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Cenozoic corals of SE Asia 165

Implications of paucity of corals in the Paleogene of SE Asia: plate

tectonics or Centre of Origin?

Moyra E. J. Wilson1 and Brian R. Rosen2

SE Asia Research Group, Royal Holloway University of London, Egham, Surrey, TW20 0EX, UK
Department of Palaeontology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK

Key words: scleractinian corals, evolution, carbonates, Indo-West Pacific, Cenozoic, Mesozoic, tectonics

Abstract We compare these patterns with the three main models of

high diversity in the Indo-West Pacific Centre (Centres of
Origin, Accumulation and Survival). During the Paleogene
The modern Indo-West Pacific centre of marine diversity,
the region was not a centre of any kind. From the Neogene
embracing SE Asia, has the highest zooxanthellate coral di-
onwards, no single model is applicable, but a combination of
versity in the world, and is characterised by abundant coral
all three models is preferred. Comparison with Mesozoic
reefs. In contrast, field and literature studies show that in
coral data and the history of the region suggests that the
Paleogene carbonates of SE Asia, corals are rare, and exten-
patterns discussed here also apply to the Late Triassic and
sive coral reefs have not been reported. Corals and reefs of
Late Jurassic. As with the Neogene, these were times when
this age may have been better developed in the mid-Pacific,
blocks rifted from Gondwana moved across the tropics to
especially in the late Eocene, but the most diverse regions at
dock against Asia. We therefore emphasize the role of plate
this time were in Europe, and to a lesser degree, the Carib-
tectonics in controlling regional high diversity patterns of
bean. The apparent scarcity of Paleogene corals in SE Asia is
zooxanthellate corals. Since corals and coral reefs provide
surprising in view of the long-standing theory that the Indo-
habitats for a myriad of marine organisms, the biogeographic
West Pacific is a Centre of Origin. The question therefore
history of Cenozoic reef corals has implications for the evolu-
arises whether this ‘Paleogene gap’ is a sampling problem,
tion of tropical reef ecosystems.
and if not what are its causes and implications?
A review of SE Asian Tertiary carbonates in their tectonic
context shows that many shallow-water Eocene-Oligocene
carbonates were dominated by larger benthic foraminifera Introduction
and coralline algae. A new Eocene coral fauna (12 spp.) from
Sulawesi accords with earlier views that there was little en-
demicity in SE Asia during the Paleogene. The general pau- The waters of SE Asia contain the highest
city of corals in carbonate platform deposits confirms that the marine faunal diversity in the world (Stehli,
‘Paleogene gap’ is real, and not an artefact. Since SE Asia, 1968; Briggs, 1974; Paulay, 1997), explanation
particularly the western part, has remained in, or close to, of which has long attracted the attention of bio-
tropical latitudes throughout the Tertiary, climatic reasons
cannot account for this. Although local ecological factors
geographers. Reef-dwelling scleractinian corals
would have been important, the tectonics point to a degree typify this pattern, and the highest species
of geographical isolation from other coral rich regions as a diversity (c.400-500, Veron, 1995) and 55% of
key factor. These patterns changed dramatically, however, in the world’s coral reefs (Muller, 1995) occur in
the earliest Neogene, as Australian fragments collided with these waters (Figs.1 and 2).
SE Asia. Collision led to decreased isolation of the region and
generation of numerous shallow-water areas, with diverse
The most enduring explanation of these kinds
and abundant zooxanthellate corals occurring in the Miocene of high diversity foci is that they represent Cen-
of SE Asia, similarly to the present day. The geographical tres of Origin (evolutionary source or ‘cradle’:
complexity of SE Asia appears to have favoured localized Ekman, 1953; Stehli and Wells, 1971; Briggs,
isolation and origination of new taxa, though the fauna also 1974, 1992; Zinsmeister and Emerson, 1981;
consists of older relicts and taxa which migrated into the
region from elsewhere. All these processes contributed to Veron, 1995). Although detailed definitions and
making the Indo-West Pacific the richest region for corals interpretations of Centres of Origin vary (Rosen,
from the Neogene onwards. 1984, 1985), they all assume that taxonomic rich-

Biogeography and Geological Evolution of SE Asia, pp. 165-195

Edited by Robert Hall and Jeremy D. Holloway
© 1998 Backhuys Publishers, Leiden, The Netherlands
166 M. E. J. Wilson and B. R. Rosen



1-3 4-8 9-15 16-24 25-35 36-48 49-63 64-74

Numbers of zooxanthellate coral genera

Fig.1. Diversity (numbers of genera) of Recent zooxanthellate corals. Localities and data are mostly based on unpublished
compilation by Professor John W. Wells. Map after Rosen (1984) and see this work for locality data.

ness has resulted from a concentration of evolu- gions, but over time, their geographical ranges
tionary appearances over tens of millions of changed to coincide in the present high diver-
years (Table 1). Alternatively, the Indo-West Pa- sity focus (Table 1; Ladd, 1960; Newell, 1971;
cific high diversity foci may represent a Centre McCoy and Heck, 1976; Kay, 1984; Wallace,
of Accumulation (evolutionary sinks or ‘muse- 1998; Rosen, 1984, 1988; Jokiel and Martinelli,
ums’), whereby species originate in other re- 1992; Pandolfi, 1992). A third Centre of Survival


Source Sink
Centre of Origin Centre of Centre of Survival

WITHIN Speciation rates high low low

THE FOCAL Extinction rates low low low
REGION Immigration rates low high low

OUTSIDE Speciation rates low low low

THE FOCAL Extinction rates high high high
REGION Immigration rates high low low

PREDICTED Diversity conspicuously Increasing high No particular pattern of

DIVERSITY higher in the centre diversity through time change in high diversity
PATTERNS than in outside regions, in the centre, relative to in the centre, through
THROUGH and of greater duration outside regions. time, but diversity
GEOLOGICAL TIME than species turnover elsewhere diminishing
IN FOSSIL RECORD timescale (< 30 Ma) through time.

OLDEST RECORDS Mostly within centre No particular pattern or No particular pattern

mostly outside centre

Table 1. The three main models for the origin of the Indo-West Pacific high diversity centre. See text for references. The table
shows the main evolutionary characteristics within and beyond the centre, together with predicted patterns for fossil record.
‘High’ and ‘Low’ convey relative not absolute rates. The classic arguments about patterns of oldest/youngest and/or most primitive/
derived taxa are reduced here to what can be reasonably easily investigated in the fossil record, especially in the context of this
paper. Related ideas about competitive displacement are omitted. The dichotomy, “cradle or museum?” was posed by Stebbins
(1974) for the tropics as a whole, but is here applied regionally to a feature within the tropics.
Cenozoic corals of SE Asia 167

120°E 0°




0 5000
Tropical carbonates with reefal corals km at Equator

Fig.2. Present day distributions of tropical carbonates with zooxanthellate corals from Nelson (1988).

or refuge model postulates that the fauna of SE particular fungiid patterns sufficiently to de-
Asia is a vestige of a formerly rich, cosmopolitan velop robust biogeographical hypotheses.
distribution of broadly Cretaceous origin (Table Veron’s (1995) theory of ‘surface circulation vi-
1; McCoy and Heck, 1976). Other ideas mostly cariance’, though drawing on fossil evidence
incorporate aspects of the three main hypoth- from age-and-area patterns and evolutionary
eses and vary in having an ecological to a his- rates, is not critically dependent on the fossil
torical emphasis (Rosen 1988; Crame, 1992; record. A regional palaeontological approach is
Veron, 1995; Palumbi, 1996; Paulay, 1997). limited to pioneering endemicity work by Gerth
This paper concentrates on the three main al- (especially 1925, 1930), and brief observations
ternatives (Table 1). The models are essentially by Umbgrove (1946) and Rosen (1988). Rosen
historical, yet paradoxically, little regional pal- and Smith (1988) subsequently analysed ende-
aeontological or historical evidence has been micity patterns of reef corals, worldwide, for the
assimilated into theories of coral evolution in SE early Miocene. This modest body of work con-
Asia. The main way in which palaeontology has trasts sharply with the extensive investigations
been applied to the history of the Indo-West Pa- of the much smaller and less diverse Caribbean
cific corals is through age-and-area studies, i.e., fauna (e.g., Budd et al., 1992, 1994, 1996).
region by region analysis of global stratigraphic If the Indo-West Pacific focus was indeed a
ages of extant taxa (Stehli and Wells, 1971; Centre of Origin, it should be recognizable in
Rosen, 1984; Veron, 1995), although, as shown the fossil record as a region of persistent high
below, this approach is flawed. Cladistic bio- diversity (Table 1) lasting significantly longer
geographical studies on extant corals have now than the <30 Ma timescale of species turnover.
also commenced (Pandolfi, 1992; Hoeksema, Although the fossil record must be incomplete,
1989, 1993; Wallace et al., 1991) but so far, only it can provide constraints, particularly when
Hoeksema (1989) has incorporated fossil evi- compared with evidence from living species,
dence. He was however unable to resolve his and it provides historical clues from large scale
168 M. E. J. Wilson and B. R. Rosen

spatio-temporal patterns (Table 1). This paper patterns discussed later in this paper.
therefore examines the Cenozoic fossil coral Scleractinians secrete hard calcareous skel-
record of this focal region, and interprets this in etons, and those under discussion mostly be-
the light of new knowledge of the tectonics and long to a broad ecological group that is confined
stratigraphical distribution of carbonates and to warm shallow waters often on, or close to,
coral facies. coral reefs. Today most of these scleractinian
corals host symbiotic unicellular algae
(zooxanthellae) within their soft parts and
Background hence are ‘zooxanthellate corals’ or ‘z-corals’.
The algae require sunlight to photosynthesise
Regional terminology and this restricts the corals to the euphotic zone
(about 100 m depth). Scleractinians mainly oc-
As a physical region, SE Asia broadly corre- cur within a low latitude circum-global band
sponds to Ekman’s (1953) ‘Indo-Malayan’ bio- between 35-40oN and about 35oS (Fig.2). The
geographical region, within which a triangular lowest mean annual surface sea-water tempera-
area formed by the Philippines, the Malay pe- tures are about 16oC, although species tolerance
ninsula and New Guinea shows the richest spe- may be lower (Rosen, 1984; Veron, 1995), and
cies diversity for numerous shallow marine the richest faunas are generally found in the
groups of organisms (Stehli, 1968; Briggs, 1974, warmest waters (Wells, 1954a; Rosen, 1984;
1992). Richness of corals progressively declines Veron, 1995; Fraser and Currie, 1996).
in all directions with increasing distance from Superimposed on this latitudinal pattern are
this central triangular area (Fig.1; Rosen, 1984 – regional ecological and historical biogeographi-
frequency plot of coral genera; Veron 1995 – in- cal factors, the most important of which is avail-
terpolated ‘contour’ plots of coral species). In ability of habitable areas, itself a combination of
this paper, we use ‘SE Asia’ for the marine re- numerous factors including temperature, nutri-
gion of the SE Asian mainland, the Australian ent flux, substrate, water depth and quality. Z-
margins and the areas between these, and use coral distributions are therefore patchy in space
the same term for the Cenozoic equivalents of and time, with modern examples found espe-
this region (Figs.2-8). This is preferred to ‘Indo- cially in clear, shallow conditions along land-
Malayan’ which presumes a rigorous biogeo- mass margins, and particularly in archipelagos.
graphical definition, criteria for which may not Although z-corals are common reefal inhabit-
have applied in the past. ants, the temperature limit of reefs (minimum of
The marine fauna of SE Asia falls within a 18oC; Veron, 1995) is slightly warmer than that
much larger biogeographical region, the ‘Indo- of z-corals, and the distribution of reefs and z-
West Pacific’ (Ekman, 1953; Paulay, 1997), corals do not correspond precisely. Contrary to
which stretches from the Red Sea and western widespread belief, fossil z-coral faunas are often
Indian Ocean to the eastern fringe of the main found in both reefal formations and bedded de-
archipelagos of the mid-Pacific. Across the east- posits, the latter being often silty, marly or even
ern Pacific is the East Pacific Barrier, with the pyroclastic.
East Pacific region beyond. The combined Indo- Corals mostly disperse by planktonic gametes
West Pacific and East Pacific constitute the and larvae, and the potential migration dis-
‘Indo-Pacific’. The SE Asian area of high diver- tances and biogeographic patterns of corals de-
sity is also referred to loosely as the ‘Indo-West pend on numerous factors including duration of
Pacific centre’ or ‘focus’, though the boundaries larval phase, life history patterns, water condi-
are not rigidly defined, and to the east and tions, substrate factors, coastal and sea-floor
southeast, the westernmost Pacific and northeast configuration, hydrodynamics and ocean cur-
Australia are often included (Fig.1). Tectonically rents (Jackson, 1986; Harrison and Wallace,
therefore, this centre lies across several plate 1990; Done et al., 1996).
boundaries, covering parts of the Indo-Australian, Populations of living corals tend to consist
Eurasian, Philippine Sea and Pacific plates. largely of either z-corals, or azooxanthellate cor-
als (az-corals), with z-corals dominating in warm
shallow conditions (Stanley and Cairns, 1988;
Zooxanthellate corals Coates and Jackson, 1987). Mixed assemblages
also occur, so simple ‘either/or’ assumptions
Some knowledge of scleractinian coral ecology cannot be made about the composition of past
is essential to understand the significance of faunas. Hence, for present purposes, the z-coral
Cenozoic corals of SE Asia 169

