QL

155

.S63

no. 82-

11.73

t

Biological Report 82 (11-W

August 1987

Library

fialional Wefkmfs Research Center

u. S. F!sh and Wiidlife Service

?BFte_7rm~O.~C Zkmlevard

Lafayette. La. 70506 TR EL-82-4

Species Profiles: Life Histories and

Environmental Requirements of Coastal Fishes

and Invertebrates (South Florida)

REEF-BUILDING CORALS

Fish and Wildlife Service

U.S. Department of the Interior

Coastal Ecology Group

Waterways Experiment Station

U.S. Army Corps of Engineers
Bio. Rept. 82(11.73) Species Profile: (South Florida) Reef-Building Corals

Erratum

P2. left column, paragraph 4, sentence 3 should read:

Both species of Montastraea, but no species of Acropora, occur on

the Texas Flower Garden Banks (Bright and Pequegnat 1974; Bright et al.

1984), and in Bermuda (La'borel, 1966; Sterrer 1986). Neither Acropora

nor Montastraea occurs on the Florida Middle Grounds (Hopkins et al. 1977).

Hopkins, T..S., D. R. Blizzard, S. A. Brawley, S. A. Earle, D. E. Grimm,

D. K. Gilbert, P. G. Johnson, E. H. Livingston, C. H. Lutz, J. K. Shaw,

and B. B. Shaw. 1977. A preliminary characterization of the biotic

components of composite strip transects on the Florida Middlegrounds,

Northeastern Gulf of Mexico. Proc. Third International Coral Reef

Symp. (Miami, FL) 1:31-37.
Biological Report 82(11.73)

TR EL-82-4

August 1987

Species Profiles: Life Histories and Environmental

Requirements of Coastal Fishes and Invertebrates (South Florida)

REEF-BUILDING CORALS

James W. Porter

Zoology Department

University of Georgia

Athens, GA 30602

Project Manager

Carroll Cordes

Project Officer

David P. Moran

U.S. Fish and Wildlife Service

National Wetlands Research Center

1010 Gause Boulevard

Slidell, LA 70458

Performed for

Coastal Ecology Group

U.S. Army Corps of Engineers

Waterways Experiment Station

Vicksburg, MS 39180

and

U.S. Department of the Interior

Fish and Wildlife Service

Research and Development

National Wetlands Research Center

Washington, DC 20240
This series should be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories

and environmental requirements of coast3 fishes and invertebrates. U.S. Fish

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile can be cited as follows:

Porter, J.W. 1987. Species profiles: life histories and environmental

requirements of coastal fishes and invertebrates (south Florida)--reef-building

corals. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.73). U.S. Army Corps of

Engineers, TR EL-82-4. 23 pp. *
w

PREFACE

This species profile is one of a series on coastal aquatic organisms,

principally fish, of sport, commercial, or ecological importance. The profiles

are designed to provide coastal managers, engineers, and biologists with a brief

comprehensive sketch of the biological characteristics and environmental

requirements of the species and to describe how populations of the species may be

expected to react to environmental changes caused by coastal development. Each

profile has sections on taxonomy, life history, ecological role, environmental

requirements, and economic importance, if applicable. A three-ring binder is

used for this series so that new profiles can be added as they are prepared.

This project is jointly planned and financed by the U.S. Army Corps of Engineers

and the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one of

the following addresses.

Information Transfer Specialist

National Coastal Ecosystems Team

U.S. Fish and Wildlife Service

NASA-Slide11 Computer Complex

1010 Gause Boulevard

Slidell, LA 70458

or

U.S. Army Engineer Waterways Experiment Station

Attention: WESER-C

Post Office Box 631

Vicksburg, MS 39180

W

1

iii
CONVERSION TABLE

Metric to U.S. Customary

Multiply FY

millimeters (mm) 0.03937

centimeters (cm) 0.3937

meters (m) 3.281

meters (m) 0.5468

kilometers (km) 0.6214

kilometers (km) 0.5396

square meters (mz) 10.76

square kilometers (km2) 0.3861

hectares (ha) 2.471

liters (1) 0.2642

cubic meters (m3) 35.31

cubic meters (ms) 0.0008110

milligrams (mg) 0.00003527

grams (9) 0.03527

kilograms (kg) 2.205

metric tons (t) 2,205.O

metric tons (t) 1.102

kilocalories (kcal) 3.968

Celsius degrees ("C) 1.8('C) + 32

micro Einsteins per square meter per second (pE m-'s-l) 6.02 x 10"

micro Einsteins per square meter per second (LIE m-%-l) 4.67

micro Einsteins oer square meter per second (uE m-'s-l) 51.22

U.S. Customary to Metric

Multiply

inches

inches

feet (ft)

fathoms

statute miles (mi)

nautical miles (nmi)

EY

25.40

2.54

0.3048

1.829

1.609

1.852

To Obtain

millimeters

centimeters

meters

meters

kilometers

kilometers

square feet (ft') 0.0929 square meters

square miles (mi2) 2.590 square kilometers

acres 0.4047 hectares

gallons (gal)

cubic feet (ft')

acre-feet

3.785

0.02831

1,233X

28,350.

28.35

0.4536

0.00045

0.9072

liters

cubic meters

cubic meters

ounces (oz)

ounces (oz)

pounds (lb)

pounds (lb)

short tons (ton)

milligrams

grams

kilograms

metric tons

metric tons

British thermal units (Btu) 0.2520

Fahrenheit degrees (OF) 0.5556 (OF - 32)

photons m-2s-' 1.661 x lo-'"

foot candles (fc) 0.21413

lux (lx) 0.01952

To Obtain

inches

inches

feet

fathoms

statute miles

nautical miles

square feet

square miles

acres

gallons

cubic feet

acre-feet

ounces

ounces

pounds

pounds

short tons

British thermal units

Fahrenheit degrees

photons m-2s-1

foot candles

lux

kilocalories

Celsius degrees

micro Einsteins per square meter per second

micro tinsteins per square meter per second

micro Einsteins per square meter per second

iv
CONTENTS

Page

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......... vi

PREFACE ................................................. . . .

CONVERSION TABLE ........................................ . . .

ACKNOWLEDGMENTS ......................................... . . .

NOMENCLATURE, TAXONOMY, AND GEOGRAPHIC RANGE ........

MORPHOLOGY AND IDENTIFICATION AIDS ..................

REASON FOR INCLUSION IN SERIES ......................

LIFE HISTORY ........................................

Spawning ..........................................

