Hard Substrata Community Patterns, 1-120 M, North Jamaica

Hard Substrata Community Patterns, 1-120 M, North Jamaica
Author(s): W. David Liddell and Sharon L. Ohlhorst
Reviewed work(s):
Source: PALAIOS, Vol. 3, No. 4 (Aug., 1988), pp. 413-423
Published by: SEPM Society for Sedimentary Geology
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MARINEHARDSUBSTRATECOMMUNITIES
413
HardSubstrataCommunity
Patterns,1-120 M,
NorthJamaica1
W. DAVID LIDDELL
Department
ofGeology,
UtahStateUniversity,
Logan,UT 84322-0705
SHARONL. OHLHORST
Department
ofFisheriesand Wildlife,
UtahStateUniversity,
Logan, UT 84322-5210
(see Milliman,
1973; Collin,1978),studiesofdeeperenvironments,particularly
thosebelow60 m, are muchrarer(GinsTropicalmarinecommunities
fromshallow-water
(<30 m) burgandJames,1973; Hartman,1973; Lang, 1974; Lang et
1979; Frickeand Meischner,
are oftendominatedby hermatypical., 1975;Jamesand Ginsburg,
carbonateenvironments
conscleractinian
coralswithlesseramountsof crustosecoralline 1985; and Littleret al., 1985) because of physiologic
algaeand endolithic
demosponges.
Livingcoveris typically
high straintson SCUBA divingand the expense of submersible
as manyofthemarinesediments
and
coastofJamaicaand at time.Thisis unfortunate,
(80-100%7p).
Alongthenorth-central
manywestern
Atlanticsites,communities
existing
below55 m biotapreservedin the rock recordare likelyto represent
a vertical
tooverhanging
thedeep depthsin excess of 30-60 m. Further,the comparisonof
wallofreeflimestone,
inhabit
existing
undergreatlydiffering
ambientenvironforereef,whichextendsto approximately
130 m. At 60 m the communities
ments
(shallow
and
deep)
will
help
elucidate
the important
resembles
scleracthat
of
shallower
water,
although
community
on community
structure.
andencrusting
and erectdemospongescontrols
tiniansarelessabundant
Deeper sitesdiffer
fromshallower
reefsitesina numberof
are muchmoreabundant.Corallinealgaeand macroalgae
are
parameters,most notably,lightand disturbance
also important
space occupantsat 60 m and livingcover important
sedimentation
rates,and abundanceof low-angle
65%. Encrusting
approaches
spongesand coralline,
filamen- intensities,
mayinfluence
the structure
of
in themiddleregionofthe substrata.These parameters
tous,and macroalgae
predominate
ina variety
ofways:1) Changesinboththe
40% ofthe deep communities
deepforereef.A low-diversity
assemblage
occupying
and spectralcharacteristics
of lightoccur withinand dominated
substratum
and endo- intensity
encrusting
bydiminutive
lithic?demosponges
and largelyendolithic
algae creasingdepth.Inasmuchas lightmay be consideredas a
filamentous
suchas algae, mostcorals,and
occursfrom100-130 m, thelowerlimitofthedeepforereef. resourceforreeforganisms
changesmight
be
structure
and zonationon theshallowerreefsis manysponges(Huston,1985),depth-related
Community
thecomposition
ofbenthic
anddiversity
mostnotably expectedto influence
controlled
bya number
ofbioticandabiotic
factors,
predationlgrazing,
lightintensity,
and turbulence.
On thedeep assemblages.2) Many types of physical(e.g., dailywindin
are greatlyreduced.While generatedwaves and, rarer,stormevents,fluctuations
forereef,grazingand turbulence
etc.) and biotic(predation/grazing)
disturbance
reduced
inintensity,
light
continues
toexert
a strong
influence
on temperature,
withincreasing
the
depth.Considering
zonation.Sedimentation
also exertsan arereducedinintensity
community
bathymetric
role assignedto disturbance
in the regulation
of
on thespatialdistribution
control
important
ofthedeepfore-reefimportant
of
(Connell,1978 and manyothers),the diversity
biotawiththemostdiverseassemblages
in areas diversity
flourishing
might
be expectedto differ
from
considerably
protectedfrom
sediment.
Despitea regime
ofreduced
disturbancedeepersettings
the Caribbean,mostsites in
indeepwater,community
constant
to shallowersites. 3) Throughout
diversity
remainsrelatively
excess of approximately
50-60m exhibita verticalto nearly
a depthof90-100 m.
vertical
submarine
profile
due to thecutting
ofseacliffs
during
theWisconsin
lowstillstand
at 100 m belowpresentmeansea
level (Goreauand Land, 1974; Liddelland Ohlhorst,1981).
INTRODUCTION
The lack of extensiveareas of low-anglehardsubstratain
Althoughhundredsof studies of tropicalshallow-waterdeeper waterinfluences
the typesof organismscapable of
fromtheWesternAtlantic
exist attaching
(<30 m) marinecommunities
to and growingon the deep reef.4) Finally,when
low-anglesurfacesdo occur on the deep reef, burialby
is a greatproblemfortheepibenthos.
no. 387 fromthe DiscoveryBay MarineLaboratoryof the sedirnent
'Contribution
This studyextendstheknowledge
ofthe shallowJamaican
of the West Indies
University
PALAIOS,1988, V. 3, p. 413-423
Copyright? 1988, The Society of Economic Paleontologistsand Mineralogists
0883-1351/88/0003-0413/$3.00
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All use subject to JSTOR Terms and Conditions
414
LIDDELL & OHLHORST
FORE
REEF
REEF CREST
BACK REEF
A. pl/mota
100m
terrace
escarpment
40
A. corv/cornis
* Zone
slope
~~\
5Z\_ >,
,
*
=
45
Fore Reef
~~~~~~East
deep
fore
reef
West Fore Reef/
j ;p
30
22
15
Zone
R
ear
Barrentflat Zone
.
Mixed/ Zone;
Buttressl
agoon
Zone
Shore
Zone
Zone
5
0.5
-6 0
- 68
-L
F__
7 5
90
---106
----1
MONTEG DI
Y SDISCOVERY
BAY,
20
200.
