L

QL

155

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no.82-

11.110

BiologiCal Report 82(11.110)

August 1989

Library

National Wet!ands Research Center

U. S. Fish and Wildllfe Servfce

700 Cajifundome Boulevard

TR EL-82-4

Species Profiles: Life Histories and

Environmental Requirements of Coastal Fishes

and Invertebrates (South Florida)

BLACK, RED, AND NASSAU GROUPEFJS

Fish and Wildlife Service

Coastal Ecology Group

Waterwavs ExDeriment Station

U.S. Department of the Interior U.S. Army Corps of Engineers

Biological Report 82(11.110)

TR EL-82-4

August 1989

Species Profiles: Life Histories and Environmental Requirements

of Coastal Fishes and Invertebrates (South Florida)

BLACK, RED, AND NASSAU GROUPERS

Darryl E. Jory and Edwin S. Iversen

University of Miami

Rosenstiel School of Marine and Atmospheric Science

Division of Biology and Living Resources

4600 Rickenbacker Causeway

Miami, FL 33149

Project Officer

David 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
DISCLAIMER

The mention of trade names in this report does not constitute endorsement nor recommendation for use by the U.S. Fish and Wildlife Service or the

Federal Government.

This series may be referenced as follows:

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

and environmental requirements of coastalfishes and invertebrates. U.S. Fish

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

This profile may be cited as follows:

Jory, D.E., and E.S. Iversen. 1989. Species profiles: life histories and

environmental requirements of coastal fishes and invertebrates (south

Florida)--black, red, and Nassau groupers. U.S. Fish Wildl. Serv. Biol. Rep.

82(11.110). U.S. Army Corps of Engineers, TR EL-82-4. 21 pp.
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 Wetlands Research Center

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

iii
CONVERSION TABLE

Metric to U.S. Customary

Multiply

millimeters (mm)

centimeters (cm)

meters(m) ’ ’

meters (m)

kilometers (km)

kilometers (km)

square meters (m2)

square kilometers (km2)

hectares (ha)

liters (I)

cubic meters (m3)

cubic meters (m3)

0.03937

0.3937

3.281

0.5468

0.6214

0.5396

10.76

0.3861

2.471

0.2642

35.31

0.0008110

milligrams (mg)

grams (9)

kilograms (kg)

metric tons (t)

metric tons (1)

kilocalories (kcal)

Celsius degrees (“C)

0.00003527

0.03527

2.205

2205.0

1.102

3.968

1.8(%) + 32

U.S. Customary to Metric

inches 25.40 millimeters

inches 2.54 centimeters

feet (ft) 0.3048 meters

fathoms 1.829 meters

statute miles (mi) 1.609 kilometers

nautical miles (nmi) 1.852 kilometers

square feet (ft2)

square miles (mi2)

acres

0.0929 square meters

2.590 square kilometers

0.4047 hectares

gallons (gal)

cubic feet (ft3)

acre-feet

3.785 liters

0.02831 cubic meters

1233.0 cubic meters

ounces (02) 28350.0

ounces (oz) 28.35

pounds (lb) 0.4536

pounds (lb) 0.00045

short tons (ton) 0.9072

British thermal units (Btu) 0.2520

Fahrenheit degrees (OF) 0.5556 (“F - 32)

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

milligrams

grams

kilograms

metric tons

metric tons

kilocalories

Celsius degrees

iV
CONTENTS

Page

iii

iv

vi

:

2

;

4

5

5

:

6

:

8

9

9"

::

10

11

::

13

13

15

15

17

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

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

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

NOMENCLATURE/TAXONOMY .......................................................

REASONS FOR INCLUSION IN THE SERIES .........................................

GEOGRAPHIC RANGE ............................................................

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

Morphological Characteristics ..............................................

Morphological Differences .................................................

Color and Piqmentation ....................................................

Size........:......... ....................................................

L IFE HISTORY ................................................................

Habitat ...................................................................

Sex Reversal and Spawning .................................................

Eggs and Larvae ...........................................................

Juveniles .................................................................

Movement and Migration ....................................................

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

FISHERIES ...................................................................

Recreational and Commercial Fisheries .....................................

Marketing .................................................................

Regulations ...............................................................

Artificial Reefs ..........................................................

Contaminants ............................................................

ECOLOGICAL ROLE .............................................................

Food and Feeding Habits ...................................................

Competitors and Predators .................................................

Parasites and Diseases ....................................................

Ciguatera (Tropical Fish Poisoning) .......................................

Behavior ..................................................................

REFERENCES .... ............................................................

V
c

ACKNOWLEDGMENTS

We are grateful for the useful references contributed by C. S. Manooch III

of the National Marine Fisheries Service, and for his thorough review. S.

Bortone of the University of West Florida also reviewed the manuscript. E. Snell

and R. Vaught of the National Marine Fisheries Service provided catch data and

pertinent literature respectively. B.E. Luckhurst also made available useful

literature. J. Iversen reviewed the general draft and made numerous useful

comments. G. A. Maury prepared the cover and Figure 1. V.R. Restrepo, E.

Lahman, J.F. Carranza, and I. Paez kindly assisted with word processing and

greatly facilitated the completion of this report.

vi
RED

NOMENCLATURE/TAXONOMY

Figure 1. Black, red, and Nassau groupers.

BLACK, RED, AND NASSAU GROUPERS

REASONS FOR INCLUSION IN THE SERIES

Scientific name . . . . . . . . . ..Mycteroperca

bonaci Poey, 1860

Preferred common name....Black grouper

Other common names . ..Marbled rockfish,

black rockfish, snider grouper, carbarita, junefish (Figure 1)

Scientific name . . . . . . . . . . ..Epinephelus

morio Valenciennes, 1828

Preferred common name......Red grouper

(Figure 1)

Scientific name . . . . . . . . . . ..Epinephelus

striatus Bloch, 1792

Preferred common name...Nassau grouper

Other common names . . . . . . . . . . ..Grouper.

rockfish, hamlet (Figure 1)

Class ................... ..Osteichthye s

Order......................Perciforme s

Family......................Serranida e

Black, red, and Nassau groupers

are actively sought by both commercial

and sport fishermen throughout their

geographic ranges. The red grouper is

one of the most abundant of the 17

species of grouper caught in Florida by

commercial and recreational fishermen.

