Novak*, M.J., W.D. Liddell and D. Torruco.

Proceedings of the Seventh International Coral Reef Symposium, Guam, 1992, Vol. 1
Sedimentology and Community Structure of Reefs of the
Yucatan Peninsula, Mexico
Dept of Geology, Utah State Univ, Logan, UT 84322-4505 USA
Dept of Geology, Utah State Univ, Logan, UT 84322-4505 USA
Centro de Investigaciones y de Estudios Avanzados del IPN, Merida, Yucatan 97310 MEXICO
Abstract. Holocene carbonate sediments from
Mexican reefs in the ~aribbeanand Gulf of Mexico
display variation in constituent composition, texture and mineralogy which is related to their location on the reefs. Samples were collected at barrier reef environments at Akumal and Chemuyil,
on the northeast coast of the Yucatan Peninsula; at
the oceanic atoll of Chinchono, off the southeast
coast of the Peninsula; and at the shelf atoll of Alacranes, in the Gulf of Mexico. Samples were collected over a depth range of 1-40 m, encompassing
back reef, shallow fore reef and deeper fore reef
environments.
G~~~~~~
than 90% of the sediment (by weight) is
contained in the interval of 0.125-2.0 mm, with
mean grain size (Mz) approaching 0.5 mm at all
sites. Mz was found to decrease with increasing
depth at three of the four sites. Reef sediments are
moderately to poorly-sorted and typically become
more poorly-sorted with increasing depth. Sedimerits collected
reef
(grooves) are
consistently more well-sorted than those from the
reef interstices (spurs).
Lagoon facies are dominated by Halimeda with
lesser amounts of coral and coralline algae. In Contrast, shallow and deep fore reef facies are dominated by coral, with lesser amounts of Halimeda
and coralline algae. Q-mode cluster analysis of constituent data enables the delineation of lagoon (back
reef), shallow reef (< 10-1 5 m) and deep reef (>1015 m) lithofacies.
Mineralogically, the sediment is chiefly composed of one or more of the polymorphs of CaCO,
(96.3-99.896) with only a small Percentage of insoluble (non-carbonate) material. The non-carbonate fraction of the sediment is dominated by organics (0.1-3.296) with lesser amounts of clay and
amorphous silica (0.0-0.496).
Quantitative analysis of Mexican reef sediments
shows that strikingly similar depth-related zonations are present at sites which occur in significantly
different geomorphic environments and are separated by greater than 500 km. These sedimentological zonations reflect the community composition
of the living reefs.
Introduction
The Yucatan Peninsula (Fig. 1) is a relatively remote area which has only recently received significant study. Studies of Holocene carbonate sedimentation along the Caribbean Coast of the
Yucatan Peninsula have primarily been restricted
to Isla Mujeres (Folk 1967a) and Cozumel (Spaw
1978). The atoll Alacranes in the Gulf of Mexico
has received considerably more study (Kornicker
and Boyd 1962; Folk and Robles 1964; Hoskins
266
Novak et al.
1966; Folk 1967b; Logan 1969). The only other area
in the vicinity of the Yucatan Peninsula which has
received significant study of Holocene sediments is
Belize. Wantland and Pusey (1 975) provide information about sedimentation, as well as reef structure, morphology and ecology along the coast of
Belize. More recently, Macintyre et al. (1986) conducted a detailed sedimentological and biological
survey of the Tobacco reef sediment apron on the
central Belizean coast and Mazzullo et al. (1992)
provide data on a lagoonal patch reef from the
northern Belize reef. Most previous studies of reef
sedimentation in the vicinity of the Yucatan Peninsula have been qualitative, rather than quantitative, in nature and few have addressed the relationship existing between carbonate sediment
composition and the organisms responsible for sediment production.
Studies of reef biota and reef structure have been
conducted at various locations along the Caribbean
coast of the Yucatan Peninsula in northeastern
Quintana Roo (Ekdale 1974; Jordan 1979a, b, 1989;
Jordan et al. 1981; Fenner 1988), the Banco Chinchorro (Chavez and Hidalgo 1984; Jordan and Martin 1987) and Belize (Wantland and Pusey 1975;
Rutzler and MacIntryre 1982; Macintyre et al.
1986). Atolls from the Gulf of Mexico have been
described by Farrell et al. (1983), Chavez et al.
(1985) and Liddell and Ohlhorst (1988).
