Biogeography of the fauna of French Polynesia: diversification within and between a series of hot spot archipelagos.
ABSTRACT The islands of French Polynesia cover an area the size of Europe, though total land area is smaller than Rhode Island. Each hot spot archipelago (Societies, Marquesas, Australs) is chronologically arranged. With the advent of molecular techniques, relatively precise estimations of timing and source of colonization have become feasible. We compile data for the region, first examining colonization (some lineages dispersed from the west, others from the east). Within archipelagos, blackflies (Simulium) provide the best example of adaptive radiation in the Societies, though a similar radiation occurs in weevils (Rhyncogonus). Both lineages indicate that Tahiti hosts the highest diversity. The more remote Marquesas show clear examples of adaptive radiation in birds, arthropods and snails. The Austral Islands, though generally depauperate, host astonishing diversity on the single island of Rapa, while lineages on other islands are generally widespread but with large genetic distances between islands. More recent human colonization has changed the face of Polynesian biogeography. Molecular markers highlight the rapidity of Polynesian human (plus commensal) migrations and the importance of admixture from other populations during the period of prehistoric human voyages. However, recent increase in traffic has brought many new, invasive species to the region, with the future of the indigenous biota uncertain.
- SourceAvailable from: Christophe Thébaud[Show abstract] [Hide abstract]
ABSTRACT: The study of islands as model systems has played an important role in the development of evolutionary and ecological theory. The 50th anniversary of MacArthur and Wilson's (December 1963) article, ‘An equilibrium theory of insular zoogeography’, was a recent milestone for this theme. Since 1963, island systems have provided new insights into the formation of ecological communities. Here, building on such developments, we highlight prospects for research on islands to improve our understanding of the ecology and evolution of communities in general. Throughout, we emphasise how attributes of islands combine to provide unusual research opportunities, the implications of which stretch far beyond islands. Molecular tools and increasing data acquisition now permit re-assessment of some fundamental issues that interested MacArthur and Wilson. These include the formation of ecological networks, species abundance distributions, and the contribution of evolution to community assembly. We also extend our prospects to other fields of ecology and evolution – understanding ecosystem functioning, speciation and diversification – frequently employing assets of oceanic islands in inferring the geographic area within which evolution has occurred, and potential barriers to gene flow. Although island-based theory is continually being enriched, incorporating non-equilibrium dynamics is identified as a major challenge for the future.Ecology Letters 12/2014; · 13.04 Impact Factor
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ABSTRACT: Background Partulid tree snails are endemic to Pacific high islands and have experienced extraordinary rates of extinction in recent decades. Although they collectively range across a 10,000 km swath of Oceania, half of the family¿s total species diversity is endemic to a single Eastern Pacific hot spot archipelago (the Society Islands) and all three partulid genera display highly distinctive distributions. Our goal was to investigate broad scale (range wide) and fine scale (within¿Society Islands) molecular phylogenetic relationships of the two widespread genera, Partula and Samoana. What can such data tell us regarding the genesis of such divergent generic distribution patterns, and nominal species diversity levels across Oceania?ResultsMuseum, captive (zoo) and contemporary field specimens enabled us to genotype 54 of the ~120 recognized species, including many extinct or extirpated taxa, from 14 archipelagoes. The genera Partula and Samoana are products of very distinct diversification processes. Originating at the western edge of the familial range, the derived genus Samoana is a relatively recent arrival in the far eastern archipelagoes (Society, Austral, Marquesas) where it exhibits a stepping¿stone phylogenetic pattern and has proven adept at both intra¿and inter¿ archipelago colonization. The pronounced east¿west geographic disjunction exhibited by the genus Partula stems from a much older long-distance dispersal event and its high taxonomic diversity in the Society Islands is a product of a long history of within¿archipelago diversification.Conclusions The central importance of isolation for partulid lineage persistence and diversification is evident in time-calibrated phylogenetic trees that show that remote archipelagoes least impacted by continental biotas bear the oldest clades and/or the most speciose radiations. In contemporary Oceania, that isolation is being progressively undermined and these tree snails are now directly exposed to introduced continental predators throughout the family¿s range. Persistence of partulids in the wild will require proactive exclusion of alien predators in at least some designated refuge islands.BMC Evolutionary Biology 09/2014; 14(1):202. · 3.41 Impact Factor
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ABSTRACT: 1. The large flightless grasshopper Acrostira bellamyi Uvarov, endemic to the island of La Gomera (Canary Islands), inhabits two different environments: the xeric euphorb shrubland, as is typical for congeneric Canarian species, and the humid laurel forest, a novel habitat for the genus.2. We investigate genetic, morphological, and ecological variation among individuals of A. bellamyi from the two habitats. DNA sequence data were used to evaluate whether grasshoppers from the two environments represent distinct lineages. Morphological and trophic analyses were performed to assess phenotypic differentiation between the two different habitats.3. Population genetic analyses support the hypothesis that the euphorb shrubland is the ancestral habitat for this species. Female laurel forest specimens are larger than those inhabiting the euphorb shrubland, and some external body parts exhibit significant morphometric differences between the two populations. Diet of shrubland individuals is completely different from that of laurel forest individuals. Although in each habitat they consume the most abundant plants, individuals are able to select food plants, which appear to be explained by their nutrient content.4. Our results suggest that A. bellamyi has colonised laurel forest from shrubland, and that this habitat shift has resulted in genetic, morphological, and ecological changes, perhaps as an adaptation to this new habitat.Ecological Entomology 09/2014; x(x):in press. · 1.97 Impact Factor
Biogeography of the fauna of French Polynesia:
diversification within and between a series
of hot spot archipelagos
Rosemary G. Gillespie1,*, Elin M. Claridge2and Sara L. Goodacre3
1Department of Environmental Science, University of California, 137 Mulford Hall,
Berkeley, CA 94720-3114, USA
2UC Berkeley Gump Research Station, BP 244, Maharepa, Moorea 98728, French Polynesia
3Institute of Genetics, University of Nottingham, Nottingham NG7 2UH, UK
The islands of French Polynesia cover an area the size of Europe, though total land area is smaller
than Rhode Island. Each hot spot archipelago (Societies, Marquesas, Australs) is chronologically
arranged. With the advent of molecular techniques, relatively precise estimations of timing and
source of colonization have become feasible. We compile data for the region, first examining
colonization (some lineages dispersed from the west, others from the east). Within archipelagos,
blackflies (Simulium) provide the best example of adaptive radiation in the Societies, though a similar
radiation occurs in weevils (Rhyncogonus). Both lineages indicate that Tahiti hosts the highest
diversity. The more remote Marquesas show clear examples of adaptive radiation in birds, arthropods
and snails.The Austral Islands, though generally depauperate, host astonishing diversityon the single
island of Rapa, while lineages on other islands are generally widespread but with large genetic
distances between islands. More recent human colonization has changed the face of Polynesian
biogeography. Molecular markers highlight the rapidity of Polynesian human (plus commensal)
migrations and the importance of admixture from other populations during the period of prehistoric
human voyages. However, recent increase in traffic has brought many new, invasive species to the
region, with the future of the indigenous biota uncertain.