Fecundity .........................................

Larvae and Juveniles ..............................

GROWTH CHARACTERISTICS ..............................

POPULATION CHARACTERISTICS ..........................

Depth Distribution ................................

Response to Hurricanes ............................

FISHERY .............................................

ECOLOGICAL-ROLE .....................................

Photosynthetic Capability .........................

Interspecific Competition .........................

Predators .........................................

Sediment Rejection Behavior .......................

Disease ...........................................

ENVIRONMENTAL REQUIREMENTS ..........................

Temperature .......................................

Salinity ..........................................

Dissolved Oxygen ..................................

Substrate .........................................

Solar Irradiance ..................................

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LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

V

ACKNOWLEDGMENTS

I thank Alex Heller for drawing Figures la - d, and Lawrence Castanares for

discussions on the systematics of these coral species. Reviews were provided by

Dr. Judith C. Lang, University of Texas, Austin, and Dr. Alina Szmant-Froelich,

University of Miami, Florida. This report is Contribution No. 303 from the Discovery Bay Marine Laboratory, Discovery Bay, Jamaica.

vii
4 Elkhorn coral w

Acropora palmata

B Staghorn coral

Acropora cervicornis

Figure 1. Major reef-building corals in the Caribbean.

m depth; San Bias, Atlantic Panama.

(A) Specimen UGA 512; 2

History, University of Georgia.

Cnidaria Collection, Museum of Natural

Curacao.

(B) Specimen UGA 292; 4 m depth; Jan-Thiel

Cnidaria Collection, Museum of Natural History, University of Georgia:

viii
C Common star coral

Montastraea annularis

D Large star coral

Montastraea cavernosa

(C) Specimen UGA 379; 2 m depth; Discovery Bay, Jamaica. Cnidaria Collection,

Museum of Natural History, University of Georgia. (D) Specimen UGA 1154; 15 m

depth; Discovery Bay, Jamaica. Cnidaria Collection, Museum of Natural History,

University of Georgia. Drawings by Alex Heller.

REEF-BUILDING CORALS

Information is presented on four Preferred common name...Elkhorn coral

major Caribbean reef-building corals, (Figure la)

mentioned individually where relevant Class . . . . . . . . . . . . . . . . . . . . . . ..Anthozoa

data exist. The species are compared Order . . . . . . . . . . . . . . . . . . ..Scleractinia

and contrasted to highlight their Suborder . . . . . . . . . . . . . . . ..Astrocoeniia

similarities and differences. Family . . . . . . . . . . . . . . . . . . ..Acroporidae

NOMENCLATURE, TAXONOMY, AND GEOGRAPHIC

RANGE

1. Specific name.. .Acropora palmata Class . . . . . . . . . . . . . . . . . . . . . . ..Anthozoa

(Lamark) Order . . . . . . . . . . . . . . . . . . ..Scleractinia

2. Specific name...Acropora cervicornis (Lamark)

Preferred common name...Staghorn coral

(Figure lb)

1
Suborder ............... ..Astrocoenii a

Family .................. ..Acroporida e

3. Specific name Montastraea annularis (Ellis and Solander)

Preferred common name . . . Common star

coral (Figure lc)

Class . . . . . . . . . . . . . . . . . . . . . . ..Anthozoa

Order . . . . . . . . . . . . . . . . . . ..Scleractinia

Suborder . . . . . . . . . . . . . . . . . . . . ..Faviina

Family . . . . . . . . . . . . . . . . . . . . . ..Faviidae

4. Specific name..Montastraea cavernosa (Linnaeus)

Preferred common name . . . . ..Large star

coral (Figure Id)

Class . . . . . . . . . . . . . . . . . . . . . . ..Anthozoa

Order . . . . . . . . . . . . . . . . . . ..Scleractinia

Suborder . . . . . . . . . . . . . . . . . . . . ..Faviina

Family . . . . . . . . . . . . . . . . . . . . . ..Faviidae

All four of these corals are found

throughout shallow coastal waters of

the Caribbean and the western tropical

Atlantic (Smith 1948). Montastraea

cavernosa is also found in Brazil

(Laborel 1967) and the eastern

tropical Atlantic (Laborel 1974). All

four species are common in southern

Florida (L. Agassiz 1852, 1869, 1880;

A. Agassiz 1888; Vaughan 1911; Shinn

1963; Ginsburg and Shinn 1964;

Kissling 1965; Marszalek et al. 1977;

Davis 1982; Halas 1985). Both species

of Montastraea, but no species of

Acropora, occur in the Florida Middle

Grounds, the Texas Flower Garden Banks

(Bright and Pequegnat 1974; Bright et

al. 1984), and in Bermuda (Laborel

1966; Sterrer 1986).

Goldbera (1973) reported that

isolated colonies -of Acropora cervicornis, Montastraea annularis, and M.

cavernosa occur in deep water

(16-30 m) as far north as Palm Beach,

Florida. at latitude 26O3' N (Fiaure

2). Acropora palmata appears firsi at

Fowey Rocks, Florida (lat. 25O37' N),

which is also the location of the

first major appearance of all of the

species in shallow water (Burns 1985).

Reef development by these species

begins slightly farther south, in

Biscayne National Park, Florida, at

25'25' N latitude. Fully developed

Acropora palmata and A. cervicornis

reef zones are found in Key Largo

National Marine Sanctuary at Carysfort

Reef, Florida (lat. 25O20' N), and

south of there (Jaap 1984; White and

Porter 1985). Little is known about

coral distribution in the turbid

waters off the west coast of Florida

(Figure 2).

Contrasting depth ranges and zones

of maximum development for these

species in Jamaica (Goreau and Wells

1967) are shown in Table 1.

MORPHOLOGY AND IDENTIFICATION AIDS

Both species of Acropora are

branching (Figures la and b), whereas

both species of Montastraea are mound

shaped (Figures lc and d). The blades

are flattened and palm shaped in A.

palmata in_rough water), bit

narrow

(raonyder

cylindrical in .

cervicornis. Branches emerge at acute

angles and generally in the same plane

of growth as the parent blade in 4.

palmata, whereas each blade emerges

more at right angles, and generally in

a different plane of growth from most

of the other blades, in 4.

cervicornis. Blades of 4. palmata are

greater than 0.5 m in length, whereas

unbifurcated blades of A. cervicornis

rarely are. Coloration is highly

variable in 4. palmata, ranging from

very light tan to brown, whereas it is

brownish yellow and more uniform in

colonies of A. cervicornis. Polyps

(the individual units of live coral

that make up a colony) of both A.

palmata and A. cervicornis are about

0.1 cm in di>meter. Acropora cervicornis has a single apical pollin

the growth axis.

