Journal of Biogeography (J. Biogeogr.

) (2004) 31, 1893–1908

ORIGINAL Species diversity and endemism of five

ARTICLE
major Malesian islands: diversity–area
relationships
Marco C. Roos1*, Paul J. A. Keßler1, S. Robbert Gradstein2 and Pieter Baas1

1
National Herbarium of the Netherlands,
University of Leiden branch, Leiden, The
Netherlands and 2Department of Systematic
Botany, Albrecht von Haller Institute of Plant
Sciences, Göttingen, Germany
*Correspondence: Marco C. Roos, National
Herbarium of the Netherlands, University of
Leiden branch, PO Box 9514, 2300 RA Leiden,
The Netherlands.
E-mail: roos@nhn.leidenuniv.nl
ABSTRACT
Aim A comparison of biodiversity patterns within Malesia in relation to surface
area.
Location Analysis of the patterns in species richness and endemism of vascular
plants in the five major Malesian islands, i.e. Java, Sulawesi, Sumatra, Borneo and
New Guinea.
Methods Available data on species richness and species ranges in correlation with
the surface area of the respective islands were examined in this work. Estimations
of total species numbers for these islands are presented based on extrapolation of
all available published Flora Malesiana information and recent checklists, all in all
comprising 12,000 different species. The regression analysis of overall species
richness and endemism were studied for all species together as well as for different
plant families to compare the fit with the Arrhenius species–area model.
Results The five islands form a series of independent areas of increasing size
suited for an analysis of the species–area relationships at the regional scale. All
species taken together and those of families with even distribution throughout
Malesia show significant species–area relationships. Non-significant relationships
were detected in families with western or eastern-centred Malesian distribution
patterns. Relationships between number of endemic species and surface area are
significant for all species and for the majority of the families with significant
species–area relationships.
Main conclusions Species–area relationships of families appear to be dependent
on species number. Families with high numbers of species usually have a
significant species–area relationship whereas small families have not. For the
families that display an eastern or western Malesian centred pattern, a historical
biogeographical explanation should be invoked. Island surface area appears to be
a predictor for island percent endemism in Malesian vascular plants. None of the
islands appears to be a hotspot of endemism nor of species diversity, as no
significant departure from the Arrhenius model was noted for any of them.
Keywords
Malesia, islands, species richness, endemism, species–area relations.

Following field research on the woody plant diversity in

INTRODUCTION
Geographical patterns in plant diversity are basic to the
recognition, circumscription and understanding of areas of
high overall biodiversity (Mittermeier et al., 1999). With
c. 41,000 species of vascular plants, the Malesian region, the
archipelago between mainland Asia and Australia (Fig. 1), is
among the most species-rich areas world-wide (Roos, 1993).
Sulawesi by the second author (e.g. Keßler et al., 2002;
Pitopang et al., 2004; Kessler et al., in press), the question
arose what the position of the island of Sulawesi is in Malesia
with respect to its woody plant diversity. We have tackled this
question by comparing species richness and endemism of
Sulawesi with that of the other major islands in the Malesian
region, i.e. Java, Sumatra, Borneo and New Guinea. Together

ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi 1893

M. C. Roos et al.
30
20
10
Sulawesi
0 Borneo New Guinea
Sumatra
–10 Java
90 100 110 120 130 140
they form a series of independent islands of increasing surface
area within one archipelago (instead of nested sampling; this is
irrespective of Java, Sumatra and Borneo having been part of
the Sunda subcontinent in the past) and are therefore well-
suited for an analysis of species–area relationships, more
specifically the Arrhenius equation1 (e.g. Rosenzweig, 1995).
Species–area relationship, or the notion that larger areas
have more species, have been frequently studied (e.g. Rose-
nzweig, 1995) and several explanations have been proposed for
this phenomenon (Connor & McCoy, 2001): (1) the ‘Habitat
diversity’ hypothesis2), (2) the ‘Area per se’ hypothesis3, (3) the
‘Passive sampling’ hypothesis4; and additionally (4) the
‘Speciation rate’ hypothesis5 (Losos & Schluter, 2000). Often
the first hypothesis is regarded the best predicting model (Fox
& Fox, 2000; Triantis et al., 2003). Although at first sight our
data may not be detailed enough to differentiate between these
possible causing factors, we may be able to at least indicate
which factors might have been the most determining ones.
Regarding habitat heterogeneity, there is no geomorphologi-
cal and topographical evidence indicating substantial habitat
differences among the five selected islands, with the exception
of the alpine zone in New Guinea, which is higher and more
extensive than on any of the other islands. Reasons for assuming
major differences in speciation, extinction and dispersal among
the five main islands of the Malesian archipelago are also
1
log S ¼ z log A + log c or S ¼ cAz (S ¼ number of species,
A ¼ area, c ¼ constant coefficient, z ¼ parameter related to the eco-
logical and geographical conditions of the area).
2
Large areas house more species than small areas because they have a
greater variety (heterogeneity) of habitats.
3
Large areas have more species than small areas because less species go
locally extinct in large areas.
4
Large areas are more likely to receive more colonists than small areas.
5
Large areas show higher rates of speciation than small areas (for
islands larger than 3000 km2).
1894
150 160 Figure 1 Map of Malesia (1:50,000,000).
lacking. All islands with the exception of Sulawesi are compar-
able regarding geological histories and distances to the main-
land. Therefore, it is expected that the impact of dispersal on the
present biodiversity patterns is comparable for the four islands.
Sulawesi, however, has been isolated for most of its geological
past and is situated far from both the Asiatic as well as the
Australian mainland. But at the same time it is at cross-roads
between the Sunda Shelf (Java, Sumatra, Borneo) in the west
and the Sahul Shelf (New Guinea) in the east (Whitmore, 1991).
This leads to the expectation that the species–area patterns to
be found will follow the equilibrium model of island bio-
geography of MacArthur and Wilson (see Rosenzweig, 1995).
Species numbers of individual families may show deviating
patterns as a result of family specific biogeographical histories.
MATERIALS AND METHODS
The data on the plant diversity of Java, Sumatra, Borneo,
Sulawesi and New Guinea have been retrieved from Flora
Malesiana (Series I, Spermatophyta (1940–2002) & Series II,
Pteridophyta (1959–2002), all hitherto published volumes) and
recent checklists (e.g. Johns, 1987; Hay, 1995; Keßler et al., 1995,
2002; Van Welzen, 1997b) (Table 1). Figures for estimated total
number of vascular plant species of Malesia and percent
endemism for the major Malesian islands, based on information
on c. 15% of the whole flora, were obtained from Roos (1993)
and Van Welzen (1997a). In spite of the large amount of data
available, knowledge of the plant species diversity in Malesia is
still very uneven. Of 120 woody plant families recorded from
Sulawesi, only 45% have been treated in Flora Malesiana.
Taxonomic revisions or regional overviews of many important
families are still lacking, including, for example, the Rubiaceae.
Plant collection density, expressed as the number of collections
per 100 km2, varies from c. 200 for Java to c. 35 for Borneo
(Ashton, 1989; Burley, 1994) and less than 25 for Sulawesi
(Whitten et al., 1987; P.J.A. Keßler, unpubl. data), New Guinea
(Conn, 1994) and Sumatra (Ashton, 1989); cf. Thailand: 50
(Parnell et al., 2003). When accepting a minimum of 100
Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd
 Species diversity and endemism of five major Malesian islands

