Journal of Biogeography (J. Biogeogr.) (2010) 37, 1239–1250
ORIGINAL
ARTICLE
The endemic plant families and the palms
of New Caledonia: a biogeographical
analysis
Michael Heads*
Buffalo Museum of Science, 1020 Humboldt
Parkway, Buffalo, NY 14211-1293, USA
*Correspondence: Michael Heads, Buffalo
Museum of Science, 1020 Humboldt Parkway,
Buffalo, NY 14211-1293, USA.
E-mail: [email protected]
ABSTRACT
This paper provides a panbiogeographical analysis of the endemic plant families
and the palms of New Caledonia. There are three endemic plant families in New
Caledonia and several genera that were previously recognized as endemic families.
Of these taxa, some are sister to widespread Northern Hemisphere or global
groups (Canacomyrica, Austrotaxus, Amborella). The others belong to trans-Indian Ocean groups (Strasburgeria), trans-tropical Pacific groups (Oncotheca) or
Tasman Sea/Coral Sea groups (Phelline, Paracryphia) that are sister to widespread
Northern Hemisphere or global groups. In palms, the four clades show allopatric
regional connections in, respectively: (1) western Indonesia, Malaysia and Thailand; (2) Vanuatu/Fiji and the southern Ryukyu Islands near Taiwan; (3) the
western Tasman/Coral Sea (eastern Australia, New Guinea and the Solomon
Islands); and (4) the eastern Tasman/Coral Sea (Lord Howe and Norfolk Islands,
New Zealand, Vanuatu, Fiji and the Solomon Islands). The four clades thus
belong to different centres of endemism that overlap in New Caledonia.
The patterns are attributed not to chance dispersal and adaptive radiation but to
the different histories of the eight terranes that fused to produce modern
New Caledonia. Trans-tropical Pacific connections can be related to the Cretaceous igneous plateaus that formed in the central Pacific and were carried, with
plate movement, west to the Solomon Islands and New Zealand, and east to
Colombia and the Caribbean.
Keywords
Ecology, limestone, New Caledonia, panbiogeography, rain forest, tectonics,
ultramafic, vicariance.
INTRODUCTION
The flora and fauna of New Caledonia are of special interest
through their high levels of diversity, their endemism and their
far-flung geographical affinities. In seed-plants, for example,
there is probably no other region of comparable area in the
world with such a rich and peculiar flora, one which ‘has good
claim to be considered the most remarkable in the world’
(Thorne, 1965, p. 1; cf. Good, 1974). Apart from the endemic
families, discussed below, 105 of the 711 genera (15%) and
2324 of the 3004 indigenous species (77%) are endemic (Jaffré
et al., 2001). It must also be one of the smallest areas in
the world with an endemic bird family (Rhynochetidae) and
there is also an endemic liverwort family (Perssoniellaceae)
(Crandall-Stotler et al., 2009).
Distribution patterns of plant and animal taxa within New
Caledonia and the tectonic development of the region are
ª 2010 Blackwell Publishing Ltd
discussed elsewhere (Heads, 2008a,b, 2009a). This paper
provides an analysis of various plant groups and their global
affinities. The patterns are related to the tectonic history of the
region but, as indicated in the next section, this does not mean
simply interpreting biological patterns in the light of geological
reconstructions. Many aspects of the geology are still unclear,
and data from biogeography constitute a separate source of
evidence that may help resolve tectonic problems. A good
example is the West Caledonian fault. Some geologists deny that
it exists, while others regard it as a major structure. Biogeographical evidence supports the latter view (Heads, 2008b).
The panbiogeographical approach used in this paper
involves an analysis of distribution patterns and a synthesis
of biogeography and geology. In particular, this method
assumes that phylogenetic and geographical breaks are caused
by vicariance, not by the physical movement (dispersal) of taxa
(Heads, 2009b). Physical movement may lead to subsequent
www.blackwellpublishing.com/jbi
doi:10.1111/j.1365-2699.2010.02292.x
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M. Heads
overlap but invoking long-distance dispersal and founder effect
speciation to explain phylogenesis is unnecessary. The New
Caledonian groups studied in this paper were selected as they
include some of the most distinctive clades in the country.
Analyses of their distributions have revealed shared patterns
and these are attributed to common tectonic and phylogenetic
causes rather than to independent dispersal events.
TECTONIC HISTORY
New Caledonia is not simply a fragment of Gondwana but is a
complex mosaic of eight allochthonous terranes – fault-bounded
crustal blocks with independent histories (Heads, 2008a). The
four New Caledonian basement terranes formed from island arcderived and arc-associated material which accumulated in the
pre-Pacific Ocean, not in Gondwana. They amalgamated and
were accreted to Gondwana (eastern Australia) in the Late
Jurassic/Early Cretaceous, but in the Late Cretaceous they
separated from Australia with the opening of the Tasman Sea
and the break-up of Gondwana. A convergent margin with
associated subduction developed near the island and an Eocene
collision of the basement terranes with an island arc arriving
from the north-east – possibly the Loyalty Ridge – is of special
biogeographical interest in connection with New Caledonia–
central Pacific affinities. Thus New Caledonia is a geological
composite formed from Gondwanan and Pacific components, a
similar pattern to that seen in New Guinea and New Zealand.