Keywords: Society Islands; Tahiti; Marquesas Islands; Austral Islands; adaptive radiation; dispersal
The Pacific Ocean contains approximately 30 000
islands (more than the rest of the world’s oceans
combined; Spiess 2007). The most remote islands are
those of Polynesia, notably the Hawaiian Islands, and
the five archipelagos of French Polynesia: the Society,
Marquesas, Austral, Gambier, and Tuamotu Islands.
French Polynesia has a total land area of approximately
3660 km2(less than one-quarter of the size of the
Hawaiian Islands) and is spread over 5 million km2
(similar to the size of Europe) in the South Pacific. For
most terrestrial species, these islands represent tiny
specks of suitable habitat separated by vast distances,
conditions that can provide successful colonists the
opportunity for adaptive radiation.
Biologists have been attracted to the region since
Sir Joseph Banks, as naturalist on the Endeavour
(1768–1771), first returned with an intriguing collection
of specimens (Whitehead 1969; Diment et al. 1984).
The Hawaiian Islands, in particular, have been the focus
of intensive scientific research, which has led to insights
into patterns of diversification and processes of
community and ecosystem development. French Poly-
nesia, by comparison, has received much less attention.
Even Darwin (1859), during his voyage on the Beagle,
had little to say about the terrestrial fauna of Tahiti,
being more concerned by the process of coral reef
formation. His experience aptly reflects the general
situation; scientific research has mostly focused on
marine environments. This is partly due to the
inaccessibility of the mountainous regions, but also the
general perception that, overall, the region is not species
rich, lacking many groups that are common to
continental faunas. However, French Polynesia exhibits
levels of diversity comparable to those of the Hawaiian
Islands when total island area is taken into account.
Until recently, information on arthropods was
largely confined to a series of articles published by the
Bishop Museum (Honolulu), as a result of the Pacific
Entomological Survey in the 1920s and 1930s
(Adamson 1936, 1939). Information from this survey,
while important in laying a foundation for future
research, is both limited and dated for the following
reasons. (i) In archipelagos other than the Marquesas,
collecting localities were limited to a few islands and
sites owing to logistical difficulties (the Marquesas
were relatively well known owing to the efforts of a
local entomologist, G. LeBronnec, although for any
Phil. Trans. R. Soc. B (2008) 363, 3335–3346
Published online 5 September 2008
One contribution of 15 to a Theme Issue ‘Evolution on Pacific
islands: Darwin’s legacy’.
*Author for correspondence (firstname.lastname@example.org).
This journal is q 2008 The Royal Society
one group collections are patchy). (ii) Native and
non-native elements were mixed together in most of
the publications, often without precise locality infor-
mation. Accordingly, useful biogeographic information
is limited. For most taxa, there is much more
information on cosmopolitan species, which are
found in lowland areas and probably represent recent
introductions. The native fauna generally appears to be
confined to higher elevations, and for many taxonomic
groups it remains largely unknown and undescribed.
For terrestrial molluscs, work has been largely
limited to one group, the partulids of the Society and
Marquesas Islands, with early studies over a century
ago (Pilsbry 1900). This early work, which showed
remarkable similarities among species at a large
geographical scale, led to the widespread acceptance
of a hypothesis that the remote islands of Polynesia
were remnants of a Late Palaeozoic or Early Mesozoic
mid-Pacific continent (Gregory 1928). Although this
idea was debunked as geological understanding of the
area increased, it dominated the literature for much of
the early part of the last century. Additional influential
malacological research in the South Pacific was that of
Crampton (1925, 1932), who demonstrated the
Mendelian inheritance of visible polymorphisms, such
as shell colour and chirality, and attempted to calculate
the rate of evolution from the observed changes in
morph frequency, that had occurred in the time
between two field trips. This work, which showed the
value of these Polynesian organisms in studying
evolutionary processes, arguably had a wider effect
through its influence on other evolutionary biologists
such as Dobzhansky and Mayr.
While overall understanding of the South Pacific is
still in its infancy relative to that of the Hawaiian
Islands, considerable advances have been made in
recent years. Here, we compile information generated
to date, focusing on molecular phylogenetic studies of
terrestrial taxa, and attempt to elucidate some general
biogeographic patterns in the region.
2. COLONIZATION OF THE ISLANDS
Once the idea of a mid-Pacific continent had been
dismissed, with geological evidence showing that the
islands were formed de novo from different hot
spots (Nunn 1994), the prevailing paradigm in Pacific
biogeography was that the biota of the central Pacific is
predominantly derived from sources on the western
Pacific Rim, either from continental islands, such
as New Guinea, or from continental regions, namely
lineages that occur in the eastern Pacific are generally
thought to have used intervening archipelagos as
stepping stones for eastward dispersal (e.g. Zimmerman
1948). As a result, lineages occurring on increasingly
on islands to the west, resulting in lineage ‘attenuation’
(Gillespie& Roderick 2002).Recentmolecularevidence
has shown the pattern in some lineages of plants, in
which some diversification has occurred on the different
islands of Polynesia, right out to Hawaii (Wright et al.