FIGURE
i-Index mapofDiscovery
Bay,Jamaica,area showing
study FIGURE 2-Reef profile along A-A' (Fig. 1) showing study sites
sites. A-A'is Watertower-Zingorro
traverse,B-B' is MooringOne (modifiedfromLiddell et al., 1984b).
traverse(modified
fromLiddellet al., 1984b).
reefs,arguably
amongthemoststudiedreefsintheworld,to wave-cutseacliffs
formedduringPleistocenelow stillstands
considerably
greaterdepths(up to 120 m, whichrepresents (GoreauandLand,1974; Liddelland Ohlhorst,
1981).
the lower limitof widespreadsuitablehard substratafor
METHODS
colonization
by encrusting
and endolithic
benthos).Specific
goalsareto document
hardsubstrata
community
zonation
over
Field
therangeof 1-120m, examinedepth-related
trendsin diversityoverthisrange,anddetermine
theeffects
offactorssuch
Shallow-water
censusdatawerecollectedovertherangeof
as disturbance,
lightintensity,
site microtopography,
and 1-30 m by diversusingSCUBA. A chainmethod(Porter,
sediment
accumulation
on community
structure
anddiversity. 1972) inwhicha chainwas drapedoverthereefandoriented
These willenablethedevelopment
ofmodelsfordeeper-water perpendicular
to the depthgradient
and the numberof chain
carbonate-producing
communities
and thetestingofhypothe- linksoccurring
over each substratum
categoryrecordedwas
ses relating
community
diversity
to disturbance
intensity.
used between0.5 and 5 m (Huston,1985). A linearpoint
method(LucasandSeber,1977;Eberhardt,
intercept
1978),in
whicha 10 m linewithpointslocatedevery20 cmwas draped
STUDY LOCALITY
overthereefandtheidentity
oftheorganisms
or substratum
beneatheachpointrecorded,
was usedbetween
DiscoveryBay lies on thenorth-central
coastofJamaicaat typeoccurring
Lat. 18?30'NandLong.77?20'W.The well-developed
fringing15 and 30 m (Liddelland Ohlhorst,1987). Up to 10 parallel
andoriented
to the
perpendicular
reefsoccurring
alongthiscoastdisplay
a striking,
depth-relatedlines,spacedat 1 mintervals
were observedat each site (Fig. 2). At 45 m
macrobiotic
zonationwhichhas been describedin several depthgradient,
papers (Goreau, 1959; Goreau and Goreau, 1973; Kinzie, diversusingSCUBA conductedphototransectswhichwere
to thedepthgradient.Colortranspar1973; Lang, 1974; Bonemand Stanley,1977; Liddellet al., orientedperpendicular
of0.14 m2areasweretaken
bystrobelight,
1984a,b;Huston,1985; andLiddellandOhlhorst,
1987). The encies,illuminated
at
1
m
intervals.
fringing
reefsstudiedoccuron theWestFore Reefand are
A Perrysubmersible
(PC-8), ownedand operatedby Reknownas Watertower
andZingorro
(A-A') andMooringOne
indata
Ltd.,GrandCayman,was utilized
(B-B', Figs. 1-2). Huston's(1985andunpublished)
0.5 and5 m searchSubmersibles
dataandLiddellandOhlhorst's
(1987)15mdatawerecollected collectionover the range of 53-120 m on the verticalto
ofthedeep forereef.The submersescarpment
on Watertower
Reef,whichis locatedimmediately
east ofand overhanging
were used to conduct
merges withZingorronear the reef crest. The followingible'sexternalcameraand strobelight
inwhichcolortransparencies
of0.14 m2areas
successionof structural/geomorphic
zones wouldbe encoun- phototransects
teredalongan onshoreto offshore
traverseacross theWest weretakenat 1 m spacingsat each depth.
Fore Reef:backreef,reefcrest,fore-reef
terrace,fore-reef
Laboratory
escarpment,
fore-reef
slope,deep forereef,andislandslope.
Abruptbreaksin slope occurring
alongthisprofile(e.g. the
Photocensusdatawere processedby projecting
transparfore-reef
escarpment
and deep forereef)are interpreted
as enciesat naturalsize ontoa screenwitha fixedarrayofpoints
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HARDSUBSTRATACOMMUNITYPATTERNS
415
TABLEI-Bathymetricdistribution
of majortaxonomicgroupsand site diversity*
Site'
W2
W2
W3
62.0
58.0
Other
2.6
Cnidaria
6.0
3.8
0.9
1.2
2.3
(?1.4) (?0.8) (?2.0) (?2.0)
Boring
Sponges6
-
0.0
0.0
0.0
Sclerosponges
0.0
Coralline 33.0
Algae
0.5
5
15
Depth(m)
N (points) (4782) (1265) (606)
Z3
Z3
22
30
(453) (244)
Z4
45
(526)
M4
45
(526)
Ms
53
(530)
Z5
M5
M5
Z5
M5
Z5
M5
M5
Z5
60
(515)
60
(531)
68
(537)
75
(514)
75
(531)
90
(516)
90
(540)
106
(535)
120
(527)
17.2
15.6
(?9.8) (?11.0)
8.9
(?8.8)
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.9
