Phil. Trans. R. Soc.

B (2008) 363, 3309–3317

doi:10.1098/rstb.2008.0122
Published online 2 September 2008

Review

New Caledonia: a very old Darwinian island?

Philippe Grandcolas1,*, Jérôme Murienne2, Tony Robillard1,
Laure Desutter-Grandcolas1, Hervé Jourdan3, Eric Guilbert1
and Louis Deharveng1
1
UMR 5202 CNRS, Département Systématique et Evolution, Muséum national d’Histoire naturelle,
45 rue Buffon, 75005 Paris, France
2
Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology,
Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA
3
Centre de Biologie et de Gestion des Populations, INRA, and Laboratoire de Zoologie Appliquée,
IRD, 98948 Nouméa, Nouvelle-Calédonie
New Caledonia has generally been considered a continental island, the biota of which largely dates
back to Gondwanan times owing to its geological origin and the presence of phylogenetic relicts. This
view is contradicted by geological evidence indicating long Palaeocene and Eocene submersions and
by recent biogeographic and phylogenetic studies, with molecular or geophysical dating placing the
biota no older than the Oligocene. Phylogenetic relicts do not provide conclusive information in this
respect, as their presence cannot be explained by simple hypotheses but requires assumption of many
ad hoc extinction events. The implication of this new scenario is that all the New Caledonian biota
colonized the island since 37 Ma Local richness can be explained by local radiation and adaptation
after colonization but also by many dispersal events, often repeated within the same groups of
organisms. Local microendemism is another remarkable feature of the biota. It seems to be related to
recent speciation mediated by climate, orography, soil type and perhaps unbalanced biotic
interactions created by colonization disharmonies. New Caledonia must be considered as a very
old Darwinian island, a concept that offers many more fascinating opportunities of study.
Keywords: New Caledonia; biogeography; phylogenetics; endemism; dispersal; adaptation

1. INTRODUCTION effects of invasive species (Gargominy et al. 1996;

New Caledonia is a large and megadiverse tropical Jourdan et al. 2001; Keith 2005; Beauvais et al. 2006;
island in the southwest Pacific, with distinctive Pascal et al. 2008). However, an alternative view,
characteristics that make it a remarkable natural presented earlier but gaining far less traction, empha-
laboratory of evolution. Owing to its geological sized the absence of certain groups such as mammals
continental origin and the presence of apparent and continental beetles and of clear geological evidence
phylogenetic relicts, New Caledonia has long been for Palaeocene marine transgression, and suggested
considered a Gondwanan refuge where archaic groups that the biota was much more recent ( Jeannel 1942;
have survived for 80 Ma (Holloway 1979; Morat Faivre et al. 1955; Darlington 1957).
1993a). Reflecting this, the amazing New Caledonian Convincing geological evidence now suggests that
species richness has been explained by local, long-term the second hypothesis is closer to the truth. New
cladogenesis rather than rapid speciation after recent Caledonia’s biodiversity is not that of a continental
island colonization (e.g. Morat 1993b). This Gond- island that has retained many ancient groups since its
wanan view became widespread during the past few separation from the northeastern margin of Australia,
decades (Lowry 1998; Lowry et al. 2005; Murienne ca 80 Ma, but an oceanic island with a composite biota
et al. 2005) and has often been invoked as a reason to dominated by neoendemism and disharmonic coloni-
study the diverse New Caledonian biota (e.g. Mittermeier zation, a ‘Darwinian’ island (Gillespie & Roderick 2002).
et al. 1996; Pagel 2003). New Caledonia was also The question now for biologists is not so much whether
characterized as a biodiversity hot spot owing to its high the biota is Gondwanan and ancient but when and in
species richness and level of endemism, and the what manner the modern biota assembled. Such
conservation issues raised by nickel mining, anthro- questions can be addressed by modern phylogenetic
pogenic burning and forest logging (Bouchet et al. approaches in the context of an accurate geological
1998; Myers et al. 2000), as well as the deleterious framework ( Trewick et al. 2007). This is now possible
for New Caledonia since the island has been the subject
* Author for correspondence ( pg@mnhn.fr). of increasing molecular phylogenetic and geological
One contribution of 15 to a Theme Issue ‘Evolution on Pacific studies. We first briefly review the geological evidence
islands: Darwin’s legacy’. that the biota is relatively young and then turn to the

