Journal of Biogeography (J. Biogeogr.) (2004) 31, 1381–1444
ORIGINAL
ARTICLE
Pollen-based reconstructions of biome
distributions for Australia, Southeast Asia
and the Pacific (SEAPAC region) at 0, 6000
and 18,000 14C yr BP
Elizabeth J. Pickett1,2, Sandy P. Harrison1*, Geoff Hope3, Kate Harle4, John R.
Dodson5, A. Peter Kershaw6, I. Colin Prentice1, John Backhouse5, Eric A.
Colhoun7, Donna D’Costa6, John Flenley8, John Grindrod6, Simon Haberle9,
Cleve Hassell10, Christine Kenyon11, Mike Macphail6, Helene Martin12,
Anthony H. Martin13, Merna McKenzie6, Jane C. Newsome14, Daniel Penny6,
Jocelyn Powell15, J. Ian Raine16, Wendy Southern17, Janelle Stevenson18, JeanPierre Sutra1,19, Ian Thomas11, Sander van der Kaars6 and Jerome Ward20
1
Max Planck Institute for Biogeochemistry,
Jena, Germany, 2PO Box 20, Walpole, Western
Australia 6398, Australia, 3Department of
Prehistory, Research School of Pacific Studies,
Australian National University, Canberra,
ACT, Australia, 4Environmental
Radiochemistry Laboratory, Australian
Nuclear Science and Technology Organisation,
Menai, NSW, Australia, 5School of Earth and
Geographical Sciences, The University of
Western Australia, Crawley, Western
Australia, Australia, 6Centre for Palynology
and Palaeoecology, School of Geography and
Environmental Science, Monash University,
Clayton, Victoria, Australia, 7Department of
Geography and Environmental Science, The
University of Newcastle, University Drive,
Callaghan, NSW, Australia, 8Geography
Programme, School of People, Environment
and Planning, Massey University, Palmerston
North, New Zealand, 9Resource Management
in Asia-Pacific Program, Research School of
Pacific and Asian Studies, Australian National
University, Canberra, ACT, Australia, 10School
of Plant Biology/Faculty of Natural and
Agricultural Sciences, The University of
Western Australia, Crawley, Western
Australia, Australia, 11Department of
Geography and Environmental Science, The
University of Melbourne, Parkville, Victoria,
Australia, 12Biological, Earth and
Environmental Sciences, The University of New
South Wales, Sydney, NSW, Australia, 132A
Birdwood St, Ryde, NSW 2112, Australia,
14
Murdoch School of Environmental Science,
Murdoch University, South Street, Murdoch,
Western Australia, Australia, 15Hawkesbury
ABSTRACT
Aim This paper documents reconstructions of the vegetation patterns in
Australia, Southeast Asia and the Pacific (SEAPAC region) in the midHolocene and at the last glacial maximum (LGM).
Methods Vegetation patterns were reconstructed from pollen data using an
objective biomization scheme based on plant functional types. The biomization
scheme was first tested using 535 modern pollen samples from 377 sites, and then
applied unchanged to fossil pollen samples dating to 6000 ± 500 or
18,000 ± 1000 14C yr bp.
Results 1. Tests using surface pollen sample sites showed that the biomization
scheme is capable of reproducing the modern broad-scale patterns of vegetation
distribution. The north–south gradient in temperature, reflected in transitions
from cool evergreen needleleaf forest in the extreme south through temperate rain
forest or wet sclerophyll forest (WSFW) and into tropical forests, is well
reconstructed. The transitions from xerophytic through sclerophyll woodlands
and open forests to closed-canopy forests, which reflect the gradient in plant
available moisture from the continental interior towards the coast, are
reconstructed with less geographical precision but nevertheless the broad-scale
pattern emerges. 2. Differences between the modern and mid-Holocene
vegetation patterns in mainland Australia are comparatively small and reflect
changes in moisture availability rather than temperature. In south-eastern
Australia some sites show a shift towards more moisture-stressed vegetation in
the mid-Holocene with xerophytic woods/scrub and temperate sclerophyll
woodland and shrubland at sites characterized today by WSFW or warmtemperate rain forest (WTRF). However, sites in the Snowy Mountains, on the
Southern Tablelands and east of the Great Dividing Range have more moisturedemanding vegetation in the mid-Holocene than today. South-western Australia
was slightly drier than today. The single site in north-western Australia also shows
conditions drier than today in the mid-Holocene. Changes in the tropics are also
comparatively small, but the presence of WTRF and tropical deciduous broadleaf
forest and woodland in the mid-Holocene, in sites occupied today by cooltemperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic
vegetation in the south and tropical deciduous broadleaf forest and woodland in
the north indicate drier conditions across mainland Australia at the LGM. None
ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi
1381
E. J. Pickett et al.
Nepean Catchment Management Trust,
Windsor, NSW, Australia, 16Institute of
Geological and Nuclear Sciences, Lower Hutt,
New Zealand, 178 Noala St, Aranda, Canberra,
ACT 2614, Australia, 18Department of
Archaeology and Natural History, Research
School of Pacific Studies, Australian National
University, Canberra, ACT, Australia, 1978 Av.
Daumesnil, F-75012 Paris, France and 20Box
2592, Auburn, AL 36831, USA.
*Correspondence: Sandy P. Harrison, School of
Geographical Sciences, University of Bristol,
Bristol BS8 1SS, UK.
E-mail: [email protected]
of these changes are informative about the degree of cooling. However the
evidence from the tropics, showing lowering of the treeline and forest belts,
indicates that conditions were between 1 and 9 C (depending on elevation)
colder. The encroachment of tropical deciduous broadleaf forest and woodland
into lowland evergreen broadleaf forest implies greater aridity.
Main conclusions This study provides the first continental-scale
reconstruction of mid-Holocene and LGM vegetation patterns from Australia,
Southeast Asia and the Pacific (SEAPAC region) using an objective biomization
scheme. These data will provide a benchmark for evaluation of palaeoclimate
simulations within the framework of the Palaeoclimate Modelling
Intercomparison Project.
Keywords
Biomization, palaeovegetation patterns, palaeoclimate reconstructions, plant
functional types, drought-limited biomes, mid-Holocene, last glacial maximum,
temperate rain forests.
INTRODUCTION
The region encompassing Southeast Asia, Australasia and the
Pacific (here referred to as SEAPAC) is climatically and
physiographically diverse. The region stretches from the
equatorial tropics via subtropical deserts to the mid-latitude
temperate belt, encompassing within each zone a range from
continental to maritime climates. Physiographically, the region
includes both ancient stable landscapes and active mountain
forming regions. Despite this diversity, SEAPAC can be
considered as a biogeographical unit because of the floristic
affinities between the vegetation of different areas. SEAPAC
encompasses the intersection between the species-rich IndoMalesian (van Steenis, 1979; Nelson, 1981; Hope, 1996) and
the Gondwanan floral realms (Barlow, 1981, 1994). Understanding the role of long-term climatic changes on the
evolution of this diverse flora has motivated the reconstruction
of vegetation history on a variety of time-scales. Although
many kinds of palaeoenvironmental evidence can be used for
this purpose, the most abundant source of information
(particularly for the last glacial–interglacial cycle, which is
the focus of the present paper) comes from pollen records.
However, the very diversity of the flora makes it difficult to
make objective comparisons between vegetation records from
different subregions. None of the previous palaeovegetation
syntheses (e.g. Bowler, 1982; CLIMANZ, 1983; Dodson, 1989;
Kershaw et al., 1991; van der Kaars, 1991; Markgraf et al.,
1992; Harrison & Dodson, 1993; Kershaw et al., 1994;
Hubbard, 1995a,b; Colhoun, 1996; Flenley, 1996) encompasses
the whole of the SEAPAC region.
The concept of plant functional types (PFTs) provides one
approach to handling taxonomic diversity, as it groups
together species that share common attributes with respect
to, e.g. life-form, phenology and bioclimatic limits (Prentice
et al., 1992a; Steffen, 1996; Smith et al., 1997). Broad-scale
1382
vegetation types (biomes) arise out of the association of
different PFTs. Thus specific biomes can be recognized in
different regions in spite of differences in taxonomic composition. The concepts of PFTs and biomes underlie the
Palaeovegetation Mapping Project (BIOME 6000: Prentice &
Webb, 1998). The aim of BIOME 6000 is to produce global
vegetation maps for the mid-Holocene (6000 yr bp or 6 ka)
and last glacial maximum (LGM, c. 18,000 14C yr bp or 18 ka,
equivalent to 21,000 calendar yr bp) using an objective method
based on the allocation of taxa to PFTs, and PFTs to biomes.
The resulting maps provide a tool for the evaluation of climate
and earth system models (e.g. Texier et al., 1997; Harrison
et al., 1998; Jolly et al., 1998a; Broström et al., 1998; Kutzbach
et al., 1998; Prentice et al., 1998; Joussaume et al., 1999,
Kohfeld & Harrison, 2000).
With the exception of a few sites from southern Africa, the
Southern Hemisphere is not represented in the current version
of the BIOME 6000 synthesis (Prentice et al., 2000) The
emphasis on Northern Hemisphere data in the initial phase of
BIOME 6000 reflects the abundance of pollen records from,
e.g. Europe and North America, which made these regions
ideal for testing the methodology. The aim of this paper is to
present a compilation of pollen data from the SEAPAC region,
to reconstruct biome maps for 0, 6 and 18 ka, and to interpret
these maps in terms of changing palaeoclimates, and thus to
extend the current BIOME 6000 synthesis.
SEAPAC region
The SEAPAC region extends from 17 N to 45 S and 100 E
to 180 E and includes the Sunda-Sahul region of Sulawesi,
Java, Sumatra, Borneo and Thailand, Papua New Guinea
(PNG), Australia, the Melanesian islands including the Solomon Islands, New Caledonia, Fiji and Vanuatu, and Polynesia
including Hawaii in the north and Easter Island to the east.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
The climate of the northern part of the region is dominated
by the seasonal migration of the inter-tropical convergence
zone (ITCZ), which shifts from 10–15 N in June–August
(Austral winter) to 10–12 S in December–February (Austral
summer) (Linacre & Hobbs, 1977). To the south, the climate is
dominated by the descending air of the subtropical belt of high
pressure and divergence. The mean latitude of the subtropical
high pressure belt (STA, or subtropical anticyclone), which is
characterized by the occurrence of travelling anticyclones,
shifts seasonally from c. 29–32 S in Austral winter to c. 37–
38 S in Austral summer (Russell, 1893; Kidson, 1925;
Karelsky, 1954; Pittock, 1973). The winds leaving the anticyclone spiral on the equatorward side to form the south-easterly
trade winds (tropical easterlies) that influence the Pacific
Islands, northern Queensland and PNG during the Austral
winter. In Austral summer, with the southward displacement
of the ITCZ and the STA, onshore winds forming part of the
southern monsoon influence the southern tropics and can even
penetrate into the interior of Australia (Pittock, 1975, 1978).
The seasonal migration of the STA also influences the climate
of the southernmost part of the region, allowing the southern
westerlies to migrate northwards during Austral winter
(Gentilli, 1971). These large-scale circulation features determine the seasonal distribution of precipitation across the
region (see e.g. Harrison & Dodson, 1993). Thus, the
equatorial belt (dominated by the ITCZ throughout the year)
is characterized by small seasonal variations in rainfall and
temperature. The southern tropics and northern Australia are
characterized by a summer rainfall maximum. The continental
interior is characterized by highly irregular, but predominantly
summer, rainfall. The southernmost part of mainland Australia
has winter rainfall, and only the southernmost part of the
region (i.e. Tasmania) lies far enough south to receive
precipitation from the westerlies throughout the year.
Two major floristic domains are represented in the region:
the Indo-Malesian floral domain (van Steenis, 1979; Nelson,
1981; Hope, 1996; Flenley, 1979, 1998) which includes both
tropical and cosmopolitan species, and the Gondwanan floral
domain (Barlow, 1981, 1994) which is dominated by sclerophyll species. The tropical rain forests and seasonal forests of
the Sunda-Sahul region, characterized by the families Elaeocarpaceae, Moraceae and Meliaceae, lie in the core of the IndoMalesian floral domain. The core of the Gondwanan floral
domain, characterized by the families Myrtaceae, Proteaceae
and Epacridaceae, occurs in southern Australia. In between,
the vegetation consists of an admixture of the two floras with
the relative importance of one over the other varying as a
function of increasing distance both latitudinally and eastwards from the core regions (Hope, 1996; Mueller-Dombois &
Fosberg, 1998). Thus, elements of the Indo-Malesian flora
persist in the subtropical forest belts of PNG and into northern
Australia, but are increasingly admixed with elements of the
Gondwanan sclerophyll vegetation southwards. Indo-Malesian
floral elements are also represented in the islands of the
Western Pacific, admixed with Gondwanan elements including
Eucalyptus and Epacridaceae, although the species diversity of
both floral domains decreases eastwards (Mueller-Dombois &
Fosberg, 1998).
An obvious climatic gradient influencing vegetation patterns
within the SEAPAC region is temperature. Thus, there is a
latitudinal zonation from tropical rain forests in the far north,
through subtropical seasonal or dry forests into warmtemperate and cool-temperate rain forests (CTRF) in the far
south. This gradient parallels the gradient from primarily
Indo-Malesian, through admixed, to primarily Gondwanan
floras. However, the primarily west–east gradients related to
moisture availability are a more important climatic control
within the subtropical and temperate zones. The continental
interior of Australia is arid, with mean annual precipitation as
low as 100 mm year)1. Both the amount and reliability of
rainfall increases towards the eastern and southern coasts. The
vegetation of the arid interior is characterized by grasslands
and shrublands, which grade into open woodlands with
increasing moisture availability, while forests are confined to
the coastal regions (Carnahan, 1997). Rainfall is high
(c. 5000 mm year)1) on the islands of the western Pacific
and gradually decreases further east; this steep gradient results
in a transition from forest-dominated to non-forest vegetation
(Mueller-Dombois & Fosberg, 1998). The decrease in precipitation across the Pacific parallels the gradual decrease in
species diversity.
Topographic gradients associated with the Great Dividing
Range clearly influence climatic gradients and hence the
vegetation zonation in eastern Australia. However, topography
has a more important role in the extensive highland regions in
the tropics, and the topographic control on local climate is a
particularly important factor underlying the complex vegetation patterns characteristic of Borneo, Sumatra and PNG.
Transitions from tropical rain forest at sea level, through
montane forests into grassland or tundra at elevations above
c. 3000 m are common on many tropical mountains (e.g. Mt
Wilhelm: Hope & Peterson, 1975).
The modern vegetation patterns of the SEAPAC region
reflect the interaction among these floristic, climatic and
topographic gradients. The same gradients must also have been
important throughout the history of the region. There has been
considerable work on the long-term vegetation history of the
SEAPAC region, capitalizing on the existence of pollen records
covering the last interglacial–glacial cycle from both the tropics
and the stable ancient landscapes of the temperate region.
Examples include the studies at Lynch’s Crater (Kershaw,
1976), Lake Wangoom (Edney et al., 1990; Harle et al., 1999),
Lake George (Singh et al., 1981; Singh & Geissler, 1985), Kosipe
(Hope, 1982), and Lac Suprin (Hope, 1996). There are
considerably more sites providing data on the changes in
vegetation patterns during and since the LGM. However,
climatic constraints on the occurrence of lakes and peatbogs
mean that pollen records are largely restricted to well-watered
coastal regions and the tropics (e.g. Bowler, 1982; CLIMANZ,
1983; Dodson, 1989; Kershaw et al., 1991; van der Kaars, 1991;
Markgraf et al., 1992; Harrison & Dodson, 1993; Kershaw et al.,
1994; Hubbard, 1995a,b; Colhoun, 1996; Flenley, 1996). New
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1383
E. J. Pickett et al.
sources of information, including studies of stick-nest rat
middens (Pearson & Dodson, 1993; McCarthy et al., 1996;
Pearson, 1999) and isotope analyses of emu shells (Miller et al.,
1997, 1999; Johnson et al., 1999), could provide information
about vegetation changes in the more arid parts of the SEAPAC
region. Unfortunately, there are relatively few such studies at
the present time, and we have therefore not drawn on these
potential sources of information in the present analysis.
METHODS
Modern pollen and vegetation data
Modern pollen assemblages were obtained for 377 sites
(Table 1). About a third of the site records were derived from
the Indo-Pacific Pollen Database (INDOPAC: Hope et al.,
1999), which includes data from India, Southeast Asia,
Australia, New Zealand and Oceania. A further 31% of the
records come from the Southeast Australian Pollen Database
(SEAPD: D’Costa & Kershaw, 1997; Kershaw, 1998). The
remaining 35% of the records were obtained from the original
authors. A number of these records have not been previously
published.
Despite the large number of sites in the SEAPAC data base,
the spatial distribution of the modern surface samples is
uneven. There are surface samples available from the coastal
regions of the SW, SE, N and NW of Australia but none from
the continental interior. This reflects poor pollen preservation
in arid environments, and the lack of suitable lake or swamp
sites. There are surface samples from upland regions of PNG,
New Caledonia, Borneo, Sumatra, Irian Jaya, Indonesia,
Solomon Islands and Thailand, but relatively few from the
tropical lowlands. The availability of samples from upland
regions reflects the widely held perception that mountain
vegetation belts are highly sensitive to climatic change (e.g.
Flenley et al., 1976; Walker & Flenley, 1979; Powell, 1982;
Flenley, 1996). The comparative lack of data from the tropical
lowlands is further compounded by logistical access problems.
The modern pollen samples were obtained from a number of
sources including moss polsters (9%), pollen traps (5%), surface
soil samples (26%), lacustrine or swamp core tops (24%) and
other sediment samples (36%). The pollen source area sampled
by moss polsters or soil samples beneath the tree canopy is
several orders of magnitude smaller than the source area
sampled by, e.g. lacustrine core tops (Prentice, 1985; Chen,
1987; Kershaw & Strickland, 1990; Kodela, 1990; Crowley et al.,
1994; Kershaw & Bulman, 1994; Sugita, 1994). In forested
regions, lacustrine samples are more likely to provide a
reconstruction of regional vegetation. However, moss polsters,
surface soil samples and other sediment samples could be more
representative of the regional vegetation under certain conditions, for example in open vegetation or when the vegetation is
dominated by poor pollen producers. It is not always clear which
type of sample will most faithfully reflect the regional vegetation
(see e.g. Bigelow et al., 2003) and we have therefore included all
types of modern pollen samples in our analyses.
1384
Raw counts were obtained for the majority of the samples
(c. 65%). The SEAPD data were available as raw counts but
some of the minor taxa had already been excluded. A further
31% of the samples were available as per cent counts. Only 4%
of the surface samples were digitized from published pollen
diagrams. Pollen abundances estimated by digitization are
generally accurate to within 10% (Williams et al., 2000) and
this degree of uncertainty is unlikely to affect the resultant
biome allocation. A more important constraint on the use of
digitized data is the fact that published pollen diagrams do not
include all of the minor taxa. Although digitized data provide a
reasonably accurate means of reconstructing forest vegetation
(Prentice et al., 1996; Tarasov et al., 1998), minor taxa can be
important in reconstructing non-forest vegetation types and
therefore complete raw pollen counts appear to provide a
better discrimination among the various non-forest and
xerophytic vegetation types (Jolly et al., 1998a, Yu et al.,
1998, 2000). For this reason we used digitized data only for
sites in key regions where the raw counts were not available.
The exclusion of minor taxa from the SEAPD data set could
potentially cause problems with the SEAPAC biomization.
However, the majority of the SEAPD samples are from forested
vegetation types and therefore the absence of minor taxa is less
likely to have affected the biome allocation.
The SEAPAC data set only includes counts for pollen taxa
that are used in the biomization procedure. Aquatic pollen and
spores, and non-native taxa, were excluded. Taxa that were
only represented by a single grain in one site were also
excluded. No attempt was made to reduce or enhance the
taxonomic resolution of the original data. Thus the taxa list
includes species where these were distinguished by the original
authors. As a result, a total of 695 taxa were used in the
SEAPAC biomization.
Fossil pollen data
The time slices for the biomization procedure were defined as
6000 ± 50 and 18,000 ± 1000 14C yr bp on the radiocarbon
time-scale (6 and 18 ka, respectively). The sites encompass a
wide range of environments including lakes, peat bogs, caves,
and organic deposits associated with fluvial or dune environments. One record for 6 ka was obtained from a marine core.
Samples were selected on the basis of an age model constructed
by linear interpolation between the radiocarbon dates designated by the original authors to be reliable. Multiple samples
were selected when more than one sample fell within the
designated window.
The data set for 6 ka consists of 114 sites (Table 2), of which
53% were available as raw counts, 42% as per cent counts, and
5% were digitized from published diagrams. Multiple samples
were obtained for 47% of the sites. The data set for 18 ka
consists of 33 sites (Table 2). Raw pollen counts were available
for 73% of the 18 ka sites, 27% were available as per cent
counts. Duplicate samples were obtained for 52% of the sites.
The SEAPAC data sets for 6 and 18 ka represent a substantial
improvement, both in terms of number of sites and the range
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
2750
4
45
240
3550
1
1524
1524
990
1080
1500
3700
35
)5.972 143.153
Vanuatu )20.234 169.782
Anumon Swamp Vanuatu )20.158 169.823
Australia )30.483 115.433
)5.815 144.988
Australia )32.002 115.503
Australia )37.833 146.283
Australia )17.250 145.917
)5.750 145.013
Baw Baw Track
Baw Baw Village Australia )37.850 146.267
Australia )42.667 146.633
Australia )36.518 149.500
Badmenangidongwa
Barker Swamp
Beattie’s Tarn
Bega Swamp
Bellenden Ker
Bendenumbun
Bibra Lake (BL2) Australia )32.100 115.833
Badgingarra NP
Anouwe 2
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
PNG
PNG
PNG
60–70
330
Australia )31.753 128.083
Australia )30.067 115.567
Airstrip
Alexander
Morrison NP
Aluaipugua
(core 2)
960
100
Adamson’s Peak Australia )43.350 146.817
Aire Crossing
Australia )38.733 143.450
Site name
Site
type
Surrounding
vegetation
Mallee heath
Mosaic heathlands
Count
Count
Count
Moss Microphyll fern thicket
polster (high altitude)
Swamp Shrubby tussock
grasslands
Swamp Eucalyptus marginata
woodland
%Count Lake
Low open forest
Count Swamp Tall open forest
%Count Swamp Subalpine woodland
Trap
Swamp Closed forest
Swamp Mid/upper montane
Nothofagus
and mixed forest
Swamp Closed forest
Soil
Trap
Eucalyptus loxophleba
woodland
Count Swamp Coprosma, Poa,
Dacrycarpus, Cyathea
Count Swamp Acanthocarpus-Stipa low
dense heath, with
thickets of Callitris
%Count Swamp Subalpine woodland
Count
Count
Count
Count
Count
Count
0–31
0–2
0–12
0–13.5
0–5
n/a
0–7.5
0–10.1
Modern
2.9–5.0
0–5.5
0–18.5
Modern
Modern
1
1
3
1
1
3
2
1
3
2
3
1
1
1
1
7
3
2 tephra 2
2
48
n/a
n/a
3 + top
1
0
3
3
0
0
1
n/a
XERO CTRF
WSFW CTRF
SEAPD
SEAPD
Macphail (1979, 1986)
McKenzie (unpubl. data),
D’Costa & Kershaw (1997)
Martin (1973)
Pickett (1997)
References
2D
WTRF; INDOPAC Corlett (1984b)
TDFO
Pickett (1997)
DSFW DSFW;
DSFW;
DSFW
STEP
WTRF WTRF; INDOPAC Hope (1976)
WTRF
Backhouse (1993)
Modern XERO XERO;
XERO;
XERO
n/a
DSFW DSFW SEAPD
Strickland (unpubl. data),
D’Costa & Kershaw (1997)
n/a
DSFW DSFW SEAPD
Strickland (unpubl. data),
D’Costa & Kershaw (1997)
7
DSFW CTRF SEAPD
Macphail (1979, 1986)
1C
DSFW DSFW; INDOPAC Polach & Singh (1980),
Green et al. (1988)
DSFW;
DSFW
WTRF WTRF
Kershaw (1973a)
Haberle (1994)
TDFO;
CTRF;
CTRF
n/a
WTRF WTRF; INDOPAC Hope & Spriggs (1982),
WTRF
Hope et al. (1999)
Hope et al. (1999)
n/a
WTRF WTRF;
TDFO;
WTRF
Modern DSFW DSFW
Pickett (1997)
CTRF
Modern XERO DSFW
Modern XERO DSFW
7
n/a
Biomes
Age range No. of
No. of
14
Recon. Data base
(kyr)
C dates duplicates DC 0 ka Obs.
%Count Swamp Open heath–closed heath 0–10
%Count Swamp Closed Eucalyptus forest n/a
Latitude Longitude Altitude Sample
Country ()
()
(m)
type
Table 1 Characteristics of the surface pollen samples from the SEAPAC region. The sites are arranged alphabetically by name. Latitude and longitude are given in decimal degrees, with N and E
conventionally indicated as +, and S and W as ). The vegetation descriptions are derived from the publications, and the observed (Obs.) biome call is derived from the natural vegetation map in
the Atlas of Australian Resources (Carnahan, 1997). Dating control (DC) follows the COHMAP dating scheme, as described by Yu & Harrison (1995). The codes used for reconstructed biomes are
defined in Table 5
Palaeovegetation patterns for Australia and Southeast Asia
1385
1386
1500
1890
760
1
100
100
100
100
100
1170
440
4
9
)32.050 151.467
)36.417 148.317
)17.167 145.600
)36.418 150.10
)35.017 116.633
)35.033 116.650
)35.033 116.650
)35.033 116.650
)35.033 116.650
)31.868 151.517
)37.433 144.600
Australia
Australia
Australia
Australia
Australia
Boggy Lake
Boggy Lake
Boggy Lake - heath Australia
Boggy Lake - scrub Australia
Australia
Blue Lake
(Snowy Mts)
Boar Pocket Rd
Boggy Lake - forest Australia
Australia
Black Swamp
Bobundarra Swamp Australia
Fiji Islands )18.067 178.533
)38.217 141.30
Big Heathy Swamp Australia
Boggy Swamp
Bolobek Peatland
Bonatoa Bog
Australia
860
)41.383 145.633
Australia
Big Dam Marsh
Boomer Swamp,
Core 12
710
)41.685 147.683
Country
Site
type
Surrounding
vegetation
%Count Swamp Woodland
%Count Swamp Open woodlandgrassland
Count
Swamp Disturbed tropical
evergreen forest
Eucalyptus mixed
open forest
Scrub
Eucalyptus
diversicolor forest
Heath
Swamp Coastal heath
Swamp Coastal heath
Moss
polster
Count
Moss
polster
Count
Moss
polster
Digitized Swamp
Count
Count
Count
Digitized Swamp Eucalyptus
pauciflora-E.
dalrympleana open
forest
Count
Lake
Alpine heath and
scrub/low shrubland
Count
Moss Wet sclerophyll forest
polster
Count
Swamp
%Count Swamp Tall open forest
%Count Swamp Open forest
Latitude Longitude Altitude Sample
()
()
(m)
type
Site name
Table 1 continued
0–6
0–4.4
n/a
0–9.3
Modern
Modern
Modern
0–4.4
0–6.8
0–7.27
Modern
0–13.34
0–8.6
n/a
1
1
1
3
3
3
1
1
3
1
1
2
1
4 (1 not 3
used)
n/a
12 (2 not 3
used)
0
0
0
1 + top
3 + top
3
0
16
3 + top
n/a
2
Thomas (unpubl. data),
D’Costa & Kershaw (1997)
Dodson (1987),
Dodson et al. (1986)
Becker (unpubl. data)
DSFW DSFW;
DSFW;
DSFW
DSFW DSFW SEAPD
D’Costa & Kershaw (1997)
Dodson (1987), Dodson
et al. (1986)
Churchill (1968)
Churchill (1968)
TRFO TRFO; INDOPAC Southern (1986), Hope
et al. (1999)
TRFO;
TDFO
Modern DSFW DSFW SEAPD
Head (1983, 1988)
5D
n/a
1D
Modern XERO DSFW
Modern XERO TDFO
TRFO STEP; INDOPAC Coddington (1983)
TRFO;
STEP
Pickett (1990), Newsome &
1C
XERO TDFO;
Pickett (1993)
TDFO;
DSFW
Churchill (1968)
4C
XERO DSFW;
DSFW;
WTRF
Modern WSFW DSFW
Churchill (1968)
Kershaw (1973b)
TUND DSFW INDOPAC Raine (1974, 1982)
DSFW DSFW;
DSFW;
DSFW
Modern WSFW WTRF
n/a
1C
n/a
DSFW DSFW; SEAPD
DSFW
DSFW WSFW SEAPD
Biomes
Age range No. of
No. of
14
Recon. Data base References
(kyr)
C dates duplicates DC 0 ka Obs.
