Coral Reefs

I n this set of notes, coral reefs are given their own chapter. These natural systems are of particular interest to marine
geology programs lying between 20o north and south of the equator. If you have no particular interest in reefs, skip this
chapter. Carbonate buildups are treated in the continental margin presentation.

Coral Reef Concepts

What is a reef

W •
e start with the elements upon which there is general agreement. First,

coral reefs owe their origins to the biological secretion of calcium carbonate by living organisms,
• they are rigid structures that stand above the surrounding sea floor,
• they exert some control on local oceanographic processes.

The primary disagreement concerns the concept of framework in reef building. The reef-forming organisms generally
possess massive skeletons that are the building blocks of the reef. Many workers hold that in a "true reef" these are
organized into an in-place and interlocking meshwork that provides the rigidity of the reef. Conversely, any feature
lacking this interconnecting framework is not a reef.

The problem is that when ancient reefs are examined, many of them are not the organized structures that are central to this
model. They have recognizable organisms, but these are loosely packed and often "floating" in a detrital matrix. In these
cases it is assumed that either:

• no "real reef" existed at these sites, or

• the reef core has somehow been removed by erosion and all that remains are piles of debris that were peripheral to
the reef, but not actually a part of it.

This pattern has occurred with sufficient regularity to raise questions about how (and why) modern and ancient reefs
might be fundamentally different from one another. Reef organisms have certainly evolved through time; perhaps this has
changed the ways in which reefs have functioned. Sea-water chemistry and the shape of ocean basins have likewise
changed - many ancient reefs existed in broad, shallow seaways and intracontinental seas, in contrast to the more-exposed
and steep platform margins of today. If all these factors have conspired to fundamentally change reefs over the span of
geologic time, then modern reefs become very poor models for their ancient counterparts. We will show that this is not the
case. In an analogy by Ginsburg, reefs are likened to Shakespearean plays in which an ever-changing cast acts out a
timeless plot.

Reef classification

F rom a geological perspective, only biological construction and topographic relief are directly observable in the rock
record. Accordingly, terms have been proposed for biologically produced deposits that, in outcrop, are moundlike
(bioherms) or else occur as discontinuous sheets or lenses parallel to the beds above and below . Dunham proposed a
genetic terminology to distinguish between reefs similar to those constructed by modern corals (ecologic reefs) and those
that are identifiable only as "reef-like" structures in the rock record (stratigraphic reefs). The former are presumed to be
dominated by in-place and interlocking organisms, as in Wilson's "Organic Framework Reefs." The latter might be "true
reefs," but they could also be the product of differential compaction along a bed or the stacking of several biostromes into
a feature that only appears to have had relief at the time of deposition.

To circumvent some of these interpretive problems, we can use the more-generic and descriptive term "buildup,"
describing modern coral reefs as "encrusted skeletal buildups." All other deposits are relegated to some other category that

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implies less internal organization or rigidity (i.e. "loose skeletal buildup" --> detritus near to, but not within a reef; "sorted,
abraded skeletal buildup" --> a hydromechanical deposit of skeletal debris, probably located some distance from the reef).

With the development of lightweight drilling systems, the scientific community gained access to the interior of modern
reefs. With each new set of cores, it has become increasingly apparent that the interior of modern coral reefs are
dominated not by in-place and interlocking framework, but rather by variable assemblages of both whole and broken
corals mixed in with sediment and debris derived from their breakdown. The relative abundance of recognizable coral
recovered from cores of modern reefs is less than 20%. The other material in Caribbean reefs is sediment and open voids.
These data are drawn from reefs occurring in several different oceanographic settings, and show that recognizable coral
comprises only a small part of the reef mass. If we consider that only a portion of that recovered coral is in place, then the
importance of in-place and interlocking framework is put into a much different perspective.

This is not to say that interlocking framework never occurs nor that it is impossible for a single reef to be constructed
largely of in-place material. However, most of what anyone would agree are true "reefs" as they swim over them are not
organized collections of in-place and interlocking corals. In extreme cases, they are disorganized "garbage piles", only
indirectly controlled by the original pattern of corals. Most often, they lie somewhere in-between along a continuum.
Their rigidity is only partly derived from interlocking framework; it is equally the result of broken bits falling into stable
positions, as well as the secondary encrustation and cementation that bind the entire mess together.

The simplest way to integrate these emerging concepts into existing models would be to expand the limits of "true reefs"
within the classifications of Heckel or Wilson. This approach could lead to considerable confusion, however, given
traditional views about the importance of in-place framework in reef development. This necessitates the distasteful
exercise of constructing yet another reef classification.

Carbonate buildups can be classified along a continuum from those dominated by in-place skeletons to loose piles of
hydromechanically deposited sediment. Any classification of ancient reefs viewed in core or outcrop should be tied to the
internal character of modern reefs. To reinforce the point that the elements contained within a "reef" need not be either in
place or interlocking, we will use the term framework element to refer to recognizable members of the constructor guild
(including encrusting algae where it dominates the fabric) that remain within the specific environment in which they lived.
Based on the above discussion, the primary criteria for classification as a reef are:

• It is constructed of material of biological origin

• It is a rigid structure, owing to either interlocked and in place framework elements or of reworked framework
elements bound together by secondary encrustation or cementation

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 • It stands topographically above the surrounding seascape and, therefore, exerts at least local control on
oceanographic processes
• The majority of the framework elements were formed in an environment similar to the one in which they were
ultimately deposited.

As an example, we will consider the margin of Salt River submarine canyon on St. Croix in the Caribbean Sea . The reef
is comprised of several genera of branching, massive and platy corals, all bound together by secondary encrusters
(primarily coralline algae) and submarine cements. Many of the individual framework elements have been so reworked
that their origins are difficult to infer, except in thin section. Also, the seemingly random pattern of dates between the
youngest parts of the reef and its surface. reflects the constant slumping and redeposition of individual corals and large
blocks throughout its accretionary history. There is virtually no original framework that is demonstrably in place within
the reef.

Similarly, a shelf-edge reef off southwestern Puerto Rico is comprised largely of toppled and encrusted Acropora palmata
branches. The pattern of radiocarbon dates within this reef similarly implies considerable reworking of the coral colonies
that comprise the structure. Despite this continual reorganization, however, individual framework elements fall into an
organized pattern that reflects shoaling of the reef through time (i.e. the environment of deposition is essentially the same
as that in which the organisms lived). It is likely that many, if not most, modern reefs contain significant quantities of

3
reworked materials within their interior and that reefs dominated by the in-place and interlocking framework of traditional
models are in fact quite rare.

