The Galápagos Islands: biogeographic patterns and geology

In the traditional biogeographic model, the Galápagos Islands appeared a few million years ago in a sea where no other islands existed and were colonized from areas outside the region. However, recent work has shown that the Galápagos hotspot is 139 million years old (Early Cretaceous), and so groups are likely to have survived at the hotspot by dispersal of populations onto new islands from older ones. This process of metapopulation dynamics means that species can persist indefinitely in an oceanic region, as long as new islands are being produced. Metapopulations can also undergo vicariance into two metapopulations, for example at active island arcs that are rifted by transform faults. We reviewed the geographic relationships of Galápagos groups and found 10 biogeographic patterns that are shared by at least two groups. Each of the patterns coincides spatially with a major tectonic structure; these structures include: the East Pacific Rise; west Pacific and American subduction zones; large igneous plateaus in the Pacific; Alisitos terrane (Baja California), Guerrero terrane (western Mexico); rifting of North and South America; formation of the Caribbean Plateau by the Galápagos hotspot, and its eastward movement; accretion of Galápagos hotspot tracks; Andean uplift; and displacement on the Romeral fault system. All these geological features were active in the Cretaceous, suggesting that geological change at that time caused vicariance in widespread ancestors. The present distributions are explicable if ancestors survived as metapopulations occupying both the Galápagos hotspot and other regions before differentiating, more or less in situ.     

Genetic diversity and connectivity of deep‐sea hydrothermal vent metapopulations

Deep‐sea hydrothermal vents provide ephemeral habitats for animal communities that depend on chemosynthetic primary production. Sporadic volcanic and tectonic events destroy local vent fields and create new ones. Ongoing dispersal and cycles of extirpation and colonization affect the levels and distribution of genetic diversity in vent metapopulations. Several species exhibit evidence for stepping‐stone dispersal along relatively linear, oceanic, ridge axes. Other species exhibit very high rates of gene flow, although natural barriers associated with variation in depth, deep‐ocean currents, and lateral offsets of ridge axes often subdivide populations. Various degrees of impedance to dispersal across such boundaries are products of species‐specific life histories and behaviours. Though unrelated to the size of a species range, levels of genetic diversity appear to correspond with the number of active vent localities that a species occupies within its range. Pioneer species that rapidly colonize nascent vents tend to be less subdivided and more diverse genetically than species that are slow to establish colonies at vents. Understanding the diversity and connectivity of vent metapopulations provides essential information for designing deep‐sea preserves in regions that are under consideration for submarine mining of precious metals.     

Pliocene climatic change in insular Southeast Asia as an engine of diversification in Ficedula flycatchers

Aims Insular Southeast Asia and adjacent regions are geographically complex, and were dramatically affected by both Pliocene and Pleistocene changes in climate, sea level and geology. These circumstances allow the testing of several biogeographical hypotheses regarding species distribution patterns and phylogeny. Avian species in this area present a challenge to biogeographers, as many are less hindered by barriers that may block the movements of other species. Widely distributed Southeast Asian avian lineages, of which there are many, have been generally neglected. Ficedula flycatchers are distributed across Eurasia, but are most diverse within southern Asia and Southeast Asian and Indo‐Australian islands. We tested the roles of vicariance, dispersal and the evolution of migratory behaviours as mechanisms of speciation within the Ficedula flycatchers, with a focus on species distributed in insular Southeast Asia. Methods Using a published molecular phylogeny of Ficedula flycatchers, we reconstructed ancestral geographical areas using dispersal vicariance analysis, weighted ancestral area analysis, and a maximum likelihood method. We evaluated the evolution of migratory behaviours using maximum likelihood ancestral character state reconstruction. Speciation timing estimates were calculated via local molecular clock methods. Results Ficedula originated in southern mainland Asia, c. 6.5 Ma. Our analyses indicate that two lineages within Ficedula independently and contemporaneously colonized insular Southeast Asia and Indo‐Australia, c. 5 Ma. The potential impact of vicariance due to rising sea levels is difficult to assess in these early colonization events because the ancestral areas to these clades are reconstructed as oceanic islands. Within each of these clades, inter‐island dispersal was critical to species’ diversification across oceanic and continental islands. Furthermore, Pliocene and Pleistocene climatic change may have caused the disjunct island distributions between several pairs of sister taxa. Both vicariance and dispersal shaped the distributions of continental species. Main conclusions This study presents the first evaluation, for Ficedula, of the importance of vicariance and dispersal in shaping distributions, particularly across insular Southeast Asia and Indo‐Australia. Although vicariant speciation may have initially separated the island clades from mainland ancestors, speciation within these clades was driven primarily by dispersal. Our results contribute to the emerging body of literature concluding that dynamic geological processes and climatic change throughout the Pliocene and Pleistocene have been important factors in faunal diversification across continental and oceanic islands.     