Many authors have attributed New Caledonian endemism to
the long isolation of the island and the variety of habitat types
present. These factors may account for the survival of an
archaic biota but they do not, on their own, explain its origin.
This is more likely to be related to the different histories of the
terranes that make up modern New Caledonia (Heads, 2008a).
For example, New Caledonia is exceptionally rich in marine
molluscs (Marshall, 2001; Bouchet et al., 2002) and the
country is the world centre of diversity for families such as
the Volutomitridae (Bouchet & Kantor, 2003). Marshall (2001,
p. 41) proposed that ‘Since the Melanesian arc [including New
Caledonia] is situated at current or former [convergent]
boundaries of the Australian and Pacific lithospheric plates,
species richness there is probably due at least partly to
progressive accumulation of taxa transported on the plates’.
This process, with taxa being ‘scraped off’ at a subduction
zone, would account for many biogeographical phenomena in
and around New Caledonia. The strata that the current
endemics originally occupied may have disappeared. Large
structures in the region, such as the South Loyalty Basin
(between Grande Terre – the New Caledonian mainland – and
the Loyalty Islands), have been largely destroyed by subduction
but were probably among the prior areas of endemism that
have left traces in the extant flora and fauna.
Oceanic sediments deposited in the Palaeogene are widespread in New Caledonia. Authors such as Pole (1994) have
accepted the idea that New Caledonia was completely
submerged by Palaeogene marine transgressions and so have
supported an ‘entirely long-distance dispersal’ explanation for
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the biota of the island. Nevertheless, without a stratum of a
single age completely covering the island, total submersion is
difficult to prove from geological data and in such a dynamic
region tectonic uplift and subsidence can be very local. With
respect to biogeography, the Australian eucalypts are represented in New Caledonia by Arillastrum, and Ladiges et al.
(2003) noted that long-distance dispersal is unlikely, due to the
limited dispersal capacity of the seeds. Ladiges et al. (2003,
quoting Russell-Smith et al., 1993) concluded that the presence in New Caledonia of ancient angiosperms, such as
Amborella and Winteraceae, and the high levels of endemism
there also imply that ‘emergent land existed in the vicinity of
New Caledonia throughout its rifted Late Cretaceous and
Cainozoic history’. Authors such as Morat et al. (1984), de
Laubenfels (1996), Lowry (1998), Jolivet (2008) and Jolivet &
Verma (2008a,b) have agreed that complete submergence is
unlikely, based on biogeographical evidence. Very small
tropical islands can preserve diverse biotas and so could have
provided effective microrefugia.
Murienne (2009, p. 1433) argued that phylogenies and
molecular dating provide the ‘necessary framework’ for
analysing New Caledonian biodiversity. He concluded that
the different hypotheses (island refugia during flooding, longdistance dispersal after flooding) cannot be separated on the
basis of the phylogenies alone and so he relied on molecular
dating studies. These have indicated that most New Caledonian groups are of Cenozoic age (cf. Figure 2 in Grandcolas
et al., 2008) and Murienne (2009) argued that for these groups
the ‘only valid interpretation’ is long-distance dispersal.
Nevertheless, the two methods employed in the dating studies
(treating the oldest fossils and clock dates derived from these –
minimum ages – as maximum ages; using the age of the young
islands currently emergent on the old Loyalty Ridge to date the
taxa there) will both give systematic underestimates of clade
age (Heads, 2008a). Murienne (2009) concluded that ‘the
presence of numerous relict groups in New Caledonia remains
puzzling’ (p. 1434), but this is only because the clades were
assumed to be so young. Likewise, while Grandcolas et al.
(2008) accepted that the New Caledonian biota was wiped out
by ‘total submersion’ during Palaeogene flooding and is the
result of long-distance dispersal since 37 Ma, they concluded
that the ‘Phylogenetic relicts remain puzzlingly enigmatic…’
(p. 3312). Neither Murienne (2009) nor Grandcolas et al.
(2008) mentioned the New Caledonian terranes or their
history, although this may provide a simple explanation for
what are otherwise enigmas in the biota.
THE ENDEMIC PLANT FAMILIES
The New Caledonian flora is well known for its endemic
families of angiosperms and, until recently, four or five were
often cited. Molecular studies have clarified their relationships
and now only three are accepted (Amborellaceae, Oncothecaceae and Phellinaceae; Stevens, 2009), but this is still an
unprecedented number for such a small area. Whatever rank
the groups are given, the endemic families and the other
Journal of Biogeography 37, 1239–1250
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Biogeography of New Caledonian plants
endemics sometimes regarded as families are well-documented
and their affinities can be used to illustrate some of the main
biogeographical connections of the island’s biota.
