Download Priority Effects in the Recruitment of Juvenile Coral Reef Fishes

Ecology, 64(6), 1983, pp. 1508-1513
<1983 by the Ecological Society of America
PRIORITY EFFECTS IN THE RECRUITMENT
OF JUVENILE CORAL REEF FISHES'
MYRA J. SHULMAN2
Department of Zoology, University of Washington, Seattle, Washington 98195 USA
JOHN C. OGDEN
West Indies Laboratory, Teague Bay, St. Croix, United States Virgin Islands 00820
JOHN
P.
EBERSOLE
Department of Biology, University of Massachusetts, Boston, Massachusetts 02125 USA
WILLIAM
N.
MCFARLAND
Section of Ecology and Systematics, Cornell University, Ithaca, New York 14853 USA
STEVEN
L.
MILLER
Department of Botany, University of Maine, Orono, Maine 04473 USA
NANCY
G.
WOLF
Section of Ecology and Systematics, Cornell University, Ithaca, New York 14853, USA
Abstract. Competing models of community structure in assemblages of coral reef fishes have
suggested that (1) these assemblages are structured by deterministic interactions between species, or
between species and resources, or (2) the composition of these assemblages are determined by highly
variable settlement from planktonic larvae. We examined interactions among newly recruited juvenile
fishes and between juvenile fishes and transplanted resident damselfish on artificial reefs in St. Croix,
United States Virgin Islands. Two kinds of priority effects occurred: (I) recruitment of three species
of settling juveniles significantly decreased in the presence of the territorial damselfish, and (2) prior
settlement of a juvenile predator lowered successful recruitment of two juvenile prey species. The
first effect increases determinism in the structure of coral reef fish assemblages, while the second
decreases their predictability.
Key words: artificial reefs; Caribbean; community structure; competition; coral reeffish assemblages; interference competition; predation; recruitment.
INTRODUCTION
Priority effects, in which the presence of one species
in a habitat decreases the probability of invasion by
another, have been hypothesized as important in
maintaining high regional species diversity and determining local species distributions (Levins and Culver
1971, Sutherland 1974, Connell and Slatyer 1977). One
species can lower the juvenile recruitment of another
in several possible ways. (I) Competition: adult and
subadult residents or settling juveniles can actively interfere with juvenile settlement through aggression (interference competition) or passively influence settlement by altering or preempting resources (exploitative
competition). (2) Predation: predatory subadult and
adult residents can decrease recruitment directly, by
preying on juveniles, or evolutionarily, by selecting for
settlement in areas not occupied by predators. Similarly, predatory juveniles settling first can prevent subsequent successful recruitment by prey species. These
I Manuscript received 3 November 1981; revised I September 1982' accepted 8 September 1982; final version received 15 October 1982.
2 Present address: Smithsonian Tropical Research
Institute, APO Miami 34002.
mechanisms are not mutually exclusive, and there are
examples of at least two of them operating together
(Sutherland 1974).
The existence of priority effects among coral reef
fishes would have significant implications for our understanding of the processes controlling these assemblages, a subject of much current controversy. The
controversy centers on whether fish assemblages are
structured by deterministic competitive and predatory
interactions or by stochastic recruitment of juveniles
into unpredictably available space (Sale 1977, 1980,
Smith 1978, Gladfelter et al. 1980, Anderson et al. 1981,
Ogden and Ebersole 1981). Priority effects between residents (adults and subadults) and juveniles, in which
residents control which species recruit into the local
habitat, suggest that assemblage composition is more
deterministic. Juvenile-juvenile priority effects, in
which order of settlement determines local species
composition, suggest an unpredictable assemblage
structure, strongly controlled by the vagaries of settlement of juveniles from the plankton. The importance of order of settlement in determining community
structure has been shown for sessile organisms in experimental space-limited systems (Sutherland 1974,
Sutherland and Karlson 1977) but has not yet been
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions
December 1983
RECRUITMENT OF JUVENILE CORAL REEF FISHES
documented for mobile species. In this study, we report the existence of two kinds of priority effects among
coral reef fish assemblages, a system involving mobile
species with strong behavioral interactions.
0
METHODS
0.