3309 This journal is q 2008 The Royal Society

3310 P. Grandcolas et al. Review. New Caledonia as a Darwinian island

(a) type of rock indicating deep submersion as its structure

is shaped by high pressure (Paris 1981a,b; Aitchison
et al. 1998; Pelletier 2006). During the Eocene, the
continental crust in the New Caledonia area was
tectonically active, being in collision with the Loyalty
Islands arc, and obduction at this time placed a layer of
oceanic crust (lithosphere) over the submerged con-
tinental crust. New Caledonia emerged during an
Oligocene lithosphere extension phase, uplifting with a
cover of lithospheric ultramafic rocks (Paris 1981a,b;
Aitchison et al. 1995; Cluzel et al. 2001; Crawford et al.
2003; Pelletier 2006; Schellart et al. 2006). The present-
(b) day mountains are relatively old since they are the
product of complex orogenesis from Oligocene time, as
indicated by several series of lateritic beds ranging from
sea level to mountain tops (Chevillotte et al. 2006). In
the period prior to subaerial emergence of New
Caledonia, other islands might have existed on the
Norfolk or the Loyalty Ridges but without any
relationships or continuity with New Caledonia (Paris
1981a,b; Herzer et al. 1997; Meffre et al. 2006). Smaller
islands such as Norfolk or the Loyalty Islands uplifted
and emerged much later, 3.7 and 2 Ma as a result of
> 1000 m volcanism and lithosphere flexure, respectively (Dubois
500 – 1000 m et al. 1974).
These successive geological events have had import-
200 – 500 m ant consequences for the evolution of the biota. First,
0 – 200 m more than 20 Ma of total submersion from the
Palaeocene to the Eocene make it difficult to retain
Figure 1. (a) Distribution of ultramafic rocks (shaded areas) the notion that a Gondwanan biota has survived locally
in New Caledonia. (b) Orography in New Caledonia showing (Murienne et al. 2005; Murienne 2006; Pelletier
several chains of mountains, peaking at more than 1600 m in 2006). Even if this biota persisted on emerged lands
the north and south. in the region, the occurrence of which is speculative,
phylogenetic patterns recently deciphered in an effort they had to disperse back to New Caledonia in the
to answer these questions. Oligocene. Second, ultramafic rocks obducted in the
Eocene onto most of the New Caledonian mainland
have given rise to an extensive area of metalliferous
2. GEOLOGICAL HISTORY AND EVOLUTION soils. Though subjected to several erosion cycles, they
OF THE BIOTA remain more extensive in the south. Being poor in
Oriented northwest to southeast roughly between nutrients and rich in metals (mainly nickel and copper),
latitudes 20 and 228 S, the island is 1220 km from they are highly stressing substrates for many organisms
Australia, 1700 km from New Zealand and approxi- and could have strongly constrained biotic evolution.
mately 400 km from the islands of Vanuatu. It is Third, there is no evidence for direct exchange with
16 890 km2 in area, with an elongate shape 500 km New Zealand but only the possibility of stepping-stone
long and 50 km wide. Mountain ranges are complex dispersal after the Oligocene emersion, since part of the
and dissected by many rivers, with tablelands and peaks Norfolk Ridge was deep below sea level, deeper than
reaching elevations of more than 1600 m (figure 1). the extent of major sea-level fluctuations. The
Most of the island is covered by wet evergreen forests terrestrial biota on neighbouring islands (Norfolk and
with anthropogenic savannahs dominating at low the Loyalty Islands) is even more recent. Even though
elevations. A few small fragments of sclerophyll dry local volcanism has produced palaeo-islands there, reef
forest remain on the western coast and shrubby structures and underlying layers indicate an unam-
vegetation (‘maquis minier’) occurs on metalliferous biguous period of total submersion before their recent
soils, mostly in the south. emersion (Dubois et al. 1974; Paris 1981a,b).
New Caledonia formed from part of a continental Owing to these three geological constraints, New
fragment that began separating from Australia ca 83 Ma Caledonia is a remarkable palaeogeographic model as it
as the Tasman Sea began to form (Brothers & Lillie presents a combination of continental and oceanic
1988; Neall & Trewick 2008). The New Caledonian features. In spite of a Gondwanan origin, it has under-
area was subsequently subject to a series of dramatic gone many recent tectonic events. Its elongated shape on
geological events (Paris 1981a,b). In the Palaeocene, the Norfolk Ridge made it roughly parallel to the
the part of Zealandia that subsequently became subduction/obduction fronts that dramatically affected
New Caledonia experienced a lengthy submersion in it, precluding a situation in which part of the island
deep water, as evidenced by marine sedimentation remained above the ocean surface while the rest was
(Globigerina limestone) and the formation of fine- submerged. This strongly constrains biogeographic
grained black chert (‘phtanites’ of French authors), a hypotheses, arguing for Oligocene recolonization after

