G
its leader; in the development of the scientific programme;
GARDINER, JOHN STANLEY (1872–1946)
Barbara E. Brown
University of Newcastle upon Tyne, Durham, UK
Definition
An eminent British zoologist who is primarily recognized
for his early work on reefs throughout the Indian Ocean
but who was also a major influence on the1928–1929
Great Barrier Reef Expedition from its inception through
to review of its resultant publications.
Professor Stanley Gardiner was best known for his
extensive expeditions into the Indian Ocean to study the
flora and fauna of coral reefs and islands in the early
1900s. As a young man, he joined the Royal Society Expe-
dition to Funafuti in the Ellice Islands in 1896 to assist the
deep boring of an atoll and test the “subsidence theory” of
reef origin proposed by Charles Darwin. On this trip, he
also visited nearby Rotuma and Fiji carrying out prelimi-
nary experiments on coral nutrition and growth, the results
of which made him one of the first scientists to question
the role of coral symbiotic algae in these processes. Dur-
ing 1899–1900, 1905, and 1908, Gardiner undertook sev-
eral expeditions into the Indian Ocean working in the
Laccadives, Maldives, Chagos Archipelago, Farquhar,
Providence, St. Pierre, and Mauritius. He was one of first
scientists to provide detailed descriptions of the reefs of
the Indian Ocean together with taxonomic details of their
constituent corals as well as comments on their form and
possible modes of origin.
Less well known is the fact that he was a key figure in
the instigation of the 1928–1929 Great Barrier Reef Expe-
dition; in the funding and appointment of C.M. Yonge as
David Hopley (ed.), Encyclopedia of Modern Coral Reefs, DOI 10.1007/978-90-481-2639-2,
# Springer Science+Business Media B.V. 2011
and in evaluation of scientific papers which resulted from
the expedition’s activities.
Gardiner was elected to the Fellowship of the Royal
Society of London in 1908 and in 1909 became Professor
of Zoology at Cambridge. In 1929, he was awarded the
Agassiz Medal of the National Academy of Sciences of
the United States and while in the country he delivered
a series of lectures on coral reefs at the Lowell Institute
in Boston which were published a year later in a book enti-
tled “Coral reefs and atolls.” In 1936, he was awarded
the Linnean Medal of the Linnean Society and in 1945
the Darwin Medal of the Royal Society of London.
Throughout his life, he promoted coral reef interests
both within his Department and abroad. At home, he
provided a laboratory for fellow reef scientists such as
R.B. Seymour Sewell and Cyril Crossland while person-
ally supervising George Matthai’s anatomical treatise on
soft tissues of reef corals. He entertained overseas visitors
in Cambridge and during his and his student’s visits to the
United States engaged actively with Thomas Wayland
Vaughan, Alfred Goldsborough Mayer, and Alexander
Agassiz. His legacy to reef science was much more than
descriptive studies from the Indian Ocean and included
scientific and administrative steerage of the Great Barrier
Reef Expedition which set the scene for a new phase in
the development of reef science in the twentieth century.
Bibliography
Gardiner, J. S., 1898. The coral reefs of Funafuti, Rotuma and Fiji
together with some notes on the structure and formation of coral
reefs in general. In Proceedings of the Cambridge Philosophical
Society, Vol. 9, pp. 417–503.
452 GENERAL EVOLUTION OF CARBONATE REEFS
Gardiner, J. S., 1907–1936. Reports of the Percy Sladen Trust Expe-
dition to the Indian Ocean in 1905. Transactions of the Linnean
Society of London, Vol. 2, pp. 12–19.
Gardiner, J. S., 1931. Photosynthesis and solution in formation of
coral reefs. Nature, 127, 857–858.
Cross-references
Indian Ocean Reefs
GENERAL EVOLUTION OF CARBONATE REEFS
Rachel Wood
University of Edinburgh, Edinburgh, UK
Introduction
While modern coral reefs are sites of high biodiversity
forming impressive wave-resistant structures in clear trop-
ical waters, many other organisms are known to form
diverse types of reefs in temperate, low-energy settings
or in deep waters.
Reefs have developed on Earth for over 3.5 billion
years, and a tremendous diversity of communities of
skeletal and nonskeletal metazoans, plants, algae, and
microbes as well as inorganic calcium carbonate precipita-
tion processes have contributed to their formation during
this time. Reefs have shown zonation in response to envi-
ronmental gradients from their inception, and metazoan
reefs have been differentiated into open surface and cryp-
tic reef communities for their entire
550 million-year
history (Wood et al., 2002).
Ancient reefs were biologically constructed reliefs that
grew on the seafloor that are now expressed as massive bod-
ies of calcium carbonate rock surrounded by bedded strata
(Figures 1 and 2). Some formed small isolated structures;
others became sufficiently large so as to influence the
regional oceanographic and sedimentological environment.
However, as sites of in situ biological growth, they often
a b
General Evolution of Carbonate Reefs, Figure 1 (a) An extensive ancient fringing reef complex: the Permian Capitan Reef, Texas,
USA. (b) Isolated patch reef, late Carboniferous (Pennsylvanian), Sacramento Mountains, New Mexico, USA. Field of view = 2 m.
record abundant palaeoecological information lost from
other types of communities, as well as data that can be used
to unravel changing climate and oceanography over evolu-
tionary timescales.
Given the diversity of ancient reefs and settings, a strict
uniformitarian approach to understanding reef evolution
has limited utility. Indeed, for decades there has been an
ongoing debate as to what constitutes an ancient reef
(see James, 1983; Fagerstrom, 1987; Geldsetzer et al.,
1988; Wood, 1999; Stanley, 2001 for summaries).
A general term often used for ancient reefs with no conno-
tations as to processes of formation, size, or shape is car-
bonate buildup.
Many ancient reefs had fundamentally different ecologi-
cal and environmental demands compared to modern coral
reefs. But all reefs, regardless of the organisms that form
them or where they grow, are controlled by factors that
enable the biological occupation of space on the seafloor.
All reefs show the recurring processes of (1) in situ bio-
logical fixation of carbonate by organic assemblages of
microbes, algae, and metazoans, (2) development of inter-
nal cavity systems during growth, (3) synsedimentary lith-
ification, and (4) bioerosion (Figure 2). Reef evolution
has recently been reviewed by James and Wood (2010),
and the following review has been based in part upon
this work.
Biological fixation of carbonate
Calcareous metazoans, algae, and microbes can either
individually or in combination form a reef. The relative
importance and abundance of each has changed through
geologic time, leading to the wide array of ancient reef
fabrics present in the geological record.
Calcareous metazoans
All invertebrate taxa of calcareous metazoans have formed
reefs at some stage in the past, but the most important
groups are sponges, corals, bryozoans, and bivalves. Reef
metazoans are often colonial or clonal and are capable of
 GENERAL EVOLUTION OF CARBONATE REEFS 453

Facies

Organic
growth
Core
Forereef Flank

Sedimentation Destruction

Shallow
Cementation
Predator Water
Reef
Branching Calcareous
algae
Sponge
Grazer
Boring
Deep
Water
Platy and
Reef
Massive encrusting
Bryozoan
and Domal
Cavity with
internal sediment Crinoid
Metazoan Automicrite
Reef
Mosaic

Calcimicrobe

General Evolution of Carbonate Reefs, Figure 2 A series of sketches illustrating (a) Cross-sectional geometry of a typical reef as
exposed in outcrop. (b) Complexity of interrelationships between processes that control reef composition. (c) Main attributes of
shallow-water reefs and deep-water mounds. (From James and Wood, 2010.)

surviving and regrowing or even multiplying after damage
to their living tissues. All are obligate calcifiers in that the
organism controls biomineralization.
Reef calcareous metazoans can be either heterotrophs
or mixotrophs. Heterotrophs consume other organisms
whereas mixotrophs are heterotrophs that also contain
symbiotic microorganisms, generally photosynthetic
cyanobacteria or single-celled algae.
Sponges including hexactinellids with siliceous spic-
ules alone and those with calcareous skeletons were major
reef builders in the past. Calcified sponges belong to sev-
eral different groups including calcareans, demosponges,
stromatoporoids (a group of sponges of probably mixed
affinity that are distantly related to various modern calci-
fied sponges), archaeocyaths, and chaetetids. Corals com-
prise both tabulates and rugosans from the Cambrian/
Ordovician to the end Permian and scleractinians from
the mid-Triassic onward. The other important organisms
are bryozoans that, even though relatively small and deli-
cate, are critical elements of many reefs and often act as
scaffolds between which in situ microbially induced lime
mud (micrite) can form and cements precipitate.
Although we cannot be certain about the physiological
tolerances of many ancient fossil reef builders, some gen-
eralities are possible. In contrast to those organisms that
contained phototrophic symbionts, heterotrophs would
have been limited by the availability of light and nutrients.
On the basis of isotopic (Stanley and Swart, 1995), and
growth-form evidence, scleractinian corals probably pos-
sessed zooxanthellae by the late Triassic. Tabulate corals
were a diverse group of organisms and it is not clear
whether they contained similar symbionts. Rudist
bivalves formed low-relief aggregations and were massive
producers of sediment in the Cretaceous, but some of the
most important groups did not appear to have symbionts
(Steuber, 1999). Fossil sponges, both spiculate and soft
bodied, and calcified sponges (such as stromatoporoids
and archaeocyaths) are poorly understood and their asso-
ciation with photosymbionts is uncertain (Wood, 1999).
Calcareous algae
Calcareous algae, both red (coralline and squamaracean)
and green (codiacean and udoteacean), are obligate
454 GENERAL EVOLUTION OF CARBONATE REEFS
calcifiers. They can form reefs as well as binding and sta-
bilizing reef frameworks being prolific producers of sedi-
ment. Such algae probably occupied these niches in the
past. Calcareous algae have been part of the carbonate fac-
tory since the Ordovician but were not significant in reef
communities until the early Carboniferous (Mississip-
pian). Phylloid (leaf-like) algae were conspicuous reef
builders in the late Palaeozoic.
Microbes
Microbes are highly diverse single-celled prokaryotes and
eukaryotes. Carbonate precipitation associated with them
can be inferred by the presence of often encrusting micro-
crystalline micrite (lime mud) with peloidal, clotted or
laminated fabrics, or by calcified microbial sheaths
(calcimicrobes).
Microbes are non-obligate calcifiers in that they induce
carbonate precipitation by their metabolic processes or
postmortem chemistry: they are incapable of calcification
under unfavorable conditions such as low carbonate satu-
ration. The attributes of microbial textures are therefore
thought to depend upon seawater carbonate saturation
and postmortem preservation.
Calcification often takes place associated with a biofilm
consisting of layers of organic matter (EPS – extracellular
polysaccharide) that contains a variety of living and dead
microbial communities within a matrix of degrading
organic matter and mucilage. The precipitated microcrys-
talline micrite is sometimes called automicrite because it is
produced in-place, generally on a substrate, and it can be
an important binding or cementing agent.
It is also possible that some micrite formation in reefs
relies mainly on early diagenetic organomineralization
processes to aid calcite nucleation on dead organic matter,
particularly via molecules derived from decayed sponges.
The presence of abundant, reactive Ca2þ-binding fulvic
acids have been confirmed in well-preserved Cretaceous
mud mounds and modern coral reef caves.
Automicrite is the main constructor of layered stromat-
olites and clotted thrombolites. Stromatolites, composed
of automicrite and synsedimentary cement, were the
only reef builders throughout the Precambrian and were
augmented by thrombolites and calcimicrobes in the
Neoproterozoic. They have continued to be a variable
contributor to reefs throughout the Phanerozoic.
Calcimicrobes, once established in the Neoproterozoic,
have continued to be a significant and sometimes domi-
nant element of reefs throughout the Phanerozoic, particu-
larly the Paleozoic. Whereas automicrite is essentially an
encrusting or binding element in most reefs, often within
cavities, it may nevertheless contribute substantially to
the reef rock volume.
The most common calcimicrobes are tubular
(Girvanella, Rothpletzella), subspherical (Renalcis), or
branching (Epiphyton). They are generally interpreted
as sheaths of bacteria that have undergone variably
taphonomic alteration. Many other similar structures of
complex affinity such as the Mesozoic Tubiphytes (proba-
bly a foraminfer–cyanobacteria association) have also
been placed in this category. Calcimicrobes were
also important sediment producers.
Internal cavity systems
Much of most modern and ancient reefs are void space as
metazoan-algal-microbe growth geometries often produce
interconnected cavities, even in those reefs almost wholly
composed of mud. These cavities are often shaded from
light and suffer less predation or grazing and reduced
wave energy compared to the open reef surface. Cavities
are often populated by cryptic organisms that encrust
walls and hang down from ceilings. They range from pho-
tic organisms near the openings to relatively fragile het-
erotrophs in the dark, lightless interiors. Automicrite
often preferentially coats inner surfaces of cavities and
encrusts the cryptic community. Cavities are also sites of
fine-grained sediment accumulation, material composed
of skeletal debris and mud that forms a geopetal inter-
nal sediment fill. Reef cavities first appeared in the
Neoproterozoic coincident with calcimicrobes and
thrombolites. Thereafter, they were a significant element
of most reefs.
Synsedimentary lithification
Most reefs are lithified immediately below the living sur-
face by a variety of calcareous precipitates. Much of such
cement is microcrystalline but other cements are spectacu-
lar, growing as millimeter of centimeter-sized crystal
arrays from the walls and ceilings of cavities or between
sediment grains. Such cements not only confer rigidity
to reefs but also occlude much original pore space.
The location of the reef determines the amount of
cementation. Platform-margin and down-slope reefs usu-
ally contain marine cements, particularly in high-energy
areas, whereas lagoonal or inner-ramp reefs rarely do
and so may be unlithified.
Synsedimentary cementation is an attribute of reefs of all
ages but the type and mineralogy is controlled in part by
changing seawater chemistry. The most intensive ancient
cementation documented is found in Precambrian reefs,
where some are volumetrically dominated by cements.
The amount of crystalline cement in reefs decreases with
the appearance of calcareous organisms in the Cambrian
and then again following the evolution of calcareous plank-
ton in the Jurassic: Many Palaeozoic reefs contain conspic-
uously more synsedimentary cement than Jurassic and
younger examples.
Bioerosion and predation
Living calcareous reef builders and reef rock are grazed
upon by a variety of bioeroders, particularly limpets, echi-
noids, and fish that not only consume soft parts but also
remove reef carbonate, which is reduced to fine sediment.
Although present in the Palaeozoic, this component of in-
place reef removal, bioerosion, and sediment production
 GENERAL EVOLUTION OF CARBONATE REEFS
began in earnest in the Mesozoic and has increased dra-
matically ever since (see entry Bioerosion).
Geologic history of reefs
The Phanerozoic witnessed major turnovers of reef biotas,
mass and minor extinction events, and profound changes
in the chemistry of seawater. The evolving history of reefs
has been reviewed extensively by Fagerstrom (1987),
Wood (1999), Stanley (2001), Kiessling et al. (2002), and
James and Wood (2010), and is summarized in Figure 3.
Reefs have broadly evolved from dense, stromatolite-
dominated reefs of the Archean, with the appearance of
thrombolites and calcimicrobes in the Neoproterozoic,
forming the first reef cavities, to the successive rise of various
skeletal metazoans from the earliest Phanerozoic onward.
During geological periods when large calcareous skele-
tal and microbial structures were common, reefs grew as
fringing reefs landward, as patch reefs across platforms,
and as barrier reefs along platform margins. Coeval
mounds developed in either quiet water settings across
the platform, or on ramps, or on the slope.
The formation of large, shallow-water, flat-topped
platforms coincide with the periods of skeletal reef devel-
opment because skeletal reefs are characterized by well-
cemented carbonate buttresses along their seaward
margin. These ramparts absorbed waves and swells, thus
allowing more tranquil, protected lagoonal facies to
develop across the platform interiors. When skeletal reefs
were absent, carbonate ramps were the norm, where reef
mounds and mud mounds were the only buildups, which
either grew leeward of sand shoals or more commonly
on deep-water ramps and slopes.
The great diversity of ancient reefs present a continuum
of shared ecologies and sedimentary characteristics, but
they can be divided into the broad end members of (1) skel-
etal reefs, (2) skeletal–microbial reefs, (3) microbial reefs,
and (4) mud mounds (James and Wood, 2010; Figure 4).
Skeletal reefs
Shallow reefs
Skeletal, shallow reefs, where skeletons dominate most of
the rock volume, are restricted in geologic time to the
(1) middle Ordovician to late Devonian, (2) late Jurassic,
and (3) Cenozoic.
Although synsedimentary cements, microbes, and
automicrite were present, they are usually of subsidiary
importance, except during the intervals of the Palaeozoic.
Internal cavities, which may comprise as much as 30% of
the reef volume, can be sites of diverse cryptic communities,
calcimicrobial growth, and internal sediment. Automicrite is
present on, around, and between skeletons. Synsedimentary
cement can be spectacular, with the largest amounts found
in the reefs at the platform margin. Such reefs were built
by calcified sponges (stromatoporoids), tabulate corals, or
scleractinian corals.
These reefs, with their large skeletons and relatively
rapid growth rates, are amongst the most extensive in the
455
rock record. Diversity is lowest in reef crest and deep-
water environments, with both environments favoring
sheet-like skeletal morphologies. The highest diversity
of skeletons and shapes, and therefore rock types, occur
at intermediate depths.
Growth forms and skeletal diversity are strongly con-
trolled by hydrodynamic energy. Unlike modern corals,
Palaeozoic stromatoporoid sponges and tabulate corals,
even though they often grew as laminar forms, sat on or
were anchored within the sediment (Figure 5). Large, tab-
ular forms that were solidly rooted in the sediment
inhabited high-energy zones such as the reef crest, but
since stromatoporoids did not typically have an encrusting
habit, it is doubtful that they were successful builders in
the surf zone. Tabular stromatoporoids, locally bound
together by calcimicrobes, automicrite, or cement, are also
known to have occurred in energetic waters. Domal, bul-
bous, and dendroid forms occupied quiet water zones,
either below wave base or in sheltered areas of the back
reef and the lagoon (such as delicate stick-like
amphiporoids). These skeletons were locally reworked
during storms and redeposited as rubble units associated
with peritidal settings.
Deep-water skeletal reefs
Today, deep-water skeletal reefs grow worldwide in water
depths of between 250 and 1,500 m, where temperatures
range from 4 C to 12 C (Freiwald and Roberts, 2005).
They are constructed by scleractinian corals that lack
photosymbionts, especially the branching form Lophelia
pertusa. These reefs are muddy and unlithified and range
up to 5 km long and 40 m high, with generally steep sides
up to 160 m thick. Many are located in areas of elevated
nutrients such as oceanic fronts or upwelling; others are
sited on the top of cold hydrocarbon seeps.
Cold-seep reefs are part of a lineage of similar struc-
tures that extends back to the Silurian, but there are very
few corals from methane seep communities older than
the late Eocene. The rock record of such skeletal reefs is
not extensive. The oldest known is Triassic, but they are
not numerous until the Cenozoic, where an especially
good record exists in north-west Europe and the Mediter-
ranean region. Pre-Cenozoic deep-water reefs seem to be
either skeletal–microbial mounds or mud mounds.
Mixed skeletal–microbial reefs
Many ancient reefs bear two important constituents: small
or delicate skeletal metazoans that are either heterotrophs
or autotrophs (e.g., phylloid algae) and a microbial com-
ponent of calcimicrobes or inferred microbialite (Webb,
1996; Wood, 1999). These structures with a mixed consor-
tium of organisms are unlike most modern skeletal reefs
and have been called reef mounds or biogenic mounds
(James and Bourque, 1992). Such reefs often contain
extensive framework cavities, with their own distinctive
cryptic biotas and abundant synsedimentary cements
(Figure 6b).
456 GENERAL EVOLUTION OF CARBONATE REEFS

Periods Buildups Major biotic elements

0
Corals +
Cenozoic 8 C
Calcareous Algae

Rudists + Corals
100 Calcareous
Cret. 7 Rudists K
algae
Corals + Rudists
Stromatoporoids
Jurassic 6 Corals sponges
JR
microbiolite
200
Corals + Calcisponges
Triassic TR
Calcisponges, Tubiphytes, bryozoans,
corals, tubular forminifera,
Permian P
m
microbiolite
5
300 Penn. Phylloid algae Bryozoans P
Bryozoans, calcareous algae
Geologic time in millions of years

Miss. M
Calcimicrobes Microbiolite

Devon. Calcimicrobes D
400 Stromatoporoids corals
Silurian 4 microbiolite sponges S
Stromatoporoids Bryozoans
Ord. + covrals sponges O
Sponges Strom-thrombolites
500
3 Strom-thrombolites
Cambrian Calcimicrobes C
Archaeocyaths

600 Stromatolites
Thrombolites
calcimicrobes

1,000 *
Protero. 2
Stromatolites P

2,000 Stromatolites

Archean ? 1 Stromatolites A
3,000

Skeletal reefs Mud mounds

Skeletal-microbial reefs Extinction events

(biogenic mounds)

General Evolution of Carbonate Reefs, Figure 3 Reefs through time, illustrating periods when skeletal, skeletal–microbial reefs, and
mud mounds were important. Numbers indicate different associations of reef- and mound-building biota. Arrows – signal major
extinction events. * – scale change. (From James and Wood, 2010.)

Autotroph-microbial or heterotrophic-microbial com- Autotroph dominated

munities dominated ancient mixed reefs in shallow waters Ancient examples of mixed autotroph reefs include Penn-
of the photic zone. No direct modern analogs are known sylvanian to early Permian phylloid algal mounds. These
for either of these systems. algae are probably of mixed affinities and grew as laminar,
 GENERAL EVOLUTION OF CARBONATE REEFS 457

Reef fabric

Skeletal Skeletal- Microbial

reefs microbial reefs
reefs
a c e
Shallow
water

Neritic

b d f

Deep
water

Skeletal Skeletal- Mud

reefs microbial mounds
reefs
Large skeletons Stromatolite
(corals; Stromatoporoids)

Small skeletons Thrombolite

(non-photic corals) Automicrite
Invertebrate skeletons calcimicrobes
and calcareous algae synsedimentary cement
Platy skeletons Internal cavity
(bryozoans) geopetal sediment
synsedimentary cement
Small articulated skeletons
(crinoids, brachiopods, bivalves) Spicules (sponges)

General Evolution of Carbonate Reefs, Figure 4 A modified classification to describe reef carbonates. (From James and Wood,
2010, after Embry and Klovan, 1971.)

cup-, bowl-, or upright leaf-like forms that may have been
encrusted by automicrite. Such reefs are usually low-relief
isolated structures, appear to have grown in shallow
waters, and are often associated with grainy sediments that
indicate growth under fairly energetic conditions with
extensive flanking beds. These mounds were typically
the source of significant skeletal debris that formed exten-
sive flank beds. Halimeda mounds have been reported
from the late Miocene and may have been stabilized by
the rapid lithification of micrite and microbial crusts
(Martín et al., 1997).
Heterotroph dominated
Heterotroph reefs are common in the ancient record.
Lower Cambrian reefs occur as small isolated mounds,
consisting of archaeocyath sponges and calcimicrobes,
particularly Renalcis and Epiphyton (Figure 7a and b).
Lower Ordovician reefs were likewise dominated by
calcimicrobes, microbialite and, locally, corals.
Some back-reef shallow-water Frasnian (late Devo-
nian) reefs exhibit large skeletal metazoans (particularly
stromatoporoid sponges and subsidiary corals), as well
as extensive fenestral microbialite and calcimicrobes, both
as free-standing mounds and as secondary encrustations
within cryptic habitats (Figures 5b and 6a).
Shallow-water Lower Carboniferous (Mississippian)
reefs are formed by endemic communities composed of
laminated microbial mounds with a rich encrusting open
surface and cryptic fauna dominated by algae, bryozoans,
corals, and sponges (Mundy, 1994; Webb, 1994; Ahr
et al., 2003).
The mid-to-late Permian is characterized by extensive
fringing reefs with a well-developed zonation. They con-
sist in part of a primary framework of frondose bryozoans
and calcified sponges (many of whom were cavity
dwellers) that were bound by extensive crusts of lam-
inated automicrite (Figure 6b). Tubiphytes is common,
together with various encrusting algae including
Archeolithoporella. Volumetrically, many of these reefs
were dominated by sediment and synsedimentary cement,
or automicrite. Such successions are inferred to have
formed within open cavity systems with freely circulating
seawater, in response to decreasing light and energy condi-
tions as the reef was progressively buried. Upper Jurassic
458 GENERAL EVOLUTION OF CARBONATE REEFS

1
15
14
a 13
1

11 12
11
12

4
10 2
3
4 4
6
1m
6
2 12
5 7
5

b 1

General Evolution of Carbonate Reefs, Figure 5 (a) Large stromatoporoid sponge, growing over soft sediment. Late Devonian
(Frasnian) Windjana Gorge, Canning Basin, Western Australia; scale = 10 cm. (b) Reconstruction of back-reef community, Late
Devonian (Frasnian) Windjana Gorge, Canning Basin, Western Australia. 1. Stromatolites, 2. Domal stromatoporoid, 3. foliaceous
stromatoporoid, 4. Renalcis, 5. fibrous cement, 6. internal sediment, 7. platy stromatoporoid, 8. crinoids, 9. branching stromatoporoid,
10. laminar stromatoporoid, 11. encrusting stromatoporoid, 12. microbialite, 13. clastic sediment, 14. gastropods, 15. Oncolites. (From
Wood, 1999; Copyright J. Sibbick.)

shallow-water reefs were dominated by hexactinellid and
lithistid spiculate sponges, coated by automicrite and
thrombolites, and encrusted by a variety of biota.
Deep-water skeletal–microbial reefs
Deep-water, heterotroph-mixed skeletal–microbial reef
communities are known from the Silurian to Devonian,
Jurassic, and Cretaceous. They were very different from
modern skeletal deep-water reefs. Muddy reefs with
Stromatactis (Figure 8), lithistid sponges, and
calcimicrobes were widespread during the Silurian, and
typically show a vertical zonation that culminates in
shallow-water stromatoporoid sponge-rich communities.
Similar communities, with the addition of microbialites,
stromatolites, and receptaculitids, formed deep-water
reefs in the mid-to-late Devonian.
 GENERAL EVOLUTION OF CARBONATE REEFS 459

a b

General Evolution of Carbonate Reefs, Figure 6 (a) Late Devonian (Frasnian) reef, showing a large branching stromatoporoid
sponge, encrusted by microbialite and Renalcis. Cavities are filled with synsedimentary sediment and cement. Windjana Gorge,
Canning Basin, Western Australia; scale = 5 cm. (b) Pendent sponges growing downward attached to a bryozoan frond. The whole
framework has been encrusted with microbialite. Remaining cavity space is filled with synsedimentary cement botryoids. Permian
Capitan Reef, McKittrick Canyon, Guadalupe Mountains, Texas, USA. Field of view = 6 cm.

Upper Jurassic deep-water mixed reefs were
constructed by thrombolitic–stromatolitic columns or
mounds associated with Tubiphytes, the worm Terebella,
and hexactinellid and lithistid sponges. The platy
scleractinian coral Microsolena, which is thought to have
fed in part heterotrophically, is found in low-light settings,
such as shallow turbid or deep-water environments.
Microbial reefs
Modern stromatolites are relatively rare and appear to
grow in areas of abundant calcium carbonate saturation
and where faster-growing algae and seaweeds are
excluded due to some environmental stress such as active
tidal currents, low nutrients, or high salinity (Figure 9b).
Microbial reefs dominated much of earth history, from
the Archean until Middle Ordovician, but they are only
found in younger Phanerozoic facies, where most inverte-
brates have been excluded by environmental stresses.
They are built primarily by stromatolites and thrombolites
that have a wide variety of morphologies. Precambrian
reefs throughout the Archean, early, and middle Protero-
zoic are essentially a series of stacked stromatolites and,
to a lesser extent, thrombolites (Grotzinger and James,
2000). They have neither cavities nor inter-microbial
synsedimentary cements but exhibit a wide variety of del-
icate branching, lamellar, hemispherical, and conical mor-
phologies that are thought to be due to a combination of
different microbial communities, hydrodynamics, and
sedimentation rates. These microbialites range in size
from centimeter to decimeter, with some large structures
up to 8 m high and 30 m long.
Shallow-water, high-energy Proterozoic buildups were
formed by isolated stromatolitic domes, linked domes,
columnar stromatolites, and their elongate equivalents.
Stromatolite fragments were ripped off during storms
and deposited as cross-bedded gravels locally surrounded
by individual shallow reefs. Reefs at platform margins
show a strong zonation of microbialite growth forms that
mimic facies zonations of metazoan reefs. Isolated deep
subtidal and slope reefs (probably below fairweather wave
base) were built primarily by conical stromatolites.
There seems to be an evolution of microbial structures
through the Precambrian. Archean and Palaeoproterozoic
reefs were mainly synsedimentary precipitates (possibly
microbially mediated) with microbial layers. Microbes
appear to become more prominent in Paleoproterozoic
and Mesoproterozoic structures. A profound change, how-
ever, took place in the Neoproterozoic with the increased
importance of thrombolites and the appearance of
calcimicrobes as reef builders (Figure 9a). This resulted
in the earliest formation of cavities complete with internal
sediment and synsedimentary cement, features that would
become a prominent attribute of all subsequent Phanero-
zoic reefs. This community had the ability to construct
reefs with up to 100 m of relief above the sea floor in
deeper water environments (Turner et al., 1997). Toward
the end of the Neoproterozoic, the first skeletal inverte-
brates (e.g., Cloudina, Namacalathus, and Namapoikia)
began to populate the surfaces of these reefs and were
entombed by the microbial precipitates.
Mud mounds
Mud mounds can form large, often over 100-m high and
400-m wide, isolated, steep-sided structures that consist
of more than 80% of a fine-grained carbonate (micrite)
(Figure 8b). This micrite has both in situ and detrital origin
but often shows accretionary structures constructed by
successive phases, known as polygenetic muds
(“polymuds”), which form on open surfaces and within
semi-enclosed cavities (Monty et al., 1995). Such
polymud fabrics produce complex, three-dimensional
accumulations that in turn form open frameworks that
460 GENERAL EVOLUTION OF CARBONATE REEFS

4 5
2

6
3
2
7
7

6
8
9

10 11
14
12
13
15 cm
10

General Evolution of Carbonate Reefs, Figure 7 (a) Growth cavity in archaeocyath skeletal–microbial reef mound. The cavity is
formed by the calcimicrobe Renalcis and populated by pendent, solitary archaeocyaths. Lower Cambrian, Siberia, Russia; scale = 5 mm.
(b) Reconstruction of lower cambrian archaeocyath community. 1. Renalcis, 2. branching archaeocyath sponges, 3. solitary cup-shaped
archaeiocyath sponges, 4. chancelloriid, 5. radiocyaths, 6. small archaeocyath sponges, 7. “coralomorphs,” 8. large achaeocyath,
9. fibrous cement, 10. microburrows, 11. cryptic archaeocyaths, 12, cribricyaths, 13. trilobite trackway, 14. cement, 15. internal
sediment. (From Wood, 1999; Copyright J. Sibbick.)

may subsequently be filled with mud or synsedimentary
cement. Many mounds also display a rich attached meta-
zoan biota of crinoids, tabulate corals, brachiopods, trilo-
bites, sponges, ostracodes, and bryozoans.
The clotted, peloidal or laminated textures, and the
encrusting or frame-forming habit argue for an in situ micro-
bial or organomineralic origin, augmented by a rapid
synsedimentary lithification. This origin is supported by
moderate rates of carbonate accumulation (0.2–0.8 m/
1,000 years), and the steepness (35–40 ) of mound flanks.
Abundant Stromatactis cavities that parallel the accretionary
mound surfaces (Figure 8a) suggest a close relationship
between mound formation and internal sediment filled voids,
cementation, and carbonate production (Monty et al., 1995).
 GENERAL EVOLUTION OF CARBONATE REEFS 461

a b

General Evolution of Carbonate Reefs, Figure 8 (a) Irregular masses of calcite (Stromatactis) that were originally internal
cavities partly filled with internal sediment and then occluded by synsedimentary cement in a late Devonian (Famennian), Windjana
Gorge, Canning Basin, Western Australia; scale = 5 cm. (b) Muleshoe Mud Mound, Mississippian, Sacramento Mountains, New Mexico,
USA.

a b

General Evolution of Carbonate Reefs, Figure 9 (a) Neoproterozoic thrombolites, Nama Basin, Namibia. (b) Living stromatolites in
Shark Bay, Western Australia.