The following paper integrates sedimentological
characteristics with the distribution of biotic communities across a broad range of environments
(back reef and shallow to deep fore reef) in an attempt to define the role organisms play in the development of sedimentary facies. This study will
also evaluate similarities and differences existing
between bamer reefs and atolls of the Mexican Caribbean and Gulf of Mexico. Finally, data from this
study will allow comparisons to be made with results obtained from fringing reef environments at
Discovery Bay, Jamaica (Boss and Liddell 1987;
Liddell ,et al. 1987).
Methods
Study Locality
Field work was conducted along the Gulf and Caribbean coasts of the Yucatan Peninsula (Fig.. l).
Sites include the bamer reefs adjacent to Akumal
and Chemuyil on the Caribbean coast in NE Quintana Roo (20°38' N Lat, 87'2 1' W Long). Sampling
at Akumal was conducted at the Parque Ecologico
Akumal. Sampling at Playa Chemuyil was conducted off the southern end of the small bay. This
site is approximately 5 km south of Akumal.
Additional field work was conducted at the atolls
of Banco Chinchorro (18°47'-18023' N Lat,
87O14'-87O27' W Long) off southeastern Q. Roo
and at Alacranes (22'24'-22O27' N Lat, 89O32'89O48' W Long) in the southern Gulf of Mexico.
GULF OF MEXICO
Sampling
Fig. 1. Location of study sites along the Yucatan Peninsula, Mexico (also refer to Fig. 3 for profiles along the section lines shown).
Sampling was conducted along traverses oriented
perpendicular to the reef crest, at depth intervals
of 5 m and across a bathymetric range of 0-40 m.
Holocene sediments were collected via SCUBA by
shallow coring of unconsolidated material. At least
two sediment samples were taken at each depth
interval. Samples were collected from both sand
channels and reef lobes when sufficient sediment
was available from both environments.
Communities were sampled by underwater photography. Ten meter lines were randomly placed
parallel to the reef crest at each study depth and
the area beneath each 50 cm increment point was
photographed. Approximately 36 color transparencies were taken at each site. Each transparency
covered an area of approximately 0.25 m2. Photocensus data were processed by projecting transparencies at natural size onto a grid with 25 equally
Sedimentology and Community Structure of Reefs
spaced points and recording the identity of each
organism at each point (planar point intercept
method).
Results
Sediment Analysis
The major component of sediments at bamer reef
sites are highly comminuted fragments of coral
(3 1.3%-56.0%), plates of the calcareous green alga,
Halimeda (1.2%-3 1.1%) and coralline algae (2.8%20.4%). The remainder ofthe sediment is composed
of other taxonomic groups (Foraminifera, molluscs
and echinoderms; Tables 1-2). Sediments from
Gulf of Mexico and Caribbean atolls are also dominated by coral fragments (10.1%-64.2%), Halimeda (0.0%-60.3%), coralline algae (2.6%-23.8%)
and lesser amounts of other taxonomic groups (Tables 3-4). Relative proportions do differ somewhat
between the different reef types.
Constituent Particle Composition
Sediments were impregnated with casting resin,
thin-sectioned and point-counted (Tables 1-4).
Rarefaction analysis indicated that 250-300 points
were necessary to adequately describe the samples.
Gross textural characteristics were determined
by dry sieving between -2 and 4 phi, at 112 phi
intervals. Weight percent data were used to construct cumulative frequency curves and to determine mean grain size (Mz) and sorting (IGSD).
Carbonate/noncarbonate ratios for select samples were determined by acid digestion. Organic
and non-organic portions of the insoluble fractions
were determined by heating residues in an oven at
450" C for 8-12 hours, until a constant weight was
achieved, to destroy (ash) the organic fraction.
Texture
Due to page constraints, only a brief summary of
textural data is presented (refer to Novak 1992 for
further details). Sediments from coastal bamer
reefs have a mean grain size (Mz) in the medium
sand range, between 0.4-0.5 mm. Mz generally de-,
creases with depth over the shallow fore reef and
deep fore reef. Sediments are moderately to poorlysorted with a tendency to become progressively
more poorlylsorted with depth. Sediments collected
Cluster Analysis
Hierarchical cluster analysis was utilized to delineate associations between samples (Kovach 1990).
The Euclidean Distance coefficient and the Unweighted Pair-Group Method with Simple Arithmetic Averages (UPGMA) algorithm were employed in the analyses.
Table 1. Mean constituent composition (%) for sediments from Akumal, Mexico.