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
5 %Count
15 %Count
750 %Count
80 Count
900 Count
Australia )38.301 141.383
Australia )40.835 145.133
Australia )42.217 146.567
Australia )37.750 140.750
Bridgewater
Lake, Core A
Broadmeadows
Swamp
Brown Marsh
Brownes Lake
Bunya Mountains 6 Australia )26.850 151.567
0–30
0–6.8
0–1.5
0–7.5
0–9.3
0–10.8
0–27
Wooded grassland
Mixed Eucalyptus open
forest
Open forest-tall open
forest
Open forest
Australia )31.917 151.400 1230 Digitized Swamp
Swamp
Swamp
Australia )32.083 151.450 1100
235 %Count
Australia )37.448 145.692
Australia )37.333 146.750 1280 %Count
Burraga Swamp
centre
Butchers Swamp
Buxton
Caledonia Fen
0
2
3
2
5
10 (1 not
used)
5
n/a
0–25
0–12
0–11.3
0–2.1
0–12
1
2 + top
Kenyon (1989),
Kershaw (1998)
Hope (1976), Hope &
Peterson (1975)
XERO WTRF;
TDFO;
WTRF
DSFW DSFW SEAPD Dodson (1983, 1986)
DSFW DSFW SEAPD Dodson (1983, 1986)
DSFW WTRF
6 1D
1 1C
1 n/a
1 1D
Kershaw (1976)
Kershaw (1976)
SEAPD Binder (1978),
Binder & Kershaw (1978)
CTRF TDFO;
Dodson et al. (1994b)
TDFO;
TDFO;
TDFO;
TDFO;
TDFO
Dodson (1987),
DSFW DSFW;
Dodson et al. (1986)
DSFW;
DSFW
DSFW DSFW SEAPD McKenzie (1989),
McKenzie & Busby (1992)
DSFW DSFW SEAPD Joyce (1979), Kershaw et al.
(1983), Kershaw (1998)
DSFW DSFW
1 Modern WTRF TDFO
1 Modern TRFO WTRF
DSFW DSFW
SEAPD van de Geer
et al. (1986)
1 7
DSFW DSFW SEAPD Macphail (1979, 1986),
Macphail & Hope
(1984/1985)
Dodson (1975b)
3 Modern XERO DSFW;
DSFW;
XERO
1 Modern TRFO WTRF
Kershaw (1976)
1 7
Churchill (1968)
DSFW TDFO;
DSFW;
TDFO
1 Modern XERO DSFW SEAPD Head (1988)
3
1 1C
1 1C
3
1 n/a
14 ( 4 not 3 1D
used)
4
2 + top
Modern 0
Modern 0
Modern 0
0–0.2
Low open forest-open heath 0–9
Tall open forest
Woodland
Mallee heath
Australia )36.783 146.767 1330
600
Swamp
Swamp
Swamp
Swamp
Swamp
Bunyip Bog
Bunya Mountains 8 Australia )26.850 151.567
Bunya Mountains 7 Australia )26.850 151.567 1050
0 Count
Australia )34.833 119.400
Bremer Bay Swamp
Swamp
Swamp
Treeline scrub and
subalpine tussock
grassland
Woodland
Woodland
Open forest
Moss
Low mesophyll vine forest
polster
and Araucarian emergents
Count
Moss polster Low microphyll vine
forest and Araucaria
Count
Moss polster Simple Eucalyptus vine
thicket and Araucaria
%Count Swamp
Subalpine woodland-open
forest
Digitized Swamp
Nothofagus moorei rain
forest/mixed Eucalyptus
forest
700 %Count
700 %Count
Australia )34.800 149.500
Australia )34.750 149.520
Breadalbane NW
Breadalbane SE
Swamp
)5.783 145.035 3910 Count
PNG
Brass Tarn
Swamp
130 Count
Boulder Flat Swamp Australia )37.467 148.967
Palaeovegetation patterns for Australia and Southeast Asia
1387
1388
90
1755
4
1350
760
180
760
900
Cotter Source Bog Australia )35.967 148.817
Australia )38.935 146.267
Australia )36.733 146.783
Australia )17.183 145.600
Australia )42.284 145.667
Australia )17.000 145.550
Australia )37.250 148.833
Cotter’s Lake
Crystal Bog 1
Danbulla
Darwin Crater
Davies Creek
Delegate River/
Tea Tree Swamp
1955
Australia )36.417 148.283
Australia )29.867 115.100
29
500
450
Australia )38.626 143.323
Australia )17.067 145.367
Australia )17.067 145.367
Chapple Vale
Chewko Swamp
Chewko Water
Storage
Club Lake core 1
Coolimba
15
Australia )40.501 144.833
Cave Bay Cave
140
1370
Australia )37.850 146.317
Australia )38.300 145.033
70
Australia )37.550 143.400
Carlisle Perched
Lake
Cascade Bog
Cobrico Swamp
1100
Camerons Lagoon Australia )41.969 146.683
Site name
Lake
Tussock grassland
low shrubland
Surrounding
vegetation
Swamp Alpine herbfield and
tall heath
0–6.7
0–6
Modern
Modern
Swamp Open forest
Moss Mixed notophyll vine Modern
polster forest and Agathis
%Count Swamp Open eucalypt forest 0–12
Count
Moss Simple notophyll vine Modern
polster forest
%Count Swamp Scrub-closed forest
5–350
Count
0–15
0–4.4
Eucalyptus erythocorys Modern
woodland
Swamp
0–5.7
Trap
%Count Swamp Subalpine woodland
Count
Count
Count
n/a
n/a
3
1
1
1
1
1
1
1
3
3
3 (1 not 1
used)
0
1
0
3
1
2
7 (1 not 1
used)
+ 1 on
section
4+8
1
210
Pb
0
1
5 + top
0
0
2
1
1
3
DSFW DSFW SEAPD
WSFW DSFW SEAPD
7
DSFW DSFW SEAPD
Modern TRFO WTRF
7
Ladd (1979b,c)
Colhoun & van de
Geer (1988)
Kershaw (1973b)
DSFW XERO; INDOPAC Hope & O’Dea
(unpubl. data)
XERO;
XERO
7
DSFW XERO; INDOPAC Hope (1974)
DSFW;
XERO
n/a
XERO DSFW SEAPD
Williams (1980, 1982),
Kershaw (1998)
Modern WTRF WTRF
Kershaw (1973b)
Yezdani (1970), Dodson
et al. (1994a)
Pickett (1997)
XERO DSFW INDOPAC Martin (1986)
Modern DSFW DSFW
1C
7
Modern XERO DSFW; INDOPAC Thomas & Hope (1994)
TUND;
TUND
n/a
DSFW DSFW SEAPD
Bottomley (1994),
D’Costa & Kershaw (1997)
n/a
DSFW DSFW SEAPD
Strickland (1980),
D’Costa & Kershaw (1997)
4D
XERO XERO; INDOPAC Hope (1978)
DSFW
1D
DSFW DSFW SEAPD
McKenzie & Kershaw (1997)
Modern DSFW DSFW
Kershaw (1976)
Modern DSFW DSFW
Kershaw (1976)
Biomes
No. of
No. of
14
Recon. Data base References
C dates duplicates DC 0 ka Obs.
0–8, 14.7–28 9
n/a
n/a
0–8
Age range
(kyr)
%Count Swamp Open forest-woodland 0–9
Count
Swamp Coastal open
scrub-heath
%Count Swamp Tall open forest
Count Soil
Eucalypt woodland
Count Soil
Eucalypt woodland
Count
%Count Swamp Subalpine woodland
%Count Swamp Tall open forest
Count
Latitude Longitude Altitude Sample Site
Country ()
()
(m)
type
type
Table 1 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
144.183
144.183
144.000
144.000
144.183
144.183
144.183
123.612
116.883
116.867
116.817
146.233
146.583
143.992
)5.650
)5.650
)5.750
)5.750
)5.650
)5.650
)5.650
)19.670
)32.783
)32.817
)32.083
)41.717
)42.683
)39.644
PNG
PNG
PNG
PNG
PNG
PNG
PNG
Australia
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Australia
Australia
Australia
Australia
Australia
Australia
Dublin Bog
Eagle Tarn
Egg Lagoon
Core 2
Dryandra State
Forest (13)
Dryandra State
Forest (14)
Dryandra State
Forest (15)
Draepi Swamp
40A
Draepi Swamp
WII29
Draepi Swamp 12
Draepi Swamp 13
Draepi Swamp 159
Draepi Swamp 40B
Draepi WiiPW
Dragon Tree Soak
145.583
)17.133
Australia
Downfall Creek
131.367
)30.950
Australia
Diprose Cave
148.483
)36.383
Australia
Digger’s Creek
Bog
146.233
)41.500
Australia
Den Plain a1b
146.233
)41.500
Australia
Den Plain a 2
146.233
)41.500
Australia
Den Plain a 1
1033
17
575
320
300
360
1885
1885
1885
1885
1885
200
1885
1885
700
100
1690
230
230
230
%Count
%Count
%Count
Count
Count
Count
Count
Count
Count
Count
Count
Digitized
Count
Count
Count
Count
Count
%Count
%Count
%Count
Lake
Swamp
Swamp
Trap
Trap
Trap
Swamp
Swamp
Soil
Soil
Soil
Swamp
Soil
Moss
polster
Soil
Soil
Swamp
Swamp
Swamp
Swamp
Eucalyptus wandoo and
E. accedens woodland
Dryandra open
woodland
Eucalyptus marginata,
E. wandoo and E.
accedens open woodland
Tall open forest (wet
sclerophyll and open
mountain forest) with
scrub in immediate
vicinity of site
Low open forest
Open-scrub heath
Dense grassland and
Acacia shrubs
Saltbush-bluebush low
shrubland
Complex mesophyll
vine forest
Eucalyptus pauciflora/
niphrophila woodland
Grassland-open forest
Grassland-open forest
Grassland-open forest
0–12
39.4 to > 55
0–13.6
Modern
Modern
Modern
0–6.4
Modern
Modern
0–10.5
n/a
n/a
n/a
3
n/a
8
0
0
0
2
0
0
5
n/a
n/a
n/a
1
1
1
1
1
1
1
1
1
1
1
5
1
1
1
1
3
1
1
1
7
n/a
7
Modern
Modern
Modern
7
Modern
Modern
1D
n/a
n/a
n/a
DSFW
XERO
WSFW
DSFW
DSFW
DSFW
TDFO
TDFO
TDFO
TDFO
TDFO
XERO
TDFO
TDFO
WTRF
XERO
DSFW
DSFW
DSFW
DSFW
CTRF
DSFW
DSFW
DSFW
DSFW
TDFO
TDFO
WTRF
TDFO
TDFO
XERO;
STEP;
XERO;
XERO;
XERO
DSFW
TDFO
TDFO
WTRF
DSFW;
DSFW;
DSFW
XERO
DSFW
DSFW
DSFW
SEAPD
SEAPD
SEAPD
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
SEAPD
SEAPD
SEAPD
Macphail (1979)
D’Costa & Kershaw (1997)
Colhoun et al. (1991)
Pickett (1997)
Pickett (1997)
Pickett (1997)
Powell (unpubl.
data, 1968)
Powell (unpubl.
data, 1968)
Powell (unpubl. data)
Powell (unpubl. data)
Powell (unpubl. data)
Powell (unpubl. data)
Powell (unpubl. data)
Pedersen (1983),
Wyrwoll et al. (1986)
Kershaw (1973b)
Martin (1973)
Moss (1994), D’Costa &
Kershaw (1997)
Moss (1994), D’Costa &
Kershaw (1997)
Moss (1994), D’Costa &
Kershaw (1997)
Martin (1999)
Palaeovegetation patterns for Australia and Southeast Asia
1389
1390
0 Count
8 Count
)34.292 119.458
)34.317 115.150
Gordon Inlet
GS3 FRNP
Governor Bog
Greens Bush
Gunung Kerinci
Plot 1
Gunung Kerinci
Plot 2
)42.184 145.650
)38.433 144.933
)1.067 100.767
Australia
Australia
Sumatra
(Indonesia)
Sumatra
(Indonesia)
)1.067 100.767
)34.292 119.458
Australia
)5.800 145.037
)3.599 132.164
G5–2-053P
Indonesia
(Seram Trench)
Giwimawi
PNG
Swamp Secondary grassland,
with forest nearby
0 Count Swamp Deltaic vegetation/open
mallee shrubland
180 %Count Swamp Scrub-closed forest
160 %Count Swamp Open woodland
3315 Count Soil
Ericaceous forest (upper
montane rain forest)
3010 Count Soil
Ericaceous forest (upper
montane rain forest)
3900 Count
Marine n/a
220 %Count Swamp Open forest-woodland
)37.583 142.317
Freshwater Lake Australia
Marine
60 %Count Swamp
Swamp
)41.067 147.133
Micronesia
Swamp
Swamp
Swamp
Forester Marsh Australia
Fool Swamp
4 Count
2 Count
)2.933 151.467
PNG
Fissoa River
9.486 138.087
190 Count
)18.761 169.001
Vanuatu
Fitzgerald Inlet Australia
Core 2
Flinders Bay
Australia
Swamp
350 Count
)28.250 153.167
Australia
Enoggra State
Forest
Evoran Pond
Mixed mesophyll
evergreen vine forest
Complex notophyll vine
forest and Araucaria
Acacia spirobis open
woodland, secondary
disturbance
Terminalia-Elaeocarpus
forest (lowland tropical
closed forest)
Deltaic vegetation/open
mallee shrubland
Eucalyptus calophylla,
Banksia grandis and Agonis
flexuosa heath-woodland
Savanna with Pandanus,
Melaleuca, Verban and
Gleichenia
Tall open forest
60 Count
)17.800 146.000
Australia
Moss
polster
Moss
polster
Swamp
17 %Count Swamp Open-scrub heath
)39.644 143.992
Australia
Egg Lagoon
Core 3
El Arish
Surrounding
vegetation
Country
Latitude Longitude Altitude Sample Site
()
()
(m)
type
type
Site name
Table 1 continued
Modern
0–13
n/a
Modern
Modern
0–1.5
0-ca14
0–7
0–4.4
0–5.3
0–2
0–7
0–2.5
0–2.46
Modern
39.4 to
> 55
Modern
1
1
3
3
1
3
3
1
1
1
0
2
n/a
0
0
1
1
1
1
1
XERO DSFW SEAPD
Kershaw (1976)
D’Costa & Kershaw
(1997)
Kershaw (1973b)
TDFO TDFO
DSFW DSFW SEAPD
XERO DSFW;
DSFW;
DSFW
TDFO TDFO;
TDFO;
TDFO
DSFW DSFW SEAPD
Thomas (unpubl. data),
D’Costa & Kershaw (1997)
Kershaw & McKenzie
(unpubl. data), D’Costa &
Kershaw (1997)
van der Kaars (1991, 1995)
Dodson & Intoh (1999)
Churchill (1968)
Modern CTRF WTRF
7
XERO WSFW SEAPD
n/a
DSFW DSFW SEAPD
Modern CTRF WTRF
Colhoun et al. (1991)
Jenkins (1992)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
WTRF WTRF; INDOPAC Corlett (1984b)
WTRF
Modern XERO XERO
Hassell (unpubl. data)
2D
7
n/a
n/a
1D
1C
TDFO WTRF; INDOPAC Hope et al. (1999)
TUND;
WTRF
n/a
TRFO WTRF; INDOPAC Hope et al. (1999)
TSFO;
TRFO
Modern XERO XERO
Hassell (unpubl. data)
n/a
Modern WTRF WTRF
Modern TRFO TDFO
n/a
Biomes
No. of
duplicates DC 0 ka Obs. Recon. Data base References
0 (isotrope 1
stratig.)
1
2
n/a
n/a
3
1 + top
1
2
2
0
0
n/a
Age range No. of
14
(kyr)
C dates
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
PNG
PNG
PNG
PNG
Australia
Australia
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
PNG
PNG
PNG
PNG
PNG
PNG
PNG
HF10
HF15
HF2
HF3
HF5
HF6
HF7
3330
3640
3700
3395
750
1130
)5.750 144.000 2585 Count Soil
)5.750 144.000 2400 Count Soil
)5.750 144.000 2740 Count Soil
)5.750 144.000 3055 Count Soil
)5.750 144.000 3190 Count Soil
Soil
Soil
Soil
Soil
Soil
Moss
polster
)5.750 144.000 2280 Count Soil
)5.750 144.000 2740 Count Soil
144.000
144.000
144.000
144.000
145.333
145.450
High altitude mixed forest
Tussock grassland
Tussock grassland
Tussock grassland
Eucalypt woodland
Simple notophyll-microphyll
vine forest
Regrowth beech forest
Quintinia, Ericaceae and
Epacridaceae mixed forest
Quintinia, Ericaceae and
Epacridaceae mixed forest
Quintinia, Ericaceae and
Epacridaceae mixed forest
Nothofagus, Castanopsis, Lauraceae
and Drimys beech forest
Nothofagus, Castanopsis, Lauraceae
and Drimys beech forest
Nothofagus, Castanopsis, Lauraceae
and Drimys beech forest
Count
Count
Count
Count
Count
Count
)5.750
)5.750
)5.750
)5.750
)17.000
)17.350
Sumatra (Indonesia) )1.067 100.767 2240
Gunung Kerinci SS9
Hagen Summit 12
Hagen Summit 3
Hagen Summit 4
Hagen Summit 9
Hastie Swamp
Herberton Range
Sumatra (Indonesia) )1.067 100.767 2395
Gunung Kerinci SS8
)5.833 142.783 1650
Sumatra (Indonesia) )1.067 100.767 2545
Gunung Kerinci SS7
PNG
Sumatra (Indonesia) )1.067 100.767 2855
Gunung Kerinci SS6
Haeapugua, Core 6
Sumatra (Indonesia) )1.067 100.767 3070
Gunung Kerinci SS5
)5.797 145.042 3740
Sumatra (Indonesia) )1.067 100.767 3175
Gunung Kerinci SS4
PNG
Sumatra (Indonesia) )1.067 100.767 1935
Gunung Kerinci SS11
Guruanglakugl
Sumatra (Indonesia) )1.067 100.767 2085
Gunung Kerinci SS10
Ericaceous forest (upper montane
rain forest)
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Soil
Lower montane forest/marsh,
swamp/dryland crops
Count Swamp Short grass bog/sedge fen/tussock,
open tropical forest nearby
Count Swamp Lower montane forest
Gunung Kerinci Plot 3 Sumatra (Indonesia) )1.067 100.767 2700 Count Soil
2 tephra
1
1
1
1
1
1
1
2 2D
1 Modern WTRF TDFO
1 Modern WTRF WTRF
1 Modern WTRF WTRF
1 Modern WTRF TDFO
1 Modern WTRF TRFO
1 Modern WTRF WTRF
1 Modern WTRF TDFO
1 Modern WTRF TDFO
1 Modern CTRF WTRF
INDOPAC
INDOPAC
INDOPAC
INDOPAC
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Kershaw (1976)
Kershaw (1973b)
Haberle (1993, 1998),
Haberle et al. (1999)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
Newsome (1985),
Stuijts et al. (1988)
INDOPAC Corlett (1984b)
WTRF TDFO INDOPAC Powell (1970)
WTRF TDFO INDOPAC Powell (1970)
WTRF TDFO INDOPAC Powell (1970)
CTRF TDFO INDOPAC Powell (1970)
CTRF TDFO INDOPAC Powell (1970)
CTRF WTRF INDOPAC Powell (1970)
CTRF TDFO INDOPAC Powell (1970)
TDFO TDFO;
TDFO
WTRF TDFO;
0–29
11 (1 not 3 1D
TDFO;
used) + 2
TDFO
tephra
1
CTRF TDFO
1
STEP TDFO
1
STEP TDFO
1
STEP TDFO
Modern 0
1 Modern DSFW TDFO
Modern 0
1 Modern WTRF TDFO
0–1.2
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Palaeovegetation patterns for Australia and Southeast Asia
1391
1392
149.133
149.133
141.800
)37.100
)37.100
)36.800
Australia
Australia
Australia
Jacksons Bog A
Jacksons Bog B
Jaka Lake
1
5
145.417
116.067
)31.950
21.425 )157.75
)156.6
)17.517
Australia
Australia
Hawaii
(USA)
Hawaii
(USA)
Australia
Kaban/Kalunga
Waterhole
Kalamunda NP
Kawainui Marsh
Core B
Kealia Pond
Killer Bog
)32.067
20.7
100
116.083
)31.867
John Forrest NP (2) Australia
151.550 1320
900
160
116.083
)31.867
200
140
750
750
137.217 3630
137.383 3580
2
John Forrest NP (1) Australia
Ijomba
Irian Jaya
)3.985
(Indonesia)
Indonesia
)4.033
Hogayaku Lake
147.70
Australia
Hidden Swamp
)37.967
Country
Surrounding
vegetation
Swamp Open Eucalypt forest
Swamp Cyathea tree ferns
with grassland
Swamp Disturbed subalpine
forest and treefern
Count
Count
Count
Count
Count
Count
Count
Eucalyptus marginata
and E. calophylla open
woodland
Eucalyptus wandoo
woodland
Eucalypt woodland
Casuarina woodland
Modern
Modern
Modern
0–12
0–13
0–13.9
0–6.5
0–7.5
Swamp Nothofagus moorei rain
0–8.3
forest/Eucalyptus
pauciflora-E. dalrympleana
open forest
3
3
1
1
1
1
1
1
3
3
2
1
DSFW
DSFW
DSFW
CTRF
1C
Pickett (1997)
Kershaw (1976)
Pickett (1997)
WTRF XERO; INDOPAC Ward & Athens
(unpubl. data)
WTRF;
XERO
WTRF DSFW;
Dodson (1987),
DSFW;
Dodson et al. (1986)
DSFW
TDFO TDFO; INDOPAC Ward & Athens
(unpubl. data)
XERO;
TDFO
Modern DSFW DSFW
Modern DSFW TDFO
Modern DSFW TDFO
Modern DSFW
n/a
n/a
7
TUND
Hooley
et al. (1980)
XERO; INDOPAC Hope (unpubl. data)
XERO
CTRF; INDOPAC Hope & Peterson
(1975, 1976),
TUND;
Haberle et al. (2001)
TUND
DSFW; INDOPAC Southern (1982),
Kershaw (1998)
DSFW;
DSFW
DSFW SEAPD
Southern (1982),
Kershaw (1998)
DSFW SEAPD
McKenzie
(unpubl. data),
D’Costa & Kershaw
(1997)
TDFO
Pickett (1997)
Modern DSFW DSFW SEAPD
Biomes
No. of
duplicates DC 0 ka Obs. Recon. Data base References
10
3
(4 not used)
6
5
0
0
0
0
n/a
4
3
2
2 + top
Age range No. of
14
(kyr)
C dates
Modern
Eucalyptus marginata
and E. calophylla open
woodland
0–3.48
Swamp Brachiaria mutica,
Schoenoplectus
and saw grass with ferns,
forest nearby
Swamp
0–4.92
Trap
Soil
Trap
Trap
%Count Lake
%Count Swamp Open Eucalypt forest
Count
Count
Count
%Count Swamp Open forest-woodland
Latitude Longitude Altitude Sample Site
()
()
(m)
type
type
Site name
Table 1 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Swamp Castanopsis mixed
montane forest
Swamp Mangroves, Melaleuca
shrub and pasture
Trap
Mixed mesophyll
vine forest
Swamp Dune Eucalyptus
woodland
)8.467 147.200 1960 Count
PNG
PNG
Komanimambuno
Kosipe
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
35 Count
75 %Count Lake
20 %Count Swamp Open forest-woodland n/a
60 %Count Lake
40 %Count Lake
5 %Count Swamp Grassland/disturbed
0–5.2
934 %Count Swamp Closed forest–tall open n/a
forest
8 %Count Swamp Open forest
0 to > 6
1 %Count Lake
Closed forest
40 %Count Swamp Coastal open
scrub-heath
)32.008 115.817
)38.250 142.450
)38.167 147.300
)38.067 141.833
)35.050 141.733
)37.833 148.500
)41.667 145.950
)42.502 145.675
)39.602 143.957
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Lake Banganup
(BGP4)
Lake Cartcarong
Lake Coleman
Lake Condah
(Core 1b)
Lake Crosby
Lake Curlip
Lake Dove
Lake Elusive
Lake Fiddler
Lake Flannigan
)37.750 149.450
235 Count
New Caledonia )22.292 166.968
Lac Xere Wapo
Tall shrubland
4
0–11
0–31
0–8
0–4
1 n/a
1 n/a
1
1 n/a
5 + top 1 1C
n/a
1 n/a
n/a
2 + top 1 1C
n/a
1 n/a
1
2 + top 1 2C
n/a
n/a
2
1 7
2 7
5 + top 3 1D
3 7
1
DSFW TDFO
1 Modern XERO XERO
WSFW WSFW SEAPD
DSFW DSFW SEAPD
van de Geer
et al. (1991)
Kershaw (1973a)
Martin (1973)
Ladd et al. (1992)
TDFO
DSFW SEAPD
CTRF SEAPD
DSFW SEAPD
CTRF
XERO
CTRF SEAPD
DSFW SEAPD
DSFW DSFW SEAPD
STEP
CTRF
XERO
DSFW DSFW SEAPD
DSFW DSFW SEAPD
McKenzie (unpubl. data),
D’Costa & Kershaw (1997)
Ladd (1978)
Dyson (1995),
D’Costa & Kershaw (1997)
Kenyon (unpubl. data),
Kershaw (1998)
Harle et al. (1999)
D’Costa & Kershaw (1997)
R. Walkley (unpubl. data),
D’Costa & Kershaw (1997)
Penny (1992), D’Costa
& Kershaw (1997)
Head (1989)
Pickett (1997)
DSFW; INDOPAC Martin (1994)
DSFW;
DSFW
WTRF; INDOPAC Hope (1996), Hope
TRFO
& Pask (1998)
WTRF INDOPAC Hope & Pask (1998)
DSFW DSFW SEAPD
XERO
XERO
XERO
XERO
WTRF WTRF; INDOPAC Hope (1976), Hope
& Peterson (1975)
WTRF;
WTRF
5 + top 2 2C
WTRF TDFO; INDOPAC Hope (1982), Haberle
TDFO
et al. (2001)
3
2 5D
XERO TDFO; INDOPAC Hope et al. (1999)
TDFO
0
1 Modern TRFO TDFO
Kershaw (1973b)
4
0
2 + top 1 1C
1 + top 1 1C
Modern, 25–80 1
0–33.3
0–8
Open woodland/forest 0–8.2
Open woodland
Gymnostoma maquis
shrublands
Lake
Gymnostoma maquis
shrublands
Swamp Banksia woodlands
235 Count
New Caledonia )22.300 166.983
Lac Suprin
Lake
2 Count
)34.017 151.167
Kurnell Fen Core 1 Australia
Modern
425 Count
)16.817 145.633
Australia
Kuranda
0–6.5
2 Count
0 to > 30
0–21.76
Modern
0–2.5, 6.4–10
0–0.6
Koumac Northeast New Caledonia )20.650 164.283
)5.800 145.067 2740 Count
Eucalypt woodland
Saltbush-bluebush
low shrubland
Swamp Montane-closed forest
)18.483 145.033
)31.117 130.633
Australia
Australia
Soil
Soil
700 Count
100 Count
)42.150 146.650
Australia
King River
Railway Bridge
Kinrara Swamp
Knowels Cave
)39.785 147.883
15 %Count Swamp Eucalyptus nitida
open forest, heath
or scrub
200 %Count Swamp Closed forest
Australia
Killiecrankie
Palaeovegetation patterns for Australia and Southeast Asia
1393
1394
1300 %Count Swamp Subalpine woodland
3 + top %Count Swamp Tall open forest
780 Count Swamp Araucaria, lower
montane forest
875 %Count Lake
Subalpine-closed forest
150 %Count Lake
Open woodland
80 %Count Lake
Open scrub-heath
)37.150 147.933
)38.784 143.467
)2.550 140.600
Australia
Australia
Australia
Lake Mountain
Lake Purrembete
Lake Ranfurlie E.