C,
The present study was conducted utilizing the facilities of the underwater habitat Hydrolab operated
by the National Oceanographic and Atmospheric
Administration on St. Croix, United States Virgin Islands. Hydrolab is located in Salt River Canyon, a 70100 m wide submarine canyon flanked by two coral
reef walls. Thirty artificial reefs, each consisting of 11
concrete construction blocks placed in a pyramid two
blocks wide and three blocks high, were built on the
sand/seagrass (Halophila decipiens) floor of the canyon at depths of 16-20 m. Ten replicate reefs were
constructed at each of three different times: 4 wk
("'old" reefs), 2 wk ("intermediate" reefs), and l/2 d
(''young' reefs) prior to the start of the 7-13 July 1979
Hydrolab mission. The reefs were placed in a grid pattern, with a minimum distance of 5 m between them.
Because of the small size and regular structure of these
reefs, all fish species could be seen with relative ease.
All the fishes on the artificial reefs were visually censused by two observers approximately twice weekly
before the mission and several times daily during the
mission. Weekly censuses were continued afterwards,
until the artificial reefs were destroyed by Hurricanes
David and Frederick in late August and early September 1979.
RESU LTS
Small artificial reefs, isolated by sand or seagrass
from natural reefs, attract a fish fauna similar to that
found on small isolated patch reefs (Talbot et al. 1978)
and similarly situated reefs constructed from coral
rubble (Shulman 1982). Compared to major reefs, all
these small habitats show higher rates of successful
recruitment by juveniles due to lower encounter rates
with reef-associated predators and the increased shelter available from the seagrasses and algae surrounding the habitat (Shulman 1982). In addition, certain
fishes, particularly those closely associated with particular coral habitats as adults, rarely recruit to isolated artificial reefs (Shulman 1982).
A total of 44 species colonized the artificial reefs;
26 of these were found on at least five reefs (Table 1).
The colonization curves (Fig. I) show that numbers of
individuals and species increase monotonically to an
apparent asymptote on young and intermediate reefs
but peak early and then decrease on the old reefs.
Each set of reefs was censused 12 d after construction. Using these 12-d censuses, comparisons among
the three sets of reefs could be made that were independent of the amount of time the reefs were available
for colonization. Grunts (Haemulidae) were excluded
from analyses involving total numbers of recruits be-
1509
4)5
0
z
25
4) 20.
/
CL
0
5-a
7 JUN
22 JUN
7 JUL
22 JUL
6 AUG
21 AUG
Date (1979)
FIG. 1. Colonization curves of species and individuals
(excludinggrunts)on the three sets of artificialreefs. Points
represent means of all 10 reefs in each set. (O = old reefs;
* = intermediatereefs; O = young reefs.)
cause they far outnumbered all other species combined; analyses that included grunts would reflect only
the trends in that family.
There are significant differences among the three
sets of reefs in both numbers of species and numbers
of individuals (excluding grunts) recruited in the first
12 d after each set was built (ANOVA, P < .001).
Young reefs were colonized by the fewest individuals
and species, and old reefs by the most. This may represent a seasonal decrease in number of juvenile fish
settling from the plankton (Munro et al. 1973), or it
may be due to a dilution effect, there being only 10
reefs present to which fish could recruit during the
12 d following construction of the old reefs, but 20 d
for the intermediate and 30 d for the young reefs.
The reefs were censused intensively during the 6 d
of the Hydrolab mission and censused again 6 d later.
Because the intervals between these censuses were
short, and therefore relatively few recruits could colonize and then disappear without being observed, the
data from this period were used to examine net recruitment rates and interspecific effects on recruitment
patterns.
At any given time, reefs that had been in the water
longer had higher resident fish populations (Fig. 1).
During the period of intensive censusing, net recruitment rates (changes in number of individuals or species
from one census to the next) to the three differentaged sets of reefs were compared. The youngest reefs,
with the lowest resident populations, showed the high-
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions
MYRA J. SHULMAN ET AL.
1510
TABLE
1.
Species colonizingthe reefs.
Number
of reefs
occupied
by
species
Species
2
10
Gv mnothorax moringa (spotted moray)
Clupeidae(herring)
7
3
8
8
I
Svnodus intermedius (sand diver)
Adiorvx coruscus (reef squirrelfish)
Mvripristis jacobus (blackbar soldierfish)
Diplectrum sp.