Table 1 Data on treated plant species in

Treated no. Endemics Extrapolated Extrapolated
Flora Malesiana and recent checklists, with
Area of species found Endemism total no. of no. of
extrapolations for the total number for
(·1000 km2) (2002) (2002) (%) species endemics
Malesia
Java 140 1696 165 9.7 5809 565
Sulawesi 185 1765 217 12.3 6045 743
Sumatra 475 3098 373 12.0 10,611 1278
Borneo 740 4211 1194 28.4 14,423 4089
New Guinea 865 4046 1886 46.6 13,858 6460
Malesia 3040 12,000 4250 35.4 41,100 14,556

The proportion of the species treated until 2002 (third column) in combination with the edu-
cated guess of a total of 41,100 species in Malesia is used to extrapolate and estimate the total
number of species per island (sixth column); the latter estimations in combination with the
percent endemism of taxa treated until 2002 (fifth column) are used to extrapolate and estimate
the total number of endemic species per island (seventh column).

collections per 100 km2 for a flora to be adequately known region (‘Pan-Malesian range’; Table 2, last column). Taken
(Whitten et al., 1987), only the island of Java has been together, they include over 30% of the woody species of
adequately collected. For this reason, we have restricted our Sulawesi and c. 30% of the whole Malesian flora (woody and
analysis to selected families (Tables 2 and 3). A much more non-woody). To obtain a broader-based picture of plant
detailed study using specimen databases has recently started. diversity of the five islands, we have additionally included
Families were chosen based on their size, geographical range, selected herbaceous families for which solid taxonomic data
importance in Sulawesi in terms of species richness and were available. Our dataset should be considered representative
abundance (Keßler et al., 2002), and availability of adequate of the Malesian flora, including some of the largest families
taxonomic treatments. The families analysed each have min- and covering a broad variety of life-forms, ranging from herbs
imally 40 species and are distributed throughout the Malesian to emergent trees, from terrestrial to epiphytic, from auto-
trophic to hemi-parasitic and from lowland to alpine taxa.
Table 2 Species diversity of selected families in the major The total number of vascular plant species (ALL) was used as
islands of the Malesian region (based on recent Flora Malesiana reference for the patterns of the different families (Tables 2 and 3).
treatments and/or information of specialists)
Table 3 Endemic species diversity of selected families
New (endemism unknown in Annonaceae, Lauraceae and Urticaceae)
Java Sulawesi Sumatra Borneo Guinea Malesia
New
Area 140 185 475 740 865 3040 Java Sulawesi Sumatra Borneo Guinea Malesia
(·1000 km2)
Annonaceae 61 50 300 500 400 850 Area (·1000 km2) 140 185 475 740 865 3040
Lauraceae 73 100 200 300 247 700 Annonaceae na na na na na na
Urticaceae 19 34 35 50 40 250 Lauraceae na na na na na na
Anacardiaceae 15 26 48 95 34 151 Urticaceae na na na na na na
Podocarpaceae 6 19 17 21 39 61 Anacardiaceae 0 0 1 21 17 104
Dipterocarpaceae 8 7 96 267 15 390 Podocarpaceae 0 0 0 12 9 43
Euphorbiaceae 209 300 314 440 461 1593 Dipterocarpaceae 2 2 11 155 11 334
Fagaceae 26 9 55 100 30 180 Euphorbiaceae 48 49 52 151 335 1000
Meliaceae 39 50 88 120 82 223 Fagaceae 4 0 5 31 20 157
Myristicaceae 15 18 61 110 138 335 Meliaceae 0 2 4 21 46 134
Araceae 83 46 156 252 126 728 Myristicaceae 1 12 7 69 112 305
Rosaceae 28 19 32 31 46 111 Araceae 36 13 99 194 90 639
Nepenthaceae 1 8 26 34 11 83 Rosaceae 2 0 2 5 30 73
Cunoniaceae 1 8 2 3 31 40 Nepenthaceae 1 4 18 25 7 75
Boraginaceae 20 11 10 14 37 65 Cunoniaceae 0 5 0 1 22 37
Loranthaceae 32 23 37 58 59 200 Boraginaceae 2 1 1 1 20 39
Mimosaceae 32 29 33 37 66 150 Loranthaceae 5 2 3 20 35 145
Polypodiaceae 55 62 77 77 73 183 Mimosaceae 0 3 2 11 36 96
Sapindaceae 31 35 39 62 145 235 Polypodiaceae 0 9 8 16 20 107
Caesalpiniaceae 42 24 62 90 42 197 Sapindaceae 0 5 0 17 95 172
ALL 1696 1765 3098 4211 4046 12,000 Caesalpiniaceae 1 2 6 36 17 130
ALL 165 217 373 1194 1886 4250
ALL ¼ all woody and non-woody taxa for which adequate informa-
tion is available (see also text). na: Not available. For further explanation see Table 2.