(?3.1)
2.3
(?1.5)
3.5
(?4.5)
1.6
(?1.5)
0.7
(?0.9)
2.7
0.8
1.0
(?2.6) (?0.9) (?1.1)
0.2
(?0.4)
0.9
(?1.2)
0.0
0.4
20.0
6.9 10.3
(?5.6) (?3.2) (?9.2) (?0.7)
0.4
(?0.5)
0.4
(?0.8)
0.4
(?0.8)
0.0
0.4
(?0.7)
0.0
0.0
0.0
1.7
2.2
6.2
4.6
(?1.7) (?1.6) (?6.8) (?2.8)
8.4
(?4.6)
15.6
(?5.0)
13.9
(?6.0)
20.1
(?7.2)
20.5
(?7.7)
17.0 16.7 27.5
19.4
(?4.6) (?4.5) (?5.5) (?10.1)
21.4
(?5.9)
20.0
(?7.0)
0.0
0.0
0.0
0.8
(?1.2)
0.0
3.2
(?2.7)
2.8
2.4
6.7
(?3.0) (?2.4) (?3.7)
1.5
(?2.0)
0.0
0.0
36.0
3.2 15.1
13.3 25.6
(?3.4) (?5.9) (?3.8) (?5.1)
5.3
(?2.9)
11.6
(?2.9)
10.9
(?5.1)
14.0
(?6.6)
13.4
(?3.8)
19.0 10.1 18.3
(?4.9) (?3.4) (?4.5)
6.1
(?2.7)
1.1
(?1.6)
0.7
(?0.9)
Filamentous2.1
Algae
0.5
5.2 12.0
3.6
4.1
(?2.8) (?3.7) (?3.5) (?2.9)
3.8
(?1.9)
4.9
(?3.2)
1.8
(?1.3)
7.1
(?3.4)
15.6
(?5.6)
6.6
16.0 16.0
(?3.5) (?5.9) (?4.4)
20.9
(?5.3)
16.6
(?7.7)
15.0
(?5.2)
Macroalgae 0.1
0.0
2.7
2.8
8.2 43.1
(?3.2) (?1.3) (?7.9) (?9.3)
26.4
(?6.0)
12.7
(?5.1)
12.4
(?5.2)
17.0
(?6.8)
16.6
(?6.9)
14.6 11.4
(?5.3) (?6.0)
0.0
0.4
(?0.5)
0.0
0.0
-
1.1
4.4
0.8
2.9
(?1.0) (?2.1) (?5.0) (?1.3)
0.2
(?0.4)
0.9
(?1.2)
0.2
(?0.4)
0.0
0.0
0.0
0.0
0.0
0.0
0.6
(?0.7)
1.2
(?0.8)
0.8
(?0.9)
1.0
(?1.1)
0.0
1.2
1.9
2.8
(?1.4) (?1.6) (?2.2)
0.4
(?0.7)
2.5
(?1.6)
1.9
(?2.7)
Corals
Sponges
Gypsina6
-
0.0
35.9 27.9 58.9 13.3
21.5
(?3.2) (?5.1) (?9.7) (?8.3) (?11.9)
0.0
0.0
0.0
0.0
0.0
0.0
Speciesno.
H'
J'
5
37
10
38
31
37
0.85 1.37 2.51 2.55 2.64 2.23
0.53 0.59 0.69 0.70 0.77 0.62
95.7
100.0
-
BareHard Substrata
Living
Cover
Sediment
0.0
0.0
Misc.
44
2.89
0.76
70.4
83.8 81.8 94.7 83.3
(?3.4) (?3.8) (?5.8) (?8.7) (?12.5)
45
3.16
0.83
66.9
(?9.6)
39
2.84
0.78
34
2.91
0.82
34
2.69
0.76
70.4
60.3
69.7
(?9.9) (?10.6) (?11.5)
0.0
0.0
0.0
0.0
0.0
39
36
43
2.93 2.63 2.98
0.80 0.73 0.79
29
2.30
0.68
40
2.60
0.70
21
1.82
0.60
63.8 59.3 72.2
(?9.2) (?8.8) (?6.8)
42.5
37.6
48.9
(?9.4) (?10.5) (?11.5)
0.0
19.6
19.6
3.8 16.8
0.8
5.8
19.8
20.8
23.4
25.5 39.7 11.1
(?2.4) (?4.2) (?1.4) (?6.4) (?12.8) (?10.2) (?10.1) (?10.6) (?12.6) (?11.3) (?9.0) (?6.9)
54.1
49.3
52.3
(?9.5) (?11.7) (?13.9)
0.0
12.4
1.3
4.5
10.9
(?3.4) (?1.1) (?4.5) (?4.9)
10.0
(?3.8)
13.6
(?5.5)
19.9
(?8.6)
9.5
(?4.4)
6.2
(?3.6)
10.6
1.0 16.7
(?5.8) (?1.1) (?5.6)
1.9
(?1.7)
5.2
(?4.0)
8.3
(?4.6)
inparentheses.
intervals
*95%confidence
Z = Zingorro(deep forereefbeginsat approx.60 m), M = MooringOne (deep forereefbeginsat approx.50 m).
'W = Watertower,
2Datafrom
Huston(1985andunpublished),
chainmethod.
LiddellandOhlhorst
3Datafrom
(1987),LPI method.
4Diverphototransects.
5Submersible
phototransects.
from
at 0.5 and5 m,butnotseparated
coralline
6Almost
rubble
certainly
present
algaeencrusted
byHuston(1985andunpublished).
ofthe made; thus,some speciesmayhave been incorrectly
(27 points,each witha 10 cmspacing)andtheidentity
"split"
categoryoccurringat each point whileotherswereincorrectly
combined.
organismor substratum
Bathymetric
trendsin
method;Kinzieand Snider, diversity,
recorded(planarpointintercept
livingcover,anddistribution
ofparticular
taxawere
community
com- testedwiththeSpearmanRankCorrelation
1978). Censusdatawereused to determine
Coefficient
(SRC).
at each depth(Table 1). Clusteranalysis(normalized
position(largertaxa) and diversity
Eucidean distancecoefficient,
(H', nat.log; Shan- unweighted
diversity
Speciesnumber(S), dominance
centroid
algorithm,
Systatversion3.0) was usedto
nonand Weaver,1948), and evenness J',nat. log; Pielou, discernzonationpatterns.
valuesso
as diversity
indices.Diversity
1966),wereemployed
Fore-reefsites are characterized
by low waterturbidity
The filamentous(lightattenuation
coefficient
0.06/mfortheupper30 m ofthe
obtainedmustbe viewedas approximations.
thatcould water colunm,Brakel, 1979). Althoughthe attenuation
aggregations
represents
multispecies
algaecategory
(a
fromtheslides.Also,spongespeciessepara- combination
notbe identified
of absorption
and scattering)
of sunlight
by sea
in theupper5 m ofthewatercolumn,the
havenotyetbeen wateris nonlinear
as spiculepreparations
tionsare tentative,
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LIDDELL& OHLHORST
416
and deeper
bothshallow,now disturbed,
used in describing
reefsites.