Epinephelus guttatus (red hind)
Serranus baldwini (lantern bass)
Serranus tabacarius (tobaccofish)
Serranus tigrinus (harlequinbass)
Rvpticus subbifrenatus (spotted soapfish)
Apogon binotatus (barred cardinalfish)
Apogon maculatus (flamefish)
Apogon townsendi (belted cardinalfish)
Phaeoptyx conklini (freckled cardinalfish)
Malacanthus plumieri (sand tilefish)
Lutjanus buccanella (blackfin snapper)
Lutjanus mahogoni (mahogany snapper)
Ocv'urus chrvsurus (yellowtail snapper)
Haemulon aurolineatum (tomtate)
Haemulon flav~olineatum (frenchgrunt)
Hoemulonmelanurum(cottonwick)
17
3
4
4
22
9
30
18
25
15
30
I
Equetus lanceolatus (jackknife fish)
Odontoscion dentex (reef croaker)
Equetus acuminatus (high-hat)
Pseudupeneus maculatus (spotted goatfish)
Pomacanthus paru (french angelfish)
Chaetodon sedentarius (reef butterflyfish)
Chromis cyaneus (blue chromis)
Eupomacentrus partitus (bicolor damselfish)
Eupomacentrus v'ariabilis (cocoa damselfish)
Halichoeres biv'ittatus (slippery dick)
Thalassoma bifasciatum (bluehead wrasse)
Callioni'mus bairdi (lancer dragonet)
Corvphopterus glaucofraenum (bridled goby)
Gnatholepis thompsoni (goldspot goby)
Acanthurus bahianus (ocean surgeon)
Acanthurus chirurgus (doctorfish)
Acanthurus coeruleus (blue tang)
Scorpaena plumieri (spotted scorpionfish)
Bothus lunatus (peacock flounder)
3
10
1I
2
3
16
I
4
4
14
8
14
24
25
9
4
7
13
Balistidae
I
24
1
Canthigaster rostrata (sharpnose puffer)
Sphoeroides spengleri (bandtail puffer)
est rate of net recruitment of individuals (excluding or
including grunts) and species, with intermediate and
old reefs having significantly lower net recruitment rates
(ANOVA, P < .05). This difference is reflected in the
colonization curves for both individuals and species,
which appear to reach an asymptote after I mo (Fig.
1).
There were also species-specific differences in net
recruitment to different-aged reefs; grunts (Haemulidae), tobaccofish
(Serranus
tabacarius),
cardinal fish
(Apogonidae), and sharpnose puffers (Canthigaster
rostrata) showed significantly higher net recruitment
to young, relatively unoccupied reefs, while no such
significant difference was found for mahogany snappers (Lutjanus
inahogoni),
blackfin snappers
(Lutja-
Ecology, Vol. 64, No. 6
nus buccanella), yellowtail snappers (Ocyurus chrysurus), high-hats (Equetus acuminatus),
and
surgeonfish (Acanthurus bahianus and A. chirurgus).
These results may reflect differences between species
in ability to settle and survive on occupied reefs, or
important differences in recruitment strategies.
To determine the effect of a resident on recruitment,
one adult beaugregory (Eupomacentrus leucostictus),
a territorial damselfish, was transplanted onto each of
5 of the 10 replicates for the three sets of reefs on 7
July 1979. Successful recruitment of surgeonfishes
(Acanthurus bahianus and A. chirurgus) and reef butterflyfish (Chaetodon sedentarius) to reefs with beaugregories present was significantly lower relative to
reefs that lacked beaugregories (Tables 2 and 3).
The most common group of piscivorous fish recruiting to the reefs were snappers of the genus Lutjanus.
Juveniles of two species were present (mahogany, L.
mahogoni; blackfin, L. buccanella), the mahogany
snapper being more abundant. These snappers were
observed to prey on grunts and presumably attacked
other small juvenile fishes. Grunts showed a nonrandom recruitment pattern relative to the presence or
absence of Lutjanus during the first 12 d after the reefs
were built. Reefs on which snappers were present
showed a significant decrease (of 20%) in the numbers
of grunts subsequently recruited, while reefs that lacked
snappers recruited grunts in numbers slightly greater
than expected (x2 test, P < .001).