2000; Gemmill et al. 2001), and in arthropods such as
broad-nosed Rhyncogonus weevils, which have sister
genera occurring on the western Pacific Rim and which
appear to have colonized island chains in a conservative
stepping stone pattern (Claridge et al. submitted). Other
molecular studies indicate that the blackfly genus
Simulium (Simuliidae) originated in Australasia and has
a stepping stone manner, diversifying prolifically in the
Partula land snails also appear to have colonized from
west to east, with large radiations in the Society and
Marquesas Islands, although the apparent retention of
ancestral lineages within species complicates the
interpretation of colonization history from molecular
the subjects of molecular phylogenetic studies, but
taxonomic affiliations strongly suggest that they may
have western origins and colonized the more remote
islands of the Pacific progressively from the less isolated
Overall, however, few lineages have been shown to
demonstrate a stepping stone model and an increasing
number of molecular phylogenies indicate that several
eastern Polynesian lineages colonized remote Oceania
from the east (the Americas). This has been demon-
strated for numerous plant lineages in the Hawaiian
Islands (Eggens et al. 2007), with some of these sharing
affinities with the Marquesas (Ganders et al. 2000) and
the Societies (Cuenoud et al. 2000). All Hawaiian
spider radiations studied show affinities with the
Americas, with two of these—jumping spiders (Salt-
icidae; Arnedo & Gillespie 2006) and crab spiders
(Thomisidae; Garb & Gillespie 2006)—including the
Society and Marquesas Islands in a large central Pacific
lineage. This American element in the eastern Poly-
nesian fauna should not be surprising, especially as
both air and ocean currents provide opportunities for
colonization of the central Pacific from either direction
(Reverdin et al. 1994; Jokiel & Cox 2003). We might
expect to see a continental American element on
central Pacific islands proportional to the distance of
island groups from the continental source.
The pertinent question may then be: why does the
French Polynesian biota not show as much affinity with
the Americas as does that of the Hawaiian Islands? An
explanation may be the relative distances from the
(Mexico) is the closest mainland to the Marquesas, it
is approximately 5000 km to the northeast, while only
approximately 3200 km east of the Hawaiian chain (the
Societies and Australs are slightly closer to Australia
than South America). There may also be geological
reasons that colonization from the east is more unlikely:
the central Pacific is moving westward, away from the
active spreading centre. Therefore, older landmasses lie
to the west, making them more likely sources of
colonization for younger islands to the east. Evidence
of biogeographic affinities with the eastern Pacific may
also have been obscured by the subduction of the
eastern Pacific plate and anyisland chains at the eastern
plate margin, beneath the Americas. Because land
masses in the western Pacific are drifting westwards,
colonization will generally be from older hot spot
islands west of the younger islands.
3336 R. G. Gillespie et al. Review. Biogeography of French Polynesian fauna
Phil. Trans. R. Soc. B (2008)
Irrespective of exactly where the original colonists
come from, the islands of French Polynesia, as other
remote islands, are characterized by a ‘disharmonic’
biota: many entire groups, notably mammals, are
absent from the native biota. There are no frogs.
Among lizards, their status in the Pacific is unclear.
There are five species of geckos in French Polynesia,
which are almost uniform genetically and hence almost
certainly of very recent origin in the area (Fisher 1997).
Also, three species of widespread Pacific geckos are
considered native in French Polynesia: Nactus pelagicus,
Lepidodactylus sp. and Gehyra oceanica. Evidence that
G. oceanica is native is based largely on a north–south
oceanic divide in genetic structure; however, human-
mediated dispersal cannot be ruled out. At the same
time, groups, such as many birds, arthropods and
snails, can be diverse with high levels of endemism.
3. AFFINITIES WITHIN THE CENTRAL PACIFIC
Biological similarities across the islands of the central
insects (Meyrick 1935a,b), spiders (Berland 1942) and
plants (Guillaumin 1928; Campbell 1933), were
initially explained as the result of a former land
connection across the Pacific Ocean. However, it is
now clear that colonization of the islands occurred
through transoceanic dispersal. Similarities among
the biotas of different archipelagos can be attributed
to two types of dispersal (Gillespie 2002): (i) jumping
from one island group to the next, as described above,
or (ii) long-distance colonization by only the most
dispersive organisms that can readily colonize repeat-
edly and independently from a mainland source,
with similarity across archipelagos explained by con-
vergent adaptation to similar ecological opportunities.
For example, reed warblers have colonized nearly all
the islands of the Marquesas and look remarkably
similar across the islands. However, molecular data
indicate that the Marquesas reed warbler includes
two independent lineages: the northern Marquesas
reed warbler, closely related to the Tuamotu reed
warbler, and the southern Marquesas reed warbler,
sister to that of Kiribati (Cibois et al. 2007). Both
colonizations occurred ca 0.6 Ma, more recently than
the formation of the islands, and suggest that the
bird is a ‘supertramp’, colonizing remote islands easily
Although a few lineages have colonized all the
islands of the central Pacific from either the east or
the west, others include elements that have colonized
part of the Pacific from the west, and part from the
east. The questions are: where do they meet and how
do they interact where they meet? This has been
examined in crab spiders (Thomisidae; Garb &
Gillespie 2006). Two of the common genera in the
central Pacific are Misumenops and Diaea. Both genera
are worldwide in distribution but Misumenops has
its highest diversity in the Americas (65% of species),
though with some representation in Asia (15%)
and none in Australia/New Guinea; Diaea has its
highest diversity in Australia/New Guinea (77%
species) and very few species in the Americas (2%).
A third genus, Mecaphesa, has four endemic Hawaiian
species (Simon 1900). Misumenops is widely distributed
in eastern Polynesia (Hawaiian, Society, Marquesas
and Austral Islands), while representatives of Diaea
occupy parts of western Polynesia (Samoa, Tonga and
New Zealand) and Melanesia (Fiji, New Caledonia,
New Guinea, Solomon Islands and Vanuatu; Platnick
2008). Molecular data have shown that the Austral
Islands species, Misumenops rapaensis, is far more
closely related to Diaea spp. from western Polynesian
and Melanesia than to other species of Misumenops
from eastern Polynesia. The narrow and well-defined
break between fauna originating from the east and west
is interesting because the eastern fauna covers 4500 km
from Hawaii to the Societies and extends to North
America, yet does not reach the Australs, which are
only 600 km south of Tahiti. Moreover, the ranges of
the two lineages abut, but do not overlap. The ability of
either group to become successfully established in any
of these islands may depend on the order of their
arrival. Indeed, it is likely that many taxa that have
colonized any of the remote islands successfully have
the potential for further colonization, with this
potential seldom being realized owing to historical
precedence of earlier colonists.