These ideas about reefs can be integrated into a general classification of carbonate buildups . "Reefs" can be comprised of
mostly in-place skeletal material (Primary Framework Reefs) or else can be dominated by reworked "framework
elements" that are held together in a rigid structure only by cementation or encrustation (Secondary Framework Reefs). In
the latter, the surface may be veneered by in-place organisms, but the internal fabric will likely seem chaotic, with the
individual framework elements mixed or "floating" within a matrix of sediment, cement or smaller detritus. This is much
more similar to "reefs" that dominate the fossil record.

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The best, and perhaps only examples of primary framework reefs are the massive algal ridges that occur along high-
energy margins in both the Caribbean and Pacific. Most of the Caribbean examples, however, have a nucleus of reworked
Acorapora palmata and head corals upon which the ultimate algal cap formed. The classification as primary or secondary-
framework-reefs thus becomes a matter of which part of the reef you are looking at. Furthermore, the in-place elements of
these features are comprised mostly of organisms normally relegated to binding the reef, not the framework constructors.

Most modern reefs fall along the continuum between the end-members of the reef axis in the classification figure, and are
therefore secondary framework reefs. The debris that comprises a large part of the reef's internal fabric is derived from the
biological breakdown of the larger framework elements by organisms seeking either food or shelter (the bioeroders).
Contrary to common opinion, a significant proportion of this material may be fine-grained, as reef framework also serves
as a baffle, trapping fine-grained material that is produced within the reef. The rigid structure of the reef serves to isolate
this material from the high wave energy that dominates at the surface.

Hydromechanical buildups are constructed by sediments that have been moved by currents from their point of origin to a
site more suitable for deposition. In this regard, they behave much like sand bars and spits described in siliclastic systems;
their location and size are controlled by sediment supply and local current patterns.

Some hydromechanical buildups owe their existence to the baffling action of upright organisms. As such, their occurrence
is largely related to the distribution of organisms that act to disrupt water flow and trap sediment. These baffled buildups
are intermediate between "reefs" and hydromechanical buildups, and reflect the mixed nature of the factors responsible
(currents and organisms). Examples include the mud mounds in Florida Bay, which are built up around seagrass.
Epibionts that grow on the grass blades and the upright calcareous algae that typically occupy grassy environments (i.e.
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Halimeda, Penecillis) contribute to the sediment budget of the features along with sediment moved in from elsewhere in
the bay. The baffling action of grass blades traps sediment that is typically finer than that along the adjacent bottom. Thus,
baffled buildups are often finer-grained than the surrounding benthos. Porites tolerates a high-sediment environment and
is part of the biomass.

Related in their process of formation are

the stromatolites that span the geologic
time scale from the Paleozoic to the
present. These features are developed by
cyanobacteria that form mats. The mats in
turn trap sediment. In the Paleozoic and
Mesozoic, stromatolites formed across a
wide morphologic spectrum from flat-lying
or gently undulating beds to domal or
columnar features of considerable relief.
Occurring in a transitional zone of our
classification, stromatolitic mounds are
generally considered as specialized baffled
buildups.

Paleozoic stromatolites dominated the

tropical seascape before the evolution of
herbivorous grazers. Today, stromatolites
are rare and exist only in a few areas where
grazers are discouraged by some outside
physical factor. For example, meter-high
stromatolites in the northern Bahamas are
alternately buried and exhumed by
carbonate sand bodies in an area of active
bedform migration.

Off-reef buildups are special features that

owe their origin to gravitational transport of
sediment and debris from the reef to some
downslope site of deposition. Usually
formed along steep escarpments, off-reef
buildups form near the base of the slope.
Unlike sedimentary fans found at the distal
ends of terrestrial fed submarine canyons,
off-reef buildups usually occur as broad
aprons along the edge of the basin. The
depth difference or distance between the
reef and an off-reef buildup depends upon
the magnitude of the underlying topography. As an example, sands found near the base of the northern insular slope of St.
Croix contain 97% shallow-water material, mostly derived from the reefs. Hubbard, et al., 1981 Water depth is roughly
2,500 meters. In most areas, the difference is much smaller.

The type of reef formed by a particular group of organisms need not remain constant through time. For example, the
rudists, large clams that dominated reefs during the Cretaceous, played very different roles in reef formation throughout
their evolution. The earliest rudists sat either on or in the bottom, exerting control more like bafflers. In some instances,
rudists may have been passive inhabitants of the bottom. Through time, rudists progressively increased both their colonial
affinities and their ability to produce an integrated and cemented structure. This would have been more analogous to
oyster reefs of today. At that point, they constituted more of a traditional framework element and may have even produced
primary framework reefs in some instances.

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Groups of organisms can produce varied buildups across either space or time. This is further complicated by the evolution
of reef dwellers and builders and the increasing pitfalls of taxonomic uniformitarianism as we go back further in the fossil
record. Nevertheless, by understanding the likely associations between various groups of organisms, the effects that they
can have on local depositional processes, and the controls that can be exerted by underlying physical factors, we stand the
best chance of constructing a realistic classification that adequately deals with both modern carbonate buildups and their
fossil counterparts. If the reader comes away with nothing else, the following two ideas should be retained:

• True "reefs" need not be comprised of in-place and interlocking framework

• Process and not product are the common denominator that we seek in linking modern carbonate buildups to
examples in the rock record

Reef types

V arious criteria have been used to classify reefs; the most accepted approach is morphological grouping. The shape
and location of reefs are controlled by the bottom topography upon which they formed, interactions among the
resident biota, and physical processes. Darwin discussed three main types of reefs - barrier reefs, fringing reefs and atolls -
still part of most classifications today. Fringing reefs occur adjacent to land with little or no separation from shore. A low
input of terrigenous sediment is important, and the best-developed fringing reefs occur off shorelines where rainfall is
low, there is little relief, or else the hillsides are stabilized by heavy vegetation. In recent years, clear cutting of forests and
poor land management have impacted fringing reefs more than any other type. Barrier reefs are separated from the
shoreline by a moderately deep (usually) body of water - the lagoon. The reef may form at the shelf edge, or it may be
located more inshore, usually localized on an antecedent break in slope.