A similar result was obtained for the relationship
between snappers and high-hats (Equetus acuminatus). High-hats only settled on "full-moon reefs" (see
below); of these 20 reefs, 9 were colonized by highhats in the 12 d following construction. No juvenile
snappers were present on those 9 reefs before or during the settlement of high-hats; 6 of the remaining II
reefs were occupied by snappers, thus demonstrating
a statistically significant effect of snapper presence on
high-hat recruitment (binomial test, P < .05).
The timing of recruitment of planktonic larvae is
important if order of settlement affects species coexistence. Old reefs and young reefs were built within
3 d of full moons, and intermediate reefs within 3 d of
a new moon. The recruitment data from the first 7 d
after each set of reefs was built were used to determine
lunar periodicity in settlement. The 7-d period represents recruitment during one-quarter of a lunar cycle.
The numbers of fishes settling on old and young ("fullmoon") reefs were compared with those settling on
intermediate ("new-moon") reefs (Table 4). The data,
analyzed with the median test (P determined from
Fisher's exact test) showed significantly higher recruitment of mahogany snappers and Diplectrum sp.
to new-moon reefs as compared to full-moon reefs
(P < .05). The data are suggestive for greater settlement on new-moon reefs by blackfin snappers, doctorfish, and sharpnose puffers (P < . 10). Only one
species, the high-hat, showed significantly higher re-
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions
December 1983
TABLE
2.
RECRUITMENT OF JUVENILE CORAL REEF FISHES
1511
Effect of the presence of an adult beaugregory (Eupomacentrus leucostictus) on the recruitment of acanthurids.
Number of reefs on which:
Acanthurid settles and stays
more than one census
Beaugregory present
Beaugregory absent
No acanthurid stays
more than one census
Observed
Expected
Observed
Expected
4.5*
14.0
7.4
11.1
7.5*
4.0
4.6
6.9
18.5
2
11.5
= 4.94; df =
Number
of reefs
12t
18
30
1;P < .05
* On one reef, an acanthurid was present before the beaugregory transplant and remained on the reef for more than two
censuses. No more acanthurids arrived; the result therefore was split between the two categories.
t Three of the 15 beaugregories transplanted onto the reefs disappeared in the first few days.
cruitment to full-moon reefs (P < .05), settling only
on those reefs. There is also very strong evidence that
French grunts (Haemulon flaiolineatum) have semimonthly peaks in recruitment (W. N. McFarland, personal observation).
DiSCUSSION
Our study suggests that at least two kinds of priority
effects exist in juvenile coral reef fish recruitment. The
underlying mechanisms producing these two effects
are very different, as are their implications for community structure.
The presence of an adult resident (beaugregory) decreases the settlement of juveniles of three other
species. This territorial damselfish is extremely aggressive, particularly towards potential trophic competitors and egg predators (Eggs are laid in territories
of male beaugregories.) (Ebersole 1977). On our artificial reefs, the presence of the herbivorous beaugregories was associated with a significant decrease in
successful recruitment of surgeonfishes, a family of
herbivores. The presence of beaugregories was also
correlated with lowered recruitment of reef butterflyfish, a member of a family known to include fish eggs
in its diet (Birkeland and Neudecker 1981).
The second kind of priority effect occurs between
juvenile piscivores and their prey. The order of settlement is critical; if snappers settle on a reef first, the
numbers of successfully settling grunts and high-hats
are likely to be reduced. The reverse, however, is not
true; snappers do settle on reefs already occupied by
their prey. It is likely that the prey quickly outgrow
the size-range in which they are susceptible to predation by newly settling snappers, thus allowing coexistence of these species in areas where the prey settle first. This situation is analogous in mechanism and
effect to size-limited predation found in temperate intertidal and freshwater communities (Dodson 1970,
Paine 1976, Zaret 1980).
Our results are similar to those found in a number
of studies on sessile marine communities. Order of
settlement can determine the species composition in
fouling communities (Sutherland 1974, Sutherland and
Karlson 1977), and depending on the species identity
of the resident and the larvae, residents may prevent
or decrease larval recruitment (Sutherland and Karlson 1977, Sutherland 1978, Kay and Keough 1981).
The suppression of juvenile recruitment by residents
is in accordance with Connell and Slatyer's (1977) inhibition model of plant succession, a model which has
also been supported by research on succession in algal
3. Effect of the presence of adult beaugregories (Eupomacentrus leucostictus) on the recruitment of reef butterflyfish
(Chaetodon sedentarius).