Phil. Trans. R. Soc. B (2008)

 Review. New Caledonia as a Darwinian island P. Grandcolas et al. 3311

the very long Palaeocene and Eocene submersions. After Howe Island and New Zealand ( Trewick et al.
emersion, its isolated position between two deep oceanic in press). The New Caledonian sandalwood, Santalum
basins calls for rather simple explanations of its current austrocaledonicum, also differentiated recently, in this case
biological diversity in terms of dispersal. In this geological from Vanuatu species 1–1.5 Ma (Harbaugh & Baldwin
context, the biota could be old, even though resulting 2007). Other cases are different since the sister-
from recolonization, and could have been shaped by group relationships involve large and remote
orogenesis and extensive metalliferous soils over 37 Ma, regions such as Indo-Malaysia (eneopterine crickets:
a far longer time than on many other oceanic islands. Desutter-Grandcolas & Robillard 2006; Robillard &
Keeping in mind this remarkable geological framework, Desutter-Grandcolas 2006), Australia and Africa
we next examine phylogenetic evidence separately to (Proteaceae: Barker et al. 2007), or are poorly resolved
maintain logical independence between the two sources (Araliaceae: Eibl et al. 2001), which leaves the dating
of evidence and to avoid circular reasoning, following the question more open. For the Proteaceae, the dating
careful argument of Waters & Craw (2006). (43–25 Ma) does not concur with a scenario of
Gondwanan vicariance, which is otherwise supported
by the tree topology (Barker et al. 2007). The New
3. ANCIENT RADIATIONS OR REPEATED Caledonian freshwater shrimp genus Paratya (Page et al.
DISPERSAL? 2005) and galaxiid fishes (Waters et al. 2000), sisters
Answering this classical question is fundamental to respectively to an Australian and a New Zealand group,
understanding the evolution of biodiversity in New are dated as younger than 12 and 9 Ma, respectively.
Caledonia and makes it possible to distinguish regional The freshwater galaxiids, supposedly unable to disperse
endemism (groups restricted to the New Caledonia over the sea, were often considered a relict taxon, even
mainland as a whole) from local endemism (groups though the occurrence of marine larvae is pervasive
restricted to certain locations in New Caledonia), an in this group (Waters et al. 2000).
often confused issue for this island (Murienne 2006). Infrequent and distinct colonization events have been
Most studies citing the high rates of endemism (often inferred in diving beetles (Balke et al. 2004, 2007a,b)
close to 100%) in many groups of New Caledonian and cockroaches (Murienne 2006; Murienne et al.
organisms actually refer to regional endemism in press a). In both, the occurrence of a few distinct
examined in the context of large-scale phylogenetic clades in New Caledonia is an argument in favour of
studies (e.g. Morat et al. 1986; Chazeau 1993; Lowry dispersal, at least for explaining the origin of one of the
1998). Such studies often reveal that within certain clades. In these cases, molecular dating indicates recent
New Caledonian groups, multiple species are nested origins (14 and 9 Ma or 8.3 Ma, respectively).
within larger clades with taxa from Australia, New We have not yet examined the case of the most
Zealand or New Guinea, calling for explanations in remarkable supposedly relict groups, which are often
terms of recent dispersal. If, conversely, large New referred to when arguing for a Gondwanan origin of the
Caledonian clades are sister groups to taxa from other New Caledonian biota. New Caledonia harbours some
regions, dating the sister-group dichotomy is the only taxa that are deeply embedded in the phylogenies of a
way to assess whether dispersal is again the explanation number of large groups (but also lacks others such as
or if a vicariance hypothesis can be supported, even in Onychophora). The most famous is the endemic
the face of strong geological evidence to the contrary. Amborella trichopoda, the sole member of the Ambor-
Dating can be obtained from a molecular hypothesis ellaceae and the sister group of all other flowering
with careful calibration of substitution rates or a plants (Mathews & Donoghue 1999; Parkinson et al.
geological hypothesis based on some physical measure- 1999; Qiu et al. 1999; Soltis et al. 1999). Another
ments (e.g. K–Ar dating for some islands). remarkable example is the emblematic flightless bird,
The clades that have been studied so far, however, do the kagu (Rhynochetos jubatus), the closest relatives of
not show very clear-cut results in this respect. In several which occur in New Zealand and South America
groups, there is evidence for multiple nested relation- (Cracraft 2000; Fain & Houde 2004). A further
ships involving taxa from New Caledonia and other example is the New Caledonian subfamily Troglosir-
regions, e.g. Nothofagus (Fagaceae; Swenson et al. 2001; onidae (Opiliones), which is the sister group of taxa
Cook & Crisp 2005), Sapotaceae (Bartish et al. 2005) from South America and West Africa and only distantly
and Meryta (Araliaceae; Tronchet et al. 2005). Bartish related to Australian and the New Zealand taxa (Boyer
et al. (2005) dated the oldest New Caledonian species in et al. 2007). According to molecular dating, these
their study to 32.4 Ma with a molecular clock, corre- Opiliones diversified recently in New Caledonia
sponding to the Oligocene–Miocene transition. (28–49 Ma) even though the divergence from South
Other groups show a single origin of their New America and West Africa is much older (124–221 Ma),
Caledonian species. However, ages were not always the contrast between the two dates suggesting some
evaluated by molecular dating in these studies but the extinction (Boyer et al. 2007). These deeply rooted and
geographical patterns implied are nonetheless very therefore relatively old groups, occurring in distant
different from one another. In Araucaria (Araucariaceae), parts of the world, are frequently considered as relicts
the New Caledonian radiation is sister to the species and used to support a Gondwanan origin for the biota
endemic to the much more recent Norfolk Island of New Caledonia. Their presence in New Caledonia—
(Setoguchi et al. 1998). Implicitly, this New Caledonian considered as a geologically old land—is assumed to
radiation also has a recent origin (3.7 Ma). The same result from survival over 80 Ma. Contrary to this
applies for Placostylus land snails, the sister group common reasoning, we submit that these groups do not
of which involves both the recently emerged Lord provide much biogeographic and temporal information