Mud mounds were often established in deep waters
below the photic zone as indicated by the absence of algae
and cyanobacterial activity. Locally, however, they show
shallowing-upward facies signatures, characteristic of
shallow depths and significant depositional energies. In
the Frasnian “récifs rouges” of the Belgian Ardennes, the
basal facies is composed of Stromatactis-rich red lime
mudstone to wackestone with abundant sponge spicules.
This may have formed in hypoxic waters below the photic
zone. Platy corals appear higher but are overlain by a more
diversified coral, stromatoporoid, and skeletal algae
wackestone to packstone with Stromatactis. Uppermost
facies consist of a stromatoporoid, coral, and bryozoan
framework with various microbial and calcimicrobial
encrusters, which formed in photic waters above fair-
weather base and sometimes in restricted or lagoonal
environments.
Early Mississippian reefs (Waulsortian facies) also
show a similar progression of facies. The growth of the
well-known Muleshoe Mound in New Mexico
(Figure 8b) records a shift from predominantly upward
(aggradational) to lateral (progradational) growth. The
largest volume of these buildups is composed of polymud
fabrics with early cements, sponges, and large fenestrate
bryozoans. The framework is composed of rigid micrite
masses with rounded, bulbous shapes, and thrombolitic
fabrics that are lined by early marine cements. In the
uppermost facies, both the microbialites and large bryo-
zoans grew with a pronounced high-angle orientation into
currents. Crinoid-rich flanking grainstone beds draped the
reef slopes. In places, fenestrate bryozoans built a delicate
framework that formed a limestone containing up to 90%
of early fibrous cement.
The location and initiation of mud-mound formation
appears to be mediated by environmental factors that dif-
fer from those of shallow skeletal reefs. Indeed, there
may therefore be real differences in the style of primary
production and organic matter recycling between these
462 GENERAL EVOLUTION OF CARBONATE REEFS

different reef systems. Deep-water mud mounds are com- (Budd and Kievman, 1994). With this as yet unexplained
monly found in groups, suggesting that their formation rise to dominance of branching Acropora, and a
was environmentally mediated. Several environmental corresponding decline in massive, domal corals, coral reef
triggers have been proposed: the episodic formation of communities with a completely modern aspect appeared
nutrified water masses reduced sediment supply during about 0.5 Ma. Except for the extinction of Pocillopora in
platform drowning or localized oxygen depletion of sea- the Caribbean at about 60,000 years (ka), the patterns of
water. Some mud mounds are aligned along faults or fis- community membership and dominance of coral species
sures, which may have acted as conduits for appear to have been highly predictable for at least the past
hydrothermal fluids. For example, the microspars and bra- 125,000 years (Kyr) (Pandolfi and Jackson, 2001).
chiopods of Muleshoe mound have been shown to have
higher d13C values than non-mound samples, possibly Response to long-term environmental change
related to methanogenic fermentation.
Climate and changing seawater chemistry are important
Mud-mound formation began in the Palaeoproterozoic
controls on the history of reef growth, in terms of both
(probably Neoproterozoic) and extended until the Mio-
complex feedback mechanisms that govern skeletal min-
cene, but they are mostly a Palaeozoic phenomenon, with
eralogy and hence promote some groups over others, and
widespread mud-mound formation occurring during the
controlling the styles of early lithification.
early Cambrian, the lower Ordovician, the late Devonian,
Tectonics ultimately drives many aspects of reef growth
and the Mississippian, which was dominated by
such as sea level, circulation patterns, climate, and evolv-
Waulsortian mounds.
ing seawater chemistry change over long timescales
(Figure 10). Reefs and carbonate platforms commonly ini-
Origin of the modern coral reef ecosystem tiate on preexisting highs such as horst blocks or salt
domes during the early stages of rifting. Reefs are most
The modern coral reef ecosystem is geologically very
numerous on passive continental margins at low latitudes
young. Scleractinian corals appeared in the mid-Triassic
where they typically form barrier or fringing reefs. Similar
and had almost certainly acquired photosymbionts by the
carbonate shelves with reefs can form around the margins
late Triassic at the latest (Stanley and Swart, 1995). The
of shallow intracratonic basins, where they can be
ancestral stock of the modern reef biota appeared in
interbedded with evaporites. Such basins may be prone
the middle Jurassic but was largely suppressed by the pro-
to isolation from the open ocean, leading to infilling by
lific growth of calcitic rudist bivalves in the Cretaceous,
extensive evaporites.
only to rebound after the end-Cretaceous extinction. There
Rates of thermal subsistence on passive margins are
is considerable debate, however, as to whether this coral-
slow and predictable and are similar to those found in
dominated biota was able to construct spectacular large
intracratonic basins. Reef geometries are thus generally
reefs like those of the modern day before the Cenozoic.
compound to progradational. By contrast, reef-forming
Most modern coral genera appeared in the Eocene–
toward the cratonward side of foreland basins is relatively
Miocene, and many extant species extend back no further
rapid and reef geometries more aggradational. Finally,
than the Pliocene. Modern reef fish appeared in the Eocene,
reefs forming in strike-slip basins and on thrust complexes
but the oldest record of parrotfish (scarid) remains are from
are affected by the vagaries of local tectonic movements,
Miocene sediments dated at 14 million years ago (Ma).
which may be highly episodic but involve substantial dis-
During the Oligocene, the compression of climatic belts
placements and are thus highly unpredictable.
and the rise of the Isthmus of Panama created two distinct
Epeiric seas, those that formed by the extensive
regions of reef growth to the Caribbean and Indo-Pacific.
flooding of continents during globally high sea levels,
As a probable result of climatic cooling or habitat loss, a
have negligible basin floor topography. Water depths
major episode of coral faunal turnover ensued between
rarely exceeded 10 m, such that shallow subtidal and inter-
4and 1 Ma in the Caribbean (Budd et al., 1994). Extinction
tidal sediments dominate, with many episodes of expo-
of genera in the Pocilloporidae and Agaricidae was most
sure. Epeiric seas could support patch reefs, often
marked, but many of these genera continued to persist in
elongated because of wind waves, storms, or tides. Due
the Indo-Pacific. A similar differential extinction coinci-
to the limited accommodation space on epeiric platforms,
dent with corals, removed many fish, including all large
progradation is the dominant depositional process,
excavating scarids, herbivorous siganids, and plantivorous
resulting in stacked shallowing-upward sequences or
caesionids from Atlantic reefs (Bellwood, 1997).
biostromes.
Although acroporid corals appeared in the Eocene,
pocilloporids appear to have dominated Caribbean reefs
from 5to 6 Ma, but following a 1-million year (Myr) tran- Climate
sition period of mixed acroporid–pocilloporid assem- Climate is a major controlling force on the evolution of reefs
blages, acroporids became dominant in reef communities on both short and long timescales, as the latitudinal range
in the early Pleistocene (approx. 1.6 Ma). Acroporids of carbonate-producing species is largely governed by tem-
may not, however, have achieved levels of extreme perature and carbonate supersaturation. Reef growth there-
abundance until the late Pleistocene (approx. 0.5 Ma) fore shows cyclicity at all scales in response to short-term
 GENERAL EVOLUTION OF CARBONATE REEFS 463

C O S D M P Pm Tr J K Pg N
6
300 Sea Aragonite and
level high-Mg calcite 5

Mg/Ca mole ratio

200

Sea level (m)

4

100 3

0 2

Mg/Ca 1
–100 Calcite

0
a 550 500 450 400 350 300 250 200 150 100 50 0 Ma

A ? Calcite sea Aragonite sea ? Calcite sea ? A

b
Ice Greenhouse Icehouse Greenhouse Ice
c
Rudists Bivalves
Rugosans
Tabulates Scleractinians Corals
Heliozoans
Chaetetids
Dominant Demosponges
reef builders Inozoans
Sphinctozoans Sponges
Sclerosponges
Stromatoporoids
Phylloids
Skeletal mineralogy Ancestral corallines Algae
Calcite Aragonite Tubiphytes Corallines
High-Mg calcite Solenopores

C O S D M P Pm Tr J K Pg N
d

General Evolution of Carbonate Reefs, Figure 10 (a) Correspondence between changing ocean chemistry and carbonate
mineralogy through time as a function the of Mg/Ca mole ratio in seawater (Stanley and Hardie, 1998) and general global sea-level
change (Vail et al., 1977); the boundary between the fields of calcite (<4 mole% MgCO3), and high-Mg Calcite (>4 mole% MgCO3) and
aragonite is the horizontal line at Mg/Ca = 2; (b) the different dominant nonskeletal mineralogies precipitated in seawater through
time (Sandberg, 1983); (c) the different global climatic and oceanographic periods (Fischer, 1982); (d) mineralogy of different reef-
building organisms (Stanley and Hardie, 1998). (From James and Wood, 2010.)

oscillations (e.g., Milankovitch and glacial–interglacial
cycles) as well as to longer-term climatic intervals driven
by slower, tectonically driven processes.
Global climate has oscillated through greenhouse and
icehouse phases, in concert with aragonite and calcite
seas, respectively (Figure 10). During icehouse times of
continental glaciation (e.g., Pennsylvanian-early Permian,
Miocene-Pleistocene), high-frequency sequences on car-
bonate platforms were generated by eustatic sea-level
changes of 50–100 m. Subaerial exposure, unfilled
accommodation space, and conspicuous regional discon-
formities were common. Aggrading reef growth in a
keep-up mode is dominant in icehouse times, as rates of
production barely match those of accommodation space
increase. These large changes in sea level caused ramps
to have steep gradients and platform tops to have signifi-
cant depositional relief, characterized by pinnacle reefs
and erosional topography. Icehouse reefs are typically
dominated by heterotrophs or autotrophs with aragonitic
or high-Mg mineralogies.
During greenhouse times with little global ice (e.g., late
Cambrian-early Ordovician, Devonian, Triassic, Creta-
ceous), reef sequences were generated by small (possibly
<10 m or so) sea-level fluctuations. Reef cycles typically
consist of very shallow-water facies, with regional-scale
tidal flat caps and minor disconformities. Greenhouse reefs
are often either compound or progradtional as reef growth
could easily match or outpace the fastest rates of sea-level
rise. TST reefs, with plenty of accommodation space were
continuously in catch-up or keep-up mode, and this part
464 GENERAL EVOLUTION OF CARBONATE REEFS
of the reef is normally the thickest but narrowest. Ramps
often have very low gradients and platforms tend to be
progradational, with minor topography. Greenhouse reefs
can also have a more extensive range as carbonate settings
extend into higher latitudes due to elevated global tempera-
tures. Composition may also be more uniform, often dom-
inated by low-Mg calcite skeletal components.
Carbonate saturation
Living coral reefs are restricted to shallow-water tropical
and subtropical environments characterized by warm tem-
peratures, high light intensities, and also high aragonite
supersaturation. The growth of coral skeletons and other
calcifying organisms precipitates carbonate ions, forcing a
re-equilibration of the bicarbonate-dominated marine inor-
ganic carbon system which also creates a source of carbon
dioxide. A coral reef therefore represents the net accumula-
tion of CaCO3 and a source of CO2, which in turn probably
leads to feedback-controls on the global carbon cycle.
Carbonate saturation has probably varied dramatically
throughout geological time. It appears that Precambrian
oceans were highly oversaturated, but this may have
declined around the beginning of the Phanerozoic with
the appearance of numerous calcareous invertebrates. Since
then, Ca2þ in seawater has broadly followed sea-level
change such that Ca2þ levels were reasonably high during
the Cambrian-Mississippian and the Jurassic to Cretaceous.
It has been suggested that the geological distribution of
microbialites might be controlled by physicochemical fac-
tors, including the saturation state of seawater and/or
global temperature distribution (Webb, 1996). This may
also explain the decline in abundance of reef microbialite
after the Jurassic due to the increase of pelagic carbonate
that led to a reduced seawater saturation state. This would
have lowered supersaturation levels below a threshold for
abundant automicrite formation, thus restricting its forma-
tion to cryptic reef habitats, where abnormal chemistries
could have existed. Such a situation might also explain
the absence of Stromatactis in the late Phanerozoic.
Evolving ocean chemistry
The dominant form of precipitated crystalline CaCO3 has
oscillated during the geological past, with both inorganic
and organic production of aragonite and high-Mg calcite
dominating carbonate formation during cool periods (ice-
house), and low-Mg calcite predominating during warm
periods (greenhouse) (Figure 10). Such mineralogical
shifts are interpreted as markers for major changes in sea-
water chemistry.
Stanley and Hardie (1998) have proposed that the shifts
in the Mg:Ca ratio have controlled the predominance of
calcite versus aragonite secretors, particularly reef
builders, due to the inhibiting effect of high Mg2þ concen-
tration on calcite secretion. They suggest that the changing
Mg:Ca ratio of seawater has been controlled by variations
in the rate of production of oceanic crust, because oceanic
hydrothermal alteration is a major sink for Mg and an
important source of Ca. Experimental work has subse-
quently confirmed the profound influence of Mg:Ca sea-
water ratios on modern reef builders, including
scleractinian corals (Ries et al., 2006) and the calcareous
green alga Halimeda.
Scleractinian corals were, for example, important reef
builders in the Jurassic, but they did not build extensive
reefs during the greenhouse period (calcite seas) of the Cre-
taceous. During this period, species diversity remained high
but their abundance on carbonate platforms was low com-
pared to the Jurassic. Distribution shifted to outer platform
settings and higher latitudes (
35–45 N). There are many
hypotheses offered to explain these observations, including
the high temperatures, restricted circulation, and unstable
sediment conditions of Cretaceous platforms, and in partic-
ular the favoring of calcite-producing rudist bivalves over
aragonite corals (Wood, 1999). Rudist bivalves with their
outer calcitic skeletons underwent a dramatic radiation in
the late Cretaceous when the Mg:Ca ratio of seawater
reached exceptionally low values so favouring calcite over
aragonite secretors (Steuber, 2002).
Major shifts in the dominant composition of carbonate
skeletal particles through geological time also mirrors in
part these proposed changes in seawater chemistry and cli-
mate (Figure 10d), but mass extinctions also play a role by
triggering changes in the predominant form of CaCO3 pro-
duced by marine calcifiers (Kiessling et al., 2008).
Changes in the abundance of aragonitic organisms follow-
ing mass extinction events appear to have been predomi-
nantly driven by selective recovery rather than selective
extinction.
Evidence is also persuasive that the changing seawater
chemistry has influenced the style of early diagenesis in
carbonate regimes, particularly in reefs. The mineralogy
of early marine reef cements also seems to follow the same
secular changes. For example, aragonitic botryoids are
known exclusively from phases of aragonite seas (early
Cambrian, mid-Carboniferous to early Jurassic, and mid-
late Cenozoic), whereas radiaxial fibrous calcite is com-
mon in reefs that grew in calcite seas (particularly the
Ordovician to Devonian); it is virtually unknown from
aragonite seas of the Cenozoic. Enhanced rates of calcite
cementation during calcite seas due to the elevated abun-
dance of calcium ions may have promoted rapid lithifica-
tion of the reef framework and aided preservation of
cryptic biota that were otherwise vulnerable to distur-
bance, but this has yet to be documented.
The rise of predation and bioerosion
Many researchers have summarized the importance of her-
bivores and large marine vertebrates to the healthy func-
tioning of coral reefs, and this importance is
corroborated by analysis of the fossil record (see Wood,
1999, 2002). A dramatic escalation of new organisms with
innovative and destructive feeding methods occurred from
the mid-Jurassic to Miocene (summarized in Vermeij,
1987). In particular, the arrival of piscine herbivores had
 GENERAL EVOLUTION OF CARBONATE REEFS 465

the potential to fundamentally alter the dynamics of reef
and other benthic marine communities.
In general, grazers and carnivores throughout the
Palaeozoic and early Mesozoic were relatively small indi-
viduals with limited foraging ranges incapable of excavat-
ing calcareous substrates. A radiation during the Devonian
of durophagous, mobile predators has been proposed by
Signor and Brett (1984), but these forms probably relied
upon manipulation only to crush or ingest (Harper and
Skelton, 1993). By the early Mesozoic, sessile organisms
had to contend with an increasing battery of novel and
more advanced feeding methods as well as sediment dis-
ruption due to deep bioturbating activity (see summary
in Vermeij, 1987). Most notable was the rise of efficient
excavation behaviors.
Bioerosion notably increased in intensity from the mid-
to-late Jurassic. A radiation of borers occurred from the
Triassic onward, with deep borers (capable of penetration
greater than 50 mm) appearing from the Jurassic. Clionid
sponges – one of the major bioeroders on modern coral
reefs – had become abundant by the latest Jurassic. The
first live borers are known from the Eocene (Krumm and
Jones, 1993), as are fishes similar to modern reef faunas
(Bellwood, 1996). The ability for substantial excavation
of hard substrata over large areas increased considerably
from the latest Cretaceous-early Tertiary when deep-
grazing limpets, camerodont sea urchins, and especially
the reef fishes appeared. The complex pharyngeal appara-
tus of labrids was present at this time, and major labrid
clades were already differentiated (Bellwood, 1997).
Balistids first appeared in the Oligocene, and the oldest
scarid fossil capable of deep excavation currently known
is from the Miocene (14 Ma) (Bellwood and Schulz,
1991). It seems likely that sometime during the Oligo-
cene–Miocene, reef bioerosion gained a modern caste.
The abundance of reef fishes is assumed to be of great
importance on coral reefs, as evidenced by the dramatic
increase of algal growth as a result of their decline on
Jamaican reefs. Tropical marine hard substrata are usually
sparsely vegetated, but a rich algal flora develops when her-
bivorous fish are excluded and/or nutrient input increases.
Grazers not only allow the dominance of corals and coral-
line algae on coral reefs, they also contribute notably to car-
bonate sediment production and redistribution, algal ridge
formation, and the maintenance of overall diversity. Like
other predators, they can also ameliorate the effects of com-
petition and may combine with physical controls to pro-
duce the characteristic zonation of modern coral reefs.
The response of reefs to predation has been reviewed by
Wood (2002, in press). Table 1 outlines the major causes
and indirect effects of predation, particularly herbivory,
on coral reef communities. From this, we might be able
to make a series of predictions concerning changes in
post-Palaeozoic reef community ecology based on their
rise to abundance in the fossil record. In the sections fol-
lowing, these predictions are tested in order to determine
to what extent the evolutionary history of reef-bioerosion
and bioturbation has been bound with the ecology and
taphonomy of reef ecosystems.
Response to increased biological disturbance
Only skeletal anatomy and morphology, spatial distribu-
tion, and skeletal attack or breakage, and regeneration
might be detected – or inferred – in the fossil record of

General Evolution of Carbonate Reefs, Table 1 Predicted changes in reef community ecology and taphonomy based on the rise
to abundance of new predatory methods and endoliths as evidenced in the fossil record. (After Wood, 1999, 2002)

Event Prediction Timing

The rise of A shift to more conspicuous, well-defended macroalgae (coralline Late Mesozoic-Eocene
macroherbivores algae) on reefs
The rise of specialized Increase in diversity and retardation of dominance; reducing or Late Mesozoic-Eocene
predators preventing competition
Limiting of foraging ranges Late
Mesozoic-Eocene
Zonation: interaction of physical controls with differential effects Eocene
of damselfish in the survival of different coral species
The rise of excavatory A shift to organisms with deterrent traits and those that tolerate partial Late Mesozoic to Miocene
grazers and predators mortality
Increase in multiserial, branching corals Cretaceous onward
Increase in the diversity of the cryptos and other spatial refugia Jurassic onward
Algal ridge formation by coralline algae Eocene
Rise of intense bioerosion Reduced reef framework preservation Late Mesozoic to Miocene
and endoliths
Sediment grain size reduction Late Mesozoic to Miocene
An increase in skeletal sediment production Late Jurassic
Increase in multiserial scleractinian corals Throughout the history of
the group
The rise of parrotfish Formation of sediment aprons Miocene
Thick coralline algal crusts Miocene
Reduction in rate of reef progradation Miocene
466 GENERAL EVOLUTION OF CARBONATE REEFS
reef organisms. The appearance of these herbivores
paralleled profound changes in reef ecology, including
the rise of well-defended, highly tolerant coralline algae
(Steneck, 1985), a notable increase in branching corals
since the late Cretaceous (Jackson and McKinney,
1991), and the loss of many functional organisms that
prove to be intolerant to excavatory attack (Table 1). This
suggests a cause–effect system, where adaptation to pred-
atory attack has been intimately bound to the origin and
assembly of modern reefs.
Secure attachment to a hard substrate
Organisms without secure attachment to a stable substrate
are susceptible to the effects of all disturbances, including
bioturbation, as inversion and burial will cause death.
Most modern suspension feeders require a hard substrate,
even if these are only isolated patches within areas of
unstable, soft substrate.
Possession of an edge zone in all but the most primitive
scleractinian corals allows them to gain permanent attach-
ment to a stable substrate. As a result, scleractinian corals
dominate modern reef framework environments, espe-
cially those in high-energy settings, where there is also
an abundance of wave-swept, extensive hard substrata
for colonization. Permanent attachment also allows the
development of very large branching morphologies.
Cambrian archaeocyath sponges usually bore small
holdfasts that enabled limited attachment to hard sub-
strates (Figure 7a). But many mid-to- late Palaeozoic reefs
were dominated by large, sheet-like invertebrates
(stromatoporoid sponges, tabulate and rugose corals, and
trepostome and cystoporate bryozoans) that were initially
attached to small, ephemeral skeletal debris and then grew
over the surrounding sediment (Figure 5). Small,
branching forms (some stromatoporoids and bryozoans)
lacking extensive attachment sites were also common,
and they were presumably partially rooted in soft
sediment.
The late Paleozoic decline of immobile epifauna coin-
cides with the rise of major bulldozing taxa, which passed
through the end-Permian extinction unscathed. This
coincidence must remain conjectural until tested
experimentally.
Resistance to partial mortality
Predation that actively excavates underlying skeleton
often results only in partial mortality, that is, sublethal
damage. In such cases, the capacity of the prey to heal or
replace damaged areas of soft tissue becomes critical to
survival. Strategies that rely upon herbivores/predators
to remove competing algae therefore often entail the loss
of the prey’s own tissues.
In coralline algae, a protective outer epithallus and con-
ceptacles that contain reproductive structures have been
demonstrated to protect the delicate reproductive anatomy
from intensive grazing (Steneck, 1982). Many coralline
algae can also tolerate intense herbivory due to their abil-
ity to rapidly regenerate removed material. Thickened
crusts are more tolerant to attack than thin encrusting or
branching forms (Steneck, 1985), but in modern reefs,
the dominance of a particular growth form appears to be
a trade-off between the cost of investment in increased
defence, and the reduction in growth rate or competitive
ability. As a result, thickened crusts dominate only in areas
of high wave energy and biological disturbance.
After the Eocene, herbivore-susceptible, delicately
branched coralline algae reduced in abundance in the tro-
pics, the proportion of thickened encrusting forms
increased, and the first algal ridges appeared – all coinci-
dent with the rise of excavatory herbivorous fish.
Many sessile reef organisms possess a modular or colo-
nial habit where partial predation and boring may remove
either individual or a few modules, or large areas may be
cleared of living tissue, sometimes together with the exca-
vation of underlying skeleton. But the modular organiza-
tion also reduces soft tissue to a relatively thin veneer
over a larger basal skeleton. This not only decreases acces-
sibility and the ease of prey manipulation by predators but
also minimizes the tissue biomass while maximizing the
cost of collection. For example, in a typical domal colony
of Porites, only about 0.5% of the colony’s radius is occu-
pied by soft tissue (Rosen, 1986). In branching and platy
colony forms, the relative proportion of skeleton is even
higher.
Cambrian archaeocyath sponges show a steady and
marked increase in the proportion of complex modular
forms during their history (Wood et al., 1992), as do
scleractinian corals since the mid-Triassic, which appears
to be uninterrupted by the end-Cretaceous extinction event
(Coates and Jackson, 1985).
Regeneration after breakage
Some morphologies are more resistant to breakage than
others. For example, colonies with closely spaced
branches can make predator access difficult by forming
hidden, protected areas. The flattening of branch termina-
tions can also offer greater resistance to all forms of break-
age and shearing, and this character is found in erect
species of bryozoans, gorgonian corals and stylasterine
corals.
A multiserial modular organization, however, in addi-
tion to promoting architectural diversity and flexibility,
also allows compartmentalization of damage and enables
some colonies to regenerate from fragments. Most signif-
icantly, branching corals also show tremendous powers of
regeneration: Acropora palmata has one of the highest
rates recorded. Indeed, unlike massive, platy or encrusting
forms, damage to branching corals often leads to an imme-
diate increase in growth rate so causing an increase in size
rather than simply repairing damaged tissue. The staghorn
coral (Acropora cervicornis) is able to re-anchor frag-
ments and rapidly regenerate and grow, often fusing with
other colonies. Such branching corals have turned adver-
sity into considerable advantage, and appear to flourish
because, and not in spite, of breakage.
 GENERAL EVOLUTION OF CARBONATE REEFS
The percentage of scleractinian erect species decreased
until the late Cretaceous (Turonian), but increased mark-
edly – particularly in multiserial forms with inferred rates
of rapid regeneration – after that time. This spectacular
rise of various morphologies of branching forms was coin-
cident with the appearance of new groups of predatory
excavators.
All families of modern scleractinian corals that domi-
nate reefs today spread throughout the Tethys Sea during
the Eocene. The poritids, their relatives the actinids, and
the favids (which had survived the Cretaceous extinction)
dominate most coral reef communities throughout much
of the Cenozoic (McCall et al., 1994). Although branching
acroporoids appeared in the Eocene, they did not dominate
reefs until early Pleistocene. The rise of this group – with
its particularly remarkable powers of regeneration from
fragmentation and rapid growth – would then seem to be
independent of any known changes in predation style.
Patterns of sediment removal and storage
Unlike modern reefs, anecdotal evidence suggests that
a greater proportion of reef framework is preserved in situ
before the Jurassic (Wood, 1999). Many Palaeozoic reefs
commonly preserve intact reef frameworks, even of fragile
biota such as frondose bryozoans (Figure 6b) or platy
stromatoporoid sponges. Such preservation was aided, in
part, by abundant and probably rapid synsedimentary lith-
ification, particularly cementation.
We might predict that substantial aprons of sediment
may not have been present on pre-Eocene reefs before
the evolution of reef fish, and likewise in the absence of
the grain size reduction activities of clionid sponges, echi-
noids, fish, mean sediment grain size may have been more
coarse prior to the late Jurassic, perhaps resulting in
a reduced net loss of carbonate to the system through the
removal of fines. It is possible also that the modern style
of coral reef lagoon may also not have appeared until the
late Jurassic or later (Wood, 1999, 2010).
The role of mass extinctions
Reefs have long been thought to be highly susceptible to
mass extinction events, with major reef-building biota typ-
ically going largely or wholly extinct. Metazoan reefs may
take some 2–8 Myr to recover after such events, some-
times longer than other communities, and post-extinction
reefs are often composed of entirely new groups of skele-
tal metazoans.
In some cases, either calcimicrobes or microbialites can
dominate post-extinction reefs, but these elements were
often already part of the pre-extinction reef community.
Shallow-water Frasnian (late Devonian) reefs show abun-
dant large skeletal metazoans (particularly stromatoporoid
sponges, and subsidiary corals) as well as extensive fenes-
tral microbialite and calcimicrobes both as free-standing
mounds (Figure 5b). Succeeding Famennian reefs grow-
ing in the aftermath of a mass extinction were composed
of frame-building microbialite and calcimicrobes (e.g.,
Renalcis and Rothplezella) and a diverse community of
467
spiculate sponges. The first metazoan reefs to form after
the end-Permian extinction in the Anisian (Lower Trias-
sic) bore large framework cavities, populated with the cal-
cified microbe Cladogirvanella.
Tropical shallow marine carbonate production has been
noted to fall after some mass extinctions (e.g., the late
Devonian and end Permian), but not others (end Creta-
ceous). Changes in seawater saturation or chemistry may
also explain the response of reef and other carbonate-
producing biota to mass extinctions. When skeletal groups
first appear, they tended to adopt the mineralogy favored
by ambient seawater chemistry, but subsequent extinction
events may have preferentially removed those groups and
selected for new skeletal biota whose skeletal mineralogy
reflected the new ocean chemistry (Kiessling et al., 2008).
Summary
Reefs have shown some unifying features over their
3.5-billion-year history. Reefs have shown zonation in
response to environmental gradients from their inception,
and metazoan reefs have been differentiated into open
surface and cryptic reef communities for their entire