Back Reef
~ocation
Depth
Shallow Fore Reef
Deep Fore Reef
Beach
lm
3m
5m
10m
llm
20m
25m
30m
35m
40m
Sample Size (n)
2
2
6
2
2
2
2
2
2
2
2
Bottom Type
S
S
S
S
S
S
S
S
S
S
S
31.3
9.0
20.0
3.6
1.1
11.5
1.7
2.5
4.9
1.9
1.5
0.2
0.5
0.0
2.4
7.8
1.4
36.8
21.2
6.2
0.0
1.2
5.4
3.4
5.4
2.2
0.2
5.0
0.4
2.0
0.0
0.4
9.2
1.0
42.3
6.9
11.4
3.6
3.2
5.5
1.4
1.6
7.2
1.8
2.0
0.1
0.2
0.0
5.1
6.7
1.1
42.5
1.6
13.0
15.2
4.4
0.7
1.1
0.0
15.0
0.6
0.9
0.0
0.0
0.0
0.9
3.7
0.4
32.7
12.9
10.2
2.5
2.7
13.1
2.7
1.7
6.6
0.7
2.8
0.3
I .O
0.0
2.5
6.5
1.5
41.2
7.5
5.7
1.5
0.3
13.4
3.3
2.4
5.0
0.0
3.0
0.0
3.0
0.0
3.0
10.1
0.6
33.2
31.1
4.5
1.1
0.8
6.3
1.4
0.9
3.7
0.5
3.1
0.2
0.0
0.0
2.1
4.0
7.0
37.2
27.6
5.4
0.2
2.4
6.6
2.8
0.8
2.2
0.0
3.0
0.0
0.4
0.0
3.2
5.8
2.4
40.7
18.7
5.3
1.7
1.0
4.0
1.3
0.3
8.7
1.7
4.7
0.0
0.0
0.0
4.7
6.3
1.0
33.0
12.3
8.0
2.0
3.3
10.2
1.7
2.4
8.1
0.6
4.2
0.0
1.6
0.0
2.6
6.4
3.7
36.4
22.6
4.4
1.0
1.4
5.6
2.2
1.8
7.2
0.2
1.8
0.0
1.8
0.0
4.4
8.2
1.2
Constituent
Coral
Halimeda
Coralline Algae
Homoirema
Gypsina
Miliolina
Textulariina
Rotaliina
Bivalve
Gastropod
Echinoderm
Sponge Spicule
Gorgonian
Bryozoan
Pellet
Micritized Grain
Unidentified
Bottom type: R
=
Reef Framework; S
=
Unstable Sand.
Novak et al.
268
Table 2. Mean constituent composition (%) for sediments from Chemuyil, Mexico.
Back Reef
Location
Depth
Shallow Fore Reef
Deep Fore Reef
Beach
Im
Im
5m
8m
15m
14m
25m
30m
35m
Sample Size (n)
2
2
2
2
2
3
1
2
2
3
Bottom Type
S
S
S
S
S
S
R
S
S
S
Constituents
Coral
Halimeda
Coralline Algae
Homolrema
Gypsina
Miliolina
Textulariina
Rotaliina
Bivalve
Gastropod
Echinoderm
Gorgonian
Sponge Spicule
Bryozoan
Pellet
Cryptocrystalline
Unidentified
Bottom Type: R
=
Reef Framework; S
=
Unstable Sand.
Table 3. Mean constituent composition (%) for sediments from Banco Chinchorro, Mexico.
Location
Depth
Back Reef
Shallow Fore Reef
Deep Fore Reef
'
EL0.5m
EL4m
Im
5m
10m
15m
15m
22m
22m
28m
28m
Sample Size (n)
4
2
2
2
2
2
2
2
2
2
2
Bottom Type
S
S
S
S
S
S
R
S
R
S
R
Constituents
Coral
Halimeda
Coralline Algae
Homolrema
Gypsina
Miliolina
Textulariina
Rotaliina
Bivalve
Gastropod
Echinoderm
Sponge Spicule
Gorgonian
Bryozoan
Pellets
Cryptocrystalline
Unidentified
EL
=
East Lagoon. Bottom Type: R
=
Reef Framework; S
in reef grooves exhibit better sorting than sediments
collected on reef lobes (spurs). Mz for the atolls of
Alacranes and Chinchorro is 0.6 and 0.7 mm, respectively, which falls in the coarse sand size range.