The 1984 grouper landings for Florida

totaled about 10 million lb, worth $14

million. The south Florida region

(from Citrus around to Brevard Counties) contributed more than 8 million

lb (81% of the total, worth more than

$11 million (about 79% of the total).

The 1984 Florida finfish landings were

worth about $57.5 million, of which

groupers contributed $14 million or

nearly 25% of the total--making them

the most valuable marine finfish group

in Florida. However, despite their

economic importance, available information on their life histories in the

south Florida region--particularly

for black and Nassau groupers--is

1
grossly inadequate for effective management of the fishery. Beaumariage

and Bullock (1976) wrote that fewer

than two dozen pertinent studies had

then been published on the biology of

groupers, and that almost half of

these were based on tagging programs

that described movement patterns. According to these authors "This paucity

of data exemplifies the need for similar life histories studies, if sound

management policies are to be adopted...."

These three species of groupers

are an important component of the

highly diverse reef fish community,

occupying positions near the top of

the food webs. Their biology, ecology,

and exploitation are integral components of coral reef fisheries and

management of reef fisheries (Bohnsack

1982; Bannerot 1984). Any significant

change in their numbers could affect

the balance of reef ecosystems (May et

al. 1979). Preservation of coral reefs

is essential for the perpetuation of

harvestable grouper populations.

In south Florida, ciguatera (poisoning from consumption of tropical

fishes) is endemic. Many of the

reported cases from black grouper are

really from incorrectly identified fish

smuggled in from the Bahamas, where

ciguatera is much more common (Jones

Bohnsack, National Marine Fisheries

Service, Miami, FL; pers. comm.). Life

history data, especially on feeding

habits, of the species implicated are

needed to help prevent ciguatera. The

toxin causes gastrointestinal, cardiovascular, and neurological disturbances

resulting in prolonged disability and

long and expensive recovery periods

(de Sylva and Higman 1980; Poli 1982).

GEOGRAPHIC RANGE

The black, red, and Nassau groupers range from New England and

Bermuda to southeastern Brazil,

including the Bahamas, Gulf of Mexico

and all of the Caribbean (Bohlke and

Chaplin 1968; Smith 1971; Fischer

1978), although the three species are

rare north of Florida. Reports of

these species north of the Carolinas

are probably a result of larval

transport by currents, as suggested by

Thompson and Munro (1978) for other

grouper species.

The black grouper is

the Florida Keys (Randall

the Bahamas, and off

Venezuela (Cervigon 1966);

abundant in

1968), in

Cuba and

it is reportedly rare in the eastern Gulf of

Mexico (Smitp2leq_ttoal. 1975), the Virgin

Islands, Rico, and the

Colombian Caribbean (Dahl 1971).

The red grouper is primarily a

continental species, having the widest

distribution of all western central

Atlantic groupers (Roe 1976). It is

found mostly in broad shelf areas; its

center of abundance is in the Florida

shelf and the eastern Gulf of Mexico

(Moe 1969). It is also abundant in the

Colombian Caribbean (Dahl 1971), and

off northeastern Venezuela (Cervigon

1966), but uncommon in the West Indies

(Randall 1968).

The Nassau grouper is primarily

insular species very common in

t"ze West Indies (R.&da11 1968), the

Bahamas (Bohlke and Chaplin 1968),

southern Gulf of Mexico (Fischer

1978), and the Colombian Caribbean

(Dahl 1971). In Venezuela, it is common in the Archipelago Los Roques but

rare in northeastern islands such as

Margarita, Cache, and Cubagua

(Cervigon 1966). Red and Nassau groupers occur sympatrically in the Florida Keys (Figure 2), although their

local distribution is essentially disjunct (Moe 1969).

MORPHOLOGY AND IDENTIFICATION AIDS

Morphological Characteristics

Smith (1971) provided the following morphological descriptions.

3
- L

ATLANTIC

OCEAN

\

t7

8

I

TERSBURG

ALM BEACH

FLORIDA t

MIAMI

d-PE-ROMANO

o,_

MILES

Area of distribution 'I50-~

100

1

0 50 100

KILOMETERS

Figure 2. Approximate distribution of the black, red, and Nassau groupers in south

Florida.

3
Black grouper. "Large species of

Mycteroperca with robust body, relatively large scales, and rounded

preopercle. Dorsal fin XI, 17; anal

fin III, 12(13); pectoral fins 17;

gill rakers 20-26. Posterior nostril

not enlarged. Vertical fins without

exserted rays. First three dorsal

spines low, not forming an elevated

lobe. Gill rakers are moderate in

length. The exposed surface of the

maxilla is scaled. The upper lip is

broad, as wide as or wider than the

shaft of the maxilla at the midpoint

of the supramaxillary groove."

Red grouper. "Moderate-sized species of Epinephelus with large eyes,

small scales, robust body. Nostrils

subequal, posterior little larger than

anterior. Dorsal fin XI, 16-17 with

interspinous membrane not notched.

Second dorsal spine longest. Vertical

fins angulate in large fish. Anal fin

III, 9; pectoral fins 17; gill rakers

23-25."

Nassau grouper. "Moderate-sized

species of Epinephelus with large

eyes, medium-sized scales, robust

body. Nostrils subequal, posterior

slightly enlarged, comma-shaped in

adults. Dorsal fin XI, 16-17 with

interspinous membranes notched; anal

fin modally III, 8; pectoral fins 18;

gill rakers 24-25. Vertical fins

rounded."

Morphological Differences

Morphological differences among

grouper species were cited by Fischer

(1978).

Black grouper. This grouper can be

distinguished from species of the

genus Epinephelus by its more elon-'

gated body, which is not deepest at

the origin of the dorsal fin, and in

having 12, sometimes 13, soft anal-fin

rays (8-9 in Epinephelus spp.). It

differs from other species of

Mycteroperca in having a gently rounded preopercle, with no definite lobe

and only a slightly emarginated notch,

and from other serranids in having the

bases of the soft dorsal and anal fins

covered by scales and thick skin.