550-million-year history (Wood et al., 2002).
Tectonics drives many aspects that control reef growth
including sea level, circulation patterns, climate, and
evolving seawater chemistry change over long timescales.
Reefs have always been most numerous on passive conti-
nental margins at low latitudes where they typically form
prograding or aggrading barriers (with growth during
transgressive and highstand periods and erosion during
longer low stand periods), or fringing reefs. By contrast,
reef-forming toward the cratonward side of foreland
basins is relatively rapid and reef geometries more aggra-
dational. Finally, reefs forming in strike-slip basins and on
thrust complexes are affected by the vagaries of local tec-
tonic movements, which may be highly episodic but
involve substantial displacements and are thus highly
unpredictable (James and Wood, 2010).
Climate is a major controlling force on the evolution of
reefs on both short and long timescales, as the latitudinal
range of carbonate-producing species is largely governed
by temperature and carbonate supersaturation. Reef growth
therefore shows cyclicity at all scales in response to short-
term oscillations (e.g., Milankovitch and glacial–interglacial
cycles) as well as to longer-term climatic intervals, driven by
slower, tectonically driven processes.
Global climate has oscillated through greenhouse and
icehouse phases, in concert with aragonite and calcite seas,
respectively. During icehouse times of continental glacia-
tion, high eustatic sea-level changes created aggrading
reef growth, common subaerial exposure and conspicuous
regional disconformities in reef complexes. These large
changes in sea level caused ramps to have steep gradients,
and platform tops to have significant depositional relief
characterized by pinnacle reefs and erosional topography.
Icehouse reefs are typically dominated by heterotrophs or
autotrophs with aragonitic or high-Mg mineralogies.
468 GENERAL EVOLUTION OF CARBONATE REEFS
During greenhouse times, with little global ice, reef
sequences were generated by small sea-level fluctuations.
Reef cycles typically consist of very shallow-water facies,
with regional-scale tidal flat caps and minor disconfor-
mities. Greenhouse reefs are often either compound or
progradational as reef growth could easily match or
outpace the fastest rates of sea-level rise. Ramps often
have very low gradients, and platforms tend to be
progradational, with minor topography. Greenhouse reefs
can also have a more extensive range as carbonate settings
extend into higher latitudes due to elevated global temper-
atures. Composition may also be more uniform, often
dominated by low-Mg calcite skeletal components.
Biological disturbance, that is bioturbation, boring, and
excavatory predation and herbivory, has clearly escalated
since the Mesozoic. Reef biotas have responded with the
proliferation of traits with proven anti-predatory benefits,
particularly regeneration after partial mortality. Indeed
some modern dominant reef taxa, such as branching corals
and coralline algae, appear not only to thrive, but actually
require conditions of considerable disturbance for their
survival in shallow tropical seas (Wood, 1999, 2002).
Many modern reefs are largely reduced to rubble and
sand via physical abrasion and bioerosion. Modern coral
reefs are also dominated by branching forms due to their
high diversity and abundance, propensity to proliferate
via fragmentation, and resilience to taphonomic destruc-
tion. Many reefs prior to the Jurassic show the preservation
of intact, in situ frameworks. Most epibenthic, soft-
sediment dwelling organisms typical of Paleozoic reefs
appear to have become largely absent from shallow marine
tropical reef biotas during the late Paleozoic to early
Mesozoic, perhaps due to intolerance of the rise of deep
burrowing taxa and excavatory attack (Wood, 2010).
Scleractinian corals, in particular branching taxa, show
a marked increase in the proportion of forms with complex
modularity from the Eocene onward, even though corals
displayed the full range of morphological forms and
corallite size by the Late Triassic. Highly defended, thick
crusts in coralline algae become more dominant, and
branching forms also become noticeably less conspicuous
on reefs from the Eocene onward. This major reorganiza-
tion of the coral reef ecosystem coincides with the rapid
appearance and radiation of herbivorous and coral-eating
reef fish, but this remain to be tested experimentally
(Wood, 2002).
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Cross-references
Accommodation Space
Aragonite
Back-Stepping
Bioerosion
Calcite
Climate Change and Coral Reefs
Devonian Reef Complexes of the Canning Basin
Dolomitization
Diagenesis
Fossil Coralline Algae
Framestone
Mass Extinctions, Anoxic Events and Ocean Acidification
Oil and Gas Reservoirs and Coral Reefs
Permian Capitan Reef System
Porosity Variability In Limestone Sequences
Sea Level Change and Its Effect on Reef Growth
Stromatolites
Submarine Lithification
Taphonomy
Tethys Ocean
GEOMORPHIC ZONATION
Paul Blanchon
National Autonomous University of Mexico, Cancun,
Mexico
Introduction
The coral reefs of the Atlantic, the Caribbean and the Indo-
Pacific do not differ fundamentally in their structural forms,
their habitats and the interaction of their species, even though
the organisms occupying specific ecological roles vary
greatly between oceans and even between individual reefs.
Goreau et al. (1979, p. 135)
Despite the similarity between the geomorphology of
coral reefs noted by the Goreaus and others (Wells,
1957; Stoddart, 1969), much of the emphasis in character-
izing coral-reef zonation has been ecological using either
statistically defined coral assemblages or more simply,
470 GEOMORPHIC ZONATION
the morphology of the corals themselves (Wells, 1957;
Rosen, 1975; Geister, 1977; Pichon, 1978; Chappell,
1980; Done, 1982; Done, 1983). Most of these studies
have found that shallow coral assemblages show
a distinct zonation as wave energy, and hydrodynamic dis-
turbance varies with depth and margin exposure. For
example, Done (1982) found that shallow communities
changed with a decreasing cross-shelf wave exposure in
the central Great Barrier Reef. Individual reefs also
showed the same pattern, with shallow-wave-adapted
communities being replaced along leeward margins by
upward shifts in the positions of deeper communities.
Growth form of corals also show similar zonation patterns
in response to depth and exposure-related variation in
wave energy (Pichon, 1978; Chappell, 1980; Madin and
Connely, 2006). Such zonation patterns have led to
a clear distinction between rough-water reefs exposed to
trade-wind-, swell-, and storm-generated wave fields and
calm-water reefs protected from them (Geister, 1977,
1980).
A comparison of zonation schemes, however, shows
that the coral assemblages in each zone can be highly var-
iable on both inter-reefal and regional scale, and there is
a significant overlap between the members of each assem-
blage zone (e.g., Done, 1982). Despite this “coral-scale”
variability, the geomorphology of these reefs is very sim-
ilar. Indeed, equally similar reef structures are produced
in areas with order-of-magnitude differences in species
abundances, and even where reefs have depauperate
species numbers, their basic geomorphology is identical
(e.g., Glynn et al., 1996). This consistency implies that
gradients in wave exposure not only control the main
environmental tolerances of shallow coral assemblages
but also the form of the structures they build (Graus
et al., 1977; Graus et al., 1984, 1985; Graus and
Macintyre, 1989). It therefore follows that reefs might be
better classified according to their geomorphic form
(Wells, 1957).
Until very recently, a synoptic view of reef geomor-
phology showing the impact of varying ecological pro-
cesses has not been possible, and previous attempts to
summarize information have had to rely on parochial
descriptions of what was accessible or visible at the time
(e.g., Hopley, 1982; Guilcher, 1988). Following this
approach, vast areas of modern reefs have been inade-
quately characterized and, with some exceptions, quanti-
tative differences and the similarities between different
regions have gone undocumented. This deficient situation
is about to change. The recent advent of widely available
orthorectified satellite imagery (Google Earth) and rapid
development and cost reduction in geographic informa-
tion systems, multi-beam sonar, LiDAR, and other
remote-sensing technologies are making large-scale mor-
phometric quantification of reefs feasible (Naseer and
Hatcher, 2001; Andrefouet et al., 2001; Storlazzi et al.,
2003; Naseer and Hatcher, 2004). As such, we are on the
cusp of being able to provide a more holistic view of mod-
ern coral-reef form and a development that will provide
timely insight into the resilience and resistance of reefs
to human-induced global environmental change.
In this entry, I examine satellite imagery using Google
Earth along with local descriptions of reefs and attempt
to summarize what is presently known about the geomor-
phic zonation of well-ordered, rough-water reefs, outline
their formative processes, and assess their development.
Standard geomorphic reef zonation
A comparison of well-ordered, rough-water reefs shows
a remarkable degree of conformity between the size and
character of their major morphological features
(Figure 1). All of them consist of five first-order or “stan-
dard” zones: a calm reef lagoon, with varying number of
patch reefs; a largely barren, shallow reef flat strewn with
detritus; a narrow, gently sloping reef front exposed to
waves and swells; a steeper reef slope bearing corals to as
deep as 100 m; and a less steeply inclined fore-reef apron,
where coral and reef detritus accumulate. The boundaries
between these standard zones are delineated by simple
slope breaks. The break between the lagoon and the reef flat
is either a sand slope or a small reef scarp. The break
between the level reef flat and the gently sloping reef front
occurs at the reef crest. Similarly, the shelf break separates
the gently sloping reef front from the steeper reef slope, and
the break between the reef slope and fore reef is the transi-
tion into slopes that reflects the angle of sediment repose.
Each of these standard and easily defined reef zones can
in turn be subdivided into commonly recurring second-
order zones or subzones, which may not be present on all
reefs and, if they are present, may vary between margin,
area, and region. As a consequence, their boundaries can
be more difficult to define.
Lagoon
With their calm, stable environments, and depth ranges
that are usually within the range of coral growth, lagoons
potentially represent an ideal and extensive habitat for
reefs (Figure 2). The actual variation in lagoonal reef
development however is striking, and even in areas where
lagoonal reefs are prolific, such as the Tuamotus, there are
examples where reefs are completely absent (Adjeroud
et al., 2000; Paulay and Kerr, 2001).
Patches, knolls, and pinnacles
Lagoons with significant reef development show two
main types of subcircular structures, patch reefs and
knolls. Patch reefs reach mean sea level and rise from
substrates <20 m deep; where the substrate is deeper,
they are termed pinnacles. Knolls are submerged reef
patches but can also develop into pinnacles where they
have >20 m relief (Stoddart, 1969). Patches, knolls, and
pinnacles can develop in a dispersed pattern or can
become connected to form linear ridges with various
arrangements (e.g., Maxwell, 1968; Hopley, 1982).
Dispersed patches or knolls have a density continuum
from widely spaced to a point where they connect with
 GEOMORPHIC ZONATION 471

Caribbean
Trade dominated
Belize

Patch Sand Coralgal Gravel Spur and Sand Chute and

and knoll flat flat rim groove terrace buttress

Shelf break
Sand slope

Scarp
Crest
Reef Reef Reef Reef
lagoon flat front slope

Shelf break
Sand slope

Scarp
Patch Sand Coralgal Algal Crest Spur and Sand Chute and
and knoll flat flat rim groove terrace buttress

Marshalls Swell dominated

Arno

Geomorphic Zonation, Figure 1 Geomorphic zonation of rough-water reefs. Both swell- and trade-wind-dominated reefs develop
five standard reef zones: lagoon, reef flat, reef front, reef slope, and fore reef (not shown). These zones are delineated by simple slope
breaks: the sand slope between lagoon and reef flat, the reef crest between the reef flat and reef front, the shelf break between the
reef front and reef slope, and the break between the subvertical lower slope and the talus cones of the fore-reef sediment. These
standard zones are divided into several widely occurring subzones, which vary with respect to margin orientation, area, and region.
Images copyright of Google Earth, DigitalGlobe and GeoEye, 2010.

neighbors (Figure 2). Connected patches or knolls can
further develop into ridges with sinuous, stellate, or retic-
ulate arrangements, with the latter closing off distinct
areas of deeper water, producing a cellular pattern of reef
development. This cellular structure develops at various
scales and can produce cells as large as 1,000 m in diam-
eter (Figure 3).
Many lagoonal reefs have diverse coral assemblages,
which can develop a distinctive windward/leeward as well
as depth zonation (Wells, 1957). In Tikehau, for example,
emergent patch reefs are topped by skeletal sand and
cobble-sizedcoral gravel. The windward side has large
head corals of 6 m, and leeward and deeper sides are
covered by branching acroporids to depths of 15 m
(Intes and Caillart, 1994). In addition, there can be lateral
gradients in coral zonation within individual lagoons as
a consequence of various environmental gradients (Paulay
and Kerr, 2001).
An obvious control on lagoonal reef development is the
degree of water exchange with the open ocean. During
World War II, causeway construction blocked spillways
over reef flats and isolated lagoonal sections, causing mass
mortality of corals (Maragos, 1993). This led to the reali-
zation that a dominant exchange mechanism was related
to reef flat currents driven by wave setup and not necessar-
ily to tidal exchange (Atkinson et al., 1981; Andrews and
Pickard, 1990). This has been most clearly demonstrated
by Callaghan et al. (2006), who showed that during most
472 GEOMORPHIC ZONATION

Geomorphic Zonation, Figure 2 Lagoonal reef development
showing the varying arrangements and densities of patch reefs
(emergent) and knolls (submerged). Images copyright of Google
Earth, DigitalGlobe and GeoEye, 2010.
Geomorphic Zonation, Figure 3 Reticulate reef development
in lagoons shown at the same scale to emphasize the varying
scales of cells enclosed by reticulate networks. In the case of
Houtman-Abrolhos, subsurface data show that the networks are
the result of coral growth during the Holocene. Images
copyright of Google Earth, DigitalGlobe and GeoEye, 2010.
 GEOMORPHIC ZONATION
of the tidal cycle, incident wave energy on the windward
margin pumps water onto reef flat and into the lagoon,
while water exits over the leeward reef flat and through
channels. Wave pumping causes lagoon water levels to
remain above the ocean at all tidal phases and to rise sig-
nificantly during periods of enhanced wave activity (see
also Kraines et al., 1999). As a consequence, exchange is
directly related to the proportion of reef flat that is sub-
merged or dissected by channels and, where tidal channels
are narrow or absent, island formation and sediment depo-
sition on the reef flat can play a key role (Kench and
Mclean, 2004). The impact of water exchange and wave
pumping on lagoonal reef development has been clearly
illustrated in the Tuamotus, where Adjeroud et al.
(2000) reported that reef pinnacles were more abundant
and diverse in bigger atolls, which had either large chan-
nels or a high proportion of submerged reef flat. Con-
versely, they found that smaller atolls with reduced
exchange due to reef-flat island formation were typified
by low pinnacle abundance and diversity.
Cellular reefs
Reef development in lagoons can also be self-limiting due
to a negative feedback between growth and circulation.
For example, prolific development of pinnacles in protected
lagoonal areas of the Houtman Abrolhos reef complex, off
the western Australian coast, has produced a cellular reef
pattern, as shown in Figure 3 (Collins et al., 1996; Wyrwoll
et al., 2006). The reef tops have a dense cover of branching
acroporids, which only extend 7–10 m down the steep
walls of the cells. In some areas, the walls overhang and
show collapse scars, which expose open frameworks of
the same corals; the floors of the cells are between 10-
and 30-m deep and covered with a thick skeletal sand and
branch-coral gravel. Their water columns are stagnant and
stratified and do not support coral growth below 10 m. Drill
cores and seismic data confirm that these cellular reefs are
composed of branching acroporid frameworks that devel-
oped during the last 7–10 ka over a flat Pleistocene reef ter-
race at
40-m depth (Collins et al., 1996; Wyrwoll et al.,
2006). These studies in the Houtman-Abrolhos show that
the prolific growth of branching acroporids has led to the
coalescence of isolated pinnacles into reticulate structures
that have enclosed deeper cells, causing restricted circula-
tion and self-limitation of coral growth. Reticulate and cel-
lular reef development is therefore one of the several
examples of self-organization in coral reefs and is an emer-
gent property of the synergy between rapid lagoonal reef
growth and circulation.
Formation of lagoonal reefs
The development of lagoonal reefs, particularly cellular
reef forms, has been persistently linked to antecedent karst
due to its similarity to polygonal and cockpit-cone forms
that develop in humid tropical karst terrains (MacNeill,
1954; Purdy, 1974; Purkis, 2010). However, as alluded
to above, drilling and seismic profiling has consistently
473
shown that reef forms are growth induced, either develop-
ing from scratch during the present interglacial or continu-
ing atop development that took place during previous
interglacial stages (Harvey and Hopley, 1981; Hopley,
1982; Smith et al., 1998). On Cocos-Keeling, seismic data
collected by Searle (1994) showed that the modern reticu-
late reef development was largely growth induced but was
influenced by a combination of fossil reef growth as well
as karst-induced collapse structures in the underlying sub-
strate. In other areas, positive relief provided by previous
episodes of interglacial reef growth has likely played
a negative role in modern lagoonal reef development.
For instance, the rims of Pacific atolls are often
multigenerational reef structures, which developed during
previous interglacials (e.g., Szabo et al., 1985); where
these rims are continuous, they must have prevented
lagoonal reef development for significant time intervals
until the sea level overtopped them and initiated their
modern structure
8 ka ago. This delay in development
may explain the anomalous paucity of reefs in some of
the deeper modern atoll lagoons.
For other forms of lagoonal reef development however,
antecedent substrates have provided little to no guidance
for subsequent events. On John Brewer Reef on the
Great Barrier Reef (GBR), Walbran (1994) obtained seis-
mic profiles showing that both lagoonal patch reefs and
the reef rim are underlain by a flat Pleistocene surface at
20-m depth. The same was also found at Heron and Sykes
reefs in the Capricorn Group by Smith et al. (1998) and in
the Houtman-Abrolhos reef complex by Collins et al.
(1996). Modern reef growth in lagoons and other zones
is clearly capable of generating its own structures with lit-
tle help from antecedent templates of any kind.
Reef flat
In contrast to the high-relief patches and pinnacles of the
lagoon, reef flats are remarkably level surfaces and their
low-tide elevation varies little for hundreds of meters. If
there is any gradient at all, it tends to slope back from
the outer reef flat toward the inner, and its lagoonal margin
water depths are no more than 1–3 m. In the absence of
islands and storm ramparts, the width of open reef flats
in atoll groups such as the Marshalls and Tuvalu is
between 500 and 1,000 m, but exceeds this in the
Tuamotus and Maldives where flats with widths of 2,000
m are common. The presence of islands and storm
ramparts however tends to significantly reduce reef-flat
width by impeding sediment transport into the lagoon.
As recognized by Hopley (1982), open reef flats consist
of two common subzones: an outer “living zone” com-
posed of an algal rim and a coral algal flat and an inner
sand zone mantled by thin sheets of shifting sediment
(Figure 1).
Algal rims
Algal rims are usually less than 50-m wide and form the
highest part of the reef flat. Two types of algal rim are
474 GEOMORPHIC ZONATION

common and have been described from the Marshalls,

Tuamotus, Tonga, and the southern Maldives (Tracy
et al., 1948; Ladd et al., 1950; Wells, 1951; Newell,
1954; Wells, 1957; Stoddart, 1966; Nunn, 1993). The first
type is an elevated
1-m-high ridge (above mean low
water) fronting the most exposed reef sections, normally
associated with islands (ridges as high as 2.5 m have been
reported from the southern Tuamotus were storm swell is
severe; Salvat and Salvat, 1992). This high ridge type is
commonly dissected by surge channels, 1-m wide and as
much as 3-m deep, which are inshore extensions of reef-
front grooves that have been narrowed by corallines
(Figure 4). These channels extend back into the reef flat
for 30–100 m and drain water pumped onto the flat under
large swells or strong trade wind conditions. The channels
can become completely roofed over and develop blow-
holes surrounded by terraced coralline mounds that rise
to as much as 1 m above the surrounding reef flat. The reg-
ular spacing of channels divides the algal ridge into but-
tresses, which form the heads of reef-front spurs (Tracey
et al., 1948). The second type of algal rim develops on
open reef flats and consists of a broader arch that rises less
than 0.5 m but is not dissected by surge channels. Less
commonly, rims consist of groups of pillars, piers, or but-
tresses that have been roofed over by corallines to form
room-and-pillar structures (Tracey et al., 1948). Sheltered
pockets and caverns in these complex structures can be
capacious and may support limited coral communities.
But in general, algal rim substrates are dominated by crus-
tose corallines, particularly Porolithon onkoides, which
requires continuous disturbance from wave surge to pre-
vent competitive exclusion by fleshy algae (Littler and
Doty, 1975). General rates of accretion of this coralline
have been measured up to 10 mm/year (Steneck, 1986;
Adey and Vassar, 1975), but these are clearly sufficient
to build and maintain algal rims.

Coralgal flat
A good proportion of the reef-flat zone is covered by
a coralgal flat, which is an intertidal platform composed
mostly of coral gravel, crustose corallines, and scattered
corals. Coral cover tends to rapidly diminish away from
the rim due to the increasing temperature and salinity var-
iations in the shallow waters of the reef-flat interior (Wells,
1951). The highest coral cover is usually found in a broad
trough developed immediately behind the low algal
mound type along open flats (Wells, 1957). Here luxuriant
patches of coral cover 50% of the surface and develop up
to low tide. Between corals, the gravel substrate is stabi-
lized by crustose corallines to form a smooth pavement.
In front of islands, where there is less coral cover, the
trough is clearly seen to be crossed by rimmed, partially Geomorphic Zonation, Figure 4 Reef flats forming in front of
roofed, or filled surge channels whose spacing reflects that islands, showing algal ridges, back-ridge troughs cut by surge
of the reef-front grooves (Figure 4). Behind the trough is channels, and the coralgal zone. Surge channels align with reef-
a largely barren surface covered only by a few centimeters front grooves and provide evidence of reef-flat progradation. In
the Marshalls, this has occurred in jumps producing wide
of water at low tide. In places, several generations of troughs. Images copyright of Google Earth, DigitalGlobe and
eroded algal-ridge remnants protrude above the flat GeoEye, 2010.
 GEOMORPHIC ZONATION
surface by as much as 1 m and clearly represent former
positions of the algal rim (Ladd et al., 1950; Scoffin
et al., 1985). Also protruding are boulders of coral frame-
work eroded from the reef front and thrown up onto the
reef flat during cyclones. These become welded to its sur-
face by corallines and often develop an intertidal notch
produced by intense bioerosion (Stoddart, 1969).
On other swell-dominated reefs, troughs are absent and
the coralgal flat behind the rim consists of a very distinc-
tive striated zone, especially in areas with high tidal ranges
such as the GBR and western Indian Ocean. This consists
of elongate strips of reef flat consolidated by corallines,
sometimes colonized by corals, and separated by furrows
of unconsolidated skeletal sand and gravel, which deepen
and widen inshore (Hopley, 1982). Although coral cover
varies, the furrows are persistent features, which can
extend across the entire coralgal zone. At Mayotte, sand
and gravel stripes clearly continue in alignment with the
reef-front spur and groove, and have a height differential
of 10–50 cm. At Mahé in the Seychelles, 2–3-m-high sta-
bilized gravel ridges narrow inshore and are separated by
1–2-m-deep furrows, which widen and deepen inshore
(Guilcher, 1988).
Sand flat
The innermost sides of open reef flats, adjacent to the
lagoon, are dominated by shifting sand that is either
washed back from the seaward rim or, when winds switch
direction during storms or monsoons, from the lagoon
itself. The predominant movement of sediment in this
zone occurs during tropical cyclones when large amounts
of sand covering islands in the reef-front reservoir is
remobilized and deposited as spill-over lobes, which pro-
grade into the lagoon, forming a distinctive sand slope.
This general sediment movement into the lagoon is
interrupted by islands, which provide shelter from sedi-
mentation and foster patch-reef development along this
boundary. These patches can become well developed and
form a linear reef with a steep lagoon-side scarp.
Islands and storm deposits
Low-lying sand islands, and their associated channels
(hoas) and gravel ramparts, constitute a large proportion
of swell-dominated reef flats and form features which
are dynamic in both space and time. Although these fea-
tures are formed by a combination of storm and
fairweather processes (Kench and Brander, 2006), storms
assume a greater importance because they generate gravel
through the destruction of corals, remobilize inactive sed-
iment deposits on islands and in sub-wavebase reservoirs,
and provide a continuous force on geological timescales.
For these reasons, storms likely play a major role in initi-
ating island formation and causing major subsequent
changes (Baines et al., 1974; Bayliss-Smith, 1988).
Reports of the impacts of tropical cyclones indicate that,
in addition to erosion, three kinds of reef-flat deposit are
common: outer-flat gravel bars, coastal ramparts along
475
existing islands, and aerially extensive gravel sheets over
both the flat and islands (McKee, 1959; Blumenstock,
1961; Baines et al., 1974). The outer-flat gravel bars are
composed of cobble to boulder-sized coral clasts, which
are eroded from the reef front and typically deposited
20–30 m inshore from the algal rim. Under fairweather
conditions, these gravel bars rapidly migrate inshore and
are eventually left as marine cemented lenses of gravel
spread out over the reef flat
100 m from the outer rim;
these are known as conglomerate platforms (Hopley,
1982). Such immobile lenses commonly provide a locus
for subsequent sedimentation, and storms can add exten-
sive shore-attached gravel ramparts for considerable dis-
tances along and between platforms (Baines et al.,
1974). However, the most volumetrically important reef-
flat deposits consist of thin but aerially extensive sheets
of finer-grained cobble to pebble gravels that can become
rapidly stabilized either in island settings by marine
vadose cements to form “cay rock,” or on open reef flats
themselves by crustose corallines (Newell and Bloom,
1970; Marshall and Jacobson, 1985). Successive deposi-
tion and cementation of gravel lenses and sheets can pro-
duce island platforms that reach 2 m or more above
present mean sea level. Dating of these platforms in sev-
eral areas has yielded anomalously old ages and, as
a result, it has been claimed that they represent widespread
reef-flat development during a regionally higher mid-
Holocene sea level (e.g., Dickinson, 2004) despite the fact
that clasts in these deposits have large age ranges (e.g.,
Scoffin et al., 1985).
Formation of reef flats
The paucity of active coral growth and the flatness and
low-tide elevation of many Indo-Pacific reef flats have
been a continued source of confusion and, as alluded to,
the age and origin of gravel deposits have provoked partic-
ular controversy (e.g., Newell and Bloom, 1970).
A persistent idea is that reef flats have been eroded to their
present low-tide level by a slight fall in regional sea level
over the last several 1,000 years (e.g., Cloud, 1952).
A variation of this has been advocated by Pirazzoli and
Montaggioni (1988) and Woodroffe (2008), who
suggested that the relative sea-level fall during past few
millennia has produced many emergent reef flats, which
are developed at elevations too high to be suitable for coral
growth. This implies that reef flats were constructed by
coral framework prior to the inferred sea-level fall; yet,
drill cores through reef flats consistently show that they
are not composed of thick framework but of detrital
deposits (Henny et al., 1974; Marshall and Jacobson,
1985). Hence, the present reef-flat surface is a reflection
of its interior, and an absence of corals and low-tide eleva-
tion does not require an erosional explanation. In addition,
elevation data used to infer higher sea level have not
accounted for elevated water levels on reef flats due to
wave pumping and ponding of water behind storm
ramparts, and elevations used from relict algal ridges are
476 GEOMORPHIC ZONATION
commonly no higher than their modern counterparts. And
even where they are, possible changes in wave climate in
controlling their elevation rather than sea level are not
taken into account. Moreover, in all of the reef-flat studies
claiming regional sea-level highstands, the error range of
the sea-level indicator is the same magnitude as the sea-
level change itself (Woodroffe and Horton, 2005). Thus,
claims that modern reef flats are somehow palimpsest
and have been modified by falling sea level lack adequate
support.
Other analyses of reef-flat form and zonation have
assumed that flats did not develop until reefs caught up
with sea level, despite the fact that the evidence for reef
catch-up in most areas is equivocal due to inadequate core
coverage (Blanchon and Blakeway, 2003). For example,
Hopley (1982) proposed a retrogressive model of reef-flat
development on the Great Barrier Reef in which presumed
catch-up reefs progressively developed a zoned reef flat
through time as reef framework and sediment migrated
into the lagoon. A prediction of this hypothesis is that
deposits should get younger toward the lagoon; yet, drill
cores consistently show the opposite, with reef-flat ages
increasing toward the lagoon in almost all the areas stud-
ied in sufficient detail (Easton and Olson, 1976; Pirazzoli
et al., 1987; Randall and Siegrist, 1988; Takahashi et al.,
1988; Kan et al., 1997a; Scoffin and Le Tissier, 1988;
Woodroffe et al., 2000).
This widely documented pattern of inshore reef-flat
ageing is more consistent with progradation, whereby
the algal rim builds seaward out over the reef front through
time (Ladd et al., 1950; Pirazzoli et al., 1987). Indeed,
such a pattern would not only explain why reef flats are
flat, with no space for framework accretion, but it would
also account for many of their morphological features.
For example, the widespread occurrence of back-rim
troughs and surge channels extending 100 m or more into
the reef flat interior is a key indication of this process
(Figure 4). Imagery from the Marshall Islands in particular
indicate that back-rim troughs represent shallow spur and
groove that were abandoned due to algal-rim development
along their leading edges (see Bikini and Rongelap, in
Figure 4). The jump in the rim position to the spur front
consequently left a shallow trough between the old and
new rim that provided space for coral growth. In many
areas, coral growth and gravel deposition rapidly filled
this trough and only surge channels remain as evidence
of its existence. In this way, reef flats can prograde rapidly
and produce an extensive flat surface devoid of coral
growth that is punctuated only by abandoned algal rims
and boulder thrown up during storms. Why algal rims
are periodically abandoned and jump seaward is related
to the dynamics of spur-and-groove development, which
is outlined below.
Reef front
The reef front zone extends from the reef crest to the shelf
break which, in many areas, occurs between depths of
20 and 50 m. This shallow zone is perhaps the most dis-
tinctive of all reef environments and consists of narrow
annular subzones, which show remarkable concordance
between regions. As shown in Figure 1, reef fronts on
swell- and trade-dominated reefs generally consist of three
common subzones: a shallow spur-and-groove zone,
which extends down to a mid-shelf slope break around
10 m, followed by a relatively low-gradient sand terrace
at 15–25 m, which passes into a chute and buttress zone
developed over the shelf break and down the reef slope.
In all of these subzones, reef development is molded by
wave scour during both fairweather and storms and forms
a characteristic cross-shelf sequence of elongate spur sys-
tems. In shallow water, scour is also responsible for sculp-
turing exposed bedrock surfaces into ridges and furrows,
with ridges becoming enhanced by coral colonization
(Allen, 1982; Blanchon and Jones, 1995). Unfortunately,
due to the rigors of collecting data in such wave-swept
areas, reef development in these subzones is poorly known
and core data exists only for the greater Caribbean with no
reports for any Indo-Pacific reef fronts.
Spur and groove
Spur and groove is a general term for regularly spaced
arrays of buttresses and channels, which extend across
the shelf (Figure 5). Two types of shallow spur and groove
are common: one dominated by coralline algae and the
other by corals. These systems are not mutually exclusive
however and their distribution and form is controlled by
wave energy (Munk and Sargent, 1954; Roberts, 1974;
Sheppard, 1982; Blanchon and Jones, 1997; Storlazzi
et al., 2003). Both coral and algal spurs can be present
along the same high-energy swell-dominated reef front:
a shallow high-energy surf-zone system, dominated by
crustose coralline algae (see Ujelang and Oeno,
Figure 5), and a lower-energy coral-dominated system that
either develops downslope in slightly deeper waters (see
Nonouti and Diego-Garcia, in Figure 5) or in an equivalent
position along the leeward margin (Sheppard, 1981;
Glynn, 1996). Furthermore, the frequency of spurs shows
a correlation with water depth (and hence wave energy):
surf-zone systems have a spacing of one spur every
6–10 m, whereas in deeper-water systems, such as the
chute and buttress zone (see below), spurs are spaced
every 30 m or more, largely as a result of greater spur
widths and wavelengths (Blanchon and Jones, 1997;
Storlazzi et al., 2003). Surf-zone spur systems extend
seaward from the reef crest for 50–100 m dependent upon
the shelf slope, whereas deeper water spur systems can be
more extensive, and submerged terraces can be covered by
coral spurs for several kilometers.
In high-energy swell-affected reefs, shallow spur-and-
groove systems are dominated by coralline algae and
extend seaward from algal rims parallel to wave orthogo-
nals (Munk and Sargent, 1954; Sargent and Austin,
1954). Spurs are commonly 4–8-m wide and have steep,
sometimes overhanging, sides; intervening grooves are
 GEOMORPHIC ZONATION
Geomorphic Zonation, Figure 5 Shallow reef-front spur and
groove. In the cases of Nonouti and Diego-Garcia, algal spurs
pass into coral-covered spurs further downslope, whereas in the
cases of Ujelang and Oeno, algal spurs pass into a bedrock
sculptured into ridges and furrows by storm-wave scour. Images
copyright of Google Earth, DigitalGlobe and GeoEye, 2010.
477
1–3-m wide and 2–8-m deep and are commonly floored
with coral gravel that is moved only during storms. At their
seaward ends, spurs have an amplitude of 5–8 m and ter-
minate in waters 5–10-m deep (Tracey et al., 1948; Ladd
et al., 1950; Cloud, 1952; Newell, 1956; MacNeil, 1972;
Pichon, 1978; Sheppard, 1981; Intes and Caillart, 1994;
Glynn et al., 1996). Spurs narrow seaward and can be
straight-sided with a smooth surface or more irregular
and covered by protuberances (Figures 4 and 5). Near surf
base, the surface of spurs are covered only by patches of
slow-growing crustose corallines or fleshy algal turfs,
whereas below it small corals with reptant growth forms
are common between the corallines (e.g., Intes and
Caillart, 1992; Glynn et al., 1996). In the Marshall Islands,
Munk and Sargent (1954) found that the grooves showed
a bimodal frequency distribution that corresponded to the
northeast trades and, during the summer, to swell from
the southeast trades of the southern hemisphere. Also their
depth and length were proportional to the wave period, so
that the wave up-rush traveled the length of the surge chan-
nel and impeded up-rush of the next wave on its return.
This, together with friction along the sides of the channels,
was sufficient to dissipate wave energy and make spur and
groove an effective natural breakwater.
In lower energy trade-wind reefs protected from swells,
spur systems also form parallel to wave orthogonals but
are covered by corals and composed of coral framework
and extend seaward from gravel-dominated rims (Shinn
et al., 1981, 1982; Blanchon and Jones, 1997). The funda-
mental difference between algal- and coral-spur systems is
simply that coral spurs lack a thick, extensive cover of cor-
allines, and this allows storms to regularly prune their coral
cover and supply the crest and flat with copious amounts
of coral gravel (Blanchon and Jones, 1995, 1997). Coral-
line algae only play a significant role in stabilizing and
binding this gravel (e.g., Macintyre et al., 2001).
Formation of spur and grooves
Although the development of algal spur systems is poorly
known due to the absence of core data, limited excavations
and drilling on the outer reef flat support a growth origin
(Henny et al., 1974; Marshall and Jacobson, 1985; Szabo
et al., 1985; Glynn et al., 1996; Kan et al., 1997b). At
Clipperton Atoll in the eastern Pacific, for example, shal-
low excavations into the ends and sides of algal spurs
revealed a 1-m-thick base of Pocillopora sp. framework
overgrown by crustose corallines, which was overlain in
places by similarly overgrown colonies of massive Porites
lobata (Glynn et al., 1996). This same Pocillopora–
Porites assemblage formed coral spurs directly downslope
in front of the algal spurs, indicating a developmental pro-
gression from intermittent coral growth, storm distur-
bance, and coralline overgrowth through time. This
process is supported by several observations. First, reptant
colonies of Pocillopora are scattered over the deeper parts
of the algal spurs between 3 and 7 m, which indicates that
corals can grow on spurs up to the base of the surf zone
478 GEOMORPHIC ZONATION

(Sheppard, 1982; Intes and Calliart, 1994). Second, coral

growth in front of the algal spurs is largely dead and over-
grown with crustose corallines (Glynn et al., 1996), indi-
cating that the framework in these areas is rapidly
stabilized and might be more resistant to removal or prun-
ing during storms. These two observations imply that the
distal spur ends have the highest potential for lateral and
vertical accretion, which can proceed rapidly until they
intersect with surf base at
3 m. From there, thick crusts
of Porolithon onkoides, previously excluded due to fleshy
algal overgrowth and surface grazing, causes a continued
rapid accretion into the surf zone, leading to the upgrowth
of a new algal ridge (Tracey et al., 1948). In this way, the
process of distal spur upgrowth could not only facilitate
spur development but also might drive reef-flat
progradation over time. It also explains the problematic
occurrence of double reef crests in several areas (Hopley,
1982; Guilcher, 1988).
More subsurface data is available for coral spur
systems – mostly those from the greater Caribbean (Shinn
1963; Shinn et al., 1982a, b). In both Florida and Belize,
for example, blasting and drilling indicate that shallow
spur systems are composed of interlocking branched
colonies of the surf-zone coral Acropora palmata, thickly
encrusted by corallines (Shinn, 1963). Although this
resembles the structure of algal spurs, the basal age of
the coral spurs shows that they are older at their seaward
end and get younger toward the reef crest, suggesting that
they retrograded over their own back-reef deposits as sea
level rose (Shinn et al., 1982a). Although the age structure
of algal spurs is still unknown, ridge progradation indi-
cates that spurs develop in the opposite direction and pro-
grade seaward. If confirmed, this would be a major
difference between these two types of spur system.