Atolls are roughly circular in plan with a central lagoon that contains no significant land mass. The central lagoon is often
deep (less than 25 m), but this is not a prerequisite. If land does exist, it sits atop a part of the encircling reef and is
comprised solely of carbonate material derived from the reef. As originally defined for Pacific reefs, the term implies a
specific genetic origin around a volcanic island. Caribbean and Atlantic atoll-like reefs are not of this type, and tend to
form around isolated highs formed by local tectonics.
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If we decoupling reef type from tectonic history (i.e. a barrier reef is a morphologic entity, separate from its tectonic
regime) Darwin’s linking of reef type solely to subsidence is not a problem. To distinguish barrier from fringing reef, a
limit must be set on lagoonal dimensions in fringing vs. barrier reefs. Implied in Darwin's definition is the idea that the
lagoon is sufficiently large to permit open circulation behind the reef. The reef serves as a "barrier" that clearly separates
lagoonal processes from those of the open ocean. Based on this, we pragmatically make the split between barrier and
fringing reefs at a point when the lagoon reaches 500 m in width and 5 m in depth. In a natural setting, and in the absence
of significant upland clearing of vegetation, a lagoon of this magnitude can substantially isolate the reef from direct
impact by terrestrial runoff. Furthermore, circulation within the lagoon is distinctly removed from that of the open ocean
beyond.

Patch reefs are smaller features, roughly equant in plan

view. While they have generally reached sea level, this is
not necessarily so. Usually, patch reefs occur within the lagoon behind the barrier or atoll rim. On occasion, however, they
can occur on the open shelf as pinnacles. Modern examples of exposed (i.e. non-lagoonal) patch reefs occur off the north
coast of St. Croix in the Caribbean Sea. Numerous small reefs, 10-20 m across, rise out of 10-15 m of water. Their fabric
of broken and piled-up coral branches has led to the local name "haystacks".

Submerged shelf-edge reefs are Caribbean platform

margins that presently sit in water depths greater than
10-15 meters after being flooded by rising sea level
6,000 - 10,000 years ago. Since then, they have not been
able to offset the effects of ever-deepening water, and
many of them have been left behind. While coral and
other calcifying organisms occur along most of these
margins, they are not producing carbonate at a rate
sufficient for the reef to "catch up" with sea level.

Equally problematic are reefs that occur on wider

shelves (more than 5 km), and fall between the criteria
for either barrier or patch reefs. They are similar to patch
reefs in shape, but they are usually larger, more linear,
and are aligned in roughly shore-parallel. They exist near sea level and, in some instances, have emerged to form islands.
The sediments behind the reef (landward) are similar to those seaward reflecting the absence of lagoonal conditions.
Because they usually occur along either insular or continental shelves, they are classified as shelf reefs.

The nature of shelf reefs changes from shore to the shelf edge. More-seaward reefs are exposed to higher wave energy.
Those closer to shore come more under the influence of terrestrial sedimentation. For example, on the southern coast of
Puerto Rico, the inner-shelf reefs are often subjected to fine-grained sediments derived from the adjacent hillsides . As a
result, they are mostly mud mounds with scattered corals. In some instances, they have been stabilized by mangroves and
have built small islands. The mid-shelf reefs are subject to the effects of open-ocean circulation and more wave action.
Accordingly, coral cover is higher and the benthic-community structure is more complex.

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Morphology of Caribbean reefs

T he morphologic terminology developed for Discovery

Bay, Jamaica is often referred to as "typical" for
Caribbean reefs . While variations occur from place to
place, this general scheme is a reasonable place to start.

Shallow Reef

T he crest of the main reef is generally emergent in

Pacific reefs at low tide, but may be below the surface
in Caribbean reefs. The seaward edge of the reef crest
takes the brunt of the incoming wave energy. The reef can
reduce incoming wave energy by up to 97%. As waves
break, water is washed across the reef crest and into the
lagoon, driving lagoonal circulation.

Because of the modification of wave forces across the

reef crest, the backreef is an environment of totally
different physical processes, ecology and sediment
characteristics. Sediments and rubble from the reef
crest are dumped behind the crest, widening the
backreef flat through time. The outer reefs of the
Great Barrier Reef have been at sea level for nearly
6,000 years. Hence, the wide backreef flats often
exhibit distinctive front-to-back zonation. By
comparison, Caribbean reef flats have only recently
reached sea level and are narrower. While zonation is
9
less pronounced, there is a general transition from branching corals and the hydrozoan Millepora near the front of the crest
to sand flats and Thalassia landward. The shallow back reef may have a shallow Porites reef flat immediately behind the
crest and numerous small patch reefs in a sand apron. The corals are generally well adapted to the high levels of
sedimentation to which they are regularly subjected. In the Caribbean, the dominant corals include Porites porites and
several head corals, especially Montastrea annularis, Porites asteroides and species of Diploria.

Forereef

T he forereef extends seaward and downward from the reef crest. It is the most complex of the reef zones, owing to the
large depth gradient over which it occurs. In many areas, the forereef is organized into a set of en-echelon reef
promontories and sand channels, termed "spur-and-groove" topography.

Spur-and-groove is common in both

modern and ancient reefs. The term was
originally coined from Indo-Pacific
examples formed by erosion of the algal
rim just below the surf zone. More
recently, examples have been described
from the Caribbean that appear to be
the result of accretion by Acropora
palmata under the influence of strong
wave surge. Both the coral branches
and the intervening sand channels are
oriented parallel to the dominant wave-
approach direction. The channels serve
as primary conduits for sediment export
from the reef..

Forereef Slope and Deep Forereef

T he forereef slope is the least consistent of any of the reef zones, in either its occurrence or character. At many sites, it
is totally absent and the forereef drops from shallow water to oceanic depths. Near Discovery Bay, it occurs as a
gently sloping sand plain, separating the shallower reef from its deeper counterpart by several tens of meters. A nearly
identical feature occurs along the southern margin of Little Bahama Bank and many other Caribbean sites. Its persistent
depth throughout the region (ca. 30-35 m) may imply control by an underlying terrace cut at a lower stage of sea level.

Where a forereef slope is present, the deep forereef usually occurs as a well-defined ridge near the platform margin.
Otherwise, it is simply a down-dip extension of the forereef. When occurring separate from the shallower reef zones, the
location of the deep forereef is probably controlled by both the break in slope and the existence of an antecedent high left
by a previous reef. The character of the reef surface is often similar to the spur-and-groove topography except that the
scale of both the reef promontories and the intervening channels is generally larger.