Red qrouper. This grouper can be

distinguished from other species of

the genus Epinephelus by its dorsal

fin, in which the second spine is the

longest and the interspinous membrane

is not notched. In E. nigritus, E.

flavolimbatus, r. mystacinus, and '.

niveatus, the pelvic fins are longer

than the pectoral fins and are inserted anterior to the pectoral-fin base,

whereas in the red grouper the pelvic

fins are shorter than the pectorals

and are inserted slightly behind the

ventral end of the pectoral fin base.

The red grouper differs from

Mycteroperca species in having a less

elongated body, and in having nine

soft anal-fin rays. The red grouper

differs from other serranids in

having a more robust body, having the

bases of the dorsal and anal fins

covered by scales and thick skin, and

in having 11 dorsal fin spines.

Nassau grouper. This grouper can be

distinguished from other species in

the genus Epinephelus by the third

spine of the dorsal fin which is

longer than the second, and in having

a slightly indented interspinous membrane. The caudal fin is slightly

emarginated. In Nassau groupers the

pelvic fins are shorter than the pectorals and are inserted below or

behind the ventral end of the pectoral

fin base. In E. mystacinus, E.

nigritus, E. flarolimbatus, and E.

niveatus, tTe pelvic fins are longer

than the pectorals and are inserted

anterior to the ventral end of the

base of the pectoral fins. In addition, &. mystacinus has greatly enlarged, equal-sized posterior nostrils, whereas in the Nassau grouper

the nostrils are subequal. The Nassau

grouper can be distinguished from

Mycteroperca spp. by its less elongated body and eight soft anal fin

rays. Other serranid species have

less robust bodies, soft dorsal and

4
anal fin bases that are not covered by

scales or thick skin, and 10 or fewer

dorsal fin spines.

Color and Pigmentation

As a rule, groupers can undergo

rapid and dramatic changes in color

and pattern, which are under nervous

and hormonal control. The extent of

change depends on their surroundings

and the grouper's own activity. Life

color can be used to distinguish different species, but specific color and

distinctive markings such as bars and

spots fade soon after the fish dies,

and undergo further discoloration when

the fish is preserved in Formalin or

alcohol.

The following coloration and pattern descriptions were compiled from

Rivas (1964), Cervigon (1966), Biihlke

and Chaplin (1968), Fischer (1978),

and the Source Document for the

Snapper-Grouper Fishery of the South

Atlantic Region (1983).

Black grouper. This species can

change its body coloration and color

pattern over a wide range, from light

tan or gray ground color with rows of

rectangular darker blotches, to dark

reddish gray with short dark bands, to

whitish with a few dusky bars on the

fin margins. In general appearance the

black grouper resembles the yellowfin

grouper (I. venenosa) and the gag

(M. microlepis), but the black grouper

can be distinguished by its straighter

posterior caudal margin and rows of

rectangular dark blotches on the body.

Blotches are larger and more

quadrangular and regularly aligned in

the black grouper than in the yellowfin grouper; in addition, each pectoral fin of the black grouper has a

narrow orange margin, whereas each of

those in the yellowfin grouper has a

broad yellow margin that sharply contrasts with the spotted basal portion.

Red grouper. This grouper has one

of most variable color patterns among

fishes. The body generally is

uniformly brownish red with a lighter

ventral coloration and a transient

pattern of whitish spots. While inactive it may have a banded pattern

that matches its surroundings and is

similar to that of the Nassau grouper.

Nassau grouper. In this species the

body around color can be tawny to

pi&i& and red, with five dark vertical bars; the third and fourth bars

divide above the lateral line and

their adjacent branches join to form a

w-shaped mark. The Nassau and red

groupers are somewhat similar in general appearance, but can be distinguished because the Nassau grouper has

a black saddle on top of the caudal

peduncle, distinctive black spots

below and behind the eyes, and a

characteristic "tuning fork" shaped

mark on top of the head, all of which

are lacking in the red grouper.

Moe (1963) reported the collection of a xanthic specimen of red

grouper off Fort Myers, Florida, that

agreed in general morphology and

meristic counts with published

descriptions of the red grouper, and

appeared normal in all respects except

coloration. Overall body coloration

was a brilliant orange-yellow on the

dorsal and upper lateral surfaces,

shading into reddish-pink on the lower

lateral and ventral surfaces.

Size

Black groupers can reach total

lengths greater than 1 m and weights

over 65 kg, but most of those caught

range up to 70 cm TL and about 26 kg

in weight. During the 1977 and 1978

Metropolitan South Florida Fishing

Tournaments, several black groupers of

22 to 36 kg were caught in the Florida

Keys (Official Entry Forms, unpubl.).

Male red groupers are

up to 72 cm TL and

reported

females up to 70 cm

TL; most fish in commercial catches

range from 45 to 70 cm TL. Nassau

groupers can grow to about 1.2 m TL

5
and 20 kg, but most of those marketed

weigh 2 to 10 kg (Bohlke and Chaplin

1968; Fischer 1978).

LIFE HISTORY

Habitat

Like most species of groupers,

the black, red, and Nassau groupers

are secretive, occupying caves, ledges

and crevices on reefs and shipwrecks

(Smith 1961); the larger fish generally occupy the lower part of the depth

range (Thompson and Munro 1978).

The black grouper lives over

rocky bottoms, coral reefs, and dropoff walls; fish longer than about 65

cm TL are generally restricted to

water deeper than 20 m and small young

of the year are usually in shallower

water (Fischer 1978). During monthly

collections on a shallow grass flat on

Matecumbe Key, Florida, Springer and

McErlean (1962) reported collecting

what appeared to be two young black

groupers (20 and 24 mm in standard

length, SL). In Venezuela, Cervigon

(1966) reported that black groupers up

to 350 cm SL frequently occur over

muddy bottoms of mangrove-fringed

coastal lagoons, and that fish longer

than 65 cm SL are generally at depths

greater than about 20 m. Moe (1966)

reported this species to a depth of

151 m, at bottom temperatures of 16 to

28 OC (mean 20 oC).

The red grouper occurs mainly

over rocky bottoms, but also lives

over muddy bottoms (Cervigon

Fischer 1978).