The plant groups concerned are all woody, and Jaffré (1995)
noted that three of them, Paracryphia, Phellinaceae and
Strasburgeria, are well represented in dense, evergreen, montane forest found above c. 1000 m elevation. Ixerba, related to
Strasburgeria, inhabits similar higher-elevation forest in northern New Zealand. Amborella, recorded from 200–1000 m a.s.l.
in New Caledonia, is also absent from the lowest-elevation rain
forest.
1. Amborellaceae. This family comprises one species,
Amborella trichopoda, endemic to New Caledonia. It is the
sister-group (or with Nymphaeales the sister-group) of all
other angiosperms (Brown, 1999; Soltis & Soltis, 2004; Müller
et al., 2006; Qiu et al., 2006). The biogeographical pattern is
similar in both the two alternative relationships, as Nymphaeales are almost cosmopolitan (but are absent in New Zealand
and New Caledonia). Bergthorsson et al. (2004) reported
massive horizontal transfer of mitochondrial genes into
Amborella from other land plants (including three mosses).
2. Strasburgeria (Crossosomatales) (Fig. 1). The genus
comprises one species, Strasburgeria robusta, endemic to New
Caledonia. The affinities of this plant were formerly unclear
and it was treated as the monotypic Strasburgeriaceae, but it is
now known to be sister to another monotype, Ixerba of
northern North Island, New Zealand (Cameron, 2003). The
two species form a clade in Crossosomatales with Geissolomataceae (monotypic) of South Africa and Aphloiaceae
(monotypic) of south-eastern Africa, Madagascar, the Mascarenes and the Seychelles (Stevens, 2009). This group, based
around the Indian Ocean is sister to a neatly vicariant group
centred around the northern Pacific: Crossosomataceae, Stachyuraceae, Guamatelaceae and Staphyleaceae.
As well as the genetic sequences that link Strasburgeria and
Ixerba, Cameron (2003) also noted the strikingly similar wood,
the persistent sepals and the clawed petals of the two genera.
In addition, personal observations made on material in the
herbarium of the Institut de Recherche pour le Développement (IRD), Nouméa (January 2002), showed that the
gynoecia are similar. The style in both is truncate, persistent
and not clearly demarcated from the ovary, while the ovary in
both has five longitudinal flanges and is star-shaped in cross
section. The style of Strasburgeria is also sometimes more or
less twisted, as in Ixerba, although it shows no sign of the
deflexed apex seen in that genus. The gynoecium of the Cape
endemic Geissoloma (Geissolomataceae) is similar and shows
comparable torsion (Figure 3I in Dahlgren & Rao, 1969). In
both Strasburgeria and Ixerba the ovules are anatropous and
descending, with the micropyles upwards and outwards
(epitropous). Matthews & Endress (2005) also noted the
presence of an aril (at least in the ovule) and other features
which ally the two families, as well as features that link the two
with Geissoloma (e.g. similar unicellular, T-shaped perianth
hairs). There are differences between Strasburgeria and Ixerba;
for example, the fruit is indihiscent in the former and
dehiscent in the latter, and Strasburgeria has a very high
chromosome number, 2n = 500, compared with 2n = 50 in
Ixerba (Oginuma et al., 2006). Nevertheless, the two genera
can easily be accommodated in one family. Xeronemataceae
(Asparagales) is another plant family restricted to New
Caledonia and New Zealand.
Fossil pollen (Bluffopollis scabratus) from the Cenozoic of
New Caledonia, western and southern Australia, Tasmania and
New Zealand is ‘directly comparable’ to Strasburgeria pollen
(Cameron, 2003), although it is only half the size (Jarzen &
Pocknall, 1993). As Cameron (2003, p. 428) concluded, this
indicates that ‘the common ancestor of Strasburgeria and
Figure 1 Distribution of Crossosomatales (from Stevens, 2009). Cro, Crossosomataceae; Gua, Guamatelaceae. Fossil pollen (Bluffopollis
scabratus) attributed to Strasburgeria extends the range to western Australia, southern Australia, Tasmania and southern New Zealand.
Journal of Biogeography 37, 1239–1250
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M. Heads
Figure 2 Distribution of Oncothecaceae and Metteniusaceae; these form either a basal clade or the first and second basal branches in the
cosmopolitan clade Lamiales, Solanales and Gentianales (González & Rudall, 2007; González et al., 2007).
Ixerba most likely evolved prior to the break-up of Gondwana’.
Miocene leaf material from southern New Zealand shows a
‘good match’ with Strasburgeria (Pole, 2008), but was not
compared with Ixerba.
3. Oncothecaceae (Fig. 2). This family, with Oncotheca
balansae and Oncotheca humboldtiana in New Caledonia, and
the Metteniusaceae of Costa Rica to Ecuador, form the first
and second basal branches in the cosmopolitan clade Lamiales,
Solanales and Gentianales (Fig. 2; González & Rudall, 2007;
González et al., 2007). This topology indicates a trans-tropical
Pacific sequence of initial differentiation events in a global
ancestor: (i) New Caledonia versus the rest, and (ii) Colombia/
Panama versus the rest. However, the statistical support is
weak and the two groups may instead be sisters, giving a transtropical Pacific clade basal in a large group of asterids.