The Reef Wall

P erhaps the most dramatic feature of the deep forereef is the "reef wall". At depths ranging from 50 to 85 meters around
the Caribbean, the forereef slope rolls over to a vertical or, in some places, overhanging precipice . The role of active
accretion at this depth is not well understood, owing to its remoteness. Along the high-energy margin of the Great Barrier
Reef, coral cover is apparently limited to a very thin veneer over the antecedent Pleistocene reef front. Along the front of
the Belize barrier reef, several episodes of reef-wall accretion have probably occurred. On the north coast of St. Croix,
accumulation of reefal debris has resulted in significant (greater than 26 m) lateral accretion in water depths of up to 30 m,
primarily the result of repeated slumps stacked one in front of another.

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Caribbean reef zonation

E arly discussions of reef zonation were based on Caribbean reefs and, therefore, reflect their species composition. A
profile across a "typical" Caribbean reef shows both morphological and species zonation. The following is a
generalized description of species patterns keyed to that profile.

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The Acropora palmata Zone

A long the front of moderate- to high-

energy reefs, the dominance of
Acropora palmata to depths of 5-10 m is
primarily a response to wave energy.
While, not as strong as head corals,
Acropora palmata orients its branches to
minimize tensional loading - the higher
the current flow, the lower is the profile
presented by the branches of the colony.
Acropora palmata is capable of rapid
growth, which elevates the colony above
the traction carpet of shifting sediment
that, can "sandblast" polyps sitting closer
to the bed. Because Acropora palmata is extremely sensitive to sedimentation and has no physiological mechanism for
sediment removal, wave-induced surge performs this important function for the coral.

The Massive Coral Zone

U sually starting at depths of 5 - 10 m, head corals increase in importance; notable among these are Montastrea
Annularis, Montastrea cavernosa, Porites astreoides, Colpophyllia natans and species of Diploria. In some areas,
the branching coral Acropora cervicornis can occur as a narrow belt between this and the shallower Acropora palmata
zone. Both their depth of occurrence and their massive nature make head corals in this zone more resistant to periodic
disturbance by storms. As a result, this zone is often the best represented in cores from Holocene reefs throughout the
Caribbean region.

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Platy Coral Zone

A t depth, most corals are platelike , an

adaptation that concentrates the photo-
receptive algae contained within their
tissues along the upper surface of the
colony. In shallow water, this fragile shape
is disadvantageous from a structural
standpoint. In deeper water, however,
physical breakage is low and the need to
gather light dominates. The primary species
at depth vary from place to place, but
usually include members of the genus
Agaricia and flattened colonies of
Montastrea annularis.

Coral Biology
C oral growth is only part of a complex equation that describes the history of modern reefs. Equally important are the
subsequent grinding away of the reef by organisms seeking food or shelter, and the redistribution of the resulting
sediment. We will focus on corals because they contribute the greatest volume to the "carbonate budget" of modern reefs.
We ignore other organisms not because they are unimportant, but rather to keep the discussion a bit simpler.

Fagerstrom proposed that we consider the relative roles of the organisms involved in the accretionary process of reef
building. He identified five basic "guilds" into which reef organisms can be placed:

• constructors, • bafflers,
• binders, • destroyers and
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 • dwellers.

The constructors provide the building blocks of the reef, whatever their ultimate fate; they can be overgrown and bound
together by algae, forams and other members of the binder guild.

Bafflers are those organisms that affect accretion by interrupting the flow of water, thereby encouraging sedimentation.
Destroyers include grazers and borers that break down the primary framework in various ways. Dwellers are usually
passive inhabitants that contribute to the ecologic diversity of the reef but often have little to do with the actual
accretionary process, except to help "fill in the spaces" within the reef interior.

Coral growth - the constructive stage

T he most diverse and abundant components of modern reefs are the stony corals. In effect they are both plant and
animal. The individual polyp is an invertebrate of the phylum Cnidaria. The body is soft and tubular, with an oral
opening surrounded by six (or multiples
of six) tentacles. At night, these are
extended to entrap plankton . The body
wall consists of three layers, the
innermost of which (gastrodermis)
contains photosynthetic dinoflagellates
called zooxanthellae . The details of the
relationship between these symbionts
and the coral polyp are not completely
understood, but their importance in
meeting the coral's metabolic demands
is well established. Nearly 90% of the
carbon fixed by zooxanthellae is
released to the coral host primarily as
glycerol. Nitrogen and phosphorous
derived from captured plankton are
shared between symbiont and host. The
contribution made to the calcification
process is of pivotal importance to this
discussion. While this link is well
known, the precise pathways along
which it occurs remain the subject of
considerable discussion.

Corals vary in their dependence upon photosynthesis. Those with larger polyps are well adapted to the active capture of
plankton from the water column. Porter, 1976 Corals whose polyps have high surface areas relative to their volume are
morphologically adapted to more efficient light reception. This relative dependence on photosynthesis plays perhaps the
greatest role in determining reef zonation and depth-related patterns of calcification in modern coral reefs.

Terminology

N ot all modern corals possess zooxanthellae. All deep-water corals, and some in shallow water rely solely on a regular
supply of plankton to meet their energy demands. The two groups (those with and without algal symbionts) differ
greatly in their ability to produce calcium carbonate and, therefore, to build reefs.

We prefer to use the terms zooxanthellate and non-zooxanthellate to distinguish between the two groups. The terms
hermatypic (mound-building) and ahermatypic are often used, and have unfortunately become synonymous with
zooxanthellate and non-zooxanthellate to many authors. This is a critical error, because many deep-water corals that are
totally lacking in zooxanthellae, are quite capable of building mounds. Conversely, there are corals that are part of the
mound-building process and do not contain endosymbionts. We, therefore, return the terms to their root origins. Any coral
that has built reef-like topography (i.e. a bioherm) is considered as hermatypic. Regardless of its ability to build such a
14
structure, it is histologically classified as zooxanthellate or non-zooxanthellate (alternately, azooxanthellate) based on the
presence or absence of algal symbionts.
Carbonate Production by Corals

T he world ocean is saturated with the three major polymorphs of calcium carbonate (aragonite, calcite and magnesian-
calcite). Yet, free precipitation in seawater is rare. As a result, biologically mediated carbonate production by corals is
the most important contributor to the carbonate budget of modern reefs. Coral polyps absorb calcium ions from seawater
and move them to the site of calcification, where they are deposited as aragonite. Minute crystals are formed in the outer
cell layer of the polyp and pass to the skeleton where they act as nuclei for continued growth.