Mid-shelf scarp and sand terrace

The area downslope from the shallow spur-and-groove
system in water
8–10-m deep, consists of a gently slop-
ing bedrock terrace, which is either covered with coral
spurs or is a bare rock ground, that has been sculptured
by wave scour into low ridges and shallow furrows (e.g.,
Blanchon and Jones, 1995). This upper rock terrace is
commonly terminated by a mid-shelf slope break before
flattening out again into a lower sand-covered terrace,
which extends out to the shelf edge (Figures 1 and 6). Both
the slope break and sand terrace are frequently traversed
by a system of widely spaced coral spurs between which
thick deposits of skeletal sand and gravel accumulate.
Being below the fairweather wave base, this sediment is
only mobilized during storms, which ensures that the coral
growth survives only on the elevated spurs. Coral gravel
deposited at the end of the spurs is sheltered to some extent Geomorphic Zonation, Figure 6 Reef-front mid-shelf scarp and
sand terrace (or “10-fathom” terrace). Scarp best seen where reef
from wave scour, and subsequent colonization by corals growth is limited and commonly has an intertidal notch (as at
during post-storm conditions ensures that spurs maintain Grand Cayman). Their presence around uplifting islands, such as
their alignment with wave orthogonols and gradually Barbados, indicates an early Holocene sea-level origin. Images
extend out across the shelf. Where the sand terrace is copyright of Google Earth, DigitalGlobe and GeoEye, 2010.
 GEOMORPHIC ZONATION
narrow, less sediment accumulates and more coral devel-
opment is possible. Conversely, where the terrace is wide,
a larger sand reservoir can prohibit spur development
completely.
The sand terrace is a prominent feature seen in imagery
from all oceans (Figure 6) and has been widely reported as
the “ten fathom” terrace (Emery, 1961; Garrison and
McMaster, 1966). Where it is not covered by excessive
reef development, the terrace has been documented in
roughly the same 20  5 m depth range from the following
areas: the Marshall Islands (Tracy et al., 1948; Emery
et al., 1954), the Carolline Islands (Tracey, 1968), the
Tuamotus and Society Islands (Newell, 1956; Chevalier
et al., 1968), Johnson Island in the central Pacific (Keat-
ing, 1985), Clipperton in the east Pacific (Glynn et al.,
1996), the Solomon Islands (Stoddart, 1969), the Cocos-
Keeling Islands and the Maldives in the Indian Ocean
(Williams, 1994; Bianchi et al., 1997), the Bahamas and
Caymans in the Caribbean (Blanchon and Jones, 1995;
Newell, 1951), and Bermuda in the Atlantic (Stanley and
Swift, 1968). In short, the slope break and the sand-
covered 10-fathom terrace are ubiquitous geomorphologi-
cal features and clearly represent the terrace and riser
(cliff) of an abandoned submerged shoreline (Blanchon
and Jones, 1995).
Formation of terrace-scarp reef-front features
Hopley (1982) argued that similar terraces on the Great
Barrier Reef represented common levels of reef growth
during glacial interstadials. A similar incremental expla-
nation was suggested by Paulay and McEdward (1990),
who simulated reef development over the last 125 ka.
Both dismissed marine erosion as insignificant based on
micro-erosion-meter measurements of fairweather coastal
erosion (Spencer, 1985). However, mathematical model-
ing using more realistic erosion rates has demonstrated
that marine planation is capable of producing terraces in
all but the most resistant lithologies in less than 5 ka
(Trenhaile, 1989). Furthermore, the presence of the sand
terrace around islands with a history of uplift during the
Pleistocene such as Barbados in the Caribbean and raised
atolls in the Vanuatu chain, south Pacific, indicates that it
must have developed during the postglacial rise in sea
level (Blanchon and Jones, 1995). In fact, several drowned
early Holocene breakwater reefs have been discovered
associated with the terrace (Lighty et al., 1978; Blanchon
et al., 2002). The crest of an 8-ka old reef was discovered
around Grand Cayman along the inner edge of the terrace
at a depth of
20 m, and its elevation corresponded with
the position of an intertidal notch preserved in the mid-
shelf scarp on the other side of the island (Blanchon
et al., 2002). A correlation with similar elevations of
drowned breakwater reefs, notches, and terraces devel-
oped around other islands in the Caribbean led Blanchon
et al. (2002) to propose that these reefs and their shorelines
were drowned by a meltwater pulse (Mwp-1c) during
the early Holocene that was associated with the final
479
disintegration of the Laurentide Ice Sheet. The timing of
this pulse is also consistent with further discoveries of
drowned reefs in the Gulf of Carpentaria (Harris et al.,
2008) and has been confirmed in several other non-reefal
records of early Holocene sea level (e.g., Hijima and
Cohen, 2010).
Much of the emphasis in these various explanations has
been placed on the terrace-forming process. Yet, it is diffi-
cult to prove from these data alone if the terrace resulted
from marine erosion or from incremental reef accretion.
Also, it is no surprise that the terrace elevation differs
slightly between reefs, given the variation in sediment
and reef deposition during the Holocene; however, neither
of these factors is particularly important to explain. This is
because the notched scarp at the head of the terrace clearly
shows it formed the shoreline behind reef crests that
drowned during the 8-ka meltwater pulse. As a conse-
quence, the presence of this scarp separating terraces in
reef-front zones in other areas indicates that they too were
subject to same shoreline development
8 ka ago. The
surprising thing is that the notched scarp survived
shoreface erosion during the subsequent sea-level rise
and went on to influence reef development (Blanchon
and Jones, 1995). Indeed, the preservation of notched
scarps and abandoned reef crests in different oceans is tes-
tament to the sudden acceleration of sea level during that
8-ka meltwater pulse. It also explains why modern break-
water reef development in these areas did not start until
after that 8-ka event (Blanchon et al., 2002).
Reef slope
In many areas, the reef slope is the most poorly known of
the major geomorphic zones. It represents the outermost
zone of reef construction and can be arbitrarily divided
into two subzones: an upper subzone from the shelf break
to the limit of effective coral-framework development
at
60 m, and a lower subzone down to 120 m or more
that may be dominated by a sciaphyllic framework of
sclerosponges, coralline algae, and microbiallites. The
geomorphology of this lower subzone has been modified
by reef accretion but still bears a strong imprint of sea-
level change during the last glacial–interglacial hemicycle
(e.g., Anderson, 1998).
Shelf-edge chute and buttress (shelf-edge reefs)
Slope breaks associated with the mid-shelf scarp and shelf
edge are often sites of preferential reef development
because sediment bypasses these steeper areas on its way
to depocenters on sand terrace and at the base of the reef
slope, respectively; this is known as the “edge effect”
(Porter, 1972). The shelf-edge site in particular is com-
monly associated with spectacular reef development (as
shown in Figures 1 and 7), which has been described from
various locations as either, sill reefs, shelf-edge reefs, or
mesophotic reef systems (Goreau and Goreau, 1973;
Burke, 1982; Blanchon and Jones, 1997; Lesser et al.,
2009). This reef system develops from the brow of the
480 GEOMORPHIC ZONATION
Geomorphic Zonation, Figure 7 Submerged shelf-edge reef
development. Accretion produces an array of buttresses with
spurs that extend back over the sand terrace (see Grand
Cayman) and down subvertical walls covered by platy corals to
60-m depth. Below that, platy coral cover is reduced and
sclerosponges become more abundant. Images copyright of
Google Earth, DigitalGlobe and GeoEye, 2010.
shelf edge at
25 m down the reef slope to
60 m and can
extend laterally for tens of kilometers (e.g., Burke, 1982;
Blanchon and Jones, 1997). These extensive shelf-edge
reefs have generally been considered to be part of the main
“reef complex” and have not been widely considered to be
separate entities. However, several observations suggest
otherwise: First, shelf-edge reefs commonly form along
margins where breakwater reefs are absent (Blanchon
and Jones, 1997). Second, they have been reported along
the edge of much wider shelves where they are widely sep-
arated from breakwater reef complexes (e.g., Harris and
Davies, 1989). In most areas however, these structures
are juxtaposed along narrow shelves and, although
influenced by similar processes, respond independently,
producing distinctive reef architectures and biotopes
(Figure 7).
One of the few characterizations of a shelf-edge reef
system has come from the Caribbean Island of Grand Cay-
man (Blanchon and Jones, 1997). There, shelf-edge reef
development extends around the full perimeter of the
island (87 km) and forms an array of coral-covered but-
tresses separated by sediment floored chutes and canyons,
which are aligned perpendicular to the shore (Figures 6
and 7). Individual buttresses consist of three elements:
a subvertical front wall colonized by platy coral that
plunges over the shelf edge; a dome-shaped crown cov-
ered by large head corals; and a shoreward-projecting spur
covered by branching corals. On windward margins, these
buttresses have an amplitude of about 10 m and their
crowns rise to a depth of
22 m. In the most-exposed shelf
sections, spurs have a frequency of one every 27 m and
extend across the full width of the sand terrace to merge
with the spur system covering the mid-shelf scarp. Along
the less-exposed sections however, the spur length
decreases and their frequency is reduced to one spur every
75 m. These trends also exist along more protected mar-
gins, and the spur frequency decreases in response to shel-
ter from the ambient wave field. In addition, crowns rise
into waters of 16 m and buttress amplitudes are reduced
to
7 m (Blanchon and Jones, 1997). Along the leeward
margin, buttresses merge to form an almost continuous sill
reef dominated by branching corals and little spur devel-
opment. This sill rises into waters as shallow as 12 m.
Formation of shelf-edge chutes and buttresses
Roberts (1974) found a correlation between shelf-edge
buttress-spur orientation and vectorial properties of the
wave field affecting Grand Cayman. From this, he
conjectured that these deep-water spur systems had
inherited the form of shallow spur-and-groove structures
that were drowned as the sea level rose during the Holo-
cene (Roberts et al., 1975). This was contested by
Blanchon and Jones (1997), who showed that the lower
frequency and more widespread distribution of shelf-edge
spurs was inconsistent with this inheritance hypothesis.
Instead, they suggested that the covariance between but-
tress area, amplitude, spur frequency, and local margin
 GEOMORPHIC ZONATION
orientation was more consistent with the varying destruc-
tive intensity of hurricane wave scour and approach angle
(Blanchon and Jones, 1997; Blanchon, 1997). In other
words, these shelf-edge reefs were produced by significant
reef accretion in deeper waters. Although no drilling has
yet proved this around Grand Cayman, cores from an
extensive shelf-edge reef off Carrie Bow Cay, on the
Belize Barrier reef, recovered an 11.6-m-thick section of
domal and branching framework that developed over the
last 5 ka (Macintyre et al., 1981). Similarly, 9 m of domal
framework that had developed in the last 7 ka was encoun-
tered on a shelf-edge reef off the south coast of Barbados
(Fairbanks, 1989). Off St. Croix, horizontal drilling into
the front wall of a shelf-edge reef buttress by Hubbard
et al. (1985) recovered 6 m of lateral accretion consisting
of alternating detritus and domal framework. These thick-
nesses of framework are clearly sufficient to account for
the amplitude of the shelf-edge reef buttresses and spurs
in these areas and therefore support the contention that
the shelf-edge reefs are independent structures controlled
by ambient environmental processes (Macintyre et al.,
1981; Blanchon and Jones, 1997).
Lower reef slope
The upper section of the reef slope to
60-m depth is
dominated by a cover of platy corals associated with
shelf-edge reef development; however, in the lower sec-
tion of the slope below 60 m, this coral coverage declines
rapidly, and by 100 m, generally forms a cover of less than
1%. In areas investigated by submersibles, this lower
slope usually consists of a near-vertical wall with an irreg-
ular surface consisting of discontinuous, sediment-draped
ledges and blind caves dissected by vertical chutes and
gullies (Ginsburg et al., 1991; Grammer and Ginsburg,
1992). This irregular substrate is colonized by a
sciaphyllic assemblage of sclerosponges, crustose cor-
alline algae, microbialites, and platy corals, and their
zonation and distribution are controlled by the light
gradient, falling sediment from the shelf above, and
the margin orientation (Goreau and Land, 1974; Land
and Moore, 1977; Sheppard, 1982; Colin et al., 1986;
Liddell and Ohlhorst, 1988; Ginsburg et al., 1991;
Lesser et al., 2009). By far, the most thoroughly inves-
tigated lower reef slope is that along the north coast of
Jamaica (Hartman and Goreau, 1970; Goreau and Land,
1974; Lang et al., 1975; Land and Moore, 1977; Liddel
and Ohlhorst, 1988). In this area, the depth zone
between 60 and 100 m consists of a variable cover of
sclerosponges, with reports ranging from 7% to 25%
and as much as 50% on the walls and ceilings of caves.
In fact, in a cluster of caves between 80 and 90 m,
Lang et al. (1975) reported sclerosponges as large as
1 m in diameter. Furthermore, a half-meter-deep blast
site at 105 m exposed the sclerosponge framework with
interstitial skeletal sediment cemented by high-
magnesian calcite (Land and Moore, 1977). Similar
abundances of sclerosponges have also been reported
481
from the Bahamas, where Ginsburg et al. (1991) esti-
mated a cover of 5–25% in the same depth range.
Formation of the reef slope
There is generally a very little subsurface data available
concerning the development of reef-slope environments.
The tendency of modern reefal margins to form steep pre-
cipitous slopes must relate to both accretion and erosion
during rapid sea-level changes during glacial–interglacial
cycles. For example, Blanchon and Jones (1997)
suggested that the pattern of vertical and lateral accretion
in shelf-edge reefs may explain the steep precipitous upper
slope development that is common in many reefal mar-
gins. However, the vertical walls of the deeper lower slope
must be related to the long phases of sea-level lowstands at
125 m or more that are associated with glacial stages
(Land and Moore, 1977). Although these cliffs are pres-
ently covered by coral and sclerosponge frameworks and
are likely undergoing some small amount of lateral accre-
tion (e.g., Ginsburg et al., 1991), cliff recession rates dur-
ing the extended duration of glacial lowstands must have
been substantial. Therefore, the basic geomorphology of
these vertical substrates is likely erosional – a claim that
is supported by widespread reports of giant blocks of wall
rock on forereef depositional slopes.
Summary and conclusions
Rough-water coral reefs that develop in trade-wind-,
swell-, and storm-dominated seas consist of five standard
geomorphic zones: a back-reef lagoon, a reef flat, a reef
front, a reef slope, and a fore-reef talus apron. These zones
are separated by simple slope breaks: back-reef lagoons
terminate at the sandslope break, reef flats terminate at
the reef-crest break, reef fronts terminate at the shelf-edge
break, and reef slopes terminate at the fore-reef talus
break.
In lagoons, reef development can be extensive and reefs
develop as dispersed knolls or patches that commonly
merge to form ridges; these in turn can connect with other
ridges to form complex reticulate networks that impede
water circulation and self-limit their own development.
In many lagoons, water circulation is a function of wind
stress and exchange with the open ocean, with the latter
process being controlled by a wave-pumping mechanism
over open reef flats. Atolls with large sections of open reef
flats therefore have better lagoonal reef development than
smaller atolls, where the exchange system is more vulner-
able to blockage by island formation in space and time.
Seismic data show that the lagoonal reef form and devel-
opment is not, as often claimed, inherited from a karstic
substrate but entirely growth induced either from simple
vertical and lateral accretion during the Holocene or from
compound growth over previous interglacials.
Reef flats by contrast are remarkably flat intertidal plat-
forms that can be several kilometers wide. They consist of
an algal rim, an outer coralgal flat, an inner sand flat, and
a variable cover of low-lying detrital islands. The algal
482 GEOMORPHIC ZONATION
rim, generated by the surf-adapted coralline Porolithon
onkoides, forms the highest part of the reef flat and con-
sists of either an elevated ridge fronting islands along the
most exposed reef sections or a lower mound along more
open flats. Behind the rim is an intertidal coralgal plat-
form, veneered by a rapidly decreasing cover of corals
and an increasing proportion of coral gravel stabilized by
coralline algae. In its outer part, this platform can be punc-
tuated by one or multiple remnants of former algal ridges
whose age increases inshore or by large blocks of reef
framework thrown up from the reef front during storms.
In addition, the coralgal flat is commonly dissected by
surge channels, which extend back into the reef flat in-line
with reef-front grooves. Some flats are crossed by corals
organized into narrow, parallel ridges that are separated
by gravel-filled furrows, which also align with reef-front
grooves. These various types of coralgal flats either grade
into an inner flat dominated by shifting sands or pass
directly into cobble rampart-fronted islands composed of
a mix of storm-lain sand and gravels that can reach
4 m or more above the surrounding flat. Although stabi-
lized by marine and meteoric cements, these gravel islands
yield clasts with a large spread of ages, suggesting that
they are commonly reworked by storms.
In explanations of reef-flat development, it has been
argued that elevated islands, remnant algal ridges, and
the extremely flat, barren nature of reef flats are a com-
bined result of a late Holocene regional fall in sea level,
which led to erosional planation of the surface to its pre-
sent level. Several authors, however, have noted that these
same features are consistent with processes that are pres-
ently active with a stable sea level. In fact, many of the
key features of reef flats, such as their monotonous flat-
ness, aligned features, and remnant ridges, can be
accounted for by the simple seaward progradation of the
algal rim out over the reef-front spur-and-groove zone
through time.
Spur-and-groove development in shallow, rough-water
reefs consists of two types: one with spurs covered by
crustose coralline algae and the other with spurs covered
by corals. On swell-dominated reefs, algal spurs extend
seaward from the algal rim parallel to wave orthogonals
and form high-frequency, steep-sided spur arrays, whose
length is proportional to wave energy and the gradient of
the seaward shelf. Coral spurs have a similar form and fre-
quency but develop in lower wave-energy regimes either
downslope from algal spurs or in an equivalent position
along the leeward margin. All subsurface evidence col-
lected so far point to a growth origin for both spur types,
with algal spurs nucleating on downslope coral growth
causing a seaward progradation through time, and coral
spurs developing downslope from the crest and
retrograding upslope through time.
The platform downslope of the shallow spur-and-
groove terminates at about 10 m in a mid-shelf slope
break that commonly takes the form of a small notched
scarp (where it has not been covered by coral spurs or
sedimentation). This scarp forms the riser to a distinct
sand-covered terrace, known as the 10-fathom terrace,
which slopes from 20 to 30 m out to the shelf edge. This
10-fathom terrace and its scarp has been reported from
all oceans and even around uplifting islands such as
Barbados, and has become a depocenter for sediment
accumulation and reef development during the Holocene.
Discoveries of several early-Holocene breakwater reefs on
the terrace prove that the notched scarp and terrace couplet
represents a cliffed shoreline that formed around the same
time and was drowned by a meter-scale sea-level jump

8 ka ago.
The shelf break at the terrace edge is commonly a site of
extensive submerged reef development, which has pro-
duced an array of sand chutes and steep-sided buttresses
that form parallel to wave orthogonals. The buttresses,
which can rise to 15 m and have amplitudes of 10 m or
more, extend back over the terrace as a series of tapering
spurs. To seaward, the subvertical face of the buttresses
extends down the reef slope to
60 m and has a dense
cover of platy corals. Covariance between buttress area,
amplitude, spur frequency, and local margin orientation
indicates that the architecture of these shelf-edge reefs is
controlled by hurricane wave scour and approach angle.
Other suggestions that they developed over drowned
breakwater reefs are not supported by drilling, which
shows that shelf-edge reefs have produced greater than
12 m of vertical growth and 6 m of lateral accretion in
the last 5 ka.
The reef slope beneath shelf edge reefs is commonly
vertical below 60 m and is covered by an assemblage of
sciaphyllic sclerosponges, crustose corallines, and
microbialites. Although this assemblage is actively pro-
ducing framework, the morphology of the deep reef slope
has been significantly influenced by cliff collapse and
mass wasting during the long intervals of glacial
lowstands of sea level.
The remarkable similarity of reef geomorphology and
architecture between and within oceans is a clear testa-
ment to their capacity for self-organization (Hatcher,
1997; Dizon and Yap, 2006). In other words, many of
the features of these large-scale structures have emerged
spontaneously from ecological processes operating at
smaller scales. For example, the negative feedback
between the simple vertical and lateral reef accretion and
water circulation has produced extensive networks of cel-
lular reefs in many lagoonal systems. Likewise, the simple
interaction between wave scour and the colonization and
growth of corals and crustose coralline algae has produced
well-ordered spur systems with similar spacings at both
local (windward–leeward), regional (trade or swell) and
inter-oceanic scales. Finally, the interaction between her-
bivory and coralline accretion along the seaward edge of
algal spurs may be responsible for intermittent seaward
advances of reef flats through time.
Perversely, many of those responsible for interpreting
large-scale reef development still cling to the paradigm
that modern reefs are somewhat of a geological oddity
and that their form and architecture are merely
 GEOMORPHIC ZONATION
a reflection of the underlying antecedent substrate –
a position that would surely have Darwin turning in his
grave. Although examples do exist of this substrate con-
trol, scaling-up of the concept to entire reef systems denies
abundant evidence that reefs have built impressively thick,
well-structured edifices during the last 14 ka, which have
maintained their breakwater position as post-glacial sea
level rose 120 m or more from its glacial low-stand. An
understanding of the interactions and feedbacks that con-
trol reef organization during this development is key to
assessing resilience and resistance of modern reef systems
in the face of human-induced global environmental
change.
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486 GLACIAL CONTROL HYPOTHESIS

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Marine Ecology and Paleoecology 1. Ecology, New York: Geo- land was subject to general movement and Eduard Suess’s
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Marine Geology, 170, 331–346.
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changes in the Indo-Pacific. Journal of Asian Earth Sciences, Semper, Wharton, Guppy, and Gardiner).
25, 29–43. Daly visited Hawaii in 1909 and observed the narrow-
Wyrwoll, K. H., Zhu, Z. R., Collins, L. B., and Hatcher, B. G., 2006. ness of the encircling reefs which he inferred as an indica-
Origin of blue hole structures in coral reefs:Houtman Abrolhos, tor of young age; he saw evidence for recent glaciation on
Western Australia. Journal of Coastal Research, 22, 202–208. the volcanic slopes of Mauna Kea, indicating lower glacial
temperatures; and he noted the closeness of contemporary
Cross-references sea surface temperatures in the northern winter at Hawaii
Algal Rims to the threshold temperature for reef growth. These obser-
Blowholes vations were built into the glacial control hypothesis, first
Boulder Zone/Ramparts published in 1910 and reaching their full development in
Double and Triple Reef Fronts the Silliman lectures delivered at Yale University and
El Niño, La Niña, and ENSO published in 1934 in a book entitled “The Changing World
Fore Reef/Reef Front of the Ice Age.” W. M. Davis saw Daly’s hypothesis as
Hydrodynamics of Coral Reef Systems
Lagoons diametrically opposed to Darwinian subsidence theory
Megablocks and submitted it to sustained intellectual attack (e.g.,
Moats Davis, 1918, 1919, 1923, 1928) but, as Stoddart (1973)
Patch Reefs: Lidar Morphometric Analysis has pointed out, Darwin was concerned with understand-
Platforms (Cemented) ing large-scale reef structure, whereas Daly was more
Reef Flats concerned with explaining reef surface morphology.
Reticulated Reefs
Spurs and Grooves Daly’s key fact, backed up by an analysis of bathymet-
Submerged Reefs ric charts of reef areas, was the global accordance and pla-
Wave Set-Up nar nature of atoll lagoon floors and continental shelves in
the reef seas within a limited depth range of 55–90 m. He
argued that during the glacial periods, sea level fell, sea
surface temperatures declined, and water turbidity
GLACIAL CONTROL HYPOTHESIS increased. These processes, perhaps exacerbated by
increased storminess and wave attack, resulted in coral
Tom Spencer mortality, the removal of dead coral framework, and the
University of Cambridge, Cambridge, UK erosion of volcanic shores no longer protected by healthy
reef structures. The outcome was the production of
Synonyms beveled platforms on volcanic islands and older limestone
basements. As sea level rose and sea surface temperatures
Glacial control theory increased with the onset of interglacial conditions, then
coral growth resumed, preferentially on the platform mar-
Definition gins (where environmental conditions were most favor-
Hypothesis developed by R. A. Daly to argue for youthful, able for coral growth) to result in typical atoll and barrier
postglacial development of atolls and barrier reefs on plat- reef morphologies (Figure 1).
forms beveled during low sea surface temperatures and
sea levels of glacial periods.
Tests of the hypothesis
‘Glacial control’ of coral reef development The hypothesis encompasses several elements all of which
Development of the hypothesis are amenable to test as a result of late twentieth/early
The glacial control hypothesis, developed and defended twenty-first century developments in Quaternary science.
through a series of lectures, scientific papers, and book First, what was the degree of sea-level fall during the gla-
chapters between 1910 and 1948 by R. A. Daly, was an cial periods? Second, what was the degree of sea temper-
exploration of the effects of Pleistocene climate and sea- ature cooling during the glacial periods? And third, how
level change on coral reef evolution and morphology. easy is it to bevel a reef structure and produce a smooth
The hypothesis should be seen in the context of Nathaniel platform?
 GLACIAL CONTROL HYPOTHESIS 487

Glacial Control Hypothesis, Figure 1 Sections illustrating aspects of the glacial control hypothesis of coral reefs. Diagonal shading
indicates stable basement. 1: eustatic rise of sea level during the last interglacial stage giving new coral growth (in black), largely
at the platform margin; 2: sea-level fall and water temperature decrease during the onset of the last glacial period, during which
corals die and reefs are eroded; 3: at the Last Glacial Maximum reefs are destroyed and the bank is eroded to below the lower sea
level; 4: the postglacial rise in sea level, together with a temperature increase, allows renewed coral growth (in black), particularly at
the outer reef margins; 5: the present features of an atoll encircling a flat-floored lagoon. Source: Daly (1934).