Although the pattern is seldom discussed, trans-tropical Pacific
affinities are common (e.g. Polygalaceae tribe Moutabeeae,
comprising Balgoya of New Caledonia, Eriandra of New
Guinea and the Solomon Islands and three other genera of
northern South America). The Pacific pattern in Oncotheca–
Metteniusa contrasts with the Indian Ocean (Gondwana)
affinity in Strasburgeriaceae and relatives.
The localities of Oncothecaeae and Metteniusaceae may
reflect the Cretaceous emplacement of large igneous plateaus
in the central Pacific and their subsequent dispersal to the west
(the Hikurangi plateau of New Zealand, the Ontong Java
plateau of the Solomon Islands, etc.) and to the east (plateaus
of Colombia and the Caribbean) (Heads, 2009a). Many of the
localities of Metteniusaceae occur west of the Romeral fault
zone (running through Cali and Medellı́n), the boundary
between autochthonous crust of the South American plate, to
the east, and accreted terranes, to the west.
4. Phellinaceae (Asterales). This family comprises the genus
Phelline, with 12 extant species in New Caledonia and records
of fossil leaf cuticle in Miocene strata of southern New Zealand
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Figure 3 Distribution of the Phelline complex: Phellinaceae,
Argophyllaceae and Alseuosmiacae. Distributions from Stevens
(2009) and Australia’s Virtual Herbarium (2009). P, fossil record
of Phelline in New Zealand (Pole, 2010).
(Pole, 2010) (Fig. 3). The family is sister to Argophyllaceae
(Argophyllum and Corokia), distributed in Queensland south
to the ‘McPherson–Macleay Overlap’ (Heads, 2009a), Lord
Howe Island, New Zealand, Chatham Islands and Rapa Island
(van Balgooy, 1966). The two families are sister to the
Alseuosmiaceae, of eastern Australia, New Guinea, New
Caledonia and New Zealand (van Balgooy, 1993; Tirel, 1996;
Kårehed, 2002).
Alseuosmiaceae and Argophyllaceae overlap in northeastern Queensland, New Caledonia and New Zealand, but
Alseusomiaceae has a ‘centre of gravity’ to the west of
Argophyllaceae, with records in western New Guinea and
south-eastern Australia, and in New Zealand it only occurs
west of the Alpine Fault (south to Mount Ellery, Jackson’s Bay;
pers. obs.). The partial allopatry of the two families probably
reflects initial vicariance. All three families of this Tasman/
Journal of Biogeography 37, 1239–1250
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Biogeography of New Caledonian plants
Figure 4 Distribution of Stylidiaceae sensu lato: Stylidium (including Oreostylidium) and Forstera (including Phyllachne), and their sistergroup Donatia. Distributions from Wagstaff & Wege (2002).
Coral Sea ‘Phelline complex’ occur together only in New
Caledonia and, including the fossil material, in New Zealand.
In contrast, the related Stylidiaceae sensu lato comprises
three genera, Stylidium, Forstera and Donatia, which all overlap
in Tasmania and New Zealand (Fig. 4). This group shows
significant allopatry with the Phelline complex as it is absent
from New Caledonia, central New Guinea and central Polynesia, but does occur in Southeast Asia, Tasmania, the far
south of New Zealand (Stewart Island and the sub-Antarctic
islands) and Patagonia, where the Phelline complex is
unknown.
The bulk of the Asterales comprises a clade with three
subclades: (Phelline complex (Stylidiaceae sensu lato (Asteraceae + Goodeniaceae + Calyceraceae + Menyanthaceae))).
The third subclade forms a very diverse, global group. (The
phylogeny as indicated is from Kårehed et al., 1999; Stevens,
2009, shows the three subclades as an unresolved trichotomy.)
Thus the phylogeny of the whole group involved a world-wide
ancestor that underwent: (i) a split between the Phelline
complex, centred on New Caledonia/New Zealand terranes,
and the remaining groups; (ii) a split between Stylidiaceae,
centred on Tasmania/South Island terranes, and the remaining
groups; (iii) subsequent regional overlap of the three clades;
and (iv) differentiation within the groups. The high diversity
of the Phelline complex in New Caledonia is correlated with
the anomalous low diversity of Asteraceae there (only eight
endemic species) and this pattern may be the direct result of
phylogeny rather than ecology.
Other New Caledonian taxa that were formerly treated as
endemic families include the following listed below.
5. Paracryphia. This genus comprises Paracryphia alticola
endemic to New Caledonia; there are also records of fossil
cuticle from the New Zealand Miocene (Pole, 2010) (Fig. 5).