The chemical process is a complicated one in which the building blocks for calcium carbonate can be provided from
several sources. The most important processes in the marine system can be described by the formula:

Ca++ + 2HCO3- <=> CaCO3 + CO2 + H2O

The vigor with which aragonite will form is thus related to the abundance of free calcium (Ca++) and HCO3-. The addition
of CO2 to water ultimately makes both of these available through the following process:

CO2 + H2O <=> H2CO3 <=> H+ + HCO3- + Ca++

Free H+, left over from the calcification process lowers the pH (i.e. makes the solution acidic).
Conversely, dissolution of carbonate will increase pH. The ability of various organisms to
regulate pH within their tissues, and drive the reaction toward the precipitation of aragonite,
may be an important factor in biologically-mediated calcification.

Marine organisms secrete all three calcium carbonate polymorphs. Aragonite forms a crystal
with a more open structure (orthorhombic) and, therefore, is more susceptible to chemical
breakdown than calcite and magnesian calcite with stronger crystal bonds (hexagonal crystals).
The only difference between the latter two is the inclusion of magnesium as an impurity within
the crystal lattice (mg-calcite is defined as any calcite containing greater than 4 mole-percent
magnesium).

Coral Growth Rate

T he growth rate of modern corals varies among species, but in general, declines in deeper
water. The highest growth rates are associated with shallow-water, branching corals (i.e.
Acropora palmata, Acropora cervicornis), followed by "finger" corals (i.e. Porites porites),
"head" corals (i.e. Montastrea annularis), and finally platy corals (i.e. Agaricia spp). Within
species, growth rate responds to a variety of factors including temperature, sedimentation,
nutrient levels and light.

Growth rates of individual colonies have been measured by weighing, volumetric

determination, staining with Alizarin Red dye and direct measurement along inert pins placed
in the coral for reference. The most commonly used technique has been X-radiography . Corals
secrete skeletal material of varying density depending on temperature change, light level and
intensity of reproduction. While the precise link between density banding and various physical-
oceanographic conditions is still a point of debate, the regular pattern that is visible on X-rays
provides a calendar upon which the development of an individual colony can be charted. In
Montastrea annularis, the banding pattern has been shown to usually be annual, and is likely
linked to seasonal changes in water temperature as well as reproductive patterns. During
periods of warmer temperatures, the coral lays down a denser band that is reflected in the
darkened band on an X-ray positive.

Bioerosion - the destructive stage

15
W hile corals and coralline algae are capable of producing massive structures over time, most other organisms living
in and on the reef counter that process in their quest for food (grazers) or shelter (borers). This process is bioerosion
which has been recognized as a major factor in both the biological and geological development of reefs.

Grazers and Predators

A ll dead surfaces of the reef are rapidly overgrown by a thin film of filamentous green algae. These form broad algal
turfs that are a favorite diet of many fishes and urchins. Some algae bore tiny but ubiquitous holes into the reef
surface. These endolithic algae can weaken the substrate, making it more susceptible to damage by grazers. As these
organisms die, their borings are usually filled by mud-sized sediment (micrite). Repeated episodes of algal infestation,
micritic infilling and cementation forms a thin and
usually darkened rind around the edges of virtually all
carbonate grains. This reworked "skin" is called a
micrite envelope.

While some grazers (i.e. damselfish) selectively pluck

turfs from the substrate, and actively " farm " the turfs
within their territories, most grazers are less selective.
Some have evolved specialized systems that allow
them to ingest wholesale patches of turf along with
sections of the supporting substrate. Parrotfish bite off
pieces of substrate and pass them through a rasping
structure, the pharyngeal mill, which produces a
mixture of algae and sediment. The algae are digested,
and the remainder is passed through the gut, mostly as
sand. Urchins similarly rasp away substrate along with
the algae they ingest. The sedimentary
by-product is a roughly equal mixture of
sand and mud. While most of these
grazers attack dead and algal-covered
substrates, some are known to also feed
on live coral. Along the Great Barrier
Reef, the Crown-of-Thorns starfish
(Acanthaster plancii) has been the focus
of national concern each time its
population reaches epidemic proportions
and devastates large areas of live coral. In
the Caribbean, coralliophyla (coral
devoring snails) are becoming larger and
more common and the number and size of fire
worms is increasing. Both of these feed on
coral.

Borers

M any organisms degrade the reef

structure in the process of creating
homes. A variety of worms, bivalves and
sponges bore into the reef for shelter. Of
these, sponges are generally the most
pervasive. In the Caribbean, several species of
the genus Cliona aggressively attack dead
substrate and can, on occasion, totally destroy
any evidence of original structure. Cliona
excavates its gallery by chemically dissolving
16
bits of carbonate away from the surrounding walls. The result is a silt-size chip that has a diagnostic shape when viewed
in SEM. The same high nutrient levels that might damage reef corals initially will encourage the subsequent infestation by
sponges that break down the structure that has been built. Also, high nutrients encourage algae that attract grazers and can
occupy available space faster than coral larvae.

In many localities, the dominance of Cliona is matched by boring bivalves. Most important among these is the genus
Lithophaga and several boring chitons. Lithophaga can reach 30 cm in length and, in isolated instances, over 50
individuals per cubic meter can be found within a patch of reef. In addition to destroying substrate, the resulting borings
significantly reduce the resistance of the overall structure to other forms of biological breakdown and physical damage.

Rates of Bioerosion

W hile grazers occasionally leave scrapes and gouges that can be identified in the geologic record, it is impossible to
determine the volume of material that has been removed. In contrast, infaunal bioeroders leave an excellent record
of their presence. Each borer produces a gallery that is distinctive in both its size and its shape. Furthermore, the
abundance of various gallery types can be directly converted into volumes of carbonate removed.

Our best estimates of bioerosion come from controlled experiments in both the laboratory and the field. Based on these,
grazers appear to be responsible for better than half of the bioerosion in Caribbean reefs. At many locations, urchins
produce larger amounts of sediment (up to 5 kg/m2-yr; avg ~ 2kg/m2-yr, equally split between sand and mud. The relative
importance of sponge boring was determined for St. Croix with rates averaging near 1.25 kg/m2-yr, with 90% of this being
mud.

Physical Determinants
W hile organisms have evolved through time, individual
processes to which they respond have not. Sea-level rise and
fall, tectonic change, wave energy and a host of other physical-
oceanographic factors all respond to basic laws of physics that have
remained constant. Therefore, recognizing the nature of these
processes and the signatures that they produce in modern reefs may
provide our best chance to understand the evolution of fossil reefs
through time.