Multiple overseas dispersal in amphibians

Amphibians are thought to be unable to disperse over ocean barriers because they do not tolerate the osmotic stress of salt water. Their distribution patterns have therefore generally been explained by vicariance biogeography. Here, we present compelling evidence for overseas dispersal of frogs in the Indian Ocean region based on the discovery of two endemic species on Mayotte. This island belongs to the Comoro archipelago, which is entirely volcanic and surrounded by sea depths of more than 3500 m. This constitutes the first observation of endemic amphibians on oceanic islands that did not have any past physical contact to other land masses. The two species of frogs had previously been thought to be nonendemic and introduced from Madagascar, but clearly represent new species based on their morphological and genetic differentiation. They belong to the genera Mantidactylus and Boophis in the family Mantellidae that is otherwise restricted to Madagascar, and are distinguished by morphology and mitochondrial and nuclear DNA sequences from mantellid species occurring in Madagascar. This discovery permits us to update and test molecular clocks for frogs distributed in this region. The new calibrations are in agreement with previous rate estimates and indicate two further Cenozoic transmarine dispersal events that had previously been interpreted as vicariance: hyperoliid frogs from Africa to Madagascar (Heterixalus) and from Madagascar to the Seychelles islands (Tachycnemis). Our results provide the strongest evidence so far that overseas dispersal of amphibians exists and is no rare exception, although vicariance certainly retains much of its importance in explaining amphibian biogeography.     

Why be amphidromous: expatrial dispersal and the place of source and sink population dynamics?

Amphidromous fishes are found predominantly on the tropical and subtropical islands of the globe and there are few amphidromous species on continents. I suggest that this idiosyncratic distribution relates in part to problems in self-recruitment on islands that are often young or volcanic, and which may have streams with ephemeral flows across relatively short times scales. Amphidromy provides the ability to invade new habitats as these become available either on newly emergent (often volcanic) islands, or following perturbation after stream dewatering or the impacts of volcanism on older islands as a consequence of expatrial dispersal. Source/sink population dynamics may also be involved with islands ‘downstream’ in oceanic current systems behaving as sinks, with little or no self-recruitment. Streams in steep topography seem to be favoured by amphidromous species, perhaps because they provide more rapid transport to sea of the tiny, newly hatched larvae.     

Towards a ‘Sea‐Level Sensitive’ dynamic model: impact of island ontogeny and glacio‐eustasy on global patterns of marine island biogeography