1966;

In the eastern Gulf of

Mexico it has been reported as occurring only over rocky reef bottoms at

depths of 3 to 122 m, frequently occupying crevices, ledges, and caverns

in limestone reefs (Moe 1969). Juveniles may be widely dispersed over

hard bottoms at depths of at least 37

m, being extremely cryptic and thus

relatively invulnerable to most collecting gear. Fish 1 to 6 years old

(under 50 cm SL) commonly inhabit

nearshore reefs (Beaumariage and

Bullock 1976). It has been reported

to depths of 189 m, at bottom temperatures from 15 to 30 oC, but most are

collected at 19 to 25 oC (Roe 1976).

The Nassau grouper is generally

found near high-relief coral reefs and

rocky bottoms. from the shoreline to a

depth of at least 90 m (Fischer 1978;

Bannerot 1984). Cervigon (1966) reported that large fish occur at depths

greater than about 50 m. In Bermuda,

Bardach (1958) reported a marked size

segregation: fish less than 40 cm SL

were on inshore banks and larger ones

were on offshore banks. The species

may be resistant to changing salinities: in the old New York Aquarium,

several Nassau groupers reportedly

lived for several years in water that

occasionally became almost fresh and

was rather polluted (Townsend 1905).

Sex Reversal and Spawning

Most serranids are protogynous

hermaphrodites (fish are first females

and then change into males), and the

sex of an individual cannot be

accurately determined unless it is

ripe. There is d

tion by sex in b

pers, but not

(Bannerot 1984).

Smith (1959

cal and field

fferential distribuack and Nassau grouin red groupers

reported histologiobservations that

strongly suggested protogynous hermaphroditism in black groupers. The

black grouper has been reported to

spawn off Puerto Rico in February

(Erdman 1956), and has been observed

in spawning condition in the Campeche

Bank area in July and August (Smith

1961). Off Bermuda, spawning extends

from May to early August; females may

weigh up to 22.7 kg, but larger fish

are usually males (Smith 1971).

Red groupers change from female

to male between the ages of 5 and 10

years, at a rate of about 15% annually* The transition occurs at any

length greater than about 275 nnn SL,

d ’

6

&
but is most common after 500 mm SL is

reached. Females are mature at 4-6

years, but reach their greatest reproductive potential at 8-12 years of

age. Males do not compose more than

10% of a year class until after age 9

(over 500 mm SL), and the sex ratio is

not equal until about age 15, or 625

mm SL (Moe 1969; Beaumariage and

Bullock 1976). Males are reproductively significant in the population in

age groups 10 years old and older

(Beaumariage and Bullock 1976).

Female red groupers usually do

not spawn until they are about 450 mm

SL. Off the west coast of Florida,

spawning peaks in April and May in

waters 20-90 m deep and at water temperatures between I9 and 21 OC. Gonada1 activity has been observed as

early as January, and culminates in

spawning in late spring. The only

environmental factor that correlates

significantly with gonadal development

is photoperiod (Moe 1969).

The Nassau grouper changes from

female to male at a length between 300

and 800 mm, which presumably indicates

that an individual fish can spend more

than one spawning season as a functional female. Off Bermuda, 3.2 kg

fish are just reaching maturity

(Bardach and Menzel 1957). Spawning

near Bermuda occurs between early May

and mid-August (Smith 1971). In the

Caribbean, ripe fish were collected

between February and May, and sexually

inactive ones were collected in

October and November (Munro et al.

1973). Nassau groupers form spawning

aggregations of as many as 100,000

fish off Bimini, Bahamas, for 1 week

during the full moon in January (Smith

1972). Similar aggregations have been

reported by Miller (1984) in Belize;

by Bannerot (1984) off the southern

Berry Islands, Bahamas (both also

during the full moon in January); by

Burnett-Herkes (1975) off Bermuda; and

by Olsen and LaPlace (1978) in the

Virgin Islands. Nassau groupers have

been observed to spawn at night in

tanks, with no formation of mating

pairs (Manday and Fernandez 1966).

Eggs and Larvae

Groupers produce planktonic eggs

that are fertilized externally, and

predators and currents affect their

survival (Smith 1961; Colin 1982).

Fecundity has been estimated to be

over 5 million eggs at 805 mm SL in

black groupers; over 785,000 at 445 mm

SL for Nassau groupers (Smith 1961);

and. 1,469,OOO (range 312,000-

5,735,700) for red groupers 495 to 667

mm SL (Moe 1969). Calculated individual egg weights ranged from 12 to 22

mg.

Moe (1969) described the eggs of

the red grouper as having no filaments

or other appendages, containing an oil

droplet, and being less than 1 mm in

diameter. Manday and Fernandez (1966)

reported that the eggs of the Nassau

grouper were between 0.9 and 1.024 mm

in diameter, and described the species' embryological and larval development until the yolk sac was

absorbed.

The distribution of grouper larvae is poorly known. Larval red grouper probably leave the plankton and

become benthic at about 20-25 mm SL

(Beaumariage and Bullock 1976).

Johnson and Keener (1984) reported that the different patterns of

serration on the long second dorsal

and pelvic fin spines that characterize grouper larvae are consistent

features that can be used to identify

larvae of certain species groups.

These patterns can identify larvae of

some grouper species at lengths as

short as 5-6 mm, in lieu of ranges of

meristic counts, where considerable

overlap exists among many American

grouper species. In the genus

Mycteroperca, which includes the black

grouper, species separation of the

larvae based on spinelet morphology is

not possible. However, larvae of the

7
genus can be distinguished from larvae

of other grouper genera by the following characteristics: "higher number

of anal soft rays (10-13, usually 11,

vs. T-10, usually 8 or 9), characteristic spine morphology...and presence

of a cleithral pigment spot." Larvae

of the red grouoer, the red hind

(Epinephelus guttatus), and the

speckled hind (E. drummondhayi) can be

separated from larvae of other

epinepheline species with the exception of the mutton hamlet (E., afer),

but cannot be distinguis?ied among

themselves because they have similar

body form, spinelet morphology and

relative spine length. Larvae of the

Nassau grouper and the rock hind (&.

adscensionis) can be separated from

all other western Atlantic grouper

larvae by their fin ray counts, with

the exception of the jewfish (E.

itajara), which has a particular spine

morphology.