Daly (1910) originally postulated a sea-level fall of
46 m at the glacial maximum which he then revised to
a figure of 60–70 m, having dismissed the estimates of
sea-level fall of 150 m as “excessive” (Daly, 1934,
p. 165). It is now clear from drilling submerged Acropora
palmata reefs offshore from the south coast of Barbados
that sea level at the last glacial maximum (18 ka BP)
was 121  5 m (Fairbanks, 1989).
For glacial sea surface temperature, Daly suggested
a typical cooling of 5 C, with a maximum potential fall of
10 C. Quantitative reconstructions of last glacial maximum
sea surface temperature, based on the microfossil and geo-
chemical analyses of deep-sea cores, were first developed
by the Climate Long-Range Investigation, Mapping, and
Prediction (CLIMAP) project in the 1970s and 1980s
(CLIMAP project members, 1976) and have recently been
updated by the Multiproxy Approach for the Reconstruc-
tion of the Glacial Ocean Surface (MARGO) project.
MARGO project members (2009) report an average mean
annual cooling at the last glacial maximum of 1.7  1 C
for the tropical ocean between 15 N and 15 S. This can
be disaggregated regionally into a cooling of 2.9  1.3 C
in the Atlantic Ocean, 1.4  0.7 C in the Indian Ocean,
and 1.2  1.1 C in the Pacific Ocean. There were, however,
considerable temperature variations in the latter region:
cooling of 1–3 C in the western Pacific Ocean warm pool,
cooling of 5–6 C in the equatorial eastern boundary current
along the Chilean coast but warming of 1–2 C in the sub-
tropical gyres in both hemispheres. Daly claimed that “over
wide stretches of the tropical seas the reef corals were exter-
minated or greatly weakened in their reef-building power”
(Daly, 1934, p. 159). However, if the known falls in temper-
ature are subtracted from the present patterns of ocean tem-
perature isotherms for the coldest month, and the isotherm
of 20 C is taken as the limit to effective reef growth, then
the change in the area available for reef growth during the
glacial periods can be illustrated (Slaymaker et al., 2009;
Figure 2). This analysis shows that while Daly’s
488 GLACIAL CONTROL HYPOTHESIS
Glacial Control Hypothesis, Figure 2 Changing tropical ocean temperatures, Last Glacial Maximum to present, and implications for
coral reef growth (from Slaymaker et al., 2009).
speculations on the magnitude of temperature fall were of
the correct order of magnitude, his interpretation of their
likely impact was incorrect. It was only on the very margins
of the reef seas and in the area of the eastern boundary cur-
rents that sea temperature cooling was sufficient to cause
a likely cessation of reef growth. Interestingly, these mar-
ginal belts were the one area where W. M. Davis was pre-
pared to accept the occurrence of coral platforms of low-
level abrasion, these being the only locations in the reef seas
where cliffed volcanic island slopes can be seen on the mar-
gins of barrier reef lagoons (Davis, 1915, 1928).
Daly estimated that 50–200,000 years was available for
platform abrasion and leveling in the periods of low sea
level during the glaciations. This inference is broadly cor-
rect; environmental reconstructions from ice-cores and
marine sediments in deep-sea cores show that lower than
present sea levels have typically characterized 88,000 years
of the characteristic 100,000-year long glacial–interglacial
cycles of the last 500,000 years (e.g., Imbrie et al., 1993;
Petit et al., 1999). While accepting that young volcanic
islands were composed of erosionally resistant lava flows,
Daly believed that most volcanic and coral islands were
“composed of generally weak material” (Daly, 1934, p.
177) and that erosion rates of coral limestones would be
comparable to the 0.3–1.0 m a1 retreat rates of the chalk
cliffs in coastal southern England and northern France and
that pyroclastic deposits and shelf sediments might be
removed at rates comparable to the retreat rates of ca.
2 m a1 of the glacial cliffs on the UK Yorkshire coast of
the North Sea. Thus, there appeared to be no difficulty in
beveling even large volcanic islands and extensive reef
deposits in the time available, particularly if material had
been weakened by previous phases of weathering and ero-
sion under earlier glaciations. Recent quantitative measure-
ments using the microerosion meter technique on shore
platforms, over a range of materials and in different process
environments around the world suggest, however, that ero-
sion rates are several orders of magnitude lower than the
rates used by Daly, with a mean rate of surface lowering
of 1.5 mm a1 (Stephenson and Finlayson, 2009). Within
this dataset, the erosional behavior of presently raised reef
limestones provides an apt modern day analog for the reac-
tion of reefs exposed by glacial falls in sea level. Such sur-
faces have been comprehensively investigated in two
locations: at meso- to macrotidal Aldabra Atoll, western
Indian Ocean by S. T. Trudgill (Trudgill, 1976a, b, 1983;
Viles and Trudgill 1984) and at microtidal Grand Cayman
Island, West Indies by T. Spencer (1985a–c). At Aldabra,
mean intertidal rates of surface lowering were 1.79 mm a1,
varying between 1 mm a1 on low energy coasts to
4–7 mm a1 on exposed coasts. In the supratidal zone, the
mean rate of subaerial erosion was 0.26 mm a1. On Grand
Cayman, the mean intertidal erosion rate was 1.23 mm a1,
varying between 0.45 and 2.43 mm a1 on reef-protected
and exposed coasts, respectively (with bioprotection
reducing rates to 0.22 mm a1 on the most exposed surf
platforms). Subaerial downwearing was between 0.45
and 0.97 mm a1. If these rates were characteristic of
 GLACIAL CONTROL HYPOTHESIS
the Pleistocene, then intertidal erosion would require 0.5–
1.0 million years to plane a reef 1 km in width, well in
excess of the time available in each glacial–interglacial
cycle. Furthermore, shoreline retreat is an episodic rather
than a continuous process in consolidated reef limestones,
proceeding by alternating phases of intertidal notch cutting
and then collapse. Next, intertidal backwearing is only part
of the process of limestone removal, the other main process
being solutional lowering of the exposed surface. On inland
limestone substrates on Grand Cayman, mean subaerial ero-
sion has been measured at 0.37 mm a1, with subsoil and
submangrove rates of 0.16 mm a1 and 0.21 mm a1,
respectively. Thus if reefs were emergent by 125 m at the
glacial maximum, then it would take over 300,000 years
to remove such a column of limestone at the highest of these
mean rates of lowering. Finally, the heterogeneity of lithifi-
cation in reef limestones is reflected in the fact that erosion
is extremely localized and gives rise to a pitted and cavern-
ous microtopography rather than undifferentiated planation;
thus, erosion is often accompanied by the preservation of
original depositional reefal surfaces.
Alternative explanations for the morphology of
modern reef surfaces
It is clear from modern process studies that once reefs are
formed they are difficult to remove by marine and subaerial
erosion processes. In addition, the refining of the magni-
tudes and duration of ocean temperature change and sea-
level fall following the late twentieth-century advances in
environmental reconstruction do not provide the contexts
within which wholesale reef removal can be realistically
achieved. Given that present sea levels have been at or near
to their present levels for at most the last 6,000 years and in
some cases for only the last 1,000 years (only 0.6–0.1% of
the last glacial–interglacial cycle), these conclusions
suggest that the topography of modern reefs is often
inherited from underlying, older reef surfaces (see also
Chapter Antecedent Platforms). On the basis of bathymetric
charts, Daly argued for the smoothness of these reef base-
ments. Davis (1928) argued that such smoothness was the
product of effective lagoon infilling during the upgrowth
of marginal reefs. However, more recent detailed lagoon
and shelf floor mapping, both of bottom morphology and,
using seismic techniques, of bedrock surfaces below reefal
sediments, has shown many basements to be characterized
by highly irregular bedrock surfaces (Bloom, 1974), often
organized into characteristic limestone erosional, or karstic,
forms. Topographic features range in scale from small
solutional features (e.g., solutional pipes in the Florida
Keys; Dodd and Siemers, 1971) to marginal raised rims
around saucer-shaped reef platforms (MacNeil, 1954; Har-
vey, 1977), “knoll and basin” topography in atoll lagoons
(e.g., Chuuk lagoon, Federated States of Micronesia; Shep-
ard, 1970), and “blue holes” in back-reef environments
(e.g., Lighthouse Reef blue hole, Belize Barrier Reef;
Stoddart, 1962), to large scale regional karst landscapes.
The most comprehensive exposition of karstic control on
489
reef landscapes was provided by E. G. Purdy who, on the
basis of seismic profiling and shallow drilling, argued that
the regional morphology of the Belize Barrier Reef reflects
a drowned doline karst in the drier northern barrier reef and
on the central and southern barrier reef and a drowned tower
karst in the lagoon and a flooded basement cockpit karst on
the barrier reef margin itself. These latter two morphologies
have resulted from the submergence of a karst marginal
plain, supplied by solutionally aggressive drainage waters
from the noncarbonate Maya Mountains. The underlying
karstic control is clear on the Belize Barrier Reef because
of the interaction of the depth below present sea level of
the basement and the sea-level history at this location;
Holocene reef development has been dominated by vertical,
“catch-up” behavior and thus postglacial reef growth has
exacerbated the underlying pre-Holocene topography
(Purdy, 1974).
Elsewhere, the relations between basement depth, sea-
level history, and postglacial reef growth modes (give-up
vs. catch-up vs. keep-up) have been more complicated
and given rise to a wider range of contemporary reef
forms; in cases where the basement has been close to pre-
sent sea level, reef growth may have completely filled the
accommodation space and thus obscured any antecedent
topography (Hopley, 1982). In some locations, the under-
lying antecedent topography is not a karst erosion surface
but a depositional one and thus reef topography reflects
colonization of stabilized carbonate shoal or reef top storm
deposits (e.g., Garrett and Scoffin, 1977) or reef growth on
underlying river levee (Choi and Ginsburg, 1982) or del-
taic deposits (e.g., Maxwell, 1970).
Conclusions
R. A. Daly’s “Glacial Control Hypothesis,” developed in
the first quarter of the twentieth century, importantly
engaged coral reef science with the question as to how
reefs had reacted to Pleistocene changes of ocean temper-
ature and sea level. Developments in Quaternary Science
and in process geomorphology have, however, not
supported Daly’s assumptions about the rapidity at which
volcanic materials and coral reef limestones can be
removed by erosion and thus the ease with which reefal
surfaces can be planed to smooth lagoon floors and reefal
shelves during the glacial phases of the Pleistocene.
Rather modern reef topographies often reflect the complex
underlying topography of much older, pre-Holocene sur-
faces and the erosional and depositional process histories
to which they have been subjected. Antecedent topo-
graphic control is, however, variable and relates to the
degree to which subsequent reef growth has either accen-
tuated or obliterated the basement surfaces from which
modern reefs have developed.
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Cross-references
Antecedent Platforms
Belize Barrier and Atoll Reefs
Blue Hole
Climate Change and Coral Reefs
Daly, Reginald Aldworth (1871–1957)
Darwin, Charles (1809–1882)
Davis, William Morris (1850–1934)
Last Glacial Lowstand and Shelf Exposure
 GLACIO-HYDRO ISOSTASY
MacNeil, F. Stearns
Postglacial Trangression
Sea Level Change and Its Effect on Reef Growth
Solution Unconformities
Subsidence Hypothesis of Reef Development
Thickness of Holocene Reefs
GLACIO-HYDRO ISOSTASY
Kurt Lambeck
The Australian National University, Canberra ACT,
Australia
Synonyms
Eustatic sea level; Glacial rebound; Postglacial sea level
Definition
Glacio-hydro isostasy refers to the earth’s response to
changes in surface ice and water loading during glacial
cycles. During the growth of an ice sheet, the crust is
loaded in the areas of glaciation and the crust beneath
and near the ice sheet subsides. During the decay phase
of an ice sheet, the crust is unloaded and rebounds. This
is the glacio-isostasy component. But as water is extracted
from, or added into, the oceans during ice growth and
decay, the oceanic crust also responds to the changing
water load. This response is called the hydro isostasy.
The two combined components, not uncoupled, are
referred to as glacio-hydro isostasy.
Introduction
The glacio-hydro isostatic effect is manifested in geologi-
cal, geophysical, and geodetic observations. It is seen in
the geological record as a complex spatial and temporal
pattern of raised shorelines (Figure 1) in areas of former
glaciation (primarily the glacio-isostatic signal) as well
as along coastlines far from the former ice margins (pri-
marily the hydro-isostatic signal). It is also seen in geo-
detic observations: as displacements of the crust, both
vertical and horizontal; as sea-level change in tide gauge
and satellite altimeter data; as changes in gravity measured
by surface instrumentation and by satellite sensors and
orbital perturbations; and as anomalies in the earth’s rota-
tion. It has implications for understanding sea-level
change, paleogeographic reconstructions of coastal envi-
ronments, archaeology, the stress state of the crust, and
the computation of precise satellite orbits.
Models for glacio-hydro isostasy
A zero-order description for isostatic response to loading
is a model of local response of a crust–lithosphere that is
unable to support shear stresses and that overlies a fluid
mantle of zero viscosity. An ice load of H m thick and of
density ri would displace the crust beneath it by
491
Hri/rmantle or
25–30% of the thickness of the ice or
water load. This model ignores the strength of the essen-
tially elastic crust–lithosphere as well as the viscosity of
the mantle, but it provides a useful order of magnitude
estimate of the maximum displacement that may occur.
Thus, beneath large ice sheets, the maximum crustal dis-
placement may approach a kilometer and the seafloor
within ocean basins may be displaced by 30–40 m for
a globally averaged sea-level change of
120 m.
An improved model is one of regional isostasy in which
the load is supported by an elastic crust–lithosphere layer
that overlies a zero viscosity mantle. For loads of horizon-
tal dimensions much greater than the lithospheric thick-
ness, the displacement at the center is nearly equal to
that predicted by the local model but at the edge of the load
it is only about one half of this amount because the load is
partly supported by the flexural strength of the litho-
sphere. Thus, along continental margins, the hydro-
isostatic signal may reach 15–20 m for the above global
sea-level change of
120 m. But this is still only an order
of magnitude estimate and to fully describe the global pat-
tern of sea-level change, more physically realistic models,
that involve the mantle viscosity, are required.
Realistic models of the glacio-hydro-isostatic phenom-
enon include the following features:
 Physically realistic representation of the planet’s
response function, or rheology, to surface loading on
timescales of thousands of years. The observational evi-
dence indicates that the earth is still responding today to
the unloading of the last great ice sheets such that the
model requires elastic as well as viscous elements in
the response function. For both mathematical expedi-
ency and because it provides a satisfactory explanation
of the observational evidence, a linear viscoelastic
response is usually assumed. Both depth and lateral var-
iability in the rheology needs to be considered.
 A realistic description of the evolution of the ice sheets
during the full glacial cycle. A detailed description of
the individual ice sheets is required (on a 0.5 grid
and preferably smaller) for the evaluation of the
glacio-isostatic component, but for the hydro-isostatic
component it is the total rate at which water is added
or removed from the oceans that is the significant
requirement.
 A detailed description of the ocean basin to ensure that
meltwater from the ice sheets is properly distributed.
This distribution has to satisfy the requirement that the
ocean surface is an equipotential surface at all times
and that the ice–water mass is conserved. Through time,
the ocean basin deforms, the shorelines migrate and the
ice sheets displace water on the shelves. The ocean
basin deforms in response to both the ice and water
loads and this introduces a coupling between the glacio-
and hydro-components of the isostatic response.
 The ability to compute the deformation and the mass
distribution of the earth–ocean–ice system through time
and to relate this to the observational evidence for
492 GLACIO-HYDRO ISOSTASY

Glacio-Hydro Isostasy, Figure 1 Representative sea-level curves for different localities around the world. The time and sea-level
scales are not the same for all examples. (After Lambeck and Chappell [2001]).

sea-level change and shoreline migration, crustal defor- cycle, the earth’s rotation, centrifugal force, and the
mation and gravity change, and the global inertia tensor. gravitational equipotential surfaces are also modified.
 The feedback that occurs through the earth’s rotation.  High spatial resolution solutions are required. Sea-
As the inertia tensor is modified during the glacial level changes above the threshold of observational
 GLACIO-HYDRO ISOSTASY
accuracies can occur, for example, over distances of
10 km or less across former ice margins or continental
margins and the numerical solutions must be able to
represent this.
The formulation of the glacio-hydro-isostatic rebound is
well established but the definition of the rheological
parameters or aspects of the ice models themselves are
often less well known and are mostly inferred from the
analysis of the rebound evidence. Thus, for descriptive
purposes, the formulation serves as an interpolation device
between fragmentary observational evidence that also
allows for a degree of extrapolation beyond the limits of
the time–space dimensions defined by the observational
data.
The sea-level response
In the context of coral reefs, the most important conse-
quence of the glacio-hydro-isostatic response is sea-level
change. Schematically, the equation describing sea-level
change, Δzrsl (j,t), can be written as
Dzrsl ðj;t Þ ¼ Dzesl ðt Þ þ DzI ðj;tÞ þ DzT ðj;tÞ þ DzO ðj;t Þ
with
DzI ðj;tÞ ¼ DzIg ðj;t Þ þ DIh ðj;tÞ
Δzrsl (j, t) represents the change at location j of the sea
surface relative to land at time t compared to its present
position at time tP. The first term on the right-hand side
represents the ice-volume equivalent sea-level contribu-
tion (ESL) and is a measure of the globally averaged
sea-level change due to a change in ice volume. It is
defined as
Z
r 1 dVi dt
Dzesl ðtÞ ¼  i
ro Ao ðtÞ dt
t B(tP). The original shoreline will now occur at A0 , at
Vi is the ice volume at time t and includes the ice mass on
continents and grounded on continental shelves. Ao(t) is
the ocean surface area and is defined by the shoreline
and the grounding lines of any ice on the shelves, both at
time t. ri, rο are the average densities of ice and ocean
water. In the absence of any other factors that lead to
changes in ocean volume, the ice-volume equivalent sea
level is equal to eustatic sea level.
The second term, DzI(j,t), is the combined isostatic
contribution. It is schematically divided here into two
contributions: the glacio-isostatic part DzI–g(j,t), and
the hydro-isostatic part DzI–h(j,t). Both terms include
contributions from the deformation of the crust under
the time-dependent surface load and from the change in
the gravitational potential of the earth–ice–water system
(sometimes referred to as geoid changes). Coupling
between the two isostatic terms arises from the deforma-
tion of the ocean basins and from the change in gravita-
tional attraction by the ice. This is ignored in this
493
schematic description but not in the actual models
(Lambeck et al., 2003; Mitrovica and Milne, 2003).
The third term, DzΤ(j,t), allows for any tectonic contri-
butions to sea level. Many reefs occur in areas of tectonic
uplift or subsidence and the separation of these effects
from the isostatic signals remains a major challenge.
A frequently used criterion is to examine whether there
is evidence for a fossil Last Interglacial reef at the same
location. If this occurs within a few meters of present sea
level, then the site can be considered to have long-term
tectonic stability to better than 0.1 mm/year. Alternatively,
if the sea-level behavior can be calibrated against sites of
known tectonic stability, the comparison of model predic-
tions and observations establishes rates of tectonic uplift
or subsidence.
The fourth term, DzΟ(j,t), includes any oceanographic
or meteorological factors that cause sea level to depart
from a long-term equipotential surface. These are gener-
ally treated as perturbations in high-resolution analyses
of recent and Holocene sea-level change.
Despite most modern coral reefs occurring far from
the ice margins of the last glaciation – far-field sites – both
components of the isostatic response occur. The glacio-
isostatic response here is characterized by a broad wave-
length response, reflecting global-scale mantle flow
driven by the growth and decay of the large ice sheets,
and the sea-level response is not sensitive to the details
of the ice sheets. More important here is the hydro-
isostatic component and this is strongly dependent on
the changes in the distribution of water around the site.
Figure 2 illustrates sea-level change at a continental
margin site. At a time t, the land surface is at a – a0 and
the shoreline is at A(t). Between t and the present, tP, land
uplift has occurred by an amount ur and the new land sur-
face is at b  b0 . The ocean volume has also increased so
as to raise the ocean level relative to the center of mass
of the earth from zA to zB and the present shoreline is at
Δz(t) below the present sea level. The relative sea-level
change is therefore
DzðtÞ ¼ ðzA  zB Þ  ur
and the change in elevation of any point (e.g., from C to C0
on land or from D to D0 offshore) measured with respect to
the sea level of the epoch is
hðt Þ ¼ hðtP Þ  ½ðzA  zB Þ  ur  ¼ hðtP Þ  Dzðt Þ
This formulation is independent of the cause of land
movement or ocean volume change but for the glacio-
hydro-isostatic contribution only, the land movement
component will depend on the response of the earth’s sur-
face to the changing ice–water load and the displacement
of the ocean surface will depend on how any meltwater is
distributed into the ocean basins and on the changing grav-
itational attraction of the solid earth and the ice on the
water surface. In the case of glacio-hydro isostasy ur and
the radial displacement of the ocean surface (zA  zB).
494 GLACIO-HYDRO ISOSTASY
Glacio-Hydro Isostasy, Figure 2 Schematic representation of relative sea-level change Dz in an environment of land uplift ur and
a rise in the radial distance of the ocean surface (zA  zB). (See text for details).
Consider the following simple model. At the end of
a long glacial period, tA, a volume of ice has been added
rapidly into the ocean at a constant rate until a time tB.
The ocean volume then remains constant until the present
tP (Figure 3a). During the deglaciation the meltwater loads
the ocean floor and induces flow in the mantle away from
the areas of loading to beneath the continents and to com-
pensate for the concomitant rebound beneath the ice
sheets. The resulting subsidence of the seafloor will
include an elastic response followed by a viscous response
that continues after the load has stabilized and will still
occur today. The ocean has to fill the void created by the
subsiding seafloor and the sea surface, in a first approxi-
mation, follows the seafloor. In the middle of the ocean
basins, the maximum subsidence can be roughly approxi-
mated by the local isostatic effect. At the continental mar-
gin, because the water loading occurs to one side only, the
subsidence will be about 50% of that in mid-ocean, largely
because of the flexural strength of the lithosphere. At far-
offshore island sites, there is no change because the island
subsides with the seafloor. Inland from the coast, within
a deeply indented gulf, for example, where the crust is
being progressively dragged down less by the ocean load,
this change will approach ur (Figure 3b).
The resulting spatial variability in the hydro-isostatic
response will then have the following characteristics
(Figure 3c):
 A small amplitude highstand near the coast with
a maximum at the time meltwater addition ceased and
with a magnitude that depends on the distance from
the coast.
 A rise of sea level prior to the maximum highstand that
occurs later with distance from the coast.
 Relative sea level at the LGM that varies across the
shelf and that approaches the ice-volume equivalent
value at some distance from the coast.
This scenario is an incomplete description of the relative
sea-level change across the continental shelf since the
glacio-isostatic contribution has been ignored, as has the
change in gravitational potential during the deglaciation
and subsequent relaxation phase and any change in water
load caused by the migration of shorelines during the sea-
level rise. Of these neglects the most important one is the
glacio-isostatic factor, including the gravitational part. At
the time of maximum glaciation a broad deformational
bulge develops around the ice sheet and this slowly sub-
sides during and after the deglaciation phase. When this
bulge occurs in an ocean environment, additional space
is created that draws water into it and away from the far-
field locations where the glacio-isostatic signal becomes
one of a falling sea level (Figure 3d). This contribution
varies relatively slowly across the far field and the pre-
dominant spatial variability across a reef system will come
from the hydro isostasy. The combined effect at the far-
field sites is that a small amplitude highstand will also
occur for many ocean island sites (Figure 3e). It is some-
times stated that ocean islands act as dipsticks and provide
a direct measure of the changes in ocean volume. But this
is a first approximation and in reality this occurs only at
a few sites where the combination of all the factors con-
tributing to the isostatic perturbation cancels out. These
simple illustrations also point to a second common
misconception, that it is the loading of the shelves that
constitutes the hydro-isostatic correction. But this is one
part of the water load only and the principal part comes
from the long-wavelength load of the entire ocean floor.
 GLACIO-HYDRO ISOSTASY 495

Glacio-Hydro Isostasy, Figure 3 Schematic components of the contributions to relative sea level at far-field sites across a continental
margin. (a) The ice-volume equivalent sea-level (ESL) component. (b) The hydro-isostatic component at three locations across the
margin. The inland site would correspond, for example, to the head of a deeply indented gulf. (c) The combined ESL and hydro-
isostatic contributions. (d) The glacio-hydro-isostatic contribution at the far-field locations. This part varies only slowly across the
area. (e) The total glacio-hydro-isostatic change in sea level.

Some examples
Figure 4 illustrates predictions based on the complete the-
ory, for sites equidistantly spaced across the Great Barrier
Reef (Queensland, Australia) from within the inner Broad
Sound, across the outer Reef and to Marion Reef in the
Coral Sea. They are based on nominal ice and earth
models that have been found to describe well the far-field
behavior of sea level during Holocene time and are repre-
sentative of a wide range of plausible models. In this
model calculation, it has been assumed that all melting
ceased abruptly at 6,000 (C14) years BP and this results
in the sharp highstand at this time. If this cessation was
less abrupt then this peak becomes broader and its shape
is diagnostic of changes in late Holocene ocean volumes.
This example illustrates that with increasing distance from
the continental coastline:
 the mid-Holocene highstand decreases in amplitude
from a few meters maximum at the ‘inland’ sites;
 the oldest corals occurring at sea level become increas-
ingly younger;
496 GLACIO-HYDRO ISOSTASY

 the pre-mid Holocene rise in sea level increasingly lags

Glacio-Hydro Isostasy, Figure 4 Predicted sea-level change for
the complete glacio-hydro-isostastic rebound along a section at
a spacing of ~35 km across the Great Barrier Reef (Australia) from
the inner side of Broad Sound to Marion Reef within the Coral
Sea. These results are for nominal earth parameters and ice
sheets that cease all melting at 6,000 14C years before present.
the globally averaged sea-level rise; and
 the Last Glacial Maximum sea levels increase in depth.
Across the Great Barrier Reef, adjacent islands and conti-
nental shoreline, the observational evidence generally
confirms these trends although it has not always been
interpreted in terms of the differential isostatic response.
For example, the observation of a progressive decrease
with distance from the shore of the oldest coral ages
found at the upper limits of growth, has sometimes been
interpreted as a result of outer reefs being less capable of
keeping up with sea-level rise than inner reefs.
The amplitudes of the highstand and the lags in the
response are dependent on the rheological parameters
describing the earth’s mantle and lithosphere so that the
matching of observations and predictions enables these
parameters to be estimated for the region of analysis.
Glacio-hydro-isostatic studies of the formerly glaciated
(near-field) regions likewise provide information on the
earth’s response function as well as on the dimensions
and timing of the former ice sheets, particularly during
their final decay phase. By an iterative process between
far- and near-field regions, it becomes possible to separate
the earth- and ice-model parameters that quantify the
rebound formulation and thus to develop a predictive
global model for that part of sea-level change associated
with the glacial cycles. These models are best developed

Glacio-Hydro Isostasy, Figure 5 Glacio-hydro-isostatic sea-level variation across the Japan Sea and adjacent areas at two epochs.
(a) 20,000 (cal) years before present, at the time of the maximum glaciation. The contour interval is 5 m. (b) 7,000 (cal) years before
present, at the time of the mid-Holocene highstand formation. The contour interval for sea-level change is 1 m. The red contours are
negative and the orange contours are positive.
 GLOBAL OCEAN CIRCULATION AND CORAL REEFS
for the interval after the last glacial maximum but they
have also been extended to earlier periods, particularly
the Last Interglacial. In the context of coral reefs from this
latter period, it is important to recognize that these iso-
static processes also occur and that the elevations of Last
Interglacial sea levels will exhibit similar patterns of spa-
tial variability.
Figure 5 illustrates the predicted spatial variability for
one region, the Japan Sea, that, in the absence of tecton-
ics, is representative of most far-field regions. The results
are for two epochs, the time of the last glacial maximum
(LGM) and the time of the mid-Holocene highstand but
the spatial variability is similar for all other epochs.
Thus, even within relatively small regions sea level can-
not be represented by a single curve. At the LGM the
predicted sea levels vary by
30 m, being significantly
shallower in the Korea Strait, for example, than in the
middle of the Japan Sea where it exceeds the ESL value.
Likewise, the Holocene highstand amplitudes are
predicted to occur up to 2 m above sea level in some
locations but not in northern Hokkaido or the Kurile
Islands.
Conclusion
The phenomenon of glacio-hydro isostasy means that
even in the absence of vertical tectonics, sea level at any
time is spatially variable, that it can be rising in one loca-
tion while falling elsewhere. This variability is a conse-
quence of the earth’s deformation under the changing
load and to the changes in the gravity field. The mapping
of this variability therefore provides information on both
the earth’s rheology and on the history of the ice sheets.
But it also needs to be known if sea level is used as
a reference for vertical tectonics or, for example, if reef
response to changing sea level is to be investigated. The
theory and implementation of the glacio-hydro-isostasy
theory is sufficiently well developed for it to provide
a sound basis for interpolating between the limited
sea-level observations for the time since maximum glacia-
tion and for reconstructing paleo shorelines and water
depths.
Bibliography
Lambeck, K., and Chappell, J., 2001. Sea level change through the
last glacial cycle. Science, 292, 679–686.
Lambeck, K., Purcell, A., Johnston, P., Nakada, M., and Yokoyama, Y.,
2003. Water-load definition in the glacio-hydro-isostatic sea-level
equation. Quaternary Science Reviews, 22, 309–318.
Mitrovica, J. X., and Milne, G. A., 2003. On post-glacial sea level: I.
General theory. Geophysical Journal International, 154,
253–267.
Cross-references
Last Glacial Lowstand and Shelf Exposure
Sea Level Change and Its Effect on Reef Growth
Volcanic Loading and Isostasy
497
GLOBAL OCEAN CIRCULATION AND CORAL REEFS
John E. N. Veron
Coral Reef Research, QLD, Townsville, Australia
Synonyms
Global scale currents
Definition
Ocean circulation denotes ocean-scale current systems
that change seasonally but persist from one year to the
next.
Introduction
Reefs develop in most parts of the world where reef-
building (zooxanthellate) corals grow and where the ocean
is warm. This geographic template is created by the
world’s great ocean currents (Figure 1). Currents therefore
play a key role in the geographic extent of reef formation
as well as in the dispersal of reef biota. They are also crit-
ical determinants of the pathways of species evolution.
The historical role of ocean circulations as drivers of
reef and faunal distributions is closely linked to other
drivers, not only temperature but also sea-levels and
the configuration of the continents. Two very different
scales – global and regional – operate simultaneously.
Global scales over geological time provide the template
of both coral reef distributions and zooxanthellate coral
distribution. Regional scales over millennia or less control
details of the distributions of corals and have a short-term,
ecologically driven influence on reef development (see
below).
Origins of modern circulation patterns
At the start of the Cenozoic Era, ocean circulations were
dominated by a circum-global equatorial path via the
Tethys Sea and Central American Seaway. There was no
circulation around Antarctica (the remnants of Gondwana)
and all oceans were warm and sluggish, with no strong lat-
itudinal temperature gradients as we have today. This pat-
tern persisted until South America rifted free of Antarctica
during the Late Eocene–Early Oligocene 40–35 million
years ago (mya), and Drake Passage opened. At this point,
for the first time since Gondwana formed, the Southern
Ocean began circulating around Antarctica, moving east-
ward in response to the direction of rotation of the Earth
and checked only by the coastlines of the three southern
continents.
The formation of the Antarctic Circumpolar Current
was the critical event that gave birth to modern circulation
patterns. Where the three continents of South America,
Africa, and (to a lesser extent) Australia project into its
path, cold water is deflected north along the western mar-
gin of each continent. The currents then move westward in
the tropics where they are warmed, and return south along
498 GLOBAL OCEAN CIRCULATION AND CORAL REEFS
Global Ocean Circulation and Coral Reefs, Figure 1 The global pattern of ocean circulation (after Veron, 2000).
the eastern side of each continent. These three anticlock-
wise circulations of the Southern Hemisphere oceans are
in turn linked to clockwise circulations in the Northern
Hemisphere in latitudinally zoned gyres (Figure 1). In
effect, the opening of Drake Passage allowed the sluggish
tropical circulations of the early Cenozoic to be replaced
with stronger circulations, powered by the Antarctic Cir-
cumpolar Current, creating the latitudinal temperature
gradients of today. It also formed an insulating blanket
around Antarctica causing, at the time of formation, the
surface temperature of the Southern Ocean to plunge by
10–15 C (Frakes et al., 1992).
During the latter half of the Cenozoic, there were two
fundamental modifications to this pattern (Adams,
1981). The first was the widening of the North Atlantic,
as, until this happened, the North Atlantic did not regulate
global climates via the Ocean Conveyor and Gulf Stream
as it now does. The second was the progressive reduction
in size of the Tethys Sea. The Tethys, the evolutionary cra-
dle of most tropical marine life, waxed and waned
throughout the Early to Middle Cenozoic according to
continental movements and sea-level changes, to be
finally choked off during the Late Miocene (10 mya),
a process which severed all tropical connection between
the Atlantic and Indian Ocean (Figure 2).
There was one final event crucial to ocean circulation
before Ice Age cycles took control: the formation of the
Isthmus of Panama in place of the old Central American
Seaway. This was a gradual process completed in
3.4 mya (Coates et al., 1992), which greatly enhanced
the flow of the Ocean Conveyor to the North Atlantic
(Burton et al., 1997) and allowed El Niño Southern Oscil-
lation cycles to establish.
Ocean circulation and biogeographic patterns
The distribution of corals and reefs during the Early Ceno-
zoic essentially involved a transfer of biodiversity from
the Old World of the Tethys Sea to the New World of the
Caribbean and Central American Seaway. Through this
process, coral diversity around the Central American Sea-
way reached a peak during the Oligocene and extensive
reef development also spread throughout the central
Indo-West Pacific, which had become clearly distinct
from the Tethys.
By Early Miocene, the distribution of reefs was similar
to that of today, with the major exception of the Tethys Sea
which was held open only by a narrow passage between
the Indian Ocean and proto-Mediterranean (Figure 2).
The biogeographic patterns of corals at this time is
unknown, but by then the Indo-West Pacific had inherited
much of the diversity that had originally evolved in the
Tethys and Caribbean (Veron, 1995). The Miocene ended
5.3 mya with the extinction of the last remnants of the
ancient fauna of the Tethys. Late Miocene reef remains
are very common in the western tropical Pacific (Kiessling
and Flügel, 2002), with outcrops extending as far south as
New Zealand (Hayward, 1977).
 GLOBAL OCEAN CIRCULATION AND CORAL REEFS 499

Global Ocean Circulation and Coral Reefs, Figure 2 Positions of the continents during the Middle Miocene (around 15 mya).
Extensive reef remains (in black) are scattered around the world and as far south as New Zealand (after Veron, 2008).