BackReefto IslandSlopeTraverse
The backreefor lagoonis an area ofreducedcoralgrowth
by macroalgaeand high
due to highsedimentproduction
ofthe
activity
causedbytheburrowing
resuspension
sediment
Callianassa(Allerand Dodge, 1974). The reef
ghostshrimp
crestextendsfromthesurfaceseawardto a depthofapprox7 m andconsistsoflargepalmatecoloniesofthecoral
imately
to the
perpendicular
palmata,withbranchesoriented
Acropora
sites(1-5 m)exhibit
fore-reef
waves.The shallowest
incoming
grazing
by
andbiotic(largely
(turbulence)
highratesofphysical
fish)disturand,to a muchlesserdegreeat Jamaica,
urchins
adaptations
bance. Few corals possess the biomechanical
(Shinn,1963; Grauset al., 1977)necessaryto surviveinthis
low.
is, accordingly,
environment
and coralspeciesdiversity
by the coralA. palmata,crustose
The faunais dominated
FIGURE 3-Fore-reef slope site at 45 m. Note A) plating coral coralline
diversity
cionidsponges;totalfaunal
algae,andboring
morphologies,B) gorgonians, and C) demosponges. Field of view is also low (Table 1).
approximately3 m.
terraceextendsfrom7 to 15 mandconsistsof
The fore-reef
to thereef
perpendicular
largeparallellobesofcoraloriented
crestandextending
seawardat a moderate(15-20?) slopefor
by sand
200 m. These lobes are interrupted
approximately
These
in
topography.
resulting
a
spur-and-groove
channels,
at greaterdepths
a constant
approaches
coefficient
attenuation
m
the
above
to
5
rise
up
lobes
or
of
coral
often
spurs
coeffiBrakel's(1979) attenuation
1977). Therefore,
Jerlov,
in
(450 +) escarpment
a
steep
sand
and
terminate
surrounding
valuesto a depthof
irradiance
cientwas used to extrapolate
siteson theouter
from15 to 25 m. Fifteen-meter
extending
120 m in thewatercolumn.This exerciseyieldeda 1.0% of
terraceare dominatedby corals
portionof the fore-reef
0.05%
m
and
a
65
at
value
approximately
surfaceillumination
including
mounds(Montastrea
diversemorphologies,
bythisprocedure exhibiting
valuesgenerated
valueat 110 m. Irradiance
fingers
(Po(Porites
astreoides),
encrusting
sheets
annularis),
forin situ quantumirradiance
are, of course,no substitute
to
In
addition
cervicornis).
(Acropora
and
branches
ritesporites),
andmustbe regardedwithcaution.
measurements
cionids)andcrustosecoralline
corals,boringsponges(largely
Noncrustose
algal
community
components.
algaeareimportant
RESULTS
and grazingby urchinsstill
biomassis verylow. Turbulence
at thisdepth.
on thereefcommunity
exertsignificant
effects
are greatly
diversities
outthatshallowreefs Coralspeciesandtotalmacrocommunity
to beginbypointing
It is appropriate
theywere once increasedover5 m sites.
stableenvironments
are farfromthe highiy
slope,whichbeginsat
for
The landward
ofthefore-reef
portion
coastofJamaica,
The reefsofthenorth-central
considered.
eventsthis 25-30 m depth,usuallyconsistsofa low-angle(5-20?) sand
two severe disturbance
example,have suffered
scattered
coralheads.Coralcoverincreases
Allen,whichstruckthenorth "moat"containing
was Hurricane
decade.The first
parallellobes and occasional
coast in August,1980, and caused extensivedamageto the by 30-35 m, oftenforming
andLiddell, pinnacleswhichmay rise up to 10 m above the sand. At
shallowreefs(see Woodleyet al., 1981;Ohlhorst
35 m the slopeincreasesto between45-60?.
The second approximately
1981; andLiddellet al., 1984bfordescriptions).
byplatecorals,
slopeat 30-45 m is dominated
of the urchingrazer,Diadema The fore-reef
event,the mass mortality
annularis,and variousAgariciaspecies
occurredin August,1983 whilethe reefswere largelyMontastrea
antillarum,
and
suchas Lobofora
Thereduction (Fig. 3). Demospongesandmacroalgae,
thehurricane.
a successionfollowing
undergoing
membersof the benthos,while
inurchin
densitiesfrom14/M2at somesitesto nearly0.0/M2 Halimeda, are important
and crustosecorallinealgae and boringcionidspongesare less
increasein filamentous
resultedin a rapidand dramatic
of
thanat shallower
depths.The declineinabundance
biomasswhichoccurredat the expenseof abundant
erectmacroalgal
to a lackof
slopeis attributed
otherbenthos,suchas corals,boringsponges,and crustose thesegroupson thefore-reef
generated
dueto theabsenceofturbulence
corallinealgae (Hugheset al., 1985; Liddelland Ohlhorst, suitablesubstrata,
with
forms
coralgrowth
ofplate-like
didnotoccuringreatnumbers debrisandtheprevalence
as theurchin
1986). Inasmuch
1987). In
at depthsin excess of 20 m, the effectsof the urchinmass littleexposed,bare skeleton(Liddelland Ohlhorst,
ofmacroalgae
overthisrangemay
thepredominance
to the shallowerreefs.As ofJuly, addition,
were restricted
mortality
influence
the abundanceof crustosecorallines.It
1987, the shallowreefshave stillnot recoveredfromthis negatively
coralsfromthe
The data?wearepresent- shouldbe notedthatboringspongesattacking
(personalobservation).
perturbation
ofplateswouldnotbe recordedusingtheabove
inghereinforshallowreefsiteswerecollectedpriorto these undersurface
andgrazingbyurchins
is muchreduced
thepresenttensewillbe methods.Turbulence
events.Forconsistency,
disturbance
This content downloaded on Fri, 15 Feb 2013 15:44:31 PM
All use subject to JSTOR Terms and Conditions
417
HARDSUBSTRATACOMMUNITYPATTERNS
Nt
B
A-~~~~~~~-
A'
FIGURE4-Deep fore-reefsite at 55 m. Note A) large, platingcoral
Agaricia,B) macroalgae Lobofora,and C) Halimeda. Field of view is
approximately35 cm.