40 %Count Lake
)34.183 142.117
Shrubland
140 %Count Swamp Open woodlandgrassland
)38.267 143.217
)37.469 145.875
140 Count
)1.183 127.483
Indonesia
Lake Majo
n/a
3 + top
17
0–8.5
0–8.5
1 + top
3
5 + top
5 + top
0–4.5
3
0 to > 35 7 (1 not
used)
0–9.5
7 + top
0–9.7
4 + top
0–15
0–10
0–16
0–128
1
1
1
2
1
4
1
1
1
1
2
1
3
1
1
1D
5D
1C
1C
2C
1C
1C
7
n/a
2C
4D
INDOPAC Singh
& Geissler (1985)
DSFW DSFW SEAPD
Yezdani (1970),
D’Costa
& Kershaw (1997)
CTRF TDFO; INDOPAC Haberle et al. (2001)
TDFO;
TDFO
DSFW DSFW SEAPD
Kershaw & McKenzie
(unpubl. data),
Kershaw (1998)
DSFW CTRF SEAPD
Head & Stuart (1980)
WTRF WTRF; INDOPAC Hope et al. (1988),
WTRF
Hope & Tulip (1994)
CTRF CTRF SEAPD
Anker et al. (2001)
DSFW DSFW SEAPD
Dodson (1974a)
XERO DSFW SEAPD
McKenzie (unpubl.
data), D’Costa
& Kershaw (1997)
DSFW DSFW SEAPD
Dodson (1974b)
Dodson (1974b)
DSFW DSFW;
DSFW;
DSFW;
DSFW
TDFO TDFO; INDOPAC Hope (1993), Haberle
TDFO
et al. (2001)
DSFW DSFW SEAPD
McKenzie (1997),
McKenzie
& Busby (1992)
XERO DSFW SEAPD
Penny (unpubl.
data), D’Costa
& Kershaw (1997)
XERO DSFW SEAPD
McKenzie
(unpubl. data),
D’Costa
& Kershaw (1997)
DSFW STEP
Biomes
Age range No. of
No. of
14
(kyr)
C dates duplicates DC 0 ka Obs. Recon. Data base References
Swamp Gymnostoma–Pandanus 0–4.27
scrub and sedgeland
1440 %Count Swamp Subalpine woodland
0–6.5
150 %Count Swamp Woodland
150 Count Swamp Woodland
)37.650 140.583
)37.650 140.583
Subalpine rain
forest/grassland
Australia
Australia
Australia
Lake Hill
Lake
3120 Count
)4.117 138.700
Lake Leake
Lake Leake (core LE)
Indonesia
Lake Habbema
)38.217 143.100
)41.850 145.533
)38.200 142.867
)35.717 142.367
Australia
Lake Gnotuk
Surrounding
vegetation
Swamp Eucalypt woodland
and grasslands
200 %Count Lake
Open woodlandgrassland
673 Count
Site
type
)35.083 149.417
Latitude Longitude Altitude Sample
()
()
(m)
type
Australia
Irian Jaya
(Indonesia)
Lake Johnston, Core JA Australia
Lake Keilambete
Australia
Lake Lascelles
Australia
Australia
Lake George
Lake Hordern Core A
Lake Hordorli
Country
Site name
Table 1 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Lake Terang
Lake Tiberias
Lake Turangmoroke
Lake
Lake
Lake
Lake
Open woodland
Open forest
Open woodland-grassland
Open scrub-heath
Closed forest
Open woodland
Open forest-woodland
)38.250 142.917 130 %Count Swamp
)42.367 147.367 442 %Count Swamp
)37.700 142.750 200 %Count Swamp
)35.250
)42.267
)38.350
)38.100
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Australia
)32.000 115.500 < 25 Digitized Swamp
2 %Count Swamp
Lighthouse Swamp
)35.767 138.067
Australia
Lashmar’s Lagoon
Acanthocarpus-Stipa low
dense heath
Open woodland
0–5
0–7
est.
5
3
)8.384 147.367 3640 Count
PNG
Laravita Tarn
0–13
)35.483 149.068 578 Count
Australia
Lanyon House
Alpine grassland
Swamp
)28.200 153.167 600 Count
Lamington National Park 3 Australia
0 (sed corr.)
3 + top
8 (1 not used)
n/a
0 (poll) + top
0 (corr.)
Moss polster Microphyll moss forest
Modern 0
Moss polster Simple notophyll evergreen Modern 0
vine forest (lower montane)
Moss polster Complex notophyll
Modern 0
vine forest
Soil
Savanna woodland, pasture,
former agricultural field
0–10
0–12
0–43
n/a
0–20
0–10
0– c. 75 3 + top
DSFW DSFW SEAPD
7
4C
7
n/a
XERO
CTRF
DSFW
DSFW
DSFW
CTRF
DSFW
DSFW
1
1 n/a
1
8
Kershaw (1976)
Salas (1981),
D’Costa
& Kershaw (1997)
D’Costa (1989),
D’Costa
& Kershaw (1995)
Macphail
& Jackson (1978)
Crowley
& Kershaw (1994)
Luly (1993)
Macphail (1979)
Edney et al. (1990)
Reid (1989),
D’Costa
& Kershaw (1997)
Kershaw (1976)
Kershaw (1976)
Pola (1993),
Colhoun
et al. (1999)
Southern (1986),
Hope et al. (1999)
XERO XERO; INDOPAC Hope & O’Dea
(unpubl. data)
XERO;
XERO;
XERO;
XERO;
XERO;
DSFW;
XERO
TUND XERO INDOPAC Hope (1980),
Hope & O’Dea
(unpubl. data)
DSFW DSFW SEAPD
Clark (1983a,b),
D’Costa
& Kershaw (1997)
XERO XERO
Churchill
(1960, 1968)
1 Modern TRFO WTRF
SEAPD
SEAPD
SEAPD
SEAPD
XERO DSFW SEAPD
1 Modern WTRF CTRF
1 Modern WTRF WTRF
1
1
1
1
1 1D
1 Modern DSFW DSFW SEAPD
1
1 Modern WSFW COCO SEAPD
4 (2 not used)
+ 3AMS (1 not used)
+ 6 U/Th (3 not used)
0–14.3 6
3 n/a
TDFO TDFO; INDOPAC
WTRF;
WTRF
0–1.5 1
1 n/a
DSFW DSFW SEAPD
0–70
)28.200 153.167 900 Count
)28.200 153.167 750 Count
142.867 17 %Count Lake
145.867 560 %Count Lake
142.600 100 %Count Lake
147.300 20 %Count Swamp
Open forest
)37.583 146.833 915 %Count Swamp
Swamp
Closed forest, degraded
to heath in immediate
vicinity of lake
Lamington National Park 1 Australia
Lamington National Park 2 Australia
Tyrrell (Site 2)
Vera
Wangoom
Wellington
Australia
Lake Tali Karng
Fiji Islands )16.817 179.933 820 Count
Lake Tagamaucia
)41.883 145.600 516 %Count Lake
Australia
Lake Selina
Palaeovegetation patterns for Australia and Southeast Asia
1395
1396
147.683
141.083
151.417
145.617
144.167 1580
144.167 1580
147.817 950
146.767
145.850
146.083
148.520
148.167
145.900
146.471
148.067 1170
)37.967
)38.084
)2.933
)17.600
)5.733
)5.733
)41.367
)38.433
)17.633
)43.267
)35.334
)39.985
)17.417
)38.251
)37.117
Australia
Loch Sport Swamp Australia
Long Swamp
Australia
Lowelak Swamp
PNG
Australia
PNG
PNG
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Maalan turnoff
Manton Core 1
W-E Transect
Manton Core 6
Mathinna Plain
McKenzie Road Bog Australia
Australia
Loch McNess
Swamp
McNamee Creek
Melaleuca Inlet
Micalong Swamp
Middle Patriarch
Swamp
Mirriwinni
Morwell Swamp
Moss Bed Lake
40
60
20
980
10
150
100
520
1
2
30
115.667 < 20
)31.533
Hawaii (USA)
Liliha Core 4
21.35 )157.967
Country
Swamp
Surrounding
vegetation
0–10.1
0–13.07
0–11.8
Modern
0–5
n/a
Modern
0–7.5
0–6
0–2.4
0–8.9
0–460
n/a
2 + top
0
3 + top
6
4
0
1
n/a
0
2 + top
2
n/a
2
1
1
1
1
1
3
1
1
1
1
1
1
1
1
1
2
2
3
DSFW DSFW SEAPD
Modern DSFW DSFW SEAPD
n/a
Thomas (1995)
Lloyd (1991), Lloyd &
Kershaw (1997)
Kershaw & McKenzie
(unpubl. data),
Kershaw (1998)
Kershaw (1973b)
DSFW DSFW; INDOPAC Kemp (unpubl. data)
DSFW;
DSFW
DSFW DSFW SEAPD
Ladd et al. (1992)
STEP WSFW SEAPD
Modern TRFO WTRF
1D
5D
TRFO TDFO INDOPAC Powell (1970)
STEP DSFW SEAPD
Thomas (unpubl.
data), D’Costa &
Kershaw (1997)
n/a
DSFW DSFW SEAPD
Robertson (1986),
D’Costa & Kershaw
(1997)
Modern TRFO WTRF
Kershaw (1973b)
n/a
Kershaw (1973b)
Hooley et al. (1980)
Head (1988)
Hope et al. (1999)
STEP TDFO INDOPAC Powell (1970)
Modern DSFW DSFW SEAPD
Modern DSFW DSFW SEAPD
n/a
TDFO TRFO; INDOPAC
TDFO
Modern TRFO TDFO
TDFO STEP; INDOPAC Ward (unpubl. data)
STEP;
STEP
DSFW DSFW;
Newsome & Pickett
DSFW
(1993)
Biomes
Age range No. of
No. of
14
(kyr)
C dates duplicates DC 0 ka Obs. Recon. Data base References
Moss Complex mesophyll vine Modern
polster forest (lowland rain forest)
%Count Swamp Open forest, dominated
0–9.5
by Eucalyptus
%Count Swamp Open forest
0–7
Count
%Count Swamp Open forest-heath
Moss Complex mesophyll vine
polster forest
%Count Swamp Buttongrass moorlandgrassland
Count Swamp Eucalyptus dalrympleana
open forest
Count
%Count Swamp Open forest-woodland
Swamp Eucalyptus erythocorys,
E. marginata and Banksia
woodlands
%Count Swamp Open forest-woodland
%Count Swamp Open forest
Count Swamp Disturbed lowland tropical
rain forest and grassland
Count Moss Complex mesophyll vine
polster forest
Count Soil
Regrowth shrubs in
grassland/disturbed
Count Soil
Swampland
%Count Swamp Buttongrass moorlandgrassland
Count
Count
Latitude Longitude Altitude Sample Site
()
()
(m)
type
type
Site name
Table 1 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
)37.833 140.750
)39.017 146.367
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Mt Glorius
Mt. Gambier
(Blue Lake)
Mt. Latrobe
Murrawijinie No. 1
Murrawijinie No. 3
Mustering Flat
Myalup Swamp
Nadrau Swamp
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
680 Count
Fiji Islands )17.750 177.883
17.183 103.033
17.100 102.933
Australia
Australia
Nong Han
Thailand
Kumphawapi (Core 3)
Thailand
Australia
Australia
Australia
Nardello’s Lagoon
Newall Creek
Nong Pa Kho
North Torbreck
Oaks Creek
Peppertree Marsh
610 %Count Swamp Open forest
890 %Count Swamp Tall open forest–closed
forest
)41.883 147.833
2 %Count Swamp Coastal heath-tall
shrubland
550 Count
Soil
Eucalypt woodland
140 %Count section Closed forest (temperate
rain forest)
170 Count
Lake
Dry to mixed deciduous
forests dominated by
Dipterocarpaceae
175 Count
Swamp Dry to mixed deciduous
forests dominated by
Dipterocarpaceae
564 %Count Swamp Tall open forest
)37.586 146.021
)37.481 146.947
)17.117 145.417
)42.150 145.517
Australia
Naracoopa
)39.883 144.083
0–11.3
Swamp Eucalyptus marginata
heath-woodlands
Swamp Lower montane forest
10 Count
)33.117 115.717
n/a
n/a
1
n/a
0–6.8
0–12
0–38.3
0–14
n/a
2 + top
1 + top
1 n/a
1 1D
1 1D
DSFW
DSFW
DSFW
TSFO
TSFO
WTRF
XERO
DSFW
DSFW
XERO
XERO
SEAPD
SEAPD
SEAPD
Strickland (unpubl. data),
Kershaw (1998)
Churchill (1968)
Martin (1973)
Howard & Hope (1970),
D’Costa & Kershaw (1997)
Martin (1973)
Salas & Kershaw (1982)
Kershaw (1973a)
Dodson & Wilson (1975)
Strickland (unpubl. data),
D’Costa & Kershaw (1997)
INDOPAC Powell (1970)
INDOPAC Powell (1970)
SEAPD
Strickland (unpubl. data),
D’Costa & Kershaw (1997)
SEAPD
van de Geer et al. (1986)
SEAPD
SEAPD
DSFW
DSFW
DSFW
TDFO
XERO
SEAPD
SEAPD
SEAPD
McKenzie (1989),
McKenzie & Busby (1992)
McKenzie (1989),
McKenzie & Busby (1992)
Becker (unpubl. data),
D’Costa & Kershaw (1997)
Penny (1998)
Penny (1998, 1999),
Kealhofer & Penny (1998)
TDFO; INDOPAC Southern (1986),
Hope et al. (1999)
TUND;
TUND
1 n/a
XERO WSFW SEAPD
Grindrod (unpubl. data),
D’Costa & Kershaw (1997)
1 Modern DSFW DSFW
Kershaw (1976)
1 7
WSFW CTRF
SEAPD
van de Geer et al. (1989)
3 n/a
1 7
DSFW
1 Modern XERO
1 n/a
DSFW
WTRF
DSFW
TDFO
TDFO
CTRF
DSFW
DSFW
WSFW CTRF
DSFW
WTRF
XERO
TSFO
STEP
DSFW
XERO
DSFW
1 Modern XERO
1 n/a
1
1
1 7
1
1
1 n/a
1 1D
1 n/a
8 (1 not 1 1C
used) +
top
3
1 7
Modern 0
0–21
2
n/a
0–2.6
1
n/a
Modern 0
Modern 0
0–6
n/a
597 %Count Swamp Closed forest–tall
open forest
)31.317 130.900 100 Count
Soil
Saltbush-bluebush
low shrubland
)31.383 130.950 100 Count
Soil
Saltbush-bluebush
low shrubland
)37.800 146.300 1448 %Count Swamp Subalpine woodland
)28.200 153.167
0–85
3
35 %Count Swamp Coastal shrublandswamp forest
650 Count
Moss
Complex notophyll
polster vine forest
100 %Count Swamp Open forest
)40.917 145.083
Australia
Mowbray Swamp
3
n/a
n/a
0–7.5
0–2.5
)6.000 145.750 2190 Count
Soil
Oak forest
)6.000 145.750 1980 Count
Soil
Regrowth and grasslands
)37.817 146.283 1448 %Count Swamp Subalpine woodland
0–2
PNG
PNG
Australia
Mount Oga 1
Mount Oga 2
Mount Whitelaw
)37.600 140.317 150 %Count Swamp Scrub and marsh
)37.867 146.333 1372 %Count Swamp Subalpine woodland
Australia
Australia
Mount Burr Swamp
Mount Kernot
Palaeovegetation patterns for Australia and Southeast Asia
1397
1398
Sumatra
(Indonesia)
Plot 10
Sumatra
(Indonesia)
Plot 11
Sumatra
(Indonesia)
Plot 12
Sumatra
(Indonesia)
Plot 13
Sumatra
(Indonesia)
Plot 16
Sumatra
(Indonesia)
Plot 17
Sumatra
(Indonesia)
Plot 18
Sumatra
(Indonesia)
Plot 19
Sumatra
(Indonesia)
Plot 2
Sumatra
(Indonesia)
Plot 20
Sumatra (Indonesia)
Plot 24
Sumatra (Indonesia)
Plot 3
Sumatra (Indonesia)
Plot 4
Sumatra (Indonesia)
Plot 5
Sumatra (Indonesia)
Plot 6
Sumatra (Indonesia)
Plot 7
Sumatra (Indonesia)
Plot 8
Sumatra (Indonesia)
Plot 9
Sumatra (Indonesia)
Plum (Plum New Caledonia
Swamp,
Core 2)
Australia
Phil’s Hill
Plot 1
Country
Site name
Table 1 continued
1535
1535
1535
1535
1440
1440
1430
1450
1535
1970
1535
1750
1780
1535
1535
1535
1535
1535
10
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067 100.767
)1.067
)1.067
)1.067
)1.067
)1.067
)1.067
)1.067
)1.067
)1.067
)22.267
100.767
100.767
100.767
100.767
100.767
100.767
100.767
100.767
100.767
166.617
1640
)1.067 100.767
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Moss
polster
Moss polster
Moss polster
Moss polster
Moss polster
Soil
Soil
Soil
Soil
Soil
Swamp
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Moss
polster
Soil
0–7
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
12 (2 not 3
used)
n/a
2D
n/a
WTRF
WTRF
WTRF
WTRF
TDFO
TDFO
TDFO
TDFO
TDFO
XERO
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Newsome (1985)
Hope (unpubl.
data), D’Costa &
Kershaw (1997)
Newsome (1985)
WTRF
Newsome (1985)
WTRF
Newsome (1985)
TDFO
Newsome (1985)
WTRF
Newsome (1985)
TDFO
Newsome (1985)
TDFO
Newsome (1985)
TDFO
Newsome (1985)
TDFO
Newsome (1985)
TDFO
Newsome (1985)
WTRF; INDOPAC Stevenson et al.
TDFO;
(2001)
WTRF
TDFO WTRF
TDFO TDFO
TDFO TDFO
TDFO TUND
TDFO TUND
TDFO TDFO
TDFO TDFO
TDFO TDFO
TDFO TDFO
WSFW WTRF
XERO DSFW SEAPD
Biomes
Age range No. of
No. of
14
Recon. Data base References
(kyr)
C dates duplicates DC 0 ka Obs.
Forest
Forest
Forest
Forest
Low regrowth/disturbed
Low regrowth/disturbed
Low regrowth/disturbed
Low regrowth/disturbed
Low regrowth/disturbed
Cyperaceous maquis and 0 to
Melaleuca woodland
> 23
Low regrowth/
disturbed
Low regrowth/
disturbed
Low regrowth/
disturbed
Low regrowth/
disturbed
Non-forest
regrowth
Non-forest
regrowth
Non-forest
regrowth
Non-forest
regrowth
Forest/disturbed
Forest/disturbed
Open scrub-heath
30
)40.367 148.267
%Count Swamp
Surrounding
vegetation
Latitude Longitude Altitude Sample Site
()
()
(m)
type
type
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Rottnest
Island (RS)
115.500
< 25 Count
Swamp Acanthocarpus-Stipa low
dense heath
Swamp Eucalyptus open forest
)32.000
Australia
)41.292
)41.300
1445 Count
148.883
)35.701
Rooty Breaks
Swamp
Rotten Swamp
Australia
Australia
Ringarooma
River Site 2
)37.334
Australia
Ringarooma
River Site 1
)41.296
Australia
Australia
Ringarooma
Humus 2
)41.283
Swamp Grasslands
Swamp Grasslands
Swamp Grasslands
Swamp Closed forest
147.450 870–900 Digitized Swamp Temperate rain forest and
Nothofagus cunninghamii
closed forest
147.400 870–900 Digitized Swamp Temperate rain forest and
Nothofagus cunninghamii
closed forest
147.433 870–900 Digitized Swamp Temperate rain forest and
Nothofagus cunninghamii
closed forest
147.417 870–900 Digitized Swamp Temperate rain forest and
Nothofagus cunninghamii
closed forest
148.833
1100 %Count Swamp Tall open forest
Australia
75 Count
Easter
)27.133 )109.283
Island (Chile)
Rano Raraku
RRA3
Ringarooma
Humus 1
110 Count
109.433
790
Easter
)27.183
Island (Chile)
145.583
)17.300
Quincan Crater Australia
168 %Count Swamp Open forest-tall open forest
Rano Kao
145.703
)37.859
Australia
Powelltown
425 Count
145.417
)37.500
Australia
Poley Creek
Easter
)27.087 )109.400
Island (Chile)
151.417
)31.967
Polblue Swamp Australia
1+
8 210Pb
10 +
6 210Pb
5 (1 not
used)
3
1 + top
2
3
Modern 0
0–7.4
0–5.5
0–1
0–1
3
DSFW DSFW
SEAPD
Ladd (1979d)
Dodson et al. (1998)
Dodson et al. (1998)
Dodson et al. (1998)
Dodson et al. (1998)
INDOPAC Flenley et al. (1991)
INDOPAC Flenley et al. (1991)
INDOPAC Flenley et al. (1991)
SEAPD
Pittock (1989),
Kershaw (1998)
McKenzie (1989),
McKenzie & Busby (1992)
Kershaw (1971, 1973a)
Dodson (1987), Dodson
et al. (1986)
Colhoun (1992)
DSFW TUND; INDOPAC Clark (1986), Clark &
Hope (unpubl. data)
DSFW;
DSFW
1 Modern XERO DSFW
Churchill (1960, 1968)
3
1
3
3
3
3
3
3
SEAPD
DSFW DSFW;
DSFW;
DSFW
DSFW WTRF SEAPD
XERO CTRF
WTRF WTRF;
WTRF;
WTRF
2C
STEP TDFO;
XERO;
XERO
3D
STEP TDFO;
XERO;
TDFO
2D
STEP XERO;
TDFO;
XERO
1D
CTRF DSFW;
DSFW;
DSFW
1D
CTRF DSFW;
DSFW;
DSFW
1D
CTRF CTRF;
CTRF;
DSFW
1D
CTRF DSFW;
DSFW;
DSFW
Modern DSFW DSFW
1 1D
1 n/a
3 1D
1 2C
11 (1 not 3
used)
4 (1 not
used)
2 + top
n/a
5
3 + top
Modern 1 +
15 210Pb
0–0.2
0–35.3
0–1.4
0–37.7
0–7.3
0–6
0–12
620 %Count Swamp Closed heath-sedgeland,
with closed temperate
rain forest nearby
0–5.5
1450 Digitized Swamp Eucalyptus open forest/
Leptospermum shrubland/
grassland
750 %Count Swamp Tall open forest- open forest 0–15
Rano Aroi
145.550
)41.883
Poets Hill Lake Australia
Palaeovegetation patterns for Australia and Southeast Asia
1399
1400
150
220
1280
< 20
240
Thailand 17.200 104.183
Australia )37.583 142.317
Australia )32.067 151.567
Australia )34.300 115.283
Australia )37.167 142.350
Sakon Nakhon
Salt Lake
Sapphire Swamp
Scott River Swamp
Sheet of Water
3680
3680
3680
3710
3760
)5.750 145.017
)5.750
)5.750
)5.750
)5.750
Site 20 Mt. Wilhelm PNG
Site
Site
Site
Site
145.017
145.017
145.017
145.017
3360
)5.750 145.017
Site 18 Mt. Wilhelm PNG
PNG
PNG
PNG
PNG
3510
)5.750 145.017
Site 17 Mt. Wilhelm PNG
Wilhelm
Wilhelm
Wilhelm
Wilhelm
3570
3490
3470
)5.750 145.017
)5.750 145.017
)5.750 145.017
Site 13 Mt. Wilhelm PNG
Site 14 Mt. Wilhelm PNG
Site 15 Mt. Wilhelm PNG
Mt.
Mt.
Mt.
Mt.
3570
)5.750 145.017
Site 12 Mt. Wilhelm PNG
21
22
23
25
3570
3530
)5.750 145.017
)5.750 145.017
Site 10 Mt. Wilhelm PNG
Site 11 Mt. Wilhelm PNG
Surrounding
vegetation
Swamp Open scrub
Swamp Eucalyptus fastigata
open forest
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Deschampsia forest
Shrub-rich grassland
(Gleichenia, Deschampsia, Poa)
Shrub-rich grassland
(Deschampsia klossii)
Shrub-rich grassland
Shrub-rich tussock grassland
Shrub-rich tussock grassland
with Cyathea
Forest, mangrove/Coprosma
and Poa tussock grassland
Shrub-rich Deschampsia klossii
tussock grassland
Ranunculus, Gentiana, Potentilla,
Deschampsia klossii tussock
grassland
Shrub-rich tussock grassland
Shrub-rich grassland
Coprosma/Poa tussock grassland
Deschampsia klossii tussock
grassland
%Count Swamp Open forest
Count
Count
Count Swamp Tropical savanna
%Count Swamp Open forest-woodland
Swamp Eucalyptus botryoides
open forest
Ryans Swamp 2
Count
Australia )35.150 150.650
Site name
1
Latitude Longitude Altitude Sample Site
Country ()
()
(m)
type
type
Table 1 continued
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
Modern
n/a
0–9.3
0–0.23
0–1
0–4.5
0
0
0
0
0
0
0
0
0
0
0
0
0
n/a
2 + top
1
1
3
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3
3
1
1
3
Modern
Modern
Modern
Modern
STEP
STEP
STEP
STEP
TDFO
WTRF
WTRF
WTRF
INDOPAC
INDOPAC
INDOPAC
INDOPAC
Corlett
Corlett
Corlett
Corlett
(1984a)
(1984a)
(1984a)
(1984a)
Modern STEP TDFO INDOPAC Corlett (1984a)
Modern STEP TDFO INDOPAC Corlett (1984a)
Modern CTRF WTRF INDOPAC Corlett (1984a)
Modern STEP TUND INDOPAC Corlett (1984a)
Modern STEP WTRF INDOPAC Corlett (1984a)
Modern STEP WTRF INDOPAC Corlett (1984a)
Modern STEP TDFO INDOPAC Corlett (1984a)
DSFW DSFW; INDOPAC Radclyffe (unpubl.
data)
DSFW;
DSFW
4D
TDFO XERO
Penny (1998)
DSFW DSFW SEAPD
McKenzie (unpubl.
data), D’Costa &
Kershaw (1997)
Dodson (1987),
1D
DSFW DSFW;
Dodson et al. (1986)
DSFW;
DSFW
Churchill (1968)
1C
XERO DSFW;
XERO;
XERO
n/a
DSFW DSFW SEAPD
McKenzie (unpubl.
data), D’Costa &
Kershaw (1997)
Modern CTRF TDFO INDOPAC Corlett (1984a)
Modern STEP WTRF INDOPAC Corlett (1984a)
Biomes
Age range No. of
No. of
14
(kyr)
C dates duplicates DC 0 ka Obs. Recon. Data base References
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
0
0
0
0
0
0
Australia )37.444 145.807
)5.782 145.033
Storm Creek
Summit Bog
PNG
0 to > 4 1 + top 2
Australia )40.550 144.750
Stockyard Swamp
4420 Count
Swamp Alpine tussock grassland (tundra)
1177 %Count Swamp Open forest-tall open forest
Swamp Eucalypt woodland and pasture
Australia )31.467 129.700 70–100 Count Soil
Mallee heath
Australia )37.133 146.500
1650 %Count Swamp Subalpine woodland
Stencils Cave
Stirling Moss
65 Count
Modern 0
n/a
n/a
885 %Count Swamp Tall open forest
0–10.7
0–17.5
1
1
1
3 (1 not 3
used)
3 + top 1
n/a
CTRF
STEP
CTRF
TDFO
STEP
(1984a)
(1984a)
(1984a)
(1984a)
(unpubl. data)
(1984a,b)
TUND
WTRF
WTRF
TDFO
WTRF
DSFW WSFW SEAPD
DSFW DSFW SEAPD
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
(1984a,b)
(1984a,b)
(1984a,b)
(1984a,b)
(1984a,b)
Hassell (unpubl. data)
Kershaw (1973b)
Corlett
Corlett
Corlett
Corlett
Corlett
WTRF INDOPAC Corlett (1984a,b)
WTRF INDOPAC Corlett (1984a,b)
Colhoun et al. (1992)
McKenzie (1989),
McKenzie & Busby (1992)
n/a
DSFW DSFW SEAPD
Becker (unpubl. data),
D’Costa & Kershaw (1997)
Modern XERO XERO
Martin (1973)
n/a
DSFW DSFW SEAPD
McKenzie (unpubl. data),
D’Costa & Kershaw (1997)
Modern DSFW DSFW; INDOPAC Hope (1999)
DSFW
1D
DSFW DSFW SEAPD
McKenzie (1997),
McKenzie & Busby (1992)
7
TUND WTRF; INDOPAC Hope (1976)
COCO;
WTRF
2
1 7
1 + top 1 1D
Australia )41.900 147.850
Modern
Modern
Modern
Modern
Modern
Corlett
Corlett
Corlett
Corlett
Hassell
INDOPAC Corlett
INDOPAC
INDOPAC
INDOPAC
INDOPAC
TDFO INDOPAC Corlett (1984a,b)
WTRF INDOPAC Corlett (1984a,b)
WTRF INDOPAC Corlett (1984a,b)
WTRF
WTRF
WTRF
WTRF
XERO
WTRF
WTRF INDOPAC Corlett (1984a)
1 Modern XERO XERO
1 Modern TRFO TDFO
1
1
1
1
1
1 Modern STEP
Snow Hill Marshes
n/a
STEP
STEP
STEP
STEP
XERO
STEP
1 Modern STEP
Australia )42.184 145.633
Australia )37.391 145.928
0–12
0–12
Modern
Modern
Modern
Modern
Modern
Modern
1 Modern STEP
1 Modern STEP
1 Modern STEP
1
1
1
1
1
1
1 Modern STEP
Smelter Creek
Snobs Creek
0
0
0
0
0
Modern 0
Modern 0
Modern
Modern
Modern
Modern
Modern
Modern 0
Modern 0
Modern 0
Modern 0
Modern 0
Modern
Modern
Modern
Modern
Modern
Modern
Modern 0
Trap
Moss
polster
210 %Count Swamp Scrub-closed forest
930 %Count Swamp Tall open forest
127 Count
1040 Count
Soil
Soil
Soil
Soil
Soil
Brachypodium-Carex fen,
Deschampsia klossii tussock grassland
Tussock grassland
Coprosma/Poa tussock grassland
Deschampsia klossii tussock grassland
Deschampsia klossii tussock grassland
Open mallee shrubland
Deschampsia klossii tussock
grassland/disturbed
Tussock grassland/disturbed
Scirpus crassiusculus fen/disturbed
Coprosma/Poa tussock
grassland/disturbed
Tussock grassland (Carex sp.)/
disturbed
Deschampsia klossii tussock
and fen/disturbed
Forest
Tussock grassland/disturbed
Coprosma papuensis and forest
Styphelia, Coprosma, Gaultheria
Coprsoma, Poa, Monostachya,
Ranunculus, Astelia/disturbed
Open mallee shrubland
Mixed notophyll vine forest
Australia )33.021 119.915
Australia )17.517 145.567
Count
Count
Count
Count
Count
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Trap
Soil
Soil
Site 63 FRNP
Sluice Creek
145.017
145.017
145.017
145.017
145.017
3750
3590
3580
3700
3800
PNG
PNG
PNG
PNG
PNG
)5.750
)5.750
)5.750
)5.750
)5.750
Wilhelm
Wilhelm
Wilhelm
Wilhelm
Wilhelm
Site
Site
Site
Site
Site
Mt.