Polyps of Montastraea annularis

average 0.3 cm in diameter (mediumsized among coral polyps); those of M.

cavernosa average 1.0 cm (large for

Caribbean corals). In both species,

polyp diameter increases and polyp

density decreases with increasing

2
A TLANTIC

CANAVERAL OCEAN

GULF O F

MEXICO

Isolated colonies of

Montastraea spp.

Isolated colonies of Acropora

cervicornis and Montastraea spp.

FLORIDA

MILES

0 50 100

Lateral distribution of -e___ ,

Acropora and Montastraea spp. 0 50 100

KILOMETERS

lgure L. Loastal distribution of the elkhorn, staghorn, common star, and large

star corals.

contour;

Montastraea exists in water depths below 1 m and out to the 40-m

Acropora exists on reefs from the surface to 20 m depth.

3
Table 1. Patterns of depth distribution, abundance, and ecology of Montastraea

annularis, M. cavernosa, Acropora palmata, and - A. cervicornis.

Species

Extensionb

Depth (m)a

Resisrate SpawningC SpatC tance to

Range optimum (cm/yr> SexC season (No./m*) hurricane

Acropora palmata O-15 o-7 5.00-9.50 H Aug 0.03 M

Acropora cervicornis O-30 3-20 10.00-26.40 H Aug 0.01-0.30 L

Montastraea annularis O-80 3-40 0.68-2.40 H Aug-Sept 0.45-1.04 H

Montastraea cavernosa O-90 lo-40 0.35-0.57 D Sept 0.14-1.09 H

ZData adapted from Goreau and Wells (1967).

See GROWTH CHARACTERISTICS section of the text for references.

ZData adapted from Szmant-Froelich (1985,

L = low, M = medium, and H = high.

1986).

depth (Dustan 1975, 1979; Porter et

al. 1987). Although colonies of both

species are mound shaped, those of M.

annularis that exceed 0.5 m in diameter tend to break into distinct lobes

in shallow water; no such tendency

exists in large colonies of M.

cavernosa. Colonies of M. annularTs

have a great diversity of shapes in

shallow water. Colonies of M.

annularis sometimes become 4-5 m Tn

diameter, but those of M. cavernosa

rarely exceed 1 m. As water depth

increases, colonies of both

Montastraea species flatten and become

platelike (Dustan 1975, 1979; Graus

and Macintyre 1976). At depths of

30-40 m in Jamaica, most colonies of

M. annularis

rosettes of

grow as flattened

overlappinq colonies.

Colonies of M. cavern&a do not

flatten completely at depths less than

60 m.

REASON FOR INCLUSION IN THE SERIES

Coral reefs, which are highly

attractive to tourists, are the most

diverse marine community on earth.

Their existence depends on the successful growth and maintenance of a

complex, three-dimensional structure &

in shallow water. This topographically complex form is built primarily by

hermatypic corals (those corals bearing symbiotic zooxanthellae). The

four species reviewed here are among

the primary corals involved in this

building process (Lighty et al. 1982).

As stated by Jaap (1984), in

southern Florida "the two stony corals

most responsible for reef building are

Acropora palmata (elkhorn coral) and

Montastraea annularis (star coral)."

LIFE HISTORY

Spawning

The four species reviewed here spawn

annually in the fall. They all

discharge their gametes into the water

column, rather than brooding their

larvae (Szmant-Froelich 1986).

Montastraea annularis is a simultaneous hermaphroditeceggs and sperm

4
are present in the same individual at

the same time). In Puerto Rico,

oogenesis begins in mid-May, and

spermatogenesis begins during mid-July

(Szmant-Froelich 1985). The annual

spawning period is short, occurring

immediately after the full moon in

late August or September. Colonies

larger than 51 cm2 (estimated age, 4-5

years) are reproductive; smaller

colonies, or pieces of colonies below

this size, are not (Szmant-Froelich

1985).

Montastraea cavernosa is dioecious.

The gametogenic period is similar to

that described above for M. annularis.

Spawning also occurs annually in late

August or September (Szmant-Froelich

1986). Neither minimum colony size

nor age at first reproduction is

known.

Acropora palmata and A. cervicornis

do not differ substantially in

reproductive biology. Like Montastraea annularis, they are simultaneous hermaphrodites in which the

development period is longer for eggs

than for sperm. They differ from y.

annularis in having a longer period of

egg development of 10 months. As in

both species of Montastraea, there is

a short spawning season in August. In

Puerto Rican populations of Acropora,

spawning is synchronous and occurs 6

days after the full moon in August

(Szmant-Froelich 1986). The relation

between fragmentation and reproduction

has not been examined.

Fecundity

Annual egg production per square

centimeter of coral tissue is 720 to

2,016 in Montastraea annularis and 288

to 576 in M. cavernosa (SzmantFroelich 1986). Because eggs of M.

annularis are slightly smaller than

those of y. cavernosa (300 and 350 pm,

respectively), the annual egg mass

produced per unit area of living coral

tissue by M. cavernosa is double that

produced by E. annularis.

Annual egg production by both

Acropora palmata and A. cervicornis is

600 to 800 eggs per cm2. The eggs are

similar in size to those of

Montastraea annularis in being about

300 pm in diameter.

Larvae and Juveniles

Little is known about the planula

larvae of the four coral species

treated here (Bak et al. 1977;

Sammarco 1980; Rylaarsdam 1983).

Settlement patterns are also poorly

understood for any of the species

reviewed here, but have been studied

on reefs in Florida (Dustan 1977), St.

Croix (Rogers et al. 1984),

Netherlands Antilles (Bak and Engel

1979), and Jamaica (Rylaarsdam 1983).

In all of these studies, coral recruitment was measured as the number

of small corals (usually fewer than 4

or 5 cm in diameter) per unit area.

These numbers cannot be translated

into the numbers of larvae settled per

square meter per year because it is

not known what the early growth rates

are, nor what patterns of settlement

and mortality produce the observed

counts of "juveniles."

In St. Croix (Rogers et al. 1984),

juveniles of Acropora palmata

occurred at densities of O.l-0.3/m2 at

depths of 3-9 m. Bak and Engel (1979)

found similar densities (0.13/m2) at

similar depths.

Juveniles of Acropora cervicornis

occurred at densities of O.Ol-0.30/m2

at a depth of 9 m (Rogers et al.