Global Ocean Circulation and Coral Reefs, Figure 3 The role of poleward currents in latitudinal reef distribution along eastern
Australia. The East Australian Current disperses coral larvae to the world’s highest latitude atolls (Elizabeth and Middleton Reefs) and
highest latitude reef (Lord Howe Island). South of the Great Barrier Reef, there are abundant corals but no reef development at the
few shallow coastal localities – Flinders Reef (a sandstone outcrop) and the Solitary Islands – due to temperature and other
environmental constraints (after Veron, 2000).
500 GLOBAL OCEAN CIRCULATION AND CORAL REEFS

At this time, corals had yet to undergo one more major
change in their global pattern of diversity. Throughout
much of the Miocene (24–5.3 mya), but certainly by the
Late Miocene, the eastern Atlantic had become too cold
for reef corals as a result of the cold current plying along
the west African coast (Esteban, 1980). The Atlantic thus
became an isolating barrier separating the Caribbean from
the Tethys where once it was a corridor.
Ocean circulation and reef formation
It is only at oceanwide scales that there is only a clear cor-
relation between the distribution of corals and the distribu-
tion of reefs, although surface currents play a dominant
role in both. The essential reason is that good reef develop-
ment can occur in regions of low coral diversity: reef for-
mation does not depend on a high coral diversity (Glynn
et al., 1996).
With rare exceptions (carbonate platforms built by
vermetid molluscs, serpulid worms, and non-scleractinian
corals), the presence of zooxanthellate corals is essential
for reef development. Thus, the oceanographic history
of any major part of the world, as described above,
determines the gross pattern of both reef development
and zooxanthellate coral occurrence. Both require
shallow substrata, minimal quantities of mud of terrestrial
origin, high light levels, high salinity, a suitable tem-
perature regime, and Oaragonite of 3.8 or higher (see

Global Ocean Circulation and Coral Reefs, Figure 4 The role of surface currents in dispersal of corals along the eastern Australian
coast (a) and the islands of Japan (b). Dendrograms show similarity of species composition occurring at each site indicated. These
include reef communities, non-reef communities, and outlying regions (after Veron, 1995).
 GLOBAL OCEAN CIRCULATION AND CORAL REEFS 501

Chapter “Corals – environmental controls on growth”). Of
these constraints, all but temperature applies equally to
corals and reefs on regional scales.
Temperature is more limiting for reefs than corals
because of an ecological constraint: most corals can toler-
ate temperatures as low as 14 C for extended periods, but
consolidated reefs will not accrete under 18 C because at
that temperature corals cease to be competitive with
macro-algae in most parts of the world (Veron and
Minchin, 1992). Along subtropical continental coastlines
where at least some zooxanthellate corals occur, reefs
extend to a latitudinal limit defined by temperature which,
in turn, is controlled by the strength of poleward currents
(for example, Figure 3). In open oceans remote from con-
tinents, the same constraint applies where suitable sub-
strates exist, but as poleward currents are weaker, the
latitudinal extent of reef development is less.
One further aspect of the role of temperature in control-
ling the latitudinal range of reefs is that reefs can develop
episodically, surviving as geological structures during
intervals of emergence and cold (such as were created by
the glacial cycles of the Pleistocene) and growing during
interglacial warm intervals.
Surface currents and dispersal
There is a wealth of information from experiments, genet-
ics, taxonomy, and biogeography that coral larvae are able
to make very long ocean journeys, some enduring weeks,
even months, on the sea surface before settling (Rich-
mond, 1987). Species endurance therefore depends on
the speed and direction of currents as well as the existence
of suitable substrates.
Survival during these voyages depends on time, there-
fore on the distance traveled, and speed.
Corals are dispersed along continental coastlines of
eastern and western Australia by the East Australian Cur-
rent and Leeuwin Current (respectively) in the Southern
Hemisphere and along the islands of Japan in the
north by the Kuroshio. In each case, there is an orderly
“drop-out” sequence where species numbers progressively
attenuate (Veron, 1995). Each of these boundary currents
varies in position and strength, each variation potentially
causing changes in the number and variety of propagules
reaching high-latitude reefs. Importantly, as each current
flows poleward, propagules entrapped in them are on non-
return journeys. As a result, these currents overwhelmingly
dominate the nature and diversity of coral communities
found today (Figures 3, 4a, b; Harriott and Banks, 2002).
Surface currents also play a part in defining the global
center of coral diversity, the so-called “Coral Triangle”
(see Chapter “East Indies Triangle of Biodiversity”;
Figure 5).
There are multiple reasons why the Coral Triangle
should be so diverse relative to other reef regions (Veron
et al., 2009): (1) The Coral Triangle acts as a ‘catch-all’
for larvae moving towards the region, entrained in both
the South Equatorial Current and the North Equatorial
Current (Jokiel and Martinelli, 1992; Veron, 1995).
(2) Dispersion occurs away from the Coral Triangle leav-
ing that region biodiverse relative to other regions where
attenuation has occurred. (3) Complex eddies created by
the Indonesian Through-flow (Gordon and Fine, 1996)
drive genetic mixing which constantly changes with wind,
season, and (over geological time) sea level. Genetic
mixing of this nature creates genetic heterogeneity
through vicariance and reticulate evolution.
Ocean circulation and evolution
Evolution in corals is exceedingly slow, whereas long-
distance dispersal can be rapid – even within the time of
a single generation. The two processes are therefore only
tentatively linked, not only because of time but also
because genetic pathways are constantly changed by vari-
ations in ocean currents.
The distance traveled by coral larvae and probably that
of most other major invertebrate taxa can indeed be so vast
that the majority of species found in, for example, the
Great Barrier Reef also occur within the Coral Triangle

Global Ocean Circulation and Coral Reefs, Figure 5 Contours of coral diversity (species richness). Species disperse from regions of
high diversity, notably the Coral Triangle, to other Indo-Pacific regions. Many regions of the world, including most of the Atlantic
(but not the Caribbean) and far eastern Pacific, have few, if any, corals and no reefs. However, over distances of a thousand kilometers,
the occurrence of both corals and reefs is highly predictable (after Veron, 2008).
502 GLOBAL OCEAN CIRCULATION AND CORAL REEFS

and a high proportion are even recognizable as far away as

the Red Sea.
The relative strength and direction of the currents and
the endurance of the larvae are key aspects of these spatial
patterns – not just the patterns of today, but those of all
times past, for as long as these species have existed.
Today, most major currents are generally stable and move
in predictable directions. However, over geological time,
these same currents have changed their strengths and
directions repeatedly. This has created changing geo-
graphic patterns of genetic mixing quite unlike genetic
pathways normally found on land. The result, over geo-
logical time, is a template of constant change, which in
turn is the mechanism that drives reticulate evolution
(see Chapter Scleractinia, Evolution and Taxonomy).
The Kuroshio, the world’s strongest continental bound-
ary current, well illustrates change over time in both
strength and direction. In historical time (decades to centu-
ries), this current has repeatedly changed its path south
of mainland Japan (Figure 6a), each time causing major
redistributions of benthic fauna as well as local extinc-
tions. Over geological intervals of 100,000s of years, com-
binations of glacial sea-level changes and tectonic uplifts
have created enormous changes to the Japanese coastline
and, consequently, the path of the Kuroshio (Figure 6b).
Variations between these two extremes not only produced
changes in distribution and diversity of most marine biota
but also changes in their genetic compositions (Figure 6).

Meso- and microscale circulation and environments

Most marine life undergoes dispersion, either long-
distance (noted above) or within single reefs during larval
stages in their life cycles, and most benthic and neritic life
becomes established on particular areas of reef according
to local environmental cues. At these mesoscales, currents
are also the conveyors of food for filter – and detritus –
feeding taxa, and oxygen and nutrients for all taxa.
At yet finer scales, currents (generated by tides and waves)
determine the growth form of branching corals and
thereby the three-dimensional nature of most types of
shallow reef habitats. These aspects of ocean currents are
outside the scope of this article.
Global Ocean Circulation and Coral Reefs, Figure 6 Changing
ocean circulation patterns of the Kurioshio. Over historical times
(a), major changes have occurred in the path of the Kuroshio
south of mainland Japan. Some of these are seen in fossil coral
outcrops (arrowed). Over 100,000s years, the coastline has
changed radically from sea-level changes and tectonic uplift,
creating far greater changes to the Kuroshio (after Kimura, 1991).
Variations between these extremes have created a constantly
changing template for evolutionary change.

Summary
The global distribution of reefs is a product of the major
patterns of ocean circulation that have existed since Early
Cenozoic which, in turn, were dependent on continental
movements. Reef distributions are also controlled by the
presence or absence or reef-building corals and therefore
the physical parameters which control the growth of
zooxanthellate corals. Of these, temperature is critical,
the lower boundary of which is different for reefs and
corals. Currents are also the pathways of dispersal of most
marine taxa and are thus the principal drivers of evolution-
ary change. On finer scales, currents are an intimate part of
reef ecological processes and morphologies.
Bibliography
Adams, C. G., 1981. An outline of Tertiary paleogeography. In
Cocks, L. R. M. (ed.), The Evolving Earth. Cambridge, UK: Brit-
ish Museum (Natural History) and Cambridge University Press,
Vol. 14, pp. 221–235.
Burton, K. W., Ling, H. F., and O’Nions, R. K., 1997. Closure of the
central American isthmus and its effect on deep-water formation
in the North Atlantic. Nature, 386, 382–385.
Coates, A. G., Jackson, J. B. C., Collins, L. S., Cronin, T. M.,
Dowsett, H. J., Bybell, L. M., Jung, P., and Obando, J. A.,
1992. Closure of the Isthmus of Panama: The Near-Shore Marine
Record of Costa Rica and Western Panama. Geological Society
of America Bulletin, 104(7), 814–828.
 GREAT BARRIER REEF COMMITTEE
Esteban, M., 1980. Significance of the Upper Miocene Coral Reefs
of the Western Mediterranean. Palaeogeography, Palaeocli-
matology, Palaeoecology, 29, 169–188.
Frakes, L. A., Francis, J. E., and Syktus, J. I., (1992) Climate Modes
of the Phanerozoic. Cambridge, UK: Cambridge University Press.
Glynn, P. W., Veron, J. E. N., and Wellington, G. M., 1996.
Clipperton Atoll (eastern Pacific): oceanography, geomorphol-
ogy, reef-building coral ecology and biogeography. Coral Reefs,
15, 71–99.
Harriott, V. J., and Banks, S. A., 2002. Latitudinal variation in
coral communities in eastern Australia: a qualitative biophysical
model of factors regulating coral reefs. Coral Reefs, 21, 83–94.
Hayward, B. W., 1977. Lower Miocene Corals from the Waitakere
Ranges, North Auckland. New Zealand Journal of the Royal
Society of New Zealand, 7, 99–111.
Jokiel, P. L., and Martinelli, F. J., 1992. The vortex model of coral
reef biogeography. Journal of Biogeography, 19, 449–458.
Kiessling, W., and Flügel, E., 2002. Phanerozoic reef patterns.
Society of Economic Paleontologists and Mineralogists. Special
Publication. Special Publications, Vol. 72, pp. 77–92.
Kimura, M., 1991. Quaternary land bridges of the Ryukyu arc
detected on seismic reflecting profiles. Spec, 35, 109–117
Vol. U. Tohoku (in Japanese).
Richmond, R. H., 1987. Energetics, competency, and long-distance
dispersal of planula larvae of the coral Pocillopora damicornis.
Marine Biology, 93, 527–533.
Veron, J. E. N. et al., 2009. Delineating the Coral Triangle. Galaxea.
Veron, J. E. N., 1995. Corals in Space and Time. New York: Cornell
University Press.
Veron, J. E. N., 2000. Corals of the World. Australian Institute of
Marine Science Vol. 3.
Veron, J. E. N., and Minchin, P. R., 1992. Correlations between sea
surface temperature, circulation patterns and the distribution of
hermatypic corals of Japan. Continental Shelf Research, 12,
835–857.
Cross-references
Corals: Environmental Controls on Growth
Scleractinia, Evolution and Taxonomy
East Indies Triangle of Biodiversity
Lagoon Circulation
Tethys Ocean
Reef Interconnectivity/Larval Dispersal
GREAT BARRIER REEF COMMITTEE
David Hopley
James Cook University, Townsville, Queensland,
Australia
In 1922, the Queensland branch of the Royal Geographi-
cal Society of Australia set up the Great Barrier Reef Com-
mittee following an address to the Society by the Professor
of Geology and Mineralogy at the University of Queens-
land, Henry Richards. The address, entitled “Problems of
the Great Barrier Reef” highlighted the need for scientific
investigation of the Reef and followed a visit by Richards
to America to attend the Pan-Pacific Scientific Conference
where, among others, he met W.M. Davis (Hill, 1985a, b;
Jones, 1974; Bowen and Bowen, 2002).
503
Membership of the Committee was through invited
nominations from the Government and Academic institu-
tions in Australia and elsewhere. However, disagreements
with the parent body, the Royal Geographical Society led
to a separation of the Committee in 1924 although it
incongruously maintained the title of Committee. Initial
funding came from the Queensland Government and Uni-
versities of Sydney and Queensland and further valuable
support came from the provision of places on survey ves-
sels of the fledgling Australian Navy and the Common-
wealth steamer that serviced the many lighthouses on the
reefs and islands.
Early research was much influenced by Richards’ geo-
logical background with, for example, Sydney University
graduates undertaking work into coastal landforms and
sea level, published in the independent Transactions of
the Great Barrier Reef Committee. However, Richards
had larger targets in mind including investigations into
the foundations of the Reef to see if Darwinian type
subsidence was applicable. In 1926, a hole was drilled to
180 m on Michaelmas Cay, and in 1937 further drilling
on Heron Island to 240 m was undertaken. Results were
inconclusive as to the proving or otherwise of the
nineteenth century hypotheses of Darwin, Dana, and
others but did provide useful information, which was
being quoted even 50 years later (e.g., Lloyd, 1973).
Another early aim of the Committee was to set up
a research station on one of the reef islands but as this
was not immediately possible, an alternative strategy
was adopted and that was to promote an expedition based
on a reef island for at least one full cycle of seasons. Dis-
cussions were held with various scientific bodies, espe-
cially in England and support came especially from the
British Association for the Advancement of Science. The
cay on Low Isles, north of Cairns, was the chosen site
where 23 scientists were based under the leadership of
(Sir) Maurice Yonge in 1928–1929. A variety of studies,
including zoological, botanical, sedimentological etc.,
were undertaken and the large number of reports were
published by the British Museum of Natural History. They
have remained the basis for coral reef research ever since.
In addition to the main expedition, the Royal Geo-
graphical Society supported a smaller study of mainly
islands of the southern half of the Great Barrier Reef
(GBR). This was led by Professor J.A. Steers of
Cambridge University and accompanied by Michael
Spender and was followed by a second 4-month expedi-
tion mainly to previously unvisited islands of the Reef in
1936. Their results were published in the Geographical
Journal and Reports of the Great Barrier Reef Committee
(Steers, 1929, 1937, 1938; Spender, 1930). This work
did much to highlight the value of reef islands as
a source of geomorphological information on coral reefs
and established Cambridge University’s Geography
Department as a source of reef researchers.
Establishment of a permanent research station on the
GBR was not possible until after World War II. Initially
a relatively small facility, it was set up in 1951 on Heron
504 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
Island at the southern end of the GBR and close to the cap-
ital city of Brisbane. This has subsequently become one of
the major reef research facilities in the world.
During the second half of the twentieth century, the
Committee was very active in conservation issues of the
GBR, something which Richards had originally seen as
being important. Scientific evidence about the crown-of-
thorns starfish and oil drilling on the Reef, and the need
to create the Great Barrier Reef Marine Park Authority
was presented to a number of government commissions.
The Committee also played a part in the organization of
the Second International Coral Reef Symposium in 1973,
which uniquely was held aboard the cruise ship “Marco
Polo” travelling from Brisbane to Lizard Island and return.
Also in the same year, the Committee was involved in the
organization of a further expedition to the northern GBR.
Enthusiastic support came from Sir Maurice Yonge and
Professor Alfred Steers from the 1928–1929 expedition.
Major input came from the Royal Society of London and
from the Universities of Queensland (Queensland Univer-
sity and the new James Cook University of North Queens-
land). The leader of the expedition was Dr. David Stoddart
of Cambridge University. Again, significant data was
obtained especially from the reef islands and comparison
with results from 1928–1929 to 1936 allowed some
assessment of the dynamics of the islands. Results of this
work were published in 1978 in the Philosophical Trans-
actions of the Royal Society, London A, 291 and B, 184.
In 1988, after 66 years as an unattached committee with
nominated membership, the Great Barrier Reef Commit-
tee became the Australian Coral Reef Society (ACRS)
with subscription membership. However, continuity is
recognized and this makes it the oldest organization
concerned with coral reefs in the world.
Bibliography
Bowen, J., and Bowen, M., 2002. The Great Barrier Reef, History,
Science and Heritage. Cambridge: Cambridge University Press,
454 pp.
Hill, D., 1985a. The Great Barrier Reef Committee 1922–1982: the
first thirty years. Historical Records of Australian Science, 6(1),
1–18.
Hill, D., 1985b. The Great Barrier Reef Committee 1922–1982: the
last three decades. Historical Records of Australian Science,
6(2), 195–221.
Jones, O. A., 1974. The Great Barrier Reef Committee - 1922–
1973. Proceedings Second International Coral Reef Symposium,
2, 733–737.
Lloyd, A. R., 1973. Foraminifera of the Great Barrier Reef bores. In
Jones, O. A., and Endean, R. (eds.), Biology and Geology of
Coral Reefs, 1, Geology, 1, 347–366.
Spender, M. A., 1930. Island reefs of the Queensland coast. Geo-
graphical Journal, 76, 194–214, 273–297.
Steers, J. A., 1929. The Queensland coast and the Great Barrier
Reef. Geographical Journal, 74, 232–257, 341–370.
Steers, J. A., 1937. The coral islands and associated features of the
Great Barrier Reefs. Geographical Journal, 89, 1–28, 119–146.
Steers, J. A., 1938. Detailed notes on the islands surveyed and
examined by the Geographical Expedition to the Great Barrier
Reef in 1936. Reports of the Great Barrier Reef Committee, 4,
51–96.
Cross-references
Daly, Reginald Aldworth (1871–1957)
Dana, James Dwight (1813–1895)
Darwin, Charles (1809–1882)
Davis, William Morris (1850–1934)
Great Barrier Reef: Origin, Evolution, and Modern Development
Royal Society of London
Steers, James Alfred (1899–1987)
Stoddart, David Ross (1937–)
Yonge, Sir Maurice (1899–1986)
GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND
MODERN DEVELOPMENT
Peter J. Davies
University of Sydney, Sydney, Australia
Introduction
The Great Barrier Reef (GBR) is arguably the largest coral
reef system to have ever existed on the planet. It lies off the
northeast Australian margin – a passive margin, but
a margin which is being gradually consumed as
a consequence of Australia’s inexorable drift toward the
New Guinea foreland basin and the trenches to the north.
Unless plate motions change, the Great Barrier Reef is
a temporary acquisition. The following is the story of its
development to date – one of three major reef systems to
have existed in the region since the Miocene, and the Great
Barrier Reef owes its existence to both its predecessors.
The architecture of offshore northeast Australia
The northeast Australian margin (Figure 1a) comprises the
continental shelf and a number of marginal plateaux and
rift troughs: the Eastern, Queensland, and Marion Pla-
teaux; the Pandora and Bligh Troughs; the Osprey Embay-
ment; and the Queensland and Townsville Troughs and the
Cato Trough. In addition, a zone of narrow rift basins
extend southeast beneath the shelf of the GBR from the
Queensland Trough southeast toward the Capricorn Basin
and separating the Marion Plateau from the inner and mid-
dle continental shelf. The entire margin is underlain by
modified continental crust formed as a result of fragmenta-
tion of a northeastern extension of the Tasman Fold Belt
(Gardner, 1970; Ewing et al., 1970; Falvey, 1972; Falvey
and Taylor, 1974; Taylor, 1975; Mutter, 1977; Taylor
and Falvey, 1977; Mutter and Karner, 1980; Symonds,
1983; Symonds Davies and Parisi, 1984). The main phys-
ical and structural elements of offshore Queensland are
shown in Figure 1b and briefly described below:
Eastern Fields Plateau
The Eastern Plateau is the most northern marginal plateau
of the northeast Australian continental margin (Figures 1a, b,
and 2a) and is bounded in the west by the Moresby, Pan-
dora, and Bligh Troughs, in the south by the Osprey
Embayment, in the north by the Moresby Canyon, and in
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
the east by the Coral Sea Basin (Figure 1a and b). The pla-
teau has an average depth of 1,500 m, and covers an area
of about 31,000 km2 at the 2,000 m isobath. Eastern Fields
Reef, the only modern reef on the Eastern Plateau, is about
45 km across at its widest point, and lies at the crest of the
plateau near its northern margin. Submerged and buried
reefs extend northeast from Eastern Fields Reef beneath
the Moresby Trough (Davies et al., 1988). The northern
and southern plateau margins are controlled by normal
faults; the western margin is more complex, and appears
to be a product of thrusting. The Eastern Plateau is
founded on a complex of tilt blocks bounded by
reactivated normal faults, some of which appear to have
undergone both wrench and reverse movement (Davies
et al., 1988). This deformation appears to have resulted
from Late Oligocene and Miocene tectonism caused by
the development of the New Guinea Orogen to the north
(Pigram and Davies, 1987). This late phase of structuring
is not represented on the other northeast Australian mar-
ginal plateaux.
Queensland Plateau
The Queensland Plateau is the largest marginal plateau on
the Australian continental margin and is one of the largest
features of its type in the world (Figure 1a and b). It is
bounded on the northeast by the Coral Sea Basin, on the
west by the Queensland Trough, and on the south by the
Townsville Trough. The plateau is roughly triangular in
shape, and extends over an area of about 165,000 km2.
Approximately half of the plateau surface lies above the
1,000 m isobath, with living reef systems at or near present
sea level making up 10–15% of the surface. The largest
modern reef complexes are Tregrosse and Lihou Reefs,
lying along the southern margin of the plateau
(Figure 1a). Both these complexes are nearly 100 km long
from east to west, and 50 and 25 km wide, respectively,
from north to south. The other major areas of modern reef
growth are the Coringa, Willis, and Diana complexes,
which are aligned north to south in the center of the pla-
teau, and the large isolated pinnacles of Flinders, Holmes,
Bougainville, and Osprey Reefs, which lie along the west-
ern margin of the plateau (Figure 1). In addition, drowned
reefs have been reported from at least 25 different loca-
tions (Taylor, 1977; Mutter, 1977; Davies et al., 1988).
Away from reef areas, the plateau surface is generally
smooth and slopes northward. A distinct terrace at approx-
imately 450–500 m depth connects both the Willis and
Diana reef complexes and the Tregrosse-Lihou-Coringa
reef complexes.
Marion Plateau
The 77,000 km2 area of the Marion Plateau lies directly
east of the central Great Barrier Reef, and is bounded
along its northern margin by the Townsville Trough and
along its eastern margin by the Cato Trough (Figure 1a).
The present plateau surface forms a deep water extension
of the shelf of the central Great Barrier Reef, with water
505
depths ranging from 100 m along the western border to
500 m along the eastern margin. At present, reef growth
is restricted to Marion Reef, on the northeastern corner,
and Saumarez Reef, at the southeastern corner of the pla-
teau (Figure 1a).
The Marion Plateau is bounded on three sides by rifts:
the Cato Trough to the east, the Townsville Trough to
the north, and in the west by a series of north–south-
oriented, narrow half-grabens which separate the plateau
from the Australian continent (Figure 1b). The most north-
ern of these half-grabens appear to join the confluence of
the Townsville and Queensland Troughs. In the early Ter-
tiary, therefore, the Marion Plateau formed a separate mar-
ginal plateau. The basement beneath the Marion Plateau is
a planated surface dipping gently northeast (Figure 2d and e).
The only disruption to this surface occurs in the north-
east corner where a basement high forms the pedestal on
which Marion Reef developed. Basement beneath the pla-
teau margins is steeply down-faulted into the troughs to
the north and east and was completely transgressed during
the Early Miocene, resulting in the development of an
extensive carbonate platform on which four, progressively
more, restricted reef growth phases occurred, the last of
which is seen today on Marion and Saumarez reefs. Over
most of the platform, shallow water carbonates ceased
with late upper Miocene/Pliocene subsidence.
The Swains Reef and Bunker Highs
The Swains Reef high (Figure 1a and b) is a structural high
some 200 km long and 50 km wide to the southwest of the
Marion Plateau and forming the eastern boundary of the
Capricorn Rift basin (Figure 1b). It is probably a late Cre-
taceous feature when it formed the eastern side of the
Maryborough basin, but became a prominent late Tertiary
feature when rifting jumped from the Maryborough to the
Capricorn basin and sea floor spreading unzipped north-
west in the Cato Trough. Marine conditions were
established over the Swains High probably in the late
Oligocene/early Miocene. It is thought that reefs
established in the Pleistocene. The Bunker High occurs
to the southwest of the Capricorn Basin (Figure 1a and b).
It formed an eastern volcanic source for the Cretaceous
Maryborough Basin and following rifting in the Tasman
Sea, the western edge of the Cainozoic Capricorn Basin.
It was transgressed in the Miocene with shallow water
marine calcarenites and foraminiferal limestones
establishing as precursors to reefs in the Pleistocene of this
southern region.
The Great Barrier Reef
The Great Barrier Reef is a modern reef system which
extends along the northeast Australian margin from
24.30 to 9.30 S (15 of latitude), a distance of 2,300 km
from north of Fraser Island in the south to the Torres
Shelf/Gulf of Papua in the north. There are estimated to
be in excess of 3,600 individual reefs within this area.
506 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

145 150 155

Gulf of Papua
Papua
New
Guinea

10

Coral Sea
Basin

15

Australia

Ba
rrie
r

20
Re
ef

25

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 1 (Continued)
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 507

Papuan
Basin
Papua
New Guinea

Torres
Shelf

Eastern
Plateau Papuan
Plateau

Coral Sea Basin

Great

Queensland
Plateau
Ba
rrie
r

Marion
Plateau
Re
ef

Capricorn
Basin

Tasman
Basin
b

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 1 (a) The main physical features of the northeast
Australian margin showing in particular the Great Barrier Reef, together with the other main plateaux and rifts. (b) The major
structural features of northeast Australia. The location of major drill holes are shown (BB Borabi, P Pasca, AC Anchor cay,
M Michaelmas cay, AQ Aquarius, CP Capricorn, W Wreck, H Heron, 209 DSDP 209, 811–824 ODP sites).
508 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

PHG Shelf Eastern

Plateau
Pandora
Trough

a
Queensland Plateau Marion
Plateau
Townsville Trough

b
Queensland Trough Queensland Plateau Coral Sea Basin

c
Shelf Marion Plateau Cato Trough

Buried canyon

d
Townsville Trough Marion Plateau Shelf

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 2 Schematic sections showing the generalized structure
and sedimentary relationships on (a) the Eastern Fields Plateau, (b) north to south through the Queensland Plateau (c), west to
east through the Queensland Plateau (d) east to west through the Marion Plateau and (e) north to south through the Marion Plateau.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 509

The shelf occupied by the Great Barrier Reef is
narrowest in the north along 14 S (23 km) and widest in
the south along 21 S (260 km) (Figure 3). Reefs occupy
the whole shelf in the northern region, but apart from
fringing reefs along coasts and large islands, reefs occur
largely in the mid- to outer shelf in the center and south.
The bathymetry is complex and varies regionally, but six
distinct bathymetric regions can be defined (Figures 1
and 3). Area A (Figure 3) in the extreme north (9–12 S)
is a narrow rimmed high-energy platform characterized
by a shallow narrow shelf (50–75 km wide) and a steep
continental slope. Reefs occur in four clearly delineated
groups: the reefs of the western Torres Shelf, the Warrior
line continuing northward off the end of Cape York, the
mid-shelf plethora of small patches, and the outer shelf
barrier reefs. In the region 12 and 16 S the shelf is no
more than 50 km wide except east of Cape Flattery where
it achieves 75 km. Water depths are generally shallow, the
outer shelf being the only place where water depths greater
than 50 m occur. The most prominent reef features in the
region are the outer shelf line of barrier or ribbon reefs
forming an almost continuous shelf edge between Capes
Grenville and Flattery but still prominent down to about
latitude 17 S and east of which the continental slope is
precipitous to a depth of 200–300 m, and also the gigantic
mid-shelf platform reefs, particularly, northeast of

145 150

Papua
New Guinea

10

Zone A (along 1230)

Depth (m)

SL
0
A
60

15
Cooktown
155

Zone B (along 1730)

Cairns SL
B 0
60

Zone C (along 1830)

C SL
Townsville 0

20 60
Bowen
D
Zone D (along 2030)
Mackay SL
0
60

Gladstone Zone E (along 2330) SL

E
0
60
0 60 km
25 Bundaberg

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 3 Physiographic variation in the Great Barrier Reef
province.
510 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
Princess Charlotte Bay. Area B (Figure 3) occurs between
Cape Bedford and Hinchinbrook Island (16 and 18 S),
where the shelf varies in width from 50 to 75 km, and is
bathymetrically relatively simple. Reefs generally occur
as fringing reefs along the coast and as large platform
and ribbon reefs on the mid- to outer shelf; these are sep-
arated from the coast by a channel or inner lagoon with
water depths of around 35 m. The outer shelf ribbon reefs
form a discontinuous barrier particularly in the northern
part of the area but becomes gradually more discontinuous
to the south, ceasing to exist as a line of reefs to the south
of Cairns, at latitude 18S but continuing as line shoals on
the shelf edge averaging 60 m water depth. Area
C (Figure 3) occurs between Hinchinbrook Island and
the Whitsunday Islands (18 and 20 S) where the shelf
takes a clear change in direction, changing from north–
south to north–west to south–east. The width of the shelf
also widens considerably to an average of 90–125 km.
The inner 50–80 km of the shelf is reefless except for
fringing reefs attached to high islands but eastward, the
mid- to outer shelf slopes gently from 45 to 75 m, at which
depth a marked change in slope occurs, reflecting the inner
edge of an outer shelf terrace some 20–30 km in width.
The Slashers, John Brewer, Myrmidon, Pith, Wheeler,
Keeper, Bowl, Stanley, and Viper reefs occupy this outer
shelf terrace. A drowned barrier reef complex occupies
the outer shelf of the 75 m terrace and extends to the south-
east for 200 km. Area D (Figure 3) occurs between the
Whitsunday Islands to the Swain Reefs (20 to 22 S),
the shelf continuing its northwest to southeast orientation
away from the coast giving rise to a dramatic increase in
the width of the shelf to over 300 km in the south. The
inner two thirds of the shelf are reefless except for small
fringing reefs around continental islands; eastward, how-
ever, the outer shelf is a plethora of gigantic reefs forming
the Pompey complex and smaller more dispersed reefs
forming the Swains complex. Area E (Figure 3) occurs
to the south of the Capricorn Channel (22 and 24 S)
where the shelf is less than 100 km wide, with the Capri-
corn and Bunker reefs occupying only a narrow zone on
the mid-shelf. Well-known reefs in the region include
Heron, Wreck, One Tree, Fitzroy, Fairfax, and Lady
Musgrave.
Small drowned reefs occupy the shelf edge in
80–100 m of water (Davies et al., 2004). In addition to
the reef ecosystem, Halimeda bioherms and biostromes
comprise an almost equally important ecosystem in
the northern and central, southern Great Barrier Reef
(See Halimeda Bioherms, this volume).
The rift basins
In addition to the platforms, the architecture of the north-
east Australian margin is dominated by a series of rift
basins (Figure 1a and b) which formed as a result of frag-
mentation of a northeastern extension of the Tasman fold
belt (Gardner, 1979; Ewing et al., 1970; Falvey, 1972;
Falvey and Taylor, 1974; Taylor, 1975; Mutter, 1977;
Taylor and Falvey, 1977; Mutter and Karner, 1980). The
Pandora and Bligh Troughs and the Osprey Embayment
occur in the north; the Queensland and Townsville
Troughs in the central region and the Cato trough occupies
the eastern and southern sections of the GBR. Relatively,
smaller rift basins are also known to underlie the shelf of
the Great Barrier Reef – the Papuan platform, the Halifax
basin, the Whitsunday basin, and the Capricorn basin
(Figure 1a and b). In the north the Papuan platform is
a thick (? 4–5 km) Jurassic to Recent basin containing
reefs within a fluvio-deltaic sequence representing the vast
outpourings from the Fly River in New Guinea. The
poorly defined Halifax Basin is a major structure beneath
the continental shelf to the east of Townsville and under-
lies the central outer GBR: it is a Cretaceous/Tertiary basin
covering some 60,000 km2 and a maximum thickness in
excess of 4,000 m. It lies at the intersection of the Queens-
land and Townsville Troughs. The Whitsunday Basin is
a very shallow structure containing little more than
1,000–1,500 m of Cretaceous/Tertiary sediments underly-
ing the region to the west of the Pompey and Swains reefs
and may represent a connection between the Halifax and
Capricorn basins. In the south, the Capricorn basin is
sandwiched between the Bunker and Swains Highs.
Bunker and Swains Highs
The Queensland and Townsville Troughs lie adjacent to
the central/ northern reef region. Directly adjacent to the
Great Barrier Reef (Figures 1a, b and 4a, b), the Queens-
land Trough occupies the region between the continental
shelf and the Queensland Plateau between 14 S and
17 30S. The floor of the trough slopes northward from
1,000 m at the junction with the Townsville Trough to
around 3,000 m at its junction with the Osprey Embay-
ment. Rather than being a simple graben, the Queensland
trough may have been initiated in an oblique wrench zone
caused by left-lateral movements between Australia and
the Queensland Plateau (Symonds et al., 1984). The
Townsville Trough is a west to east trough separating the
Queensland and Marion Plateaux. It is over 500 km in
length and 150 km wide. At its confluence with the
Queensland Trough, the Townsville Trough is 1,000 m
deep, deepening eastward to 3,000 m at its exit into the
Coral Sea Basin. At this eastern end, at about
Longitude 154 E, a bifurcation sends one branch south
into the Cato Trough and the other winding sinuously
north into the Coral Sea Basin. The Capricorn Basin
started in the late Cretaceous as the result of an eastward
shift of the Maryborough basin rift axis to the east con-
comitant with spreading in the Cato Trough. The basin
extends in a northwest to southeast direction and is
500 km long by 250 km wide and is bordered by the Bun-
ker and Swains Highs. This early to late Cretaceous rift
phase development preceded continental breakup and the
formation of the Coral Sea Basin by sea floor spreading
in the Paleocene (62–56 million year BP) or latest Creta-
ceous (Weissel and Watts, 1979; Symonds et al., 1984).
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 511

W
0.00
Site 821 Schematic section–
Site 820 Queensland trough transect
0.55
Site 819 0 30 km V = 36
H Site
1.10 811/8
Site 822 Upper middle Site 824
miocene
Two-way traveltime (s)

1.65
Pliocene
Site 823 Upper middle
2.20 miocene

2.75
Eocene/oligocene

3.30

3.85 Eocene/oligocene
Eocene/oligocene
4.40

4.95
WSW Queensland trough transect ENE
Site 821 Site 820 Site 819 Site 822 Site 823 Site 824 Site 825 Site 811
16 38.793’S 16 38.220’S 16 37.439’S 16 25.379’S 16 36.981’S Site 823 16 26.703’S 16 30.955’S 16 30.956’S
146 17.366’E 146 18.224’E 146 19.486’E 149 12.904’E 149 36.045’E (continued) 147 45.737’E 148 9.457’E 148 9.447’E
0 700

lwr. Upper Pliocene Pleist.