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M. C. Roos et al.

Table 4 Regression analysis of three parameters of selected Malesian plant families: (1) all species ¼ species per island in relation to
the total Malesian number of species for the respective taxa (power regression); (2) endemic species ¼ endemic species per island in relation
to the total Malesian number of endemic species for the respective taxa (power regression; exponential regression also given in case of
a much lower significance); (3) percent endemism ¼ proportion of endemic species of the total number of species per island and for
Malesia as a whole (logarithmic regression; other regression also given in case of a much lower significance)

All species Endemic species Percent endemism

Family z-value R2 Significance z-value R2 Significance c-value R2 Significance

Annonaceae 1.2861 0.935 0.007 na na na na na na

Lauraceae 0.7252 0.961 0.003 na na na na na na
Euphorbiaceae 0.3650 0.863 0.022 0.8820 0.683 0.085 0.1656 0.579 0.079
Lin. 0.905 0.013 Exp. 0.839 0.029 Power 0.596 0.072
Meliaceae 0.5089 0.845 0.027 6.0280 0.645 0.101 0.2093 0.740 0.028
Lin. 0.800 0.041
Myristicaceae 1.2444 0.996 0.000 1.9976 0.748 0.058 0.2150 0.455 0.142
Lin. 0.823 0.034
Loranthaceae 0.4360 0.786 0.045 1.1440 0.578 0.136 0.2126 0.755 0.025
Lin. 0.903 0.013 Exp. 0.749 0.058
Lin. 0.777 0.048
Polypodiaceae 0.1665 0.833 0.031 5.4439 0.509 0.176 0.1659 0.851 0.009
Lin. 0.829 0.032 Lin. 0.923 0.002
Sapindaceae 0.6322 0.675 0.088 4.8277 0.262 0.378 0.2411 0.706 0.036
Exp. 0.811 0.037
Anacardiaceae 0.6599 0.616 0.116 8.9562 0.969 0.002 0.2385 0.828 0.012
Podocarpaceae 0.6642 0.648 0.100 8.1539 0.689 0.082 0.2408 0.733 0.030
Lin. 0.673 0.089 Exp. 0.820 0.034
Caesalpiniaceae 0.3717 0.380 0.268 1.7241 0.916 0.011 0.2118 0.905 0.004
Dipterocarpaceae 1.3082 0.439 0.223 1.7468 0.646 0.101 0.2171 0.675 0.045
Fagaceae 0.7252 0.435 0.226 4.3519 0.346 0.297 0.2720 0.773 0.021
Araceae 0.6148 0.611 0.118 1.0568 0.683 0.085 0.1824 0.849 0.009
Mimosaceae 0.2870 0.518 0.171 5.7495 0.588 0.130 0.2141 0.806 0.015
Rosaceae 0.2925 0.572 0.139 4.9402 0.336 0.306 0.2224 0.671 0.046
Lin. 0.673 0.089
Urticaceae 0.3642 0.692 0.080 na na na na na na
Nepenthaceae 1.3280 0.610 0.119 1.2677 0.654 0.097 0.0230 0.020 0.788
Cunoniaceae 0.8280 0.258 0.382 4.1168 0.188 0.466 0.2198 0.411 0.170
Boraginaceae 0.2021 0.097 0.610 0.6615 0.175 0.484 0.1712 0.597 0.072
Lin. 0.617 0.064
ALL 0.5324 0.982 0.001 1.2341 0.903 0.013 0.1034 0.588 0.075
Exp. 0.978 0.001 Power 0.681 0.043
Choros model
ALL 0.3901 0.945 0.006 0.9083 0.876 0.019 0.1140 0.632 0.108
Exp. 0.974 0.002 Exp. 0.898 0.014

For comparison the figures for ALL are given following the CHOROS model of Triantis et al. (2003; see text). na, Not available.