Until recently Paracryphia was regarded as the only member of
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Figure 5 Distribution of Paracryphiales (grey) and Dipsacales
(grid pattern). Distribution data from Stevens (2009) and
Australia’s Virtual Herbarium (2009). In Dipsacales the distribution of the basal species in Viburnum is shown by an asterisk.
P, fossil record of Paracryphia in New Zealand (Pole, 2010).
Paracryphiaceae (Paracryphiales) but the family is now taken
to include Quintinia of New Zealand, New Caledonia, eastern
Australia, New Guinea and the Philippines. Sphenostemon of
New Caledonia, north-eastern Queensland, New Guinea and
Sulawesi is also closely related to, or part of, this clade (Bremer
et al., 2004; Winkworth et al., 2008; Stevens, 2009). This
south-west Pacific trio (Paracryphiales) is sister to the large
clade Dipsacales (Dipsacaceae, Caprifoliaceae, Adoxaceae,
Valerianaceae). Dipsacales are more or less cosmopolitan but
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M. Heads
are notably absent in the indigenous floras of New Zealand,
New Caledonia, the Bismarck Archipelago/Solomon Islands
and the central Pacific. As Winkworth et al. (2008) suggested, a
simple geographical split between the Paracryphiales and the
Dipsacales seems likely. This implies a break between an
eastern group centred around New Caledonia and a group
with its diversity further west, a similar pattern to that seen in
the break between the Phelline complex and the Stylidiaceae
(Figs 3 & 4). Early divisions within the Dipsacales also took
place in this general region; for example, Viburnum (160
species; Northern Hemisphere, South America, Asia) is one of
the largest genera in the order and has its basal species
(Viburnum clemensae) in Borneo (Winkworth & Donoghue,
2005). This does not mean that the species was ‘the first
diverging clade’ in the genus (Jacobs et al., 2008, p. 428) – it is
the same age as the clade it diverged from – but does indicate
that the first division within the genus occurred around here.
6. Canacomyrica. This monotypic New Caledonian endemic
genus has been treated in its own family (Doweld, 2000), but
most authors place it in Myricaceae (Macdonald, 1989;
Kubitzki, 1993; Carlquist, 2002). Recent molecular studies
have retrieved it as sister to the rest of the family (Herbert
et al., 2006). Canacomyrica is extant only in New Caledonia,
but Eocene to Miocene pollen from New Zealand has been
identified as belonging to this genus (Lee et al., 2001). The
only other member of the family, Myrica, is widespread
globally but is notably absent from New Caledonia, Australia,
eastern New Guinea, New Zealand and southern South
America.
7. The conifer Austrotaxus is another monotypic genus
endemic to New Caledonia and is sometimes treated in its own
family (Bobrov et al., 2004). Recent studies (Cheng et al.,
2000; Price, 2003) showed it to be sister of the other Taxaceae
sensu stricto, comprising Pseudotaxus (China) and Taxus
(north temperate zone, south to the Philippines, Sulawesi
and Mexico).
To summarize, the New Caledonian endemics cited here are
either sister to widespread Northern Hemisphere or even
global groups (Amborella, Canacomyrica, Austrotaxus), or form
parts of Indian Ocean (Strasburgeria), Pacific Ocean (Oncotheca) or Tasman/Coral Sea (Phelline, Paracryphia) groups that
are sister to widespread Northern Hemisphere or global
groups.
Amborella, the basal angiosperm, shows a ‘globally basal’
pattern. A similar case occurs in the orchid Pachyplectron,
endemic to New Caledonia and basal in the virtually cosmopolitan subtribe Goodyerinae (Cribb et al., 2003). This does
not mean that Amborella and Pachyplectron are the same age;
the two genera are distributed on different terranes on Grande
Terre (Aubréville et al., 1967–present; Heads, 2008a,b) and the
island is a geological and biological composite.
A basal group is just a small sister-group and so its location
does not indicate a centre of origin. The fact that the New
Caledonia endemics have diverse sister-groups that are widespread globally but not on New Caledonia does not mean, as
might be inferred, that New Caledonia is a centre of origin
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from which the widespread groups have migrated. But sistergroup relationships do indicate that the endemics did not
migrate recently to New Caledonia. These patterns also explain
why so many familiar groups that are more or less cosmopolitan are strangely absent in New Caledonia – they are
represented there by other, locally endemic taxa.
The direct relationship of many New Caledonian taxa with
trans-tropical Pacific groups could be due to some New
Caledonian terranes having originated in the central Pacific
and subsequently colliding with New Caledonia. The Bismarck
Archipelago is a similar case. Its biota is quite different from
that of nearby mainland New Guinea, although the two land
masses are now almost joined and probably will be in the near
geological future. The biological differences can be attributed
to the history of the Bismarck Archipelago basement, which
formed thousands of kilometres away from mainland New
Guinea in the central Pacific.
THE NEW CALEDONIAN PALMS
Palms are a conspicuous, well-studied group and the New
Caledonian members are especially diverse, with 10 genera
represented. The group is thus a good example for showing
differentiation around New Caledonia in the south-west
Pacific region. The molecular phylogeny presented by Baker
et al. (2009, their Fig. 2) is followed here, along with the
classification of Pintaud & Baker (2008).