Pleistocene
Pleistocene

Plioc.
wash
Upper miocene

100 800

Upper miocene
Pliocene
Pleistocene
Pleistocene

Pleistocene
Pleistocene

Upper miocene

200
Upper pliocene

900

Middle miocene

300 1000
Lower miocene
Mioc.
mid.
Depth (mbsf)

Upper pliocene

?
-lower mioc.

?Upper oligocene
upper olig.

400
?
?
Periplatform deposits
?
lower pliocene

Pelagic ooze/chalk
Sediment gravity-flow deposits
500
Dolomitic sediment
Dolomite
Unrecovered interval
Skeletal deposits
(packstone, grainstone, floatstone)
Reef deposits
(boundstone, framestone, grainstone)
600 Clay/claystone
(terrigenous silt plus clay)
Mixed pelagic/hemipelagic deposits
Basement

Unconformity

a 700

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 4 (Continued)
512 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

0.0 NNW SSE NNE SSW

Schematic section- Marion plateau
0.55 Sites 812, 814, 813 (equivalent) townsville trough trasect Sites 816, 826
Site 815
Site 818 (equivalent)
0 30 km V
– = 32 P
H
1.10 Site 817 Townsville trough
Platform
Two-way traveltime (s)
1.65 “debris”
Upper middle miocene facies
2.20 Lower Miocene

2.75 Eocene/Oligocene Paleocene

3.30 Basement/P2 Paleocene

metasediments
Syn-rift Syn-rift
3.85 section section Basement/P2
metasediments
4.40 ?
?
4.95
P = Carbonate platform
5.50

Townsville trough transect

NNW SSE
Site 812 Site 814 Site 814 Site 818 Site 817 Site 815 Site 816 Site 826
17 48.842’S 17 49.985’S 17 49.985’S 18 3.767’S 18 9.493’S 19 9.034’S 19 11.915’S 19 13.530’S
149 36.317’E 149 30.831’E 149 30.831’E 150 2.533’E 149 45.510’E 149 59.516’E 150 0.606’E 150 0.597’E
0
upper Mioc.-Pliocene Pleist.

upper Pleist.

Pleist.
Pleist.
Pleist.

Pleist.

Pleistocene

upper Pliocene

Plioc.
Pleistocene
Pliocene

lower Pliocene

wash
Pliocene
upper
Mioc.

upper
Plioc.
upper Pliocene
100
upper Miocene
middle Miocene

upper Pliocene

middle Miocene
middle Miocene
lwr. Plioc.

lower Pliocene
middle Miocene (?)

middle Miocene

upper
Mioc.

200
lwr. Plioc.

300
upper
Mioc.
Depth (mbst)

uppermost lower to middle Miocene

Periplatform deposits

Pelagic ooze/chalk
upper Miocene

400
Sediment gravity-flow deposits
Dolomitic sediment
mid. Mioc.

Dolomite
lower to

Unrecovered interval
Skeletal deposits
(packstone, grainstone, floatstone)
500 Reef deposits
(boundstone, framestone, grainstone)
Clay/claystone
(terrigenous silt plus clay)
Mixed pelagic/hemipelagic deposits

Basement

Unconformity
600

b 700

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 4 (a) Cross section of the Queensland trough and the
stratigraphy of ODP drill holes across the trough; (b) Cross section of the Townsville trough and stratigraphy of ODP drill holes across
the trough.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
A major outcome was the development of a series of
interconnected three-branch rift systems with the Towns-
ville and Queensland Troughs and the Osprey Embayment
representing failed arms of such a system (Mutter and
Karner, 1980). These troughs are the most important struc-
tural elements associated with the formation of the Great
Barrier Reef Province.
Causes of architecture
The causes of the architecture described above are rifting,
drifting, subsidence, sea-level/climate change, and colli-
sion. These are briefly summarized as follows:
Rifting
Late Cretaceous extension formed the basins described
above, all of which are rift basins proximal to carbonate
platforms. Of particular importance is the Queensland-
Townsville-Cato rift systems, but also of importance if
less developed is the rift system extending south from
the Queensland trough through the Whitsunday basin
to the Capricorn basin separating the Queensland shelf
from the Marion and Queensland plateaux. The rifting
process has clearly influenced the form of platform devel-
opment in both, a general way through influencing the
shape of the platforms and providing a substrate for shal-
low water deposition, but also in a specific way through
the influence of fault blocks as sites for early reef develop-
ment (Figures 2 and 5a, b).
Subsidence
The effect of subsidence is mainly to produce space. It can
also have the effect of termination. Quantitative subsi-
dence data for northeast Australia in four wells, Anchor
Cay 1, DSDP Site 209, Capricorn 1A, and Aquarius 1
are shown in Figure 6. This data indicates that northeast
Australia has not subsided solely as a result of uniform
post-rift thermal cooling; rapid subsidence pulses have
occurred at different times, and such pulses have clearly
initiated transgressions. In the northern region at Anchor
Cay, subsidence increased to 50 m/million years in the
Miocene and increased again to 149 m/million years in
the Pliocene. On the Queensland Plateau, at ODP site
209 subsidences doubled after the middle Miocene. In
the Capricorn Basin, major subsidence of 75 m/million
years occurred in the middle Oligocene and again in the
Pliocene with uplift in the intervening early Pliocene. In
the north and south, subsidence pulses occurred in the
Miocene and the Pliocene. Seismic data from the central
to northern shelf (Townville to north of Cairns) shows
a mid-shelf hinge of subsidence, delineating the point
from which sediment packages thicken eastward. The
northern shelf north of Cairns lies dominantly west of this
hinge line; in the central region, most reef development
occurs adjacent to the hinge line whereas southwards the
Pompey and Swain reefs occur well to the east of the hinge
line. There is fundamental control of reef growth by base-
ment subsidence.
513
Drift
Hotspot (Duncan, 1981; Wellman, 1983) and magnetos-
tratigraphic studies (Idnurm, 1985, 1986) define the basis
for reconstruction of Indian-Australian plate motion
through the Cenozoic (Figure 7a). Since the late Eocene
when northeast Australia was located between 29 and
44 S, the continent has moved almost directly northward
to its present position between 9 and 24 S. This move-
ment would have resulted in profound climatic changes
along the northeast Australian margin. Simplistically
(Figure 7), the northern tip of northeast Australia would
have entered the tropics only 25 Ma ago whilst the south-
ern part of the Great Barrier Reef would have entered the
tropics only in the last 1 Ma with substantial impact on
the likely sedimentary facies that could have formed.
However, a more detailed analysis of the effects of plate
motions on paleo-oceanography (see Davies et al., 1989,
p. 65) gives a much more precise analysis (Figure 7b) from
which the following conclusions are drawn:
1. In the earliest Eocene, temperatures were very briefly
warm enough for coral reef growth. Chaproniere
(1984) reported large foraminifera from a dredge haul
off the northwest Queensland Plateau, which he con-
cluded grew in waters with temperatures of 18–27 C.
2. Sea surface temperatures from the middle Eocene to
the middle Early Miocene (44–19 Ma) over the whole
northeast Australian margin were temperate.
3. During the early Miocene sea surface temperatures off
northeast Australia were marginal for coral reef devel-
opment. In modern parlance, they were subtropical,
comparable to southern Queensland today.
4. A rapid rise in sea surface temperatures occurred in the
early mid-Miocene. Conditions were warm enough for
coral reef growth in the northern and central GBR.
5. During the upper Miocene, tropical conditions persisted
in the north and center but warming slowed in the south
and conditions were not conducive for coral reef
growth. Subtropical conditions prevailed in the south.
6. During the Pliocene and Pleistocene, the entire north-
east Australian margin was situated within the tropics.
The consequences of these apparently simple conclusions
are substantial and fundamental for northeast Australia:
a. Any tropical platform must be underlain by a temperate
platform and that initial rifting of the margin will con-
tain temperate facies sediments.
b. The consequence of the drifting model of sediment
development is that reefs grew first in the north and,
progressively, later in the south depending on SST.
c. Tropical facies development will thin southward and
overlie a progressively southerly thickening subtropi-
cal to temperate section. Facies diachroneity is, there-
fore, a fundamental factor of a stratigraphy dominated
by plate motion through climatic belts.
The conclusions can be tested by comparing present day
facies distributions along the east Australian margin with
514 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

W
Borabi reef trend E W
Pasca reef E
0 0

1
Reflection time (s)

Reflection time (s)

2
2

3
4
4
0 Km 5
a b 0 2 km

NNW Pandora reef prospect SSE Northern Marion plateau SSE

0 410 m
Reflection time (s)

3
0 Km 10
c
1.5
Reflection time (s)

W Coringa bank E
0

490 m
Reflection time (s)

2.0
1

d
2
0 10 km 23/OQ/139

SW Western queensland plateau

0 NE

1160 m
Reflection time (s)

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 5 Seismic sections across (a) Borabi, Pasca, Eastern Fields
Plateau and Pandora Trough and (b) northern margin of the Marion Plateau showing the plateau edge reefs. Both sections show the
importance of faulted margins as the sites for large reef systems. (c) western parts of the Queensland Plateau.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 515

Time (Ma) Time (Ma)

Cret P Eocene Olig Miocene P Cret Paleoc Eocene Oligocene Miocene P
O O
70 30 10
70 50 30 10
Sea surface
0
0

Sea bottom

Depth (km)
1

Depth (km)
2
Anchor cay 1 DSDP 209
torres shelf Queensland plateau 2

3
a b

Time (Ma)
Cretaceous Paleoc Eocene Oligocene Miocene P
O
90 70 50 30 10

1
Aquarius 1
Depth (km)

marion plateau

3
c

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 6 Subsidence history for drill holes at Anchor cay, DSDP
Site 209, Aquarius and Anchor Cay.
516 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

0° South of latitude 28S – branching bryozoan/foram/

molluscan facies.
10°S A similar facies sequence occurs vertically in the Borabi
Tropical
No. 1 drill hole in the Gulf of Papua (Figure 8a). The bot-
No tom to top facies variations mirror those on the present
rt he
20°S rn
gre
a
shelf from south to north, and including the change to fore-
tb
ar r
ier Subtropical land basin clastics in the Gulf of Papua. The effects of
Latitude

ree drift, therefore, are as shown in Figure 9.

30°S f/gu
lf of
papu
a
So Sea-level variations
uth
ern
40°S gre Operating relatively in conjunction with subsidence, the
a tb
arr effects of sea-level variations on platform development
ier
Temperate ree
50°S
f on the northeast Australian margin are different on the
continental shelf and the marginal plateaux. On the shelf,
high sea levels are carbonate-dominated and low sea
60°S Q Plio Miocene Oligocene Eocene Paleocene levels are terrigenous-dominated on shelf and slope, and
0 10 20 30 40 50 60 unconformities are widespread. However, on the oceanic
a Time (Ma) plateaus where no terrigenous source occurs, high stands
30° are reef or periplatform dominated and lowstands are
Northern
dominated by unconformity development and caliche for-
great barrier reef mation on the platform and carbonate gravity deposition
25° Tropical
and peri-platform deposition on the steep slopes.
Surface water temperature (°C)

20°
15°
Phosphate
‘spike’
10°
5° Q Plio Miocene
0 10
b Time (Ma)
Great Barrier Reef: Origin, Evolution, and Modern
Development, Figure 7 (a) Projected latitudinal movement of
northeast Australia throughout the Cenozoic. The northern
boundary corresponds to Anchor Cay (presently 9 300 S) and the
southern boundary to Heron Island (presently 24 S). (b) Surface
water temperature envelope for the northeast Australian region
throughout the Cenozoic, showing periods when temperatures
were suitable for reef growth. The Miocene “phosphate” spike
which inhibited reef growth is also shown (see Riggs, 1984).
the vertical facies seen in drill core from the Gulf of Papua.
The present day sediment distribution on the outer conti-
nental shelf shows three major facies (Marshall and
Davies, 1978):
North of Latitude 24S-tropical carbonate and clastic sedi-
ments dominated by corals and Halimeda.
Latitude 24–28S – subtropical coralline algal coated
grains (rhodoliths), encrusting foramifera, bryozoa.
Collision
If, as a result of plate motion, a passive margin moves into
Subtropical a compressional tectonic regime, the carbonate platforms
developed on that margin will be profoundly affected.
They will become embroiled in mountain building and
possibly subduction. These effects are seen on the northern
Southern edge of the Australian craton, which collided with an island
great barrier reef
arc complex in the Oligocene. The sequential development
of the platform during collision is shown in Figure 10. This
convergent history, leading to foreland basin development
Temperate and mountain building, paradoxically, first led to an expan-
Oligocene Eocene Paleocene sion of the area of carbonate deposition, but was followed
20 30 40 50 60 by contraction, demise, and burial of the carbonates by
clastic detritus (Pigram and Davies, 1987; Davies et al.,
1989). This may be the fate of the whole of the GBR if
Australia’s northern motion is not arrested.
Establishment of reefs in northeast Australia
As a consequence of the processes and architecture
described above, coral reefs have been established on
three occasions (1) in the early to late mid-Miocene
(2) in the uppermost Miocene, and (3) in the Pleistocene.
Reefs first established on the Papuan shelf at Borabi
and Pasca. Seismic (Figure 5) and drill hole data
(Figure 8) show early to mid-Miocene reefs at Borabi
established on pre-reefal red algal banks and also on
a basement high at Pasca. On the Queensland Plateau
(Figures 4 and 5b) reefs established in the Middle Miocene,
once again on red algal banks (Brachert et al., 1993).
On the Marion Plateau (Pigram et al., 1992) an extensive
reef system covered over 400 km of the plateau and was
established and reestablished on the plateau and along
its northern edge (Figures 4 and 5). These reef systems
terminated at different times and for different reasons.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 517

Gulf of Papua Great Barrier Reef

Borabi-1 Pasca A1 Pasca C1 Anchor cay 1 Heron Wreck Michaelmas

0 0
Holocene
Pliocene
Pliocene to
to recent
recent
Pleistocene
1000
Late
Miocene 100

Depth (m)
miocene

Middle
Depth (m)

miocene Eocene
2000
Early
miocene Pliocene TD 183m
200
Eocene TD 2953m Mesozoic
mesozoic
3000 TD 2878m TD 223m

Reef limestone
TD 575m
Reef limestone TD 3623m Foraminiferal
limestone
4000 Cor/Bryo/foram
limestone Calcarenite

Shale TD 4267.5m
Quartz sand and sandstone

Sandstone Calcareous quartz sandstone

b
Echinoderm limestone

a Conglomerate and sandstone

Capricorn Basin
0 Aquarius 1
Capricorn 1A
Quaternary
Queensland Plateau
DSDP 209 Late
0 Pleistocene Pliocene
Early
Middle-late Late
pliocene miocene
100 Miocene Middle
Late oligocene miocene
Depth (m)

200 Late Early

1000
eocene miocene
Depth (m)

300 Middle
eocene
Late
TD 344m
oligocene
400 Limestone
Calcareous ooze
Marl
TD 1710m
Nodular chert Claystone
2000
Terrigenous sand Sandstone
c Early
Conglomerate tertiary?

Volcanics

2600
d TD 2658m 23/OQ/136

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 8 Details of drill holes on the northeast Australian margin.
518 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

Northern great barrier reef

0

Time (seconds)
Seismic insert

a 4

Reef

Tropical

North-central great barrier reef Temperate

0

Time (seconds)
Basemet

b 4
Queensland plateau
0

Time (seconds)
Central great barrier reef
0
Time (seconds)

e 3

140 145 150

c 4
Papua New Guinea

Gulf of
Papua

Ashmore-boot
10
-Portlock reefs

3000
Southern great barrier reef 4000
0
Time (seconds)

Coral Sea

200
0
Queensland

d 3 Cairns B Plateau

E
Queensland 1000
Ba 1000
Townsville rri
er
20 Marion
C 0 plateau 20
0 300 km
Re

3000
ef

Australia D
Gladstone

23/OQ/116

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 9 Aspects of the structural history and sedimentologic
framework of northeast Australia illustrated by schematic sections across the north, central and southern regions. An interpretative
north to south tie shows the thinning and thickening of tropical and temperate sediments as a consequence of drift.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 519

Eocene – 40 Ma Late miocene – 10 Ma

Abyssal plain
24
9

29 14

Abyssal plain
SW NE

Depth (km)
0 0

6 6
a c
Early miocene – 20 Ma
Pliocene – 3 Ma

7
14

12
19
Borabi trend

0
Northern great barrier reef
Depth (km)

6 0

4
b d
23/09/126

Shelf Clastic sediments

Reef Carbonate
Emergent
sediments

Slope 0 250 km

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 10 Cenozoic palaeo-oceanography of the Gulf of Papua/
northern Great Barrier Reef showing the effects of progressive foreland basin development on carbonate platforms during
(a) Eocene, (b) Early Miocene, (c) Late Miocene and (d) Pliocene. The schematic sections are orientated approximately
northeast-southwest. Approximate palaeolatitudes are shown for each section.

In the north, reef growth at Borabi and Pasca through
development of the foreland basin, and the southward
flooding of fluvio-clastics caused the eventual migration
of reefs eastward. On the Queensland Plateau, reefs termi-
nated at 13.7 Ma due to subsidence of the plateau. On the
Marion Plateau, reef growth terminated at 10.7 Ma as
a consequence of a major fall in sea level in excess of
a 150 m (Pigram et al., 1992).
Reefs reestablished in the north on Anchor Cay and
Pandora in the upper Miocene and Pliocene, but not on
the Queensland and Marion Plateaux for two reasons.
First, an upper Miocene (7 Ma) subsidence pulse affected
both plateaux (Figure 4) and secondly, oceanographic
cooling established over the region at 6 Ma (Isern et al.,
1993) inhibiting reef establishment, probably related to
the Messinian glacio-eustatic sea-level fall. Reefs never
520 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

again established on the Marion plateau except at two Barrier Reef – did not establish at this time; it occurred
places, Marion reef on the extreme northeastern edge much later.
and Suarez reef in the extreme southeast. This likely
occurred in the Pliocene as it did on the Queensland Pla-
teau (Figure 5c). This phase of growth substantially Establishment of the Great Barrier Reef
contracted on the Queensland Plateau by a third phase of The Great Barrier Reef, as we know it, started in the late
subsidence at 3.7 Ma, drowning much of the plateau and Pleistocene, much later than had been previously thought,
in the north probably initiating further eastern migration and very much later than when the main architecture of the
of the reefs at Pandora. This pulse probably also shelf had been established. The evidence for this is in the
established the main shape of the Australian continental drill holes from Ribbon 5 and ODP 820 (Figure 11a).
shelf. However, the third phase of reef growth – the Great These data are summarized as follows

Distance (Km)
0 1 2 3 4 5 6 7

a Ribbon 5
0
R1
M
R2 25 m
L
40 R3 M 50 m
L
R4 M 75 m
80
L
R5 ML
R6
M
Rd1 L
120 Rd2 Mb
Mb
Caves
R7 M at 120 m
L
R8 M
L
160 Rd3
Depth (M) below sea floor

Mb Caves at
Mb
Mb
Rd4 L 160 m
Mb
Grainstone/Packstone

200 L

Mb
L

Mb
240

280 0 6C 0+2C
Coral Ass
Lithology

Reef unit

0 A
R1 Hiatus
320 275ka
R2
B2 50
R3
Depth(M)
mbsf

R4 B2
360
B1
MUD Simplified lithology
100
In-situ coral
50 R5
framework
465ka
400 MUD
R6 Mud and B2
MUD coral rubble (Fac
Mud, clay with
MUD
minor sand (Faci 150
440 80 C

Boulder reef
b 480 c ODP 820

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 11 Stratigraphy defined for (a) Ribbon Reef 5. Ribbon 5
data also shows the reef front morphology and its correlation with ODP 820 (b) Boulder Reef, and (c) ODP Site 820.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
Ribbon Reef No. 5
Drilled in 1995, the detailed description of the corals, sed-
iments (Webster and Davies, 2003) and coralline algae
(Braga and Aguirre, 1997) in a core from Ribbon Reef 5
define the basis for understanding the establishment of
the Great Barrier Reef. The litho-logs (Figure 11a) divide
into three parts:
0–96 m – Six reef horizons comprised of two coral assem-
blages (1) robust branching Acropora (humilis, robusta,
palifera, Stylophora (pistillata) and Pocillopora
(verruclosa, damicornis) with associated massive
Goniastrea and Platygyra and (2) Massive Porites (lutea,
solida) with associated encrusting Porites and Montipora.
Boundaries between reef horizons are identified by com-
binations of prominent horizons of pale brown sediment
containing areas of calcitized plant cells, radiometric dat-
ing, negative shifts in 018 values, exposure horizons, color
changes from pink to brown, changes in algal associations
and shifts in sediment facies (Webster and Davies, 2003).
Algal associations throughout reflect shallow water tropi-
cal environments.
96–158 m – A rhodolith sequence dominated by four
rhodolith grainstone horizons, coralline algal grainstones/
packstones and two in situ coral framework horizons.
The two in situ coral framework horizons represent the
oldest reef horizons encountered in the Ribbon 5 core
and separate the rhodolith horizon as into an upper and
lower section. Changes from coral to rhodolith-dominated
horizons are reflected in changes in the coralline algal
associations and in one instance at 130 m by a sharp irreg-
ular surface and a substantial shift in the 018 record. The
algal associations define lower and upper subtropical envi-
ronments with tropical reef horizons sandwiched between,
i.e., the earliest reef growth occured on a substrate of sub-
tropical rhodolith grainstones.
158–210 m – 52 m of alternating grainstones/packstones
and packstones/wachestones. The former are comprised
of poorly sorted (rudaceous (4 mm) to medium sand)
fragmentary corals, encrusting and branching corallines,
bryozoans, foraminifers (Amphistegina, Iterostegina,
encrusting forams), molluscs, serpulids, echinoids, and
gastropods: fresh water diagenesis throughout is indicated
by interparticle spaces infilled by thick calcite rims,
druses, and syntaxial overgrowths while mouldic porosity
is developed within molluscs, corals, and especially
Halimeda. The latter are well sorted, sometimes graded
but finer grained (medium to coarse sand) with similar clasts
but with up to 50% lime mud dispersed unevenly; little fresh
water diagenesis evident. Boundaries between the
grainstones/packstones and packstones/wachestones are
sharp. Corals are subordinate to coralline algae, of which
there are two assemblages, a shallower water Lithophyllum
assemblage and a deeper water Mesophyllum assemblage;
both occur today on the warm temperate to subtropical
mid- to outer shelf of southern Queensland, south of the
Great Barrier Reef (Braga and Aguirre, 1997). The sedi-
ments are typical of the same region. The wachestones/
521
packstones are defined as outer shelf (þ60 m water depth)
highstand deposits while the grainstones are defined as
mid- to inner shelf regressive or lowstand deposits. The rel-
atively sharp boundary between the facies types is, there-
fore, “event driven” by oscillating sea levels. The fresh
water diagenesis is due to exposure in the lowstand
effecting the grainstones but not the highstand
wachestones/packstones. They are unlikely to represent
the debris flows and turbidites postulated by Braithwaite
et al. (2004).
Ocean drilling program site 820
Drilled in 1991, Site 820 forms part of a suite of holes
(819, 820, and 821) drilled immediately in front of the
Great Barrier Reef in Grafton Passage, east of Cairns
(Davies et al., 1991). While Site 821 is considered the
most complete sedimentological section, it unfortunately
suffers from intense fresh water diagenesis and Site 819
has a major hiatus in the upper part of the section. Site
820 is, therefore, considered to be by far the most com-
plete section and the basis for the interpretations shown
in Figure 11c (Davies and Peerderman, 1998). Three sed-
imentologic units are recognized:
Unit A 0–65 m composed of wachestones separated by
8 m thick sandy bioclastic packstones composed of plank-
tonic and benthic forams, mollusks, pteropods, minor
echinoids, corals, corallines, bryozoans, rhodoliths and
Halimeda. High and low sedimentation rates reflect alter-
nating high and low sea-level oscillations. Seismic imag-
ing shows aggradation with clear erosional geometries
reflecting similar sea-level oscillations.
Unit B 65–145 m composed of wachestones and thin
packstone units overlying wachestones and thick
packstone units. A major reduction in sedimentation rates
coincides with the boundary between Units AþB while
alternating high and low rates in Unit B are thought to
reflect oscillating sea levels. Seismic imaging shows the
upper wachestones and thin packstones to be aggrading
packages and the lower wachestones and thick packstones
as prograding packages, the boundary occurring at around
110 m in the core.
Unit C – between 145 and 220 m dominated by sandy
packstone beds and minor wachestones. An unconformity
may occur in this sequence while seismic imaging defines
prograding sediment geometries.
In all of the above, there are good agreements between
stratigraphic, sedimentologic, and seismic data. However,
it is when this data is tied to the isotope data (Figure 11c)
that it starts to make sense in terms of the origin of the
Great Barrier Reef. Three principal isotopic sequences
are recognized:
(1) Between 0 and 75 m (Unit A above) the signal is dom-
inated by low frequency, high amplitude variation
defined by Peerdeman et al. (1993) as related to the
orbital eccentricity frequency. This isotopic sequence
extends from isotope stage 1–8 and, therefore, has
522 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
a base at around 300,000 years. Kroon and Alexander
(1993) interpret this signal to indicate a progressive
decoupling of surface and bottom water temperatures
and raised surface water temperatures.
(2) Between 75 and 145 m (Unit B above), the signal is
characterized by a much higher frequency as well as
high amplitude. Peerderman et al. (1993) interpret this
as dominated by the obliquity signal and extending
from isotope stage 9–19 at the Brunhes/Matayama
boundary. Davies and Peerdeman (1998) divided the
section into two parts (2–1) between 75 and 110 m
(isotope stages 9–13) characterized by low frequency
high amplitude and (2–2) between 110 and 143 m
(isotope stages 13–19) characterized by a much
higher frequency and also a high-amplitude signal.
These correspond with the division in packstones
thicknesses noted above.
(3) Betweeen 145 and 220 m (Unit C above), the signal is
characterized again by a much lower frequency(per-
haps dominated by the eccentricity signal) and has
high amplitude and as such is similar to the section
above 75 m.
Peerdeman et al. (1993) consider the boundary at 75 m as
a critical event in the history of the margin. Using the dif-
ference between the planktonic O18 signal in the Site 820
core and the benthic O18 signal at the deep sea Site 677
(Shackleton and Hall, 1989), they defined sea surface
temperature variation for the northeast Australian margin
which showed a substantial rise in temperature of
around 3–5 C at around 75 m in the core (around isotope
stage 9). This temperature variation corresponds to a
change in frequency and amplitude of the isotope signal
arising from the change from obliquity to precessional
orbital cyclicity. Kroon and Alexander (1993) recognized
a similar temperature change at Site 819. Further, Jansen
et al. (1986) also recognized a fundamental change in iso-
tope frequency at the 9–8 boundary as defining a change
from lower to higher sea surface temperatures leading to
the establishment of more interglacial conditions in the
period of isotope stages 1–8. At sites 820 and 819, the
boundary at 65 m is, therefore, a globally driven change
dated at around 300,000 years. Further, the changes defined
at the B1/B2 boundary, particularly the change from
prograding to aggrading geometries (dated as just prior to
465 ka) reflects a change from shallow to deeper water, coin-
cident with major regional oceanographic reorganization in
the Southern Ocean (Kuijpers, 1989; Nelson et al., 1988).
The boundary may also mark the change from an
obliquity signal to the eccentricity signal seen in Unit A,
with Unit B1 representing the transitional warming.
Relations between Ribbon 5 core and ODP
site 820 and implications
The proposed relations and reconciliation of the two data
sets are shown in Figure 11a. The profile to the east of Rib-
bon 5 is one proven by profiling and submersible exami-
nation (Phipps et al., 1985). The fundamental change
from subtropical rhodoliths to tropical reef reflects a tem-
perature change and just such a change is seen in the iso-
topic signal at Sites 820 at a depth of 65 m. The reef
section is thus equivalent to Unit A (isotope stages 1–8)
and the rhodolith section is equivalent to Unit B1 (isotope
stages 9–13). Initial reef establishment of the coral frame-
works in the middle of the rhodolith section must, there-
fore, date at sometime after 465 ka while the firm
establishment of the main reef section above 96 m dates
from around 300,000 years. The six stacked reefs at Rib-
bon 5 correspond to sea-level highstand growth phases,
the first being either stages 8 or 9. The rhodolith section,
equivalent to B1, represents the transitional warming
suggested by the change in the obliquity signal noted
above. This would be equivalent to conditions operating
today to the south of the Great Barrier Reef on the south
Queensland shelf (Davies et al., 1991). Further the sea-
level oscillations defined by the wachestone/packstone
alternations at Site 820 and representing warming and
cooling episodes, serve also to explain the reef intercala-
tions within the rhodolith sections. Davies and Peerdeman
(1998) propose that the facies variations on that shelf and
their lateral migration during oscillating sea level, define
a model for the sub-reef rhodolith section at Ribbon 5
and explain the reef frameworks within that section.
Cross-shelf growth of the great barrier reef
Once established in the shelf edge zone, is it right to
assume that reefs quickly spread everywhere on the shelf?
The results of a second borehole at Boulder Reef
(Figure 11b) on the inner shelf and drilled during the Rib-
bon 5 campaign suggest otherwise. Four stacked reef
sections (R1–R4 in Figure 11b) sit on siliciclastic mud
and clay foundations at a depth of 34 m. The siliciclastics
extend to 86 m where drilling terminated; however, coral-
bearing units (R6 and R5) lacking a true framework occur
within this mud section and have been interpreted as low-
diversity coral communities reworked to form a rubble
(Webster and Davies, 2003). Their presence is, however,
significant. The base of the core has been tentatively dated
by ESR as 210 þ 40 ka, which suggests that the begin-
ning of reef growth on the inner shelf was much later than
on the outer shelf. Two failed attempts (R5 þ R6,
Figure 11b) were followed by the establishment (R4) of
a reef structure, after which Boulder Reef was reestablished
on three later occasions. The two failed attempts at 55
m (R5) and 63 m (R6) and the beginning of the main reef
section (R4 at 34 m) occurred on siliciclastic foundations,
testifying to the fact that the inner shelf was a turbid envi-
ronment during highstands, perhaps more turbid than
today. It is concluded, therefore, that the pre-reef environ-
ment was totally different on the inner shelf compared with
the outer shelf and that reef initiation was later on the inner
shelf compared with the outer. In both cases, however, reef
growth was a relatively recent phenomenon.
The drill core at Ribbon 5 and Boulder Reefs testify to
the youth of the Great Barrier Reef and to the fact that reefs
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
have been destroyed and regrown again on at least six
occasions on the outer shelf (Figure 11a, R1–R6) and four
occasions on the inner shelf (Figure 11b, R1–R4).
Coupled with the isotopic data from the ODP cores, it is
concluded that sea-level oscillations are the cause of the
repeated demise and subsequent reestablishment. Further,
the isotopic data also indicate that the initial establishment
of the reefs on the outer shelf was climate controlled, and
occurred on a rhodolith-dominated bank on the outer
shelf. Such subtropical red algal banks may have formed
in the way such banks occur on the south Queensland shelf
today.
The modern reef
The history of growth and development of the modern reef
has been defined largely by drilling through the reefs into
their Pleistocene substrate. Hopley et al. (2007) report 160
drill holes in 50 reefs, and 750 dates. These data indicate
that the amount of Holocene reef growth varies from 0 to
28 m (Davies and Hopley, 1983; Hopley et al., 2007).
Transects across the northern reefs east of Cooktown and
off Townsville indicate little difference in the thickness
of reef from west to east (Davies et al., 1985). Longitudi-
nally, however, down the length of the Great Barrier
Reef, Hopley et al. (2007) report an increase in thickness
from Torres Strait to south of Cairns and then a gradual
decrease in thickness to the south. They, therefore, imply
a change in the depth of the Pleistocene substrate from
north to south and invoke either long or short-term subsi-
dence to explain it. An additional process, hydroisostasy,
has also been at work to ultimately define systematic
variations in thickness (see Glacio-Hydro Isostasy, this
volume).
The composition of the Pleistocene substrate has been
shown to be reef framework on the mid- and outer shelf,
similar or identical to the modern reef. The Pleistocene
corals are, however, sometimes accompanied by thick cor-
alline algal crusts and Halimeda grainstones as seen at
One Tree Reef and Ribbon Reef 5 (Marshall, 1983;
Webster and Davies, 2003). On fringing reefs, the Pleisto-
cene is sometimes composed of mud, sand, and gravel
(Kleypas and Hopley, 1993). Soil horizons have been
reported from Fairfax and Fitzroy reefs in the southern
Great Barrier Reef, from Redbill, Hayman, and Davies
Reefs in the central region and from Ribbon Reef 5 in
the northern region. Soil development implies erosion of
the underlying limestone and it is surprising that so few
have been reported considering the number of holes
drilled. As the age of the Pleistocene substrate has been
established in the south at One Tree Reef (Davies and
Marshall, 1980; Marshall, 1983) and in the north at Rib-
bon Reef 5 (International Consortium for Great Barrier
Reef Drilling, 2001) as isotope stage 5 (around 125 ka),
then sea level is likely to have been þ5 m above present
sea level. Assuming the stage 5 reef grew to sea level, ero-
sion of up to 33 m must have occurred on reefs in the
region (assuming no subsidence). Allowing for
523
subsidence of 10 m (rapid) in the 100 ka period since
exposure of the 125 ka reef, then erosion of up to 23
m will have occurred. The Pleistocene surface on which
the Holocene reef has grown is, thus, a much modified
(some would say karstified) earlier reef.
Holocene reef start-up
Holocene sea level in the Great Barrier Reef transgressed
the foundations of most Pleistocene platforms in the time
frame 7–10,000 years BP (Hopley et al., 2007). Start-up
growth occurred after transgression was complete, with
17 out of 22 reefs dated starting growth in the time frame
7,100–9,000 years BP. On the outer shelf start-up began at
6.6–8.6 years BP; in the mid-shelf it was around 9.1–6.9
years BP, while on the inner shelf it was around 7.8–6.3
years BP.
In the Great Barrier Reef, sea level changed from
a transgressive to a stillstand condition around 6,500 years
BP. The age at which reefs reached sea level broadly
depends on the depth of their foundations and these ages
are summarized in Hopley et al. (2007). Nearly 60% of
all reefs cored and dated reached sea level in the interval
5,000–6,500 years BP meaning that nearly 60% of the
reefs studied have been subject to a shallow water energy
regime for most of the time that they have been growing.
Growth, therefore, must reflect this.
The interactions between growth and sea level prompted
Davies and coworkers (Davies and Montaggioni, 1985;
Davies et al., 1985) and Neumann and Macintyre (1985)
separately, independently, and in different oceans to define
five strategies of reef growth in the Holocene:
Keep-Up Reef growth – reefs growing and keeping pace
with sea-level change, such that reef flats developed
early. Pleistocene foundations are relatively shallow.
Katch-Up 1 Reef growth – reefs which started to grow
well after inundation but then grew so quickly that they
were able to catch-up before sea level stabilized.
Katch-Up 2 Reef growth – reefs which started to grow
well after inundation but did not grow so fast so that
catch-up occurred after sea level had stabilized.
Katch-Up 3 Reef growth – reefs which did not start to
grow until after sea level had stabilized so that the
growth was into continually shallowing water.
Screwed-Up (Give-Up) Reef Growth – reefs which died
before reaching sea level. Neumann and Macintyre
(1985) considered that these were “drowned reefs,”
but Davies and coworkers considered other factors
could also effect death (turbidity, elevated nutrients,
and subsidence).
More than one growth strategy can occur on the same reef
as local environment factors affect the growth response
(Davies and Montaggioni, 1985; Davies et al., 1985). Fur-
ther, the rates of growth are depth dependent; thus growth
approximates an “S” shaped curve – slow growth to start,
rapid growth as the reef grows into shallow water and slow
growth again as the reef is subject to high energy in shal-
low water. Plots of growth rates against paleo-water
524 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