FIGURE5-Deep fore-reefsite at 68 m. Note A) sclerosponges, B)
demosponges, C) crustose corallinealgae, and D) Halimeda. Field of
view is approximately35 cm.
4
is
coraldiversity
relativeto shallowerreefsites. Although
toshallower
sites,overallmacrorelative
decreasedsomewhat
A
to thatof15 m sites.
is equivalent
diversity
community
~~~~~~~~~~~~~~~~~~~~~~~~~~~.
~
~
~
~
~ ~ ~
at
between45 and65 m, typically
A slopebreakoccurring
"wall."
of the deep fore-reef
55 m, marksthe beginning
oftheupperpartofthedeep
thefaunalcomposition
Although
tothatofthelowerfore-reef
fore-reef
wall(55-60 m)is similar
increase
whiledemosponges
slope,coralsare less abundant
in abundanceand macroalgaeremainimportant
dramatically
(Fig. 4). By 70-75 m coralsbecome
components
community
butnotdetectedby
rare(observedbyresearchers,
extremely
photocensuses).Crustosecorallinealgae are abundantand
macroalgae(largelyvariousHalimedaspecies)are also cominmoresheltered
areas(Fig.
arecommon
mon.Sclerosponges
sites.
to thatofshallower
is similar
diversity
5). Community
The slope of the walldecreasesat a depthof 105 m and
on exposedsurfacesbecomesa major
accumulation
sediment
FIGURE6-Deep fore-reefsite at 120 m. Note A) demosponges and
Endolithic
algae and encrustingfilamentous-endolithic
problemforthe epibenthos.
algae (nearlyentire surface not occupied by
components, sponges). Field of view is approximately35 cm.
are themostimportant
community
demosponges
inabundecreasegreatly
andsclerosponges
whilemacroalgae
dance(Fig. 6).
The steep (60-90?) escarpmentends at approximately
120-130 m wherethe moregentle(20-45?) and sediment- the rangeof 0.5-30 m, declinesto 9-16% by 60 m on the
organismsare re- upperdeep fore-reef
coveredislandslope begins.Encrusting
dropsto near0.0% by
wall,and finally
stricted
to isolatedblocksoftalusderivedfromabove.
colonies
75 m, althoughrare hermatypic
and ahermatypic
in
abundance
this
The
decline
occur
below
depth.
occasionally
Composition
Trendsin Community
Bathymetric
withincreasing
(SRC, p<0.01).
depthis significant
algaeexhibita somewhatbimodaldistriLivingcoveris highest(82-100%) on the crest,fore-reef Crustosecoralline
slope (Table 1). On the bution(Fig. 7) reachinghighestabundancesat 0.5-5 m
terrace,and upper(30 m) fore-reef
terraceandat 75-90 m (18-19%)
slope(45 m)andtheupper-tomid-deep (33-36%) on thefore-reef
mid-tolowerfore-reef
slope
coverrangebetween on the deep forereef.At 30-53 m on the fore-reef
fore-reef
wall(53-70 m),valuesforliving
Overall,
algaeoccupyonly3-15% ofthesubstratum.
60-70%,whileonthelowerdeepforereef(105-120m),living coralline
correlatedwithdepth(SRC,
coverdeclinesto 38-43%. Thereis a generaldeclineoverthe corallinealgae are negatively
p<0.05).
rangeof0.5-120 m (Fig. 7; SRC, p<0.01).
Noncrustosealgae, including
both filamentous-endolithic
between28-62% over
Coralabundance(Fig. 7) fluctuates
4
This content downloaded on Fri, 15 Feb 2013 15:44:31 PM
All use subject to JSTOR Terms and Conditions
418
LIDDELL& OHLHORST
100
70
90
60
*
8
80
0
S
*
Z70
S0
~~~~~~~~~~~50
1)40
*
M60
30O
0
0~~~~~~~~~~~~~~~~2
50
10 t
*
40 |
30~~~~~~~~~~~~~~~~
,
,
,
,
,
,
, *
t
t
e
0.5M 15M 30M 53M 68M 90M 160M
0.5M 15M 30M 53M 68M 90M 120M
5M 22M 45M 60M 75M 105M
5M
50
50
40
40
22M 45M 60M 75M 105M
O~~~~~~~~~
OS30
z
o
30
12
~~~~~~~~~~~~~U)0
w
I(1)
20
2
*e0
.
0
,
0~~~~
10~~~~
10
0~~~00
0.5M 15M 30M 53M 68M 90M 120M
5M 22M 45M 60M 75M 105M
0.5M 15M 30M 53M 68M 90M 120M
SM 22M 45M 60M 75M 105M
8-
30
~~~~~~25
0
6
o
z
U
2000
0
wn
o
Z 15
U)
*
10*
2
0~~~~~~~~~~
0.5M 15SM 30M 53M 68M 90M 120M
SM 22M 4SM 60M 7SM 10OSM
00.5M
DEPTH
15SM 30M 53M 68M 90M 120M
SM 22M 4SM 60M 7SM 10OSM
ofbioticcover(percentofavailablesubstratum).
distribution
FIGURE
7-Bathymetric
This content downloaded on Fri, 15 Feb 2013 15:44:31 PM
All use subject to JSTOR Terms and Conditions
DEPTH
HARDSUBSTRATACOMMUNITYPATTERNS
419
reds)and W0.5
and,inshallow-water,
algae(greensandblue-greens
macroalgae(greensandbrowns)reachtheirpeak abundance W5
theyare stillsignificant
(30-47%) at 45 m (Fig. 7), although
(16%) at 120 m. The increaseinnon- WW15
community
components
Z 22
(SRC, p<0.05)
to 45 m is significant
crustosealgalabundance
less so (SRC, Z 30
whilethedecreasefrom45 to 120 m is slightly
ofthetwogroupsdo Z 45
p<0. 10). Shifts
intherelative
abundances
overtherangeof30- M 45
occur,withmacroalgaepredominating
algaefrom75-120 m. Macro75 m andfilamentous-endolithic
inabundance
to45 m(SRC, p<0. 05) M 53
algaeincreasesignificantly
Z 60
andthendeclineto90 m(SRC,p<O. 01). Filamentous-endolithic
depthoverthe M 60
correlated
withincreasing
algaeare positively
rangeof0.5-120 m (SRC, p<0.01).