Phil. Trans. R. Soc. B (2008)

3312 P. Grandcolas et al. Review. New Caledonia as a Darwinian island

geological events regional endemism local endemism

emersion of the Loyalty Islands sandalwoods Angustonicus (Murienne et al. 2005)

(Harbaugh & Baldwin 2007) Araucaria (Setoguchi et al.1998)
emersion of the Norfolk Island Lauraesilpha (Murienne 2006)
galaxiid (Waters et al. 2000)
Paratya (Page et al. 2005)
diving beetles (Balke et al. 2007a, b)

20

geckos (Bauer et al. 2006)

Sapotaceae (Bartish et al. 2005)
emersion of New Caledonia
37

obduction 40 Tasmantis Scincidae (Smith et al. 2007)

(submersion) Proteaceae (Barker et al. 2007)

60
fine-grained black chert
(submersion)

separation from Australia

80

Ma
Figure 2. A time scale for the major geological events including the emersion of the New Caledonia mainland, 37 Ma ago
(horizontal dotted line), and estimated ages (horizontal bars) for New Caledonian clades according to the studies cited
(confidence intervals are shown if provided by the authors). Smith et al. (2007) provided a confidence interval for the age of
scincid lizards of the whole of Tasmantis, indicated here with a vertical dotted line (the age of the New Caledonian clade itself
was not provided but is necessarily more recent). Note that all these ages inferred independently from geology have been found
to post-date the emersion of the island.