Mt.
Mt.
Mt.
Mt.
3680 Count
)5.750 145.017
Site 45 Mt. Wilhelm PNG
46
47
49
51
54
3320 Count
Count
Count
Count
Count
Count
Count
)5.750 145.017
3660
3670
3460
3330
81
3320
Site 43 Mt. Wilhelm PNG
PNG
)5.750 145.017
PNG
)5.750 145.017
PNG
)5.750 145.017
PNG
)5.750 145.017
Australia )34.188 119.414
PNG
)5.750 145.017
3740 Count
3910 Count
3680 Count
Mt. Wilhelm
Mt. Wilhelm
Mt. Wilhelm
Mt. Wilhelm
FRNP
Mt. Wilhelm
3630 Count
)5.750 145.017
)5.750 145.017
)5.750 145.017
29
30
31
34
36
36
)5.750 145.017
Site 37 Mt. Wilhelm PNG
Site 39 Mt. Wilhelm PNG
Site 41 Mt. Wilhelm PNG
Site
Site
Site
Site
Site
Site
Site 27 Mt. Wilhelm PNG
Palaeovegetation patterns for Australia and Southeast Asia
1401
1402
Swamp Epacris-Leptospernum
heath
1580
160
780
1158
1000
350
3820
1250
)4.117 138.967
)37.150 145.483
)35.668 148.033
)42.667 146.500
)41.917 145.583
)36.683 147.133
)5.787 145.008
)37.750 146.083
Irian Jaya
(Indonesia)
Australia
Australia
Australia
Australia
Australia
PNG
Supulah Hill
Tamaringa
Billabong
Tarcutta
Swamp
2
60
1160
1070
1075
1500
20
20
)39.033 146.317
)38.117 145.267
)17.117 145.567
)17.117 145.517
)37.386 145.818
)32.000 151.467
)38.317 142.367
)38.317 142.367
Australia
Tiger Snake
Australia
Swamp
Tinaroo Range Australia
Australia
Tom Burns, D Australia
Australia
Australia
Australia
Tinaroo Road
Top Swamp
Tower Hill
Main
Tower Hill
NW Crater
Tidal River
4
9.545 138.183
Micronesia
Australia
Thompson
River
Thool Swamp
Tarn Shelf
Tarn Shelf,
Mt Field
Tawonga Bog
Tegepangwa
0–3.5
0–11.5
0–7.7
0–1.2
0–10
0–9.8
0.2–9.8
Swamp Local gardens, savanna 0–2.3
with grassland
Swamp Eucalyptus delegatensis
forest/temperate
sclerophyll forest
Swamp Sedgeland-herbfield
Swamp Subalpine forest-alpine
heath
Swamp Open forest
Soil
Carpha alpina, Scripus
and Caren grass bog
Swamp Tall open forest
Tall open forest
0–32
Simple notophyllModern
microphyll vine forest
Wet sclerophyll forest Modern
%Count Swamp Open forest-woodland 0–20
Digitized Swamp Eucalyptus pauciflora 0–3
open forest
%Count Lake
Open forest-woodland 0–20
Moss
polster
Count
Moss
polster
%Count Swamp
Count
Swamp Eucalytpus obliqua tall 0–4.9
forest
%Count Swamp Woodland
0–7
Count
Count
%Count
%Count
Count
%Count
%Count
Count
1
1
1
1
2
3
1
1
2
1
1
2
1
3
3
6
1
TUND CTRF
XERO CTRF
SEAPD
SEAPD
Macphail (1979)
Macphail (1986)
XERO DSFW; INDOPAC Hope (1999)
DSFW;
DSFW
WTRF WTRF; INDOPAC Swadling & Hope (1992)
TDFO;
WTRF
WSFW DSFW SEAPD
Reid (unpubl. data),
D’Costa & Kershaw (1997)
DSFW DSFW; INDOPAC Williams (unpubl. data)
DSFW
References
1C
1D
DSFW DSFW SEAPD
DSFW DSFW SEAPD
DSFW DSFW
DSFW DSFW SEAPD
Modern WSFW WTRF
Modern WTRF WTRF
D’Costa et al. (1989)
McKenzie (1997),
McKenzie &
Busby (1992)
Dodson (1987),
Dodson et al. (1986)
D’Costa et al. (1989)
Kershaw (1973b)
Kershaw (1973b)
Modern DSFW DSFW SEAPD
Kershaw & Green (1983)
2D
TUND TDFO; INDOPAC Corlett (1984b)
TDFO
n/a
DSFW CTRF SEAPD
Strickland (unpubl. data),
Kershaw (1998)
Dodson & Intoh (1999)
1D
TDFO TDFO;
TDFO;
TDFO
7
DSFW DSFW; INDOPAC Hope (1974)
DSFW
Modern DSFW DSFW SEAPD
Aitken & Kershaw (1993)
n/a
2C
n/a
1C
Biomes
No. of
Recon. Data base
duplicates DC 0 ka Obs.
3 (1 dup) 1
+ top
6
1
1 + top
0
0
1
2
3
n/a
1 + top
2 tephra
3
2 + top
2
n/a
3
2 + top
Age range No. of
14
(kyr)
C dates
Swamp Lower montane closed 0–3.3
forest/agriculture and
regrowth forest
%Count Swamp Forest
n/a
Count
Count
5
Country
)41.117 144.667
Site name
Sundown Point Australia
Surrounding
vegetation
Site
type
Latitude Longitude Altitude Sample
()
()
(m)
type
Table 1 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
PNG
142.583
145.800
)5.667
)41.583
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
146.450
178.033
121.383
128.417 60–70 Count Soil
Mallee heath
147.633
10 %Count Swamp Low coastal heathland
115.850
115.750
115.850
116.517
)42.933
)18.417
)2.550
)31.707
)40.867
)30.317
)30.300
)30.283
)34.683
Australia
Fiji Islands
Sulawesi
(Indonesia)
146.250
)42.533
Australia
143.450
116.600
144.000
144.000
144.000
144.000
)38.317
)34.467
)5.750
)5.750
)5.750
)5.750
Australia
Australia
Australia
Australia
PNG
PNG
PNG
PNG
Watheroo NP (9)
Weld River
Swamp
West Basin
West Lake
Muir Swamp
Weylk 3
Weylk 4
Weylk 6
Weylk 7
Trap
Trap
Swamp Tropical mixed forest
Swamp
Banksia low woodlands
Eucalyptus loxophleba
woodland
240 Count Trap
Banksia low woodlands
10 Count Swamp Eucalyptus marginata,
E. calophylla and
E. diversicolor dense woodland
110 %Count Lake
Open woodland-grassland
100 Count Swamp Eucalyptus calophylla and
E. rudis woodland
1885 Count Soil
Garden area
1885 Count Soil
Garden area
1885 Count Soil
Sweet potato garden area
1885 Count Soil
Garden area
300 Count
390 Count
445 Count
1 Count
545 %Count Swamp Tall closed forest
1040 %Count Swamp Subalpine forest-alpine heath
2 Count
Swamp
169.849
)20.249
Vanuatu
3550 Count
145.053
)5.792
Swamp Prosopis, Leucaena,
Pithecellobium
Soil
Forest
PNG
2 Count
21.517 )158.250
Swamp Mid-montane mixed forest
with Castanopsis, Lithocarpus
and Araucaria/anthropogenic
grassland & gardens
230 %Count Swamp Tall open forest–closed forest
(eucalypts and cool-temperate
rain forest)
1000 Count Soil
Eucalypt woodland
2 Count Swamp Prosopis, Leucaena,
Pithecellobium
2300 Count
Hawaii (USA)
Wanteen
Australia
Waterhouse
Australia
Marsh
Watheroo NP (10) Australia
Watheroo NP (8) Australia
Wanda
Upper Lake
Wurawina
Upper Timk Lake
Mt Anne
Vunimoli Swamp
Uko’a Pond
Core 3
Umbamambuno
(Imbuka)
Umej
Tumoulin Swamp Australia
)17.550 145.450
Uko’a Pond
Hawaii (USA) 21.517 )158.250
Core 2
Tullabardine Dam Australia
Tugupugua
(Core 1)
0–10
0–7.76
Modern
0–6.8
3 + top
1
0
2
n/a
1
4
2
0
6
7 + top
CTRF;
CTRF;
CTRF
WSFW CTRF
CTRF
SEAPD
Ward (unpubl. data),
Hope et al. (1999)
Corlett (1984a,b)
Kershaw (1976)
Ward (unpubl. data),
Hope et al. (1999)
Colhoun & van de
Geer (1986)
Haberle (1998), Haberle
et al. (1999, 2001)
CTRF
COCO SEAPD
TDFO XERO INDOPAC Hope & Spriggs (1982),
Hope et al. (1999)
XERO COCO SEAPD
Macphail (1986)
1
1
1
1
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
INDOPAC
INDOPAC
INDOPAC
INDOPAC
1 Modern XERO DSFW
3 3D
DSFW WTRF;
TDFO;
WTRF
1 2C
XERO DSFW SEAPD
1 Modern DSFW DSFW
1 n/a
1 7
1 n/a
1 Modern DSFW TDFO
3 n/a
TDFO XERO; INDOPAC
TDFO;
STEP
2 n/a
TDFO TDFO; INDOPAC
STEP
1
CTRF WTRF INDOPAC
1 1C
3 4D
Powell
Powell
Powell
Powell
(1970)
(1970)
(1970)
(1970)
Gell et al. (1994)
Churchill (1968)
Pickett (1997)
Churchill (1968)
Harle (1989), D’Costa &
Kershaw (1997)
0–2.7
3
2 n/a
WTRF WTRF; INDOPAC Southern (1986),
WTRF
Hope et al. (1999)
0 to > 40 11 (4 not 3 7
TDFO TDFO; INDOPAC Hope (2001)
used)
TDFO;
TDFO
Modern 0
1 Modern XERO XERO
Martin (1973)
0–10.4
n/a
1 n/a
XERO DSFW SEAPD
Thomas (unpubl. data),
D’Costa & Kershaw (1997)
Modern 0
1 Modern XERO DSFW
Pickett (1997)
Modern 0
1 Modern DSFW DSFW
Pickett (1997)
0–9
0–10.5
0–5.5
0–4.3
Modern
0–7
0–43.8
0 to > 13 5
Palaeovegetation patterns for Australia and Southeast Asia
1403
1404
4500
10
1580
1825
1825
1825
1580
1580
1580
1825
1885
2220
1675
450
< 20
< 20
< 20
650
)5.500 143.417
Australia )37.750 149.433
144.167
144.000
144.000
144.000
144.167
144.167
144.167
144.000
144.000
144.000
144.000
143.461
PNG
)5.733
PNG
)5.750
PNG
)5.750
PNG
)5.750
PNG
)5.733
PNG
)5.733
PNG
)5.733
PNG
)5.750
PNG
)5.750
PNG
)5.750
PNG
)5.750
Australia )38.647
Australia )32.867 115.667
Australia )31.283 116.067
Australia )31.283 116.067
Australia )42.883 147.300
Wriggly
Stick Swamp
Wurup 2
Wurup 3
Wurup 4
Wurup 5
Wurup 6
Wurup 7
Wurup 8
Wurup Core 1
Wurup slopes 2
Wurup slopes 203
Wurup slopes 4
Wyelangta
Yalgorup
Yanchep NP (4)
Yanchep NP (5)
Yarlington Tier
760
Australia )17.317 145.783
Windin
PNG
20
Australia )37.783 145.050
Willsmere
Billabong
WISU
1885
3850
)5.750 144.000
)5.500 143.417
PNG
PNG
Site
type
Surrounding
vegetation
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
Count
%Count
Mixed grasses
Garden beds
Garden beds
Taro garden bed
Grassland swamp
Grassland swamp
Grassland swamp
Garden area
Regrowth forest
Oak forest
Forest edge
Closed forest (temperate
rain forest)
Trap
Eucalyptus gomphocephala
open woodland
Trap
Eucalyptus gomphocephala
open woodland
Trap
Banksia low woodlands
Swamp Dry sclerophyll Eucalyptus
forest
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Soil
Swamp
Moss Complex mesophyll
polster vine forest
Count Moss
polster
%Count Swamp Open forest
Count
Soil
Garden area
Moss Above the forest limit
polster
%Count Swamp Woodland
Count
Count
Latitude Longitude Altitude Sample
Country ()
()
(m)
type
continued
Weylk 8
WI10
Site name
Table 1
n/a
0
n/a
Modern
0–9.1
Modern
Modern
0
2 + top
0
0
0 to > 46 5 + top
n/a
Modern
n/a
1
3
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
INDOPAC
SEAPD
Pickett (1997)
Kenyon (unpubl. data),
D’Costa & Kershaw
(1997)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
Powell (1970)
McKenzie &
Kershaw (2000)
Pickett (1997)
Modern XERO DSFW
Pickett (1997)
Modern DSFW CTRF; INDOPAC Harle et al. (1993)
TUND;
DSFW
Modern DSFW DSFW
Modern DSFW DSFW
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
TDFO
CTRF
DSFW DSFW SEAPD
STEP
TDFO
TDFO
TDFO
STEP
STEP
STEP
STEP
TDFO
TSFO
WTRF
Modern WSFW
n/a
McNess (1988),
D’Costa &
Kershaw (1997)
Kershaw (1973b)
WTRF WTRF INDOPAC Flenley (1967, 1969)
WTRF
DSFW WSFW SEAPD
Modern TRFO
n/a
References
TDFO TDFO INDOPAC Powell (1970)
TUND WTRF INDOPAC Flenley (1967, 1969)
Biomes
Age range No. of
No. of
14
Recon. Data base
(kyr)
C dates duplicates DC 0 ka Obs.
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
107.728
)6.979
Bandung DPDR-II Java
1524
1080
35
1500
1890
150
1
100
1
1170
Australia )32.002 115.503
Australia )37.850 146.267
Australia )36.518 149.500
Australia )32.100 115.833
Australia )32.050 151.467
Australia )36.417 148.317
Australia )37.618 140.517
Australia )36.418 150.100
Australia )35.017 116.633
Australia )31.868 151.517
Barker Swamp
Baw Baw Village
Bega Swamp
Bibra Lake (BL2)
Black Swamp
Blue Lake
(Snowy Mts)
Blue Tea Tree
Swamp
Bobundarra
Swamp
Boggy Lake
Boggy Swamp
662
661
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Swamp 0–13.5
Swamp 0–5
Swamp 0–7.5
7
3 + top
48
n/a
3 + top
7+
5 U/Th
3
3
Swamp 0–6.8
Swamp 0–7.27
14
No C
Dates
3
3
2
3
n/a
3
3
1
3
2
n/a
yes
2C
5D
n/a
7
n/a
n/a
2C
2C
n/a
1C
5D
n/a
n/a
n/a
n/a
n/a
n/a
n/a
1
n/a
n/a
n/a
n/a
3
2
3
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
1C
5D
n/a
DSFW;
DSFW;
DSFW
DSFW;
DSFW;
DSFW
TUND;
DSFW
DSFW;
TDFO;
DSFW
DSFW;
DSFW;
DSFW
DSFW;
DSFW;
DSFW
DSFW;
DSFW;
DSFW
XERO;
DSFW;
XERO
DSFW
TDFO;
WTRF
Data base
References
DSFW
Dodson & Wilson (1975)
Dodson (1987),
Dodson et al. (1986)
Churchill (1968)
INDOPAC Coddington (1983)
SEAPD
INDOPAC Raine (1974, 1982)
Strickland (unpubl. data),
D’Costa & Kershaw (1997)
INDOPAC Polach & Singh (1980),
Green et al. (1988),
Hope et al. (2000)
Pickett (1997)
Dodson (1987),
Dodson et al. (1986)
SEAPD
Haberle (1994)
CTRF;
CTRF;
CTRF
TDFO; INDOPAC van der Kaars &
TDFO
Dam (1995, 1997)
TUND; INDOPAC van der Kaars &
Dam (1995, 1997)
TDFO;
TDFO
Backhouse (1993)
Reconstructed
No of
No of
biomes
DC
6 ka
DC 18 ka
18 ka
duplicates 6 ka duplicates 18 ka 6 ka
12 (2 not 3
used)
3 + top
3
1
0–13.34 16
Swamp 0–8.5
Lake
Digitized Swamp 0–9.3
Count
Count
%Count
Count
Count
Lake
0–31
Digitized Swamp 0–8.6
Count
%Count
Count
Count
Count
Swamp 0 to
> 135
Swamp 0 to
> 135
107.600
)6.835
Java
Bandung DPDR-I
Count
Swamp 0–18.5
2750
143.153
)5.972
PNG
Aluaipugua
(core 2)
Site
type
Country
Site name
Latitude Longitude Altitude Sample
()
()
(m)
type
Age
range
(kyr)
Table 2 Characteristics of the 6000 and 18,000 14C yr bp pollen sites from the SEAPAC region. The sites are arranged alphabetically by name. Latitude and longitude are given in decimal degrees,
with N and E conventionally indicated as +, and S and W as ). Dating control (DC) follows the COHMAP dating scheme, as described by Yu & Harrison (1995). The codes used for reconstructed
biomes are defined in Table 5
Palaeovegetation patterns for Australia and Southeast Asia
1405
1406
3910
700
700
5
755
180
1330
1260
235
1280
29
1955
1955
140
1755
)5.783 145.035
)34.800 149.500
)34.750 149.520
)38.301 141.383
)17.383 145.550
)38.250 143.117
)36.783 146.767
)31.917 151.400
)37.448 145.692
)37.333 146.750
)38.626 143.323
)36.417 148.283
)36.417 148.283
)38.300 145.033
)35.969 148.817
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Breadalbane NW
Breadalbane SE
Bridgewater
Lake, Core A
Bromfield Swamp
Bullenmerri
Bunyip Bog
Butchers Swamp
Buxton
Caledonia Fen
Chapple Vale
Club Lake
core 1
Club Lake core 2
Cobrico Swamp
Cotter Source
Bog
PNG
9
Australia
Boomer Swamp,
Core 12
Brass Tarn
Site
type
Age
range
(kyr)
Lake
Swamp 0–11
0–7.5
0–9.3
0–6.8
5
Swamp 0–6.7
5 + top
7 (1 not
used) + 1
on section
Count
Lake
6.8 to 6 (3 not
> 10 used)
%Count Swamp 0–9
4+
8 210Pb
Count
Swamp 0–5.7 2
Count
%Count Swamp 0–6
1
Digitized Swamp 0–11.3 14 (4 not
used)
%Count Swamp 0–12 2 + top
%Count Swamp 0–25
5
1
1
1
2
3
1
1
1
2
n/a
1
3
n/a
2C
4D
1C
1D
n/a
1D
3C
n/a
3C
3C
1C
1C
2D
2C
2D
n/a
n/a
n/a
n/a
n/a
1
n/a
n/a
1
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
6D
n/a
n/a
n/a
n/a
n/a
n/a
n/a
XERO
SEAPD
SEAPD
SEAPD
SEAPD
Data base
Kershaw (1973a, 1975)
Dodson (1983, 1986)
Dodson (1983, 1986)
Head (1988)
Hope (1976), Hope &
Peterson (1975)
Head (1983, 1988)
References
Dodson (1979)
Binder (1978), Binder &
Kershaw (1978)
DSFW;
Dodson (1987),
DSFW
Dodson et al. (1986)
CTRF
SEAPD
McKenzie (1989),
McKenzie & Busby
(1992)
DSFW XERO SEAPD
Joyce (1979),
Kershaw et al. (1983),
Kershaw (1998)
CTRF
SEAPD
McKenzie & Kershaw
(1997)
INDOPAC Martin (1986)
DSFW;
DSFW;
DSFW
DSFW;
INDOPAC Martin (1986)
DSFW
DSFW
SEAPD
Yezdani (1970),
Dodson et al. (1994a)
XERO
INDOPAC Hope & O’Dea
(unpubl. data)
DSFW
WTRF;
WTRF;
WTRF
WTRF;
WTRF;
WTRF;
WTRF;
TDFO
DSFW
DSFW
DSFW
DSFW
Reconstructed
No of
No of
biomes
DC
6 ka
DC 18 ka
18 ka
duplicates 6 ka duplicates 18 ka 6 ka
5
1
10 (1 not used) 1
2
1
0–10.8 5
2
No 14C
Dates
Count
Lake
8–16.1 4
%Count Swamp 0–12 2 + top
Count
%Count Lake
%Count Lake
Count
Lake
Count
%Count Swamp 0–5.6
Latitude Longitude Altitude Sample
()
()
(m)
type
)38.217 141.300
Country
Site name
Table 2 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
137.217
149.133
149.133
)4.033
)37.100
)37.100
Australia
151.367
)31.917
Jacksons Bog B
147.700
137.383
)37.967
)3.985
Australia
142.783
)5.833
PNG
Jacksons Bog A
132.164
)3.599
Indonesia
G5–2-053P
(Seram Trench)
Haeapugua,
Core 6
Indonesia
142.317
)37.585
Australia
Freshwater Lake
Ijomba
146.467
)38.950
Australia
Australia
119.458
)34.292
Australia
Fitzgerald
Inlet Core 2
Five Mile Beach
Horse Swamp
123.612
)19.670
Australia
Australia
Indonesia
148.483
)36.383
Australia
Hidden Swamp
Hogayaku Lake
148.833
)37.250
Australia
Delegate River/
Tea Tree Swamp
Digger’s Creek
Bog
Dragon Tree Soak
146.300
)39.117
Australia
100.764
)1.072
Darby Beach
146.783
)36.733
Australia
Sumatra
(Indonesia)
145.267
)38.117
Australia
Danau Di Atas
Cranbourne
Annexe (Tadpole Swamp)
Crystal Bog 1
750
750
3630
1260
2
3580
1650
Marine
220
20
0
200
1690
900
1
1535
1350
60
%Count
Count
Count
Digitized
%Count
Count
Count
Count
%Count
Count
Count
Digitized
Count
%Count
Count
Count
%Count
Count
Swamp
Swamp
Swamp
Swamp
Swamp
Lake
Swamp
Marine
Lake
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
0–12
0–13
0–13.9
0–11
0–7.5
0–6.5
0–29
0-ca14
0–7
0–13.9
0–7
0–6.4
0–10.5
0–12
0–6
0–31.5
0–15
0–9
n/a
4
3
7 (1 not
used)
0 (isotope
stratig.)