M 68
themostrestricted
rangeof M 75
(Fig. 7) exhibit
Sclerosponges
components,occurringin relatively
the major community
locationsonlyfrom60-90 m. A peak Z 75
exposed (noncryptic)
abundanceof 7% occursat 90 m. Sclerospongesare known Z 90
settingssuchas caves and theunder- M 90
fromshallowercryptic
surfacesof foliaceouscorals (Hartmanand Goreau, 1970; M 106
Jackson
et al., 1971).
Z 120
bimodaldistribution
with
a somewhat
Demospongesexhibit
10
20
0
an overallslighttrendof increasingabundancewithdepth
cionidsare
(SRC, p<0.10, Fig. 7). If, however,endolithic
NORMALIZED EUCLIDEAN
fromerector encrusting
demosponges,
considered
separately
withincreasing
depth(from FIGURE 8-Cluster dendrogram showing zonation pattern. Note
theformer
decreasesignificantly
20.0% to 0.0%; SRC p<0.01) whilethelatterincreasesignif- prominentseparation of shallow (<30 m) fore-reefsites and, to a
icantly
(from0.0 to 20.0%; SRC, p<0.01) andremainimpor- lesser degree, mid-fore-reef
slope (45 m) sites fromdeep fore-reef
to 120 m (Table 1).
tantcommunity
components
sites. Also note divisionof deep fore-reefsites intothree zones.
zonation
bathymetric
Clusteranalysisrevealsa well-defined
were delineatedwiththeir
(Fig. 8). Two majorgroupings
thedeeper
between30 and45 m. Within
boundary
occurring
while
slopesitesweremostdistinct
clusterthe45 m fore-reef
EffectofMicrotopography
intofore-reef
slope-deep
deepforereefsitesweresubdivided
Microtopographies
are inequally
distributed
overthebathyfore-reef
break(53-60 m), upperdeep forereef(68-75 m),
and lower deep fore reef (90-120 m). From onshoreto metricrangeof our study.Exposed low-anglesurfacespreoffshore
the following
groupsdominateeach zone (listedin dominateabove 60 m, whileverticalexposed or sheltered
on theupperporcorals,coralline
algae, boring (beneathoverhangs)surfacespredominate
orderofdeclining
importance):
ofthedeep
sponges (0.5-22 m); macroalgae,corals, corallinealgae tionsofthedeep forereef.On thelowerportions
inclinedexposed surfaces
(45 m); sponges, corals, macroalgae, coralline algae forereef,verticalto moderately
coralline
algae,filamentouspredominate.
(53-60 m); sponges,macroalgae,
On thedeep forereef,verticalsitesconsistently
exhibit
the
algae(90-120 m). In
algae(68-75 m); andsponge,filamentous
groupsare approxi- highestpercentageof livingcover, while low-anglesites
the threedeepest zones the dominant
cover.This
exhibit
thelowestpercentage
ofliving
consistently
matelycoequal.
effectis largelycaused by the accumulation
of sedimenton
sitestypically
have
low-angle
surfaces.Whilevertical
sheltered
TrendsinDiversity
Bathymetric
higherpercentages
oflivingcoverthanverticalexposedsites,
no consistentdifferences
existbetweenlow-anglesheltered
demo- andexposedsites.
level(corals,noncoralcnidarians,
At themacrobiota
availableto
also influences
theirradiance
etc.), species diversity Microtopography
macroalgae,
sponges,sclerosponges,
thata north-facing
andevenness(S, H', J'; Table 1, Fig. 9) is low at 0.5-5 m, the benthos.Brakel(1979) demonstrated
to increaseto 53 m (SRC, verticalsurfacewouldreceiveonly25% and a 600 slope only
increasesby 15 m, andcontinues
surface.Thus,
to 120m (SRC, p<0.05 forH' andJ' 50% oftheillumination
receivedbya horizontal
declining
p<0.05), finally
value
is notcorrelated
withpercentage theestimateddepthofthe 1.0% ofsurfaceillumination
diversity
only).Community
is relatively for the water column(65 m) must be shiftedupwardto
ofliving
totalcommunity
cover.Although
diversity
55 m fora 600 surfaceon theupperdeep fore
constantover a broadbathymetric
range(15-106 m), major approximately
ofthelargertaxonomic
slopeandto 45 mfora verticalsurface.
shiftsin species diversity
categories reeforlowerfore-reef
thedepthofthe0.05% valuemustbe shifted
declinesrapidly Similarly,
upward
occur.For example,coralspecies diversity
on from110 m in thewatercolumnto approximately
90 m fora
byan increaseinspongediversity
below50 m butis offset
verticalsurfaceandto 100 m fora 60? surface.
thedeep forereef.
This content downloaded on Fri, 15 Feb 2013 15:44:31 PM
All use subject to JSTOR Terms and Conditions
420
LIDDELL & OHLHORST
VS DEPTH
DIVERSITY
DISCUSSION
50
Controlson Community
Composition
andDiversity
.0
40 .
40*
0.~~
*
*
*
@
@
30
@
0
U)
20
10
0
o
0.5M 15M 30M 53M 68M 90M 120M
5M 22M 45M 60M 75M 105M
3.5
3.0~~~~~~
3.0
2.5
:
*
-
*
.*0
2.0
I~~~~
1.5
1.0
0
0. ,@
., . .
,
.
0.5M 15M 30M 53M 68M 90M 120M
5M 22M 45M 60M 75M 105M
1.0
.8
.8
0.
.
0
06
.6
.41
0.5M 15M 30M 53M 68M 90M 120M
5M 22M 45M 60M 75M 105M
DEPTH
FIGURE9-Bathymetric trends in diversity(all taxa). S = species
number,H' = the Shannon-Weaver(1948) information
function,J' =
evenness (Pielou, 1966).
The depth-related
bioticzonationshownby these reefsis
the resultof the complexinterplay
betweena varietyof
physical/chemical
parameters
andbioticinteractions.