component of a fossil fauna has to be identified 1996; Paulay, 1997), have mostly made little ref-
and the az-coral component removed, in order erence to this palaeontological pattern, prefer-
to filter out the potentially conflicting ecological ring indirect evidence from age-and-area stud-
and biogeographical signals of az-corals. ies. One reason for this may be an uncertainty
over how ‘real’ the gap is, and one of the main
aims of this paper is to assess this.
The ‘Paleogene gap’ in Indo-West Pacific coral
Approach and Methods
Preliminary evidence in the existing literature
suggests that the high diversity centre of the Coral records
Indo-West Pacific is relatively young, since coral
diversity in the region was lower than else- To investigate the main relevant biogeographi-
where until the Neogene. Gerth (1925; see also cal hypotheses (Table 1), we have compiled pre-
Gerth, 1930) for a similar discussion in English viously published z-coral data for SE Asia and
but without his previous data compilations and neighbouring regions, and also included an im-
map) indicated four localities with Paleogene portant new fossil coral fauna found mostly in
‘reef corals’ in Borneo, Java, Sulawesi, Papua the upper Eocene part of the Tonasa Formation
New Guinea and a dubious reefal locality in of Sulawesi (Wilson, 1995; Fig.10). Knowledge
New Zealand. In contrast, for the Neogene, he of fossil corals in SE Asia is derived largely from
identified nine coral areas within the Indone- investigations made during the days of the
sian archipelago alone, and noted a three-fold former Dutch colonial era. More recently, the
increase in generic diversity of ‘reef-building French have studied the Mesozoic of the region
corals’ from 29 in the Paleogene to 76 in the (see below), but no recent taxonomic accounts
Neogene-Quaternary. Umbgrove (1930), con- exist for the Tertiary. Taxonomic revision of the
veyed a similar picture, and in his review of the entire Tertiary scleractinian fauna is undoubt-
Tertiary fossil record of extant Indo-Pacific cor- edly necessary. Unfortunately, the published
als, made no note of any from Paleogene strata coral collections often consist of single small
(Ta-Td; East India Letter Classification, cf., samples of each species, and it is difficult to ob-
Adams, 1970). In comparison, for Te-Tf strata tain any picture of species variability from them.
(regarded then as entirely Miocene), he men- Revisionary taxonomy is outside the present
tions formations that were known to contain scope, and estimates of species numbers here
rich and well-described coral faunas. Umbgrove are based on existing nomenclature, with taxo-
(1946) subsequently suggested that increased nomic adjustments made only in obvious cases.
speciation of corals in SE Asia occurred during Moreover, the stratigraphic, palaeoenvironmen-
the late Miocene (Tg). tal and tectonic context of these corals in the old
An Oligocene to Recent generic level com- literature was poorly documented by modern
parison of the various major reef coral re- standards. Today, since exploration for natural
gions of the world (Rosen, 1988; his Table 2) resources often focuses on carbonate rocks,
showed that reef coral richness in the which commonly include the remains of corals
Oligocene was highest in the Caribbean and and other marine organisms, new, well-docu-
Mediterranean, but from the Miocene on- mented collections are being made. The nature,
wards the Indo-West Pacific centre became extent and history of carbonate development in
the richest region, increasing more than four- SE Asia is therefore becoming clearer.
fold, from 14 genera in the Oligocene to 58 in
the early Miocene (below and Fig.9).
Despite the prevalent theory that the Indo- Inference of algal symbiosis in fossil corals
West Pacific focus represents an evolutionary
centre, it is surprising that the Paleogene of this Zooxanthellae are not preserved in fossil coral
region is apparently so poor in z-corals, thus skeletons, but algal symbiosis often needs to be
implying that this centre is only a post-Paleo- inferred for palaeobiogeographical studies. A
gene phenomenon. Although evidence for this range of evidence can be used to infer symbio-
‘Paleogene coral gap’ goes back 70 years, the sis, but methods based on intrinsic features of
numerous ideas that have been developed con- the corals, including colonial characters, skeletal
cerning Indo-Pacific marine biogeography (e.g., stable isotope geochemistry, palaeoautecology
Rosen, 1988; Veron, 1995; Flessa and Jablonski, and phylogenetic relationships, are preferable
170 M. E. J. Wilson and B. R. Rosen

Table 2. Simplified scheme for designating levels of confidence in methods for inferring algal symbiosis in a fossil coral species.
Note criterion 3(a) is particularly unreliable because although most living forms in such habitats are zooxanthellate, such
habitats today also contain az-corals. With respect to 4, note that ‘nkz’ status is a neutral category, and does not signify
inference of azooxanthellate status. The latter can be inferred using criteria parallel to 1-3. Inference of symbiosis in genera
(e.g., Fig.9) can be derived from integration of respective member species, or approximately, by applying the parallel criteria at
generic level. Examples of corals in different ‘z-categories’ are shown in Figs.10 and 12. In this paper, we use criteria 1-2 to
obtain a minimum estimate of frequency of z-corals in a given fauna, and criteria 1-4 to obtain a maximum estimate (Table 3).


LEVELS' (abbreviated)

1. Species is still living today and known to be zooxanthellate z-coral

2. Species is extinct, but

(a) belongs to a genus whose living species are all z-corals z-like coral
(b) has skeletal characteristics found only in living z-corals (mainly stable isotopes, morphology)

3. Species is extinct but

(a) occurs in warm shallow-water carbonate facies z-like coral?
(b) lacks skeletal characteristics consistently found in living z-corals
(c) there is doubt about applicability of other stronger lines of evidence (i.e. 1 and 2)

4. Species is extinct and there is currently no evidence for inferring algal symbiosis, nkz-like
but symbiosis cannot be ruled out (i.e., status of algal symbiosis not known)

(Table 2; Rosen, 1998). Different methods have gional tectonics. The aim is to assess whether
different merits so corals can be grouped ac- there are significant areas in SE Asia which are
cording to a broad, albeit subjective, level of potentially rich in z-corals, but which lack coral
confidence according to method, strengthened information. Inhabitable areas for z-corals can
where applicable by corroboration from differ- then be assessed against other factors such as
ent independent methods. Since algal symbiosis taphonomic effects, climate, and biogeographi-
in fossil corals is an inference, it is more precise cal barriers.
to regard fossil examples as ‘z-like’ or ‘az-like’ Tertiary carbonate rocks of SE Asia are
rather than ‘z-corals’ and ‘az-corals’ respectively formed mostly in shallow marine environments
(Rosen and Turnsek, 1989), although the latter away from clastic input and are composed of the
pair of terms may be used for simplicity. calcareous skeletons of marine organisms. Mod-
A simplified summary of methods, criteria and ern z-corals are mostly restricted to carbonate
approach is shown in Table 2. Each group of environments and through a study of these
criteria and its respective confidence level is de- lithified sediments and their constituent compo-
noted in the third column. If inference from first nents it is possible to track the availability of
order criteria is not applicable, then second or- potential habitats for reef corals in the past.
der criteria are used, and so on, working Data on the carbonate rocks of SE Asia have
through criteria with decreasing levels of confi- been extracted from the literature and are also
dence. A given faunal list can then be parti- based on current field research by the authors
tioned into those for which symbiosis has been and other members of the London University SE
inferred (or not) according to these levels of Asia Research Group.
confidence (see captions to Tables 2 and 3). In SE Asia, local carbonate studies usually
concentrate on the sedimentological rather than
palaeontological aspects of the rocks, and taxo-
The sampling problem and reconstruction of nomic studies rarely include information on the
habitable areas sediments. However, sedimentologists usually
note the presence of corals and, although taxa
The regional geographical and stratigraphic lo- are not always identified, information on
cations of all the Paleogene coral faunas have growth forms, together with an indication of
been compared with facies distributions and re- coral abundance and diversity are commonly
Cenozoic corals of SE Asia 171

given. This is clearly not ideal for detailed bio- Brasier, 1995). They occur in similar tropical en-
geographical work, but when combined with vironments to corals and have broadly equiva-
taxonomic data, provides information on Ceno- lent global distribution patterns (Adams, 1983;
zoic z-coral distributions for the region. Hallock and Glenn, 1986; Adams et al., 1990). A
number of depth zonation schemes for larger
benthic foraminifera (Ghose, 1977; Hottinger,
Reconstructions 1983; Hallock and Glenn, 1986; Gorsel, 1988;
Racey, 1994) and other shallow marine
To track the location and nature of potentially bioclasts, such as coralline algae (review in
available habitats for corals during the Ceno- Perrin et al., 1995) have been deduced for differ-
zoic, three main carbonate rock types have been ent stages during the Cenozoic. The taxa, to-
plotted on to time slices of Hall’s (1996) recon- gether with their forms and the nature of the en-
structions between the late Eocene (about 37 closing sediment, can be used to infer
Ma) and Recent (Figs.3-8). Paleocene and lower depositional depths for carbonate sediments
Eocene carbonates are rare in SE Asia and there (Fig.11). Therefore an ancient carbonate forma-
have been no published reports of z-corals of tion in which specific larger benthic foraminif-
this age, although a Paleocene locality has been era dominate, such as robust Nummulites,
reported in Java (A. Russo, J. Pignatti, pers. would have been deposited in a shallow marine
comms., 1997). This suggests that conditions area during the Eocene or early Oligocene,
suitable for z-coral growth were sparse, and re- where one might also expect to find z-corals. For
constructions for these intervals are not shown. each carbonate formation illustrated on the
The ages of the Cenozoic carbonate formations maps (Figs.3-8), the setting (S) and the nature of
are based on a range of nannofossil, planktonic corals (C) present are briefly described.
and larger benthic foraminifera, and, in a few The limits of the maps correspond to 30oN and
cases, strontium isotope dates. Although full 20 S (Figs.3-8) – close to the latitudinal limits of

documentation is not possible here, these have the modern reef coral belt to the north, and
been gleaned from a large literature database somewhat within the belt to the south. The
and the authors’ own work, and in a few cases it ‘reefal belt’ has variously widened and nar-
has been necessary to revise ages (see Table 3). rowed through geological time in response to
The main environments illustrated on the phases of global warming and cooling. For ex-
reconstructions of relevance to marine bio- ample, the early to middle Miocene corresponds
geographers are: shallow marine carbonate to a period of global warming, and the reef belt,
depositional environments, major land areas together with distributions of z-like corals and
which would usually have been surrounded by larger benthic foraminifera, was as much as 10o
clastic shelves, and deep marine areas (below of latitude wider (Fulthorpe and Schlanger,
100-200 m water depth). 1989; Adams et al., 1990; James et al., 1996).
A wide variety of skeletal remains of shallow Davies et al. (1989) constructed an oceanic sur-
marine organisms are present in Cenozoic car- face water temperature curve for NE Australia
bonate deposits, including corals, larger benthic and suggested that the earliest part of the
foraminifera, coralline algae, Halimeda, echi- Paleocene was also a period of global warming,
noids and molluscs. However, in most SE Asian sufficient to sustain reefal coral growth.
carbonates the dominant constituent grains are
coral fragments and larger benthic foraminifera,
although coralline algae may be locally abun- Summary of Mesozoic scleractinian records
dant. Carbonate deposits have been divided in SE Asia
here into three categories on the basis of these
components, and the different types are illus- The earliest global records of scleractinian cor-
trated on the maps (Figs.3-8). These are carbon- als are Middle Triassic (Wells, 1956; Stanley,
ates where z-like corals are abundant, carbon- 1988), since when scleractinians have occurred
ates dominated by larger benthic foraminifera discontinuously in SE Asia, concentrated over
and carbonates where both z-like corals and particular time spans. Triassic corals (largely
larger benthic foraminifera are abundant. Upper Triassic) are represented on the Asian
Modern larger benthic foraminifera, similar to mainland (Yunnan and around the Burma-Thai-
many living corals, contain symbiotic algae, and land border) and in the Indonesian archipelago,
their growth is thought to be restricted to the mostly east of Borneo in Sulawesi, Buru,
euphotic zone (Haynes, 1965; Hallock, 1985; Misool, Seram, Timor and Roti (Stanley, 1988;
172 M. E. J. Wilson and B. R. Rosen