1984). Bak and Engle (1979) found no

juveniles of this species in surveys

totaling 45 m2 at appropriate depths.

Rylaarsdam (1983), however, reported

about 4.3/m2 on barren substrate 11 m

deep chosen because settled larvae

were abundant there.

Juveniles of Montastraea annularis

live at densities of 0.09-1.04/m2 at

depths of 9-27 m in St. Croix (Rogers

et al. 1984), 0.20/m2 at depths of

3-9 m in Curacao (Bak and Engel 1979),

5
and 2.33-9.32/m2 on bare substrates at

a depth of 11 m in Jamaica (Rylaarsdam

1983).

Juveniles of Montastraea cavernosa

are found at densities of 0.14-1.09/m2

at depths of 9-27 m and 0.05/m2 at

37 m on St. Croix (Rogers et al.

1984). In Curacao, densities are

0.07/m2 in 9-17 m and 0.20/m2 at

26-37 m (Bak and Engel 1979).

Although both species of Montastraea

are common as adults on the Texas

Flower Garden Banks, neither M.

annularis nor M. cavernosa recruits to

artificially -constructed settling

plates there (Baggett and Bright

1985). The genus Acropora does not

occur in either adult or juvenile form

in this locality.

Phototransects on the north coast of

Jamaica (Porter et al. 1981) revealed

no conspicuous sexual recruitment of

larvae in waters less than 20 m deep

in 1976 or 1980 for any of the four

species treated here. The difference

between our study in Jamaica and the

others mentioned above is that the

phototransect method identified only

corals that had successfully settled

and grown into the light, where they

could be recorded by the camera; in

contrast, studies by Bak and Engel

(1979), Rogers et al. (1984), and

Rylaarsdam (1983) also included

individuals that settled cryptically.

These authors commented on (1) the

disproportionate rarity of juveniles

relative to the commonness of adults

of these four species, and (2) the

rarity of juveniles of these four

species in relation to the commonness

of juveniles in other taxa of corals,

such as the lettuce corals (Agaricia).

The phototransect data from Jamaica

(Porter et al. 1981) revealed that, by

far, most recruitment of new coral

colonies in all four species occurred

by asexual rather than sexual

reproduction. In both species of

Acropora, fragmentation of adult

colonies is the most common means of

forming new colonies (Gilmore and Hall

1976; Davis 1977; Tunnicliffe 1981;

Bak and Criens 1982; Neigel and Avise

1983). Fragmentation occurs during

storms (Highsmith et al. 1980; Porter

et al. 1981; Tunicliffe 1981;

Highsmith 1982), but the susceptibility of a branch to breakage may be

enhanced by boring by clionid sponges

or lithophagan bivalves (Goreau and

Hartman 1963; Neumann 1966; MacGeachy

1975; Hudson 1977). New colonies

appear asexually in Montastraea by

processes of fission or separation

resulting from the death of intervening tissue.

GROWTH CHARACTERISTICS

Coral growth has been described in

terms of either linear extension rates

or rates of deposition of calcium

carbonate (Bak 1973; Barnes and Taylor

1973; Dodge and Thomson 1974; Dodge et

al. 1974; Buddemeir and Kinzie 1976;

Gladfelter and Monahan 1977). Both

linear extension and skeletal weight

are known to be influenced by light

(and its interaction with water depth

and water clarity) and by sea temperature. What is known of the shallow

water (~10 m) linear growth rates of

the four species of interest is

reviewed here.

Acropora cervicornis may be the most

rapidly growing coral in the world in

terms of linear extension (Chalker

1977; Chalker and Taylor 1978;

Gladfelter et al. 1978). Linear

extension rates were as great as

26.4 cm/year in Jamaica (Lewis et al.

1968), averaged 14.4 cm/year in

Barbados (Lewis et al. 1968), and

averaged between 5.1 cm/year (Vaughan

1915) and 10.0 cm/year (Shinn 1966) in

Florida. Extension rates between

5.9-10.0 cm/year were measured in St.

Croix, U.S. Virgin Islands (Gladfelter

et al. 1978; Gladfelter 1984). All of

these authors, together with J.W.

Porter (unpublished data from Panama)

recognized the extreme variability in

growth rates from branch to branch on

.

6
the same colony and from season to

season on the same branch.

The growth rate and many other

physiological parameters of Acropora

cervicornis are adversely affected by

increases in turbidity and sedimentation (Kendall et al. 1985). Acropora

palmata grows at rates of

4.7-10.2 cm/year in St. Croix

(Gladfelter and Monahan 1977;

Gladfelter et al. 1978), 6-10 cm/year

in Curacao (Bak 1976), and

5.0-9.5 cm/year it, Florida (Vaughan

1915).

Literature on the growth rate of

Montastraea annularis is extensive.

Of special interest is the use of

growth rates of this species in the

Florida Keys as an environmental

indicator of cold-water stress (Hudson

et al. 1976; Hudson 1981, 1982) and

water quality (Dodge et al. 1974).

Growth rates (cm/year) of

Montastraea annularis were reported to

be 0.81-0.90 in the Florida Keys

(Agassiz 1890), 0.46 at Key West,

Florida (Weber and White 1977), 0.68

(Vaughan 1915) and 0.50 (Macintyre and

Smith 1974) in the Dry Tortugas, 1.07

in the Florida reef tract (Hoffmeister

and Multer 1964), and 1.0-1.2 in

Florida (Hudson et al. 1976). Elsewhere in the Caribbean, rates varied:

0.47-0.67 in Jamaica (Weber and White

1977; Dustan 1975), 0.66-0.89 in St.

Croix (Gladfelter et al. 1978),

0.01-2.40 in Barbados (Lewis et al.

1968; MacGeachy 1975; Tomascik and

Sander 1985), 0.68-0.73 in Curacao

(Bak 1976), 0.50 in Panama (Weber and

White 1977), and 0.71 in Belize (Weber

and White 1977).

Growth rate decreases significantly

as water turbidity increases (Dodge et

al. 1974; Loya 1976; Dodge and Vaisnys

1977; Bak 1978). In water 2 m deep

with low sediment resuspension rates

of 0.5 mg x cm-2 x d-l, maximum growth

rates of M. annularis exceeded

1.1 cm/year and averaged 0.88 cm/year

(Dodge et al. 1974); in areas of

similar depth, but with higher

sediment resuspension rates of 1.1

mg x cm-2 x d-l, maximum growth rates

were significantly lower and did not

exceed 0.7 cm/year (average,

0.6 cm/year).