=
Unstable Sand.
Fore reef sediments from the Banco Chinchorro
exhibit a pronounced decrease in mean grain size
with depth. Sediments from Alacranes exhibit a
similar trend, although with a much poorer corre-
Sedimentology and Community Structure of Reefs
269
lation with depth. Chinchorro sediments become
progressively poorer-sorted with depth, while sediments from Alacranes exhibit no discernable
depth-related variation in sorting.
boundary between shallow fore reef and deeper fore
reef facies typically occurs at a depth of 10-15 m.
Mineralogy
Only modest correlation occurs between mean
grain size or sorting and depth. Boss and Liddell
(1987) found that data gathered by traditional size
analysis techniques were only of limited value in
carbonate depositional environments. This is due,
in part, to the in-situ generation of sediment grains,
which is occumng at all depths on the reef. Inhomogeneities in the hydraulic characteristics of the various sediment constituents also makes interpretation of energy levels or sediment transport based
on clast size tenuous.
Although significant wave surge was observed
within the reef channels, the dominant process
within the area of wave turbulence appears to be
the winnowing of fines and an increase in sorting,
as indicated by the higher degree of sorting in shallow fore reef sediments. The compositional similarity between fore reef sediments collected on reef
lobes and in reef channels observed elsewhere (Boss
and Liddell 1987)was also found by our study. This
suggests an absence of sufficiently competent directional sand-transport mechanisms under modal
wind, wave and tide conditions.
Previous research at Jamaica indicated that variation in the abundance of major sediment con-
The calcium carbonate fraction of back reef sediments for the bamer reef sites averages 99.1%, with
insoluble organics averaging 0.7% and other nonorganic insolubles averaging 0.2%. Fore reef sediments contain an average calcium carbonate fraction of 99.6%, with insoluble organics averaging
0.4% and non-organic insolubles averaging < 0.1%.
The calcium carbonate fraction of back reef sediments from Caribbean and Gulf atolls averages
98.9%, with insoluble organics averaging 0.9% and
other non-organic insolubles averaging 0.2%. Fore
reef sediments contain an average calcium carbonate fraction of 99.5%, with insoluble organics averaging 0.1% and non-organic insolubles averaging
0.4%.
Quantitative Lithofacies
Q-mode cluster analysis of epireefal sediments produced the dendrograms illustrated in Figure 2A-D.
Three well-defined groupings are represented which
reflect variation in the composition of the back reef,
shallow fore reef and deep fore reef facies. The
Discussion
Table 4. Mean constituent composition (%) for sediments from Alacranes, Mexico.
Back Reef
Location
Depth
Shallow Fore Reef
Deep Fore Reef
lm
2m
4m
N3m
S7m
NlOm
Sllm
S12m
S17m
N17m
Sample Size (n)
8
2
2
2
2.
2
2
2
2
2
Bottom Type
Constituents
Coral
Halimeda
Coralline Algae
Homotrema
Gypsina
Miliolina
Textulariina
Rotaliina
Bivalve
Gastropod
Echinoderm
Gorgonian
Sponge Spicule
Bryozoan
Pellet
Cryptocrystalline
Unidentified
S
Bottom Type: S
=
,
23.1
43.5
11.0
0.4
0.5
7.9
1.8
0.3
7.6
2.6
0.4
0.0
0.1
0.2
0.2
0.1
1.O
Unstable Sand. N
=
North transect; S
=
South transect.
Novak et al.
ID'
Fig. 2A-D. Cluster dendrograms based upon analysis of constituent particle composition and illustrating the relationship
between sediment composition and reef environment. Numbers indicate sample depths (m); R = reef; S = sand channel
or sandy lagoon; L = lagoon or back reef; WL = western lagoon; EL = eastern lagoon; BRC = back reef adjacent to reef
crest; BCH = beach).
stituents was directly related to the presence of sediment-producing organisms living on the reef at a
particular depth interval (Boss and Liddell 1987).
Coral clearly constitutes the largest portion of the
sediment at our Mexican reef sites and is also found
to be the dominant member of the biotic community. The percentages of coral, Halimeda plates
and coralline algae present in sediment at different
depths is generally wellcorrelated with the abundance of the living organisms on the reef (Fig. 3AD). Sediment constituent data from a patch reef in
the northern Belize lagoon near the border with Q.
Roo (Mazzullo et al. 1992) is very similar to that
from our sites.