The function of the spinelets is

not known. According to Johnson and

Keener (1984) they may be involved in

interspecific recognition, but this is

not likely because the spinelets are

relatively transparent and lack pigmentation. The purpose of these structures may be to repel or hook

attacking predators.

Juveniles

Juveniles of the three species of

groupers treated here are commonly

found in inshore seagrass beds in

south Florida, and are often caught in

shallower waters than those occupied

by adults. Juvenile red groupers do

not wander far away from their reefs

of residence and are scattered in low

densities over hard bottoms in water

at least 36 m deep, where they are

difficult to collect due to their

cryptic behavior (Moe 1969). Juvenile

red groupers are "exact miniatures" of

the adults in form and color, and they

were often collected in seagrass beds

in water shallower than 15 m (Smith

'1971). Small Nassau groupers are also

common in seagrass beds (Randall

1968).

Movement and Migration

Most grouper species apparently

migrate vertically as they grow, the

larger fish living at progressively

greater depths. However, adult groupers may stay in the vicinity of specific reefs for long periods. Black

groupers tagged and released in inshore reefs in the Florida Keys displayed strong home-reef specificity,

and "even a hurricane failed to disrupt their residence" (Beaumariage and

Bullock 1976).

Using evidence from tag and recapture studies of red groupers, Moe

(1969) summarized their offshore movement. During their early years they

remain in shallow water, usually 3 to

18 m deep. Later, at about 400-450 mm

SL and 4-6 years, they leave the nearshore reef environment and move to

depths greater than 36 m. Evidence

supporting the tagging and recovery

data was given by size and age

analyses of onshore and offshore

fishes. The offshore movement seems to

coincide with the onset of sexual

maturity. Commercial fishermen have

reported seasonal movement of adults

in offshore waters 27 to 91 m deep;

some schooling or group movement among

adults has been evidenced by commercial catches of tagged individuals

(Source Document for the SnapperGrouper Fishery of the South Atlantic

Region 1983). Although no alongshore

migrational patterns were noted, a few

individual red groupers moved 28 to 72

km from tagging locations.

Beaumariage and Bullock (1976)

reported that tagged Nassau groupers

in the Florida Keys, transported to

other nearby reefs, promptly returned

to their original reef, showing a

"strong home-reef specificity."

Randall (1962) reported that tagged

Nassau groupers in the Virgin Islands

moved at most about 820 m from the

site of release. From results of

8
tagging studies,

(1962)

Springer and McErlean

reported that the Nassau

grouper tends to become established in

an area.

Nassau groupers have been reported to form enormous spawning aggregations and undergo spawning migrations.

A large aggregation of Nassau groupers

was observed in the Bahamas by Smith

(1972). The fish congregated as a

school of perhaps as many as 100,000

in water about 27 m deep. Analyses of

gonad samples indicated that fish of

both sexes were present and ripe. Some

individuals in the aggregation displayed an unusual color pattern; they

were dark above and light below and

their characteristic head markings

were reversed. Similar spawning aggregations were observed by Bannerot

(1984), Miller (1984), and others.

GROWTH CHARACTERISTICS

Mean back-calculated total lengths

for black groupers aged 1, 5, 10, and

14 years were 260, 664, 975, and 1,110

mm (Manooch and Mason 1987). Growth in

length was most rapid for the first 3

to 4 years, and then gradually slowed.

The rate of growth of male and

female red groupers is believed to be

similar, though males reach a larger

ultimate size than females. Their

effective fishable life span is about

17 years, but life span may reach 30

years (Moe 1969; Beaumariage and

Bullock 1976). Data from the Schlitz

tagging program indicated that the red

grouper grows at about 5.8 to 10.3 mm

per month (Moe 1966, 1967). For Nassau

groupers in the Virgin Islands, and

based on results from tagging studies,

Randall (1962) reported the following

mean growth rates: fish 175-250 mm TL

grow about 4.55 mm/month; 251-325 mm TL

about 3.5 mm/month; and 326-451 mm TL

about 1.92 mm/month. Table 1 presents

theoretical growth parameters for red

and Nassau groupers.

FISHERIES

Recreational and Commercial Fisheries

Estimated grouper landings and

fishing effort from 1983 to 1986 by

U.S. sport fishermen are sown in Table

2. Recreational grouper fishermen use

a variety of boats in pursuit of their

sport. Boats are a necessity since

grouper catches from shore and dock

fishing are rare. Boats used vary from

3.7 m long, privately-owned skiffs with

outboard motors to "party boats," also

known as head boats, up to 26 m long

and powered by diesel engines. Party

boats take groups of fishermen to

fishing reefs for a fee. Boat numbers

have increased greatly recently.

Conventional types of fishing rods and

reels are used routinely, and occasionally electric reels are used. Minor

catches are made by spear fishermen,

who most often use scuba gear. Fish

traps were used sporadically in Florida

beginning in 1919 (Schroeder 1924).

They were prohibited in Florida State

waters in 1980, but are still legal to

use in all Federal waters.

Since 1948, commercial grouper

landings have been significantly larger

in Florida than in several other States

combined (Figure 3). Allen and Tashiro

(1976) estimated that about 96% of

commercial grouper catches were made

with handlines. Incidental catches

were made in shrimp trawls, spiny

lobster traps, fish pots, haul seines,

trammel and gill nets, and longlines.

Longlines became a major gear (in terms

of pounds landed) in the gulf grouper

fishery in the early and mid-1980's and

accounted for the increase in

commercial landings then (Goodyear

1988). Vessels are usually 7.9 to 24 m

long, and frequently use a small

steadying sail while fishing over

reefs. Some boats are multi-purpose,

used in shrimp and lobster fisheries

when not used for groupers.

Additional information on the reef

fish fisheries in the gulf is provided

by Waters (unpubl.). Fishermen, like

9
Table 1. Yield-per-recruit parameters for black, red, and Nassau groupers. N.D.

= no data available; t - age at length of first capture; t, = age at length 0; K

von Bertalanffy growth coefficient; Z = total mortality; La, = length at infinity;

;L = total length; SL = standard length.