Linear growth rates of Montastraea

cavernosa were 0.35-0.57 cm/year in

the Dry Tortugas, Florida (Vaughan

1915), and 0.32 in Key West, Florida

(Weber and White 1977). Average rates

of 0.32 in Belize, 0.35 in Jamaica,

and 0.32 in Panama were measured by

Weber and White (1977).

POPULATION CHARACTERISTICS

Depth Distribution

As shown in part by the depth ranges

and zones of maximum abundance of the

four species treated here (Table 1 as

determined in Jamaica), the basic

zonation of almost all Caribbean coral

reefs consists of Acropora palmata in

shallow water, a mixed zone of A.

cervicornis and Montastraea annularis

just below this, and a zone, below

30 m, not always developed, where

Acropora is absent and M. annularis

and M. cavernosa dominate the

horizot?tal surfaces. Species in the

genus Agaricia dominate steeply

sloping or vertical surfaces at these

greater depths.

Response to Hurricanes

Hurricanes frequently influence reef

structure. Moderate storms can be

expected in southern Florida at least

once every 5 years and severe storms

perhaps once per decade (Jaap 1984).

During the 20-year period 1966-85,

however, the occurrence of severe

storms in this region was unusually

low. After Hurricane Allen (1980),

Porter et al. (1981) demonstrated that

in water less than 10 m deep,

mortality was 95% in Acropora palmata

and 98% in A. cervicornis, but only

15% in Mont>straea annularis. (M.

cavernosa was not common enouah in

7
shallow water to enable quantification

of its response to this storm.) At

30 m there was no damaqe to A.

cervicornis, M. annularis, or M.

cavernosa. (Acropora palmata was not

found at this depth.) After the

storm, mortality was 98% in fragments

of 4. palmata (J.W. Porter, unpubl.

data) as well as in storm-generated

fragments of 4. cervicornis (Knowlton

et al. 1981)--mostly due to intense

predation from grazing snails (Coralliophila) and sea urchins, whose

populations were little affected by

the storm.

Morphologically, both species of

Montastraea, but neither species of

Acropora, are well equipped to survive

moderate storms. However, the thick

fronds of A. palmata and its interlocking branching morphology confers

some degree of protection that A.

cervicornis does not have (Ball et.

al. 1967; Stoddart 1969, 1974;

Tunnicliffe 1981; Rogers et al. 1982).

Further, A. palmata resists fracture

better than A. cxcornis because of

its continuous multiple bundles of

axial corallites in its branch core

(Constanz 1984). This may in part

explain its distribution in more

exposed areas.

From an evolutionary standpoint, it

is probably not coincidental that most

Caribbean coral species (including all

of those reviewed here) reproduce

during hurricane season when there

would be a higher availability of

newly bared substrate for successful

larval settlement.

FISHERY

Commercial coral collecting has been

banned in State of Florida waters

since 1974 (Shinn 1979; Jaap 1984).

Extension of this ban to all of the

Gulf of Mexico and the South Atlantic

has been recommended on the grounds

that low growth rates and low yields

make coral collecting similar to

mining a nonrenewable resource

(Anonymous 1982).

Poaching is almost impossible to

prove, although it is probably common.

Florida shell shops sell specimens of

Acropora palmata and A. cervicornis at

a high price. While these shops state

that the specimens for sale come from

Haiti, the high rate of sale of these

spectacular branching colonies, and

their ease of collection in Florida,

may actually increase poaching on

local coral populations.

Although Ross (1984) proposed a

classical fisheries model to estimate

maximum sustainable yield for a

branching Pacific coral, his model

did not incorporate partial mortality

and fragmentation of colonies; consequently, further refinement will

be required before it is applicable to

Caribbean Acropora.

ECOLOGICAL ROLE

Photosynthetic Capability

Reef-building corals harbor the

symbiotic dinoflagellate alga Symbiodinium microadriaticum (Taylor 1973;

Trench 1979). New evidence (Blank and

Trench 1985a, b) suggests that previously identified strains of this alga

(Schoenberg and Trench 1980) may prove

to be separate species. The alga constitutes only 5% to 15% of the dry

weight of living tissue (Muscatine and

Porter 1977), but the effect on the

colony is profound; it causes coral

heads of all of these species to produce more oxygen and fix more carbon

than they consume, under optimal sunlight conditions (Table 2). For the

last half century, a controversy has

raged about whether corals get their

food primarily from photosynthesis or

by suspension feeding on plankton.

That is to say, although they are

taxonomically animals, are they

functionally plants? Acropora corals

look like plants and were so

classified until the early 1800's.

a
Table 2. Patterns of primary production (maximum gross production (P) and

nocturnal respiration (R) as ug0, cm-*h-l) and instantaneous production/

respiration ratios (P/R) for four reef-building corals (J.W. Porter, unpubl.

data).

Acropora Acropora Montastraea Montastraea

palmata cervicornis annularis cavernosa

Depth (m) P R P/R P R P/R P R P/R P R P/R

1 89.5 14.3 6.3 34.4 7.7 4.5 109.9 30.6 3.6 62.2 26.2 2.4

10 52.9 11.8 4.5 42.6 8.2 5.2 56.9 20.6 2.8 42.6 21.3 2.0

30 37.4 6.8 5.5 55.8 16.0 3.5 43.8 14.6 3.0

50 44.7 14.0 3.2 26.9 11.6 2.3

Using polarographic oxygen chambers

at various depths, Porter (1980)

demonstrated that even at a depth of

50 m, Montastraea annularis was capable of effecting integrated 24-h production/respiration ratios (P/R) in

excess of 1.0 (indicating autotrophy,

or self-sufficiency, with respect to

carbon) on sunny days. Autotrophy in

the animal depends on the amount of

fixed carbon that is transferred to

the animal relative to the respiration

of the animal. (See Porter et al.

1987 for an extensive review of the

literature on photosynthesis and

respiratory metabolism of this

species.) However, if one integrates

natural surface irradiance, including

the seasonal effects of sun angle and

cloudiness, colonies of M. annularis

in water shallower than 10 m deep are

capable of such autotrophy on a sustained basis; colonies of this species

below that depth are not (Porter

1985). Colonies of M. annularis below

10 m must feed on particulate organic

carbon (Lewis and Price 1975) or

zooplankton (Porter 1974) to make up

the deficit. At 50 m, the annual

deficit is roughly 30% of the metabolic requirement (Porter 1985).