4. BIOGEOGRAPHIC PATTERNS WITHIN
Whatever their origin, much of the fauna of the remote
French Polynesian islands, just as the Hawaiian fauna,
may be endemic not only to an archipelago but often to
a single island and even a single area within an island.
Moreover, there can be multiple closely related species
in an archipelago (usually attributed to adaptive
radiation; Craig et al. 2001). However, patterns may
differ considerably between islands.
(a) Society Islands
Age progression in the Society Islands is in good
agreement with the fixed hot spot hypothesis (figure 1;
Clouard & Bonneville 2005). The islands extend from
Maupiti, the oldest of the current islands in the north
at 4.3 Ma (Guillou et al. 2005), to the largest and
youngest island of Tahiti at 2.0–0.5 Ma and the islet of
Mehetia (0–0.26 Ma) in the south (Clouard &
Bonneville 2005). One of the best-studied groups of
arthropods in the Society Islands—and indeed in all of
French Polynesia—is the blackfly genus Simulium
(Joy & Conn 2001; Craig 2003). In the Society
Islands, there are 31 described species of Simulium;
Tahiti has the most (29) with fewer on the older and
smaller islands of Moorea (10), Huahine (2) and Bora-
Bora (1). The two widespread species, Simulium
malardei and Simulium lotii, appear to be basal, and
all species appear to have arisen on the youngest island,
Tahiti, with back-dispersal of the highly modified
cascade-dwelling species northwest to Moorea and
Raiatea (figure 2a). The phylogenetically basal position
of the widespread species coupled with the molecular
clock calibration suggest that the Tahiti specialist
species Simulium oviceps is 1.8–2.0 Ma old, which fits
the geological framework (Clouard & Bonneville
2005), although it is also possible that some of the
species on Tahiti arose on older islands from which
Review. Biogeography of French Polynesian fauna
R. G. Gillespie et al.
Phil. Trans. R. Soc. B (2008)
they have subsequently gone extinct (Craig 2003).
Extinctions of habitat specialist species, and others, on
the older islands might be expected owing to island
erosion and concomitant loss of running water
habitats. Tahiti is the largest island at present, with
an abundance of running water habitats; however, its
diversity may simply be a temporary result of the latest
intra-island species radiation. As Tahiti ages and
erodes away, most of these species will no doubt
Larval ecology has played a major role in the
diversification of blackflies in the Society Islands, and
fine-scale partitioning of feeding niches may have
facilitated the repeated colonizations of rivers (Joy
et al. 2007). Cascade populations exhibit higher levels
of genetic subdivision than river populations, perhaps
because cascades are intrinsically more isolated from
each other than rivers. Genetic assimilation, in which
moderate levels of phenotypic plasticity promote
establishment in novel habitats, and subsequent
selection for extreme phenotypes leading to genetic
differentiation, and perhaps speciation, may have
played a major role in the radiation of blackflies in the
Society Islands by enhancing their ability to success-
fully colonize novel niches (Joy et al. 2007).
The broad-nosed weevil genus Rhyncogonus also
occurs across French Polynesia. There are 16 described
species (Van Dyke 1937) and 11 recently collected
undescribed species in the Society Islands. As in the
blackflies, Tahiti hosts the highest number of species
(14) and the smaller and older islands host fewer;
each species is endemic to a single island. Unlike the
blackflies, in which the Society Islands appear to
have been colonized once, the closest out-groups
being a lineage that includes Fiji, New Caledonia
and Micronesia/Marquesas (Craig et al. 2001), in
Rhyncogonus there appear to have been multiple
independent colonizations of Tahiti from neighbouring
island chains (Claridge 2006).
There are two lineages of long-jawed spiders
(Tetragnatha, Tetragnathidae; Gillespie 2003b), repre-
senting independent colonizations to the islands
(Gillespie 2002). Tetragnatha moua is limited to the
highest elevations of Tahiti, with a close relative
(undescribed) on the summit of Moorea (R. G.
Gillespie 2007, unpublished data), while Tetragnatha
rava and Tetragnatha tuamoaa are sister taxa that occur
lower down on both islands (figure 2b). A similar
pattern, in which species from the cloud forest (limited
to the summits of Raiatea, Moorea and Tahiti) are most
Figure 1. The main high archipelagos in French Polynesia, with geological ages indicated (Clouard & Bonneville 2005).
3338 R. G. Gillespie et al.Review. Biogeography of French Polynesian fauna
Phil. Trans. R. Soc. B (2008)
closely related to taxa in other adjacent cloud forests
rather than species farther down the mountain on the
same island (Gillespie et al. 2008; R. G. Gillespie 2007,
unpublished data), has been shown for Polynesian
plants (Meyer 2004). Likewise, taxa lower down the
mountain are related to others lower down on adjacent
islands. This is in contrast to the genetic similarity
between Partula snail species found in the same
geographical location rather than between those in
similar habitat types at different geographical locations,
a factor that is believed to be strongly associated with
historic (and/or contemporary) hybridization between
sympatric species (Goodacre 2002).
Like the arthropods, the terrestrial snails of the
genus Partula in French Polynesia also show what is
probably an important feature of Pacific Island
colonization, i.e. occasional long-distance migration
may play an important part in determining the current
distributions, since indirect estimators of gene flow
have much higher values than those predicted directly
from observed migration distances (Murray & Clarke
1984). One explanation for the discrepancy is that
direct measures fail to account for the effects of
occasional long-distance migrants. Like the blackflies,
there are fewer species of Partula on older islands
(Johnson et al. 1993).