Of each family, total species richness and endemic species
richness for each of the investigated islands as well as for the
whole of Malesia (Tables 2 and 3), were counted and plotted
against area size. These data were subjected to three regression
analyses (Table 4):
• Total number of species per island per family and overall
(ALL)
• Number of endemic species per family and overall (ALL)
• Percent endemism per family and overall per island
(endemic nisland total n
1
island )
To scale down the differences in magnitude between
families and facilitate comparisons between them, species
richness figures in the first and second analysis were presented
as percentages of total numbers per area (nisland n
1total , endemic
nisland endemic n
1 total ) [ntotal ¼ all Malesian species in the
sampling], i.e. indicating the contribution of each family and
island to total Malesian species richness. The regression
analysis was carried out using SPSS, of which the results are
presented in Table 4. Following the Arrhenius model the
species as well as the endemic species figures are analysed
according to a Power or Linear regression. The size of the
Malesian Islands analysed is far larger than 3000 km2, which
marks the division between small-sized islands and large-sized
islands according to Losos & Schluter (2000). These authors
discuss the difference of scale in relation to patterns in species–
area relations (see also the discussion of break point regression

1896 Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd

 Species diversity and endemism of five major Malesian islands

0.9
New Guinea
0.5324
0.8 y = 0.0097x
2
R = 0.9823
Borneo

Species numbers / total species numbers

0.7

0.6
Sumatra
0.5
Sulawesi
0.4

Java
0.3

0.2

0.1

Figure 2 Graph of the data on species and

0
area size of selected plant families and of all
0 200 400 600 800 1000
sampled species in Malesia. The solid line is
Area size
the power regression of ALL. The symbols
Annonaceae Lauraceae Urticaceae Anacardiaceae
indicate the species numbers for the selected Podocarpaceae Dipterocarpaceae Euphorbiaceae Fagaceae
families in five vertical columns (from left to Meliaceae Myristicaceae Araceae Rosaceae
right) representing Java, Sulawesi, Sumatra, Nepenthaceae Cunoniaceae Boraginaceae Loranthaceae
Borneo, resp. New Guinea. Area site is in all Mimosaceae Polypodiaceae Sapindaceae Caesalpin
figures ·1000 km2. ALL Power (ALL)
Percentage Endemic species / total numbers endemic species

0.7

0.6

0.5

0.4

0.3

0.2

0.1

0
0 200 400 600 800 1000
Area size
Anacardiaceae Podocarpaceae Dipterocarpaceae Euphorbiaceae
Figure 3 Graph of the data on (endemic) Fagaceae Meliaceae Myristicaceae Araceae
species and area size of ALL (for explanation Rosaceae Nepenthaceae Cunoniaceae Rafflesiaceae
see the legend of Fig. 2), showing the most Boraginaceae Loranthaceae Viscaceae Mimosaceae
relevant cq. significant regression lines (see Polypodiaceae Davalliaceae Sapindaceae Caesalpin
Table 4). ALL Exp (ALL)

in Lomolino & Weiser, 2001). This means that the Malesian species numbers are expected to increase with area size much
islands are positioned in the middle or upper part of the steeper than the species numbers, the former have been also
generally accepted sigmoid species–area curve over the whole analysed with the option exponential regression. There is no a
range of scales (Rosenzweig, 1995; Connor & McCoy, 2001; priori model for the percent endemism, which is therefore
Hubbell, 2001). Therefore, a linear or power type regression analysed with the options ‘Linear’, ‘Power’, ‘Exponential’ and
is expected according to these models. As the endemic ‘Logarithmic’.

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M. C. Roos et al.
0.5
0.45
Species numbers / total species numbers
0.4
0.35
0.3
0.25
0.2
0.15
0.1
0.05
0
0 200
all species
percent endemism
Power (endemics)
Log (percent endemism)
RESULTS AND DISCUSSION
Species richness
The calculated figures for total and endemic species richness of
vascular plants for each of the five islands are given in Table 1.
Total species richness extrapolated for Java (c. 5800 spp.) and
Sulawesi (c. 6050 spp.) are considerably higher than the figures
published by Backer & Bakhuizen van den Brink (1963–1968;
Java, 4598 native spp.) and Whitten et al. (1987; Sulawesi,
5000 spp.). For Java our figure may be too high, due to
overrepresentation of Javanese collections in the early volumes
of Flora Malesiana. Our estimation for Sulawesi is based on a
verifiable extrapolation of Flora Malesiana data, whereas
Whitten et al. (1987) do not give any source information.
Our figure for Sumatra (c. 10,600 spp.) is somewhat higher
than that of De Wilde (1989: 8–10,000) and comparable with
that of Kiew (2002: 10,000–11,200), that for Borneo (c. 14,500
spp.) is close to the upper estimate of Ashton (1989:
10–15,000) but deviates from the figures of Soepadmo (1995:
10–12,000) and Davis (1995: 20–25,000). As the basis for the
latter two figures is unknown, we cannot explain the major
difference between these estimates. Our figure for New Guinea
(c. 14,000 spp.), finally, is a bit lower than those given by Johns
(in Davis, 1995: 15–20,000) and Frodin (2001: 15–17,000) that
are both based on personal experience, but without any
quantitative source information presented.
All species taken together (ALL) show a highly significant
species–area relationship (Table 4, Figs 2–4). The same hold
for the species–area–habitat relationship as recently proposed
by Triantis et al. (2003), that encompasses a model (the choros
model) for species diversity, which embodies number of
1898
400 600 800 1000
Area size
Figure 4 Graph of the data on (endemic)
species and area size of ALL (for explanation
endemics
Power (all species) see the legend of Fig. 2), showing the most
Exp (endemics) relevant cq. significant regression lines (see
Exp (percent endemism) Table 4).
species, area and habitat diversity6 and mathematically unifies
area per se and habitat hypotheses (Table 4, Fig. 5). Crucial in
their model is the specific determination of the ‘habitat’ term,
which has to be defined based on the natural history of the
taxon studied. In applying their model to the areas included in
the present study, we adopted the main habitat types from
MacKinnon (1997; slightly adapted), as a fair surrogate for the
total number of habitat types. The number of main habitat
types increases with area size (Table 5). As the results of the
choros model of Triantis et al. (2003) and the Arrhenius
species–area model are very similar (the R2 values of the latter
seemingly are a little higher), and because of the lack of
information regarding the habitat occupation of the families,
the choros model has not been further pursued (Triantis et al.,
2003, conclusions were based on data on a much smaller
geographical scale than the islands included in the present
study).
The z-value1 of the species–area relation of ALL (0.5324) is
lower than the figure mentioned by Rosenzweig (1995) for
intercontinental tropical rain forest provinces (0.97), which is
in line with the differences in z-values between regional
(relatively low) and intercontinental (relatively high; see
Hubbell, 2001) scales. The z-value for the endemic species
(1.2341) is indeed much higher, as predicted by Harte & Kinzig
(1997); see below). In the case of endemic species the
exponential relation is even more significant (Table 4). The
z-value for ALL according to the species–area–habitat relation
(0.3901) is lower than the one of the species–area relation
which is in line with the results of Triantis et al. (2003).
6
S ¼ c(HA)z (H ¼ number of habitats).
Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd
 Species diversity and endemism of five major Malesian islands