11 (1 not
used) +
2 tephra
2 + top
2
n/a
2
1
2
3 (1 not
used)
5
1
10 (2 not
used)
3
3
1
2
2
3
1
4
3
1
1
3
1
2
2
1
2
2
1
1
n/a
n/a
2D
4C
1C
n/a
2C
7
n/a
4D
n/a
2D
1C
7
1D
1C
n/a
1C
n/a
n/a
n/a
n/a
n/a
n/a
3
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
2
1
n/a
n/a
n/a
n/a
n/a
n/a
n/a
2C
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
1D
7
n/a
WTRF;
WTRF;
WTRF
DSFW
COCO;
COCO;
COCO;
COCO
DSFW;
TDFO;
TDFO
TDFO;
TDFO
TUND;
DSFW
DSFW
TDFO
XERO;
XERO;
XERO
DSFW
DSFW;
DSFW
STEP;
STEP
XERO
DSFW;
DSFW
DSFW
TDFO;
TDFO
DSFW
DSFW
CTRF;
CTRF;
CTRF
WTRF;
WTRF
DSFW
SEAPD
INDOPAC
INDOPAC
SEAPD
INDOPAC
INDOPAC
SEAPD
INDOPAC
INDOPAC
SEAPD
INDOPAC
SEAPD
Hope & Peterson (1975,
1976), Haberle et al. (2001)
Southern (1982),
Kershaw (1998)
Southern (1982),
Kershaw (1998)
Dodson (1987),
Dodson et al. (1986)
Hooley et al. (1980)
Hope (unpubl. data)
Haberle (1993, 1998),
Haberle et al. (1999)
Kershaw & McKenzie
(unpubl. data),
D’Costa & Kershaw (1997)
van der Kaars (1991, 1995)
Ladd (1979a)
Pedersen (1983),
Wyrwoll et al. (1986)
Hassell (unpubl. data)
Martin (1999)
Ladd (1979b,c)
Williams (1980, 1982),
Kershaw (1998)
Newsome (1985),
Stuijts et al. (1988),
Newsome & Flenley, 1988
Hope (1974)
Aitken & Kershaw (1993)
Palaeovegetation patterns for Australia and Southeast Asia
1407
1408
PNG
PNG
Komanimambuno
Kosipe
147.200
)8.467
75
60
5
8
673
102
3120
1300
780
)38.250 142.450
)38.067 141.833
)37.833 148.500
)37.750 149.450
)35.083 149.417
)38.233 143.100
138.700
)4.117
)37.150 147.933
)2.553
Australia
Australia
Australia
Australia
Australia
Australia
Indonesia
Australia
Irian Jaya
(Indonesia)
Lake Condah
(Core 1b)
Lake Curlip
Lake Elusive
Lake George
Lake Gnotuk
Lake Habbema
Lake Hill
Lake Hordorli
140.550
35
)32.008 115.817
Australia
Lake Banganup
(BGP4)
Lake Cartcarong
730
)17.167 145.633
Australia
235
New Caledonia )22.300 166.983
Lake Euramoo, A
Lac Suprin
2
2
1960
1.5–9.7
0–33.3
0–8.2
Lake
Lake
0–10
0–16
0–128
Count
Lake
2
1
1
3
n/a
2
n/a
1
15 (on 1
multiple
cores)
3 + top 3
17
2 + top 1
n/a
1
2 + top 1
n/a
2
7
4
3
1
5 + top 3
0 to > 35 7 (1 not 2
used)
%Count Swamp 0–15
Count
%Count Lake
Count
%Count Lake
0–5.2
%Count Swamp 0 to > 6
%Count Lake
0–11
Swamp 0–31
Lake
Lake
Swamp 0–6.5
Swamp 0–8
%Count Lake
Count
Count
Count
Count
Count
Count
4
No 14C
Dates
5D
n/a
2C
1C
4C
4D
n/a
3C
n/a
n/a
1C
7
1C
1C
2C
4C
2
1
n/a
n/a
2
n/a
n/a
n/a
n/a
1
n/a
2
n/a
n/a
3
yes
6C
n/a
n/a
n/a
7
n/a
n/a
n/a
n/a
n/a
n/a
2D
n/a
n/a
7
3C
Data base
References
R. Walkley (unpubl. data),
Kershaw (1998)
Head (1989)
Kershaw & McKenzie
(unpubl. data),
Kershaw (1998)
WTRF; CTRF; INDOPAC Hope et al. (1988),
WTRF WSFW
Hope & Tulip (1994)
SEAPD
SEAPD
SEAPD
SEAPD
SEAPD
Ladd (1978)
Kenyon (unpubl. data),
Kershaw (1998)
TRFO; INDOPAC Singh & Geissler (1985)
TRFO
SEAPD
Yezdani (1970),
Harrison (1989),
Kershaw (1998)
INDOPAC Haberle et al. (2001)
TDFO;
TDFO;
TDFO
DSFW XERO
TDFO;
DSFW
DSFW
DSFW
WSFW
XERO
DSFW
INDOPAC Hope (1976),
Hope & Peterson (1975)
WTRF; INDOPAC Hope (1982),
Haberle et al. (2001)
CTRF;
CTRF
TDFO
INDOPAC Hope et al. (1999)
INDOPAC Martin (1994)
DSFW;
DSFW;
DSFW
WTRF; INDOPAC Hope (1996),
TSFO
Hope & Pask (1998)
WTRF;
Kershaw (1970, 1973a,
WTRF;
1974, 1979)
WTRF
DSFW DSFW
Pickett (1997)
WTRF;
TDFO
Reconstructed
No of
No of
biomes
DC
6 ka
DC 18 ka
18 ka
duplicates 6 ka duplicates 18 ka 6 ka
Swamp 0 to > 30 5 + top yes
Swamp 0–21.76
Count
145.067
)5.800
2740
Site
type
Age
range
(kyr)
Latitude Longitude Altitude Sample
()
()
(m)
type
Koumac Northeast New Caledonia )20.650 164.283
Kurnell Fen Core 1 Australia
)34.017 151.167
Country
Site name
Table 2 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
138.067
107.617
115.667
147.683
141.083
145.700
127.083
148.517
146.417
148.067
140.317
146.300
115.717
)35.767
)6.735
)31.533
)37.967
)38.084
)17.367
)31.933
)35.334
)38.250
)37.117
)37.600
)37.800
)33.117
Java
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Australia
Lembang Swamp/
Panjairan
Loch McNess Swamp
Loch Sport Swamp
Long Swamp
Lynch’s Crater A
Madura
Micalong Swamp
Morwell Swamp
Moss Bed Lake
Mount Burr
Swamp
Mustering Flat
Myalup Swamp
Australia
Lashmar’s Lagoon
147.367
)8.384
PNG
Laravita Tarn
142.867
142.600
)35.250
)38.350
Australia
Australia
Lake Tyrrell (Site 2)
Lake Wangoom
142.750
)37.700
Australia
Lake Turangmoroke
)179.933
)16.817
Fiji Island
Lake Tagamaucia
145.875
)37.469
Australia
Lake Mountain
142.867
140.583
)38.200
)37.650
Australia
Australia
Lake Keilambete
Lake Leake (core LE)
143.450
)5.451
PNG
Lake Inim M4
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
10
1448
150
1170
40
100
980
1
2
760
< 20
1200
2
3640
17
100
200
820
1440
150
150
2500
Count
%Count
%Count
%Count
%Count
Count
Count
%Count
%Count
Count
Count
Count
%Count
Count
%Count
%Count
%Count
Count
%Count
%Count
Count
Count
Lake
Swamp
Swamp
Swamp
Swamp
Cave
Swamp
Swamp
Swamp
Lake
Lake
Swamp
Lake
Lake
Lake
Lake
Lake
Lake
Swamp
Lake
Lake
Lake
0–11.3
0–6
0–7.5
0–7
0–9.5
0–8
0–13.07
0–7.5
0–6
0–190
0–8.9
0 to > 100
0–7
0–13
0–10
0–43
0–20
0–14.3
0–6.5
0–9.7
0–8.5
0 to > 7
1
n/a
3
n/a
2 + top
3
6
2 + top
2
10
2
2
5
0 (poll)
+ top
0 (sed corr.)
8 (1 not
used)
3
6
1 + top
6 (2 not
used)
4 + top
5 + top
2
1
1
1
1
1
3
1
1
2
2
n/a
1
1
1
1
1
3
1
1
3
1
3D
n/a
1C
n/a
6D
1C
n/a
1C
1D
3D
n/a
n/a
n/a
n/a
7
1C
7
n/a
3D
1C
2C
1C
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
3
n/a
2
n/a
n/a
n/a
1
1
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
7
n/a
7
n/a
n/a
n/a
7
7
n/a
n/a
n/a
n/a
n/a
TDFO;
TDFO
DSFW
DSFW
DSFW
XERO
DSFW;
DSFW;
DSFW
DSFW
DSFW;
DSFW
DSFW
XERO
WTRF;
TDFO
XERO
XERO
XERO
DSFW
TDFO;
WTRF;
WTRF
XERO
DSFW
XERO;
XERO;
XERO
DSFW
CTRF
TDFO;
TDFO;
TDFO
XERO;
WTRF
XERO
XERO
SEAPD
SEAPD
SEAPD
SEAPD
INDOPAC
SEAPD
SEAPD
INDOPAC
INDOPAC
SEAPD
INDOPAC
SEAPD
SEAPD
SEAPD
INDOPAC
SEAPD
SEAPD
SEAPD
INDOPAC
Strickland (unpubl. data),
Kershaw (1998)
Churchill (1968)
Lloyd (1991),
Lloyd & Kershaw (1997)
Kershaw & McKenzie
(unpubl. data), Kershaw (1998)
Dodson & Wilson (1975)
Martin (1973)
Kemp (unpubl. data)
Hooley et al. (1980)
Head (1988)
Kershaw (1974, 1976, 1994)
Newsome & Pickett (1993)
Hope (1980), Hope &
O’Dea (unpubl. data)
Clark (1983a,b),
D’Costa & Kershaw (1997)
van der Kaars & Dam (1995)
Luly (1993)
Edney et al. (1990)
Crowley & Kershaw (1994)
McKenzie (1997),
McKenzie & Busby (1992)
Southern (1986),
Hope et al. (1999)
Flenley (1972, 1984),
Walker & Flenley (1979)
Dodson (1974a)
Dodson (1974b, 1975a)
Palaeovegetation patterns for Australia and Southeast Asia
1409
1410
17.100
Thailand
Thailand
Australia
Australia
Nong Han
Kumphawapi
(Core 3)
Nong Pa Kho
North Lake
North Torbreck
Oaks Creek
564
610
10
1450
750
168
790
425
75
1100
1445
5
)37.481 146.947
)37.586 146.021
)22.267 166.617
)31.967 151.417
)37.500 145.417
)37.859 145.703
)17.300 145.583
)27.087 )109.400
)27.133 )109.283
)37.334 148.833
)35.701 148.883
)22.234 166.55
Australia
Plum (Plum
New
Swamp, Core 2) Caledonia
Australia
Australia
Australia
Australia
Easter Island
(Chile)
Easter Island
(Chile)
Australia
Australia
Polblue Swamp
Poley Creek
Powelltown
Quincan Crater
Rano Aroi
Saint Louis Lac
New
Caledonia
35
)32.008 115.817
Rano Raraku
RRA3
Rooty Breaks
Swamp
Rotten Swamp
175
170
100
102.933
103.033
17.183
Australia
N145
0–35
Count
Count
%Count
Count
Count
Count
%Count
%Count
2 + top
1 + top
6
8 (1 not
used) +
top
3
5
No 14C
Dates
1
1
2
0–7.3
0–35.3
Swamp 0–6.6
Swamp 0–7.4
Swamp 0–5.5
Lake
Swamp 0–37.7
Lake
Swamp 0–6
Swamp 0–15
3
1
1
1
9
3
2
2
11 (1 not 2
used)
10 +
yes
6 210Pb
1
1 + top
4 (1 not
used)
2 + top
n/a
5
1C
n/a
2D
4D
4C
2D
1C
n/a
3D
1C
1D
5D
n/a
4D
1C
3
yes
2C
yes
n/a
n/a
n/a
1
2
n/a
n/a
1
n/a
3
n/a
n/a
1
3
n/a
1
n/a
n/a
n/a
6C
7
n/a
n/a
n/a
n/a
2C
n/a
n/a
n/a
7
n/a
6C
Data base
DSFW;
DSFW
WTRF;
WTRF
WSFW
WTRF;
WTRF;
WTRF
XERO; STEP;
TDFO XERO
TDFO
DSFW
WTRF
WTRF;
WTRF;
WTRF
DSFW
CTRF
Pickett (1997)
Penny (1998)
Penny (1998, 1999),
Kealhofer & Penny (1998)
Martin (1973)
References
INDOPAC Clark (1986), Clark &
Hope (unpubl. data)
INDOPAC Stevenson (in press)
INDOPAC Flenley & King (1984),
Flenley et al. (1991)
INDOPAC Flenley & King (1984),
Flenley et al. (1991)
SEAPD
Ladd (1979d)
McKenzie (1989),
McKenzie & Busby (1992)
SEAPD
McKenzie (1989),
McKenzie & Busby (1992)
TRFO; INDOPAC Stevenson et al. (2001)
WTRF;
WTRF
Dodson (1987),
Dodson et al. (1986)
DSFW SEAPD
Pittock (1989),
Kershaw (1998)
SEAPD
McKenzie (1989),
McKenzie & Busby (1992)
Kershaw (1971, 1973a)
TDFO;
TDFO;
TDFO
DSFW; DSFW
DSFW
CTRF
SEAPD
XERO;
XERO;
XERO
XERO
Reconstructed
No of
No of
biomes
DC
6 ka
DC 18 ka
18 ka
duplicates 6 ka duplicates 18 ka 6 ka
Swamp 0 to > 23 12 (2 not 3
used)
Swamp 0–6.8
Swamp 0–12
Lake
Swamp 0–38.3
Lake
2.6 to
> 20.2
0–14
Age
range
(kyr)
Digitized Swamp 0–5.5
Count
%Count
%Count
Count
Count
Count
Count
Cave
)31.617 129.167
Country
Site
type
Latitude Longitude Altitude Sample
()
()
(m)
type
Site name
Table 2 continued
E. J. Pickett et al.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
116.517
143.450
116.600
)38.317
)34.467
Australia
Australia
Weld River
Swamp
West Basin
West Lake Muir
Swamp
)34.683
121.383
)2.550
Sulawesi
(Indonesia)
Australia
Wanda
169.849
)20.249
Vanuatu
Umej
2
)158.250
21.517
Hawaii
(USA)
142.583
)5.667
PNG
Uko’a Pond
Core 2
2300
142.367
142.367
)38.317
)38.317
Australia
Australia
Tower Hill Main
Tower Hill
NW Crater
Tugupugua
(Core 1)
145.818
)37.386
Australia
Tom Burns, D
146.317
)39.033
Australia
Tidal River
146.083
)37.750
Thompson River
147.133
100.752
)36.685
)1.051
Australia
Sumatra
(Indonesia)
Australia
Tawonga Bog
Telago Swamp
148.033
)35.668
Australia
Tarcutta Swamp
144.750
145.807
)40.550
)37.444
Australia
Australia
110
100
10
445
2
20
20
1075
2
1250
350
1535
780
65
1177
930
Stockyard Swamp
Storm Creek
145.928
)37.391
2500
Australia
143.417
)5.401
240
Snobs Creek
142.350
)37.167
Australia
< 20
PNG
115.283
)34.300
Australia
Sirunki Wabag
(A16)
Scott River
Swamp
Sheet of Water
%Count
Count
Count
Count
Count
Count
Count
%Count
%Count
%Count
Count
%Count
%Count
Count
Count
Count
%Count
%Count
Count
%Count
Count
Lake
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Lake
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
Swamp
0–10
0–7.76
0–6.8
0 to > 40
0–5.5
0–7
0 to > 13
0–11.5
0–21
0–32
0–4.9
0–11.5
0–7.7
0–9
200–9.8
0 to > 4
0–17.5
0–12
0 to > 30
n/a
0–9.3
3 + top
1
11 (4 not
used)
2
4
6
3 + top
7 (2 not
used)
5
1 + top
2
n/a
1 + top
1
2
1 + top
3 + top
1 + top
9 + top
n/a
2 + top
1
1
2
2
3
3
2
1
1
1
3
1
1
2
2
1
1
1
3
1
2
3C
4D
5D
6C
n/a
n/a
2C
1C
1C
3D
5D
n/a
4C
n/a
n/a
7
1C
5C
3C
n/a
7
n/a
n/a
n/a
1
n/a
n/a
2
n/a
1
1
n/a
n/a
n/a
n/a
n/a
n/a
1
n/a
3
n/a
n/a
n/a
n/a
n/a
7
n/a
n/a
7
n/a
2C
7
n/a
n/a
n/a
n/a
n/a
n/a
7
n/a
2C
n/a
n/a
TSFO;
TDFO;
STEP
TDFO;
TDFO;
TDFO
WTRF;
TDFO
DSFW;
DSFW
XERO
TDFO
COCO;
WTRF
DSFW
DSFW
WTRF;
WSFW;
WSFW
DSFW
DSFW;
DSFW
WTRF
TDFO;
TDFO
CTRF
DSFW
COCO
WSFW;
TDFO;
CTRF
COCO
DSFW;
DSFW
DSFW
TDFO
CTRF;
WTRF
XERO
TUND
DSFW
TRFO;
CTRF;
CTRF
SEAPD
INDOPAC
INDOPAC
INDOPAC
SEAPD
SEAPD
SEAPD
INDOPAC
SEAPD
SEAPD
INDOPAC
INDOPAC
SEAPD
SEAPD
INDOPAC
SEAPD
Gell et al. (1994)
Churchill (1968)
Churchill (1968)
Hope (2001)
Hope & Spriggs (1982),
Hope et al. (1999)
Haberle (1998),
Haberle et al. (1999),
Haberle et al. (2001)
Ward (unpubl. data),
Hope et al. (1999)
McKenzie (1997),
McKenzie & Busby (1992)
D’Costa et al. (1989)
D’Costa et al. (1989)
Strickland (unpubl. data),
Kershaw (1998)
Hope (1974)
Kershaw & Green (1983)
Newsome (1985)
McKenzie (1989),
McKenzie & Busby (1992)
Hope (1999)
McKenzie (1997),
McKenzie & Busby (1992)
Williams (unpubl. data)
McKenzie (unpubl. data),
D’Costa & Kershaw (1997)
Walker & Flenley (1979),
Flenley (1984)
Churchill (1968)
Palaeovegetation patterns for Australia and Southeast Asia
1411
1412
INDOPAC Harle et al. (1993)
Dodson (1977)
DSFW;
DSFW
DSFW;
DSFW
n/a
n/a
2C
6C
2
n/a
650
Yarlington Tier
Australia )42.883 147.300
Count
Swamp 0–9.1
6 (1 not n/a
used)
2 + top 2
Swamp 8.7–< 85
25
Count
of vegetation types sampled, on earlier continental-scale
compilations of pollen data from this region. Prior to the
present compilation, the most extensive compilation of fossil
pollen data for the region included data from only 77 sites, of
which only five included the LGM (Harrison & Dodson, 1993).
The regional compilations covering Southeast Australia
(Kershaw, 1998) and the tropical western Pacific (Hope et al.,
1999) also contain a more limited selection of sites than the
SEAPAC data sets.
Biomization
Wyrie Swamp W3 Australia )37.650 140.300
18 ka
No of 6 ka
No of 18 ka
duplicates DC 6 ka duplicates
DC 18 ka 6 ka
Site name
Table 2 continued
Country
Latitude Longitude Altitude Sample Site
()
()
(m)
type
type
Age range No 14C
Dates
(kyr)
Reconstructed
biomes
Data base
References
E. J. Pickett et al.
Conceptual basis
The biomization procedure (Prentice et al., 1996) is based on
(1) the allocation of pollen taxa to PFTs, (2) the definition of
biomes in terms of their constituent PFTs, (3) the calculation
of the affinity between a pollen spectrum and every biome and
(4) the allocation of the pollen spectrum to the biome to which
it has the highest affinity. The affinity score is calculated taking
into account both the presence of taxa characteristic of specific
PFTs and the relative abundance of these taxa. In cases where
the pollen spectrum has equal affinity with more than one
biome, a tie-breaking rule is applied to determine the biome
allocation. This tie-breaking rule gives preference to biomes
whose constituent PFTs form a subset of the suite of PFTs
characteristic of another biome.
The large-scale geographical distribution of PFTs is determined by climatic controls on plant physiology (Woodward,
1987; Prentice et al., 1992a; Haxeltine & Prentice, 1996;
Harrison et al., in press). In the SEAPAC region, the most
important climate gradients are winter temperature and
moisture availability. The length and warmth of the growing
is a secondary constraint which in this region is important only
at the highest elevations. The division of vegetation into
biomes (how many, and the precise location of boundaries) is
to a greater or lesser extent arbitrary. The biomization
procedure requires that this division is made on taxonomic
grounds, such that each biome is defined by a unique set of
PFTs. However, the PFT differentiating one biome from
another is not required to be the dominant or most abundant
life-form. Furthermore, it is rare for a PFT to occur in only one
biome, and individual PFTs generally occur in a range of
biomes. Given that the distribution of PFTs is determined by
climate, the conceptualization of the relationship among
biomes in bioclimatic space is a fundamental step in the
analysis (Fig. 1). In practice, the biomization procedure is a
test of how well this conceptual scheme works.
Application of the biomization procedure in the SEAPAC
region
Pollen taxa were assigned to PFTs (Tables 3 and 4) on the basis
of our knowledge of the ecology and biology of individual
plants, and on descriptions of the flora and vegetation given in
Jessop (1981), Petheram & Kok (1983), Brock (1993), Ashton
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Figure 1 Conceptual model of SEAPAC biome distribution in climate space.
& Attiwill (1994), Barlow (1994), Busby & Brown (1994), Gill
(1994), Gillison (1994), Johnson & Burrows (1994), Leigh
(1994), Mott & Groves (1994), Parsons (1994), Webb & Tracey
(1994), Williams & Costin (1994), Bowman & Harris (1995),
Enright (1995), Gibson et al. (1995), Jaffré (1995) and
Mueller-Dombois & Fosberg (1998). Individual taxa could be
assigned to one or several PFTs.
The SEAPAC biomization recognizes 25 PFTs (Table 3). We
considered the suite of PFTs defined in biomizations of other
regions of the world (Prentice et al., 1996; Jolly et al., 1998b;
Tarasov et al., 1998; Yu et al., 1998; Edwards et al., 2000;
Elenga et al., 2000; Takahara et al., 2000; Thompson &
Anderson, 2000; Williams et al., 2000; Yu et al., 2000; Bigelow
et al., 2003), adopting those PFTs that appeared to be
appropriate for the SEAPAC region (e.g. tropical evergreen
malacophyll broad-leaved tree), subdividing one PFT (warmtemperate evergreen broad-leaved tree), and creating four new
PFTs (evergreen sclerophyll broad-leaved tree, cool-temperate
sclerophyll broad-leaved low or high shrub, temperate malacophyll broad-leaved low or high shrub, tropical evergreen
broad-leaved low or high shrub). We have adopted a PFTnaming convention (based on life-form, leaf-form, phenology
and bioclimatic tolerance) proposed as part of a global PFT
scheme that is currently under development (Harrison et al., in
press) and was first explicitly adopted in the context of
biomization by Bigelow et al. (2003).
Warm-temperate broad-leaved evergreen trees are an important component of the SEAPAC flora. They occur where winter
temperatures are relatively warm [mean temperature of the
coldest month (MTCO) > 5 C] but still too cool for tropical
trees. Some species are characteristic of the temperate rain
forests, growing in areas with mean annual rainfall as much as
3000 mm year)1, while others are characteristic of drier environments where mean annual rainfall is < 800 mm year)1. We
therefore subdivided the warm-temperate broad-leaved evergreen tree category into three PFTs: drought-tolerant warmtemperate evergreen broad-leaved tree (dt-w-te.e.b.t), ubiquitous warm-temperate evergreen broad-leaved tree (w-te.e.b.t)
and warm-temperate evergreen broad-leaved rain forest tree (wte.e.b.raint) (Table 3). Typical taxa are Arytera in w-te.e.b.t and
Leea, Nothofagus brassii, and Poikilogyne in w-te.e.b.raint, while
dt-w-te.e.b.t is characterized by sclerophyll evergreen taxa.
Evergreen sclerophyll broad-leaved trees (e.sb.t) are highly
characteristic of the Gondwanan flora (Barlow, 1994). They are
defined as tall-medium (35–60 m) single stem, hard-leaved,
evergreen trees that are fire tolerant and well adapted to arid
environments. Typical examples include trees in the genera
Acacia, Eucalyptus and Melaleuca. Evergreen sclerophyll broadleaved trees do not occur in regions where rainfall is
< 500 mm year)1 and are characteristic of areas where mean
annual precipitation is in the range of 800–1200 mm year)1.
Cool-temperate sclerophyll broad-leaved low or high shrub
(c-te.sb.lhs) are hard-leaved plants that grow in regions where
MTCO < 5 C. They are a component of cool-temperate
forests. Typical genera are Agastachys, Cenarrhenes, Poranthera
and Securingea.
Temperate malacophyll broad-leaved low or high shrubs
(te.mb.lhs) are erect, low to medium (1–2 m), soft-leaved
plants which are generally evergreen but may be semideciduous in the tropics. They have a wide temperature range
but are drought intolerant. Characteristic genera include
Buddleia, Crotolaria, Entada and Muehlenbeckia.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1413