Lengthweight

Species t, t, K Z L 00 relationship Area Source

Black 5-7 -0.927 0.116 0.49-053 1352 W=5.55~10-~ TL3.14' Mostly Manooch and

;;$,, Mason (1987)

Red 1 -0.449 0.179 0.322 672 W=4.34~10-~ SL2.g3 West Moe (1969)

Florida

Nassau 4 -0.488 0.185 N.D. 974 w=o.1393 SL3." u s

Virgin

Olsen &

LaPlace

Islands (1978)

the fishing vessels they use, may work

only part time fishing for grouper,

often working in other fisheries or

jobs.

Manooch and Mason (1987) present a

yield-per-recruit fishery model for

black grouper.

Marketinq

Data from a telephone survey

reported by Cato and Prochaska (1976)

showed that Gulf of Mexico groupers

were usually sold by fish dealers fresh

and iced (82% by volume); 16% were

filleted and only about 2% were frozen

whole. The principal markets are in

the Southeastern United States,

relatively close to the important

fishing grounds, thereby permitting

shipments of a fresh, iced product.

Most dealers sold their fish to other

wholesalers (52%); about 35%T;En\et;

retail markets or agents.

were sold to New York agents.

Groupers probably do not contribute

substantially to the export market;

perhaps only 1% by volume of both

groupers and snappers are exported.

The strong import market for groupers,

which are imported into south Florida

from throughout the Caribbean and from

as far away as Brazil, implies a weak

export market.

The minimum size for any species

(including species such as scamp and

gag) is 18 inches total length. New

gear restrictions prohibit use of

longline nets and stab nets (also

called sink nets) off the Atlantic

coast of Florida.

Artificial Reefs

Artificial reef programs designed

to improve recreational fishing in the

ocean have become popular in recent

years; most activity in United States

waters has been in Florida. Artificial

reefs consisting of a wide variety of

materials, ranging from abandoned

vessels to designed and prefabricated

structures and even junk (debris, large

appliances, etc.), have been placed on

the coasts of Florida (greatest

activity has been in Dade County).

10
Table 2.

1983-86.'

Recreational statistics (catch and effort) for groupers in Florida,

Year

Florida (east coast) Florida (west coast)

Number of Effort (number Number of Effort (number

fish caught of trips)' fish caught of trips)2

1983 182,000 7,793,ooo 2,189,OOO 10,224,OOO

1984 187,000 9,891,OOO 2,429,ooo 11,451,ooo

1985 355,000 12,493,ooo 2,851,OOO 13,372,OOO

1986 87,000 10,298,OOO 2,196,OOO 13,346,OOO

'From National Marine Fisheries Service

2Trips for all sport fish.

(1984-87).

A controversy exists as to the best

depths for artificial reefs. Among the

recreational fishermen, there seems to

be a consensus favoring water about

73 m deep, where diving and spearfishing are discouraged and reefs tend

to attract large fish. The long-term

effect of artificial reef placement

would seem to be that recreational

catches of groupers would improve,

perhaps in some proportion to the

numbers and kinds of artificial reefs

installed (Cardozo and Hirsch 1985).

It is not known if artificial reefs

increase fish populations or just

redistribute them (Bohnsack, in press).

Contaminants

Red grouper from the Southeastern

United States had an average of 0.008

ppm DDT and undetectable levels of

PCB's. Levels for black grouper were

0.009 ppm DDT and up to 0.059 ppm PCB's

compared to other fish, low levels of

these contaminants are correlated with

low lipid content (Stout 1980).

ECOLOGICAL ROLE

Food and Feedina Habits

Groupers are unspecialized and

opportunistic carnivores, feeding on a

variety of fishes and crustaceans

(Thompson and Munro 1978) during the

day and at night. Generally, feeding

is most active at dawn and dusk

(Randall 1967). Foods include

cephalopods, crustaceans and other

invertebrates, and fishes. By opening

the mouth and rapidly dilating the gill

covers to draw in water, groupers can

generally engulf prey whole (Bardach et

al. 1958). The usual collection method

(baited handlines) may bias results of

food studies in groupers because these

fish frequently regurgitate while being

brought to the surface and may also

contain fishing bait (Randall 1967; Moe

1969). Spearfishing has been regarded

as the collection method most likely to

assure unbiased evaluations of stomach

contents (Randall 1967).

Black groupers are less intimately

associated with the bottom, and have

more slender bodies and better

developed canine teeth than do the red

or the Nassau groupers, suggesting a

predominately piscivorous diet (Randall

1967). Reported stomach contents

include clupeoid fishes (Cervigon

1966), grunts, and cornet-fish (Randall

1967). Both black and red groupers

have been reported to prey on pink

shrimp, Penaeus duorarum (Costello and

Allen 1970).

1 1
6-J l5

z 14

I

1 Florida.

2 13 c) Alabama.

Q 12 q Other states.

1964 1969 1974 1979

Year

Figure 3. U.S. grouper landings (primarily dressed weights) by States.

"Other States" includes North Carolina, South Carolina, Georgia, Alabama,

Mississippi,Louisiana, and Texas. From Allen and Tashiro (1976).

Off the Dry Tortugas, Florida, red

groupers were reported to feed on

lutjanid and sparid fishes (Gudger

1929) and on various other fishes,

octopuses, shrimps, stomatopods, and

spiny lobsters (Longley and Hildebrand

1941). Randall (1967) reported

unidentified crustaceans, crabs, and

fishes in the stomachs of two specimens

collected off the Virgin Islands. In

the stomachs of red groupers collected

by commercial and sport fishermen off

the Florida west coast, Moe (1969)

found small fish of several species,

octopuses and squids, crabs

(particularly Portunus spp. and Calapoa

SPP.), panulirid and scyllarid

lobsters, shrimps, and unidentified

crustaceans. Moe (1969) also indicated

that red groupers seemed to feed more

on invertebrates, particularly

crustaceans, than on fish, but that

feeding habits may change in large

groupers, which probably consume a

greater proportion of fish. Longley

and Hildebrand (1941) reported no

difference in feeding habits of red

groupers between day or night, but Moe

(1969)indicated that fishermen on the

west coast of Florida generally made

their better catches during daylight.