Martini et al., 1997). The Timor fauna appears to 1953; Paulay, 1997). Until the latest Paleogene,
be the richest with over 30 nominal species. The the whole Australian craton and its coastal wa-
Jurassic is represented by some Middle Jurassic, ters lay too far south to fall within the bounda-
but more extensively by Upper Jurassic faunas ries of the z-coral belt. It was not until between
(Beauvais, 1986), some, like those of Sarawak, 25 to 16 Ma (late Oligocene to early Miocene)
being quite rich (about 20 species or more). that the transition from temperate to tropical car-
They may continue in some places into the bonates, with abundant z-coral and reef devel-
lowermost Cretaceous (Beauvais and Fontaine, opment, occurred in the northern part of what is
1990). Upper Jurassic corals are also known now the Great Barrier Reef (Davies et al., 1987).
from Sumatra, Burma, Thailand and the Philip- The central part of the Great Barrier Reef did not
pines (e.g., Beauvais, 1983), whilst substantial experience a tropical climate until 15 to 10 Ma,
Middle to Upper Jurassic faunas occur in Japan and the southern part only became tropical in
(Eguchi, 1951). In contrast, although Cretaceous the last few million years (Davies et al., 1987).
carbonate platforms do occur in SE Asia (Cebu, The Indo-Pacific gateway progressively nar-
Misool, Timor and Papua New Guinea), Creta- rowed during the Cenozoic as Australia moved
ceous corals are rare throughout the region, ex- northwards (Figs.3-8), and closer to SE Asia.
cept, again in Japan. A few Cretaceous corals are This, combined with the northward drift of vari-
known from the Upper Cretaceous of western ous microcontinental blocks, and emergence of
Borneo and northern Sumatra, but rudists ap- new islands, or development of new shallow
pear to be the dominant shallow water element carbonate producing areas, would have in-
(P. Skelton, pers. comm., 1997). Overall, corals creased the potential for exchange of coral lar-
were most common in the Upper Triassic and vae with other regions and the development of
Upper Jurassic, partly reflecting times when cor- z-coral communities and coral reefs (Rosen,
als flourished globally. 1988; Pandolfi, 1992). Davies et al. (1989) noted
that the closure of this seaway would also have
restricted equatorial current flow and caused
Cenozoic plate tectonics of SE Asia and the diversion of warm tropical waters to the
implications for coral history south along NE Australia (see also Grigg, 1988).
The Indian continent drifted northwards
SE Asia is an extremely active tectonic area, and across the zone of reef coral growth during the
considerable lateral and vertical movements of Late Cretaceous and early Paleogene, and is in-
the Earth’s crust throughout the Mesozoic and ferred to have collided with the SE Asian main-
Cenozoic have affected the spatio-temporal vari- land during the Eocene (Hall, 1998 this volume).
ations in carbonate depositional environments During this time suitable habitats for z-coral de-
and hence the potential habitats for corals. Dur- velopment may have occurred on narrow
ing this time the Indo-Australian and Philippine- shelves with limited clastic input around the
Pacific plates, and a large number of smaller landmasses of India and mainland SE Asia. Ex-
microcontinental and oceanic fragments, inter- amples include the z-like coral communities
acted and collided with Sundaland, the stable known from upper Paleocene to Miocene strata
eastern margin of the SE Asian craton (Figs.3-8 along the NW side of the Indian continent in
and in Hall, 1998 this volume). The reconstruc- Kaachchh and western Pakistan (Duncan, 1880;
tions of Hall (1996), were used for Figs.3-8, but Gregory, 1930; Ghose, 1982; Rosen, 1988; re-
other recent reconstructions (Rangin et al., 1990, vised dates in Adams 1970), belonging tectoni-
Daly et al., 1991; Lee and Lawver, 1995) yield a cally to India (C. Izatt, pers. comm., 1997).
similar picture for the purposes of plotting shal- Following the Eocene collision of India with
low marine carbonate depositional environ- Asia, and the subsequent uplift of the Himalayas
ments, although variations in timing may give and other related mountain chains, large quanti-
slightly differing patterns (cf. Pandolfi, 1992). ties of clastic material were shed into the adja-
At the beginning of the Tertiary, the separa- cent seas via large delta systems. These deltas,
tion of Australia from mainland SE Asia across including those of the Irrawaddy and Ganges,
this ‘Indo-Pacific gateway’ was about 3000 km. currently inhibit reef coral growth, and form lo-
This distance, particularly if the current direc- cal coastal barriers to biogeographical exchange
tions were unfavourable, is sufficient to have of many shallow marine organisms. Substantial
been a potential biogeographical barrier to the clastic input along the southern margin of the
dispersal of z-coral larvae, comparable in scale Asian continent during the Paleogene may simi-
with the East Pacific Barrier today (Ekman, larly have inhibited z-coral growth. The closure
Cenozoic corals of SE Asia 173

of the intervening seaway may also have hin- firmed as upper Eocene, consists entirely of az-
dered the migration of shallow marine organ- like corals. This leaves a small number of pub-
isms between areas such as the Middle East and lished z-like faunas of Eocene age, but even the
SE Asia. dating of some of these is uncertain. In fact, our
There are very few islands in the modern In- new coral fauna of late Eocene (Tb) age from
dian Ocean between the Maldive-Chagos line Sulawesi (Wilson, 1995), though modest, is cur-
and Indonesia, although volcanic islands along rently the richest known z-like fauna of definite
the Ninetyeast Ridge were emergent during the Eocene age in SE Asia (Fig.10). A z-like fauna
Paleogene (Kemp and Harris, 1975). Prior to the from the Enewetak boreholes of the Marshall Is-
Paleogene, islands may have existed in the sea- lands on the Pacific plate, has an even smaller
way between India and mainland Asia. How- number of species (Table 3).
ever, following the juxtaposition of India with Chevalier et al. (1971) compiled other Eocene
Asia the islands of the Ninetyeast Ridge would ‘reefal and parareefal’ coral records including
have been among the few habitable areas for their own from New Caledonia. Most are in-
corals in the eastern Indian Ocean and the Indo- cluded in Table 3 and some are probably
Pacific gateway, and this otherwise open sea- younger than Eocene (see above). Localities
way may have formed a barrier to the survival of mentioned by these authors, not in Table 3, are:
most coral larvae. In contrast, in the western Sumatra (no references given), Sulawesi (after
and southern Pacific, numerous atolls, reefs and Bemmelen, 1970), Saipan in the Marianas (after
islands existed from the Cretaceous onwards Yabe and Sugiyama, 1935), and Makatea in the
(Menard, 1964; Schlanger and Premoli Silva, Tuamotus (after Obellianne, 1962). We have
1981; Grigg, 1988; Winterer, 1991), and z-coral failed to find the Sulawesi records in Bemmelen
communities existed there throughout the Ceno- (1970). Of Makatea, the authors say that Eocene
zoic. The mid-plate foundations of most of these corals “should exist”, as well as in “several other
shallow areas of the western Pacific originated raised islands of the Pacific” but they cited no
in a geographically restricted phase of further work. For Saipan, Yabe and Sugiyama
volcanicity during the Mesozoic (Darwin Rise/ (1935) describe only Saipania, which in our
Superswell). Since then, they have all been opinion is a chaetitid sponge, not a coral. How-
moving in unison, towards Asia, so narrowing ever, Eocene carbonates of Saipan (Matansa
deeper water areas to their west and widening Limestone and Densinyama Formation), and
those to the east (East Pacific Barrier). This must Eocene volcaniclastic rocks on Guam (Marianas)
have contributed to the development of z-coral are all reported to contain z-like corals (G.
habitats and larval exchange around SE Asia, Siegrist and D. Randall, pers. comm., 1997) and
while further diminishing potential for larval ex- require further study.
change across the East Pacific Barrier. Lower Oligocene records are even sparser
than for the Eocene (Table 3), but from the mid-
Oligocene (Td) there are signs of increasing
Cenozoic geological evolution and coral richness (SE Borneo). A silicified Oligocene
faunas coral reef is reported from Guam by Siegrist and
Randall (above). By the late Oligocene to early
Overview of Paleogene corals Miocene (Te), richer reefal coral faunas, similar
to those today, seem to have become progres-
At the start of the Paleocene, the global sively more common throughout much of the
scleractinian record mainly comprises az-like region (Fig.9).
corals, with an apparent hiatus of up to 2-3 Ma Fig.9 shows a Cenozoic compilation of ge-
before the first Tertiary records of z-like corals neric richness of z-corals for the four principal
(Rosen, 1998). In the Indo-West Pacific centre tropical marine regions (update of Table 2 in
the oldest recorded Tertiary corals are Eocene Rosen, 1988) incorporating new data from the
(Table 3), and superficially seem quite rich, al- Indo-West Pacific centre (Table 3) and other re-
though an unpublished Paleocene fauna has gions. This confirms the reality of the ‘Paleogene
been found in Java (A. Russo, pers. comm., Gap’ in terms of available coral data, though not
1997). On checking the Eocene records and necessarily its biogeographical reality, and
their associated larger benthic foraminifera (Ta- shows that substantially richer faunas occurred
ble 3), two of the richest faunas (SE Borneo and elsewhere, particularly in Europe. This pattern
Papua New Guinea) are probably Oligocene or reversed itself in the early Miocene: as the Indo-
younger, and a third (Tonga), although con- West Pacific centre became noticeably richer,
174 M. E. J. Wilson and B. R. Rosen


erL- A
spLeci- genera
E sApLecies
Dollfus, 1915 a es
SULAWESI Oligocene Gerth,1931 Neogene according to
Umbgrove, Umbgrove (1942b)
N eogene 1942b
BUTON Asphalt Umbgrove, Paleogene age ruled
deposits 1942a out by author
OL I G OC E N E M io c en e)
(to Latest Z-like corals in
PAPUA NEW Oligocene brecciated reefal
GUINEA (Boera Boera to earliest unpublished c.10 c.12 >4 c.12 Rlimestones (collected
Head, southern Formation Miocene osen & Darrell, NHM
Papua) (latest Te1- Collections; work in
5) p
Coorgarles snso
r ) t found in
PAPUA NEW outcrop;
Eocene, ?Oligocene Gregory & notwithstanding
GUINEA (Fly (Rolled probably - Trench, 1915 11 13 8 (?11) (?1103) published age, the
River, at 6o5'S and pebbles) Middle Gerth, 1931 fauna looks mostly
/ or at 5 40'S)
o ?Miocene
Neogene or of mixed
MARSHALL Borehole: No date
ISLANDS (Bikini Late Wells, 1954b unidentifiable coral fragments Age revised by Adams
Atoll) <355 m given (1970)
Oligocene -
MARSHALL Boreholes: Early 10 Age revised by Adams
ISLANDS 845-355 m Miocene Miocene Wells, 1964 8 11 7 ( ? 8) (?11) (1970)
(Enewetak Atoll) (Te)
SERAM (eastern Oligocene Dollfus, 1908 Umbgrove (1924) said
part of the island ('Sannoisea- Oligocene Umbgrove, 1 1 0 (?1) 0 (?1) coral was "probably
near Mt. Téri) n' = Early 1924 Oligocene"
Oligocene) Gerth, 1931
Dollfus (1908) stated
that Fritsch's fauna is
(Riam-Kawa and Verbeek's Mid ('Sannoisean' = Lower
Riam-Kanan area, Gamma Eocene Oligocene Fritsch, 1878 23 25 1 0
9 (?21) (?22) O ligocene).
Meratus Beds Gerth, 1931 Bemmelen (1970) cites
Complex) foraminifera that
suggest mid-Oligocene
(Td) (Adams 1970)
MARSHALL Boreholes: ? Mid Age revised by Adams
ISLANDS 931- 845 m ? Miocene Oligocene Wells, 1964 6 7 6 7 (1970), but no direct
(Enewetak Atoll) (Td) evidence for Td
MARSHALL Boreholes: ? Early Age revised by Adams
ISLANDS 976-931 m ? Miocene Oligocene Wells, 1964 3 3 3 3 (1970), but no direct
(Enewetak Atoll) (Tc) evidence for Tc
TONGA (Eua) Late Eocene Wells, 1976 14 17 0 0 Z-like corals absent
MARSHALL Boreholes: Late Eocene
ISLANDS 1220-1251 m (Tb) Wells, 1964 4 4 3 (?4) 3 (?4) A ge confirmed by
Adams (1970)
(Enewetak Atoll) Late
Late Eocene (Tb) from
Eocene Pellatispira (see
SUMBA (c.80 km Umbgrove, Adams 1970, and A.
from western Eocene (Tb) 1 2 0 ( ? 1) 0 (?2) R
end) 1943 acey, pers. comm.,
1997). See also
Bemmelen (1970)
SULAWESI Formation this paper 11 12 6 (?11) 7 (?12)
(basal part)
Mid-Eocene Gerth, 1921
JAVA (south Nanggulan Late Eocene tOoligocene Gerth, 1931 7 8 0 (?5) 0 (?5) cPoreraclissenhorizons of the
central) Beds Gerth, 1933 ot known
NEW (Ta3-Td)
CALEDONIA, SW Mid-Late Mid-Late
(Baie St Vincent Eocene ( Eocene Chevalier et 2 2 1 1
(îles Ducos and ‘Eocene C’) al., 1971
Saint Phalle); Oua (Ta3-Tb)
JAVA Jati Bunkus, Late Russo,
near reefal Paleocene Pignatti
Karangsambung limestones (Thanetian) (pers. 8 11 6 (?8) 8 (?11)
(Central Java) (Ta1) comms.,