E. J. Pickett et al.
Table 3 Definitions and characteristics of plant functional types used in the SEAPAC biomization
PFT Code
PFT name
Characteristic taxa
Description and characteristic adaptations
tr.dd.mb.t
Tropical drought-deciduous
malacophyll broad-leaved tree
Tropical evergreen malacophyll
broad-leaved tree
Cool-temperate evergreen
broad-leaved tree
Amoora, Argyrodendron,
Delarbrea
Darlingia, Kissodendron
Frost intolerant; withstands a long dry
season by shedding leaves
Frost intolerant
Nothofagus moorei,
Placospermum coriaccum
Warm-temperate evergreen
broad-leaved rain forest tree
Ubiquitous warm-temperate
evergreen broad-leaved tree
Drought-tolerant warm-temperate
evergreen broad-leaved tree
Evergreen sclerophyll
broad-leaved tree
Leea, Nothofagus brassii,
Poikilogyne
Arytera
Temperate cold-deciduous
broad-leaved tree
Cool-temperate evergreen
needle-leaved tree
Nothofagus gunii
Frost tolerant; evergreen, medium–tall
(30–60 m) trees; occur in subalpine and
upper cool-temperate forests
Medium to very tall trees (30 to > 60 m);
confined to areas c. 3000 mm year)1 rainfall
Medium to very tall trees (30 to > 60 m);
rainfall range 800 to > 3000 mm year)1
Medium to very tall trees (30 to > 60 m);
mean annual rainfall < 800 mm year)1
Evergreen hard-leaved; tall-medium (35–60 m)
single stem trees; fire tolerant; MTCO
< 12.0 C and rainfall range of 800–1200 mm year)1
Winter deciduous; requires warm growing-season
Lagarostrobus, Microstrobus,
Phyllocladus
i-te.e.n.t
Intermediate-temperate evergreen
needle-leaved tree
Agathis, Araucaria, Callitris,
Pinus, Podocarpus
tr.e.b.lhs
Tropical evergreen broad-leaved low
or high shrub
Temperate malacophyll broad-leaved
low or high shrub
Cassia, Colubrina, Gouania,
Lonicera
Buddleia, Crotolaria, Entada,
Muehlenbeckia
Cool-temperate sclerophyll
broad-leaved low or high shrub
Warm-temperate sclerophyll
broad-leaved low or high shrub
Agastachys, Cenarrhenes,
Poranthera, Securingea
Hypocalymma
ar.ds
Arctic dwarf shrub
Vaccinium
h
Heath
tr-di.fb
te-di.fb
eu-dt.fb
ar.fb
Tropical drought-intolerant forb
Temperate drought-intolerant forb
Eurythermic drought-tolerant forb
Arctic forb
Carronia, Cudrani
Apiaceae
Asteraceae, Chenopodiaceae
Astelia, Asteraceae
s
g
man
tf
eu.f
Sedge graminoid
Grass graminoid
Mangrove
Tree fern
Eurythermic fern or fern ally
Carex, Machaerina, Restionaceae
Poaceae
Avicennia, Rhizophora
Cyathea, Dicksonia
Blechnum, Drymaria, Pteridium
tr.e.mb.t
c-te.e.b.t
w-te.e.b.raint
w-te.e.b.t
dt-w-te.e.b.t
e.sb.t
te.cd.mb.t
c-te.e.n.t
te.mb.lhs
c-te.sb.lhs
w-te.sb-lhs
1414
Eucalyptus calophylla
Acacia, Eucalyptus, Melaleuca
Evergreen small-medium (< 10–30 m) trees;
occur in subalpine and upper cool-temperate
(> 600 m) forest; warm temperatures required
for budburst; frost tolerant
Small-tall (< 10–60 m) evergreen trees;
warm-temperate to temperate regions,
variable rainfall (< 500 to > 4000 mm year)1);
warm temperatures required for seed release
Soft or hard-leaved, erect, low to medium
(1–2 m) shrubs; frost sensitive, MTCO > 5 C
Erect, low-medium (1–2 m) shrubs, usually
evergreen, or semi-deciduous soft-leaved
varieties; wide temperature range,
drought tolerant
Usually evergreen hard-leaved varieties;
frost tolerant
Usually evergreen hard-leaved varieties;
MTCO > 5 C; occur in tropical rain
forest and seasonal forests or subtropical/
montane environments
Prostrate or small shrub; frost tolerant,
occurs in areas where rainfall 800–2500 mm
year)1 and MTCO c. )6.0 C
Woody shrub with narrow microsclerophyllous (hard, leathery) leaves,
usually evergreen
Frost-intolerant forb
Frost-tolerant forb
Arid or seasonally dry environments
Cold and frost tolerant, usually evergreen
and restricted to montane areas
Grow in locally wet environments
Not used in biomization
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Table 4 Assignments of pollen taxa to the plant functional types used in the SEAPAC biomization
Codes
Plant functional types
Pollen taxa
tr.dd.mb.t
Tropical drought-deciduous
malacophyll broad-leaved tree
tr.e.mb.t
Tropical evergreen malacophyll
broad-leaved tree
c-te.e.b.t
Cool-temperate evergreen
broad-leaved tree
Acalypha, Alangium, Alangium villosum, Alanguma rotundifolium, Albizzia,
Aleurites, Aleurites moluccana, Algaia, Altingia, Altingia excelsa, Amoora,
Archontophoenix, Arenga, Argyrodendron, Argyrodendron trifoliatum,
Balanops australiana, Barringtonia, Berrya, Blepharocarya involucrigera,
Bombax, Castonospora, Celtis, Cissus, Cocos nucifera, Combretaceae/
Melastomataceae, Delarbrea, Diospyros, Elaeodendron/Ophiorrhiza,
Emmenosperma, Erythrina sandwicensis, Eugenia/Tristania, Evodia,
Evodia brownwickii, Evodia xantholoides, Fagraea, Heritiera, Hypsophila,
Ilex, Ilex cymosa, Irvingbaileya, Lagerstroemia, Laportea, Longetia,
Macaranga, Macaranga tanarius, Macaranga/Mallotus, Maesa, Mallotus,
Mallotus paniculatus, Melastoma, Melia, Meliaceae, Meliaceae/Sapotaceae,
Meliosma, Melochia, Memecylon, Metrosideros, Mimosaceae, Mucuna gigantea,
Myricaceae, Neonauclea, Oleaceae, Olea paniculata, Palmae, Papilionaceae,
Paratrophis, Pipturus, Pometia, Sambucus, Sloanea, Sterculiaceae, Symplocos,
Terminalia, Trema, Viburnum
Acacia, Acacia kao, Acacia aulacocarpa, Acacia/Albizzia, Acalypha, Acanthaceae,
Acmena, Acmena/Eugenia/Syzygium, Acronychia, Adinandra, Albizzia,
Alphitonia, Anacardiaceae, Anisoptera, Antidesma, Antirhea, Apocynaceae,
Apodytes, Araliaceae, Archidendron, Archontophoenix, Arecaceae, Arenga,
Atherospermaceae, Baccaurea, Balanops, Berrya, Bischofia, Calamus, Calophyllum,
Canthium, Cardwellia, Carnarvonia, Casuarina–Gymnostoma, Celtis, Cheirodendron,
Cissus, Claoxylon, Claoxylon sandwicensis, Cleistocalyx, Clusiaceae, Codia,
Collospermum, Combretaceae/Melastomataceae, Cordyline, Cunoniaceae,
Cunoniaceae/Elaeocarpaceae, Cupaniopsis, Darlingia, Dillenia, Dipterocarpaceae,
Dipterocarpus, Distylium stellare, Doryphora, Drypetes, Dysoxylum, Ebenaceae,
Elaeocarpaceae, Elaeocarpus, Embelia, Endospermum, Engelhardtia,
Eugenia/Syzygium, Eugenia, Euodia, Euphorbiaceae, Ficus, Fagraea, Flindersia,
Flindersia pimenteliana, Freycinetia, Freycinetia arborea, Ganophyllum,
Gardeni, Gendarussa, Glochidion, Guioa, Gymnostoma, Halfordia scleroxyla,
Heritiera, Hibbertia, Hibiscus, Hymenanthera, Homalanthus, Homalium,
Ilex, Irvingbaileya, Juglandaceae, Kissodendron, Lagerstroemia, Lauraceae,
Barringtonia, Macaranga, Macaranga tanarius, Macaranga/Mallotus,
Macropanax, Maesa, Mallotus, Mallotus paniculatus, Melastoma, Melia,
Meliaceae, Melicope, Melicope barbigera, Melicope broadbentiana,
Melicope clusiifolia, Memecylon, Meliaceae/Sapotaceae, Mimosaceae,
Moraceae, Metroxylon, Mucuna gigantea, Musgravea, Myricaceae,
Myristica, Myrtaceae, Notelaea, Oleaceae, Palmae, Palaquium, Pandanus,
Pandanus ansariensis, Pandanus brosimos, Pandanus tectorius, Papilionaceae,
Parasponia, Pisonia, Pittosporum, Pittosporaceae, Planchonella, Pleomele,
Pleurandra, Polyscias, Pometia, Pouteria, Pouteria sandwicensis, Pritchardia,
Proteaceae, Prunus, Psidium, Psychotria, Pygeum turnerianum, Randia,
Rapanea, Rhamnaceae, Sapindaceae, Sapotaceae, Saurauia, Schefflera,
Scolopia, Sloanea, Sphenostemon, Stenocarpus, Sterculiaceae, Strobilanthes,
Symingtonia, Syncarpia, Synoum, Symplocos, Symplocos sessilifolia,
Tapeinospermum, Tapinosperma, Terminalia, Tetraplasandra, Theaceae,
Tieghemopanax, Trema, Tristania, Urtica, Urticaceae, Urticaceae/Moraceae,
Viburnum, Wickstromia, Wilkaea, Zanthoxylum
Acacia, Acacia/Albizzia, Anodopetalu, Araliaceae, Arenga, Aristotelia, Ascarina,
Atherosperma, Daviesia, Dodonaea, Drimys, Drimys/Bubbia/Tasmannia,
Elaeocarpaceae, Elaeocarpus, Eucalyptus, Eucryphia, Hedycarya, Homalanthus,
Monimiaceae, Metrosideros, Myrtaceae, Decaspermum, Xanthomyrtus,
Nothofagus, Notelaea, Nothofagus moorei, Oxylobium, Papilionaceae,
Pandanus, Pomaderris, Pittosporum, Pittosporaceae, Placospermum
coriaccum, Polyosma, Polyscias, Quintinia, Rapanea, Sapindaceae,
Saurauia, Schefflera, Sloanea, Tasmannia, Timonius
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1415
E. J. Pickett et al.
Table 4 continued
Codes
Plant functional types
Pollen taxa
w-te.e.b.raint
Warm-temperate evergreen
broad-leaved rain forest tree
w-te.e.b.t
Ubiquitous warm-temperate
evergreen broad-leaved tree
dt-w-te.e.b.t
Drought-tolerant
warm-temperate evergreen
broad-leaved tree
e.sb.t
Evergreen sclerophyll
broad-leaved tree
te.cd.mb.t
Temperate cold-deciduous
broad-leaved tree
Cool-temperate evergreen
needle-leaved tree
Acmena, Acmena/Eugenia/Syzygium, Albizzia, Alphitonia, Apocynaceae, Arecaceae,
Castanopsis, Castanopsis/Lithocarpus, Casuarina–Gymnostoma, Ceberiopsis,
Cheirodendron, Cissus, Claoxylon, Claoxylon sandwicensis, Cleistocalyx, Cocos nucifera,
Cupaniopsis, Dillenia, Diospyros, Dipterocarpaceae, Dipterocarpus, Distylium stellare,
Doryphora, Drimys, Drimys/Bubbia/Tasmannia, Dysoxylum, Embelia, Eucalyptus
intermedia, Eugenia/Syzygium, Eugenia, Eugenia/Tristania, Euodia, Evodiella,
Freycinetia, Freycinetia arborea, Ganophyllum, Glochidion, Guioa, Gymnostoma,
Hedycarya, Helicia, Hibiscus, Ilex, Ilex anomala, Ixora, Juglandaceae, Lauraceae,
Leea, Lithocarpus, Lithocarpus/Castanopsis, Macaranga, Macaranga/Mallotus,
Mallotus, Mallotus paniculatus, Melicope broadbentiana, Melicope, Melicope barbigera,
Melicope clusiifolia, Monimiaceae, Moraceae, Myoporaceae, Myricaceae, Nothofagus
brassii, Olea paniculata, Palmae, Pandanus, Petalostigma, Pipturus, Polyscias,
Poikilogyne, Pometia, Portulaceae, Pouteria, Pouteria sandwicensis, Pritchardia,
Prunus, Psidium, Quintinia, Rapanea, Rubiaceae, Sapotaceae, Saurauia, Schefflera,
Sphenostemon, Styrax, Synoum, Symplocos, Symplocos sessilifolia, Terminalia,
Tetraplasandra, Theaceae, Timonius, Tristania, Xanthomyrtus, Zanthoxylum
Acacia, Acanthaceae, Acronychia, Acalypha, Angophora/Bloodwood, Araliaceae,
Arecaceae, Arytera, Ascarina, Castanopsis, Castanopsis/Lithocarpus, Ceberiopsis,
Drimys, Elaeocarpaceae, Elaeocarpus, Engelhardtia, Epacris, Epacridaceae,
Epacrid/Ericaceae, Eucalyptus, Eucalyptus diveriscolor, Euphorbiaceae, Flindersia,
Gendarussa, Hymenanthera, Homalanthus, Homolanthus, Lithocarpus, Lithocarpus/
Castanopsis, Macaranga, Macaranga/Mallotus, Macropanax, Mallotus, Metrosideros,
Myrtaceae, Neonauclea, Notelaea, Papilionaceae, Pomaderris, Pittosporum,
Pittosporaceae, Psychotria, Rhamnaceae, Sapindaceae, Sterculiaceae, Strobilanthes,
Syncarpia, Tasmannia, Trema, Urticaceae, Urticaceae/Moraceae, Wickstromia
Angophora, Casuarina, Casuarinaceae, Eucalyptus bloodwood type,
Eucalyptus calophylla, Eucalyptus cornuta, Eucalyptus diveriscolor, Eucalyptus eugenie,
Eucalyptus ficifolia, Eucalyptus gomphocephela, Eucalyptus guilfoylei, Eucalyptus
jacksoni, Eucalyptus loxophleba, Eucalyptus marginata, Eucalyptus megacarpa,
Eucalyptus reantheroid, Eucalyptus wandoo, Grevillea, Gyrostemonaceae,
Leucopogon, Malvaceae, Melaleuca, Proteaceae, Santalaceae
Acacia, Agonis, Angophora, Angophora/Bloodwood, Banksia, Bursaria, Bursaria
spinosa, Calothanus, Casuarina, Casuarinaceae, Corymbia calophylla, Daviesia,
Dodonaea, Dodonaea viscosa, Eucalyptus, Eucalyptus accedens, Eucalyptus bloodwood
type, Eucalyptus cornuta, Eucalyptus ficifolia, Eucalyptus gomphocephela, Eucalyptus
loxophleba, Eucalyptus marginata, Eucalyptus megacarpa, Eucalyptus reantheroid,
Eucalyptus wandoo, Exocarpos, Flindersia, Grevillea, Grevillea glauca, Gyrostemonaceae,
Loranthaceae, Melaleuca, Myrtaceae, Nuytsia floribunda, Oxylobium, Papilionaceae,
Persoonia/Lomatia, Persoonia, Persoonia/Grevillea, Proteaceae, Santalaceae, Santalum,
Syncarpia, Xanthorrhoea
Alnus, Myricaceae, Nothofagus gunnii, Sesbania
c-te.e.n.t
i-te.e.n.t
tr.e.b.lhs
1416
Intermediate-temperate
evergreen needle-leaved
tree
Tropical evergreen
broad-leaved
low or high shrub
Athrotaxis/Diselma, Athrotaxis, Dacrydium, Dacrycarpus, Dacrycarpus imbricatus,
Gymnospermae, Lagarostrobus, Microcachrys, Microstrobos, Papuacedrus,
Phyllocladus, Podocarpus, Podocarpaceae, Podocarpus lawrencii
Agathis, Araucaria, Araucariaceae, Callitris, Callitris/Cupressaceae, Dacrydium,
Dacrycarpus, Dacrycarpus imbricatus, Gymnospermae, Neocallitropsis, Picea,
Pinus, Pinaceae, Podocarpus, Podocarpaceae, Podocarpus lawrencii
Aboutilon, Adinandra, Alyxia, Araliaceae, Ardisia, Borreria, Brownlowia, Caesalpiniaceae,
Canthium, Cassia, Celastris, Cheirodendron, Colubrina, Cordyline, Cunoniaceae,
Cunoniaceae/Elaeocarpaceae, Cyrtandra, Daphniphyllum, Dolicholobium, Dodonaea,
Garcinia, Gardenia, Goodeniaceae, Gouania, Hedyotis, Hedyotis vestitas, Helisia javanica,
Hibiscus, Hiptage, Ixora, Johnsonia, Kanaloa kahoolowensis, Lamiaceae, Leucaena, Lonicera,
Macropanax, Malvaceae, Mastixia, Medinilla, Melastoma, Melastomataceae, Memecylon,
Menispermaceae, Moraceae, Myrica, Myricaceae, Myrica javanica, Nepenthes, Nesoluma
polynesicum, Nestigis, Nestigis sandwicensis, Nyctaginaceae, Parsonsia, Petalostigma, Piper,
Platydesma, Polyscias, Portulaceae, Rauwolfia, Rhus, Schefflera, Schuurmansia, Sesbania,
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Table 4 continued
Codes
te.mb.lhs
c-te.sb.lhs
Plant functional types
Temperate malacophyll
broad-leaved low or
high shrub
Cool-temperate sclerophyll
broad-leaved
low or high shrub
w-te.sb-lhs
Warm-temperate sclerophyll
broad-leaved
low or high shrub
ar.ds
Arctic dwarf shrub
h
Heath
tr-di.fb
Tropical drought-intolerant
forb
te-di.fb
Temperate drought-intolerant
forb
Pollen taxa
Sphenostemon, Staurophyra elegans, Stenogyne, Stephania, Styphelia, Styphelia
tameiameiae, Symplocos, Symplocos sessilifolia, Tarenna fragrans, Theaceae, Tournefortia,
Uncaria/Wendlandia, Verbenaceae, Viburnum, Vitaceae, Wendlandia
Buddleia, Crotolaria incana, Entada, Gastrolobium, Hibbertia, Lonchocarpus,
Muehlenbeckia, Mussaenda, Pomaderris, Proteaceae, Rhododendron, Rutaceae, Salvia,
Styrax, Tinospora
Acacia, Agastachys, Astroloma/Acrotriche, Astroloma, Baeckea, Banksia, Banksia marginata,
Bauera, Boronia, Bossiaea, Cenarrhenes, Coprosma, Correa, Daviesia, Drimys, Epacris,
Epacridaceae, Epacridaceae/Ericaceae, Epacrid/Styphelia/Leucopogon, Ericaceae, Eriostemon,
Eucryphia, Goodeniaceae, Grevillea/Persoonia, Grevillea, Hakea, Hibbertia, Hymenanthera,
Hovea, Hovea/Bossaia, Kunzea, Lamiaceae, Leucopogon, Leptospermum, Leptospermum/Baeckea,
Lomatia, Monotoca, Myrtaceae, Opercularia, Orites, Oxylobium, Papilionaceae,
Pimelea, Poranthera, Poranthera microphylla, Prostanthera, Rhamnaceae, Richea, Richea
continentis, Rutaceae, Securingea, Sprengelia, Sprengelia incarnta, Stackhousiaceae,
Symplocaceae, Tasmannia, Telopea, Tremandraceae
Acacia, Acalypha, Adenanthos, Adenanthos/Stirlingia, Agonis, Austrobuxus/Dissiliaria,
Banksia, Banksia/Dryandra, Beaufortia, Boronia, Bossiaea, Calothanus, Calytrix, Calytrix/Agonis,
Calytrix/Verticordia, Chloanthaceae, Colobanthus, Conospermum, Conospernum/Persoonia,
Coprosma, Darwinia micropetala, Dasypogonaceae, Daviesia, Dodonaea, Dodonaea viscosa,
Ericaceae, Eriostemon, Gastrolobium, Goodeniaceae, Grevillea/Persoonia, Grevillea, Gymnostoma,
Gyrostemonaceae, Gyrostemon, Hakea, Hibbertia, Hovea, Hovea/Bossaia, Haloragodendron,
Hypocalymma, Isopogon/Petrophile, Johnsonia, Kunzea, Leptospermum laevigatum, Leptospermum
myrsinoides, Loranthaceae, Melaleuca, Myrtaceae, Macrozamia, Oxylobium, Papilionaceae,
Parinarium, Persoonia/Lomatia, Persoonia, Persoonia/Grevillea, Pimelea, Portulaceae, Proteaceae,
Helicia, Rhamnaceae, Rosaceae, Rubiaceae, Rubus, Rutaceae, Rynchosia, Sapindaceae, Sapindus,
Scholitzia, Schuurmansia, Sophora toromiro, Styphelia, Styphelia tameiameiae, Tremandraceae,
Trichospermum, Triumfetta, Verticordia, Xanthorrhoea
Asteraceae, Asteraceae-Cardaeae, Asteraceae-Tubuliflorae, Calomeria, Athrotaxis/Diselma,
Athrotaxis, Baeckea, Bossiaea, Brassicaceae/Lamiaceae, Comesperma, Epacris, Epacridaceae,
Epacridaceae/Ericaceae, Epacrid/Styphelia/Leucopogon, Eriostemon, Exocarpos, Goodeniaceae,
Hymenanthera, Lamiaceae, Leucopogon, Lomatia, Nothofagus gunnii, Orites, Parahebe, Phebalium,
Polygalaceae, Podocarpus lawrencii, Polygalaceae, Proteaceae, Helicia, Prostanthera, Richea
continentis, Richea, Rubiaceae, Rutaceae, Sprengelia, Sprengelia incarnta, Vaccinium
Acacia, Adenanthos, Adenanthos/Stirlingia, Amperea xiphocladus, Angophora, Apiaceae,
Astroloma/Acrotriche, Asteraceae, Asteraceae-Cardaeae, Asteraceae-Tubuliflorae, Astroloma,
Austrobuxus/Dissiliaria, Baeckea, Banksia/Dryandra, Bauera, Brachycome, Calorophus, Calytrix,
Calytrix/Agonis, Calytrix/Verticordia, Casuarina, Casuarinaceae, Correa, Dasypogonaceae, Epacris,
Epacridaceae, Epacridaceae/Ericaceae, Epacris obtusifolia, Epacrid/Styphelia/Leucopogon, Eriostemon,
Euphorbiaceae, Exocarpos, Grevillea, Hakea, Haloragis, Haloragis/Gonocarpus, Hibbertia,
Isopogon, Isopogon/Petrophile, Leucopogon, Leptospermum, Leptospermum/Baeckea, Lomatia,
Lomandra, Melaleuca, Monotoca, Myrtaceae, Opercularia, Orites, Oxylobium, Papilionaceae,
Parinarium, Persoonia/Lomatia, Pimelea, Platysace, Proteaceae, Rhamnaceae, Rutaceae,
Sprengelia, Sprengelia incarnta, Tetratheca, Xanthorrhoea
Angioteridaceae, Apiaceae, Asplenium nidus, Araceae, Boea, Brassia actinophylla, Burmannia,
Calocasia/Alocasia, Campanulaceae, Carronia, Cibotium, Clematis, Colysis, Commersonia,
Commelina, Convolvulaceae, Convolvulus, Cudrania, Deeringia, Dianella, Elattostachys,
Elephantopus, Ficus, Flagellaria, Gesneriaceae, Goodeniaceae, Hodgekinsonia, Hydrocera trifolia,
Hypserpa, Impatiens, Johnsonia, Laurembergia, Liliaceae, Lobelia, Malisia, Morinda, Mussaenda,
Nephrolepis, Ophiorrhiza, Ophioglossum, Plumbago zeylanica, Pothos, Pyrrosia, Rhipogonum,
Salvia, Schizaea, Schizaeaceae, Schizeilema frageosum, Stephania, Syzgium, Tinospora,
Uncaria/Wendlandia, Urticaceae, Urticaceae/Moraceae, Wilkstoemia
Acaena, Amaranthaceae, Apiaceae, Asperula, Asteraceae-Liguliflorae, Australina,
Boraginaceae, Brachycome, Campanulaceae, Chiono, Gentiana, Commersonia, Commelina,
Convolvulaceae, Caryophyllaceae, Caryophyllum, Dianella, Dichosciadium ranunculaceum,
Diplaspis, Drosera, Droseraceae, Drynaria, Emex australis, Epilobium, Euphrasia, Galium,
Gastrolobium, Geraniaceae, Gnaphalium, Gnaphalium/Ewartia, Goodeniaceae, Gunnera,
Haloragis/Gonocarpus, Hovea, Impatiens, Lamiaceae, Laxmannia, Liliaceae, Lobelia, Lomandra,
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1417
E. J. Pickett et al.
Table 4 continued
Codes
Plant functional types
eu-dt.fb
Eurythermic drought-tolerant
forb
ar.fb
Arctic forb
s
g
tf
eu.f
Sedge graminoid
Grass graminoid
Tree fern
Eurythermic fern or fern ally
Pollen taxa
Lomandra longifolia, Lomandra/Xanthorrhoea, Orchidaceae, Oxalis, Pelargonium, Pentapanax,
Plantago, Plantago major, Plantago varia, Platysace, Polygonaceae, Pueraria, Rumex, Scaevola,
Schizeilema frageosum, Sida, Samolus, Stellaria, Stylidium, Stylidiaceae, Tetratheca, Tetragonia,
Thalictrum, Tremandraceae, Trigonotis, Urticaceae, Violaceae, Wahlenbergia
Acacia, Aizoaceae, Asteraceae, Asteraceae-Cardaeae, Asteraceae-Liguliflorae, Asteraceae-Tubuliflorae,
Calomeria, Campanulaceae, Chenopodiaceae, Chenopodiaceae/Amaranthaceae, Dodonaea, Ephedra,
Eremophila, Euphorbiaceae, Goodeniaceae, Grevillea, Gyrostemonaceae, Gyrostemon,
Haemodoraceae, Hakea, Lamiaceae, Malvaceae, Myoporaceae, Nitraria, Papilionaceae, Pimelea,
Proteaceae, Solanaceae, Solanum, Vernonia arboreas, Wahlenbergia
Aciphylla, Acaena, Apiaceae, Asteraceae, Asteraceae-Cardaeae, Asteraceae-Liguliflorae,
Asteraceae-Tubuliflorae, Calomeria, Astelia, Astelia papuana, Australina, Boraginaceae, Brassicaceae,
Brassicaceae/Lamiaceae, Caltha, Calorophus, Centrolepis, Chiono/Gentiana, Chionohebe, Coprosma,
Caryophyllaceae, Caryophyllum, Drapetes, Epacridaceae/Ericaceae, Epilobium, Ericaceae, Euphrasia,
Galium, Gentiana, Gentianaceae, Gnaphalium, Gnaphalium/Ewartia, Goodeniaceae, Gunnera,
Lactuca, Lactuceae, Leptocarpus, Neopaxia, Plantago, Plantago debilis, Plantago major, Plantago
muelleri, Plantago varia, Potentilla, Ranunculus, Ranunculaceae, Restionaceae, Scrophulariaceae,
Stellaria, Trigonotis, Urtica, Urticaceae, Vernonia arboreas, Wahlenbergia
Carex, Eleocharis, Lepironia, Leptocarpus, Machaerina, Restio, Restionaceae, Xyris
Lepidium, Poaceae, Poatia
Cyathea, Cyatheaceae, Dicksonia, Marattia, treeferns
Acrostichum, Adenophorus tamariscinus, Angioteridaceae, Anopteris, Asplenium, Aspidium, Aspidium
exaltum, Aspidium molle, Belvisia, Blechnum, Botrychium, Ceratopteris thalictroides, Culcita, Culcita
dubia, Cyclosorus, Davalliaceae, Davallia, Dennstaedtiaceae, Dennstaedtia, Dicranopteris, Dryneria,
ferns, Gleicheniaceae, Gleichenia, Gleichenia linearis, Gleichenia vulcanica, Grammitidaceae,
Grammitis, Histiopteris, Humata, Huperzia selago, Hymenophyllum, Hypolepis, Lycopodium, Lygodium,
Lycopodium cernuum, Lycopodium clavatum, Lycopodidae, Lycopodium phlegmaria, Marattia,
Microsorium, Nephrolepis, Ophiorrhiza, Parkeriaceae, Pellea, Pteris, Polystichum, Polypodiaceae,
Polypodium, Polypodium pellucidum, Polypodium pallidium, Pteridophyta, Pteridium, Pteridium
aquilinum, Schizaceae, Selaginella, Selaginella plantac, Selaginella semi-caudaraus, Sphenomeris,
Stenochlaena palustris, Todea
Tropical evergreen broad-leaved low or high shrubs
(tr.e.b.lhs) are frost sensitive species confined to regions where
MTCO > 15 C. The PFT includes both soft-leaved and hardleaved evergreen varieties of erect, low to medium (1–2 m)
shrubs. Characteristic genera include Cassia, Colubrina, Gouania and Lonicera.
We initially adopted the conceptual biome classification
shown in Fig. 1. This scheme shows the relationship among
biomes in bioclimate space. As Fig. 1 shows, we defined 11
biomes (Table 5), each characterized in terms of its component
PFTs (Table 6). We adopted a standardized biome terminology,
designed to broadly conform with the classification scheme
used in the equilibrium vegetation model BIOME4 (Kaplan
et al., 2003; Bigelow et al., 2003; Ni et al., in press; Sutra et al.,
in press). However, we recognize four biomes that were not
distinguished in BIOME4: warm-temperate rain forest (WTRF),
CTRF, wet sclerophyll forest (WSFW), and temperate sclerophyll woodland and shrubland (DSFW). These four biomes are
widely recognized by ecologists as distinctive elements of the
modern landscape in the SEAPAC region (Barlow, 1981, 1994;
Hope, 1996; Mueller-Dombois & Fosberg, 1998).
Temperate rain forests were first defined (Schimper, 1903)
and extensively studied in Australia (Beadle, 1981; Webb &
1418
Tracey, 1981; Webb et al., 1984; Adam, 1992). Rain forests are
closed-canopy forests, which nevertheless include a wide range
of life-forms other than trees, and with high species diversity
both in the canopy and the understorey (Adam, 1992; Busby &
Brown, 1994; Webb & Tracey, 1994). Temperate rain forests
are distinguished from tropical rain forests because they have
fewer canopy species (Busby & Brown, 1994). Cool-temperate
rain forests (CTRF) are distinguished from WTRF by the
absence of buttress tree forms and lianas (Busby & Brown,
1994). WTRF and CTRF both occur in montane areas in the
tropics, where they are often referred to as lower and upper
montane forests, respectively (e.g. Flenley, 1996; Hope, 1996).
Warm-temperate rain forests occur in scattered pockets along
the coastal margin of north-eastern Australia, as far south as
c. 36 S, and on mountains in the Pacific islands. The southern
limit is controlled by winter temperature, as these forests do
not occur in areas subject to frost. Warm-temperate rain
forests are characterized by a mixture of tropical and temperate
trees, shrubs and forbs (including such PFTs as tropical
drought-deciduous malacophyll broad-leaved tree, tropical
evergreen broad-leaved low or high shrub, tropical droughtintolerant forb, warm-temperate evergreen broad-leaved rain
forest tree, temperate malacophyll broad-leaved low or high
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Table 5 Definition and description of the biomes used in the SEAPAC biomization
Biome code
Biome name
Definition
Characteristic species or assemblages
Equivalents
TRFO
Tropical
evergreen
broadleaf
forest
Tall (> 45 m), closed-canopy
lowland evergreen forests.
Mesophyll leaf size, MTCO
> 18C, intolerant of frost
Agathis, Alstonia, Araucaria,
Castanopsis, Celtis, Dacrydium,
Eleaocarpus, lianas, e.g. Freycinetia,
epiphytes, e.g. Astelia
TSFO
Tropical semievergreen
broadleaf
forest
Acalypha, Barringtonia, Bombax,
Celtis, Metrosideros
TDFO
Tropical deciduous
broadleaf forest
and woodland
Acacia, Casuarina, Lysiphyllum,
Melaleuca quinquenevua (‘nialouli’),
Terminalia
Gulf country of north-eastern
Australia, New Caledonia
‘nialouli savanna’, Thailand
WTRF
Warm-temperate
rain forest
Araucaria, Fagaceae, Lauraceae,
Eucalyptus
CTRF
Cool-temperate
rain forest
Tall (45 m), semi-evergreen
(inc. raingreen or semideciduous trees)
closed-canopy forests
and monsoon forests.