In areas
locatedabove wave-base(approximately
20-30 m), depthrelatedshiftsin reefcommunity
composition
are mostlikely
influenced
byenvironmental
turbulence
(Geister,1977). Light
levels willprobablynot limitcoral species diversity
on the
shallower
(<30 m)reefs(Loya,1976).Turbulence
willinteract
withthebiomechanical
characteristics
ofvariouscoralspecies
all butAcropora
(e.g., excluding
palmatafromtheveryshallow-1-5 m-fore reef). Another,more subtle,effectof
turbulence
on reefcommunity
composition
maybe thegenerationof hardsubstrataby the toppling
and fragmentation
of
coralcoloniesinshallowwater.Suchhardsubstrata
are rapidly
colonizedbycoralline
and
algaeandboringsponges(Ohlhorst
Liddell,1981; Liddellet al., 1984b), whichare important
components
of shallowreefcommunities.
The inversecorrelationsof these groupswithdepthsuggestthatthe lack of
hard substratamay accountfortheir
turbulence-generated
muchreducedabundanceon thefore-reef
slope (30-55 m).
Diademagrazing
hasbeenshowntoinfluence
themacroalgal
structure
ofcoralreefsmarkedly
community
andalso to affect
sessileepibenthic
invertebrates
(Sammarcoet al., 1974; Carpenter,1981). The increasein macroalgal
abundanceon the
fore-reef
slope is most likelyan effectof the decrease in
Diadema abundancebelow 20-30 m (Liddelland Ohlhorst,
1987). The causal factorsbehindDiadema's depthlimitation
are unknown.
The bathymetric
of
distribution
oftheintensity
predationor grazingby othergroupsis not well known,
thereis someevidencethattheeffects
although
offish(Bakus,
1969, 1972) and otherinvertebrates,
suchas the polychaete
Hermodice
and the gastropodCoralliophila(Ott and Lewis,
in shallowwater.Personalobser1972), are mostimportant
vationsat Jamaicasupportthesefindings.
On thewallofthedeep forereef,turbulence
and grazing?
are likelyto generaterelativelylittleeffecton the biota;
and sedimentation
instead,lightintensities
are likelyto be
controllers
ofcommunity
important
composition.
Lightintensitiesdeclinewithincreasing
depthandare also influenced
by
site microtopography
and shelteredor
(degreeof inclination
Sedimentation
is also influenced
exposed)andorientation.
by
sitemicrotopography,
withhighestaccumulations
on
occurring
surfaces.
exposed,low-angle
The community
of the deep forereefdiffers
composition
fromthatofshallower
greatly
reefsites.Whereastheshallow
reef(5-30 m)is dominated
crustosecoralline
byscleractinians,
algae, and endolithic
demosponges,the deep forereef(60120 m) is dominated
by erect or encrusting
demosponges,
crustosecoralline
algae,andnoncrustose
(macro-andfilamentous)algae. In termsofmacrocommunity
thedeep
diversity,
forereefis verysimilar
to shallowersites.As such,thedeep
fore-reef
datado notappearto followthepredicted
diversity
trendsof the intermediate
disturbance
as applied
hypothesis
to coralreefs.Sites withintermediate
levels of disturbance
shouldexhibitthehighestdiversities
as disturbance
prevents
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HARDSUBSTRATACOMMUNITYPATTERNS
421
exclusionby competitively
superiorspecies. The deep fore maica,coralsdidnotextendin abundancebelow60 m even
reefappearsto be an environment
ofreduceddisturbance
as thoughsuitablesubstratawere availableconsiderably
below
turbulence
and grazing,amongthe majorsourcesof distur- thisdepth.Thisrelatively
with
shallowupperlimit
is consistent
banceontheshallow-reefs,
are absentor reduced.Deep fore- thecalculatedquantum
irradiance
availableon theverysteep
reefcommunity
diversity,
however,is as highas orhigher
than deepforereef(1% leveloccurring
at 55 m). M. cavernosaand
thatofmuchshallower
sites(e.g., 0.5-30 m). Perhapsother Agariciaspecies were also the most abundantdeep-water
factors,suchas sedimentation
or detachment
fromthesteep coralsat Jamaica.
escarpmentof the deep fore reef are servingto reduce
Littleret al. (1985) concentrated
on thealgae occurring
on
competitive
exclusionand, thus, promotediversity.The thesteepfaceofa seamount
offSan SalvadorIsland,Bahamas.
to do so as diversity
is highestin From81 m (the top of the seamount)to 268 m, the algal
former,
however,is unlikely
areas littleaffected
by sedimentation
(e.g., verticalsheltered community
complexity
and spatialheterogeneity
decreased
sites). Furthermore,
whilethe lattermayaffectcoralsand withincreasing
depth.The brown,erectmacroalgaLobofora
erectdemosponges
on theupperpartofthedeep forereef,it variegatapredominated
from80-88 m, occupying
a meanof
is unlikely
to influence
thetightly
encrusting
demosponges
on 59.4%ofthesubstratum.
Halimedacopiosadominated
thealgal
themiddleandlowerportions
ofthedeep forereef.
overtherangeof117-130m, accounting
community
forup to
The deep forereefis nota uniform
environment,
however, approximately
20% coverage.Two greenalgaeoccurredabunas light
intensity
does decreasewithincreasing
depth.Iflightis dantlyfromapproximately
130-157 m and the green,rockviewedas a nutritive
resourceforphotosynthetic
organisms boringalga Ostreobium
occurredas deep as 210 m. Coralline
(cf.,Huston,1979, 1980, 1985; Tilman,1982), perhapsthe algaedominated
between210-268m andoccupiedfrom5% to
diversitiesof the deep fore-reef
communities
reflectthe over 20% of the substratum
from80-268 m. Encrusting
mediating
effectsofintermediate
resourcelevelson competi- spongeswere foundto be the predominant
organismsfrom
tionbythebenthosdespiteverylowlevelsofdisturbance. 268-520 m.