0.5

0.45

Species numbers / total species numbers

0.4

0.35

0.3

0.25

0.2

0.15

0.1

0.05

0
0 2000 4000 6000 8000 10,000 12,000
Figure 5 Graph of the data on the choros
Area size*habitats
model of (endemic) species, area size and
all species endemics
habitat types of ALL (for explanation see the percent Power (ALL species)
legend of Fig. 2), showing the most relevant Power (endemics) Exp (endemics)
cq. significant regression lines (see Table 4). Log (percent endemism) Exp (percent endemism)

Table 5 Distribution of the major habitat types in Malesia, Separate species–area regression analysis for the 20 selected
adapted after MacKinnon (1997) pan-Malesian families (Fig. 2) yielded two groups of families:
1. Families having significantly more species in larger islands,
New
hence complying with the species–area relationship (Figs 6–8)
Major vegetation types Java Sulawesi Sumatra Borneo Guinea
They are all among the 12 largest families selected. This group
Area (·1000 km2) 140 185 475 740 865 includes the woody families Annonaceae, Lauraceae, Euphor-
Lowland evergreen x x x x x biaceae, Meliaceae (although the regression line seems steeper
tropical rain forest than that of ALL) and Myristicaceae, as well as the parasitic
Montane evergreen x x x x x Loranthaceae and the epiphytic fern family Polypodiaceae, all
tropical rain forest showing the expected Power regression. The Sapindaceae are
Limestone forest x x x x x
deviating by showing an Exponential regression, whereas the
Ironwood forest x x
most significant regression for the Loranthaceae is actually
Tropical Pine forest x
Peat swamp forest x x
Linear. The Polypodiaceae shows by far the lowest z-value
Mangrove x x x x (0.1665; Table 4). This may indicate that this fern family
Heath forest x x comprises species with relatively large ranges compared with
Fresh water swamp forest x x x x the flowering plant families. As for many of the latter families
Tropical deciduous x x detailed species ranges are not available yet, we take the
monsoon forest distribution patterns of the Malesian seed plants and terrestrial
Montane deciduous forest x ferns as treated in Flora Malesiana so far as indicators. Of this
Alpine shrub vegetations x x selection c. 80% of the species show a range comprising one or
Ultrabasic rock vegetation x x x two of the selected islands, the remaining 20% occur in three
Semi-evergreen forest x x x x
to five of the islands. The Polypodiaceae species, however,
Beach forest x x
show a completely different picture as 65% occur on one to
Alpine grassland x
Lake x x x x
two islands only and 35% on three to five islands.
Savannah/grassland x 2. Families not exhibiting a significant relationship between
Total no. of habitat types 6 9 11 11 13 species richness and area size (Figs 9–11). This group includes
Area-habitat figures 840 1665 5225 8140 11,245 12 families; six of them have a western Malesian distribution
pattern (the woody families Anacardiaceae, Dipterocarpaceae,
Fagaceae and Caesalpiniaceae, and the herbaceous Araceae and
Here again the z-value of ALL endemic species–area–habitat Nepenthaceae; Fig. 10), two an eastern Malesian pattern
relation (0.9083) is more than two times higher than for ALL (Podocarpaceae and Cunoniaceae, all woody; Fig. 9), and the
species. remaining four (Urticaceae, Rosaceae, Boraginaceae and

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M. C. Roos et al.

0.7

0.6
Species numbers / total species numbers

0.5

0.4

0.3

0.2

0.1

0
0 200 400 600 800 1000
Area size Figure 6 Graph showing the regression lines
Annonaceae Lauraceae Myristicaceae of the families with a significant power
ALL Power (ALL) Power (Annonaceae) species–area relation and a z-value higher
Power (Lauraceae) Power (Myristicaceae) than that of ALL.

0.6
Species numbers / total species numbers

0.5

0.4

0.3

0.2

0.1

Figure 7 Graph showing the regression lines

0
of the families with a significant Power
0 100 200 300 400 500 600 700 800 900 1000
species–area relation and a z-value lower than
Area size
that of ALL. For the Euphorbiaceae the
Euphorbiaceae Meliaceae Polypodiaceae expected Power regression and the somewhat
ALL Power (ALL) Power (Euphorbiaceae) more significant Linear regression are
Power (Meliaceae) Power (Polypodiaceae) Lin (Euphorbiaceae) presented; both lines are almost parallel.