since their relatives are either absent from the region 4. LOCAL RADIATION AND VACANT NICHES
around New Caledonia or have a worldwide distri- The question of radiation is often considered in relation
bution. Their long time survival as relicts in New to adaptation, and this is especially meaningful in the
Caledonia is an indirect assumption requiring further context of islands, where disharmony in colonization
assumption of many extinction events in neighbouring offers evolutionary opportunities for groups to diversify
regions such as Australia or New Zealand (Grandcolas in ‘vacant’ niches (Losos et al. 1998; Gillespie 2004;
et al. in press). Gillespie et al. 2008). Undoubtedly, some of the New
None of these studies provides clear evidence for old Caledonian radiations have such adaptive back-
local diversification since most dates inferred from grounds. The most remarkable examples are the
molecular phylogenies do not pre-date the Oligocene monophyletic scincid and geckonid lizard radiations,
(figure 2). Several emblematic groups such as Araucaria the ecological diversity of which is unparalleled. These
and Nothofagus have even undergone more recent groups exhibit many remarkable foraging behaviours
radiations or colonizations of New Caledonia. Also, and morphologies (e.g. minute and giant species) on an
there is ample evidence for the occurrence of repeated island where other small vertebrates are scarce (Bauer &
dispersal in many groups. Thus, there is no strong, Sadlier 2000; Bauer et al. 2006; Smith et al. 2007).
unambiguous evidence for very old local Gondwanan Another example is the radiation of the cricket genus
radiations. Rather, many old Gondwanan groups are Agnotecous ( Eneopterinae), which includes many
represented in New Caledonian by species of quite recent species with peculiar stridulatory apparatuses that emit
origin. As a whole, phylogenetic patterns and especially songs in unusually high frequencies and with harmonic
the dating are consistent with the geological framework shifts (Robillard & Desutter-Grandcolas 2004; Robillard
presented above. Phylogenetic relicts remain puzzlingly et al. 2007). The cockroach subfamily Tryonicinae
enigmatic and their presence cannot be explained in a (Blattidae) includes the speciose genus Lauraesilpha,
simple way. the members of which exhibit a unique combination of

Phil. Trans. R. Soc. B (2008)

 Review. New Caledonia as a Darwinian island P. Grandcolas et al. 3313

behavioural traits, wood eating, intestinal ciliates and in plants (Eibl et al. 2001; Swenson et al. 2001, 2007;
solitary habits (Grandcolas 1997; Murienne 2006; Bartish et al. 2005; Bottin et al. 2005; Tronchet et al.
Murienne et al. in press a, submitted). These charac- 2005), insects (Desutter-Grandcolas & Robillard 2006;
teristics recall those of panesthiine cockroaches (Blaber- Murienne et al. 2005, in press a), land snails ( Trewick
idae), which include many wood-eating species in et al. in press) and lizards (Bauer & Sadlier 2000; Bauer
Australasia and throughout the southwest Pacific et al. 2006; Smith et al. 2007), even though some cases
(including Vanuatu and Lord Howe Islands), but with of sympatry and even syntopy have also been docu-
the exception of New Caledonia (Roth 1991). The mented in diving beetles (Balke et al. 2004, 2007b).
springtails of the genus Caledonimeria are among the Most species in these various groups are restricted to
largest species of Collembola (up to 8 mm), perhaps a single mountain or group of mountains, sometimes in
filling the niche of some other locally absent soil sympatry with species from more distantly related
arthropods (D’Haese 2003). groups in the same clade. If the molecular substitution
Concerning adaptation and vacant niches, the rates used by the authors of these studies are correctly
expanding invasion by the little fire ant (Wasmannia calibrated, microendemism is a recent feature dating
auropunctata) is a sad natural experiment that demon- back only 2 or 3 Ma. This time frame is consistent with
strates how the structure of native communities can the most commonly accepted scenario in such studies
evolve in a peculiar manner and offer evolutionary that microendemic species have established through
opportunities to colonizers on islands. Following its repeated vicariance events on neighbouring mountains
recent anthropogenic introduction, the little fire ant has subsequent to Quaternary climatic changes. The
colonized New Caledonian communities in which the classical succession of dry/cold and hot/wet periods
