and</strong> 1977, Goreau et al.<br />

1981, Rogers et al. 1984, Baggett <strong>and</strong> Bright 1985, Harriott 1985, Hughes <strong>and</strong> Jackson 1985,<br />

Sammarco 1985, Morse et al. 1988, Fisk <strong>and</strong> Harriott 1990, Richmond <strong>and</strong> Hunter 1990).<br />

Growth rates are relatively rapid, expressed as the linear extension of branches, ranging from 1.2<br />

to 4.3 inches annually, <strong>and</strong> have enabled staghorn coral to construct significant reefs in several<br />

locations throughout the Caribbean (Vaughan 1915, Jaap 1974, Adey 1978). Branching species,<br />

such as acroporid corals, grow differentially in response to light such that coral polyp growth<br />

maximizes exposure to available light (Kaniewska et al. 2009). During the 1970s there were vast<br />

fields, or thickets, of staghorn coral on many reefs. The nominal situation in 2004 was isolated<br />

branches <strong>and</strong> small thickets, 1.6 to 3.3 feet across. Growth can also occur from fragmentation<br />

<strong>and</strong> dispersal (Tunnicliffe 1981, Bak <strong>and</strong> Criens 1982). A broken branch may be carried by<br />

waves <strong>and</strong> currents to another location <strong>and</strong>, if favorable, branches grow into a new colony.<br />

Rapid growth <strong>and</strong> fragment dispersal facilitate a competitive advantage for staghorn coral<br />

relative to other coral <strong>and</strong> benthic species (Shinn 1976, Neigel <strong>and</strong> Avise 1983, Jaap et al. 1989).<br />

Larval recruitment is influenced by the type <strong>and</strong> availability of benthic substrate, with certain<br />

types of coral or rock substrates resulting in greater or lesser recruitment success (Ritson-<br />

Williams et al. 2009).<br />

Habitat<br />

Historically, staghorn coral so dominated reef systems within the 23 to 49 feet depth that the area<br />

became known as the staghorn zone. In other reef systems (Jamaica, Cayman Isl<strong>and</strong>s, Belize,<br />

<strong>and</strong> eastern Yucatan), staghorn coral was a major mid-depth (33 to 82 feet) reef-builder (Adey<br />

1977, 1978). Historically, staghorn coral was reported from depths ranging from surface to 200<br />

feet, although it is considered rare below 66 feet (Goreau <strong>and</strong> Goreau 1973). In southeastern<br />

Florida, this species historically occurred on the outer reef (52 to 66 feet), on spur, groove bank,<br />

<strong>and</strong> transitional reefs, <strong>and</strong> on octocoral-dominated hard-bottom (Goldberg 1973, Davis 1982,<br />

Jaap 1984, Wheaton <strong>and</strong> Jaap 1988). Colonies were common in back- <strong>and</strong> patch-reef habitats<br />

(Gilmore <strong>and</strong> Hall 1976, Cairns 1982). Although staghorn coral colonies are sometimes found<br />

interspersed among colonies of elkhorn coral, they are generally in deeper water or seaward of<br />

the elkhorn zone <strong>and</strong> more protected from wave action.<br />

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