Mimosaceae; Fig. 11) more complex patterns that cannot Hopkins & Hoogland (2002). The Gondwana hypothesis
easily be interpreted. explaining current biodiversity patterns of the latter two
The species richness patterns of the second group of families, families has also been proposed for the pan-tropical Dipter-
lacking a significant correlation with area size, might possibly ocarpaceae (Ashton, 1983). The distribution pattern of the
explained by historical biogeographical factors, as has already Sapindaceae does not deviate significantly from the Arrhenius
been proposed for Dipterocarpaceae by Ashton (1983), for model (significance 0.088), but shows a low number of species
Fagaceae by Soepadmo (1972), and for Podocarpaceae and on Sumatra. The aberrant diversity pattern of the Boraginaceae,
Cunoniaceae (both with highest species richness in Sulawesi finally, showing high species numbers on Java and low ones
and New Guinea) by De Laubenfels (1988) and Fortune elsewhere, is probably a collecting artefact (Veldkamp, 2001).

1900 Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd

 Species diversity and endemism of five major Malesian islands

0.7

0.6

Species numbers / total species numbers

0.5

0.4

0.3

0.2

0.1

Figure 8 Graph showing the regression lines

of the families with a most significant Linear 0

(Loranthaceae) or Exponential (Sapindaceae) 0 100 200 300 400 500 600 700 800 900 1000

species–area relation. For the former family Area size

the significant power relation is also presen- Loranthaceae Sapindaceae ALL
ted, which runs almost parallel to the Linear Power (ALL) Exp (Sapindaceae) Lin (Loranthaceae)
one. Power (Loranthaceae)

0.9

0.8

0.7
Species numbers / total species numbers

0.6

0.5

0.4

0.3

0.2

0.1

0
Figure 9 Graph showing the species–area 0 100 200 300 400 500 600 700 800 900 1000
data on the families with an eastern Malesian Area size
distribution pattern. The figures for Sulawesi
and New Guinea are relatively high. Podocarpaceae Cunoniaceae ALL

Endemism and hotspots
Endemic species–area relationship was significant for all
endemic species taken together (ALL; Table 4, Figs 3 and 4).
This indicates that concentrations of endemics do not occur
at the island level in Malesia. Either the whole of Malesia is
one hotspot or the concept of hotspots does not hold in this
region. The rationale behind hotspots (sensu Mittermeier
et al., 1999; Myers et al., 20007) is to identify cases where
diversity and, additionally, endemism departs positively in a
significant manner from an otherwise generally supported
7
Biogeographical units of quite unequal size featuring exceptional
concentrations of endemic plant and vertebrate species and experien-
cing exceptional loss of habitat.

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M. C. Roos et al.

0.8

0.7
Species numbers / total species numbers

0.6

0.5

0.4

0.3

0.2

0.1

0
Figure 10 Graph showing the species–area
0 200 400 600 800 1000
Area size
data on the families with an western Malesian
distribution pattern. The figures for Borneo
Anacardiaceae Dipterocarpaceae Fagaceae
Araceae Nepenthaceae Caesalpin are relatively high, and those of Sulawesi and
ALL New Guinea relatively low.

0.6
Species numbers / total species numbers

0.5

0.4

0.3

0.2 Figure 11 Graph showing the data on the

families with a rather complex species–area
0.1 pattern. The Urticaceae seem indifferent to
area-size, but the information on this family
0 is less referenced than the other families
0 200 400 600 800 1000 selected. The mainly herbaceous Boragina-
Area size
ceae, and the predominantly woody Rosaceae
and Mimosaceae all show relatively high fig-
Urticaceae Rosaceae Boraginaceae Mimosaceae ALL ures for Java and low ones for Borneo.

relationship with area (New Caledonia provides a good
botanical example). Mittermeier et al. (1999) define three
hotspots (i.e. Sundaland, Philippines and Wallacea) and one
wilderness area8 (Papuasia) together covering the whole of
Malesia. Consequently, Malesia is regarded a hotspot area
all over, and an endemic species–area relation can be
expected within the region indeed, which is actually found
in our study. The studies of Bibby & Adam (1992),
revealing a number of different small-sized endemic bird
areas in the region, as well as of Davis (1995), recognizing
8
Large and intact biogeographical units mainly defined by vascular
plant diversity and relatively free from transformative ecological
impact by humans.
over 100 (very) small-sized botanical hotspots throughout
Malesia, indicate that it might be possible to differentiate
between areas within the region and delineate hotspots at a
local level. According to this idea, various patterns of
endemicity should be expected, which is actually found for
a number of families in our study. However, our data do
not allow for a more detailed analysis per island. Such
analyses will be part of the recently started research project
using our growing specimen-based data bases, as mentioned
above2.
About half (eight) of the investigated families also show a
significant endemic species–area relationship (Table 4;
Figs 12–14). Five of these (Euphorbiaceae, Meliaceae, Myris-
ticaceae, Loranthaceae, Polypodiaceae) also show a significant
species–area relationship for total species numbers (see above),

1902 Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd

 Species diversity and endemism of five major Malesian islands

Percentage endemic species / total numbers endemic species

0.6

0.5

0.4

0.3

0.2

0.1

0
0 200 400 600 800 1000
Area size

Anacardiaceae Caesalpin
Figure 12 Graph showing the regression
ALL Power (Anacardiaceae)
lines of the families with a significant Power
endemic species–area relation. Power (Caesalpin) Power (ALL)