The fan palm Pritchardiopsis (subfamily Coryphoideae;
Fig. 6a) is perhaps the rarest palm in the world, with just a
few individual plants in southern Grande Terre (the three
other lineages of New Caledonian palms, discussed below, are
all widespread in Grande Terre). Pritchardiopsis is sister
to Johannesteijsmannia and Pholidocarpus, both in western
Malesia.
The other New Caledonian palms make up a remarkable
assemblage of nine genera (eight endemic) in the tribe Areceae
(subfamily Arecoideae). Areceae range from Pemba Island,
50 km off mainland Tanzania, to the Pacific islands (Samoa).
The distinctive western limit so close to the African mainland
is seen in many widespread Indo-Pacific taxa (plant, bird and
mammal examples are cited in Heads, 2006). The tribe Areceae
comprises an ‘Indian Ocean clade’ which extends east to the
Solomon Islands and Fiji but is not in New Caledonia, and a
‘Western Pacific’ clade which is very diverse in New Caledonia
and other Pacific islands, and extends west to eastern Australia,
Indonesia and Sri Lanka (Norup et al., 2006). The Indian
Ocean and the Western Pacific clades are mainly vicariant,
with secondary overlap around the margin of the Pacific plate.
The basal group in the Western Pacific clade is Loxococcus of
Sri Lanka, but this does not require the eastward dispersal that
Norup et al. (2006) proposed, only an eastward sequence of
differentiation. The New Caledonian genera have ‘invaded’ the
region by evolving there, not as the result of nuts being blown
or carried there from Sri Lanka.
The New Caledonian Areceae belong to three separate
groups of the Western Pacific clade. (Baker et al., 2009): (1)
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Biogeography of New Caledonian plants
(a)
(b)
(c)
(d)
Figure 6 Affinities of the New Caledonian palms. (a) Distribution of the Pritchardiopsis clade. (b) Distribution of the Clinosperma clade
(Clinosperma includes Lavoixia and Brongniartikentia, Cyphokentia includes Moratia). (c) Distribution of the Archontophoenix clade
(Kentiopsis includes Mackeea). (d) Distribution of the Basselinia clade (Basselinia includes Alloschmidia). Taxonomy from Pintaud & Baker
(2008), distributions from Uhl & Dransfield (1987), Dowe (1989), Dowe et al. (1996), Fuller (1999) and Pintaud et al. (2001).
the Clinosperma clade is in New Caledonia, Vanuatu/Fiji and
the southern Ryukyu Islands near Taiwan (Fig. 6b); (2) the
Archontophoenix clade is in New Caledonia, eastern Australia,
New Guinea, the Bismarck and Louisiade Archipelagos and
Bougainville Island (northern Solomon Islands) (Fig. 6c); and
(3) the Basselinia clade is in New Caledonia, Lord Howe and
Norfolk Islands, New Zealand, Vanuatu, Fiji, the Solomon
Islands and New Britain (Fig. 6d). Thus the four clades of
palms in New Caledonia have relatives in different places:
western Malesia (Pritchardiopsis clade), Ryukyu Islands (Clinosperma clade), western Tasman/Coral Sea (Archontophoenix
clade) and eastern Tasman/Coral Sea (Basselinia clade). The
simplest explanation for this allopatry is an origin of the
groups by vicariance, followed by overlap caused by tectonic
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juxtaposition of terranes in New Caledonia, local range
expansion, or both.
This process of vicariance is reflected in aspects of the
morphology. Norup et al. (2006) stressed that several genera in
Areceae have no unique characters but are distinguished
instead by different ‘combinations of widespread character
states’. This phenomenon is common in all groups – Reinert
et al. (2006) described it in mosquito genera, to cite just one
detailed study – although it is seldom acknowledged as a
general principle. This may be because it is incompatible with
the idea of hierarchical phylogeny and the idea that taxa are
defined by ‘uniquely derived characters’. Recombination of
characters would not be expected if every new mutation, form
or taxon arose at a centre of origin and spread from there by
1245
M. Heads
physical movement. It would be expected if vicariant groups
evolved as recombinations of widespread ancestral characters
already distributed over different, broad sectors of the ancestral
range before the evolution of the modern groups (cf. Heads,
2009c). It is well known that parallel evolution is ubiquitous
and recombination of characters probably takes place by
parallel evolution in the populations of a given biogeographical
sector.