local Pheidole ant species are unable to compete, unlike probably promoted allopatric speciation after range
other Pheidole species in the native South American fragmentation as the altitudinal ranges of species on
communities of W. auropunctata (Le Breton et al. 2005, mountains changed. Sea-level changes during the same
2007). The little fire ant seems to be preadapted to fill a period could also have caused a few additional
vacant or weakly preoccupied niche in New Caledonia. vicariance events between the most peripheral moun-
Every group studied has been characterized by some tains connected by low passes to the main body of the
peculiar presumed adaptation, showing that evolution- central ranges. A stronger argument than one based on
ary novelties are manifold in New Caledonia. What is molecular clocks alone can be made when a clade of
now needed is to use new molecular phylogenetic microendemic species in New Caledonia is sister to
hypotheses to document how such novelties appeared species occurring only on recently uplifted neighbour-
on the island. Did they occur repeatedly in New ing islands ( Norfolk, Lord Howe), which, as
Caledonia by convergence with taxa absent from the mentioned above, is the case for Araucaria trees
island? If so, this would substantiate the assumption of (Setoguchi et al. 1998) and Placostylus land snails
evolutionary diversification into vacant niches. (Trewick et al. in press). Another case is the cockroach
Angustonicus, two species of which occur only on the
Loyalty Islands and are sister to all those on the New
5. LOCAL ENDEMISM Caledonia mainland (Murienne et al. 2005). As argued
Microendemism is extremely high on this medium-sized by Murienne et al. (2005), this kind of relationship
island (16 890 km2) and should not be neglected by between taxa of the New Caledonia mainland and
emphasizing only larger-scale regional endemism of a neighbouring and more recent island is good
(Murienne 2006). Along this 500 km long island, evidence for recent diversification in each group since
many related species are each often restricted to a very sister groups are the same age, i.e. the age of the more
small area (often less than a few square kilometres). This recent island. Trewick (2000) reported a similar case
amazing small-scale endemism has classically been for New Zealand and the Chatham Islands. A contrary
explained by a combination of orography, soils and interpretation would require invoking either unknown
climates, diverse landscape features that result in a or extinct mainland species closely related to those on
mosaic of habitats (Chazeau 1993; Morat 1993b). Many the more recent islands, a presumption that prevents
plant distributions are clearly related to these features, any further logical biogeographic reasoning. However,
as many species are specialized for soils derived from following this presumption, some authors have
either metamorphic–granitic, calcareous or ultramafic hypothesized that palaeo-islands, pre-existing in the
substrates. Plant species are also often limited to either same place as recently uplifted islands (a geologically
dry sclerophyll forest, maquis shrubland or evergreen plausible assumption), could have harboured a
forest on different parts of the mountain ranges at member of the same clade, thus allowing for an older
different elevations, on the leeward, drier western coast age (Heads 2005; Ladiges & Cantrill 2007). Such an
or the windward, wetter eastern coast, from sea level to assumption is, however, not warranted, as there is no
approximately 1600 m (Morat 1993b; Lowry 1998). actual evidence for recently uplifted islands occurring
Until now, no attempt had been made to explain in conjunction with those palaeo-islands; and in any
microendemism with reference to historical factors and case, such a scenario would require several dispersal
speciation processes, except for some assumptions of events among those islands in the past.
climatic refuges for restricted distributions of palms Based on these same studies, speciation seems to
(Pintaud & Jaffré 2001; Pintaud et al. 2001). However, have occurred frequently between different mountains
recent phylogenetic studies provide some insight into or mountain massifs but not necessarily along altitud-
microendemism patterns and their causes. Microende- inal gradients on a single mountain, even though some
mism appears primarily related to allopatric speciation species are distributed preferentially at low or high