Pacific islands such as Tahiti and Moorea have high
ridges and steep-sided valleys, characteristic features of
volcanic islands, but the ridges are no significant barrier
to the movement of snails, which are often observed
at high altitude. Despite the absence of the current
movement barriers, marked differences are observed
T. moua Marau Tahiti
T. nitens Moorea Gump
T. punua Nuku Hiva Tekao
T. punua Nuku Hiva Tekao
T. marquesiana Nuku Hiva Tovii
T. marquesiana Nuku Hiva
T. marquesiana Nuku Hiva
T. marquesiana Nuku Hiva Tekao
T. (marquesiana) Ua Huka
T. kapua Hiva Oa Temetiu
T. kapua Hiva Oa Temetiu
0.05 substitutions per site
T. macilenta Moorea
T. laboriosa USA MA
T. maxillosa Moorea
T. mandibulata Australia
Nuku Hiva Pomarea mendozae nukuhivae
Pomarea iphis iphis
Pomarea mendozae mendozae
Ua Pou Pomarea mendozae mira
Eiao Pomarea iphis fluxa
Pomarea mendozae whitneyi
Pomarea mendozae motanensis
T. rava Tahiti Iti 39
T. rava Tahiti Iti 38
T. rava Tahiti Aorai 580m 18
T. guatemalensis USA PA 29
T. versicolor USA California 22
T. moua Tahiti Aorai 5
T. moua Tahiti Marau 40
T. moua Tahiti Marau 41
T. moua Tahiti Aorai 9
T. moua Tahiti Aorai 1
T. nitens New Zealand 85
T. nitens New Zealand 97
T. mandibulata Australia 26
T. laboriosa USA MA 31
T. rava Tahiti Belvedere 7
T. tuamoaa Moorea Vaiare-Paopao 42
T. tuamoaa Moorea 3-coconuts 50
T. tuamoaa Moorea 3-coconuts 46
T. nitens New Zealand 91
T. nitens Papua New Guinea 86
T. nitens Moorea, Gump 1
T. nitens Indonesia 32
T. nitens Puerto Rico 2
S. bellula (Ua Pou)
S. strigata (Ua Huka)
S. decussatula (Hiva Oa)
S. ganymedes (Hiva Oa)
S. ganymedes (Fatu Hiva)
S. inflata (Tahuata)
Islands of occurrence
Tahiti <--------> Bora-
Figure 2. (Caption overleaf.)
Review. Biogeography of French Polynesian fauna
R. G. Gillespie et al.
Phil. Trans. R. Soc. B (2008)
between conspecific Partula populations, which appear
highly structured in terms of shell shape, shell colour
and banding patterns (Clarke & Murray 1969; Johnson
et al. 1986) and mitochondrial DNA haplotypes
(Goodacre 2002). A leptokurtic pattern of dispersal
may explain this patchiness, which is the expected
consequence of long-distance migration, that is pre-
dicted to remain for many generations in the absence of
selection (Ibrahim et al. 1996).
(b) Marquesas Islands
The Marquesas extend from Nuku Hiva, the oldest of
the current high islands in the north at 3.7 Ma (plus
some lower islands, including Eiao, 5.5 Ma, and
Hatutaa, 4.8 Ma, farther north), to Hiva Oa (2.4 Ma),
Tahuata (1.9 Ma) andFatu Hiva (1.8 Ma),the youngest
island, in the south (figure 1; Clouard & Bonneville
2005). The arrangement of islands in a chronological
series is not strictly regular, with Ua Huka (1.9 Ma)
adjacent to the much older island of Nuku Hiva.
The best-known radiation of birds in the Marquesas
is the monarch genus Pomarea (Monarchidae), which is
endemic to the Cook, Society and Marquesas archipe-
lagos, with the most extensive diversification in the
Marquesas (figure 2c). The genus has suffered
extensive recent extinction but molecular studies have
used specimens at the American Museum of Natural
History collected during the Whitney South Sea
Expedition in the 1920s, which allowed development
of a phylogeny of the entire genus, including extinct
taxa (Cibois et al. 2004). The phylogeny is consistent
with the sequential appearance of the Marquesas
Islands. Differences between the ages of the islands
and the estimated ages of the nodes indicated
colonization 1–2 Ma after the islands emerged.
Among Tetragnatha spiders, sampling is incomplete,
but data from Nuku Hiva, Ua Huka and Hiva Oa
indicate that the genus has undergone asmall radiation,
with two sympatric species on Nuku Hiva and one
species on each of the other islands (Gillespie 2003a).
The species on Nuku Hiva are sisters and the sister
relationship between the species on Ua Huka and Hiva
Oa accords with the youth of these islands relative to
Nuku Hiva (figure 2e).
There are 22 described species of Rhyncogonus
weevils in the archipelago, three of which inhabit the
oldest islands of Eiao and Hatutaa. There was a single
colonization of the island chain. There has also been
considerable diversification within the southern island
group, particularly on Hiva Oa (seven species) and
Fatu Hiva (five), but there are fewer species in the
northern group, with Nuku Hiva, of size comparable to
Hiva Oa, having just a single species (Claridge 2006).
There is less information on the diversity and history
of blackflies in the Marquesas, with the southern
islands poorly sampled. However, the flies have
not diversified into specialized habitats, as in the
Society Islands, and the larvae of most species are
Partulid land snails of the genus Samoana are
monophyletic in the Marquesas, based on allozyme
variation, with multiple co-occurring species (Johnson
et al. 2000; figure 2d). Several Marquesan Samoana
species display a suite of characteristics (thick shells,
short tentacles and non-sticky mucus) that were
thought to be restricted to the partulid genus Partula,
drawing into question the classification of these species.
But molecular data (Johnson et al. 2000; Goodacre &
Wade 2001) showed the similarity to be the result of
independent evolution of ‘thick-shelled’ and ‘thin-
shelled’ species in Samoana. The recent discovery of a
thin-shelled Partula on Raiatea demonstrates that such
independent evolution has occurred several times and
in both directions (Burch 2007).
(c) Austral Islands
The Austral archipelago is approximately 500 km
southwest of the Society Islands (at their closest points:
between Tahiti and Rurutu) and extends over 1500 km
from the southernmost Marotiri Isles to the atoll
Maria. The Austral Islands are geologically continuous
with the Cook Islands (to the northwest), which
together were formed from repeated episodes of
vulcanism at several sites (Dickinson 1998; Bonneville
Figure 2. (Overleaf.) Phylogenetic hypotheses for different lineages on the (a,b) Society and (c–e) Marquesas Islands.