In the New Caledonian palms, the distributions of the
Pritchardiopsis and Clinosperma clades are notable for their
large disjunctions to the north-west. Another palm, Clinostigma, has a similar pattern: Samoa/Fiji, Vanuatu, Solomon
Islands, New Ireland, Caroline and Bonin islands, skirting
mainland New Guinea to the north. Uhl & Dransfield (1987,
p. 437) described this as a ‘great arc…a most interesting
distribution’. In dispersal interpretations, the arcs would be
regarded as routes of migration, perhaps along ‘stepping-stone’
islands. But the arcs are interpreted here instead as allopatric
sectors and fragments of formerly widespread ancestral ranges,
with central Pacific taxa having survived as metapopulations
on ephemeral volcanic islands. Pritchardiopsis, for example, is
represented in neighbouring Vanuatu by Licuala, in eastern
Australia by Livistona, and in the central Pacific by Pritchardia
(Fig. 7). Pritchardia has its sister-group, Copernicia, in Cuba
(20 species), Hispaniola (two species) and South America
(three species). These genera belong to tribe Trachycarpeae (18
genera) which is itself largely allopatric with Phoeniceae (the
date palms), its sister-group. Thus the evolution of the two
tribes has involved the allopatric break-up of a more or less
global ancestor into Phoeniceae, based around the Atlantic and
Indian oceans, and Trachycarpeae based around the Pacific,
with subsequent secondary overlap along the Tethys sector.
Within the tribes, differentiation of the genera (cf. Pritchardia,
Pritchardiopsis, etc.) has also been allopatric. The central
Pacific representative, Pritchardia, indicates evolution on and
around the large igneous plateaus emplaced in the Cretaceous.
These include many flat-topped seamounts that originally
formed high islands. Other Cretaceous volcanics that Pritchardia may have occupied include the Line Islands, an archipelago
4000 km long (between south-eastern Polynesia and Hawaii)
in which all the islands are now reduced to atolls (Heads,
2009a).
In Australia, Crisp et al. (2010) tested whether Livistona was
a Gondwana relict or a Miocene immigrant and relied on
fossil-calibrated dates. These are minimum ages and can only
eliminate later events as relevant, not earlier ones. Despite this,
the minimum dates were illogically transmogrified into
maximum ages, used to rule out early vicariance, and then
cited as support for a centre of origin/dispersal model. This is
not accepted here.
The pattern shown in palms, with different New Caledonian
clades having sister-groups in different areas, also occurs in
other groups. New Caledonian Diospyros (Ebenaceae) provides
a good example (Duangjai et al., 2009).
One clade (Diospyros balansae, etc.) has its sister in eastern
Australia (Cairns to Sydney), indicating a Cretaceous split with
the opening of the Tasman Sea. This clade is basal to a global
group (the entire genus except for Diospyros maingayi and
Diospyros puncticulosa). A second clade (Diospyros calciphila,
etc.) has its sister in the Hawaiian islands, indicating evolution
around the central Pacific igneous plateaus. A third clade
comprises Diospyros fasciculosa of New Caledonia and its sister
Diospyros ehretioides of Southeast Asia (cf. the affinities of the
fan palm Pritchardiopsis), also Australia, Vanuatu and Fiji. The
fourth clade (Diospyros olen) has its sister in the Solomon
Islands, Vanuatu, Fiji, Tonga and Samoa. This is explained by
terranes accreting in New Caledonia from the north-east
(Heads, 2008a).
The largely allopatric distribution of the four clades can be
explained by original vicariance of a widespread ancestor
followed by juxtaposition in New Caledonia of the four clades
with terrane accretion. The only evidence for local range
Figure 7 Distribution of the Trachycarpeae and Phoeniceae (Arecaceae). Line ‘a’ indicates the affinity of Pritchardiopsis with western
Malesian genera. Line ‘b’ indicates the affinity of Pritchardia with Copernicia of Cuba, Hispaniola and South America. Areas with fossil
records only. Distributions from Uhl & Dransfield (1987).
1246
Journal of Biogeography 37, 1239–1250
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Biogeography of New Caledonian plants
expansion is in D. ehretioides and, in all four clades, within
New Caledonia itself. The overall process is one of juxtaposition rather than radiation.
EVOLUTIONARY ECOLOGY OF THE
TRACHYCARPEAE
In traditional analyses geographical centres of origin are
assumed and their location is calculated with reference to the
location of centres of diversity, ‘basal’ clades and grades,
minimizing the number of founder dispersal events, and so on.
In the same way, an ecological centre of origin or ancestral
niche of a group is often estimated and the group is proposed
to have spread from there into other niches. In contrast,
vicariance analysis assumes that the ancestor of a group was
already widespread and so may have already occupied a wide
ecological range before the modern groups existed. In the palm
tribe Trachycarpeae, the New Caledonian Pritchardiopsis
occurs on soils derived from ultramafic rock. Other members
include the Central American Brahea and the Asian Maxburretia, Guihaia, Rhapis and Trachycarpus, all on limestone
and karst (the ecological data cited here for palms are from Uhl
& Dransfield, 1987). The close physiological, ecological and
tectonic relationship between limestone and ultramafic floras
is examined in more detail elsewhere (Heads, 2008a). Both
rock types characterize belts of subduction and accretion,
where taxa of uplifted coral reefs are pre-adapted for life on
obducted ultramafic terranes. The ecological ranges of the
other genera of Trachycarpeae are as follows.