Seasonally dry up to
5 months, rainfall meets
80–95% demand
Medium-tall (10–30 m)
discontinuous tree canopy,
semi-evergreen woodland with
xeromorphic, fire-tolerant
grasslands. Characteristic of
long dry season
Medium-tall (15–33 m)
evergreen closed forest.
Moisture-demanding,
tolerates no freezing.
Mesophyll leaf size in
tropical lower
montane forests
Medium-tall, closed/open
semi-evergreen forest.
Temperature dependent,
tolerates freezing
Lowland forests of PNG,
Queensland (Australia), New
Caledonia, Thailand, Malaysia,
Phillipines, Fiji, Solomon Islands,
Vanuatu
Seasonal forests of Queensland
(Australia) and Southeast Asia
COCO
Cool evergreen
needleleaf
forest
Subtropical rain forest
(e.g. Queensland and
northern New South Wales),
sub/lower-montane tropical
forests (750–1500 m),
e.g. PNG, Malaya,
New Caledonia
Temperate rain forest of,
e.g. NSW, Victoria, Tasmania.
Upper montane and subalpine
tropical rain forest areas
(1500– to > 3000– m), e.g. PNG,
Queensland, New Caledonia
Southern conifer forests,
e.g. montane and mainland
south-eastern Australia,
New Caledonia
WSFW
Wet sclerophyll
forest
DSFW
Temperate
sclerophyll
woodland and
shrubland
XERO
Xerophytic
woods/scrub
Medium, open forestwoodlands of mixed
conifers with smaller
understorey trees, shrubs,
herbs and grasses
Tall closed canopy
evergreen with dense
understorey. Frost tolerant;
low rainfall seasonality,
moderately drought tolerant
Medium-tall (5–30 m)
mixed-evergreen, open
forest (30–70%) to tall
woodlands with grass/shrub
understorey usually
dominated by Eucalyptus.
Characteristic of low rainfall
areas, drought tolerant
Low-medium, open canopy
evergreen woodland to tall
shrublands (< 2 m)
Agathis, Castanopsis, Dacrydium,
Dicksonia, Ericareaceae, Eucalyptus,
Ilex, Lithocarpus, Metrocidrus,
Nothofagus, Quintiana
Athrotaxis, Dacrydium, Dacrycarpus,
Lagarostrobus franklinii, Microstrobos
fizgeraldii, Phyllocladus, Podocarpus
lawrencei, with Eucalyptus emergents
Eucalyptus
Eucalyptus forests of
south-eastern and
south-western Australia
Agonis, Banksia, Casuarina, Callitris,
Dodonaea, Eucalyptus
Large area of southern Australia,
New Caledonia dry sclerophyll
forest (e.g. Noumea)
Acacia, Actinostrobus, Callitris,
Epacridaceae (e.g. Styphelia),
Myrtaceae (Eucalyptus), Proteaceae
(e.g. Banksia, Isopogon, Petrophile,
Beaupropsis)
New Caledonia maquis,
Australian mallee woodlands,
Australian mulga, kanji in
northern Australia, kwongan
of south-western Australia,
parts of South Australia
and Tasmania
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1419
E. J. Pickett et al.
Table 5 Continued
Biome code
Biome name
Definition
Characteristic species or assemblages
Equivalents
STEP
Temperate or
tropical
grassland and
xerophytic
shrubland
Shrubland or grassland, lacking
trees, and with drought-tolerant
or salt-tolerant herbs and forbs
Amaranthaceae, Atriplex, Chenopodiaceae, Poaceae
TUND
Tundra
Low (< 1 m) to dwarf shrublands
of mixed herbs, forbs, sedges and
grasses. Tolerates drought
and freezing
Carex, Empodisma, Eucalyptus,
Nothofagus gunii, Prostanthera
cuneata, Phebalium, Poa,
Podocarpus lawrencei
Semi-arid shrubland steppe,
shrub and grassland, chenopod
low open scrub, and chenopod
shrublands of South Australia,
western New South Wales,
south-east Western Australia,
Queensland and Northern
Territory (< 300 mm year)1,
< 10 C)
Alpine/subalpine shrublands of
Gippsland Highland and Snowy
Mountains (> 1370 m), Tasmania
(> 915 m), high montane areas
of PNG and Malaysia (> 3500 m)
Table 6 Assignment of plant functional types to the biomes used in the SEAPAC biomization. The order of the biomes reflects the order
used in the tie-break procedure
Codes
Biomes
Plant functional types
TRFO
tr-di.fb, tr.e.b.lhs, te.mb.lhs, i-te.e.n.t, tr.e.mb.t
WTRF
Tropical evergreen
broadleaf forest
Tropical semi-evergreen
broadleaf forest
Tropical deciduous broadleaf
forest and woodland
Warm-temperate rain forest
COCO
CTRF
WSFW
Cool evergreen needleleaf forest
Cool-temperate rain forest
Wet sclerophyll forest
STEP
Temperate or tropical grassland
and xerophytic shrubland
Xerophytic woods/scrub
Temperate sclerophyll woodland
and shrubland
Tundra
TSFO
TDFO
XERO
DSFW
TUND
tr-di.fb, w-te.sb.lhs, tr.e.b.lhs, te.mb.lhs, i-te.e.n.t,
tr.e.mb.t, tr.dd.mb.t
g, tr-di.fb, w-te.sb.lhs, tr.e.b.lhs, e.sb.t, w-te.e.b.t,
w-te.e.b.raint, tr.e.mb.t, tr.dd.mb.t
tr-di.fb, te-di.fb, w-te.sb.lhs, tr.e.b.lhs, te.mb.lhs, i-te.e.n.t,
e.sb.t, w-te.e.b.t, w-te.e.b.raint, tr.dd.mb.t, tf
te-di.fb, c-te.sb.lhs, te.mb.lhs, c-te.e.n.t, c-te.e.b.t
te-di.fb, c-te.sb.lhs, tr.e.b.lhs, te.mb.lhs, 4c-te.e.n.t, te.cd.mb.t, c-te.e.b.t, tf
te-di.fb, w-te.sb.lhs, c-te.sb.lhs, te.mb.lhs, c-te.e.n.t, i-te.e.n.t,
e.sb.t, w-te.e.b.t, tf
g, eu-dt.fb
g, eu-dt.fb, te-di.fb, h, w-te.sb.lhs, i-te.e.n.t
g, te-di.fb, h, w-te.sb.lhs, c-te.sb.lhs, te.mb.lhs, i-te.e.n.t,
e.sb.t, dt-w-te.e.b.t
g, s, ar.fb, ar.ds, te.cd.mb.t
shrub, temperate drought-intolerant forb) and the presence of
tree ferns. Thus, they are distinguished from tropical rain
forests by the presence of temperate elements, and from CTRF
by the presence of tropical elements. Cool-temperate rain
forests occur in isolated pockets along the coastal margin of
south-eastern Australia, but are most extensive in Tasmania.
As they are found in regions subject to frost, CTRF does not
include, e.g. tropical tree forms. Temperate rain forests have
not been recognized in biome analyses of other regions,
although they occur in south-eastern China, southern South
America, New Zealand and the Pacific Northwest region of
North America (Mitchell et al., 1991; Busby & Brown, 1994).
Sclerophyll forest and woodland, which are characterized by
evergreen or semi-evergreen, sclerophyll species, are typical of
large parts of the temperate zone in the SEAPAC region
(Ashton & Attiwill, 1994). In the wetter areas, where mean
1420
annual precipitation is > 1100 mm year)1, they form forests
with tall (30–65 m) trees. Tree height and the density of the
canopy decrease progressively as precipitation declines and the
sclerophyll forests become increasingly open, grading into
open woodland and semi-arid woodland. In areas where the
rainfall is < 500 mm year)1, open woodland is replaced by the
low (4–10 m), multi-stemmed trees of the mallee or the mulga
(Parsons, 1994). Both sclerophyll forest and woodland are
characterized by a diverse understorey. Eucalyptus trees are the
dominant genus throughout the temperate sclerophyll vegetation zone, but the gradual change in structural characteristics
along the rainfall gradient is paralleled by changes in the
taxonomic composition. Thus, Phyllocladus and Atherosperma
are important components of the lower strata of WSFW, while
treeferns (e.g. Cyathea, Dicksonia) are characteristic of the
understorey. The dominant canopy-forming species in the dry
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
sclerophyll forest and woodland is Eucalyptus; species from the
Proteaceae, Myrtaceae and Epacridaceae, however, are present
in the diverse understorey. Wet sclerophyll forest (Beadle,
1981) extensively in wetter areas of south-eastern and the
extreme south-west of Australia, and as a discontinuous belt
between rain forest and open sclerophyll forest in some parts
of north-eastern Australia. Dry sclerophyll woodland covers c.
25% of the Australian continent and is also common on many
of the offshore islands, extending into the southern parts of
PNG and into New Caledonia and Timor (Gillison, 1994).
Tree ferns (tf) are a significant component of several
SEAPAC biomes. In addition to the tropical forests, they are
one of the PFTs that distinguish WTRF, CTRF and WSFW
(Table 6). Tree ferns were recognized as a distinctive PFT in
the biomization of China (Yu et al., 1998, 2000; Ni et al.,
in press). In China they only contributed to the definition of
tropical biomes; in the SEAPAC region, they are characteristic
of high-rainfall areas (> 1100 mm year)1) in both tropical and
temperate zones.
Once the taxa to PFT (Table 4) and PFT to biome (Table 6)
allocations were determined, the biomization procedure was
implemented as described in Prentice et al. (1996). Biomization was carried out for all samples, including multiple samples
from individual sites. Comparison of the results from several
samples at the same site provides a measure of the robustness
of the procedure. The biomization of the modern pollen data
set was compared with descriptions of the observed vegetation
around the collection site (Table 1). Many of these descriptions reflect the local rather than the regional vegetation. While
useful for determining the degree to which the modern pollen
sample might be affected by, e.g. natural or human-induced
disturbance or reflect azonal conditions, such a comparison is
not an entirely adequate measure of the degree to which the
biomization procedure captures regional vegetation patterns.
There is no reliable vegetation map covering the whole of the
SEAPAC region, but it was possible to make use of the natural
vegetation map from the Atlas of Australian Resources
(Carnahan, 1997). This map classifies the vegetation of Australia
prior to European colonization in the eighteenth century, on the
basis of structure and the dominant component of both the
canopy and the understorey, and could be readily reclassified to
match the biomes used in the SEAPAC biomization (Fig. 2,
Figure 2 The vegetation of Australia in the 1780s prior to European colonization. Data are derived from the Australian vegetation map
(Carnahan, 1997) and have been reclassified (Table 7) to conform with the SEAPAC biome scheme.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1421
E. J. Pickett et al.
Table 7 Allocation of the vegetation codes used in the vegetation map of Australia to the biomes used in the SEAPAC biomization. The
order of the biomes reflects the order used in the tie-break procedure
SEAPAC biomes
Biome codes
Australian vegetation map code
Tropical evergreen broadleaf
forest
Tropical semi-evergreen
broadleaf forest
Tropical deciduous
broadleaf forest
and woodland
Warm-temperate rain forest
Cool evergreen needleleaf forest
Cool-temperate rain forest
Wet sclerophyll forest
Temperate or tropical grassland and
xerophytic shrubland
TRFO
xT4NL
TSFO
Not present in Australia
TDFO
eL1wS, eM2G, eM3G, eM3L, eM3S, eM3Z, epM2G, epM2S, epM3L, ewM2G, ewM2S,
mM2G, mM3G, peM2G, weM2G, weM3NL
WTRF
COCO
CTRF
WSFW
STEP
pL4NL, xM4NL
pM3NL
nM4NL
eT3L, eT3M, eT3S
aG2NL, aG3NL, cL1DtH, cL1DtHD, cL1DwSD, cL1ewS, cL1kZ, cL1tH, cL1wS,
cL1yG, crL1kZ, crL1yG, crL2G, crL2Z, daG3NL, dG3NL, ecL1DtHD, ecL1DwSD,
ecL1wS, ecL1yG, eL1aG, eL1dG, eL1DtHD, eL1DyGD, eL1kF, eL1kZ, eL1NL, eL1tG,
eL1tH, eL1wS, eL1xS, eL1xZ, eL1yG, eL1yG, epL1wS, eS1DtHD, eS1tH, eS1xZ, ewL1kF,
ewL1xZ, ewL1yG, ewS1DtHD, ewS1tH, gG3NL, gG4NL, gyG4NL, hS1tH, kF1DNL,
kF1NL, kZ1DkFD, kZ1DxFD, kZ1DyGD, kZ1NL, kZ1yG, kZ2DGD, kZ2DNL, kZ2F,
kZ2FS, kZ2G, kZ2NL, mcL1xZ, meL1yG, mL1tH, mL1yG, oL1DtHD, oL1DtHD,
pL1wS, pL1xS, qcL1kZ, qL1kZ, sG3NL, tH2NL, wcL1kZ, weL1kF, weS1tH, whS1DyGD,
whS1tH, wL1aG, wL1dG, wL1DyGD, wL1kF, wL1kZ, wL1NL, wL1yG, wS1DtH,
wS1DtHD, wS1DyGD, wS1kZ, wS1tH, wS1xZ, wS1yG, wZ1DtHD, wZ1NL, wZ1tH,
wZ1yG, xF1NL, xH2NL, xL1kZ, xL1tH, xL1xS, xL1xZ, xL1yG, xS1aG, xS1DkZD,
xS1DtHD, xS1DyGD, xS1kZ, xS1tH, xS1yG, xZ1aG, xZ1DtHD, xZ1yG, xZ2NL,
yG1DNL, yG1NL, yG2DNL, yG2NL, yG3NL, yG4NL, oL1tH
bcZ3NL, bL2Z, cL2G, cL2Z, cS3Z, cwS3Z, ebL2Z, ecL2S, ecL2Z, ecS3Z, eL2H, eL2S,
eL2Z, emL2G, epL2G, epL2S, epL2Z, eqS2Z, eS2DZD, eS2G, eS2H, eS2Z, eS3G, eS3Z,
ewL2G, meL2G, meS2Z, mL2G, mL2H, peL2S, peL2Z, wcL2DZD, wcL2Z, wcS3Z, weL2G,
weL2S, weL2Z, weS2H, wL2G, wL2H, wL2NL, wL2S, wL2Z, wpL2G, wS2DGD, wS2FS,
wS2G, wS2H, wS2Z, wS3H, wS3Z, xL2G, xL2S, xS2Z, xS3Z, xZ3G, xZ3NL, eL229
cM3L, cwM3L, ecM2L, eL3G, eL3Z, eM1aG, eM1tH, eM1wL, eM1wpL, eM1xL,
eM1yG, eM2G, eM2H, eM2L, eM2S, eM2Z, eM3G, eM3L, eM3S, eM3Z, epM2G,
epM2L, epM2S, epM3L, ewM2G, ewM2L, ewM2S, mL3G, mM2G, mM3G, peM2G,
peM2S, pM2G, pM3NL, wcM3L, weM2G, weM3NL, wL3NL, wM3L, wM3NL, xL3S,
xL3Z, xL4NL
xZ2G
Xerophytic woods/scrub
XERO
Temperate sclerophyll
woodland and shrubland
DSFW
Tundra
TUND
Table 7) for validation purposes. The biomization procedure
was applied without modification to the fossil pollen samples.
To determine whether the differences in the distribution of
modern and fossil biomes apparent on the resultant maps were
robust, we analysed biome changes at those sites with a record
for the present and either 6 or 18 ka (Table 8).
RESULTS
Reconstructed modern biomes vs. observed modern
biomes
The pollen-derived biome map (Fig. 3) shows patterns which
broadly conform with the observed patterns in vegetation
distribution (Fig. 2). Thus, the biomization procedure correctly reproduces the transition from cool-temperate forests
in the extreme south (cool evergreen needleleaf forest,
1422
CTRF), through into WSFW and into DSFW in dry areas of
south-eastern Australia, and the occurrence of isolated patches
of WTRF in the coastal regions of eastern Australia. The
gradient from DSFW into more xerophytic vegetation in
south-western Australia is also correctly reconstructed. In
north-eastern Australia, the pollen-based reconstruction shows
the patchy distribution of sclerophyll forest, WTRF and
tropical semi-evergreen broadleaf forest characteristic of the
observed vegetation patterns. The extent of tropical forests
(tropical evergreen broadleaf forest, tropical semi-evergreen
broadleaf forest, tropical deciduous broadleaf forest and
woodland) is less than might be expected from comparison
with the observed biome map. However, this appears to reflect
the large proportion of the sites (15%) which, according to the
vegetation descriptions provided by the original authors, come
from disturbed environments. There are very few sites from the
arid interior (i.e. the region characterized by xerophytic
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Table 8 Assessment of the robustness of
biome allocations at sites with multiple
samples for 0, 6 or 18 ka. Biome codes are
given in Table 5
Site
0 ka
Aluaipugua (core 2)
Anouwe 2
Anumon Swamp
Badmenangidongwa
Bandung DPDR-I
Bandung DPDR-II
Barker Swamp
Bega Swamp
Bendenumbun
Bibra Lake (BL2)
Big Dam Marsh
Black Swamp
Blue Lake (Snowy Mts)
Blue Tea Tree Swamp
Bobundarra Swamp
Boggy Lake
Boggy Swamp
Bonatoa Bog
Brass Tarn
Bremer Bay Swamp
Bromfield Swamp
Brownes Lake
Burraga Swamp centre
Butchers Swamp
Camerons Lagoon
Cave Bay Cave
Club Lake core 1
Club Lake core 2
Cotter Source Bog
Cotter’s Lake
Danau Di Atas
Darby Beach
Digger’s Creek Bog
Dragon Tree Soak
Evoran Pond
Fissoa River
Five Mile Beach
Flinders Bay Swamp
Fool Swamp
Giwimawi
Guruanglakugl
Haeapugua, Core 6
Hogayaku Lake
Horse Swamp
Ijomba
Jacksons Bog A
Kawainui Marsh Core B
Kealia Pond
Killer Bog
Komanimambuno
Kosipe
Koumac Northeast
Kurnell Fen Core 1
Lac Suprin
Lake Euramoo, A
Lake George
Lake Habbema
Lake Hordorli
CTRF 2 TDFO 1
WTRF 2
WTRF 2 TDFO 1
WTRF 2
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
XERO 3
DSFW 3
WTRF 1 TDFO 1
DSFW 3
DSFW 2
DSFW 3
STEP 2 TRFO 1
DSFW 2 WTRF 1
DSFW 3
TRFO 2 TDFO 1
WTRF 2 TDFO 1
TDFO 2 DSFW 1
6 ka
18 ka
CTRF 3
TDFO 1 WTRF 1
XERO 2 DSFW 1
DSFW 3
TDFO 2
TDFO 2 TUND 1
DSFW 3
DSFW 3
TUND 1 DSFW 1
DSFW 2 TDFO 1
DSFW 3
DSFW 3
WTRF 4 TDFO 1
WTRF 3
DSFW 2 XERO 1
TDFO 6
DSFW 3
TUND 2 DSFW 1
XERO 1 DSFW 1
DSFW 2
DSFW 3
DSFW 2
XERO 3
XERO 2 DSFW 1
DSFW 3
XERO 4 STEP 1
WTRF 2 TUND 1
WTRF 1 TSFO 1 TRFO 1
TDFO 2
DSFW 2
DSFW 2
STEP 2
WTRF 2
XERO 3
DSFW 3
TDFO 3
WTRF 2
TDFO 2
TDFO 3
XERO 2
TUND 2 CTRF 1
DSFW 3
TDFO 2 XERO 1
XERO 2 WTRF 1
DSFW 3
WTRF 3
TDFO 2
TDFO 2
DSFW 3
WTRF 1 TRFO 1
TDFO 3
WTRF 2
WTRF 3
COCO 4
TDFO 2 DSFW 1
TDFO 2
TUND 1 DSFW 1
CTRF 3
WTRF 1 TDFO 1
CTRF 2 WTRF 1
DSFW 3
WTRF 1 TSFO 1
WTRF 3
TDFO 1 DSFW 1
TDFO 3
WTRF 2
Not used
CTRF 1 WSFW 1
1423
E. J. Pickett et al.
Site
0 ka
6 ka
Lake Leake (core LE)
Lake Majo
Lake Tagamaucia
Lanyon House
Lembang Swamp/Panjairan
Liliha Core 4
Loch McNess Swamp
Lowelak Swamp
Lynch’s Crater A
DSFW 4
TDFO 2
WTRF 2 TDFO 1
XERO 7 DSFW 1
XERO 3
Rano Kao
Rano Raraku RRA3
Ringarooma Humus 1
Ringarooma Humus 2
Ringarooma River Site 1
Ringarooma River Site 2
Rotten Swamp
Ryans Swamp 2
Saint Louis Lac
Sapphire Swamp
Scott River Swamp
Sirunki Wabag (A16)
TDFO 2 XERO 1
XERO 2 TDFO 1
DSFW 3
DSFW 3
CTRF 2 DSFW 1
DSFW 3
DSFW 2 TUND 1
DSFW 3
Stockyard Swamp
Summit Bog
Sundown Point
Supulah Hill
Tarcutta Swamp
Tegepangwa
Telago Swamp
Thool Swamp
Tidal River
DSFW 2
WTRF 2 COCO 1
DSFW 3
WTRF 2 TDFO 1
DSFW 2
TDFO 2
Uko’a Pond Core 2
Uko’a Pond Core 3
Umej
Vunimoli Swamp
Wanda
Weld River Swamp
Wyrie Swamp W3
Yarlington Tier
1424
Table 8 continued
WTRF 2 TDFO 1
XERO 1 WTRF 1
STEP 3
DSFW 2
TRFO 1 TDFO 1
Micalong Swamp
DSFW 3
Myalup Swamp
Nadrau Swamp
TUND 2 TDFO 1
Nong Han Kumphawapi (Core 3)
Nong Pa Kho
North Lake
Plum (Plum Swamp, Core 2)
WTRF 2 TDFO 1
Polblue Swamp
DSFW 3
Quincan Crater
WTRF 3
Rano Aroi
XERO 2 TDFO 1
Tugupugua (Core 1)
18 ka
DSFW 2
WTRF 1
TDFO 1
DSFW 3
TDFO 2
TDFO 3
XERO 3
TDFO 3
DSFW 2
WTRF 3
WTRF 3
XERO 1
TDFO 1
WTRF 2 TRFO 1
STEP 1 XERO 1
DSFW 2
WTRF 2
DSFW 3
XERO 2 DSFW 1
DSFW 2
WSFW 1
TDFO 1
CTRF 1
CTRF 2 TRFO 1
DSFW 2
TDFO 2
TDFO 3
DSFW 2
WSFW 2
WTRF 1
CTRF 3
COCO 1
CTRF 1 WTRF 1
WTRF 1
XERO 1 TDFO 1 STEP 1 TSFO 1
TDFO 1
STEP 1
TDFO 1 STEP 1
TDFO 3
WTRF 2
TDFO 3
WTRF 1 TDFO 1
WTRF 2 TDFO 1
DSFW 2
DSFW 2
CTRF 1 TUND 1 DSFW 1DSFW 2
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Figure 3 Modern biomes reconstructed from surface pollen data. In cases where multiple samples from a site yielded different biome
reconstructions, the biome given by most of the samples is plotted. In cases where it was impossible to determine a robust biome, the site is
plotted as an open circle. The biome reconstructions for all the samples from a site are shown in Table 8, permitting an evaluation of the
robustness of the mapped reconstructions to be made.
woods/scrub or temperate or tropical grassland and xerophytic shrubland vegetation). This lack of data makes it
difficult to evaluate how well the biomization procedure
captures the arid biomes characteristic of a large part of the
SEAPAC region.
The robustness of the modern biome reconstructions can be
assessed by the fact that at 55% of the 83 sites with multiple
samples, the biomization procedure consistently yields the
same result (Table 8). However, there are regional differences
in the robustness of the biomization: only 48% of sites from
the tropics yield robust results, while 63% of sites from the
extratropical Pacific islands and mainland Australia yield
consistent results. Most of the sites with multiple biome
assignments, both in the tropics and the extratropics, are from
small swamps or bogs, surrounded by relatively open vegetation, where the regional pollen rain is likely to be variably
affected by local flowering with consequent dilution of the
PFTs characteristic of the regional forest vegetation. Some sites
with multiple biome assignments in the tropics are in regions
that are heavily impacted by human activities. Thus, at least
some of the samples from these sites are likely to reflect local
disturbance rather than regional vegetation. The youngest
samples reflect the recent expansion of agriculture. The two
archaeological sites in the modern data set (Lanyon House and
Cave Bay Cave) both yield non-robust biome reconstructions,
presumably because the pollen input to such sites is far from
natural.
Quantitative comparisons of the reconstructed and observed
biomes (at the location of the pollen sites) indicate that 56% of
the 227 sites with robust biome reconstructions from Australia
in the modern pollen data set are correctly attributed by the
biomization procedure (Table 9a). The distribution of DSFW
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1425
E. J. Pickett et al.
Table 9 Comparison of biomes as predicted by the SEAPAC biomization procedure and observed biomes: (a) for mainland Australia and
(b) for the greater SEAPAC region. The order of the biomes reflects the order used in the tie-break procedure. Modern biomes for Australia
were derived from the Australian vegetation map (Figure 1, Table 7) and observed biomes for the greater SEAPAC region were derived from
original site descriptions given in Table 1. Sites with biomization results that are non-robust (see Table 8) are not included in these
comparisons
Reconstructed biomes
(a)
TRFO
TSFO
TDFO
WTRF
COCO
CTRF
WSFW
STEP
XERO
DSFW
TUND
Total obs.
% Correct
(b)
TRFO
TSFO
TDFO
WTRF
COCO
CTRF
WSFW
STEP
XERO
DSFW
TUND
Total obs.
% Correct
TRFO
TSFO
TDFO
4
7
WTRF
COCO
2
8
WSFW
STEP
1
XERO
3
2
1
5
1
1
3
4
2
10
50
13
8
3
0
15
30
1
55
27
6
7
1
16
18
1
3
1
5
1
1
3
2
1
1
12
0
0
n/a
0
n/a
1
1
12
67
0
n/a
DSFW
TUND
7
3
5
3
1
2
99
1
120
83
2
0
1
1
3
34
4
11
15
1
2
2
2
50
1
1
2
1
4
0
2
43
79
1
29
52
2
0
n/a
(83%), WTRF (67%), and CTRF (50%) are correctly predicted
most often. Quantitative comparisons of the reconstructed and
observed biomes (at the location of the pollen sites) from the
remainder of the SEAPAC region, indicate that 37% of the 142
sites in the modern pollen data set are correctly attributed by
the biomization procedure (Table 9b). The distribution of
tropical deciduous broadleaf forest and woodland (79%),
WTRF (52%) and tropical evergreen broadleaf forest (50%)
are correctly predicted most often. The persistent problems in
the modern biomization procedure provide guidelines about
the likely accuracy of the fossil biome assignments. In
particular:
1. The occurrence of DSFW is overestimated, at the expense
of both temperate rain forests (WTRF, CTRF) and WSFW at
the wetter end of the range, and xerophytic woods/scrub
(XERO) towards the arid end of the range. This appears to
reflect the overwhelming dominance of Eucalyptus in all of
these biomes. Eucalyptus spp. are prolific pollen producers,
forming the uppermost canopy layer in the vegetation and thus
releasing pollen well above the ground. Being small, the pollen
1426
CTRF
2
17
12
1
0
1
37
0
2
4
0
0
n/a
5
0
is also readily transported long distances by the wind (e.g.
Dodson, 1983; Kershaw & Strickland, 1990). In contrast, the
PFTs which serve to distinguish DSFW from temperate rain
forests and WSFW (i.e. tree ferns, grasses, heath) and from
xerophytic woods/scrub (i.e. eurythermic drought-tolerant
forbs) are generally present in low abundances, are often low
pollen producers, are confined to the lower strata of the
vegetation, and are often not wind pollinated (Keighery, 1982;
Dodson, 1983; Wills, 1989; Kodela, 1990; Boyd, 1992; Kershaw
et al., 1994).
2. The occurrence of tropical deciduous broadleaf forest and
woodland (TDFO) appears to be overestimated in the modern
biomization. This is partly due to the misclassification of
tropical semi-evergreen broadleaf forest (TSFO) and CTRF/
WTRF, to tropical deciduous broadleaf forest and woodland,
but more importantly due to the misclassification of sites from
more arid biomes (DSFW, xerophytic woods/scrub, and
temperate or tropical grassland and xerophytic shrubland).