(a) Phylogenetic hypothesis for blackflies, Simulium (Simuliidae), from the Society Islands showing distribution and shifts in
larval habitat (redrawn from Craig et al. 2001). Islands within the Society chain where each species occurs are shown (Simuliun
adamsoni is the only species in the Society Island clade that does not occur in the Societies, as indicated by the black bars).
Habitats range from streams, of various types, to rivers, cascades and madicolous flow (thin films of water). Line width is
proportional to the number of different habitats used. The phylogeny suggests a single shift to the specialized cascade habitat
with three losses (white bars on branches) and one independent gain earlier in the phylogeny (black bar to Simulium fossatiae).
(b) Phylogenetic hypothesis of spiders, genus Tetragnatha (Araneae, Tetragnathidae), in the Society Islands. The data are based
on sequences (approx. 750 bp) of mitochondrial COI DNA (GenBank accession numbers EU796899–EU796932). Details of
methods are provided by Gillespie (2002). Analysis was by parsimony, maximum likelihood and Bayesian estimates of
likelihood, and values beside nodes indicate bootstrap support (above node) and posterior probabilities (below node).
(c) Phylogenetic tree for the Pomarea monarch flycatchers mapped on the Marquesas Islands (branch lengths not proportional to
sequence evolution; redrawn from Cibois et al. 2004). Taxa endemic to other Polynesian archipelagos are connected to the
Marquesan topology with a dashed line. Three taxa, Pomarea iphis fluxa, Pomarea mendozae nukuhivae and Pomarea iphis iphis,
are basal in the tree (relative positions uncertain). All other taxa form a tight clade consistent with the age and proximity of the
islands. For estimating the age of lineages, the three best-supported nodes were used, labelled as 1 (separation of the basal
Marquesan monarchs), 2 (divergence between the basal Marquesan taxa and the remaining taxa) and 3 (divergence between
species on Hiva Oa versus Tahuata and Fatu Hiva). Node 1 was estimated at 3.0–3.3 Ma, node 2 at 1.6–1.8 Ma and node 3 at
0.41–0.45 Ma. (d) Fitch tree based on genetic distances (allozymes) among samples and species of Samoana snails from the
Marquesas and Society Islands. Numbers at each node indicate the number of times that clade appeared in 100 iterations in the
bootstrap analysis (redrawn from Johnson et al. 2000). (e) Phylogenetic hypothesis of Tetragnatha spiders in the Marquesas. Data
and analysis as in (b).
3340R. G. Gillespie et al. Review. Biogeography of French Polynesian fauna
Phil. Trans. R. Soc. B (2008)
et al. 2002). Potassium–Argon (K–Ar) dating indicates
that the Cook–Austral chain began forming ca
20–30 Ma (Keating 1987; Munschy et al. 1998) with
ages of the Australs ranging from 4.3 Ma (Marotiri
Isles) to 15.7 Ma (Maria), and the main islands ranging
from 4.5 to 12.2 Ma (Clouard & Bonneville 2005;
figure 1). The Austral Islands, like the Societies and
Marquesas, are sequentially ordered from southeast to
northwest by increasing age, as a result of the north-
westward movement of the Pacific tectonic plate over
stationary volcanic plumes, decreasing in age from
northwest to southeast. However, there was secondary
volcanic activity beneath Rurutu ca 1–2 Ma, which
is also associated with secondary uplift of the island
and the neighbouring island of Rimatara. It is possible
that both islands were subaerial prior to their recent
uplift. Thus, the progression rule here would predict
colonization from west to east down the chain, but
with possible secondary recolonization of Rurutu
Although generally taxonomically depauperate, the
Austral Islands host a surprising number of endemic
species, in particular on Rapa (Claridge et al. in press).
For example, among beetles, the genus Miocalles,
a group of tiny flightless weevils, has undergone an
astonishing radiation on Rapa with 67 described
species in an area of just 40 km2, with diversification
accompanied by striking morphological and ecological
differentiation (Paulay 1985). The weevil genus
Rhyncogonus is also particularly diverse in the Australs,
with 22 described species, half of these on Rapa,
though three previously undescribed species have been
collected recently on Raivavae, and the oldest high
island, Rimatara, just 8 km2and 83 m high, supports
an endemic species (Van Dyke 1937; Claridge 2006).
Studies by Clarke (1971) on the Lepidoptera Rapa
resulted in the description of a number of diverse
groups. Recent phylogenetic studies (Craig et al. 2001;
Wright et al. 2001; Mitchell & Heenan 2002) have
incorporated a few species from some of these islands in
the context of broader biogeographic analyses of
relationships among different sets of islands in the
Pacific Basin. However, there have been no thorough
phylogenetic treatments of these groups and much
work remains to elucidate the enigmatic diversity of
Similar stories are emerging of sequential coloniza-
tion of islands by spiders, with large genetic distances
M. rapaensis occurs throughout the Austral Islands,
with large genetic distances between islands: the
uncorrected distance between Rurutu and Tubuai is
8.4 per cent, nearly as much as the maximal distance
across all 16 included Hawaiian taxa (Garb & Gillespie
2006; figure 3). Molecular clock estimates indicate that
the split between the Tubuai clade and the clade
672 M. rapaensis, Raivavae
673 M. rapaensis, Raivavae
677 M. rapaensis, Rapa
678 M. rapaensis, Rapa
679 M. rapaensis, Rapa
683 M. rapaensis, Rapa
685 M. rapaensis, Rapa
674 M. rapaensis, Raivavae
675 M. rapaensis, Raivavae
676 M. rapaensis, Raivavae
682 M. rapaensis, Rapa
680 M. rapaensis, Rapa
684 M. rapaensis, Rapa
681 M. rapaensis, Rapa
665 M. rapaensis, Tubuai
671 M. rapaensis, Tubuai
664 M. rapaensis, Tubuai
669 M. rapaensis, Tubuai
670 M. rapaensis, Tubuai
666 M. rapaensis, Tubuai
667 M. rapaensis, Tubuai
668 M. rapaensis, Tubuai
663 M. rapaensis, Rurutu
662 M. rapaensis, Rurutu
649 M. rapaensis, Rurutu
699 Diaea sp., Fiji
697 Diaea sp., Fiji
696 Diaea sp., Fiji
Raivavae + Rapa
Figure 3. Phylogenetic hypotheses for crab spiders, M. rapaensis (Thomisidae), in the Austral Islands inferred from molecular
genetic analysis (Garb & Gillespie 2006). Numbers above branches refer to parsimony bootstrap values from 1000 replicates
(asterisks indicate less than 50% bootstrap support) followed by decay indices. Numbers below the branch indicate posterior