Table 3. Summary of Paleogene coral faunas of the Indo-West Pacific centre including SE Asia. Ages based on foraminifera unless
indicated. Revised ages show that the Eocene is even more poorly represented than previously realised. By the late Oligocene z-
like corals were relatively common and the pattern is more Miocene-like, Te1-4 formations are therefore a selection only. For the
period before the late Oligocene the data are as complete as possible apart from a few problematic or unstudied cases (see text). Top
two rows show younger faunas which have been included because of their supposed Paleogene age. Minimum values of z-corals are
in bold and maximum values are in parentheses (see caption to Table 2). Paleogene Indo-West Pacific data for Fig.9 are based on
net totals of faunas in this table, except for the Oligocene (based on data from Kalimantan and the Marshall Islands).
Cenozoic corals of SE Asia 175

Foraminifera dominated carbonates

z-like corals & foramanifera present

z-like corals dominate

Land areas
S: platform
C: not known

37 Ma Late Eocene

S: mixed clastic
carbonate shelf
C: some Melinau
S: platform
C: rare

Ritan Davao
S: faulted highs
C: not known

S: clastic shelf Makale
C: reefal corals S: isolated platform
Gunung Putih C: few corals Cabariohan
S: platform
C: not known Tonasa
S: isolated platform
C: rare reefal corals

1000 km at equator
S: platform/shelfal
C: rare

Fig.3. Late Eocene time slice from the reconstructions of Hall (1996) with carbonate deposits shown. The nature of the carbonate
deposits (S) and any coral records (C) are shown. Abbreviations used in Figs.3-9 for corals records are: Br. branching, Pl. platy,
M. massive, Ab. abundant, Div. diverse. Names shown are formations. Fig.8 shows present-day geography.

the Mediterranean declined. Since then, the number of basins occurred around Sundaland
Indo-West Pacific centre has remained the rich- during the Paleogene (Hall, 1998 this volume),
est z-coral region. Throughout the Cenozoic, the and those to the east quickly became marine
Caribbean, by contrast, has remained relatively and carbonates developed (Fig.3). In compari-
constant and moderately rich. The picture in SW son, mainland SE Asia, Sumatra and much of
Asia is unclear because of lack of adequate data, Borneo (Fig.3) was a land area which was shed-
but Paleogene z-like faunas, such as those from ding considerable clastic material into the sur-
western Pakistan, together with unpublished rounding seas. Along the southern margin of this
coral collections in The Natural History Mu- land area, the shelf would have been narrow,
seum, London, from Oman, Iran and Somalia close to the adjacent subduction trench (Fig.3),
suggest that this region was also richer than SE and this, together with the amount of clastic in-
Asia during the Paleogene. put, would have hindered carbonate produc-
tion. The distance between shallow marine ar-
eas across the Indo-Pacific gateway was greater
Late Eocene (Tb) than 2000 km in the late Eocene.
In eastern SE Asia carbonate development oc-
Carbonate deposition was common on curred mainly as extensive shallow-water plat-
microcontinental blocks in the eastern part of SE forms, such as in Sulawesi (Wilson, 1995), the
Asia (Fig.3) where emergent land areas were Melinau Limestone of Sarawak (Adams, 1965)
small and clastic input was low. Initiation of a and New Guinea (Koesoemadinata, 1978). The
176 M. E. J. Wilson and B. R. Rosen

margins of these carbonate platforms varied some clastic input, and did not form reefal
from steep escarpments to gently dipping buildups (Wilson, 1995).
ramps, but reefal buildups have not been re- These SE Asian z-coral faunas are paltry com-
ported (Wilson, 1995). In western SE Asia, pared with the rich European faunas of Eocene
shelfal carbonates or localised areas of carbon- age, such as those known from former Yugosla-
ate deposition occurred where clastic input was via with 85 z-like species, or the 42 z-like spe-
insufficient to hinder carbonate production. cies known from Eocene of Panama (interpreta-
The Eocene carbonates are dominated by tion of data from Felix, 1925, and from Budd et
larger benthic foraminifera (Fig.11) and in some al., 1992, respectively). Moreover, apparent
cases coralline algae, while corals are extremely paucity of corals in SE Asia should also be seen
rare (Adams, 1965; Kohar, 1985; Siemers et al., in the context of Ladd’s statement (in Wells,
1992; Wilson, 1995). Within the carbonates, 1976) that upper Eocene limestones have been
there occur a wide variety of larger benthic reported from the Pacific at Palau, the Marianas,
foraminifera showing various adaptations, the Marshalls, Fiji and Tonga, across a distance
which suggest they occupied the full range of of 6560 km (see also Schlanger and Premoli
habitats available on the platforms within the Silva, 1981; Winterer, 1991). While some of
euphotic zone (Adams, 1965; Wilson, 1995). these, as in the Marshalls (Table 3), and
The first well-dated, published appearance of Marianas (G. Siegrist and D. Randall, pers.
z-like corals in the Indo-West Pacific centre oc- comm., 1997) contain z-like corals, this cannot
curred in the late Eocene. There is a possibility be assumed in all cases, since Eocene corals
of older (middle Eocene) z-like corals occurring from Eua, Tonga, (Table 3) are entirely az-like. It
in New Caledonia and Java (Table 3), but the may be that a largely concealed upper Eocene
age range of these very limited faunas also in- sequence with z-like corals is now submerged,
cludes the late Eocene. The Nanggulan Beds are or lies buried beneath younger volcanics or
mixed clastic-carbonate deposits and show no limestones throughout much of the Pacific. Thus
evidence of any reefal buildups or of in situ whereas the ‘Paleogene gap’ is a sampling real-
growth fabrics. An enigmatic Sumba locality ity for the Eocene of SE Asia, it is possibly a sam-
whose place-name we have been unable to lo- pling artefact for Pacific part of the Indo-West
cate on modern maps, is of late Eocene age, but Pacific centre.
contains a single genus which may not be z-like.
Following revision of the ages of Paleogene
corals in SE Asia (Table 3), the only known z- Early (Tc) to middle (Td) Oligocene
like corals of definite late Eocene age in SE Asia
are from the basal part of one area of the Tonasa The early and middle Oligocene were similar to
Formation in south Sulawesi (Fig.10 and Table the late Eocene with carbonates common on mi-
3). The fauna is small, consisting of 12 species, 7 crocontinental blocks in eastern Indonesia, but
z-like and 5 z-like?, mostly known also from the rare around the margins of mainland SE Asia
Paleogene of western Pakistan (Ranikot, Khirthar (Fig.4). An open seaway between the Pacific
and Nari series) and Europe, and therefore not and Indian Oceans was still present and was
strongly endemic. It includes small solitary dis- about 1,400 km wide by the end of the middle
coidal (z-like?) forms such as Cycloseris subcren- Oligocene (Td). However, as Australia and the
ulata and Trochocyathus ? nummiformis from associated microcontinental blocks moved
the lowermost beds of the formation at the Doi northwards, large carbonate platforms devel-
doi coal quarry. Well-preserved massive colo- oped in New Guinea (Fig.4, Wilson et al., 1993),
nial forms found about 2 km away, slightly and the distance across this gateway decreased,
higher in the succession, include Favia pedun- altering the configuration of ocean currents
culata (z-like), Astrocoenia bistellata (z-like?), (Grigg, 1988) and increasing the possibility of
and a striking four-lobed flabellate z-like form, biogeographical exchange of marine biotas.
Trachyphyllia indica. Specimens of all of these Carbonates were also deposited around the
show flattened bases in response to a soft margins of the developing South China Sea
substrate. There are also colonies of Stylocoenia (Holloway, 1982). Their distribution was similar
sp. (z-like?) which, with their unique spheroidal to the late Eocene, with platform and shelfal car-
corallith habit and stout stilt-like spines (but bonates dominating to the east and west respec-
eroded), are also soft-substrate dwellers (Darga, tively. Larger benthic foraminifera again domi-
1991). The corals from the basal part of the nate in most of these deposits (Fig.11), and
Tonasa Formation occurred where there was there is no evidence for any reefal buildups.
Cenozoic corals of SE Asia 177

Foraminifera dominated carbonates

z-like corals & foraminifera present

z-like corals dominate

Land areas

S: platform
C: not known 29 Ma early Oligocene

S: shelfal
C: some Melinau
S: platform
C: rare

S: clastic shelf
C: reefal corals Lutak
Berai Cabariohan S: shelfal
S: shelfal C: few
C: some Br. & Pl.

Gunung Putih
S: platform Salodik
C: not known Tonasa S: slope?
S: platform C: rare
C: rare Br.
S: shelfal Faumai
C: rare S: shelfal Biri
C: not recorded S: shelfal
C: not recorded

1000 km at equator

Waripi Mendi Group

S: shelfal S: carbonate/clastic ramp
C: rare C: generally absent

Fig.4. Early/middle Oligocene time slice from reconstructions of Hall (1996) with carbonate deposits shown; abbreviations on Fig.3.

Corals are rare in the lower Oligocene (Tc) of bean (Fig.9). No other published z-like fauna is
SE Asia, and those that do occur are described known from Td strata in SE Asia, but its richness
as thin branching forms, such as Acropora from relative to older Paleogene localities hints at an
south Sulawesi (Fig.10l). Some massive and increase in suitable areas for z-corals at this time
thick branching forms have been reported from in parts of SE Asia. A locality in Seram with just a
shelf margin deposits in south Kalimantan single coral record is dated as Oligocene (Table
(Saller et al., 1993). On the Pacific plate, a fauna 3) but its exact age is not known. On the Pacific
possibly of this age is known from the plate, a modest z-fauna of possible mid-
Enewetak boreholes (Table 3). The richest Oligocene age is known from the Enewetak
Paleogene z-like fauna in SE Asia is that de- boreholes (Table 3). Grigg (1988) discussed the
scribed by Fritsch (1878) from the Tanjung For- Cenozoic coral record of the mid- and northern
mation of SE Borneo (Table 3). Fritsch and later tropical Pacific and emphasized that z-corals
workers (e.g., Gerth, 1925; 1930) regarded this started to become more widespread in the early
fauna as Eocene, although the associated for- Oligocene, the date of their first appearance in
aminifera, Nummulites fichteli and Lepidocyclina the Hawaii-Emperor chain.
sp. (Bemmelen, 1970), point to an Oligocene
(Td) age. The corals consist of only nine clearly
z-like species, but if other corals of less certain Late Oligocene (Te1-4)
z-status are included, the total increases to 22.
This is small when compared with the rich con- Carbonates were still common in eastern Indo-
temporaneous faunas of Europe and the Carib- nesia (Fig.5), and some of the basins, such as
178 M. E. J. Wilson and B. R. Rosen

Foraminifera dominated carbonates

z-like corals & foraminifera present

z-like corals dominate

Land areas

S: platform/pinnacles
C: abundant towards top 26 Ma late Oligocene

S: platform
C: minor

Berai Mountain
S: shelfal Maid
C: abundant Taballar Cebu
S: shelfal
at margin S: platform C: rare Br. S: shelfal
C: reefal margins C: rare Br. & M.