In examination of production rates,

respiration rates, and instantaneous

P/R ratios for the four species of

corals under examination here, Table 2

reveals several trends. For all

species, both production and respiration rates decrease with increasing

depth (Davies 1977, 1980; Porter et

al. 1987). Furthermore, at any

given depth, the P/R ratios of

branching Acropora are always greater

than the same ratios for the moundshaped corals Montastraea. This

higher P/R ratio is reached because

the Acropora species here have a

combination of higher production rates

(Chalker and Taylor 1978) and consistently lower respiration rates

than the Montastraea species

(Table 2).

As of 1986, the developing evidence

suggests that, although the symbiotic

association is an obligate one for

these and other coral species, corals

exist on a nutritional cline. The

cline ranges from those like the

Acropora, which are almost totally

dependent on sunlight for survival, to

those like Montastraea, which are less

dependent on sunlight and more

dependent on zooplankton for nourishment (Porter 1976; Lewis 1977). In

this context it is useful to note that

A. palmata drops out of the reef at

the depth where its 24-h P/R ratio

9
falls below 1.0 (J.W. Porter, unpubl.

data). Mouth sizes and tentacle

lengths also correlate with

morphological (surface/volume) and

physiological (P/R) ratios (Porter

1976). Polyps of M. cavernosa

(Richardson et al. 1979) feed on

zooplankton much more effectively than

do 'polyps of A. cervicornis (Porter

1974).

The management implications of these

basic biological data are clear: the

branching Acropora species are much

more susceptible to increases in water

turbidity than are the mound-shaped

Montastraea (Dallmeyer et al. 1982).

Dredging or pollution that reduces the

long-term water clarity can reduce P/R

ratios below unity for all four

species. Species of Acropora may not

be able to compensate with an

alternate food supply, such as zooplankton, which is used by Montastraea

colonies. One expected result is that

the depth distribution of Acropora

will be truncated in turbid water

(Griffin 1974; Johannes 1975; Loya

1976; Dodge and Vaisnys 1977; Bak

1978; Kendall et al. 1985). Under

conditions of turbidity, Loya (1976)

noted a reduction in the densities of

colonies of Montastraea annularis and

an increase in the densities of

colonies of M. cavernosa.

Interspecific Competition

Lang (1973) described the ability

of reef corals to extend their

mesenterial digestive filaments onto a

neighboring coral species and digest

away its living tissue. A dominance

hierarchy exists within the coral

species. The defense hierarchy for

the four species examined here is

Montastraea annularis > p1. cavernosa >

Acropora palmata A . > cervicornis

(Lang 1973). Species-in the family

Acroporidae are generally at the

bottom of the pecking order, whereas

species in the family Faviidae are

generally near the top. Lang (1973)

has speculated that the superior

defense abilities exhibited by favid

corals like Montastraea may be a

compensatory mechanism for the

superior overgrowth capabilities

of the acroporid species. These

allelochemical mechanisms may be of

special significance in periods

between hurricanes, because catastrophic storms always favor the

massive morphology of Montastraea over

the branching morphology of Acropora.

Although the attack patterns of

extracoelenteric destruction were

initially thought to be a fixed

property of each species (Lang 1973),

P. Dustan (Department of Biology,

College of Charleston, Charleston,

S.C.) unpubl. observations) identified

several forms of M. annularis which

destroy each other- In addition, the

pecking order for the 65 species of

hermatypic coral species in the

Caribbean is not identical for all

localities (Lang 1973).

Predators

All four species reviewed here are

subject to predation, but "plagues" of

coral predators such as Indo-Pacific

Acanthaster (the crown-of-thorns

starfish) observed by Endean (1976)

have not been described for the

Caribbean.

A major coral predator is the

polychaete bristle worm Hermodice

carunculata (Glynn 1962; Marsden 1962;

Ebbs 1966; Lizama and Blanquet 1975;

Antonius 1977). This predator impales

itself on the long branches of A,

cervicornis and denudes the branch

tips of their living tissue. The

predation marks show up as scars

several centimeters long on 4. cervicornis. Hermodice also digests tissue

of A. palmata and both Montastraea

spec?es; on these forms the predation

scars appear as white patches.

The coral-shell gastropod Coralliophila abbreviata is also axr

predator on all four species of corals

treated here (Rylaarsdam 1983;

C. Rosesmyth, Department of Zoology,

10
University of the West Indies,

Kingston, Jamaica; unpubl. data). Its

preferred food is Acropora cervicornis, but it also feeds extenmy

on Montastraea annularis. After

Hurricane Allen, when Acropora populations were decimated by storm damage,

C.

the

abbreviata significantly reduced

Acropora populations further.

Populations of Acropora that were

healthy before the storm could sustain

a low level of snail predation. This

same predation level after the storm

decimated the few remaining Acropora

colonies (Knowlton et al. 1981).

Although not widely documented, the

long-spined sea urchin Diadema

antillarum is known to eat Acropora

(Bak and Van Eys 1975; Sammarco 1980,

1985). It does so actively when

starved, but may also do so

adventitiously while feeding on

turf algae growing next to coral

tissue.

Hermit crabs such as Petrochirus

diogenes and Paguristes sp. live

between the branches of both species

of Acropora (Gilchrist 1985). They

sometimes cause extensive damage, and

can decimate their host species

locally when their population numbers

are high.

Finally, the three-spot damselfish

Pomacentrus planifrons Cuvier establishes alqal qardens on branchina

Acropora when it is available and on

Montastraea annularis when Acropora is

rare (Thresher 1976; Brawley and Adey

1977; Kaufman 1977; Itzkowitz 1978;

Williams 1978; Sammarco and Carleton

1982). The fish nips off the living

coral tissue and weeds the garden as

algae begin to grow on the bared coral

skeleton.

Predation of any kind may be

repaired by tissue regeneration over

the dead region, or the area may be

colonized by other epibiota. These

dead regions may allow settlement by

boring sponges and bivalves that can

further weaken the skeleton (MacGeachy

1975; Bak et al. 1977)

Sediment Rejection Behavior

Sediment rejection is an important

behavioral characteristic influencing

the growth, survival, and distribution

of reef corals. The four species

considered have differing abilities to

shed sediment. Hubbard (1973) and

Hubbard and Pocock (1972) ranked them

in terms of their ability to remove

silt (particles t62 urn in diameter),

fine sediments (62-250 pm), and coarse

sediments (250-2,000 pm). Acropora

palmata and A. cervicornis were unable

to remove coarse sediments and only

weakly able to remove fine sediments.