For purposes of our discussion, these controls are divided into three
groups, based on the scale at which they are most important. It
should be kept in mind throughout the following discussion that
these scale-based groupings are somewhat artificial and that most
of the controls discussed below exert some influence at all scales.

Macro-Scale Controls

M acro-scale controls are those that can operate globally or at

least over very large areas. Most important among these are
tectonics and sea-level rise and fall.

Tectonics

T he importance of tectonics was recognized by the earliest reef

workers. Mojsisovics concluded that great upheavals of the
Earth's crust must have been responsible for fossil reefs found in
the Italian Alps. It was noted before Darwin that many fossil reefs
had attained thicknesses of hundreds to thousands of meters, far

17
greater than the known depth range of modern corals (ca. 100 meters). It was argued that this was the result of gradual
accretion by the reef under the influence of constant subsidence. As the "floor" of the system slowly sank, the reef
accreted to maintain the reef crest near sea level.

Based on his examination of Pacific atolls, Darwin argued that the world-wide distribution of reefs could be explained
solely by understanding the tectonics involved. He proposed that Indo-Pacific fringing reefs, barrier reefs and atolls
represented an evolutionary continuum tied to the subsidence of a volcanic core at the nucleus of the system. He further
argued that fringing reefs like those in the eastern Caribbean Sea, formed in areas of tectonic stability and that barrier
reefs and atolls were reflective of significant subsidence.

While Darwin's ideas about the genetic sequence of fringing reefs to atolls in the Pacific were later confirmed by deep
cores through Enewetak and Bikini Atolls, our growing knowledge of global tectonics has shown that Darwin's ideas
about subsidence as a universal and sole control of reef type were flawed. Contrary to his prediction, the fringing reefs of
the eastern Caribbean exist is an extremely volatile tectonic setting. Conversely, barrier reefs often occur along trailing
continental margins that, by comparison, are tectonically stable. Nevertheless, tectonics does exert a strong influence on
large-scale patterns of reef development, and must be integrated into any realistic "reef model."

Sea Level

S uperimposed upon the long-term tectonic regime are periodic oscillations in sea level. With each waxing and waning
of sea level, reefs that live near the upper limit of sea-level rise and fall have been alternately exposed and flooded.
The result is a series of reefs, each built upon the remains of its predecessor.

We now understand that reef development is the result of not a single controlling variable, but rather a combination of
them. By identifying and averaging the patterns common to depositional sequences from many different ocean basins
through time, "seismic stratigraphy" has been used to construct a record of global sea-level change through time. Careful
analyses of the variability of this pattern from site to site can be used to infer the superimposed effects of local tectonics.
The term relative sea level refers to the change in sea level at any one locality resulting from the combined effect of the
glacio-eustatic rise and fall of the world ocean (related to the melting and freezing of polar ice) and local tectonic motions.

Keep up, catch up or give up: the reef responds - Reefs

can be generally classified into one of three groups .
Keep-up reefs have maintained their crests at or near
sea level throughout their history. The reef interior is
dominated by organisms typically found in shallow
water. Catch-up reefs were initially outpaced by rising
sea level, but have subsequently caught up, usually
after the rate of sea-level rise slowed. Their lower
(older) section is dominated by deeper-water
organisms, and is overlain by shallower-water types
that reflect shoaling of the reef.

The geometry of the reef system, when viewed in

cross section, will reflect the relative ability of reef
accretion to match the rate of sea-level rise. Catch-up
reefs or keep-up reefs that accrete at a rate equal to
that of sea-level rise will build vertically. Once
reaching sea level, many reefs also build seaward to
accommodate the imbalance between the volume of
carbonate being added to the system and the
accommodation space being created by rising sea.

Give-up reefs are those that, for whatever reason, simply stopped accreting. The condition most commonly cited as the
cause for a particular reef giving up is a sudden rise in sea level that is faster than the reef can match. When this happens,
the reef is either gradually left behind (i.e. "drowned") or else " backsteps " to a shallower and more shoreward location.

18
Over the past decade, the interest in "give-up" reefs has been heightened due to the fact that most shallow-water reefs are
capable of accreting at rates greater than even the most rapid sea level rise. This problem has particular significance in
recent discussions about global warming. It has been proposed that, due to increasing levels of "greenhouse gases" in the
atmosphere, a dramatic increase in the rate of sea-level rise may occur by the end of the century. Will present-day reefs be
able to keep pace with this rise or will they be left behind, changing the character of nearshore shelves and banks
dramatically?

Recognition of a particular type of accretionary pattern can provide valuable information on the factors that have shaped a
particular reef and its associated environs. The unraveling of depth-related sequences within individual reefs can provide
insight into the sea-level history of a particular area. Conversely, within areas where the likely patterns of sea-level
change are known, deviations from the expected suite of facies or organisms outlined above can provide useful clues
about other factors that might be in control. These principles can be applied to both modern and ancient reefs alike.

Meso-Scale Controls

M eso-scale processes are generally most important in determining the variability in reefs across a single basin. Wave
energy and regional temperature patterns head this category.

Temperature

C oral reefs are generally restricted to water between 18 and 34° C, with an optimal range of 26-28°C. This is expressed
in latitudinal patterns of coral-reef diversity. Within this range, certain corals will change their growth rates,
depending on their sensitivity to temperature.

Along the 2500-km long Great Barrier Reef of Australia, cores reveal a
record of gradual climatic shift from subtropical to tropical conditions
over the past 30 million years. This is a result of the northward (i.e.
toward the Equator) motion of the Australasian plate in response plate
tectonics. As a result of rising temperature, the benthic community and
the sediments that they produce have changed gradually through time.

19
It is important to note that most corals exist near their upper thermal limits. Therefore, even a slight increase in tropical
temperatures in the future could have a significant impact on the distribution of corals in the tropics. Temperatures only a
few degrees above normal can result in the expulsion of algal symbionts ( bleaching ) that make an important contribution
to their metabolic budget. The 3-4o C rise in temperature in the Pacific associated with the 1982-83 El Niño Event has
been linked to widespread bleaching off the western coast of Panama and the eventual devastation of the coral
community.

At the other end of the temperature spectrum, cold water that upwells off western Panama may similarly limit the
occurrence of corals. In the northern Florida Keys, the growth rates and distribution of Montastraea annularis are in large
part controlled by the periodic influx of cold water pushed out from Florida Bay during the passage of major cold fronts.