Batu Rajah
S: shelfal
C: abundant Kujung
& diverse Trankalan
S: shelfal S: platform
C: not known Poh C: Br. corals
Tonasa S: shelfal
S: platform C: few
C: rare Br.

1000 km at equator Irame

S: shelfal
C: no reefs identified
Dari & Puri
S: rimmed shelf
C: not known

Fig.5. Late Oligocene time slice from the reconstructions of Hall (1996) with carbonate deposits shown; abbreviations on Fig.3.

Java and Sumatra, started to become marine, logical literature, they tend to be thin delicate
with the development of carbonates. Carbonates branching forms, or finger-like corals (such as
also became more extensive along the eastern Acropora), although massive framework corals
side of the Sundaland craton due to continued occur in some localities forming patch reefs. The
subsidence in the region. Shelfal and platform Boera Formation of Papua New Guinea, dated
carbonates again dominated, particularly during as uppermost Oligocene to lowest Miocene (Te:
the earlier part of the late Oligocene, although Rogerson et al., 1981), contains a moderate
patch reefs and some reefal buildups may have fauna of z-like corals (Fig.12c,d) which is un-
developed later in the Oligocene. Larger benthic published but well-represented in The Natural
foraminifera, such as lepidocyclinids, hetero- History Museum, London. These are the oldest
steginids and various larger imperforate for- z-like corals known from outcrop on or around
aminifera, were still common (Fig.11). In forma- the Australian cratonic margin, and provide a
tions which pass conformably upwards into the minimum age for the arrival of the craton’s lead-
Miocene, such as those in Java, Sumatra or New ing edge in tropical waters (cf., Davies et al.,
Guinea, shelfal carbonates pass upwards into 1987). Initial identifications suggest affinities
reefal buildups in the upper part of the succes- with European Oligocene faunas. Reefal corals
sion (Cucci and Clark, 1993). However, it is of- are also common in the upper part of the upper
ten difficult to separate upper Oligocene from Oligocene in the Batu Asih limestones in Java
lower Miocene carbonates on the basis of infor- (A. Wonders, pers. comm., 1997). Quite a rich z-
mation in the literature. like fauna is emerging from the uppermost
When corals are mentioned in the sedimento- Oligocene or lowest Miocene Gomantong Lime-
Cenozoic corals of SE Asia 179

Foraminifera dominated carbonates

z-like corals & foraminifera present

z-like corals dominate

Land areas
S: platform
C: abundant at margin

21 Ma early Miocene
S: pinnacles
C: abundant & diverse
Reed bank
S: pinnacles
Belumai/Peutu C: abundant & diverse
S: shelfal buildups
C: Br. & Dig.

S: shelfal
C: Br., Pl. & Fr.
Olodano S: platform
S: shelfal buildups
Kubu C: Ab. & Div.
C: Br., Pl. & Fr. S: shelfal
C: some
S: patch reefs
C: Ab. & Div.
pre-Parigi Berai
S: shelfal Labang Kitcharo
S: shelfal S: shelfal
C: some C: Ab. & Div. Taballar C: abundant
S: shelfal
C: Ab. & Div.
Rajamandala S: shelfal & patch reefs
S: shelfal C: abundant & diverse
C: abundant & diverse Tomori
Prupuh S: shelfal
S: pinnacles C: abun. & div.
C: some Tonasa Salodik
S: isolated platform
C: rare Br.

S: shelfal
Kais C: abun. & div.
S: reefal builups on shelf Dari/Puri
C: abundant & diverse S: rimmed shelf
helf &&pinnacles
1000 km at equator C: abundant framestones
NW shelf
S: rimmed shelf
C: abundant & diverse

Fig.6. Early Miocene time slice from the reconstructions of Hall (1996) with carbonate deposits shown; abbreviations on Fig.3.

stone in Sabah (J. Noad, pers. comm. 1997) with The ages and taxonomy of this rich and his-
17 z-like genera identified so far (work in torically important SE Asian Neogene coral
progress, BRR), many of them previously known fauna are much in need of modern revision, as
from the lower Miocene elsewhere in Borneo. this is the key to more rigorous studies of the
global endemicity patterns and biogeographical
history of corals discussed by Gerth (1925, 1930)
Neogene and Rosen (1988). Nevertheless, such work is
unlikely to alter radically the existing picture of
There was a striking increase in the distribution substantial increase in z-like coral faunas which
and extent of carbonates (Fig.6), and a marked seem to have developed relatively suddenly
change from larger foraminifera to coral domi- throughout the region, from the early Neogene
nated carbonates at the beginning of the onwards. Only a few key points concerning the
Neogene. A distinct change also occurred in the Neogene are made here.
coral faunas during the earliest Miocene, within
Te time. Bemmelen’s (1970) Table 15 lists some
of the key coral faunas of Neogene age, many of Early and middle Miocene (Te5-Tf)
which consist of 30 or more z-like species. They
are far richer and more widespread than Terti- The early Miocene was a major phase of carbon-
ary faunas of pre-Te age in the same region. ate deposition both in SE Asia (Fig.6), and
These patterns are clearly reflected in the ge- throughout much of the tropics and subtropics,
neric diversity patterns of Fig.9. with reef corals occurring in much higher lati-
180 M. E. J. Wilson and B. R. Rosen

Foraminifera dominated
z-like corals & foraminifera present

z-like corals dominate

Land areas

13 Ma middle
Reed bank

S: platform Luconia
C: Ab. & Div.
S: buildups
C: Ab. & Div.

S: shelfal
C: Ab. & Div. Uling
S: shelfal
C: Ab. & Div.
S: shelfal buildups
C: Br., Pl. & Fr. Wahig
S: shelfal
Parigi C: some Fr.
S: platform Tomori
C: some S: shelfal buildups
C: Ab. & Div.

Rajamandala S: isolated platform
S: shelfal C: rare Br.
C: abundant & diverse Puwani/Gowop
S: shelfal
C: abundant & diverse
S: reefal builups on shelf Dari/Puri
C: abundant & diverse S: rimmed shelf & pinnacles
C: abundant framestones

1000 km at equator Great Barrier Reef

NW shelf S: platform & barrier reef
S: rimmed shelf C: abundant & diverse
C: abundant & diverse

Fig.7. Middle Miocene time slice from the reconstructions of Hall (1996) with carbonate deposits shown; abbreviations on Fig.3.

tudes than today, probably as a result of a glo- marked contrast to the earlier Cenozoic. Many
bal warm phase (Fulthorpe and Schlanger, 1989). modern genera, and even species (Umbgrove,
Carbonates were common in marine basins 1946; Rosen, 1984; Veron, 1995), as well as all
around the margins of mainland SE Asia, although the growth forms typical of modern reefs in the
a combination of clastic input and a narrow shelf Indo-West Pacific, occur in these Miocene reefs
restricted carbonate development along the (e.g., Fig.12a,b). The accompanying marine
southeast coast (Fig.6). Carbonate production still biota is also similar to modern reef biotas, with
occurred on microcontinental blocks in eastern SE abundant benthic foraminifera, echinoderms,
Asia (Fig.6), although more islands were emerg- molluscs, coralline and calcareous green algae.
ing due to collision-related uplift. During the mid-
Miocene (Fig.7), the area of carbonate deposition,
though still extensive and diverse, had been re- Comparison with present day
duced, due to the emergence of more land areas,
resulting partly from microcontinental collisions The modern situation (Fig.8; after Wells and
and related shedding of clastic material into adja- Sheppard, 1988) is not unlike the Miocene
cent marine areas. (Figs.6 and 7), although the extent of coral-rich
Platform and reefal carbonates were common carbonate is reduced, especially when com-
throughout the region and pinnacle reefs have pared with the lower Miocene. This is due to the
been reported from many areas (Livingstone et continued emergence of land areas and to a cor-
al., 1992). Z-like corals were widespread, abun- responding influx of clastic material into adja-
dant and diverse throughout the region, in cent coastal waters hindering carbonate produc-
Cenozoic corals of SE Asia 181

Foraminifera dominated
Coral & forams present

Corals dominate

Land areas
Mainland SE Asia

Philippine Sea Present day

South China Sea Philippines


Pacific Ocean



New Guinea
Indian Ocean


Great Barrier Reef

1000 km at equator
90°E 100°E 110°E 120°E 130°E 140°E 150°E 20°S

Fig.8. Present day distribution of shallow water depositional environments (after Wells and Sheppard, 1988); abbreviations on Fig.3.

tion. However, despite this slight reduction in tain fauna of this age being our own new finds
carbonate sedimentation, reefal development from the Tonasa Formation. Oligocene faunas
with z-corals is still common in SE Asian waters, are slightly richer than previously thought. Z-
particularly in the clearer waters of eastern In- coral richness is much higher in other regions in
donesia (Wells and Sheppard, 1988). the Paleogene, and higher in SE Asia from the
As already emphasized, modern z-corals have Neogene onwards. In the Paleogene, facies and
their highest species diversity in SE Asian wa- environments in which z-like corals could have
ters, with the whole range of morphological thrived (mainly shallow water carbonates) were
growth forms present. At least 262 species and sparse, and SE Asia was really a wide oceanic
78 genera have been recorded in Sulawesi seaway in which the dense arrangement of is-
(Moll, 1983), and Veron (1995) recorded 472 lands of the modern region did not yet exist.
species in the northern Philippines. Coral reefs There was a fundamental change at or around the
are abundant, and, of the estimated 600,000 km2 beginning of the Neogene (i.e., within Te time),
of these complex ecosystems worldwide, 25- coincident with the beginning of interaction be-
30% are located in SE Asia (Bernard, 1991). tween Australian fragments and SE Asia, leading
to a notable increase in shallow water carbonates,
widespread reefs, high z-coral abundance and
Summary of geology and coral faunas in SE Asia fourfold increase in z-coral richness.
Tectonics controlled the emergence, disap-
Eocene z-like coral faunas appear to be even pearance, and movement in and out of the z-
less rich than previously thought, the only cer- coral belt, of shallow marine areas, as well as
182 M. E. J. Wilson and B. R. Rosen