The carbonate mineralogical content of the sediments from all depths sampled show relatively little depth-related variation. Sediment insoluble content (largely organics) reaches the highest level in
the back reef sediments. Atoll reef sediments contain slightly higher percentages of non-carbonate
material than bamer reef sediments, which is most
likely attributable to the large quantity of algae
growing in the back reef environments of the atolls.
Q-mode cluster analysis of constituent particle
data demonstrates that Holocene epireefal sediments display variability in biotic composition
which is related to the community structure of the
different reef zones. The close clustering of sediment samples from sand channels and reef interstices at similar depths further supports that little
diferential ofieef transport is occumng on the fore
reef to a depth of approximately 40m. Initial cluster
analysis of sediment constituent data produced
dendrograms which grouped some shallow fore reef
samples with back reef samples. An examination of
the constituent composition of these sediments and
the percentages of individual grain types within the
samples indicated that a high percentage of Halimeda was present at both locations. Further inspection of Halimeda grain types allowed the de-
Sedimentology and Community Structure of Reefs
liillirrn~ar-a
COR~LL~HEALWEE.-
7
COPALL E A L C A E
-
Fig. 3A-D. Splndle diagrams illustrating abundance of major sedlment constituents and relative proportions of major sedlment-producing biotic groups. Sedlment sampllng s ~ t e and
s depths indicated on generalized reef profiles at top.
-?
i'
+'
Z
"..
lineation of deep- and shallow-water Halimeda
suites, based on Halimeda morphology. Similar
measures proved useful in previous studies (Boss
and Liddell 1987). The continued grouping of some
back reef samples with fore reef clusters is believed
to be a result of sediment input from patch reefs
located in the lagoonal environment.
Separation of shallow fore reef and deep fore reef
facies occurred at a depth of approximately 10-1 5
m. Inconsistent grouping of some samples from this
interval indicates the gradational nature of the fac i e ~transitions. Cluster analysis of combined data
from Akumal and Chemuyil produced dendrograms which were strikingly similar to those of the
individual sites. Similar depths were clustered together to yield the same three facies observed in
cluster runs for the individual sites (ie. back reef,
shallow fore reef and deep fore reef facies). Further,
combined data from Chinchorro and Alacranes
yielded dendrograms resembling those of the individual sites. Cluster analysis of sediment constituent data with all four localities combined produced cluster d e n d r o g r a m s with m u c h less
distinctive zonations. This indicates that coastal
banier reef sediments differ significantly from the
atoll reef sediments, although similar bathymetric
breaks between lithofacies occur in both geomorphic settings.
Conclusions
Quantitative analysis of Holocene carbonate sediments from Mexican reefs in the Caribbean and
Gulf of Mexico allows the delineation of reef zonations based on constituent composition. Q-mode
cluster analysis of epireefal sediments enables the
separation of lagoon, shallow reef ((10-1 5 m) and
deep reef (> 10- 15 m) lithofacies. These groupings
were maintained even when constituent data from
different locations (e.g. Akumal and Chemuyil, Alacranes and Chinchorro) were combined. However,
cluster analysis failed to produce clear associations
when data from bamer reefs and atolls were combined. The similarity of the distributions of sediment allochems and sediment-producing organisms
suggest that reef community structure, at least for
calcifying organisms, is potentially preservable.
Acknowledgements. Funding for this research was provided
by Fulbright and Utah State University faculty research grants
to WDL. We wish to acknowledge the following organizations
for additional financial and logistical support during field work:
The Armada de Mexico, CINVESTAV (Merida, Yucatan) and
the (Fishermen's) Cooperativa Felipe Angeles #487 (Chetumal,
Q. Roo). We would also like to thank the following individuals
for their assistance in the field: Arturo Chable, Juan Chin,
Mauricio Garduno and Enrique Rodriguez.