A synthetic model is presented to enlarge the evolutionary framework of the General Dynamic Model (GDM) and the Glacial Sensitive Model (GSM) of oceanic island biogeography from the terrestrial to the marine realm. The proposed ‘Sea‐Level Sensitive’ dynamic model (SLS) of marine island biogeography integrates historical and ecological biogeography with patterns of glacio‐eustasy, merging concepts from areas as diverse as taxonomy, biogeography, marine biology, volcanology, sedimentology, stratigraphy, palaeontology, geochronology and geomorphology. Fundamental to the SLS model is the dynamic variation of the littoral area of volcanic oceanic islands (defined as the area between the intertidal and the 50‐m isobath) in response to sea‐level oscillations driven by glacial–interglacial cycles. The following questions are considered by means of this revision: (i) what was the impact of (global) glacio‐eustatic sea‐level oscillations, particularly those of the Pleistocene glacial–interglacial episodes, on the littoral marine fauna and flora of volcanic oceanic islands? (ii) What are the main factors that explain the present littoral marine biodiversity on volcanic oceanic islands? (iii) How can differences in historical and ecological biogeography be reconciled, from a marine point of view? These questions are addressed by compiling the bathymetry of 11 Atlantic archipelagos/islands to obtain quantitative data regarding changes in the littoral area based on Pleistocene sea‐level oscillations, from 150 thousand years ago (ka) to the present. Within the framework of a model sensitive to changing sea levels, we discuss the principal factors affecting the geographical range of marine species; the relationships between modes of larval development, dispersal strategies and geographical range; the relationships between times of speciation, modes of larval development, ecological zonation and geographical range; the influence of sea‐surface temperatures and latitude on littoral marine species diversity; the effect of eustatic sea‐level changes and their impact on the littoral marine biota; island marine species–area relationships; and finally, the physical effects of island ontogeny and its associated submarine topography and marine substrate on littoral biota. Based on the SLS dynamic model, we offer a number of predictions for tropical, subtropical and temperate volcanic oceanic islands on how rates of immigration, colonization, in‐situ speciation, local disappearance, and extinction interact and affect the marine biodiversity around islands during glacials and interglacials, thus allowing future testing of the theory.     

Biogeographical affinities of the New Caledonian biota: a puzzle with 24 pieces

Aim The distributions of many New Caledonian taxa were reviewed in order to ascertain the main biogeographical connections with other areas. Location Global. Methods Panbiogeographical analysis. Results Twenty‐four areas of endemism (tracks) involving New Caledonia and different areas of Gondwana, Tethys and the central Pacific were retrieved. Most are supported by taxa of lower and higher plants, and lower and higher animals. Main conclusions Although parts of New Caledonia were attached to Gondwana for some time in the mid‐Cretaceous, most of the New Caledonian terranes formed as oceanic island arcs and sections of sea floor bearing seamounts. The flora and fauna have evolved and survived for tens of millions of years as metapopulations on ephemeral islands. Later, the biotas were juxtaposed and fused during terrane accretion. This process, together with the rifting of Gondwana, explains the biogeographical affinities of New Caledonia with parts of Gondwana, Tethys and the Pacific.     

Unravelling the peculiarities of island life: vicariance, dispersal and the diversification of the extinct and extant giant Galápagos tortoises

In isolated oceanic islands, colonization patterns are often interpreted as resulting from dispersal rather than vicariant events. Such inferences may not be appropriate when island associations change over time and new islands do not form in a simple linear trend. Further complexity in the phylogeography of ocean islands arises when dealing with endangered taxa as extinctions, uncertainty on the number of evolutionary ‘units’, and human activities can obscure the progression of colonization events. Here, we address these issues through a reconstruction of the evolutionary history of giant Galápagos tortoises, integrating DNA data from extinct and extant species with information on recent human activities and newly available geological data. Our results show that only three of the five extinct or nearly extinct species should be considered independent evolutionary units. Dispersal from mainland South America started at approximately 3.2 Ma after the emergence of the two oldest islands of San Cristobal and Española. Dispersal from older to younger islands began approximately 1.74 Ma and was followed by multiple colonizations from different sources within the archipelago. Vicariant events, spurred by island formation, coalescence, and separation, contributed to lineage diversifications on Pinzón and Floreana dating from 1.26 and 0.85 Ma. This work provides an example of how to reconstruct the history of endangered taxa in spite of extinctions and human‐mediated dispersal events and highlights the need to take into account both vicariance and dispersal when dealing with organisms from islands whose associations are not simply explained by a linear emergence model.     