Review. Biogeography of French Polynesian fauna
R. G. Gillespie et al.
Phil. Trans. R. Soc. B (2008)
comprising individuals from Raivavae and Rapa
occurred more recently (ca 2.5–3.8 Ma) than the
divergence of M. rapaensis from Rurutu and Tubuai
(estimated at 4.9–7.5 Ma), suggesting that M. rapaensis
initially colonized the Australs several million years
after the formation of Rurutu (12.7 Ma) and Tubuai
(10.4 Ma) but possibly before the emergence of
Raivavae (6.8 Ma) and Rapa (5.0 Ma). The estimated
divergences further suggest that from the initial point
of colonization (Rurutu or Tubuai), M. rapaensis did
not reach Raivavae or Rapa until the emergence of
both islands. The orb web spider Tangaroa tahitiensis
(Uloboridae) occurs throughout both the Society
and Austral Islands, again with large genetic distances
between islands (R. G. Gillespie 2007, unpublished
data), although, as with M. rapaensis, the data suggest a
more recent colonization than the geological age
The land snail fauna of the Australs is particularly
impressive, with more than 100 species on Rapa alone
(Solem 1982, 1984), but their relationship to species
elsewhere in Polynesia has not been studied in depth
using molecular methods, with one exception. The
rather atypically widely distributed Partula hyalina is
shared among the geographically distant Cook, Society
and Austral archipelagos, a distribution suggested as
being the legacy of prehistoric human transport (Lee
et al. 2007a).
5. HUMAN COLONIZATION
The human history of Polynesia is one of the most
fascinating and tractable systems in which to examine
the interactions between people and biodiversity. The
best-known scenarios for the peopling of the Pacific are
the ‘express train’ and the ‘entangled bank’ (Hurles
et al. 2003). The express train is built on the notion
of a simple spread, with a strong phylogenetic signal,
of Polynesian ancestors (Austronesians), first into
near Oceania and then into remote Oceania. The
entangled bank is a reticulate model that highlights
the importance of ongoing interaction among popu-
lations. Many intermediate models have been pro-
posed, such as the ‘slow train’ (more genetic mixing
between Austronesians and the original inhabitants
before moving into remote Oceania) and ‘slow boat’
(ultimate origin Asia, and proximate origin within
Wallacea). Mitochondrial DNA studies have detected
characteristic haplotypes, such as a 9 bp deletion at
high frequency, that predominate in Polynesia. The
mutations implyan ultimate originin Asia (‘out of Asia’
model), with subsequent movement following either
the express-train, slow-train or slow-boat models
(Melton et al. 1995). Y-chromosome studies also
show male-biased European admixture among some
Polynesian populations and raise the possibility of
sex-specific differences in prehistoric demography,
including levels of endogamy and/or migration patterns
(Su et al. 2000).
Polynesians carried a range of plants and animals
with them as they moved across the Pacific, and genetic
evidence from these has also been examined. Mito-
chondrial DNA lineages shared between Pacific rat
(Rattus exulans) populations identify homeland regions,
and disjunct variation within island rat populations
suggests separate introductions from several sources,
with origins in the Lapita and a spread into remote
Oceania. The express train to Polynesia model is
rejected, while a ‘voyaging corridor’ aspect of the
slow-boat model is supported, but with most of the
evidence for a ‘voyaging corridor triple’, a model
allowing various components of the Lapita cultural
complex to be the result of intrusion of new
components (Matisoo-Smith & Robins 2004). Other
animals examined include pigs, dogs and chickens,
although evidence from these is complicated by
more recent introduction of the same species by
European settlers. However, recent data from the pig,
Sus scrofa, suggest that a Pacific clade originated in
peninsular Southeast Asia, where they were first
domesticated. Polynesian dispersals into Oceania
appear to be exclusively associated with Pacific clade
pigs (Larson et al. 2007).
Other organisms were introduced accidentally by
early Polynesians. For example, genetic analysis of
populations across the Pacific of Lipinia noctua, a lizard
native to New Guinea, which has been transported
from near to remote Oceania, support the express-train
model of human colonization (figure 4; Austin 1999).
In addition to the standard migration routes of
Polynesians and associated organisms, recent molecu-
lar work has revealed ‘trading’ of species between
archipelagos. In particular, the white-shelled tree snail,
P. hyalina, has been found in the Society, Austral
and southern Cook Islands. Recent molecular work
has shown that P. hyalina was originally restricted to
Tahiti, but trading between archipelagos resulted in
multiple founder populations in the Australs and
southern Cooks (Lee et al. 2007a). With the recent
arrival of the alien carnivorous land snail Euglandina
rosea and subsequent devastation of island snails on
Tahiti (see below), P. hyalina is now restricted to the
southern Cooks and Australs; on Tahiti it still persists
but is threatened.
6. RECENT IMPACTS
Since the arrival of Europeans in Polynesia (late
1700s), the rate of influx of non-native species has
increased tremendously. Native plants are now largely
confined to montane forests above approximately
300 m (cloud forests and subalpine forests), while
coastal, dry lowland and low and middle elevation
valley forests have been seriously disrupted by human
activities, introduced mammals (feral goats, sheep,
cattle, horses, pigs) and invasive plants (approx. 375
alien plant species are naturalized on Tahiti, and
approx. 220 on Nuku Hiva (Florence 1993)). The
many bird extinctions on these islands are now well
known (Steadman & Rolett 1996).
The origins of more recent arrivals to French
Polynesia can best be predicted by tracing international
trade and transport routes. Most invasive species arrive
in French Polynesia via the international airport or port
on Tahiti and then gradually spread across the region,
again following transport routes (Grandgirard et al.