their regional concentration. In the Paleogene Tertiary Tethyan counterparts (e.g., Mediterra-
however, even where shallow water carbonates nean), though the Indian Ocean (with Red Sea)
did exist, they often lacked z-coral faunas, sug- is also sometimes considered separately (see
gesting that other factors (biogeography, local Veron, 1995; Flessa and Jablonski, 1996; Budd
ecology) were also important. In contrast to SE et al., 1996; Paulay, 1997; for reviews of these
Asia, there are extensive Paleogene (especially regions). We concentrate on the ideas shown in
upper Eocene) limestones in the tropical west- Table 1 for the Indo-West Pacific centre, particu-
ern Pacific, but they are mostly in the subsurface larly the extent to which patterns may have been
and have remained largely unsampled. The few affected by plate tectonics, though the influence
limestones that have been sampled contain ei- of eustasy (not discussed here) was also impor-
ther z-like or az-like coral faunas. However, tant, especially in the later Cenozoic (Schlanger,
many of these limestones lay further east than 1981; Rosen, 1984, 1988; Grigg, 1988; Veron,
now, representing perhaps a distinct region. 1995). The emphasis here on physical factors
The Cenozoic record of z-corals points to the differs from models which emphasise intrinsic
modern Indo-West Pacific centre of diversity be- evolutionary processes sustained in one region
ing a young feature, probably little older than over great lengths of time (Centres of Origin).
around the beginning of the Neogene (c.24 Ma). Flessa and Jablonski (1996) have remarked
that Rosen’s (1984,1988) physical contingency
model “implies that intensive sampling of the
Discussion Eocene, prior to global refrigeration and the
northward migration of the Australian plate,
Global biogeographical context would detect neither diversity gradient, nor age
gradient, but perhaps only the legacy of the evo-
The most challenging biogeographical problem lutionary rebound from the end-Cretaceous ex-
of the Indo-West Pacific centre is to understand tinction”. Their caveat that “few data are avail-
the origins of its marked taxonomic richness, able to test (this) idea of ... young age” reflects
models for which are shown in Table 1. This can in part the previous emphasis on age-and-area
be partly attributed to a global pattern in which studies of extant taxa. We address Flessa and
richness of numerous marine taxa increases as Jablonski’s prediction below.
latitude decreases, related to available energy In modelling the biogeography of a particular
(Stehli, 1968; Jablonski, 1993; Fraser and Currie, region, three sets of factors need to be consid-
1996; Flessa and Jablonski, 1996; Paulay, 1997). ered (Rosen, 1988): (1) originations (patterns of
This has long been known for z-corals (Yonge, evolutionary turnover); (2) distributional change
1940; Wells, 1954a), and more recently demon- (often also called ‘migrations’, ‘dispersal’ etc.,
strated quantitatively (Rosen, 1971, 1984; Fraser and meaning changes through time of biogeo-
and Currie, 1996). There is also empirical evi- graphical ranges of taxa, combined with
dence that more organisms originate in the trop- changes of biogeographical barriers); and (3)
ics than elsewhere (Jablonski, 1993; Flessa and maintenance (the sum of local-to-regional eco-
Jablonski, 1996). Temperature however cannot logical factors). This framework is used to ad-
account for the entire global pattern. Z-corals, dress the Cenozoic history of z-like corals in the
for example, are four times richer in the Indo- Indo-West Pacific and to discuss criteria for the
West Pacific centre than in the western Atlantic alternative models for the high diversity foci
(Figs.1 and 9), but the prevailing temperatures (Table 1). In particular, since the main conclu-
in both are similar (Rosen, 1984), and this rich- sion of our stratigraphic review points to high
ness ratio has remained stable over decades of diversity in the SE Asian part of the Indo-West
sampling and taxonomic study. Pacific centre being surprisingly young, we con-
In order to consider regional anomalies, and centrate on possible reasons for this.
the Indo-West Pacific centre in particular, it is
useful to resolve diversity (and endemicity) pat-
terns into two geographical components, ‘latitu- Maintenance factors
dinal’ (N-S) and ‘longitudinal’ (E-W) (Rosen,
1975). The strongest endemicity patterns are Although the paucity of Paleogene z-corals in SE
longitudinal. Four regions have dominated lon- Asia can be largely explained by sparseness of
gitudinal discussion (Gerth, 1925, 1930; Paulay, inhabitable areas (i.e., shallow water carbon-
1997): the Indo-West Pacific, the eastern Pacific, ates) across a wide seaway, this does not ac-
western Atlantic and eastern Atlantic, and their count for the enigmatic absence of z-corals from
Cenozoic corals of SE Asia 183

most of the few shallow water carbonates that Distributional change

did exist. Larger benthic foraminifera are usu-
ally the dominant constituents in these carbon- The factors of distributional change most rel-
ates, and today they inhabit very similar shal- evant to the models in Table 1 are immigration
low-marine tropical environments to corals. rates. Low rates may have contributed to the
Temperature and latitudinal effects cannot ex- Paleogene paucity of z-corals in SE Asia, and
plain the lack of corals, especially since these higher immigration rates to the subsequent
Paleogene habitats clearly lay within the global surge in their abundance and diversity in the
limits of the z-coral belt, and climatic conditions Neogene. There are as yet few data for address-
were particularly warm in the middle and late ing this, but Rosen and Smith (1988) used an in-
Eocene (Miller et al., 1987; Frakes et al., 1992). direct approach, analysing the history of ende-
Rapid subsidence rates, dispersal and recruit- micity patterns of corals and echinoids from dif-
ment factors, limiting chemical conditions of the ferent time slices. In the middle to late Eocene,
water, including salinity, oxygen, and nutrients, and in the early Miocene, Pacific and Australa-
and unfavourable speeds and directions of cur- sian faunas were relatively isolated from
rents, might all have been contributory causes, Tethyan and Indian Ocean faunas, and a pan-
but cannot be assessed easily in an ancient con- Indo-Pacific fauna in the modern sense did not
text. exist until these more remote elements became
Substrate control is more readily inferred, fully integrated during or after the early
since some of the foraminifera-dominated car- Miocene.
bonates represent shifting coarse-grained Further support for this comes from the tec-
shoals (Fig.11d), perhaps comparable with ooid tonic evolution of SE Asia and neighbouring re-
shoals today, and hence unfavourable for coral gions (Schlanger, 1981; Rosen and Smith, 1988;
colonisation. Water depth is also an important Rosen, 1984, 1988; Paulay, 1997). Paleogene
factor, although the taxa and growth forms of paucity of z-corals in SE Asia may reflect partial
the larger benthic foraminifera and other shal- isolation of its carbonate environments from z-
low marine bioclasts, such as coralline algae, coral areas elsewhere, though the existence of
within the Paleogene limestones studied, indi- modern examples of rich reefs in isolated set-
cate a range of water depths within the photic tings (e.g., Chagos) suggests that the factors
zone (Adams, 1970; Ghose, 1977; Hottinger, here may be more complex. Australia was not a
1983; Hallock and Glenn, 1986). Some of the reefal region as it is today, and lay over several
limestones dominated by thin flat forms of thousand kilometres away to the southeast of
larger benthic foraminifera were probably de- the Asian margins, and outside the z-coral belt.
posited below the depth of the most prolific z- To the east, on the Pacific plate, the Paleogene
coral growth (generally < 100 m, Fig.11e). How- forerunners of the modern mid-Pacific archi-
ever, in other deposits, the taxa and growth pelagos lay much further to the east than now.
forms of foraminifera, which show little signs of Apart from the Hawaii-Emperor chain, most of
reworking, indicate deposition in the upper part these archipelagos are broadly confined to the
of the photic zone (Fig.11a-d). Perhaps coral limits of the Darwin Rise/Superswell. Mid-plate
facies were once more widespread but have oceanic islands are otherwise unusual features,
been largely destroyed, though there are too and few suitable areas for corals may have oc-
few deposits with abundant coral clasts to sup- curred between the Darwin Rise/Superswell
port this. and the western Pacific convergence zones in
Whatever the proximal causes, secondary the area which has been lost due to subduction.
feedback effects would also have been impor- Thus, during the Paleogene, this intervening oce-
tant if coral communities were too scattered in anic region may have been wide enough to have
time and space to sustain frequent larval ex- caused partial or intermittent isolation of Pacific
change and recruitment between areas within areas now represented by the Eocene coral-bear-
the region. This would have led to high rates of ing limestones underlying these islands.
local extinction, and prevented longer-term de- To the west of SE Asia, the collision of the
velopment of z-coral facies. While the Paleo- Indian continent with Eurasia in the Eocene
gene paucity of z-corals in SE Asia appears to be eliminated the east-west trending Tethys sea-
a real phenomenon linked to lack of habitable way in that region, and initiated large-scale clas-
areas, no clear set of ecological factors yet ex- tic input along the surrounding coastlines. In
plains the paucity of z-corals in the few areas the tropical waters of the Indian Ocean, the few
where they might be expected. oceanic island ‘staging posts’ (Rosen, 1983),
184 M. E. J. Wilson and B. R. Rosen

(Z & AZ) (Z) (Z) (Z) (Z) (Z)
ca 1400 spp.
Modern fauna
Mess ?
MIO- Serray
CENE Langh
OLIGO- Chatt
CENE Rupel + +
ENE Lutet + + ?

Thanet ?
PALEO- Seland + + + + + +

10 Ma 100 100 100 100 100 100 100

species genera genera genera genera genera genera

Fig.9. Taxonomic richness of zooxanthellate corals from Paleocene to present day. Z - zooxanthellate and zooxanthellate-like
corals. AZ - azooxanthellate and azooxanthellate-like corals. I-WP - Indo-West Pacific. + symbols indicate interim compilation
where faunas are actually richer than shown. Where data have been compiled according to Table 2, inner shaded blocks
represent minimum estimate of z-like corals and unshaded outer blocks represent maximum estimates. See Rosen (1988) for
discussion of time-averaging of coral data. Data sources follow. GLOBAL SPECIES; GLOBAL GENERA (Z and AZ). Rosen (1998)
and sources therein. GLOBAL GENERA (Z). Eocene, Oligocene, Plio-Pleistocene and modern fauna from Veron (1995, Fig.38).
These totals were independently compiled by Veron and are not summations of regional values elsewhere in this diagram.
Miocene data from Rosen (1988, Table 2, from various sources); not adjusted for newer data incorporated in regional values
elsewhere in this diagram. CARIBBEAN GENERA i.e., Caribbean islands with adjacent mainlands of North, Central and South
America. Paleocene from Bryan (1991). Eocene: compilation of middle to upper Eocene reef-coral genera from Budd et al. (1992,
Table 1). Oligocene from Rosen (1988, Table 2, from various sources) but needs revision. Miocene to modern fauna from Budd
et al. (1994, Fig.5). MEDITERRANEAN GENERA (i.e., Mediterranean islands, and adjacent mainland mostly southern Europe and
Egypt). Paleocene: partial compilation from Rosen (1998, Table 2 and sources therein). Eocene: partial compilation for a single
‘sample region’ of southern Europe (Bosnia, Hercegovina, Croatia and ‘Dalmatia’) from Felix (1925). Inclusion of other rich
southern European faunas (e.g., Spain, France, Italy) would show Mediterranean as the richest Eocene region. Oligocene and
Plio-Pleistocene from Rosen (1988, Table 2, from various sources). Miocene from Rosen, unpublished. Modern fauna from
Zibrowius (1980, and pers. comm., April 1997). SOUTHWEST ASIAN GENERA i.e., Middle East, Iran, Pakistan, western India.
Paleocene based on Duncan (1880) and Gregory (1930) for Ranikot fauna. Lower Miocene from McCall et al. (1994), and
remainder from Rosen (1988, Table 2, from various sources). Eocene to Pleistocene faunas are rich in this region but await further
research and compilation (the same applies to the rich adjacent areas of the eastern margins of Africa). INDO-WEST PACIFIC
CENTRE GENERA i.e., SE Asia, northern Great Barrier Reef and islands of western and mid-Pacific). Paleocene: this paper (Table
3); also unidentified ‘coral fragments’ known from boreholes. Eocene and Oligocene: this paper (Table 3). Miocene and modern
fauna from Rosen (1988, Table 2, from various sources). Note that many ‘lower Miocene’ localities are of undifferentiated Te age
and may therefore include uppermost Oligocene (Te1-4) corals, but lack adequate stratigraphical detail.

such as the Ninetyeast Ridge, may have aided area during the Paleogene. This did not com-
only intermittent larval exchange of benthic pletely prevent colonisation by z-corals since
populations between SE Asia and the rich the biogeographical affinities of the Tonasa
Eocene z-like coral regions on the western side fauna indicate that biogeographical continuity
of India, the Middle East and Europe (Fig.9). between SE Asia (Sulawesi) and the western
These palaeogeographical factors, combined side of the Indian continent occurred at least
with the great width of the Indo-Pacific gateway once, perhaps via the Ninetyeast Ridge. The
between the Asian and Australian continents, faunal affinities of the contemporaneous
and scarcity of shallow areas in that gateway, Enewetak z-corals (Wells, 1964) suggest an
must have limited immigration into this gateway analogous pattern for mid-plate Pacific areas.
Cenozoic corals of SE Asia 185