Montastraea annularis and M.

cavernosa, on the other hand, were

weakly able to remove coarse sediments

and highly competent at removing fine

sediments. In al 1 four species,

ciliary movements remove silt and

clay-sized particles. Hubbard and

Pocock (1972) showed general

correlations between coral zonation

and sediment-rejection capability.

Although it is clearly an influencing factor, the differential

response to hurricanes, ordinary

storms, light gradients, turbidity,

other physical effects, and biotic

interactions are probably of greater

causal importance than sediment

rejection properties in determining

the overall vertical and lateral distribution of these species along a

reef. Further, the effect of

suspended sediments, even when successfully shed, tends to increase

respiration and decrease photosynthesis (J.W. Porter, unpubl. data).

Photosynthesis decreases because of

reduced light penetration, and P/R

ratio ’ concomitantly

(Dallmeyeyet al. 1982).

reduced

Water movement (turbulence) and gravity are

probably more important in removing

sediments from the branching Acropora

species than are their capabilities

for sloughing sediments in stagnant

water.

11
Disease

Coral diseases are not well

understood (Voss 1973; Antonius 1982;

Bak 1983; Peters 1984). Two diseases

recently received considerable attention in the Florida area. The first

is "white-band" disease, occurring in

epidemic proportions on Floridian

colonies of Acropora palmata (Jaap

1984). Gladfelter (1982) demonstrated

that this unidentified pathogen also

severely reduced populations of A.

palmata in St. Croix, U.S. Virgin

Islands. This disease also attacks A.

cervicornis (J.W. Porter, unpubl.

data).

The second disease is caused by

infections of the blue-areen alaa

Phormidium corallyticum - (Antoni&

1977, 1982; Rutzler and Santavy 1983;

Taylor 1983; Rutzler et al. 1983).

Because of its appearance as a black

ring around the infected area, it has

been called the "black-band" disease.

It kills both species of Montastraea,

but neither species of Acropora

(Rutzler et al. 1983). It may be the

same disease described by Mitchell and

Chet (1975) and Ducklow and Mitchell

(1979a, b) as

al infection.

an unidentified bacteriCancerlike ulcers have also been

described on corals (Preston 1950;

Squires 1965; Bak 1983; Peters 1984;

Peters et al. 1986).

In all cases of algal or bacterial

disease, prior stress to the coral

increases both the likelihood that an

individual colony will contract a

disease and the likelihood that it

will subsequently be killed by it

(Peters et al. 1986).

ENVIRONMENTAL REQUIREMENTS

In general, the environmental

requirements of all reef-building

corals are narrow. Clear, warm,

well-oxygenated sea waters of constant

salinity, such as that off southern

Florida (Churgin and Halminski 1974a,

b; NOAA 1981) are requisite for the

development of coral reefs. I

describe here some of the minor

differences between the four present

species.

Temperature

Acropora cervicornis and A. palmata

are hiahlv sensitive to lower than

usual temperatures (Mayer 1914, 1916;

Shinn 1976; Porter et al. 1982). In

January 1977 surface waters on the Dry

Tortugas reefs dropped to 14 OC

(Walker 1981). About 96% of all

Acropora colonies in water less than

2 m deer, died durina this cold-water

intrusion; Montastraea was too rare at

this depth to allow measurement of

mortality (Porter et al. 1982). At

13 m, where water temperatures may not

have been as low, mortality was 15% in

Acropora but only 2% in Montastraea.

In Bermuda, surface water temperatures

as low as 8 OC have been recorded

(Verrill 1902); no species of the

genus Acropora live off Bermuda. Low

water temperature probably limits the

northward extension of Acropora in

Florida. Perhaps Montastraea is found

much farther north than Acropora on

both the aulf and Atlantic coasts of

Florida (IFigure 2) because it is

capable of flourishing in deeper water

with less temperature stress.

Cold stress can also affect

Montastraea, which has a lower lethal

temperature of 13.9 OC (Mayer 1914,

1916). Cold- or warm-water stresses

occur frequently at Hens and Chickens

Reef off Plantation Key, because these

reefs lie near Florida Bay, where

temperature fluctuates greatly in

response to atmospheric conditions

(Lee and Mooers 1977). Cold stress

has been well documented in M.

annularis as distinct skeletal bands

of high density, or whole or partial

mortality of colonies (Hudson et al.

1976; Dustan 1977).

s

Although August temperatures average

29.3 OC in the Dry Tortugas and

17
.

28.7 OC at Fowey Rocks (Vaughan 1918),

water temperatures as high as 31 'C

have been recorded (Vaughan 1918).

Mayer (1918) reported that the upper

lethal temperature is 35.8 OC for $.

palmata, and 33.8 ='C for

cervicornis (Shinn 1966). TheJe

elevated temperatures would occur only

in coral habitat at the surface and

under unusual oceanographic conditions

(Jaap 1984). Jaap (1979) recorded an

example of putative heat stress at

Middle Sambo Reef, Florida, involving

bleaching and loss of symbiotic

zooxanthellae from A. palmata and A.

cervicornis. Affected specimens r-ecovered after 6 weeks.

Salinity

Corals prefer salinities between 27

and 40 ppt (Lewis et al. 1968). Both

Acropora and Montastraea can survive

l-h exposure to seawater approaching

20 PPt, such as occurred after

torrential rains in Panama in 1971

(J.W. Porter, unpubl. data), but

coral reefs never occur where oceanic

salinities are consistently below 33

PPt. Typical responses to lowered

salinity are the loss of zooxanthellae

(Goreau 1964) and the production of

mucus (Coffroth 1985). It is unclear

whether the absence of corals in

slightly brackish water is due to the

reduced salinity itself, to increases

in suspended sediments, or temperature

fluctuations often associated with

areas of inflowing freshwater (Dole

and Chambers 1918).

Dissolved Oxygen

Like all coral species, the four

species reviewed here exist where

dissolved oxygen rarely extends beyond

the range of 90% to 125% saturation

(Jaap and Wheaton 1975). Montastraea

annularis tolerates oxygen tension

above 100% (Wells and Wells 1971), and

probably most coral species are able

to tolerate very high oxygen tensions

because the coral tissue contains

algae that produce large amounts of

oxygen. As a correlate of this,

corals contain high concentrations of

superoxide dismutase, a compound which

burns off excess oxygen (Dykens and

Shick 1982). Conversely, given the

ability to produce oxygen, corals are

also able to tolerate waters with

exceedingly low oxygen tensions

(Burris et al. 1983). In closed

respirometry experiments in which the

flush-pump failed but where the oxygen

recorder continued to work (J.W.