2006; Petit et al. in press). Recent introductions to
French Polynesia include a series of predators. The sac
3342R. G. Gillespie et al.Review. Biogeography of French Polynesian fauna
Phil. Trans. R. Soc. B (2008)
spider Cheiracanthium mordax (Miturgidae), native to
Australia, is invasive in the Pacific, with almost
identical haplotypes from Micronesia through all of
French Polynesia to Hawaii (R. G. Gillespie 2007,
unpublished data). Likewise, the spider Pholcus ancoralis,
Fiji, Micronesia and all of French Polynesia; it has also
recently been reported in Hawaii (R. G. Gillespie 2007,
unpublished data).Alltheseintroductions appear tohave
been accidental, mediated through transportation routes.
An exception to the main route of accidental
introduction is the arrival of the rosy wolf snail,
E. rosea, which was deliberately introduced to French
Polynesia as part of a biological control programme
directed towards eliminating the giant African land
snail, Achatina fulica. Documentation of the spread of
this non-native species illustrates the speed at which
invasive species may become established and influence
the distribution of native species (Clarke et al. 1984).
Indeed, snails in the genus Partula epitomize rapid
extinction caused by an introduced predator, with 56 of
61 Society Island species now extinct in the wild
(Cowie 1992; Coote & Loeve 2003). However, recent
molecular work paints a more positive conservation
picture, with montane populations of Partula otaheitana
and valley populations of Partula clara/Partula hyalina
persisting, and these populations include genetic
representation of all major mitochondrial clades that
occurred historically on Tahiti (Lee et al. 2007b).
Other non-native taxa that have been studied in the
Pacific include parasites and their associated vectors.
Among parasites, the Hawaiian form of malaria is the
only lineage of malaria parasite that appears to be
et al. 2006). In a survey of birds on Moorea, Society
several introduced species (no data on native species
owing to their rarity). However, in the Marquesas, the
parasite is common in a small sample of endemic
Marquesan reed warblers (Acrocephalus mendanae),
suggesting that these birds, which are relatively recent
(1–2 Ma) colonists of the islands, may be resistant to the
parasites. On the other hand, the Pomarea flycatchers
may have succumbed to the disease.
Marked genetic structure has been demonstrated
among Tahiti and Moorea populations of mosquitoes,
in particular Aedes aegypti, which is associated with
dengue fever in humans. Here, the structure appears to
be dictated by human population density, intensity of
insecticidal control and ecological characteristics of
mosquito ecotopes (Paupy et al. 2000).
coagulata, is notorious in North America for transmitt-
ing Pierce’s disease, caused by the bacterium Xylella
fastidiosa, in grapevines. The bacterium can also infect
many other plants. The sharpshooter’s arrival in French
Polynesia (Tahiti) in July 1999 caused considerable
concern, both owing to its abundance, which results in
almost constant ‘rain’ from the excreta in some areas,
and its potential role as a vector of X. fastidiosa, which
could affect species of Metrosideros, Weinmannia,
Dodonaea, Glochidion, Hibiscus and Gardenia in the
Society Islands (Grandgirard et al. 2006). Populations
of H. coagulata are geographically structured into two
Figure 4. Evidence of the ‘express-train’ model of Polynesian island colonization by lizards (redrawn from Austin 1999).
Maximum-parsimony phylogram for Lipinia noctua; localities denoted by grey circles are all genetically distinct and represent
natural prehuman dispersal. Localities denoted by black circles are genetically similar (mean sequence divergence 0.008%) and
represent human-mediated dispersal within the past 4000 years. Dates represent approximate time of first human settlement.
Review. Biogeography of French Polynesian fauna
R. G. Gillespie et al.
Phil. Trans. R. Soc. B (2008)
groups of COI haplotypes in North America, a group of
populations from east of the Mississippi River, and a
group comprising populations from west of the
Mississippi River (Texas and California): haplotypes
from Tahiti fall in the latter group (Smith 2005).
Several general remarks can be made from this overview
of studies of such a wide range of taxa throughout
French Polynesia. First, patterns of biodiversity differ
among archipelagos within the area. This may reflect
environmental heterogeneity across the region and
differences in the degree to which island populations
are influenced by immigration, natural selection and
drift. For example, the Society Islands are characterized
by high numbers of endemic species on the youngest
island, with independent colonization from either
neighbouring islands or archipelagos playing a promi-
nent role in shaping biodiversity. By contrast, the
Australs show a somewhat different pattern, with little
evidence of ecological differentiation on the low islands
but extensive genetic divergence among islands. This
pattern might be predicted on the basis of the small size
of these islands, if size limits diversification. If so, Rapa,
the southernmost of the Australs, is an exception, with
its high level of endemism and striking examples of
adaptive radiation, despite being similar in size to others
in the Austral group.
The Marquesas show a strong signal of adaptive
radiation within the archipelago, although to a lesser
extent than in the Hawaiian Islands. Invertebrates are
characterized by a large number of monophyletic
lineages with evidence for at least some degree of
adaptive radiation and (in snails) for parallel evolution
of similar morphs on different islands.
Phylogenies for the Australs and Marquesas show
some support for the progression rule, a pattern seen in
the Hawaiian Islands, where lineage formation reflects
successive colonization of islands in the order of their
formation (Wagner & Funk 1995). However, at least for
spiders in the Australs, the age of the lineages is
considerably younger than the geological age of the
islands. There is little evidence as yet for the progression
rule in the Societies.
Another general trend is that the non-native biota
appears to be similar across much of French Polynesia.
It is interesting to observe that native communities
appear to be more intact on the Australs than on the
other higher archipelagos, at least at lower elevations.
This is somewhat surprising, given that native com-
munities are smaller and apparently more depauperate
on the Australs; it could provide insights into factors
underlying successful biological invasion. Further work
is required in order to understand the interaction
between native and non-native elements of the biota. In
summary, results to date highlight the value of the
fauna of French Polynesia for studying the different
evolutionary processes that driveadaptation, speciation
and community assembly, and the forces that govern
how species invade and become established over both
evolutionary and ecological time.
We thank Rob Cowie and Steve Trewick for organizing this
special issue. The manuscript was greatly improved by the
comments of three anonymous reviewers. This work was
supported by the Territorial Government of French Poly-
nesia, with funds from the National Science Foundation
(DEB 0451971 to R.G.G.), the Schlinger Foundation,
University of California Berkeley and the Gordon and Betty
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