The broadly contemporaneous collision of Their affinities (above) cannot rule out SE Asian,
Australia with SE Asia and closure of Tethys in European or Indian Ocean origins, although
the Middle East during the latest Oligocene to given the paucity of this fauna there is therefore
earliest Miocene (Rosen and Smith, 1988; McCall no evidence for high endemic origination rates
et al., 1994, for the coral history) had a profound in SE Asia at this time. The Oligocene picture is
effect on the palaeogeographic setting of z-coral less clear for taxonomic reasons. The main fauna
faunas in SE Asia. Taxa within the Indo-West (Tanjung; Table 3), comprises a mixture of Euro-
Pacific centre from this time onward must have pean and apparently endemic taxa. Our initial
consisted of (1) pre-Neogene, but not necessar- work on the Boera corals of Papua New Guinea
ily Indo-Pacific, relicts, (2) those of local SE (Fig.12), reveals strong European affinities, even
Asian origin and (3) taxa that originated else- though they are dated uppermost Oligocene
where within the Indo-West Pacific but which (Table 3). For a more complete picture how-
immigrated into the centre (see below). It is un- ever, revision and/or description is urgently
likely any other taxa that had originated else- needed of the important Paleogene faunas of
where could have then entered the Indo-West Pakistan (Duncan, 1880; Gregory, 1930), to-
Pacific. Meanwhile, northern Australia and New gether with those of the western Indian Ocean
Guinea had entered the reef coral belt, and the margins from Iran and the Arabian region, to
Pacific archipelagos of the Darwin Rise/ eastern Africa. Currently, however, there is no
Superswell had moved closer to Asia by over reason to change Gerth’s (1925, 1930) view that
3000 km (CGMW/UNESCO, 1990) since the late relations between the corals of SE Asia during
Eocene. These latter events must have greatly the Paleogene generally, were closer to that of
increased the extent and variety of potential the Indian and Mediterranean regions, than they
habitats for z-coral colonisation in SE Asia. Dur- were during the Neogene, and that exchange of
ing the Neogene, SE Asia became, in effect, a species took place throughout these three re-
biogeographical cross-roads and meeting point gions during the Paleogene (see also Pandolfi,
for faunal convergence of shallow marine or- 1992). Hence palaeontological evidence of origi-
ganisms from numerous different directions nation patterns give no support for SE Asia be-
(Rosen, 1988; Pandolfi, 1992) perhaps enhanced ing a Centre of Origin (or part of one) during the
by changes in oceanic current configurations at Paleogene.
this time (Grigg, 1988). The fourfold surge in z-coral diversity in the
Indo-West Pacific centre from around the early
Neogene (Fig.9) must reflect, at least in part, the
Originations dramatic geological changes. Severance of
Tethyan seaways in the Middle East at around
Rates and locations of originations are crucial this time would have confined subsequent
criteria of the different models in Table 1. Rigor- originations either to the Atlantic-Mediterranean
ous treatment however requires good knowl- or to the Indo-Pacific, respectively, and this is
edge of the phylogenetic relationships of taxa to reflected in their well-established post-Paleo-
each other and to their times and places of oc- gene endemicities. Each region became an evo-
currences. However, such work is currently con- lutionary entity, with the Indo-Pacific exhibiting
fined to studies of a few extant Indo-Pacific z- the more marked taxonomic radiation (Fig.9).
corals, and until now fossil data have not been Independent Neogene evolution in the western
incorporated. Stratigraphic distributions also of- Atlantic is shown by Agaricia species, and
fer important historical clues, but since the fossil amongst such groups as the meandrinids,
record is incomplete and there is insufficient mussids and meandroid faviids (Budd et al.,
taxonomic knowledge, they mainly provide in- 1994; Johnson, 1998). The important question is
dications of the minimum limits of former distri- whether the Indo-Pacific radiation was particu-
butions in time and space and evidence of larly confined to the Indo-West Pacific centre.
changing patterns of faunal affinities. A third The oldest record of the ecologically impor-
approach, age-and-area studies of modern taxa, tant and very diverse modern z-coral, Acropora,
has until recently dominated Indo-Pacific work. is in the Paleocene of Somalia (Carbone et al.,
Stratigraphic distributions, age-and-area studies 1994). It occurs in numerous European localities
and cladistic biogeography are discussed in this from the Eocene to Miocene, but it does not ap-
order, below. pear in either the Caribbean or the Indo-West
As emphasized already, endemic Eocene z- Pacific centre until the Oligocene (Fritsch, 1878;
corals have yet to be recorded from SE Asia. Wells, 1964; Frost, 1972; Budd et al., 1992, 1994,
186 M. E. J. Wilson and B. R. Rosen

and Fig.10l). It shows a very recent (latest are flawed because they give the illusion that
Miocene or Pliocene) marked radiation in the the age of a modern taxon found in a particular
Indo-West Pacific but not in the Caribbean area indicates that it has been in that area
(McCall et al., 1994; Budd et al., 1994). This his- throughout that time. Admittedly, Stehli and
tory has parallels with the fungiids whose most Wells did not claim this, but by plotting average
primitive genus Cycloseris, occurs in the Paleo- ages for different areas on global maps, they
cene of Somalia and western Pakistan, and in provided a visually powerful synthesis which
the Eocene (Tonasa) of SE Asia (Fig.10b,c) and compounded the illusion, leading others to in-
Europe (Hoeksema, 1989; Carbone et al., 1994). terpret such analyses in this way. For example,
Subsequent radiation occurred during the of the 18 extant Indonesian z-genera which are
Neogene in SE Asia and fungiids are now richest of Eocene age (Rosen, 1984), only 6 actually oc-
in the Indo-West Pacific centre, though in con- cur in the Eocene of SE Asia, and a further one
trast to Acropora they are not known from the (Astrocoenia) is extinct (present data). None of
Caribbean. These two cases show that the Indo- the genera appear to have been local Eocene
West Pacific centre has not been the location of endemics, so the bulk of the 18 corals must have
their origins, although it has been important in arrived in the region from elsewhere. Although
the evolutionary history of z-corals from the this line of argument assumes that the fossil
Neogene onwards. Even so, it is still question- record is reliable (cf., Briggs, 1992), it is equally
able on other grounds (see below) whether or clear that the record does not support a Centre
not the region then acted as a Centre of Origin of Origin in SE Asia at this time. Parallel argu-
for these corals from that time. ments apply to Eocene corals of the Pacific.
Perpetuation of the idea that the Indo-West Age-and-area information alone cannot be
Pacific focus was a Centre of Origin for z-corals used to infer origination patterns, but is best
can be largely attributed to the landmark age- combined with phylogenetic work as in cladistic
and-area study by Stehli and Wells (1971). In biogeography. Paradoxically though, Stehli and
this particular context, however, such analyses Wells (1971), used their age-and-area data to

Fig.10. Representative fossil scleractinian corals of Eocene and Oligocene age from the Tonasa Formation in South Sulawesi.
Specimen numbers, prefixed AZ, are those of The Natural History Museum, London. Details of inferred zooxanthellate status
are also given. Ages of Pakistan occurrences revised according to Adams (1970). So far, the upper Eocene fauna from the
Tonasa Formation consists of 12 species, 7 of which are z-like and 5 are z-like? In addition to those illustrated here, are: fungiid
sp. (z-like), Stylocoenia sp. (z-like?), and a solitary or phaceloid species (z-like?) A-C: Basal beds of the Tonasa Formation,
lowest upper Eocene (Tb), Doi doi locality. A: Trochocyathus ? nummiformis (Duncan, 1880). Transverse thin section, x1.5 (AZ
1188). Patellate to discoidal solitary coral. Modern Trochocyathus is azooxanthellate, but Tonasa specimens and Duncan’s coral
may not belong to this genus. Present species is z-like? on facies occurrence only; previously known from lower to mid-
Oligocene (Tc-d) of Pakistan. B, C: Cycloseris (Cyclolites) crenulata (Duncan, 1880) syn?: Funginellastraea (Cyclolites) heberti
(Tournouer, 1872). B: aboral surface, x1.5, C: polished transverse section, x1.5 (AZ 1202). Discoidal solitary ‘mushroom coral’.
Modern genus is a soft-substrate dweller, usually but not always zooxanthellate. Present species is z-like? on facies occurrence
only; previously known from upper Paleocene of Pakistan (Ta1) and (?) various Eocene localities in southern Europe (F. heberti).
D-K: Bulo Bunting locality, lower part of Tonasa Formation, but stratigraphically above Doi doi locality, upper Eocene (Tb). D:
Trachyphyllia (Montlivaltia) indica (Duncan, 1880). Transverse thin section, x1.2 (AZ 1182). Flabelloid pedunculate form,
consistently four-lobed in mature examples. Modern genus is a zooxanthellate soft-substrate dweller. Present species is z-like on
generic and morphological grounds; previously known from middle Eocene (Ta3) of Pakistan. E: Caulastrea (Calamophyllia)
indica (Duncan, 1880). Transverse thin section, x2 (AZ 1179). Phaceloid branching colony. Modern genus is zooxanthellate.
Present species is z-like on generic grounds; previously known from middle Eocene (Ta3) of Pakistan. F: Favites sp. Transverse
thin section, x2 (AZ 1193). Cerioid colony. Modern genus is zooxanthellate. Present form is z-like on generic grounds. G:
Diploastraea ? cf. Rhizangia? agglomerata Fritsch, 1878. Transverse thin section, x2 (AZ 1191). Astreoid colony. Modern genus is
zooxanthellate. Present species is z-like on generic and morphological grounds; previously known from mid-Oligocene (Td) of
Kalimantan (Table 3). H: Favia (Phyllocoenia) conferta (Duncan, 1864). Transverse thin section, x4 (AZ 1178). Plocoid colony.
Modern genus is zooxanthellate. Present species is z-like on generic and morphological grounds; previously known from
uncertain Tertiary horizon in Pakistan, possibly upper Oligocene to middle Miocene (Te - lower Tf). I: Astrocoenia bistellata
(Catullo, 1856). Transverse thin section, x6 (AZ 1177). Cerioid colony. Extinct genus. Present species is z-like? on facies
occurrence only; previously known from Eocene of Italy. J, K: Favia pedunculata Duncan, 1880. J: (AZ 1192) weathered calical
surface, x0.5. K: (AZ 1187) Transverse thin section, x2. Plocoid pedunculate colony. Modern genus is zooxanthellate. Present
species is z-like on generic and morphological grounds; previously known from middle Eocene to lower Oligocene (Ta3-Td) of
Pakistan. L: Tonasa-II Quarry locality, upper Oligocene (Te1-4) middle part of the Tonasa Formation. Acropora sp. Polished
longitudinal section through ramose branch, x2 (AZ 1213). Ramose branching colony with plocoid corallites. Modern genus is
zooxanthellate, widespread, and an abundant reef contributor in both Indo-West Pacific and Caribbean. Present form is z-like on
generic and morphological grounds.
Cenozoic corals of SE Asia 187




188 M. E. J. Wilson and B. R. Rosen


Fig.11. Thin section photomicrographs of Tertiary limestones dominated by larger benthic foraminifera and deposited in different
environments in the photic zone. All were photographed in plane polarized light at x10 magnification. Samples and thin sections
are held at Royal Holloway London University. A: Imperforate foraminifera bioclastic packstone (MTR114) containing soritiids,
miliolids, alveolinids (Borelis sp.), Austrotrillina sp. and other fragmented bioclasts. Oligo-Miocene sample from the Taballar
Limestone, Taballar River, Mangkalihat peninsula, East Kalimantan. The foraminifera are all typical of the shallower parts of the photic
zone and can tolerate some fluctuations in salinity. B: Probable lower Oligocene (Tc) Nummulites bioclastic pack/grainstone (GU6)
from Gua Unguk, Bengalon River area, north Kutai, East Kalimantan. Contains robust Nummulites and other fragmented and often
abraded marine bioclasts and is inferred to have been deposited in moderate to high energy in the upper part of the photic zone.
C: Lower Oligocene (Tc) Nummulites and Heterostegina bioclastic packstone (GB43) from Gua Bangki, Bengalon River area, north
Kutai, East Kalimantan. Contains moderately robust Nummulites and thin forms of Heterostegina in a micritic matrix inferred to have
been deposited under moderate to low energy conditions in moderate depths in the photic zone. D: upper Eocene (Tb) Discocyclina
and Biplanispira bioclastic packstone (BR81) from the Bengalon River area, north Kutai, East Kalimantan. Contains abundant whole
and sometimes fragmented larger benthic foraminifera, including Discocyclina sp., Biplanispira sp., Pellatispira sp., Nummulites sp.
and other fragmented marine bioclasts such as coralline algae and echinoid plates. The species of foraminifera, their growth forms and
the nature of the surrounding sediment suggests deposition in a moderate to high energy setting in shallow to moderate depths in
the photic zone, possibly as a foraminiferal shoal. E: Upper Oligocene (Td/Te1-4) Lepidocyclina and heterosteginid packstone
(MTR100) from the Taballar Limestone, Taballar River, Mangkalihat peninsula, East Kalimantan. Includes Eulepidina sp.,
Nephrolepidina sp., Heterostegina sp. and possible Spiroclypeus sp. Large, thin forms of these species of larger foraminifera are
typical of development towards the base of the photic zone.
Cenozoic corals of SE Asia 189