Porter, unpubl. data), both Acrdpora

cervicornis and M. annularis tolerated

oxygen tensions as low as 1.0 ppm for

1 -h, but both died when these-conditions persisted for 6 h.

The absence of these species in

seawater in which oxygen is less than

saturated, like their absence in water

of low salinity, probably reflects

intolerance to both depressed oxygen

levels and associated factors rather

than to low dissolved oxygen alone.

Substrate

All four of these corals must settle

and grow on a stabilized, shallow

(Table 1) sea floor. Neither the

adults nor the attached juveniles are

capable of surviving in areas of heavy

sedimentation (Hubbard and Pocock

1972). All four species can colonize

stable vertical or sloping substrates

above sandy bottoms, and large

detached fragments of Acropora

cervicornis can roll into sandy areas

and colonize them (Tunnicliffe 1981),

and thus support the development and

spread of patch reefs.

Solar Irradiance

Light intensity decreases rapidly

with increasing depth (Gordon and Dera

1969; Brake1 1979; Dustan 1982). As

judged by work in Jamaica on Acropora

(Porter 1980) and Montastraea (Porter

et al. 1987), A. palmata is most

abundant where the maximum ambient

irradiance reaches 1,200-1,800 pE

m-2s-1 and optimal integrated daily

irradiance is about 28 E m-2d-1.
Populations of A. cervicornis flourish

where peak amb:ent irradiance levels

are 300-800 uE m-2s-1 and optimal

integrated irradiance approaches

15 E m-2d-1. Montastraea annularis

populations dominate substrates over a

broader depth range but still are most

common where maximum instantaneous

light intensities are

m_2s_1

200-800 pE

and integrated values approach

lo-15 E m-2d-1. Montastraea cavernosa

is most common just below the zone of

greatest development of M. annularis.

Here peak light intensity reaches

600 pE m-2d-1 and the integrated

daily total approaches 12 E m-2d-1.

Both M. annularis and M. cavernosa are

still-common at 65 rn, where instantaneous light does not exceed

50 pE m-2s-1 and daily totals do not

exceed 2 E m-2d-1.

Solar radiation probably controls

the intensity and timing of reproductive activity in all four species.

This may be especially true for M.

annularis and M. cavernosa becauze

most of their -colonies exist well

below their light compensation depth,

where their integrated production/

respiration ratio is below 1.0 (Porter

et al. 1987).

14
LITERATURE CITED

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Agassiz, L. 1869. Florida reefs,

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15
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lEPORl DOCUMENTATION 1. “PORT No, 2 3. hClP4Wt’S kC..slon IQ.

PAGE Biological Report 82(X.73)*

, Title l d subtltl* 5. R.oorl oat*

pecies Profiles: Life Histories and Environmental Requirements of August 1987

oastal Fishes and Invertebrates (South Florida)--Reef Building '&

orals

. AuthodS) a. hfwmine O~~nlztilon R*pt. N,

James W. Porter

. hdwmlng Oqw#lxation Name and Addnss 10. Pmi=Vfask/Work Unit No.

11. CO+ttr~Ct(c) or Crmtt(G) No.

(0

2. Sgonsori”~ OrSwIizatlon N,m. and Address

-(C)

lational Wetlands Research Center U.S. Army Corps of Engineers 13. Tyw of Report L Pariod Gown,

'ish and Wildlife Service Waterways Experiment Station

J.S. Department of the Interior P.O. Box 631

dashington, DC 20240 Vicksburg, MS 39180 II.

IS. Suppl.m.ntary Notes

*U.S. Army Corps of Engineers Report No. TR EL-82-4

6. Ab8tnct (Limit: 200 words)

Species profiles are literature summaries of the taxonomy, morphology, range, life

history, and environmental requirements of coastal aquatic species. They are designed

to assist in environmental impact assessment. Four species of reef-building corals are

considered: elkhorn coral, staghorn coral, common star coral, and large star coral.

All four species spawn annually in the fall during hurricane season. Juvenile

recruitment is low in all four species. Rapid growth rates of species in the genus

Acropora (lo-20 cm/yr) contrast with slower growth rates of species in the genus

Montastraea (1.0-2.0 cm/yr), but both species of Montastraea are also important in reef

development due to their massive form and great longevity. Shallow-water colonies of

Montastraea survive hurricanes; shallow colonies of Acropora do not. Because of their

dependence on photosynthesis for all of their carbon acquisition, the Acropora species

reviewed here have a more restricted depth distribution (O-30 m) than do the Montastraea

species considered (O-70 m). All four species are subject to intense predation by the

snail predator, Coralliophila. Species of Montastraea are susceptible to infection from

blue-green algae, which produce "black band disease"; species of Acropora are

susceptible to a different, as yet unidentified pathogen, that produces "white-band"

disease. Increased water turbidity and sedimentation cause reduced growth rates and

partial or whole mortality in all four species.

7. DownwIt Analysis a. O*scripton

Growth Salinity Reefs Feeding habits

Photosynthesis Temperature Coral Sediments

Life cycles Depth Oxygen

b. ldantlfnn/Open-Ended Terms

Reef-building corals Elkhorn coral Acropora palmata

Staghorn coral Acropora cervicornis Montastraea annularis

Large star coral Montastarea cavernosa Common star coral

c. COSATI Field/Group

S. AvaIlability Statement 19. Sccurlty Class (This Repoti) 21. No. of PaSea

Unclassified 23

Unlimited release

- -

20. kcurity Class This Page) 22. price

Unclassified

). ANSI-239.18) OPTIONAL FORM 272 (4-77)

(Formerly NTIS-35)

Oep.rtm.nt oi comm.rc.
As the Nation’s principal conservation agency, the Department of

the Interior has responsibility for most of our nationally owned

public lands and natural resources. This includes fostering the

wisest use of our land and water resources, protecting our fish

and wildlife, preserving the environmental and cultural values of our

national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department assesses

our energy and mineral resources and works to assure that their

development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation

communities and for people who live in island territories under U.S.

administration.

U.S. DEPARTMENT OF THE INTERIOR

FISH AND WILDLIFE SERVICE

TAKEPRIDE

in America

UNITED STATES

DEPARTMENT OF THE INTERIOR

FISH AND WILDLIFE SERVICE

National Wetlands Research Center

NASA-Slide11 Computer Complex

1010 Gause Boulevard

Slidell, LA 70458