depths (Davies et al., 1985) show rates of >8 m/ka1 for and Davies (Reef flats this volume). Attention is given
water depths of 12–15 m, 4 m/ka1 for water depths of here, therefore, to the sand sheets which are intertidal and
20 m and corresponding low rates in shallow water. This subtidal. In Figure 13, intertidal sand sheets occur on the
is particularly well seen for the outer shelf reefs and total southern reef flat bordering the lagoon while subtidal sand
reef data set in Figure 12. It is not so well seen for the sheets occur between the rubble banks and the lagoon along
mid-shelf reefs where the shallow water rates of growth the eastern margin. The southern intertidal sand sheet
are not that much less than the mid-water-depth rates but occupies an area of 822  1,000 m2 and contains a sand
this may be due to the reduction in energy on the mid-shelf volume of 14 million tones of carbonate. A section through
reefs compared to the outer shelf reefs. Surprisingly, the sand sheet summarizes its major growth features as
growth rates change little with latitude. seen in vibrocores and drill holes and surface maps show
its essential granulometric variations (Figure 13c). In
Growth at sea level Figure 13b a lower subtidal sand overlies rubble close to
When a reef reaches close to a stabilized sea level, it comes the junction with the coral flat and is in turn overlain by
under the influence of the ambient hydrologic regime. intertidal sands as the body builds up to sea level. The sand
Three features are produced on most reef types (see is derived from the windward margin and the outer edge of
below) – the algal and coral reef flat and the sand sheets: the coral flat from where it is wave transported on every
rubble flats are found on only a few reefs. The reef flats tide toward the lagoon. The sediments of both units com-
and the rubble flats are described in detail by Thornborough prise corals, coralline algae, foraminifera, mollusks, and
minor Halimeda. The subtidal sands are characterized by
fining upward rhythms while the upper intertidal sequences
m ka–1 are characterized by coarsening upward sequences. How-
0
0 1 2 3 4 5 6 7 8 9 10 11 ever, in both units most stratification is destroyed by shrimp
bioturbation. The intertidal sand sheets are prograding into
1 (filling up) the lagoon.
2
The eastern subtidal sands occur in 3–5 m of water
along the eastern junction of the lagoon with the eastern
3 rubble flat (Figure 13a). Surface sediments are very coarse
to fine sands and show a decrease in grain size toward the
4 lagoon; cores show fining upward sequences being
5 destroyed by bioturbation. These subtidal sands occupy
an area of 538,000 m2 and contain approximately 9 million
6 tones of carbonate. They are comprised of uncemented
7
corals, gastropods, foraminifera, corallines, and bivalves
derived through bio and physical erosion of the eastern
8 rubble flats.
The distinctive sea-level signatures on almost all reefs
9 in the Great Barrier Reef are the algal/coral flats and sand
meters

10 sheets. Rubble flats occur on only the high energy reefs.

11
Types of reefs
12
The following reef types have grown in the Great Barrier
13 All GBR data Reef – fringing reefs, mid-shelf reefs (patches, crescentic,
Outer shelf lagoonal platform reefs, and flat-topped planar reefs), and
14 Mid shelf outer shelf reefs (detached reefs, deltaic reefs, ribbon
15 reefs, and giganto-reefs) (Maxwell, 1968; Hopley, 1982).
For completeness, they are briefly described below.
16 (See entries Reef Classification by Hopley (1982); Reef
17
Classification by Maxwell (1968)).
According to Hopley et al. (2007), there are 758 fring-
18 ing reefs in the Great Barrier Reef Marine Park, but
representing less than 2% of the total reef area. Most occur
19
between latitudes 20–22S and most fringing reefs show
20 a weak biologic and morphological zonation. Recent stud-
ies describe fringing reefs in turbid inner shelf environ-
Great Barrier Reef: Origin, Evolution, and Modern ments (Perry and Smithers, 2009; Smithers et al., 2006)
Development, Figure 12 Vertical growth rate data plotted comparable to the environments postulated above for
against palaeo water depths (modified after Hopley et al., 2007). Boulder Reef off Cooktown.
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 525

A B
0
0.01 mm.y.–1 Suspension
Core Load –14.6 kg. d–1
? kg (small)
Sediment m
5 fining
40
Fines 5

Note 1. Past growthy >

2. In vertical profile, growth
?m R
T
3. Facies i fine to coarse
upwards
ii coarse to fine
laterally
4. Fines thicken with time

Gravel Coarse sand

(> 2mm) 1-2 mm

a b
Sand Mud
(0.063-1mm) (<0.063)

5 10 20 50 40 50 60 70 80 90100 0 1 2
c Fraction %
d

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 13 (a) Vertical air photograph of One Tree Reef in the
southern region. (b) Section through the sand sheet summarizing its major features, and (c) granulometric variations across the sand
sheet and lagoon.

Mid-shelf reefs occur throughout the Great Barrier Reef
region although the number, shape, and complexity varies
substantially. On the basis of size and physical attributes,
they are classified as patches, crescentic, lagoonal plat-
forms, and flat-topped planar reefs. Juvenile, mature, and
senile varieties of the crescents, lagoonal, and flat-topped
reefs have been described (Davies, 1983; Hopley et al.,
2007). Critical factors in their evolution are size, depth,
and shape of the Pleistocene substrate; the length of time
they have been at sea level; and the energy in the shallow
water environment.
Outer shelf reefs occur on the shelf edge and are often
linear, and aligned parallel to the edge which can be steep
or stepped. Detached reefs occur in the northern region
eastward of the main reef edge (and therefore detached),
particularly east of the Torres Shelf where the Ashmore-
Boot-Portlock complex occurs Figure 14a. They have
grown on pinnacles of continental crust and are fronted
by very deep water in excess of 1,000 m, while the moat
between the reefs and the shelf edge can be up to 700 m
deep and infilled with prograding siliciclastics from the
Papuan platform (Figure 14a). Deltaic reefs also occur in
the extreme northern region and are made up of short nar-
row reef segments separated by passes behind which dis-
tinctive delta-like patterns have been produced by tidal
currents flooding through the passes and on which reefs
have grown mimicking the delta-like shapes of the under-
lying sand substrate (Figure 14b).
Ribbon reefs occur on the shelf edge over a distance of
700 km between 11S and 17S as a series of narrow discrete
reefs (10–28 km long) separated by passes up to
a kilometer wide. They are numbered sequentially north-
wards. Probably the best known of the Ribbons is
Ribbon 5 (Figure 14c) because there are three drill holes
 526

40
20

60

NW torres shelf SE

80
Portlock
reef

0
10
15B Boot reef
200 F
Torres 600
E
shelf Boot reef 1 T
800

1600
R
Ashmore R

00
00
reef

10
14 1200
Reflection time (s)
2 M
Ashmore slope
100 KM 0 10 km
a b
GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT

c d e

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 14 (a) Seismic sections across the Ashmore – Boot-Portlock reef complex (b) the northern deltaic
reefs (After Hopley et al., 2007), (c) Ribbon Reef 5 and (d) the large reefs of the Pompey Reef complex (After Hopley et al., 2007).
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 527

through the Holocene and the deep hole through the total
thickness of the Great Barrier Reef (Davies et al., 1985;
Webster and Davies, 2003) reported above. The giganto-
reefs of the central southern region (Pompey Reefs) are
indeed large (Figure 14d). Forming an almost continuous
barrier over a shelf edge length of 140 km, many of the
reefs are 50–100 km2 in area, and 15 km long (Hopley,
2006; Hopley et al., 2007). They are surrounded by water
depths of 60 m and oceanwards give way to a terrace
sloping down to a shelf edge at 100 m. The morphology
of this terrace is shown in Figures 14b and 15.
Growth – the facies produced
Five broad carbonate facies are recognized as occurring in
the Holocene reefs of the Great Barrier Reef. Three are
framework facies and two are detrital facies:
1. Coralline algal facies are laminated crusts, often centi-
meters thick, and/or encrustations on branching or
massive corals, composed principally of Mastophoroid
algae either Hydrolithon onkodes or Neogoniolithon
fosliei. Vermetid gastropods and encrusting foraminif-
era are constant faunal associates. The facies is best

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 15 Bathymetric profiles across the submerged shelf east
of Hydrographer’s Passage (From Harris and Davies, 1989).
528 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
developed on windward margins, but encrusting coral-
lines coat most dead or dying surfaces in all intertidal
reef environments.
2. Branching coral facies is comprised essentially of var-
ious species of Acropora, Pocillopora, and branching
Porites. Growth form varies with depth, light, and
energy. In shallow water, variations in energy define
wide differences in species composition and diversity.
Water-depth changes produce similar compositional
variations.
3. Massive (Head) coral facies is comprised mainly of
various species of Porites, Goniopora, Favia, Leptoria,
Platygyra, Symphyllia, Leptastrea, Goniastrea, and
massive Acropora. As with the branching forms, water
depth and energy defines diversity.
4. Sand facies is usually medium to coarse sand compris-
ing coral, corallines, benthic foraminifera, and minor
Halimeda. Occasionally rudaceous-sized fragments are
found. Sorting varies depending on distance from source
and degree of bioturbation. It forms large sand sheets
between coral flats and lagoons and also infills lagoons.
5. Rubble facies is comprised of massive, platey, and
branching corals varying in size from 20 kg blocks to
finger-like sticks. They are poorly sorted, occupying
large intertidal areas, sometimes cemented, often
bioeroded; also found in spurs and grooves, and at
the base of lagoonal patch reefs. They are produced
either by in situ collapse (in lagoons) or by energy-
related breakage and transport.
Two non carbonate facies have also been identified but
are minor compared with the carbonate facies. These are
siliceous muds and clays derived from rivers inputting
onto the inner shelf (Davies and Hughes, 1983; Smithers
and Larcombe, 2003), and granites as at Lizard Island con-
tributing coarse quartz and felspar to an otherwise carbon-
ate environment, but with corals actually growing on the
granites (Webster, 1994).
Growth models
Three growth models are presented below for three dis-
tinctive reef types defined above as lagoonal platforms,
crescent reefs, and ribbon barrier reefs.
The One Tree Reef Model
One Tree Reef (Figure 13) occurs in the Capricorn and
Bunker Reefs of the southern Great Barrier Reef. Surface
sampling and subsurface drilling and coring define One
Tree as the best understood reef in the whole region. Three
dimensional models show the Holocene One Tree Reef
growing atop its Pleistocene foundation and mimicking
its shape. In Figure 16a, the Pleistocene is exposed during
the lowstand. The highest part of the exposed rim is the
algal flat and coral flat because the areas of greatest ero-
sion during the lowstand would likely be the reef front
and the sand flat/lagoon margin (see Antecedent Plat-
forms, this volume). Transgression would initiate growth
on the high rim, after which growth would be to sea level
(Figure 16b). At this point, development of the reef flat,
sand sheets, lagoonal patch reefs, and the rubble sheets
will grow to the stable sea-level position (Figure 13a).
The effect of dominant energy is to produce clear con-
structional and destructional windward/leeward aspects.
Throughout its history different parts of One Tree
exhibited keep-up or catch-up strategies.
The Stanley Reef Model
Stanley Reef (latitude 19.18S) is a crescent reef in its east-
ern part and an “almost-atoll” in the west (Figure 16c). All
work conducted was on the eastern part which is both sim-
ilar and very different to One Tree Reef; similar in having
a distinct edge to the east and southeast and dissimilar in
having a wide open lagoon and a lee edge which is patchy.
The reef is highest around the rim along the eastern margin
where the Pleistocene is at nearly 15–18 m and on the lee-
ward patches at 20–22 m. A lagoonal patch reef in a lee
position showed the Pleistocene at around 20 m. The lagoon
is currently deep (þ20 m), and assuming 5–10 m of sedi-
ment, the Pleistocene in the lagoon would be at a depth of
þ30 m. During low sea level, Stanley was a crescent with
a high eastern rim decreasing in height to the west, and
a large open deep basin on the site of a former lagoon and
open lee end. Holocene growth on this antecedent surface
occurred almost simultaneously on windward and leeward
margins (Table 1). At the time of the first dated growth,
palaeowater depths on the windward margins were
10–12 m and on leeward edges 14–18 m. The first phase
of growth was vertically to sea level, after which, growth
of the windward margin replicates that seen on most reefs
(see One Tree and Ribbon 5) with the development of a
windward reef flat and sand flat. The leeward edge arrived
at sea level later than the windward margin. The next phase
of growth for this crescent reef is for the leeward patches to
coalesce to form a continuous lee edge, at which time, the
crescent reef will become a lagoonal platform reef. At this
stage, most sediment derived from the windward margin
will be trapped in the lagoon leading to an acceleration in
the infill of the lagoon (assuming enough time).
The Ribbon 5 model
Ribbon 5 (Figures 11a and 17) is a shelf edge barrier reef
east of Cooktown for which the database is more complete
than any reef in the Great Barrier Reef. It defines both the
Pleistocene and the Holocene evolution. The model,
defined by data in Figures 11a and 17, has the following
elements:
1. Initiation of reef growth on a shelf edge rhodolith bank
following a major climate change around 300,000–
350,000 years BP.
2. Destruction and regrowth of reefs successively, due to
sea-level oscillations, on eroded antecedent surfaces
with each reef initiating and growing on or close to
the position of the previous reef. A stacked succession
of reefs is thus formed (Figure 17b).
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 529

One Tree lowstand

One Tree highstand

Stanley reef

BMR4

B
JCU1
JCU2
BMR3
A

BMR
BMR2

JCU1 BMR3 BMR4 JCU2 BMR2 BMR1

0
3390±90 6270±130 5600±100
3350±100 5350±80
Channel Lagoon 5870±70 2
5250±100
3350±100 5060±100 6140±170
3950±90 6000±90 4
5260±100 6540±130
6240±90
6290±60 6

4250±90 7020±100
8
5160±120
3500±100 4740±100
3940±110 5580±150 7050±180
3410±120 6060±90 10
4950±110 5940±100
7500±120 12
6320±90
Meters
5040±110 6220±100
14
3530±110
1610±100 Plst 16
8250±180
2770±120
18

7990±110 7010±100 20
Plst
5230±120
22
Plst

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 16 (a) Low stand position shows the pedestal on which
One Tree Reef sits. (b) high stand growth of One Tree Reef on its pedestal and (c) Stanley reef - a mid shelf reef in the central reef
region. The lithologies encountered in drill holes indicate a framework-dominated windward margin and a sand dominated lee
margin. Plots of depth against age show the windward margin getting to sea level first and progressively later backwards.
530 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
Great Barrier Reef: Origin, Evolution, and Modern Develop-
ment, Table 1 Oldest drill dates and depths from Stanley Reef
and calculated sea-level and water depths
Drill hole Oldest age Depth SL position Water depth
BMR 1 7,500 12 m 2m 10 m
BMR 2 8,250 16 m 4m 12 m
BMR 3 7,990 19 m 5m 14 m
JCU 2 7,010 19 m 1m 18 m
3. Growth of the Holocene reef occurred first on the
windward margin (6,500–7,000 BP) on a Pleistocene
substrate at around –15 to 16 m, and progressively later
leewards where the oldest date for growth is around
4,500 BP at a depth of about 14 m. The windward
margin began to develop its reef flat and shed sand lee-
wards after reaching sea level around 6,000 BP. The
leeward margin holes (holes 3 þ 4) are dominated by
sands which postdate the time at which the windward
margin approached sea level.
4. Growth of the Ribbon reefs since sea level stabilized
has been due to (1) catch-up to sea level by patch reefs
that initiated late, and (2) leeward growth of the coral
flat (as described for One Tree in Reef Flat
Thornborough and Davies this volume), and substan-
tial backward shedding of sand into the leeside
environments.
Most reefs in the Holocene, with the possible exception of
fringing reefs, have undergone the same basic processes of
growth from an antecedent platform to sea level and then
growth controlled by the direction and energy of the
impinging energy. This is common in the three models
defined above. The models are however of two totally dif-
ferent reef growth environments – the shelf edge and the
mid-shelf platforms. They are almost two end members
with different characteristics produced by the same param-
eters. In both cases the dominant control is substrate, to
produce a linear outer shelf feature and crescentic/lagoonal
mid-shelf features. However, in the mid-shelf reefs, Davies
(1978), Davies and Marshall (1980), Hopley (1982),
Davies (1983), and Davies and Hopley (1983) all recog-
nized patterns of reef development with specific character-
istics which appeared to overlap and which could be
related to the depth, size, and shape of the Pleistocene sub-
strate, the timing of reaching sea level, and the total energy
within the physical environment. Hopley (1982) and
Hopley et al. (2007) show similar models for the central
reefs. In an evolutionary sense, reefs can be judged to be
juvenile, mature, and senile. Thus, crescentic reefs (Stan-
ley, Fitzroy) can grow into lagoonal platform reefs and
eventually into flat-topped platforms with live coral
growth only around the perimeter. Most importantly, the
rate of progress along the evolutionary chain is a function
of the size and depth of the platform and the energy
impinging in the stillstand system. Thus, small reefs
(Wreck in the Capricorns) will progress to senility rapidly
while Stanley is yet to achieve maturity. The three reef
types will however be dominated by both different pro-
cesses and the same processes acting at different rates. This
becomes an important issue when considering how reefs
will react to climate changes proposed for the next 50
years. As a consequence, for example of increased stormi-
ness, mature reefs (and most are) will progress more rap-
idly to senility in spite of (or rather because of) possible
increases in growth potential in areas which are not temper-
ature marginal. The same conditions affecting a senile reef
will accentuate erosion leading to possible obliteration of
such reefs, particularly if they are small in size.
Summary
The Great Barrier Reef occupies the passive margin of
northeast Australia, and is arguably the largest coral reef
system to have ever existed on the planet. It represents
one of a series of major architectural features forming
the margin – the Eastern, Queensland and Marion Pla-
teaux; the Pandora and Bligh Troughs; the Osprey Embay-
ment; and the Queensland, Townsville, and Cato troughs.
Coral reefs occur on all three plateaux in addition to the
Great Barrier Reef. Some 3,600 reefs are estimated to
form the Great Barrier Reef. The margin architecture seen
today is a consequence of rifting, drifting, subsidence, sea-
level/climate change, and continental collision. Coral
reefs established on three occasions (1) in the early to
late-mid-Miocene; (2) in the uppermost Miocene; and
(3) in the Pleistocene. The Great Barrier Reef, as it is
known, started in the late Pleistocene, as seen in drill core
data from Ribbon 5/Boulder Reefs and at ODP site 820.
On the outer shelf at Ribbon 5, drill core data show
a reef section (0–96 m) above a rhodolith-dominated sec-
tion (95–158 m) above a packstone/wachestone section
(158–210 m). Initial reef establishment of the coral frame-
works in the middle of the rhodolith section dates from
some time after 465 ka while the firm establishment of
the main reef section above 96 m dates from around
300,000 years. At Boulder Reef on the inner shelf, tenta-
tive dating places reef start-up at around 210,000 years.
Reef initiation is therefore most probably facies controlled
and not karst antecedent controlled. The ages and the suc-
cession of reefs at Ribbon 5 (six reefs) and Boulder Reef
(four reefs) also testify to the fact that reefs have been
destroyed and re-grown again on at least six occasions
on the outer shelf and four occasions on the inner shelf.
The modern Great Barrier Reef is the last expression of
regrowth following the last post-glacial transgression.
Almost certainly, antecedence has assumed greater rele-
vance through each new growth phase.
The development of the modern reef system is
documented through the largest geoscientific reef data-
base for any system in the world (160 drill holes in 50 reefs
and 750 dates). Start-up occurred 7–10,000 years BP most
often on eroded (karstified) remnants of the previous
(125 ka) reef phase. Nearly 60% of all reefs cored and
dated reached sea level in the interval 5,000–6,500 years
BP meaning that nearly 60% of the reefs studied have been
 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT 531

4 3 2 1
0

Metres
Sediment transport 10

Terrigenous Carbonate

40 CMSEC –1
80 CMSEC–1 ? 15
27 GM M3 1.57 GM M3
up to 1 M
20
Trade wind
Cyclones Branch coral facies
conditions
A
Coral head facies
Sediment facies

4 Pleistocene LST

3
B
2 C14 yrs x 103
1
0 4 8
0

10

Metres
1

20
a

Highstand
A B
1

Coralline cap
Lowstand
Branching Fm
2
Massive Fm
Rubble
Inter-tidal sand
Sub-tidal sand
Highstand Lagoon sand
3 Antecedent surface
Caliche
Vadose zone

Great Barrier Reef: Origin, Evolution, and Modern Development, Figure 17 Ribbon Reef 5 (a) Three dimensional model of Ribbon
5 down to about 25 m and showing the lithologic and radiocarbon dates from drill holes (1–4). (b) The distribution of facies
developed in a high energy reef system as a function of successive sea-level oscillations. Facies variations within the successive reefs
show backward prograding sedimentological and biological wedges.
532 GREAT BARRIER REEF: ORIGIN, EVOLUTION, AND MODERN DEVELOPMENT
subject to a shallow water energy regime for most of the
time that they have been growing; growth therefore
reflects this. Different reef growth strategies reflect the
interaction of substrate depth, sea-level rate of change,
and coral growth rate. A major change in growth and
bio- and sedimentary facies occurs when the growing reef
approaches sea level – the formation of algal and coral
flats reflect biofacies changes and sand sheets and rubble
flats reflect sedimentary facies changes.
Five broad carbonate facies are recognized; three are
framework facies and two are detrital facies:
1. Coralline algal facies. These are laminated crusts,
often centimeters thick and/or encrustations on
branching or massive corals. The facies is best devel-
oped on windward margins, but encrusting corallines
coat most dead or dying surfaces in all intertidal reef
environments.
2. Branching coral facies. Growth form varies with depth,
light, and energy. In shallow water, variations in energy
define wide differences in species composition and
diversity. Water-depth changes produce similar com-
positional variations.
3. Massive (head) facies. As with the branching forms,
water depth and energy define diversity.
4. Sand facies. These are usually medium to coarse sand
comprise coral and corallines, benthic foraminifera,
and minor Halimeda. Forms large sand sheets between
coral flats and lagoons and also infills lagoons.
5. Rubble facies. These are comprised of massive, platey,
and branching corals. Produced either by in situ col-
lapse (in lagoons) or by energy-related breakage and
transport.
Three growth models are presented, two for mid-shelf
platform reefs (One Tree and Stanley Reefs) and one for
outer shelf barrier reefs (Ribbon 5). It is concluded that
most Holocene reefs in the Great Barrier Reef, with the
possible exception of fringing reefs, have undergone the
same basic processes of growth from an antecedent plat-
form to sea level and then growth controlled by the direc-
tion and energy of the impinging energy. This is common
in the three models defined above. The models are how-
ever of two totally different reef growth environments –
the shelf edge and the mid-shelf platforms. They are
almost two end members with different characteristics
produced by the same parameters. In both cases, however,
the dominant control is substrate, to produce a linear outer
shelf feature and crescentic/lagoonal mid-shelf features.
However, in the mid-shelf reefs, patterns of reef develop-
ment are recognized which have specific characteristics
which appear to overlap and which are probably related
to the depth, size, and shape of the Pleistocene substrate,
the timing of reaching sea level, and the total energy
within the physical environment. In an evolutionary sense,
reefs can be judged to be juvenile, mature, and senile.
Thus, crescentic reefs may grow into lagoonal platform
reefs and eventually into flat-topped platforms with live
coral growth only around the perimeter. Most importantly,
the rate of progress in this evolution is a function of the
size and depth of the platform and the energy impinging
in the still stand system. Thus, small reefs (Wreck in the
Capricorns) will progress to senility rapidly while Stanley
(large reef) is yet to achieve maturity. Evolutionary pro-
gression may occur through more than one highstand
growth phase. Predicted future climate changes will accel-
erate evolutionary progression.
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Cross-references
Algal Rims
Antecedent Platforms
Barrier Reef (Ribbon Reef)
Boulder Zone/Ramparts
Fossil Coralline Algae
Fringing Reefs
General Evolution of Carbonate Reefs
Glacio-Hydro Isostasy
Halimeda Bioherms
Lagoons
Plate Tectonics
Reef Classification by Hopley (1982)
Reef Classification by Maxwell (1968)
Reef Classification, Response to Sea Level Rise
Reef Flats
Rhodoliths
River Plumes and Coral Reefs
Submerged Reefs