Wave Energy

R eef zonation is in large part a response to decreasing wave energy with depth. Geister proposed that on present-day
reefs, a decrease in the day-to-day (prevailing) energy level will trigger a systematic change in the reef-crest
community from one dominated by coralline algal ridges at the high end of the spectrum to a mixed-coral assemblage if
wave energy is very low. A primary control in this regard is the instantaneous flow velocity related to incoming waves.
Similar relationships have been proposed for fossil reefs.

Storms also play an important role in determining reef character. Three primary hurricane tracks exist in the Caribbean
Sea. One of these passes to the south of the Greater Antilles and two pass to the north. Based on these tracks and the
regional distribution of total wave energy, Caribbean reefs can be roughly divided into three types.

Areas of high prevailing wave energy and frequent hurricane disturbance (Type I: i.e. Windward Islands) are
characterized by algal ridges, comprised of storm-generated piles of broken Acropora palmata bound and capped by thick
crusts of coralline algae. High wave energy on a day-to-day basis discourages grazing by fishes that would inhibit the
accumulation of thick algal crusts under calmer conditions. High wave-energy appears to be the most important factor in
the distribution of both Caribbean and Indo-Pacific algal ridges.

Areas of moderate-to-high prevailing wave energy but infrequent disruption by storms (Type II: i.e. northern St. Croix)
are dominated by branching Acropora palmata. From this, it would appear that the important distinction is the lack of
frequent destruction by passing storms.

Areas of low prevailing wave energy and frequent storm disruption (Type III: i.e. northern Bahamas) are dominated by
open pavements with only scattered coral cover. Frequent storm disruption combined with little inhibition of grazing

20
between storms discourages the formation of either the thick algal ridges of Type I reefs or the abundant, branching cover
of Type II reefs.

In the Indo-Pacific, both wave energy and sedimentation play similar roles in reef zonation. As in the Caribbean region,
algal ridges dominate the high-energy margins of open-Pacific atolls. Along the Great Barrier Reef, the reef-crest and
forereef communities change from the high-energy reefs in clearer water near the shelf edge and in the Coral Sea to reefs
subjected to higher levels of sedimentation in calmer waters near land.

Micro-Scale Controls

M icro-scale processes are those that exert an influence within an individual reef or reef system. These include the
topography upon which the reef formed (antecedent topography), light level, local nutrient input, salinity variations
and sedimentation patterns.

Salinity

C oral reefs are limited to areas of reasonably

normal marine salinity (3.3-3.6%). Below normal
levels, carbonate buildups are progressively dominated
by vermetids, oysters, serpulids and blue-green algae.
Teichert, 1958; Heckel, 1974 Low salinity (along with
turbidity) is a primary reason why extensive coral
reefs do not occur opposite the mouths of major rivers
(i.e. the Amazon and Orinoco Rivers of northern South
America empty into seas that are otherwise suitable for
reef development). On a smaller scale, the passes
through many nearshore reefs are controlled by the
present or past locations of streams.

Antecedent Topography

L ike ancient cities, new reefs often form atop older ones. Topographically elevated areas offer significant benefits to
larval recruits, especially those sensitive to sedimentation. As sea level falls and rises again, the elevated remnants of
the last generation of reefs hold the greatest possibility for the survival of their successors. Thus, reef sequences that are
recognized in the fossil record are often not single depositional units, but rather a complex of several reefs, each localized
atop the remains of an earlier one. Many present-day reefs sit astride their Pleistocene ancestors that formed 120,000 years
earlier.

In addition to older reefs,

remnant topography can be
related to the edges of
tectonic blocks, cemented
sand bars, fossilized dunes
and even ancient river deltas.
One common controlling
factor is a process called karsting . During episodes of lowered sea level, limestone strata are dissolved by rainwater.
Remnant highs often result from the combined influence of weathering and pre-existing reefs. Because reefs are generally
reinforced by syndepositional cements formed within the their interstices, the resulting mass is more resistant to
weathering than are the muddier and less-cemented sediments of the platform interior. They serve as focal points for
recruitment once the area is reflooded by rising sea level.

Light

T he intensity and quality of incident light are probably the most-studied of the controls on coral growth and reef
accretion. Because of the importance of photosynthetic symbionts within coral tissues, skeletal growth drops
21
dramatically with depth in response to a decrease in total light, as well as a shift in the spectrum toward the blue end. The
changes of coral community structure with depth are directly related to the decrease in ambient light with increasing
depth. Montastraea annularis in the Caribbean and Porites lutea in the Pacific are the preferred subject of most coral-
growth studies. The regular annual bands that are seen in X-ray. The annual bands provide a calendar, much like tree
rings, that records the growth patterns of each colony.

Montastraea annularis possesses the ability to change its shape in response to more or less light. Grauss & Macintyre,
1982 In the Indo-Pacific region, other corals have shown similar morphologic plasticity. While recent studies have shown
that there may be at least three sub-species of Montastraea annularis, the demonstrated ability of individual colonies to
change shape in response to artificially changed depth or light level supports the importance of this plasticity to a wide
variety of conditions. It
should come as no surprise
that these more adaptable
species are the most
common among those
recovered in cores from
modern reefs.

Sedimentation

T he most important
types of sediment
stress are:

• smothering,
• shading,
• abrasion and
• inhibition of recruitment.

Smothering is the easiest to visualize.

Reefs on the downwind flanks of large
carbonate platforms can be buried by
sediment derived from the bank top.
During storms, or more recently,
nearshore dredging, the levels of
suspended sediment can increase
markedly, burying entire reefs or at least
damaging reef corals and other sediment-
sensitive biota.

Shading is probably the most important of

all the sediment-related effects. Because it
is more subtle than smothering, however,
its effects are difficult to quantify.
Despite an impressive body of literature,
little quantitative information exists on
the specific responses of reef organisms
to sediment loading. The frequency with
which coral reefs succumb to turbid water
from dredging projects or increased
runoff are totally inconsistent with
laboratory experiments that have
documented surprising tolerance by corals
to high doses of sediment over short
periods of time. The obvious factor here
22
is the effect of long-term exposure. Sediment traps on reefs can be used to measure the sediment influx over a year and the
rate of growth of Montastraea annularis. Even lower levels of stress can gradually wear down the reef's defenses if
allowed to persist long enough. This accounts for the lack of well-developed reefs along the downwind flanks of most
platforms and the increasing number of post-mortem autopsies of reefs killed by dredging or upland clearing.