272
References
Boss SK, Liddell WD (1987) Patterns of sediment composition of fringing reef facies. Sedimentology 34:77-87
Chavez EA, Hidalgo E (1984) Spatial structure of benthic
communities of Banco Chinchorro, Mexico: Advances in
Reef Sci Abstr of Papers, Joint Mtg of the Atlantic Reef
Comm and the Intl Soc Reef Stud 19-20
Chavez EA, Hidalgo E, Izaguirre MA (1985) A comparative
analysis of Yucatan coral reefs: Proc Fifth Int Coral Reef
Cong, Tahiti 3:4 10-4 16
Ekdale AA (1974) Marine molluscs from shallow-water env~ronments (0-60m) OK the northeast Yucatan coast,
Mexico. Bull Mar Sci 24:638-668
Farrell TM, D'Elia CF, Lubbers L, Pastor LI (1983) Hermatypic coral diversity and reef zonation at Cayos Arcas,
Campeche Bank, Gulf of Mexico. Atoll Res Bull 270: 1-7
Fenner DP (1988) Some leeward reefs and corals of Cozumel,
Mexico. Bull Mar Sci 42:133-144
Folk RL (1967a) Carbonate sediments of Isla Mujeres, Quintana Roo Mexico and Vicinity. 1n:Weide AE (ed) Yucatan
Field Trip Guide Book, Second Edition (Geol Soc Am,
pp. 100-123)
Folk RL (1967b) Sand cays of Alacran Reef, Yucatan, Mexico:
Morphology. J Geol 75:412-437
Folk RL, Robles R (1964) Carbonate sands of Isla Perez,
Alacran Reef Complex, Yucatan. J Geol 72255-292
Hoskins CM (1966) Coral pinnacle sedimentation, Alacran
Reef lagoon, Mexico. J Sed Petr 36:1058-1074
Jordan ED (1979a) An analysis of a gorgonian community in
a reef calcareous platform of the Caribbean coast of Mexico. Ann Inst de Cien del Mar y Limnol del Univ Nac
Aut de Mex 6:87-96
Jordan ED (1979b) Structure and composition of coral reefs
in the northeast region ofthe Yucatan Peninsula, Mexico.
Ann Inst de Cien del Mar y Limnol del Univ Nac Aut de
Mex 6:69-86
Jordan ED (1989) Gorgonian community structure and reef
zonation patterns on Yucatan coral reefs. Bull Mar Sci
45:678-696
Jordan ED, Merino M, Moreno 0 , Martin E (1981) Community structure of coral reefs in the Mexican Caribbean.
Proc Fourth Int Coral Reef Symp, Manilla 2303-308
Novak et al.
Jordan ED, Martin E (1987) Chinchorro: Morphology and
composition of a Caribbean atoll. Atoll Res Bull 310:l20
Kornicker LS, Boyd DW (1962) Shallow-water geology and
environments of Alacran Reef complex, Campeche Bank,
Mexico. Amer Assn Petr Geol Bull 46:640-673
Kovach WL (1990) M.V.S.P. Plus Version 2.0a. A multivariate statistical package for IBM PC's and compatibles.
Aberystwth, Wales
Liddell WD, Boss SK, Nelson CV, Martin RE (1987) Sedimentological and foraminifera1 characterization of shelf
and slope environments (I- 234m), North Jamaica. Roc
Third Symp Geology Bahamas 91-98
Liddell WD, Ohlhorst SL (1988) Comparison of western Atlantic coral reef communities. Proc Sixth Int Coral Reef
Symp 3:281-285
Logan BW (1969) Carbonate Sediments and Reefs, Yucatan
Shelf, Mexico. Amer Assoc Petr Geol, Mem 11, Part
1:129-198
Macintyre IG, Graus RR, Reinthal PN, Littler MM, Littler
DS (1986) The bamer reef sediment apron:Tobacco Reef,
Belize. Coral Reefs 6: 1-12
Mazzulo SJ, Anderson-Underwood KE,Burke CD, BischoK
WD (1992) Holocene coral patch reef ecology and sedimentary architecture, Northern Belize, Central America.
Palaios 7591-601
Novak MJ (1992) Sedimentology and Community Structure
of Reefs of the Yucatan Peninsula, Mexico. Unpub Master's Thesis, Utah State Univ, 96 pp
Rutzler K, MacIntyre IG (1982) The habitat distribution and
community structuie of the bamer reef complex at Came
Bow Cay, Belize. In: Rutzler K, MacIntyre IG (eds)The
Atlantic Bamer Reef Ecosystem at Came Bow Cay, Belize: Structure and Communities (Smithsonian Inst,
Washington D.C. 9-45)
Spaw RH (1978) Late Pleistocene carbonate bank deposition;
Cozumel Island, Quintana Roo, Mexico. Gulf Coast Assoc
Geol Soc, Trans 28:601-619
Wantland KF, Pusey WC (1975) Belize She1f:Carbonate Sediments, Clastic Sediments, and Ecology. Amer Assoc Petr
Geol Stud Geol 2599 pp