Phil. Trans. R. Soc. B (2008)

3314 P. Grandcolas et al. Review. New Caledonia as a Darwinian island

elevations and some altitudinally vicariant distributions 6. CONCLUSION AND FUTURE RESEARCH
are known in plants (most summits are only 1000 m DIRECTIONS
high, the highest being Mont Panié at 1628 m and Based on phylogenetic studies and geological evidence,
Mont Humboldt at 1618 m). In this respect, there is New Caledonia must be regarded as a very old
no differentiation between low- and high-elevation Darwinian island, dating to 37 Ma. The island has
populations of the same microendemic Lauraesilpha been subject to long periods of submersion in the
cockroach species, which are generally restricted to Palaeocene–Eocene, the extent of which, contrary to
one mountain or massif (Murienne et al. in press a). the situation in New Zealand, is not disputed by
Future studies will need to address this issue further by geologists (Lee et al. 2001; Pole 2001; Trewick &
focusing on the highest summits to assess whether the Morgan-Richards 2005; Trewick et al. 2007). Conse-
presence or absence of altitudinal speciation in various quently, the island’s entire biota can only have resulted
groups could be related to the local size of the gradient. from a total recolonization since the Oligocene, which
Although adaptation appears to be a salient feature of is consistent with independent dating from molecular
regional endemism, this is not the case for local phylogenetic studies.
microendemism, in which many closely related and In this context, New Caledonian diversity appears to
allopatrically distributed species apparently do not have resulted from relatively ancient adaptive diversi-
differ from each other in functional traits. For example, fications with abundant recent small-scale speciation
niche modelling among different species of the involving niche conservatism. This contrasts with large
cockroach genera Angustonicus and Lauraesilpha failed tropical forest basins, where small-scale speciation plays
to detect gross differences in microclimatic or altitudinal a minor role (Moritz et al. 2000). A parallel conclusion
parameters (Murienne 2006; Murienne et al. in press b). was drawn by Latimer et al. (2005), who compared
Similarly, niches of Nothofagus species largely overlap plant diversity in the South African fynbos with that
with regard to climate even though they have different in the Amazonian basin. In contrast to its role in the
altitudinal distributions (Read et al. 2005). An excep- New Caledonian terrestrial biota, small-scale speciation
tion to this pattern is the two ecologically segregated also seems to play a minor role in New Caledonian
sympatric sister species of diving beetles on Mont marine ecosystems and especially on the sea mounts of
Panié (Balke et al. 2007b). This probably means that the Norfolk Ridge, where diversity is more related to
microendemism and adaptation reflect complex ecological patchiness than to microendemism on each
individual sea mount (Samadi et al. 2006).
evolutionary processes that take place at various levels
An increasing number of phylogenetic studies have
in the phylogenetic hierarchy, with microendemism
made it possible to propose some preliminary general
tending to happen more distally (and thus inferred to be
conclusions about the evolution of diversity in New
more recent) and adaptive changes occurring more
Caledonia at both regional and local levels. Such
basally (and therefore regarded as more ancient).
studies should now be orientated to address several
Following the same adaptive line of reasoning,
emerging questions. First, work on estimating the time
speciation related to soil diversity and especially to
of origin of groups that may represent pre-Oligocene
metalliferous soils has often been suggested to explain
relicts should be continued to confirm or falsify the
high local richness and persistence of adapted archaic Darwinian nature of New Caledonia. Second, fossil
groups when confronted with supposedly poorly adapted diversity for many groups, including insects, must be
new immigrants (Holloway 1979; Morat 1993b; Haase & better studied, as it can shed light on past diversity and
Bouchet 1998; Lowry 1998; Setoguchi et al. 1998; ecosystem history (Antoine et al. 2006). In this regard,
Bauer & Sadlier 2000). Metalliferous soils have been some studies have revealed a recently extinct large
considered highly stressing substrates for many organ- vertebrate fauna, showing that generalizations based
isms, being poor in nutrients and rich in toxic metals only on extant faunas can sometimes be misleading
including nickel (Proctor 2003). This opinion has been (Gaffney et al. 1984; Balouet & Buffetaut 1987; Balouet &
tempered by some recent studies showing that adaptation Olson 1989). Third, efforts to understand the adaptive
to metalliferous soils in plants is pervasive in many significance of New Caledonian evolutionary novelties
groups, being either symplesiomorphic or convergent must be sought by documenting whether taxa diversified
(de Kok 2002). In insects, except for oligophagous or into so-called vacant niches (Losos et al. 1998; Gillespie
monophagous species feeding exclusively on hyperac- 2004). The amazingly small scale of speciation in New
cumulating plants (Boyd et al. 2006), diversification Caledonia is also an issue for study. In particular,
does not appear to be dependent on metalliferous soils modelling should address the question of whether
(Desutter-Grandcolas & Robillard 2006; Murienne landscape complexity combined with climatic changes
et al. submitted). is sufficient to explain the scale and amount
For all these reasons, a general explanation of New of endemism. Biotic factors possibly promoting specia-
Caledonian microendemism fits better with a model of tion also need to be considered, such as some
speciation involving niche conservatism and population remarkably low population densities, perhaps caused
divergence in environments isolated after climatic by high predation or competition pressures resulting
changes (e.g. mountains; Wiens 2004). New Caledonia from disharmonies in colonization.
is a good model for addressing such issues owing to its
We thank Raphaël Leblois, Sarah Samadi and Jérôme Sueur
complex orography and elongate shape in a subtropical
for reading the manuscript and provided insightful com-
northwest–southwest geographical position resulting ments, and Pete Lowry who also corrected our English. This
in climatic diversity and major effects of historical is a contribution from the project BIONEOCAL funded by
climate changes. the Agence Nationale de la Recherche (2008–2010) and

Phil. Trans. R. Soc. B (2008)

 Review. New Caledonia as a Darwinian island P. Grandcolas et al. 3315

funded previously through the Programme pluriformation Boyd, R. S., Wall, M. A. & Jaffré, T. 2006 Nickel levels in
‘Structure et évolution des écosystèmes’ (Muséum national arthropods associated with Ni hyperaccumulator plants
d’Histoire naturelle). from an ultramafic site in New Caledonia. Insect Sci. 13,
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