Cuban commercial boats fishing in the

Campeche Bank area reported catching

more and larger red groupers at night

than during the day (Zupanovich and

Gonzalez 1975).

Randall (1965) reported that the

stomach contents of 150 Nassau groupers

collected during daylight off the

Virgin Islands and Puerto Rico

consisted of about 53% fish, 39%

crustaceans, 5% cephalopods, 2%

gastropods, and 2% pelecypods. The

most abundant families of fish in the

diet, listed in order of occurrence,

were parrot-fishes (Scaridae), wrasses

(Labridae), damselfishes

(Pomacentridae), squirrelfishes

(Holocentridae), snappers (Lutjanidae),

and grunts (Haemulidae). Crustaceans

dominated the diet of smaller fish,

whereas fish predominated in the

stomach contents of larger Nassau

groupers (over 300 mm SL). Crustaceans

included crabs (Majidae, Portunidae,

Calappidae, Porcellanidae, and

Xanthidae), stomatopods (Squillidae),

hermit crabs (Paguridae), panulirid

lobsters, and caridean and penaeid

12
shrimp. Mollusks included octopuses and

squids, arks (Arcoida), and queen

conchs (Strombus aiqas),. Randall

(1964) suggested that despite their

lacking the dentition to cruSh the

shells, Nassau groupers may consume

queen conchs after other predators have

made soft parts available. In Bahamian

waters, however, Nassau groupers have

been observed forcefully pulling conchs

out of their shells (B. Rose,

commercial fishernan, Freeport, Bahamas;

pers. comm.) Moray eels (Muraenidae)

have been reported in stomachs of

Nassau groupers collected in the Virgin

Islands and Puerto Rico (Randall 1965)

and Jamaica (Thompson and Munro 1978).

Cervigon (1966) reported that Nassau

groupers in Venezuela fed mostly on

crustaceans.

Comoetitors and Predators

Interspecific competition for food

and shelter among groupers is likely

because of the overlap in habitat,

distribution, size, and food habits

(Thompson and Munro 1978). Adults of

most grouper species occupy high levels

in the complex trophic webs of reef

communities. These webs include many

species of generalized, opportunistic

predators (Bohnsack 1982; Bannerot

1984), including groupers, which can be

assumed to constantly compete with, and

prey upon, similar species. Other

competitors for food probably include

various species of jacks (Carangidae),

snappers (Lutjanidae), barracudas

(Sphyraenidae), and sharks

(Carcharhinidae, Sphyrinidae).

According to Thompson and Munro

(1978) "nothing is known of the causes

of natural mortality in groupers."

Small, cryptic individuals probably

fall prey to fishes such as moray eels

(Muraenidae), which can attack them in

their hiding places. Large groupers of

several species are also reported to

readily feed on smaller groupers (Smith

1961).

mentioned

ThtOhmepson and Munro (I978)

lack of published

information on grouper predators, and

suggested that groupers are probably

preyed upon by sharks and other large

fishes. Large groupers are probably

preyed upon only by certain shark

species. Only two species of sharks

were reported by Compagno (1984) to

prey on groupers: the sandbar shark

(Carcharhinus Dlumbeus) and the great

hammerhead (SDhvrna mokarran), but

other species of sharks undoubtedly

feed on groupers.

Parasites and Diseases

Black, red, and Nassau groupers are

hosts for a number of parasites (Table

3). The effect of these parasites on

the health of their hosts is unknown.

Groupers from the Tortugas Islands,

Florida, were hosts for several species

of digenetic trematodes (Manter 1947).

Overstreet (1968), who surveyed species

of fish in Biscayne Bay,Florida,reported digenetic trematodes from black

and Nassau groupers. In waters off

Campeche, Mexico, Fajer et al. (1979)

found larval cestodes and nematodes in

red groupers, and noted that males

tended to have more parasites than

females. They also observed that the

larger groupers (specimens that ranged

from 320 to 500 mm TL) tended to be

more heavily parasitized than smaller

ones--an observation that has been made

for a wide range of host species.

Observations on grouper parasites

were made by Thompson and Munro (1978)

from specimens caught around Jamaica.

Parasites were not identified and only

general groups were reported. In

Nassau groupers, they found parasitic

isopods in the nostrils, encysted

larval tapeworms commonly in the

viscera, and nematodes in the ovaries.

They observed, "Heavy infection by this

(nematode) parasite can drastically

reduce the number of eggs produced by

an individual." The gonads of male

fish were not infected.

The only report on grouper diseases

that we found in the literature was an

account by Moe (1969) of a large

skeletal tumor removed from a red

grouper. The fish was caught about

13
Table 3. Parasite species in black, red, and Nassau groupers.

Species of

parasite

Location Geographic

in host location Source

BLACK GROUPER

Digenetic trematode

Lecithochirium microstomum

Lecithochirium parvum

Postporus epinepheli

Prosorhynchus pacificus

RED GROUPER

Digenetic trematodea

Helicometra torta

Lepidapedon levenseni

Stephanostomum dentatum

Stomach II

Intestine

Intestine &

pyloric caeca

N.D.

Intestine &

caecum

Intestine

Cestoda

Callotetrarhynchus

(larvae-encysted)

sp. Muscles

Digestive tract

Liver

Abdominal cavity

Gonads

Biscayne Bay, Fl Overstreet (1968) II II

II II

II II

Tortugas, Fl

II

II

Manter (1947)

II

II

Campeche Bank Fajer et al.

(Mexico) (1979)

Digestive tract

Liver

Gonads

Nematoda

Anasakis

(larvae-encysted)

NASSAU GROUPER

Digenetic trematode

Helicometra torta N.D. Tortugas, Fl Manter (1947)

II Pyloric caeca Biscayne Bay, Fl Overstreet (1968)

Lecithochirium microstomum Stomach Biscayne Bay, Fl Overstreet (1968)

Lecithochirium parvum II Tortugas, Fl Manter (1947)

Sterrhurus musculus II Biscayne Bay, Fl Overstreet (1968)

aC.L. Smith (1961) reports

based on a report by Manter

Manter's report.