1. Pritchardia: rain forest (Hawaii), uplifted coral reef near sea
(other Pacific Islands). In Fiji, plants occur only on limestone
islets and limestone lagoon cliffs, where they grow right to the
edge of the sea and are subject to salt spray (Heads, 2006).
2. Johannesteijsmannia: undisturbed, primary rain forest;
kerangas heath forest (Johannesteijsmannia altifrons).
3. Pholidocarpus: freshwater and peat-swamp forest.
4. Licuala: mostly rain forest; Licuala calciphila is a calcicole,
Licuala spinosa occupies the landward fringe of mangrove,
Licuala paludosa is in peat swamp forest.
5. Livistona: rain forest, freshwater and peat swamp forest,
montane forest, woodland, savanna, permanent water bodies
in desert.
6. Acoeloraphe: coastal brackish swamps.
7. Serenoa (south-eastern USA): pinelands, prairies, coastal
sand-dunes, often forming dense swards; the architecture is
unusual: ‘stem subterranean and prostrate or surface creeping’.
Similar plagiotropic axes occur in many mangroves, for
example the mangrove palm Nypa, with ‘prostrate or subterranean’ stems.
8. Rhaphidophyllum: low, moist to wet areas with rich humus,
calcareous clay or sandy soils in woods or swamps. In grottos,
limestone sinks, shaded pinelands, but usually associated with
limestone.
9. Copernicia: savanna, woodland; Copernicia prunifera forms
vast stands in South America in areas subject to annual
flooding.
Journal of Biogeography 37, 1239–1250
ª 2010 Blackwell Publishing Ltd
10. Colpothrinax: open, sandy country, open forest on ridges.
11. Washingtonia: by springs and streams in deserts.
12. Chamaerops (western Mediterranean): sandy or rocky
ground usually near the sea.
The sister tribe of Trachycarpeae is Phoeniceae, with only
one genus, Phoenix (including the date palm). Species occur
around oases and watercourses in semi-arid areas with a few in
tropical monsoon areas. Phoenix paludosa is in perhumid parts
of Asia where it is confined to the landward fringe of the
mangrove.
To summarize, Trachycarpeae/Phoeniceae occupy a series of
habitats that is typical of tropical groups: back-mangrove,
limestone/ultramafic sites, monsoon forest, savanna, swamp
forest, rain forest, montane forest. This may have resulted from
a weedy group of back-mangrove and associated habitats
spreading through inland areas with the advance of Cretaceous
seas, later becoming stranded there with the retreat of the seas
and onset of Cenozoic orogeny (cf. Heads, 2006). It is often
suggested that the biogeography of a group is determined by its
ecology, but in the model proposed here a group’s current
ecology is determined by its location. Populations derived
from a widespread Trachycarpeae/Phoeniceae ancestor have
ended up being stranded in areas that became desert in Arabia,
flood plains in South America, atolls in the Pacific, and rain
forests in Southeast Asia.
CONCLUSIONS
New Caledonian terranes derived from the west (Gondwana,
i.e. Australia) include the ‘basement’ terranes. Those derived
from the east (Pacific) include the Loyalty Ridge, formerly an
active island arc. This history is reflected in the composite
nature of the New Caledonian biota, which shows connections
with areas to the west (Figs 1, 5 & 6c) and to the east (Figs 2 &
6d). Affinities with taxa to the north-west (Figs 6a,b & 7) may
reflect evolution associated with the former Tethys basins.
The old Pacific plate/Indian plate margin ran through
New Zealand, New Caledonia and New Guinea, and so this
arc represents both a tectonic and a biogeographical edge of
intercontinental significance.
In the dispersal paradigm, the standard questions asked for
any group in an area are: ‘Where is its centre of origin?’
and ‘Which route and which means of dispersal did the
group use to disperse to the area?’ For example, Keast (1996)
proposed the ‘fundamental question’: ‘When, and how,
did the archaic southern elements reach New Caledonia?’.
It is suggested here that neither the ‘archaic’ nor any
other indigenous elements ever ‘reached’ New Caledonia
using ‘means of dispersal’. They evolved there, or rather on
arc and intraplate islands associated with the different
component terranes. The groups differentiated out of already
widespread ancestors before the terranes were assembled as
modern New Caledonia in the Jurassic and the Eocene. The
different global patterns involving New Caledonian taxa
reflect areas around which differentiation took place, not
centres of origin.
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M. Heads
ACKNOWLEDGEMENTS
I am grateful to T. Jaffré for permission to examine material in
the IRD Herbarium, Nouméa.
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BIOSKETCH
Michael Heads has taught ecology and systematics at
universities in Papua New Guinea, Zimbabwe, Ghana and
Fiji. His main research interests are in tree architecture,
biogeography and the evolution of rain forest plants and
animals.
Editor: Jon Sadler
Journal of Biogeography 37, 1239–1250
ª 2010 Blackwell Publishing Ltd