The deepestoccurring
coralline
alga was at 268 m. Significantly,thisalga occurredat a lightintensity
of 0.0005% of
OtherStudiesofDeep-WaterCommunities
surfaceillumination,
considerably
belowthegenerally
accepted
marinecommunitieslower limit(1%) of the photiczone. In addition,fleshy
Studiesofdeep-water
(>60 m) tropical
werefoundto extendto a 0.05% level.AtJamaica
holisticstudiesof macroalgae
are rare. Even more so are quantitative
macrophytes,
but
deep-watercommunities.
The earliestattemptsto utilize LoboforaandHalimedaare also important
inthestudyofCaribbeandeep-water
overa muchshallowerdepthrange.AtJamaica
submersibles
communi- predominate
ties were conductedat Belize (Ginsburgand James,1973; corraline
algae were also foundto extendto greaterdepths
andspongeseventually
JamesandGinsburg,
1979)andJamaica
(Hartman,
1973;Lang, thanmacroalgae
replacedbothofthese
1974;Langet al., 1975).These studieswerenon-quantitative,
groupsas dominant,
although
the"zone"boundaries
are shifted
on the lower depth considerably
documentation
upwardsthere.
yet providedimportant
limits
ofmanytaxa.In particular,
variousalgaeandhermatypic Studiesof deep-watercommunities
Atfromnon-western
corals were foundto extendto muchgreaterdepthsthan lanticsitesincludeFrickeandSchuhmacher
(1983) andHilliscoral Colinvaux(1986). The formerexaminedthe distribution
of
previouslyrecognized.For example,the hermatypic
Agariciafragiliswas shownto extendto at least 97 m at coralsovertherangeof0-145 m alongtheSinaicoastofthe
hermatypic
coralspecies were
Jamaica
(Lang,1974)and103matBelize JamesandGinsburg, Red Sea. One hundredthirty
1979). Furthermore,
crustosecoralline
algae and filamentousfoundto occurovertherangeof10-30 m whilea totalofonly
algaewerefoundto extendto 175 m at Jamaica(Lang,1974) forty-seven
species occurredbelow 50 m and nine species
and crustosecorallinealgae to 250 m at Belize Uames and below100 m. A singlespeciesoccurredbelow110 m. At this
ofsclerosponges
as primary locality
the1% lightleveloccurredat 100 m. The occurrence
Ginsburg,
1979). The importance
was documented
framework
constructors
deep-water
byLang of such a highnumberof hermatypic
species below 70 m
et al. (1975).
contrastssharply
withtheWesternAtlantic
settings(Jamaica
Frickeand Meischner(1985) examinedthe bathymetricandBermuda).Also,no individual
coralor smallgroup(e.g.,
ofcoralsovertherangeof0-79 m at Bermudaand M. cavernosaandAgariciaspecies) dominated
thedeep Red
distribution
foundcoraldiversity
constant
(H') to be relatively
(1.50-1.75) Sea assemblageas intheWesternAtlantic.
of
withonlya slight
on the distribution
increaseovertherangeof0-39 m. Diversity Hillis-Colinvaux
(1986) concentrated
to 1.2 by49 m andfinally
to 0.0 by algae,chiefly
Halimeda,overtherangeof30-367 m on the
decreasedrathersharply
The deep-water steepforereefat Eniwetok
79 m due to a lackofsuitablehardsubstrata.
Atoll.From30-65 mthebiotawas
associationconsistedof Montastrea composed of hermatypicscleractinians,Halimeda, and
(>60 m) hermatypic
cavernosa,Agariciafragilis,and Scolymiacubensis,withM. crustosecoralline
algae.From65-110 mHalimedadominated
thegreatestamountof space and being theassemblage,although
coralswerealso present.Few or no
cavernosaoccupying
the corals extendedbelow 90 m whileHalimeda reached30%
theonlycoralto extendbelow70 m (to 78 m). Although
photiczone limit(i.e., the compensation
pointwherephoto- cover above 90 m and 10-25% cover between90-110 m,
to 0% below140 m. Finally,
decreasing
synthesisequals respiration,
normally
regardedas 1% of finally
Hillis-Colinvaux
surfaceillumination;
werenotas abundant
Ryther,1956) didnotoccuruntil100 m, notedthat,whilepresent,sclerosponges
to
coralswererestricted
to muchshallower
The deep-water
at thissiteis similar
depthsdueto lackof as atJamaica.
zonation
withmacroalgae.At Ja- thatat Jamaica,although
suitablesubstrataand competition
the transition
to the coralhine
algal
This content downloaded on Fri, 15 Feb 2013 15:44:31 PM
All use subject to JSTOR Terms and Conditions
422
LIDDELL& OHLHORST
dominated
community
occurredat a muchshallowerdepthat
Jamaica.
Implications
forStudiesofAncientHard
SubstrataCommunities
EBERHARDT,L.L., 1978,Transect
methods
forpopulation
studies:
Journal
ofWildlife
Management,
v. 42, p. 1-31.
FRICKE, H.W.,andSCHUHMACHER,
H., 1983,Thedepthlimits
ofRedSea
stonycorals: An ecophysiological
problem(a deep divingsurveyby
MarineEcology,
v. 4, p. 163-194.
submersible):
1985,Depthlimits
ofBermudan
scleraccorals:a submersible
tinian
Marine
v. 88,p. 175-187.
survey:
Biology,
This studyprovidesseveralimplications
forthe paleoecoof wave exposureon the ecological
logicalinterpretation
of ancienthardsubstratacommunities.GEISTER, J., 1977,The influence
zonation
ofCaribbean
coralreefs:Proceedings
oftheThirdInternaMost notableis the trendforskeletonized
communities
to
tional
Coral
Reef
ofMiami,
Symposium,
Rosenstiel
School,
University
in shallowwaterandless skeletonized
dominate
communities Florida,
v. 1, p. 23-30.
to dominate
in deep water.The preservedassemblagewould GINSBURG,R.N.,andJAMES,N.P., 1973,British
Honduras
bysubmarine:
indeep water,accomshowan apparent
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What could be more comforting,
what more convenientforhuman domination,than the traditional
concept of a young earth,ruled by humanwillwithindays of itsorigin.How threatening,by contrast,
withhuman habitationrestrictedto a millimicrothe notionof an almost incomprehensibleimmensity,
second at the veryend!
-Stephen Jay Gould
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