0.5
Percentage endemic species / total numbers endemic species

0.45

0.4

0.35

0.3

0.25

0.2

0.15

0.1

0.05

0
0 200 400 600 800 1000
Area size
Podocarpaceae Euphorbiaceae
Figure 13 Graph showing the regression
ALL Exp (Podocarpaceae)
lines of the families with a significant Expo-
nential endemic species–area relation. Exp (Euphorbiaceae) Exp (ALL)

the other three (Anacardiaceae, Podocarpaceae and Caesalp- It thus appears that five of six of the families exhibiting a
iniaceae) do not. Sapindaceae is the only family showing significant species–area relationship for which also data on
significant species–area relationship for all species but not for endemics are available, also show to an endemic species–area
endemic ones. Annonaceae and Lauraceae, also significant relationship (with only the Sapindaceae as exception; see
regarding total species, could not be analysed due to lack of Table 4, central column). It seems that, if a taxon displays a
data on endemism. significant relation between the size of an island and species

Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd 1903

M. C. Roos et al.

0.5
Percentage endemic species / total numbers endemic species

0.4

0.3

0.2

0.1

0
0 200 400 600 800 1000

-0.1
Area size
Meliaceae Myristicaceae Loranthaceae
Polypodiaceae ALL Lin (Meliaceae) Figure 14 Graph showing the regression
Lin (Myristicaceae) Lin (Loranthaceae) Lin (Polypodiaceae) lines of the families with a significant Linear
Power (ALL) endemic species–area relation.

0.7
Endemic species numbers / total endemic species numbers

0.6

0.5

0.4

0.3

0.2

0.1

0
0 200 400 600 800 1000
Figure 15 Graph showing a comparison
Area size
Euphorbiaceae spec Euphorbiaceae end Myristicaceae spec
between the figures for species–area and
Myristicaceae end Loranthaceae spec Loranthaceae end endemic species–area relationships of selected
Sapindaceae spec Sapindaceae end Power (Euphorb)
Power (Euphorb end)
Power (Loranth)
Power (Myrist)
Power (Loranth end)
Power (Myrist end)
Power (Sapind)
families with significant species–area
Power (Sapind end) relations.

numbers, this may be indicative of a high richness in endemic steeply correlated with area, small areas having very few
species as well. endemics but these numbers increasing quickly as area size
According to Harte & Kinzig (1997), in their theoretical increases. Such a difference is only reflected in the regression
model based on nested sets of areas, both species number and curves of the Meliaceae and Polypodiaceae, and in ALL (Figs 16
endemic species number show significant species–area rela- and 4, respectively). The two respective regression curves of the
tions. Number of endemics, however, supposedly is more families Euphorbiaceae, Myristicaceae and Loranthaceae run

1904 Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd

 Species diversity and endemism of five major Malesian islands

0.8

Endemic species numbers / total endemic species numbers

0.7

0.6

0.5

0.4

0.3

0.2

0.1

Figure 16 Graph showing a comparison 0

between the figures for species–area and 0 200 400 600 800 1000
endemic species–area relationships of selected Area size
families with significant (Power) species–area
Meliaceae spec Meliaceae end Polypodiaceae spec
relations (for clarity reasons presented in a
Polypodiaceae end Power (Meliaceae) Power (Meliaceae end)
separate graph in view of exceptional
regression lines for endemic species). Power (Polypod) Power (Polypod end)

0.9

0.8

0.7
Borneo
Endemic species / total species

0.6

0.5 Sumatra

0.4
Java Sulawesi
0.3

0.2

0.1

0
0 100 200 300 400 500 600 700 800 900 1000
- 0.1

- 0.2
Area size
Anacardiaceae Podocarpaceae Dipterocarpaceae Fagaceae
Meliaceae Araceae Rosaceae Loranthaceae
Mimosaceae Polypodiaceae Sapindaceae Caesalpin
ALL Log (Anacard) Log (Podocarp) Log (Dipterocarp)
Figure 17 Graph showing all families with a Log (Fagaceae) Log (Meliaceae) Log (Araceae) Log (Rosaceae)
significant log percent endemism regression Log (Lorantha) Log (Mimosaceae) Log (Polypodia) Log (Sapindac)
curve. Log (Caesalpin) Log (ALL)

almost parallel, whereas the Sapindaceae are deviating with a regression curves of the majority of the families are quite
very low regression curve for the endemic species (Fig. 15). similar; only the Polypodiaceae show a different angle of the
In contrast to the picture emerging regarding the endemic curve (indicating a lower increase in percent endemism, which
species–area relationship (Fig. 16), the percent endemism (i.e. may seem paradoxical in view of the exceptional steep
the proportion of the endemic species in relation to the total regression line of the endemic species–area relation in Fig. 16;
number of species) for 12 of the 17 families is significantly again, this might be related to the relative large range size of the
correlated to the size of the island (Table 4, Fig. 17). The species) and the curve of the Araceae and Dipterocarpaceae

Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd 1905

M. C. Roos et al.
shows higher average values (indicating many small-ranged
species). The finding that the majority of the pan-Malesian taxa
show higher percent endemism in larger-sized islands, irres-
pective of their total species number and distribution within the
archipelago is a rather counter-intuitive novelty as a random
pattern was expected. However, our findings mean that even
when the families are unevenly distributed over Malesia, the
proportion of endemic species per island is still related to area
size. It follows that percent endemism for pan-Malesian families
are normally highest in New Guinea and lowest in Java,
irrespective of the biogeographical history of the group. It is not
clear what the mechanism behind this pattern may be. One
possible explanation is that larger areas give colonists more
opportunities to deviate from their mother species (even if
further radiation does not occur; see Losos & Schluter, 2000). If
Heaney (2000) is correct in stating that in old archipelagos
cladogenesis on the individual islands is a more important
factor determining their respective species richness than
colonization, these endemic species should comprise endemic
clades; otherwise it is expected that each of these endemic
species should show a vicariant pattern with a sister species on
another island. This should be tested in future phylogenetic
research. Some phylogenies of particular taxa may indicate
support for Heaney’s model with regard to Borneo (Ridder-
Numan, 1996), Sulawesi (Holloway, 2003) and New Guinea
(Van Welzen, 1990; Turner, 1995; B. Gravendeel, unpublished
data; Schot, in press), whereas Haegens’ (2000) results are more
indicative for the vicariant model.
The position of Sulawesi within Malesia
Total species richness and endemic species richness of Sulawesi
show similar patterns as those of the other four Malesian
islands (Figs 5 & 6), in spite of the very different geological
history of Sulawesi and the far greater distance of the island to
the mainland (Whitmore, 1991). Whereas the islands of
Borneo, Sumatra and Java had terrestrial connections to
mainland Asia, also in the last glacial period, Sulawesi was
isolated from these islands as well as from New Guinea by deep
maritime straits (Ridder-Numan, 1996). More recently, Moss
& Wilson (1998), elaborating on the geological history of
Sulawesi, state that western Sulawesi had been accreted to
eastern Borneo by the late Cretaceous. By the early Eocene
there are indications of continuous land masses, but in the
Tertiary (Paleogene), the two areas progressively separated
again. The unusual biogeographical composition of the flora of
Sulawesi, comprising eastern and western Malesian centred
floristic elements, is probably due to the unique geological
history of the island. However, as shown above the long-term
isolation of Sulawesi is not reflected in the plant species
diversity of the island, which complies with the species–area
relation model. The hypothesized effect of isolation for the
biodiversity patterns of Sulawesi (passive sampling hypothe-
sis1), is therefore not supported by our data.
Vertebrates, on the contrary, exhibit patterns, which are
indeed expected following the island biogeography model with
1906
25–75% endemism (higher than all other islands except New
Guinea) and low total species numbers (lower than on Java;
Whitten et al., 1987; MacKinnon et al., 1996). There seems no
plausible correlation to dispersal capacity, as mammals and
birds (quite different in their mobility) show identical patterns.
The plant taxa included in our study also show differences in
dispersal capabilities (with diaspores ranging from spores to
wind and bird dispersed seeds/fruits to big or short-lived
fruits). Further research is needed to reveal what the causes
might be for these different patterns.
CONCLUSIONS
1. Plant diversity figures given in this paper are based on the
most recent and most complete sampling data available for
Malesia and are therefore more reliable than those of previous
authors. Only, the figure for the plant species richness of Java
might be excessively high, due to the relatively high plant
collection density on this island and consequently overrepre-
sentation of the collection information.
2. Species–area relationships of families appear to be depend-
ent on species number. For this reason, families with high
numbers of species usually have a significant species–area
relationship whereas small families have not.
At present, the most plausible hypotheses for explaining these
patterns in the case of the large Malesian islands is that
in larger areas the rate of speciation is higher than the
amount of immigration and extinction (Losos & Schluter,
2000). However, without any information on phylogeny it
is not possible to hypothesize anything definitive on this
matter.
For the families that display an eastern or western Malesian
centred pattern, a historical biogeographical explanation
should be invoked.
3. Percent endemism is positively area size determined, as even
species poor families show higher proportions of endemism in
larger islands.
The results of our analysis of all species together confirms the
model of Harte & Kinzig (1997) of endemic species–area
relationship.
Families with significant results for their total number of
species, almost always show also significant results regarding
endemic species–area relationships. Therefore, speciose taxa
also indicate richness in endemics.
4. No hotspot patterns have been found at island level.
Whatever factors may have played dominant roles none of the
islands shows a pattern, which significantly deviates from
what is expected from the (endemic) species–area relationship.
5. In spite of the long-term geological isolation of Sulawesi and
the far greater distance of the island to the mainland as
compared with other major Malesian islands, species diversity
patterns in vascular plants of Sulawesi are basically similar to
those of other major Malesian islands and in agreement with
species–area and endemic species–area relationships in the
region. Endemism in vascular plants is rather low (<15%),
much lower than in animals.
Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd
 Species diversity and endemism of five major Malesian islands

ACKNOWLEDGMENTS Hay, A. (1995) Checklist and botanical bibliography of the

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BIOSKETCHES
Marco C. Roos is coordinator of the international Flora
Malesiana project and has a great interest in Malesian plant
diversity and especially its biogeography.
Paul J.A. Keßler is studying Malesian tree diversity,
especially of Borneo and Sulawesi, and is an internationally
renowned expert on South East Asian Annonaceae and
Menispermaceae.
S. Robbert Gradstein is a world expert on hepatics and a
project leader of the Research Programme ‘Stability of
Rainforest Margins in Indonesia (Sulawesi)’ (STORMA) of
the German Research Foundation (GFB).
Pieter Baas is a leading expert on comparative wood
anatomy and is director of the NHN actively involved in
Malesian botany.
Journal of Biogeography 31, 1893–1908, ª 2004 Blackwell Publishing Ltd