THE RELEVANCE OF GENE FLOW IN METAPOPULATION DYNAMICS OF AN OCEANIC ISLAND ENDEMIC,OLEA EUROPAEA SUBSP. GUANCHICA

Theoretical and empirical studies suggest that geographical isolation and extinction–recolonization dynamics are two factors causing strong genetic structure in metapopulations, but their consequences in species with high dispersal abilities have not been tested at large scales. Here, we investigated the effect of population age structure and isolation by distance in the patterns of genetic diversity in a wind‐pollinated, zoochorous tree (Olea europaea subsp. guanchica) sporadically affected by volcanic events across the Canarian archipelago. Genetic variation was assessed at six nuclear microsatellites (nDNA) and six chloroplast fragments (cpDNA) in nine subpopulations sampled on four oceanic islands. Subpopulations occurring on more recent substrates were more differentiated than those on older substrates, but within‐subpopulation genetic diversity was not significantly different between age groups for any type of marker. Isolation‐by‐distance differentiation was observed for nDNA but not for cpDNA, in agreement with other metapopulation studies. Contrary to the general trend for island systems, between‐island differentiation was extremely low, and lower than differentiation between subpopulations on the same island. The pollen‐to‐seed ratio was close to one, two orders of magnitude lower than the average estimated for other wind‐pollinated, animal‐dispersed plants. Our results showed that population turnover and geographical isolation increased genetic differentiation relative to an island model at equilibrium, but overall genetic structure was unexpectedly weak for a species distributed among islands. This empirical study shows that extensive gene flow, particularly mediated by seeds, can ameliorate population subdivision resulting from extinction–recolonization dynamics and isolation by distance.     

Lauraceae fossils from a volcanic Palaeocene oceanic island,Ninetyeast Ridge,Indian Ocean: ancient long‐distance dispersal?

Aim Geological and fossil records are critical for historical biogeography studies. A plant fossil assemblage from a small, well‐dated, transient late Palaeocene island was re‐investigated with regard to regional geology and vicariance versus dispersal hypotheses. Location Deep Sea Drilling Program Leg 22, Site 214 on the Ninetyeast Ridge (NER) in the mid‐Indian Ocean region. Methods Leaf cuticular material was recovered from residues from a previous palynofloral study of Site 214 sediments during the 1970s and identified. The palynoflora was reassessed. Results The only leaf cuticular material recovered with stomata can be placed in crown‐group Lauraceae. It is confirmed that the palynoflora reflects the presence of a low‐diversity island flora in the late Palaeocene, comprising ferns and mostly herbaceous angiosperms with readily dispersible propagules, and perhaps austral podocarps. Other pollen taxa of almost certain local origin were arecoid palms and taxa related to Chloranthaceae. The strong overall similarity of the palynoflora to Australo‐Antarctic and New Zealand assemblages is also confirmed. Main conclusions Foliar fossils of Lauraceae demonstrate the occurrence of one of the world’s largest, most widely distributed woody plant families on a late Palaeocene island. The presence of plants on this island could be explained by vicariance via a vegetated Upper Cretaceous Kerguelen Plateau, in part because crown‐group Lauraceae may be at least this old. However, there are records of other taxa in the Kerguelen region that are anomalous with vicariance, plus evidence for a catastrophic biotic extinction event centred in the area in the latest Cretaceous. Plants were therefore most likely to have reached the island by means of dispersal. This suggests either the presence of presently unknown vegetated land nearby in the Kerguelen region in the late Palaeocene, or long‐distance dispersal, probably from the Australian region. The dispersal of viable seeds could have been facilitated by birds or perhaps by ocean‐surface drift with or without the assistance of ocean‐going animals. The fossils allow that even small, short‐lived islands could have acted as ‘stepping stones’ for biotic interchange between Australia and Africa, and perhaps other regions.     