The level of suspended matter in the water column has a direct effect on light penetration. The greater depth at which
corals are found in the open Pacific (>100m) likely reflects clearer water relative to that along the Great Barrier Reef or
throughout the Caribbean.
Turbidity ("cloudiness" of
the water column) exertes
a strong control on the
depths at which coral
zones change, presumably
in response to decreased
ambient light levels due to
suspended solids in the
water. Reduced light levels
can suppress coral-growth
rate, impact natural
zonation patterns and
induce wholesale mortality
if allowed to persist. The
change in percent of cover
on individual reefs in
relation to the amount of
fine-grained suspended
sediment can be a practical
guide to species resistance to sediment influx .

Abrasion by moving sediment can cause substantial damage to coral tissue, especially during storms. Hubbard, 1992 Even
under less-energetic conditions, sediment scour can exclude head corals from the reef crest as regular abrasion inhibits
their growth or kills smaller colonies.

Excess sedimentation can also discourage recruitment by coral larvae. Morelock, discussed the importance of substrate
type in larval recruitment in Puerto Rico. Roy and Smith proposed that on Fanning Island, the increased vulnerability of
young corals to sediment damage was a more important factor than sediment covering available space.

Nutrients

U ntil recently, nutrients have

generally been considered as
beneficial to reefs. However, more careful
measurements of oceanic nutrient flux
have shown that open-ocean reefs exist in
a "nutrient desert". Kinsey and Davies
demonstrated that the key to the reef's
success is its ability to efficiently utilize
these low levels of nutrients. In fact, high
nutrient levels are now considered to be
detrimental to "reef health." Hallock and
Schlager proposed that elevated nutrient
levels were responsible for widespread
reef degradation in the Cretaceous, and
suggested that nutrient availability has been greatly underrated as a primary control of reefs over large spatial and
temporal scales.

23
The role of nutrient inhibition in coral reefs is multi-faceted. On the organism level, it has been proposed that high
phosphate levels in the water can effectively shut down the calcification mechanism. At the community scale, higher
nutrient levels tend to favor sponges and algae, which can out compete corals for space and prevent larval settling. Once
the coral dies, higher levels of nutrient availability favor grazing and infestation by infaunal borers such as Cliona spp.,
which will progressively destroy the remaining skeleton and can totally remove any record of the original organism.

Man's Impact

A s population rises and exploitation of the world's coastal regions follows, the influence of man is rapidly moving
toward the top of the list of reef controls. Along a reef system off Costa Rica, Cortes and Risk proposed that coral
growth (and probably cover) has been progressively reduced by widespread agriculture and logging since the late 1950's.
In a natural experiment in Kaneohe Bay, Hawaii, reefs were severely damaged after the installation of a sewer outfall near
the reef. After the discharge was moved, the reef showed significant signs of recovery.

In a study involving transplanting of coral from sediment and nutrient impacted waters to a nearby pristine environment,
Morelock and Ramirez found very little recovery by the coral of more than 10 species.

By understanding the response of reefs to natural stress, we can better equipped to predict and mitigate damage related to
development. Much of this information will come from detailed experimentation by biologists. However, even the most
carefully designed experiment is incapable of addressing one factor that is emerging as perhaps the most important of all -
time. Over the past decade, better than half of the Acropora palmata and much Acropora cervicornis in the Caribbean has
died off , probably a result of "White Band Disease" and hurricanes. In the mid 1980's, the sea urchin Diadema antillarum
underwent an unprecedented population crash throughout the region. This grazer is an important regulator of algal turfs
that compete with corals for space on the reef. More recently, widespread coral bleaching and other disturbing changes in
reef communities noted by the scientific community have spurred congressional hearings on the "health" of the world's
reefs.

A critical question in all of this is whether these represent a sudden, man-induced decline or whether we are simply on the
downside of a natural "boom-and-bust" cycle that occurs over a period of decades or centuries. Recent concerns over
global warming only serve to reinforce the need to understand the long-term responses of coral reefs to large-scale
physical-oceanographic forcing functions.

The answers to these temporal questions may be no further away than the interior of the Holocene reefs which underwent
a similar warming 10,000 years ago. Placing recent events into the context of change on a scale of decades to centuries
also demands a greater commitment to long-term monitoring, which is unfortunately a low priority in a society that
demands quick fixes and short-term solutions. Both should be priorities for future research.

Summary & References

C oral reefs are highly diverse features, the importance of which is related to both the need to manage our present-day
natural resources and the desire to understand the evolution of life in tropical seas. Most of our models are based of
the principle that reefs are in-place and interlocked assemblages of organisms growing on the backs of their predecessors;
these are critically flawed. To adequately understand the dynamics of reefs and their structural evolution through time, we
must understand not only the processes that produce calcium carbonate, but those that break it down and redistribute it as
sedimentary debris as well.

Reefs are largely controlled by physical-oceanographic processes while, at the same time, profoundly influencing them.
Through geologic time, the individual organisms that are responsible for producing calcium carbonate have changed
markedly, but the process that have dictated their success or failure have been constant. Thus, process and not product is
the ultimate common denominator that we seek in comparing modern reefs to their ancient counterparts.

In striving to understand how ancient reefs might have functioned, the present is truly the key to the past. Many of the
critical management problems facing us today, however, require that we turn this equation around. By examining the
character of past reefs and identifying the large-scale temporal response of reefs to various natural stresses, we may be
24
able to place the rapid changes we are now witnessing into the context of longer-term change – the past may be the key to
the present... and the future.

The ever-increasing world population and its dependence on natural resources are placing new stresses on reefs at an
accelerating rate. While these systems are surprisingly resilient, increasing levels of human impact in the form of elevated
nutrients and sedimentation are taking their toll. Recent episodes of coral bleaching have brought national attention to the
perils of tropical reef systems. The outbreak of the Crown-of-Thorns starfish on the Great Barrier Reef or the sudden die-
off of the long-spined sea urchin in the Caribbean have forced us to weigh the likelihood of these resulting from man-
induced stresses versus natural
ups and downs in the
organisms that populate our
seas. The possibility that sea
level may soon rise at a rate
exceeding the ability of
present-day reefs to keep up is
further cause for great
concern. The resulting impacts
on coastal populations that
depend on the reef for food
and protection from oceanic
waves could be devastating.
Whether the goal is to better understand reefs and how they have evolved through time or to protect these valuable
ecosystems from increased degradation, placing the processes discussed in this chapter into a realistic spatial and temporal
framework is of paramount importance.

25