0 istho orus epinephely occurring in red grouper, .7-b?--No record of this trematode was found in

14
43 km southwest of Venice, Florida, in

May 1967. No information on length,

age, or sex of the fish was recorded.

Otolith abnormal growths have been

reported in red groupers by Moe (1969)

and in Nassau groupers by Thompson and

Munro (1978).

Ciauatera

Ciguatera is a toxin transmitted by

hundreds of species of tropical fishes.

It causes gastrointestinal,

cardiovascular, and neurological

disturbances in persons who eat toxic

g:ih, resulting in prolonged disability

long and expensive recovery

periods. Several species of Western

Atlantic groupers, including the back

and the red groupers, are routinely

reported to be ciguatoxic, whereas

others such as the Nassau grouper seem

to be uniformly nontoxic, throughout

their range. In south Florida

ciguatera is endemic, and of all

grouper species the black grouper is

the most often implicated; this species

is considered toxic throughout the

Caribbean, except near St. Croix, U.S.

Virgin Islands (de Sylva and Higman

1980; Poli 1982).

The occurrence of the toxin is

usually restricted to large predatory

species associated with reefs. The

larger older fish are generally more

toxic than small younger fish because

the old fish have had time to

accumulate the toxin. Even within

certain species the distribution of

toxic fish is not geographically

continuous, occurring in some areas and

not in others (Olsen et al. 1984). In

a study based on interviews (Poli

1982), "groupers" were responsible for

the largest number of recorded cases of

ciguatera intoxication in Florida

during two periods, "pre-1978" and

1978-1980. Many times the species

responsible were not identified due to

the processing procedures used for

marketing the fish.

Behavior

Groupers can generate sound in

different ways, particularly by

vibrating their thin-walled swim

bladder by sudden contraction of axial

muscles, such as the single pair of

bilateral muscles behind the opercles.

Various sounds have been attributed to

all three species: low-frequency,

high-amplitude "mooing-like" sounds,

deep booms, and sustained rumbles,

vibrant grunts (single or in rapid

series), and long noises from grinding

of teeth when fighting for food. Sound

may have many functions, including

warning, intimidation, orientation, and

recognition (Hazlett and Winn 1962;

Fish and Mowbray 1970). Nassau

groupers are diurnal or crepuscular in

their movements (Collette and Talbot

1972) and do not usually move far from

cover (Starck and Davis 1966). They

are often wary of divers and quickly

retreat into hiding places, producing

a series of grunting sounds (Hazlett

and Winn 1962). Black and Nassau

groupers were reportedly attracted by

irregularly pulsed signals during

experiments designed to attract and

film sharks off Bimini, Bahamas. The

groupers approached slowly, came to

rest in front of the sound projector,

and slowly moved away after lo-30

seconds (Myrberg et al. 1969). Randall

(1962) reported that Nassau groupers

have a tendency to re-enter fish traps;

some tagged fish were recaptured

several times, presumably re-entering

to feed on other fish in the traps.

Nassau groupers are often involved

i n symbiotic cleaning behavior at

"cleaning 'stations" on coral reefs.

Species most often reported involved

include several species of gobies

(Gobiosoma spp.) and shrimp that mainly

remove gnathiid isopods from the

bodies, fins, gill chambers and mouths

of groupers (Bohlke and McCosker 1973;

Darcy et al. 1974; Sargent and

Wagenbach 1975).

15

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17

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2 1
REPORT DOCUMENTATION I.. RE”)RT. NO.

PAGE Biological Report 82(11.1101*

/ 2.

1. T#tle and Subtitle

Species Profiles: Life Histories and Environmental Requirements

of Coastal Fishes and Invertebrates (South Florida)--Black, Red,

and Nassau Groupers

1. Author(s)

Darryl E. Jory and Edwin S. Iversen

$. Pwformin6 Orgrnizatton Name and Address

11. ContrwXfC) or Grant(G) No.

U.S. Department of the Interior U.S. Army Corps of Engineers 13. TYP* of Rwort & P*rmd coveme

Fish and Wildlife Service Waterways Experiment Station

National Wetlands Research Center P.O. Box 631

Washington, DC 20240 Vicksburg, MS 39180

F

14.

15. Supplementary Notes

- -

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

16. Abstract (Limat: 200 wards)

Black, red, and Nassau groupers (Mycteroperca bonaci, Epinephelus morio, and g.

striatus, respectively) are widely distributed on rocky bottoms and reefs along the

south Florida coast. They are the most valuable marine finfish group in Florida,

comprising about 25% of the total value of landings in 1984. The three species can be

distinguished by morphometric, meristic, and body color characteristics. Younger fish

are typically found in shallow, inshore grass beds, and larger, older fish are generally restricted to deep waters. The three species are protogynous hermaphrodites.

Sexual transition can occur at any length over about 300 mm SL. An offshore movement

apparently coincides with the onset of sexual maturity. Spawning aggregations have

been observed throughout the year, but occur mostly between late spring and early

summer. Fecundity estimates range from about 800,000 to over 5,000,OOO eggs per

female. Both the eggs and the larvae are planktonic. Their early life history is

poorly known. Larvae probably leave the plankton and become benthic at around 20-30 mm

SL. Growth rates range from about 2 to 10 mm/month. The three species are

unspecialized carnivores, feeding on a variety of fishes, crustaceans, and mollusks.

Interspecific competition for food and shelter may be common because of the overlap in

distribution, habitat, size, and food habits. For the three species, a number of

predators and parasites have been reported. Both the black and red groupers have been

implicated in ciguatera poisonings in south Florida.

1 17. Document Analysis a. Dc*cripton

Marine fishes Life cycles

Fisheries

~

Growth

Feed habits Competition

Parasites

Spawning

Predators

I

b Idsntlfieo/Opcn~Endcd Terms

Black Grouper (Mycteroperca bonaci)

Red Grouper (Epinephelus morio)

Nassau Grouper (Epinephelus striatus)

c. COSATI Field/Group

lb Avallrbility Statement

Unlimited

19. Securaty Class (Thns Retort) 2,. NO. of Pa6es

Unclassified 21 - -

20. Security Class CThls Page) 22. Prlcc

Unclassifed

(See ANSI-239.16) OPTIONAL FORM 272 M-71

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