TABLE
Number of reefs on which:
Reef butterfly arrives and
stays more than one census
Beaugregory present
Beaugregory absent
Observed
Expected
0.0
6.5t
2.6
3.9
6.5
No reef butterfly stays
more than one census
Expected
Observed
9.4
14.1
12.0
ll .5t
N umber
of reefs
12*
18
23.5
X2 = 5.53; df = 1; P < .025
* Three of the 15 beaugregories transplanted onto the reefs disappeared in the first few days.
t On one reef, a reef butterflyfish was present before the beaugregory transplant and remained on the reef for more than
two censuses. No new reef butterflyfish arrived, so the result was split between the two categories.
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions
1512
MYRA J. SHULMAN ET AL.
4. Comparison of recruitment to "new-moon" and
"full-moon" reefs the first 7 d after they were built.
TABLE
No. recruits per reef
New moon
(intermediate)
Full moon
(young + old) Median
test
Species
x
SD
X
SD
Mahogany snapper
Diplectrum sp.
Blackfin snapper
Doctorfish
Sharpnose puffer
High-hat
1.10
0.30
0.40
0.40
0.60
0.00
1.45
0.48
0.70
0.97
0.86
0.00
0.75
0.00
0.05
0.00
0.10
0.75
1.16
0.00
0.22
0.00
0.31
1.29
P
.006
.03
.09
.10
.06
.04
Ecology, Vol. 64, No. 6
ment into a habitat. The existence of priority effects
may have evolutionary significance by producing selection of potential prey for settlement before potential
predators.
It is probable that there exist other priority effects,
of varying degrees of subtlety, among juvenile coral
reef fish. Indeed, our results show that the higher the
density of the resident population, the lower the net
recruitment to a reef. It is likely that a large number
of intra- and interspecific interactions, of which the
ones elucidated in this study are examples, combine
to produce this result.
ACKNOWLEDGMENTS
communities (Sousa 1979) and terrestrial plant communities (see examples in Drury and Nisbet 1973,
Connell and Slatyer 1977).
The ecological implications of the two kinds of
priority effects found in our study are diametrically
opposed. If residents can successfully exclude certain
species from a habitat while allowing the recruitment
of others, fish assemblages become more predictable.
The composition of these assemblages can be stable if
residents allow their own species to settle while excluding others, "successional" if residents replace each
other in a transitive order, and cyclical if multispecies
priority effects are nontransitive (Jackson and Buss
1975, Buss and Jackson 1979). In contrast, the existence of juvenile-juvenile priority effects, in which an
unpredictable order of settlement of juveniles alone
determines community composition, introduces a stochastic element into the structure of fish assemblages
and increases their variability. This variability may be
prolonged by subsequent, deterministic resident-juvenile priority effects, depending on which juveniles
are permitted to recruit.
The present controversy about the structure of reef
fish assemblages centers on the degree of determinism
and stochasticity inherent in the processes that govern
this system. Data of the type presented here are critical to this issue and suggest the following conclusion:
in unoccupied habitat (created by storms, mortality,
changes in reef structure), the order of settlement, particularly of predators and prey, will partially determine
composition of the initial assemblage; in occupied
areas, residents are likely to determine which species
can possibly invade, but within this group of "permissible" invaders, order of settlement may determine
which will successfully recruit into the habitat.
The order of settlement of juveniles will be influenced by the lunar and seasonal periodicities in settlement patterns. Lunar cycles of settlement are reported
here and elsewhere; many studies have also show seasonality of spawning and settlement of reef fishes (Johannes 1978 and included references). The timing of
settlement of potential prey species relative to their
predators can be a crucial factor in successful recruit-
We thank the MannedUndersea Science and Technology
programof the National Oceanic and AtmosphericAdministrationfor financialsupportand use of the Hydrolabfacilities. The Hydrolabstaff, B. Walden, Dr. W. Shane, and R.
Catanack, and support divers, F. Pecora and M. Pacifico,
providedexcellent logisticalsupport.We thankthe manycolleagues who commented on the manuscript.This is contribution No. 74 from the West Indies Laboratory, Fairleigh
Dickinson University.
LITERATURE
CITED
Anderson, G. R. V., A. H. Ehrlich, P. R. Ehrlich, J. D.