Fig.12. Representative fossil scleractinian corals from the upper Miocene part of the Tacipi Formation, Sungai Pesing,
Ujunglamuru, south Sulawesi (A-B), and from the uppermost Oligocene to lowest Miocene (Te1-5) Boera Formation, Boera
Head, south Papua (C-D; see Table 3). A: Hydnophora astraeoides (Martin, 1880). Calical surface, x2 (AZ 1212). Continuously
meandroid colony with vestigial walls reduced to isolated monticules. Modern genus is zooxanthellate. Present species is z-like
on generic and morphological grounds; previously known from Miocene of Indonesia. B: Platygyra daedalea (Ellis and
Solander, 1786). Calical surface, x2 (AZ 1211). Meandroid colony. Modern species is zooxanthellate, so same is inferred for
present occurrence. Extant, widespread and important reef contributor, previously recorded from numerous fossil and modern
Indo-West Pacific localities from Miocene onward. C: Favites [Prionastraea] borneensis (Gerth, 1923). Polished transverse
section, x3 (AZ 1209). Cerioid colony. Modern genus is zooxanthellate. Present species z-like on generic grounds; previously
known from Neogene of Indonesia. D: Astreopora cf. meneghiniana (D’Achiardi, 1866). Polished transverse section, x2 (AZ
1210). Plocoid colony. Modern genus is zooxanthellate. Present species z-like on generic and morphological grounds; previ-
ously known from Oligocene of Italy.

support the widely held notion that Centres of in the Indo-West Pacific centre, and all but two
Origin have an intrinsic evolutionary character- of them still occur in the modern Indo-West Pa-
istic in which continuous emergence of more cific centre. Therefore most of these genera were
successful taxa pushes older taxa out from the not displaced from the centre, or if they were,
focal region. This predicts a pattern of continu- they evidently returned. This, as well as the
ous evolutionary turnover, in which (1) older paucity of the fauna is incompatible with Centre
faunas, prior to their displacement, should of Origin notions.
share the same central origin, and be more or Moreover, sympatric speciation modes aside,
less as rich, as younger faunas, and (2) a given the very commonly observed overlapping pat-
fauna of one time slice should comprise numer- terns of sibling species (and sister genera)
ous taxa that were subsequently displaced. amongst modern z-corals species in the Indo-
There is as yet no evidence however that Pacific, especially the central region (e.g.,
Tonasa and Enewetak Eocene genera originated Hoeksema, 1989, 1993; Veron, 1995; Wallace et
190 M. E. J. Wilson and B. R. Rosen

al., 1991; Wallace, 1998; Pandolfi, 1992) suggest such as Sibumasu and Indochina in the north,
that siblings must have originated in different from those Gondwana blocks to the south
areas and subsequently migrated into their ob- which would later form Borneo, Sulawesi, Nusa
served overlapping distributions (see also Tenggara and Australia (Metcalfe, 1996).
Paulay, 1997). Hence, whatever the evolution- Around the margins of Meso-Tethys, z-coral
ary characteristics of Centres of Origin, competi- facies developed in shallow marine waters and
tive displacement, in this case, does not appear have been preserved today in Upper Triassic
to be one of them. Since average longevity esti- (Stanley, 1988) and Upper Jurassic limestones in
mates for z-coral species are about 10 Ma Burma, Thailand and eastern Indonesia. Rifting
(Veron, 1995), many of these numerous overlap- during the Jurassic and Cretaceous separated
ping species must be post-Paleogene in origin, blocks, such as west Burma and west Sulawesi,
suggesting that distribution patterns in and from the northern part of the Australian craton
around the centre are predominantly Neogene by a seaway. Australia lay too far south of the z-
or younger. This is again incompatible with a coral belt to support coral growth. In contrast,
Centre of Origin model, though it is also likely the rifted blocks were drifting northwards
that some species originated through geographi- across tropical latitudes to be juxtaposed
cal isolation within and around the Indonesian against mainland SE Asia during the Jurassic to
archipelago (Pandolfi, 1992; Wallace, 1998). Late Cretaceous and the few known Cretaceous
Evidence for heterogeneous geographical ori- scleractinian corals from Borneo and Sumatra
gins of Indo-Pacific z-corals, including periph- reflect this. However, as with the Paleogene, z-
eral sources, has come not just from their over- corals are rare in the rest of the region during
lapping distribution patterns but also from the Cretaceous. Thus the ‘Paleogene data gap’
cladistic studies of the fungiids (Hoeksema, for z-corals is the continuation of an earlier pat-
1993), the siderastreids and mussids (Pandolfi, tern which goes back into the Cretaceous, and
1992), and some species of Acropora (Wallace et presumably had similar causes.
al., 1991). These studies do not include esti- The underlying issue of interest is the relative
mates of ages of origination from fossil evidence importance of plate tectonics as an evolutionary
but Pandolfi infers these from the timing of geo- influence compared with intrinsic evolutionary
logical events whose associated features corre- processes. In general, development and diver-
spond geographically to his area cladograms. sity of both Mesozoic and Cenozoic
Parallel studies in other groups (cf., Pandolfi, scleractinian z-corals in SE Asia have been ulti-
1992; Palumbi, 1996; Paulay, 1997) give similar mately controlled by the plate tectonics of the
indications of widespread heterogeneous ori- region. For the Paleogene in particular, Flessa
gins for the common marine organisms of the and Jablonski (1996) gave three criteria (above)
Indo-West Pacific centre. Reef fishes, for exam- by which plate tectonics might be judged to be
ple, apparently evolved rapidly during the first of greater importance than intrinsic evolution-
20 Ma or so of the Cenozoic and then remained ary factors in the Paleogene history of SE Asian
in relative stasis (Choat and Bellwood, 1991; coral patterns. Firstly, the lack of latitudinal di-
Bellwood 1996). Cladistic analysis of reefal versity gradients is effectively fulfilled in our
parrotfish (Scaridae) by Bellwood shows that study because the zooxanthellate occurrences
the labroid lineage which gave rise to this are so few that there is no gradient to speak of.
group, was already present by the latest early to Secondly, the lack of endemics of Eocene age in
earliest middle Eocene of Europe (i.e., most re- particular, and the European-Indian affinities of
cent date of Monte Bolca fish fauna from J. SE Asian Paleo-gene corals in general, suggest,
Pignatti, pers. comm., 1997), a pattern that has as predicted, that their average age cannot be
parallels with Acropora and the fungiids. much younger, if at all, than that of corals in
these other regions.
Thirdly, Flessa and Jablonski predict that re-
Plate tectonics and the implications of the covery from the K-T extinction would also have
Mesozoic coral record been a significant influence on SE Asian distri-
butions if plate tectonics were important. In fa
It is instructive to view the Cenozoic coral t, the paucity of both Cretaceous and Paleogene
record in the longer-term context of the coral records within the region means that K-T
Mesozoic coral record and its plate tectonic set- extinctions and recovery were largely witnessed
ting. During the Triassic and Jurassic, the large elsewhere. Extensive development of z-coral
tropical ocean of Meso-Tethys separated blocks faunas and communities did not really occur in
Cenozoic corals of SE Asia 191

SE Asia until around the earliest Neogene, the region. Local originations derived, at least in
whereas elsewhere in the tropics its earliest part, from these antecedent elements. The geo-
signs begin only 3-5 Ma into the Paleocene, with graphical complexity of the region since the
quite rich faunas established by the Thanetian Neogene favoured all three processes through
(Fig.9; Schuster, 1996; Rosen, 1998). Thus while increasing habitat heterogeneity and potential
Flessa and Jablonski’s third prediction is not ful- allopatric speciation amongst fragmented shal-
filled in their precise terms, the absence of K-T low water areas, further enhanced in the last 10
coral evidence in SE Asia, clearly reflects the in- Ma or so by the effects of glacio-eustasy and in-
fluence of plate tectonics. This also suggests that creased climatic fluctuation.
the idea that the Indo-West Pacific centre is Our fossil patterns show that a Centre of Ori-
largely a vestige or derivation of Cretaceous pat- gin model (Table 1) is completely inapplicable
terns (McCoy and Heck, 1976; Table 1) is at best to Paleogene z-corals of the SE Asian region,
an oversimplification. More generally therefore, since there was only a small non-endemic fauna
there is no need to suppose that SE Asia was here. In fact, it is misleading to think of it as a
endowed with special evolutionary processes, ‘centre’ of any kind during this time. This con-
as Centres of Origin models imply. It has been a trasts with the Neogene onwards, which super-
region in which the potential to develop rich ficially accords with a Centre of Origin model, -
communities has been controlled by the way in albeit of very short duration compared with s-
which suitable areas for them have been con- pecies turnover timescales. Moreover, the c-
tinually created, reorganized and lost, espe- ombined patterns from a range of different stud-
cially according to the timing and movements of ies show that a more appropriate model re-
blocks which have broken away from quires a combination of all three of the possi-
Gondwana, their northward passage across bilities in Table 1. A longer-term perspective
Tethys into tropical waters, and their eventual suggests that the above contrast in Paleogene
accretion to the Eurasian continent. and post-Paleogene patterns represents a cycli-
cal sequence which has occurred at least twice
before with strong z-coral developments in the
Plate tectonics or Centre of Origin? late Triassic and late Jurassic in SE Asia, coinci-
dent with times when rifted Gondwanan blocks
Two contrasting major patterns can be dis- docked against the Asian continent in the trop-
cerned: a Paleogene one in which diversity, en- ics. Thus plate tectonic processes rather than in-
demicity and origination rates of z-corals in SE trinsic evolutionary processes (like Centres of
Asia were low, and a post-Paleogene one which Origin, competitive displacement, etc.) have
is effectively the opposite. The change coincides been a major control on regional diversity pat-
with tectonic events, especially the collision of terns of z-corals, and presumably also numer-
Australia with SE Asia. During the Paleogene, ous other shallow marine organisms.
the marine region between the continental mar-
gins of SE Asia and Australia was about 3000 km
wide and, compared with the post-Neogene, Acknowledgements
there were few shallow water areas suitable for
z-corals. A much more extensive, but now The authors would like to acknowledge the fi-
largely inaccessible area of shallow water car- nancial and technical support provided by a
bonates existed in the mid-Pacific (on the Dar- number of groups during the course of this re-
win Rise/Superswell) especially during the search: namely London University SE Asia Re-
Eocene. However, the z-coral faunas of both re- search Group and its consortium of funding
gions appear to have been low diversity out- companies, The Natural History Museum, Lon-
posts of the very much richer faunas of Europe, don, and the Geological Research and Develop-
eastern Tethys and the western Indian Ocean ment Centre in Indonesia. We thank the follow-
margins. ing for helpful discussions or correspondence
The modern high diversity pattern in SE Asia on numerous aspects of the paper: Dan
and the Indo-West Pacific centre began to Bosence, Jill Darrell, Robert Hall, Alit Ascaria,
emerge around the earliest Neogene with an Dick Randall, Galt Siegrist and Carden Wallace.
apparent regional radiation of z-corals. How- Robert Hall is gratefully acknowledged for al-
ever, detailed studies are revealing that relicts, lowing us to use time slices of his plate tectonic
and migrations of taxa into the region were both models as base maps. Jill Darrell also provided
as qualitatively important as originations within curatorial help with the relevant coral collec-
192 M. E. J. Wilson and B. R. Rosen

tions, prepared the coral figures and assisted reefs. Lethaia 24: 423-438.
with the field collecting of the Boera corals Budd, A. F., Johnson, K. G. and Stemann, T. A. 1996. Plio-
Pleistocene turnover and extinctions in the Caribbean
(with David Haig and Ava Kila). The coral pho- reef-coral fauna. In Evolution and environment in tropical
tographs were taken by Phil Crabb. The com- America. pp. 168-204. Edited by J. B. C. Jackson, A. F.
ments of two reviewers, Ken Johnson and Neil Budd and A. G. Coates, University of Chicago Press,
Harbury, together with our editor Robert Hall, Chicago.
and four other readers, Johannes Pignatti, Budd, A. F., Stemann, T. A. and Johnson, K. G. 1994. Strati-
graphic distributions of genera and species of Neogene to
Alison Kay, Gustav Paulay and Andrew Racey Recent Caribbean reef corals. Journal of Paleontology 68:
greatly improved the paper. 951-977.
Budd, A. F., Stemann, T. A. and Stewart, R. H. 1992. Eocene
Caribbean reef corals: a unique fauna from the Gatuncillo
Formation of Panama. Journal of Paleontology 66: 570-
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