The misallocation of the rain forest sites largely reflects the fact
that the characteristic PFTs (e.g. tropical drought-deciduous
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
malacophyll broadleaved tree, tropical evergreen malacophyll
broad-leaved tree) are comparatively low pollen producers
(Flenley, 1973; Kershaw & Strickland, 1990). The fact that
Eucalyptus is an important component of the SEAPAC tropical
deciduous broadleaf forest and woodland as well as the biomes
DSFW, and xerophytic woods/scrub, may be a partial
explanation for the misclassification of sites from more arid
environments.
Despite these systematic biases, the modern biomization
appears to capture the observed vegetation patterns sufficiently
well to merit applying the method to fossil pollen data in order
to reconstruct biome maps for 6 and 18 ka. Furthermore, we
cannot rule out the possibility that some of the apparent
mismatches in our quantitative validation result from the use
of a vegetation map which represents Australian vegetation
prior to European colonization and is, to some extent, based
on inferences about the nature of post-colonial vegetation
changes.
Mid-Holocene biomes
The robustness of the 6 ka biome reconstructions, as measured
by the number of sites in which multiple samples yield the
same result (69%), is similar to that obtained in the modern
biomization (Table 8). The mid-Holocene vegetation patterns
(Fig. 4) are broadly similar to those of today, suggesting that
the 6 ka climate was not radically different. Nevertheless, there
are spatially coherent changes in biome distribution between
6 ka and today. Of the 93 sites with a record for both 0 and
6 ka (Table 10), 32% show a change.
The broad-scale patterns of vegetation distribution across
south-eastern Australia at 6 ka were very similar to today. In
Figure 4 Biomes reconstructed from fossil pollen data at 6 ka. In cases where multiple samples from a site yielded different biome
reconstructions, the biome given by most of the samples is plotted. In cases where it was impossible to determine a robust biome, the site is
plotted as an open circle. The biome reconstructions for all the samples from a site are shown in Table 8, permitting an evaluation of the
robustness of the mapped reconstructions to be made.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1427
E. J. Pickett et al.
Site
0 ka Biome
Aluaipugua (core 2)
Barker Swamp
Baw Baw Village
Bega Swamp
Bibra Lake (BL2)
Black Swamp
Blue Lake (Mt Gambier)
Bobundarra Swamp
Boggy Lake
Boggy Lake
Boggy Swamp
Brass Tarn
Breadalbane NW
Breadalbane SE
Bunyip Bog
Butchers Swamp
Buxton
Caledonia Fen
Chapple Vale
Club Lake core 1
Cobrico Swamp
Cotter Source Bog
Crystal Bog 1
Delegate River/Tea Tree Swamp
Digger’s Creek Bog
Dragon Tree Soak
Fitzgerald Inlet Core 2
Freshwater Lake
G5-2-053P (Seram Trench)
Haeapugua, Core 6
Hidden Swamp
Hogayaku Lake
Ijomba
Jacksons Bog A
Jacksons Bog B
Komanimambuno
Kosipe
Koumac Northeast
Kurnell Fen Core 1
Lac Suprin
Lake Banganup (BGP4)
Lake Cartcarong
Lake Curlip
Lake Elusive
Lake George
Lake Gnotuk
Lake Habbema
Lake Hill
Lake Hordorli
Lake Keilambete
Lake Leake (core LE)
Lake Mountain
Lake Tagamaucia
Lake Turangmoroke
Lake Tyrrell (Site 2)
Lake Wangoom
Lashmar’s Lagoon
Loch McNess Swamp
CTRF
XERO
DSFW
DSFW
DSFW
DSFW
DSFW
STEP
TDFO
DSFW
DSFW
WTRF
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
XERO
DSFW
DSFW
DSFW
XERO
XERO
DSFW
TDFO
TDFO
DSFW
XERO
TUND
DSFW
DSFW
WTRF
TDFO
TDFO
DSFW
WTRF/TRFO
TDFO
DSFW
DSFW
DSFW
STEP
DSFW
TDFO
DSFW
WTRF
DSFW
DSFW
DSFW
WTRF
DSFW
DSFW
DSFW
DSFW
DSFW
1428
6 ka Biome
18 ka Biome
Table 10 Changes in biomes at sites with a
record for 0 ka and either 6 or 18 ka
CTRF
XERO
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
WTRF
DSFW
DSFW
DSFW
DSFW
CTRF
DSFW
CTRF
DSFW
DSFW
XERO
DSFW
DSFW
DSFW
STEP
XERO
DSFW
TDFO
WTRF
DSFW
COCO
TDFO
TUND/DSFW
DSFW
WTRF/TDFO
XERO
DSFW
CTRF
CTRF
TDFO
DSFW
DSFW
DSFW
DSFW
WSFW
TDFO/DSFW
DSFW
TDFO
DSFW
WTRF
DSFW
XERO
DSFW
WTRF
XERO
XERO
DSFW
XERO
DSFW
WTRF/TSFO
DSFW
Not used
XERO
CTRF/WSFW
XERO
XERO
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Table 10 Continued
Site
0 ka Biome
6 ka Biome
Loch Sport Swamp
Long Swamp
Micalong Swamp
Morwell Swamp
Moss Bed Lake
Mount Burr Swamp
Mustering Flat
Myalup Swamp
Nong Han Kumphawapi (Core 3)
Nong Pa Kho
North Torbreck
Oaks Creek
Plum (Plum Swamp, Core 2)
Polblue Swamp
Poley Creek
Powelltown
Quincan Crater
Rano Aroi
Rano Raraku RRA3
Rooty Breaks Swamp
Rotten Swamp
Scott River Swamp
Sheet of Water
Snobs Creek
Stockyard Swamp
Storm Creek
Tarcutta Swamp
Tawonga Bog
Thompson River
Tidal River
Tom Burns, D
Tower Hill Main
Tower Hill NW Crater
Tugupugua (Core 1)
Uko’a Pond Core 2
Umej
Wanda
Weld River Swamp
West Basin
West Lake Muir Swamp
Yarlington Tier
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
XERO
TDFO
DSFW
DSFW
WTRF
DSFW
WTRF
DSFW
WTRF
XERO
XERO
DSFW
DSFW
XERO
DSFW
DSFW
DSFW
DSFW
DSFW
DSFW
CTRF
DSFW
DSFW
DSFW
DSFW
CTRF
XERO/TDFO/STEP
XERO
TDFO
WTRF
DSFW
DSFW
CTRF/TUND/DSFW
DSFW
XERO
DSFW
DSFW
DSFW
DSFW
DSFW
TDFO
XERO
the western part of south-eastern Australia, xerophytic
woods/scrub (XERO) occurred at 6 ka at five sites characterized today by DSFW. This difference indicates conditions
somewhat drier than today at 6 ka. Sites in the Snowy
Mountains, in the Southern Tablelands and east of the Great
Dividing Range show more moisture-demanding vegetation
at 6 ka than today. Today most of these sites are characterized by DSFW, whereas the vegetation at 6 ka was variously
WSFW (two sites), WTRF (1), CTRF (4) and cool evergreen
needleleaf forest (2).
Only three sites in Western Australia show a different biome
at 6 ka than today. Weld River Swamp (Churchill, 1961, 1968)
was characterized by less moisture-demanding vegetation
18 ka Biome
TDFO
CTRF
CTRF
WTRF
DSFW
WTRF
DSFW
WTRF
XERO/TDFO
WSFW
DSFW
DSFW
DSFW
COCO
DSFW
COCO
DSFW
WTRF
CTRF
WSFW
DSFW
DSFW
DSFW
COCO/WTRF
TSFO/TDFO/STEP
TDFO
WTRF/TDFO
DSFW
XERO
TDFO
DSFW
WTRF
DSFW
STEP/XERO
TDFO
DSFW
TUND
XERO
CTRF/WTRF
TDFO
(DSFW) at 6 ka than today. However, Scott River Swamp
(Churchill, 1961, 1968) which is classified as xerophytic
woods/scrub (XERO) today was characterized by DSFW at
6 ka. This biome shift implies wetter conditions. The single site
in north-western Australia (Dragon Tree Soak: Pedersen, 1983;
Wyrwoll et al., 1986) shows conditions drier than today at
6 ka.
In the tropical highlands of PNG, Sumatra and Irian Jaya,
sites that are occupied today by either tundra or CTRF (i.e.
upper montane forest) were characterized by the presence of
WTRF (i.e. lower montane forest) and even tropical deciduous
broadleaf forest and woodland at 6 ka. These changes imply a
rise of > 500 m in the limits of the montane forest belts and in
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1429
E. J. Pickett et al.
treeline at 6 ka. The largest shift in these limits occurs at the
highest sites.
Last glacial maximum biomes
There are 33 sites with a record for the LGM. The samples
from Lake George (Singh & Geissler, 1985) have been
omitted from further analysis because of the extremely low
pollen counts (23 and 33 grains, respectively). The robustness
of the 18 ka biome reconstructions, as measured by the
number of sites in which multiple samples yield the same
result (44%), is less than that obtained in the modern
biomization (Table 8). Thus, there is more uncertainty
attached to both the biome reconstructions and any climatic
interpretation of these data. Nevertheless, the differences
between the 18 ka (Fig. 5) and modern (Fig. 3) biome maps
are striking. Of the 22 sites with a record for both 0 and
18 ka (Table 10), 50% show a change.
Xerophytic woods/scrubs (XERO) were more extensive at
18 ka than today in south-eastern and south-western Australia.
In the north, tropical deciduous broadleaf forest and woodland
(TDFO) is recorded at 18 ka in sites characterized today by
tropical evergreen broadleaf forest (TRFO). These changes
suggest conditions drier than today at the LGM across
mainland Australia.
In the tropics, some sites that lie today within the montane
forest belts (WTRF, CTRF) were characterized by tundra at
18 ka, while WSFW occurred in sites characterized by tropical
evergreen broadleaf forest (TRFO) today. These changes reflect
a lowering of treeline and montane forest belts by between 300
and 1500 m at 18 ka. The magnitude of the downslope shifts
in vegetation belts are comparable with those reconstructed by
Figure 5 Biomes reconstructed from fossil pollen data at 18 ka. In cases where multiple samples from a site yielded different biome
reconstructions, the biome given by most of the samples is plotted. In cases where it was impossible to determine a robust biome, the site is
plotted as an open circle. The biome reconstructions for all the samples from a site are shown in Table 8, permitting an evaluation of the
robustness of the mapped reconstructions to be made.
1430
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
Farrera et al. (1999), and indicate that conditions were
between 1 and 9 C colder than today. The largest temperature
lowering occurs at highest elevation. The encroachment of
tropical deciduous broadleaf forest and woodland (TDFO)
into the tropical lowlands suggests that it was also drier than
today.
DISCUSSION
The biomization procedure
Previous applications of the biomization method to a number
of regions in the Northern Hemisphere and the tropics
(Prentice et al., 1996; Jolly et al., 1998b; Tarasov et al., 1998;
Yu et al., 1998; Edwards et al., 2000; Elenga et al., 2000;
Takahara et al., 2000; Thompson & Anderson, 2000; Williams
et al., 2000; Yu et al., 2000; Bigelow et al., 2003) have shown
that the technique works well across the entire range of global
climates and vegetation types. The SEAPAC biomization
provides a further test of the method, in a region characterized
by significant temperature gradients (from equatorial to cooltemperate zones) and precipitation gradients (from arid to
extremely wet). Our application of the method to modern
pollen data from the SEAPAC region resulted in the correct
classification of 56% of the samples from mainland Australia,
and 37% of the samples from the rest of the SEAPAC region.
The allocation of samples to biomes controlled primarily by
temperature appears to be good. However, there are considerably more difficulties in correctly predicting the boundaries
of biomes that are controlled by moisture availability, specifically the transitions from DSFW to tropical deciduous
broadleaf forest and woodland and to tropical semi-evergreen
broadleaf forest in the north, and the transitions from WSFW
to DSFW to xerophytic woods/scrub and ultimately to
grassland and xerophytic shrubland in the temperate zone.
This appears to reflect the fact that these biomes are dominated
by a few genera (e.g. Eucalyptus, Acacia) which are tolerant of a
wide range of conditions and which respond to increasing
moisture stress through changes in height, structure and
spacing.
There are a number of plausible explanations for why the
discrimination between biomes controlled by moisture availability is less good than that between biomes controlled by
temperature.
1. The temperature limits on the growth of specific PFTs are,
in general, determined by the presence/absence of specific
physiological adaptations amongst the dominant, canopyforming taxa. For example, there are physiological adaptations
that enable some trees to survive frost (e.g. dormancy, freezedrying). Tropical evergreen malacophyll broadleaf trees have
no mechanism to survive frost and therefore cannot grow in
regions subject to frost. The frost limit therefore provides an
absolute constraint on the distribution of biomes which
includes tropical evergreen malacophyll broadleaf trees, e.g.
tropical evergreen broadleaf forest and WTRF. Although
specific adaptations do exist to compensate for moisture stress
(e.g. ephemeral growth forms, water storage mechanisms), the
primary response to moisture stress appears to be expressed
through structural changes in the dominant PFTs, e.g. changes
in tree height, foliage density or canopy spacing (Specht, 1972).
These responses are largely determined by competition
amongst individuals for a limited resource (i.e. water) rather
than reflecting a physiological control on the individual itself.
Thus, specific PFTs appear to be capable of surviving under a
wide range of moisture conditions through structural plasticity. For example, evergreen sclerophyll broadleaf trees are an
important component of the vegetation in regions with mean
annual rainfall ranging from > 1100 to < 500 mm year)1. In
high rainfall areas, they are tall, single-stem trees forming
closed-canopy forests. Increasing moisture stress is registered
by increasing openness of the canopy and, in general, by a
gradual reduction in tree height.
2. Since the dominant, canopy-forming PFTs display
considerable plasticity with respect to moisture stress, discrimination between biomes at the more arid end of the
moisture gradient tends to be made on the basis of PFTs which
form the subcanopy and understorey components of the
vegetation. These PFTs are in general less abundant. Furthermore, as they release pollen within or beneath the canopy, the
pollen tends to be less widely dispersed. Additionally, some of
the characteristic species are poor pollen producers (e.g.
Acacia, Proteaceae) or rely primarily on dispersal by birds,
insects and animals (e.g. Banksia). As a result, the presence/
abundance of these PFTs in the pollen spectrum from a specific
site tends to be less consistent than the presence/abundance of
the dominant canopy-forming PFTs, with consequent effects
on the reliability of the biome allocation.
3. The difficulties caused by a reliance on non-canopy
forming PFTs to discriminate between biomes are further
compounded by the fact that pollen discrimination amongst
the non-tree taxa is limited. The grasses (Poaceae) are a classic
example of this problem. Spinifex grasslands are confined to
regions with mean annual rainfall of < 200 mm year)1 and are
replaced by tussock grasslands in more well-watered areas
(Mott & Groves, 1994). In the modern landscape, the presence
of spinifex could be used to discriminate between xerophytic
woods/scrub and DSFW in the temperate zone, or between
tropical deciduous broadleaf forest and woodland and tropical
semi-evergreen broadleaf forest in the tropics. As it is not
possible to distinguish reliably between different types of grass
on the basis of their pollen (Pedersen, 1983; Boyd, 1990), this
criterion cannot be used in pollen-based biomizations.
Our attempt to improve the discrimination of moisturelimited biomes has not been entirely successful. Nevertheless,
given that these biomes are distinctive features of the modern
vegetation landscape and occupy extensive areas of the
SEAPAC region, the attempt to discriminate between them
should not be abandoned. Analysis of the underlying causes of
the uncertainties in allocations between the moisture-limited
biomes suggest improvements could be made by:
1. Using pollen data with a better level of taxonomic
resolution for biomization. It would be possible, for example,
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1431
E. J. Pickett et al.
to improve the level of discrimination within some families
and genera that are important in the SEAPAC region. The
regular identification of different taxonomic groups within
Eucalyptus (Pike, 1956; Martin & Gadek, 1988; Pickett &
Newsome, 1997) would considerably improve our ability to
discriminate between moisture-limited biomes. Furthermore,
in cases where the level of pollen identification cannot be
improved, it might be possible to use plant macrofossils from
the same sediments to help determine which species were
present (e.g. Jackson et al., 1997).
2. Using macrofossil data to reconstruct the biomes in
regions where pollen data are sparse. Macrofossil data from
packrat middens have already been successfully used to
reconstruct arid vegetation types in western North America
(Thompson & Anderson, 2000). The stickrat midden data
from the arid interior of Australia (Pearson & Dodson, 1993;
McCarthy et al., 1996; Pearson, 1999) could be used in a
similar fashion.
3. Using carbon isotopes to distinguish between grass types.
Carbon isotope measurements have been used successfully to
discriminate between the preponderance of C3/C4 grasses in
the diets of emus, and hence to infer changes in the
distribution of these types of grasslands during the past
60 ka (Miller et al., 1997, 1999; Johnson et al., 1999). Direct
measurements of the carbon isotopic composition of plant
macrofossils and possibly even pollen could potentially
improve our discrimination of understorey components that
are key to the separation of arid zone biomes.
In addition to the attempt to make finer discriminations
between moisture-stressed biomes (e.g. with the introduction
of WSFW and DSFW), the SEAPAC biomization has introduced two new rain forest biomes (WTRF and CTRF). We are
able to discriminate between these biomes, and to distinguish
them from other biomes that are limited by temperature rather
than moisture availability, with some accuracy. Some of these
biomes occur today in other regions of the world, including
New Zealand, South America and south-eastern China.
Furthermore, it seems likely that there are structural equivalents to these biomes in the mountains of equatorial Africa.
Successful discrimination of these biomes in the SEAPAC
region suggests that attempts to reconstruct the modern and
palaeodistribution of these distinctive vegetation types in other
regions would be worthwhile.
Vegetation and climate of the SEAPAC region in the
mid-Holocene
The 6 ka biome reconstruction suggests the broad-scale
patterns of vegetation distribution across south-eastern Australia during the mid-Holocene were similar to today. Western
Australia and the western part of south-eastern Australia were
characterized by less moisture-demanding vegetation at 6 ka
than today, implying that the climate was drier. In contrast,
sites in the Snowy Mountains, in the Southern Tablelands, and
east of the Great Dividing Range show more moisturedemanding vegetation at 6 ka than today, implying wetter
1432
climate. The vegetation records from mountain sites in the
tropics indicate that the upper altitudinal limits of montane
forest belts and treeline were higher than today, implying
warmer temperatures.
These reconstructed changes are broadly consistent with
previous reconstructions of mid-Holocene vegetation patterns,
based on a more limited number of sites (e.g. Markgraf et al.,
1992; Harrison & Dodson, 1993; Kershaw, 1998). The expansion of more moisture-demanding vegetation in much of
south-eastern Australia is consistent with Markgraf et al.’s
(1992) inference that WSFW and temperate rain forest
elements were extended in the south-eastern highlands of
mainland Australia between c. 8 and 5 ka. As pointed out by
Markgraf et al. (1992), the vegetation evidence for wetter
conditions is also consistent with lake-level evidence from the
region (Markgraf et al., 1992; Harrison, 1993). Furthermore
Harrison & Dodson (1993) also show an expansion of
moisture-demanding forests in this region at 6 ka. Markgraf
et al. (1992) also suggested that temperate rain forest was
present at higher elevations in Tasmania and in northern
Queensland. We have no data to verify the upward expansion
of rain forest in Tasmania. Biome reconstructions for sites in
Queensland do not indicate a change in the extent of rain
forest. Markgraf et al. (1992) suggested that the forests and
shrublands in PNG were more extensive than today; this is
consistent with our reconstruction that the tropical montane
forest belts occurred at higher elevations than today. Both
Markgraf et al. (1992) and Harrison & Dodson (1993) suggest
that the limited amount of data available from Western
Australia implies conditions wetter than present during the
mid-Holocene. Our reconstruction shows little change in this
region compared with today, although two sites seem to
indicate conditions may have been slightly drier.
On the basis of experiments using the BIOCLIM model, Nix
& Kalma (1992) have suggested that northern Australia and
PNG had higher winter precipitation during the mid-Holocene
than today. In these experiments, the increased precipitation
occurred because the Arafura Sea, which was shallower and
warmer than today, acted as an enhanced local moisture
source. Our reconstructions suggest that the biome changes in
northern Australia and PNG are a reflection of changes in
temperature rather than precipitation. However, they are not
inconsistent with the hypothesis put forward by Nix & Kalma
(1992).
Vegetation and climate of the SEAPAC region at the
last glacial maximum
The reconstructed patterns of biome distributions at 18 ka are
strikingly different from today. Xerophytic woods/scrub and
DSFW characterized the southern part of mainland Australia.
Further north, tropical deciduous broadleaf forest and woodland is recorded in sites characterized today by tropical
evergreen broadleaf forest or WTRF. These changes suggest
that conditions may have been drier than today at the LGM
across mainland Australia. This is not consistent with
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
Palaeovegetation patterns for Australia and Southeast Asia
lake-level data from the south-east interior (where we have no
pollen records) which indicate conditions somewhat wetter
than today (Harrison & Dodson, 1993). However, vegetation
registers changes in growing-season soil moisture whereas
lakes register changes in runoff (Cheddadi et al., 1997; Farrera
et al., 1999). Thus, the changes in regional moisture conditions
recorded by vegetation data need not be congruent with the
signal derived from lakes, particularly in regions with highly
seasonal precipitation regimes (see e.g. Prentice et al., 1992b;
Farrera et al., 1999; Harrison et al., 2001).
The biome changes between LGM and present in mainland
Australia do not allow us to estimate the degree of cooling at
the LGM. However, the reconstructed biome changes in the
tropics, which indicate that treeline and the altitudinal limits
of the montane forest belts were lower than today, clearly
indicate temperatures between 1.7 and 9 C (depending on
elevation) colder than today. The encroachment of tropical
deciduous broadleaf forest and woodland into areas occupied
today by lowland tropical evergreen broadleaf forest suggests it
was also somewhat drier than today in the tropics.
Previous reconstructions of vegetation at the LGM (van
der Kaars, 1991; Markgraf et al., 1992; Harrison & Dodson,
1993) consistently show conditions more arid and colder
than today. Harrison & Dodson (1993) argue that the interval
between 15 and 18 ka was cooler and drier than any
subsequent time. Our reconstructions are consistent with,
but in general less extreme than, the changes implied by
previous syntheses of the LGM vegetation data. For example,
van der Kaars (1991) shows grassland/shrubland across
northern Australia and onto the Sunda-Sahul shelf with
woodland and open forest replacing lowland tropical forests
in southern PNG. According to this reconstruction, tropical
lowland forests were very much reduced in extent compared
with today. Our reconstruction implies a more widespread
persistence of tropical lowland forests. This apparent difference may be because the van der Kaars (1991) reconstruction
is based on pollen records from marine cores in the eastern
Indonesian region, which can be expected to blend the signals
from montane and lowland sites within the same area. As
tropical lowland forest species tend to produce less pollen
than montane equivalents, the reduction in tropical lowland
forests is likely to be exaggerated. Similarly, our reconstruction is in good agreement with Markgraf et al.’s (1992)
reconstruction of treeline lowering in PNG, savanna woodland in tropical northern Australia, and the extension of
shrubland in interior south-eastern Australia. However,
Markgraf et al. (1992) suggest that coastal regions of the
south-east (including Tasmania) were characterized by steppe
vegetation at the LGM. The biome reconstructions indicate
xerophytic woods/scrub vegetation across south-eastern Australia; there are no data from Tasmania in the current data
set.
A multi-proxy compilation of data on temperature and
moisture-balance changes in the tropics (Farrera et al., 1999)
indicates that temperatures at sea level within the SEAPAC
region were of the order of 1–2 C colder than present.
However, the lapse rate was considerably steeper than today,
resulting in much larger temperature changes (6–9 C) at high
elevation sites. These estimates are consistent with the shifts in
treeline and montane forest belts shown by our biomization
procedure, and the estimates of temperature changes made
from these shifts. The Farrera et al. (1999) compilation also
indicates that plant-available moisture was less than today in
the tropical part of the SEAPAC region, consistent with our
reconstructed biome shifts.
CONCLUSIONS
This paper represents a first attempt to map the vegetation of
the SEAPAC region from pollen data using an objective
method based on the recognition of biomes characterized by a
unique assemblage of PFTs. The method is broadly successful.
Difficulties in reliably discriminating between biomes controlled by moisture gradients largely reflect the fact that
adaptations to moisture stress tend to be expressed through
structural changes rather than species replacement. However,
some improvement in discrimination could be made if the
data available for biomization were more highly resolved
taxonomically. The regular identification of subgroups of
Eucalyptus would certainly help in this respect, as would the
use of plant macrofossils to identify the likely species-level
identification of taxa which cannot be resolved to species level
from pollen. Similarly, isotopic methods could be used to
identify whether the grasses present in a pollen assemblage
were predominantly C3 or C4 types.
We have applied the biomization technique to reconstruct
vegetation patterns at 6 and 18 ka. The changes in biome
distribution at 6 ka compared with present are small, implying
that the climate of the mid-Holocene was not significantly
different from present. The changes in biome distribution at
18 ka are more striking, and suggest that the SEAPAC region
was drier than today and, at least in the tropics, colder. These
reconstructions are consistent with earlier, subjective reconstructions of vegetation and climate changes in the SEAPAC
region.
The Southern Hemisphere is 75% ocean and thus the ocean
is the primary determinant of Southern Hemisphere regional
climates. Accurate simulation of palaeoclimates using atmospheric general circulation models (AGCMs) would therefore
require well constrained ocean reconstructions for the Southern Hemisphere whereas the available ocean data, particularly
from the extra tropics, is limited (CLIMAP Project Members,
1981; Broccoli & Marciniak, 1996; Rosell-Melé et al., 1998). As
a result AGCM palaeoclimate simulations appear to capture
the regional climates of the Southern Hemisphere less well
than those of the Northern Hemisphere (Harrison & Dodson,
1993, Markgraf, 1993). The use of coupled ocean-atmosphere
models (e.g. Hewitt & Mitchell, 1998; Braconnot et al., in
press), which explicitly simulate ocean circulation should
improve our ability to simulate Southern Hemisphere regional
palaeoclimates. The palaeovegetation reconstructions provided
here will be useful as a means of evaluating their performance.
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd
1433
E. J. Pickett et al.
ACKNOWLEDGMENTS
We thank Pat Bartlein, Gerhard Bönisch, Julie Delaney, Jed
Kaplan, Caroline Roelandt, Silvana Schott, Kerstin Sickel,
Helmut Staoubeurauch, and Natalia Tchounareva for technical
assistance, and Pat Bartlein, Bob Thompson and Rob Marchant for reviews of earlier versions of the paper. Financial
support for this work (LP) was provided by the Max Planck
Institute for Biogeochemistry and the TEMPO project. The
SEAPD data base is lodged at Monash University (http://
www.geography.monash.edu.au). The INDOPAC data base is
lodged at NOAA/NGDC (http://www.ngdc.noaa.gov/paleo).
The SEAPAC data base is part of the BIOME 6000 data base
and is available from MPI-BGC (http://www.bgc-jena.mpg.de/
bgc_prentice/). BIOME 6000 was supported by IGBP-GAIM,
IGBP-DIS, IGBP-GCTE and IGBP-PAGES.
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BIOSKETCHES
This paper is a contribution to the ongoing work of the Global
Palaeovegetation Mapping (BIOME 6000) Project. BIOME
6000 is a community-wide collaboration inaugurated in 1994
under the joint auspices of four of the elements of the International Geosphere–Biosphere Project: IGBP-GAIM, IGBPDIS, IGBP-GCTE and IGCP-PAGES. The aims of BIOME
6000 are to create fully documented pollen and plantmacrofossil data sets for 6000 and 18,000 14C yr bp, and to
use these data sets to construct global maps of major
vegetation types (biomes) using a standard objective biomization technique based on plant functional types (PFTs). The
resulting maps are designed to be used for climate model
evaluation within the framework of the Palaeoclimate Modelling Intercomparison Project (PMIP).
Journal of Biogeography 31, 1381–1444, ª 2004 Blackwell Publishing Ltd