Roughgarden, B. C. Russell, and F. H. Talbot. 1981. The
community structure of coral reef fishes. American Naturalist 117:476-495.
Birkeland, C., and S. Neudecker. 1981. Foraging behavior
of two Caribbean chaetodontids: Chaetodon capistratus
and C. aculeatus. Copeia 1981:169-178.
Buss, L. W., and J. B. C. Jackson. 1979. Competitive networks: nontransitive competitive relationships in cryptic
coral reef environments. American Naturalist 113:223-234.
Connell, J. H., and R. 0. Slatyer. 1977. Mechanisms of
succession in natural communities and their role in community stability and organization. American Naturalist 111:
1119-1144.
Dodson, S. I. 1970. Complementary feeding niches sustained by size-selective predation. Limnology and Oceanography 15:131-137.
Drury, W. B., and I. C. T. Nisbet. 1973. Succession. Journal of the Arnold Arboretum 54:331-368.
Ebersole, J. P. 1977. The adaptive significance of interspecific territoriality in the reef fish Eupomacentrus leucostictus. Ecology 58:914-920.
Gladfelter, W. B., J. C. Ogden, and E. H. Gladfelter. 1980.
Similarity and diversity among coral reef fish communities:
a comparison between tropical western Atlantic (Virgin Islands) and tropical central Pacific (Marshall Islands) patch
reefs. Ecology 61:1156-1168.
Jackson, J. B. C., and L. W. Buss. 1975. Allelopathy and
spatial competition among coral reef invertebrates. Proceedings of the National Academy of Sciences (USA) 72:
5160-5163.
Johannes, R. E. 1978. Reproductive strategies of coastal
marine fishes in the tropics. Environmental Biology of
Fishes 3:65-84.
Kay, A. M., and M. J. Keough. 1981. Occupation of patches
in the epifaunal communities of pier pilings and the bivalve
Pinna bicolor at Edithburgh, South Australia. Oecologia
(Berlin) 48:123-130.
Levins, R., and D. C. Culver. 1971. Regional coexistence
of species and coexistence between rare species. Proceedings of the National Academy of Sciences (USA) 68: 12461248.
Munro, J. L., V. C. Gaut, R. Thompson, and P. H. Reeson.
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions
December 1983
RECRUITMENT OF JUVENILE CORAL REEF FISHES
1973. The spawning seasons of Caribbean reef fish. Journal of Fish Biology 5:69-84.
Ogden, J. C., and J. P. Ebersole. 1981. Scale and community structure of coral reef fishes: a long-term study of
a large artificial reef. Marine Ecology Progress Series
4:97-103.
Paine, R. T. 1976. Size-limited predation: an observational
and comparative approach with the Mytilus-Pisaster interaction. Ecology 57:858-873.
Sale, P. F. 1977. Maintenance of high diversity of coral
reef fish communities. American Naturalist 111:337-359.
1980. The ecology of fishes on coral reefs. Oceanography Marine Biology Annual Review 18:367-421.
Shulman, M. J. 1982. The structure of coral reef fish assemblages: effects of resource limitation, predation, and
interference competition on recruitment patterns. Dissertation. University of Washington, Seattle, Washington,
USA.
1513
Smith, C. L. 1978. Coral reef fish communities: a compromise view. Environmental Biology of Fishes 3:108-128.
Sousa, W. P. 1979. Experimental investigations of disturbance and ecological succession in a rocky intertidal algal
community. Ecological Monographs 49:227-254.
Sutherland, J. P. 1974. Multiple stable points in natural
communities. American Naturalist 108:859-873.
1978. Functional roles of Schizoporella and Styela
in the fouling community at Beaufort, North Carolina.
Ecology 59:257-264.
Sutherland, J. P., and R. H. Karlson. 1977. Development
and stability of the fouling community at Beaufort, North
Carolina. Ecological Monographs 47:425-446.
Talbot, F. H., B. C. Russell, and G. Anderson. 1978. Coral
reef fish communities: unstable high-diversity systems?
Ecological Monographs 48:425-440.
Zaret, T. M. 1980. Predation and freshwater communities.
Yale University Press, New Haven, Connecticut, USA.
This content downloaded from 195.113.56.251 on Mon, 14 Apr 2014 18:21:02 PM
All use subject to JSTOR Terms and Conditions