Changing perspectives on the biogeography of the tropical South Pacific: influences of dispersal, vicariance and extinction

Aim The biogeographical patterns and drivers of diversity on oceanic islands in the tropical South Pacific (TSP) are synthesized. We use published studies to determine present patterns of diversity on TSP islands, the likely sources of the biota on these islands and how the islands were colonized. We also investigate the effect of extinctions. Location We focus on oceanic islands in the TSP. Methods We review available literature and published molecular studies. Results Examples of typical island features (e.g. gigantism, flightlessness, gender dimorphism) are common, as are adaptive radiations. Diversity decreases with increasing isolation from mainland sources and with decreasing size and age of archipelagos, corresponding well with island biogeographical expectations. Molecular studies support New Guinea/Malesia, New Caledonia and Australia as major source areas for the Pacific biota. Numerous studies support dispersal‐based scenarios, either over several 100 km (long‐distance dispersal) or over shorter distances by island‐hopping (stepping stones) and transport by human means (hitch‐hiking). Only one vicariance explanation, the eastward drift of continental fragments (shuttles) that may have contributed biota to Fiji from New Caledonia, is supported by some geological evidence, although there is no evidence for the transport of taxa on shuttle fragments. Another vicariance explanation, the existence of a major continental landmass in the Pacific within the last 100 Myr (Atlantis theory), receives little support and appears unlikely. Extinction of lineages in source areas and persistence in the TSP has probably occurred many times and has resulted in misinterpretation of biogeographical data. Main conclusions Malesia has long been considered the major source region for the biota of oceanic islands in the TSP because of shared taxa and high species diversity. However, recent molecular studies have produced compelling support for New Caledonia and Australia as alternative important source areas. They also show dispersal events, and not vicariance, to have been the major contributors to the current biota of the TSP. Past extinction events can obscure interpretations of diversity patterns.     

Vicariance and dispersal in the differentiation of vocalization in the Ryukyu Scops Owl Otus elegans

The distribution of species and species diversity can be affected by vicariance or dispersal. To understand their role in shaping species distribution and population structure these two processes must be estimated within and among populations. I analysed large‐scale variation in the call structure of the Ryukyu Scops Owl Otus elegans. This owl is distributed over a 1200‐km range, and only inhabits islands. Within this range, I studied this species across 22 continental islands of the Ryukyu Archipelago and two oceanic islands. The study aimed to assess whether there is variation in the acoustic structure of Owl hoot calls within islands, among major groups of islands and across a large area comprising a major biogeographical barrier (the Kerama Gap). The acoustic structure of calls was homogeneous within islands and among major island‐groups. Acoustic differentiation, however, increased over longer geographical distances of up to about 1200 km. The acoustic structure of hoots of the Ryukyu Scops Owl populations was clearly divided into two groups, north and south of the Kerama Gap. It is suggested that the Kerama Gap acted as a biogeographical barrier and contributed to the differentiation between the two major island‐groups. It is likely that this difference developed during the fragmentation of a widespread ancestral population by vicariant isolating events. There was also evidence of an effect of dispersal on vocal differentiation in subspecies inhabiting the two oceanic islands.     

Dispersal is fundamental to biogeography and the evolution of biodiversity on oceanic islands

Vicariance biogeography emerged several decades ago from the fusion of cladistics and plate tectonics, and quickly came to dominate historical biogeography. The field has since been largely constrained by the notion that only processes of vicariance and not dispersal offer testable patterns and refutable hypotheses, dispersal being a random process essentially adding only noise to a vicariant system. A consequence of this thinking seems to have been a focus on the biogeography of continents and continental islands, considering the biogeography of oceanic islands less worthy of scientific attention because, being dependent on stochastic dispersal, it was uninteresting. However, the importance of dispersal is increasingly being recognized, and here we stress its fundamental role in the generation of biodiversity on oceanic islands that have been created in situ , never connected to larger land masses. Historical dispersal patterns resulting in modern distributions, once considered unknowable, are now being revealed in many plant and animal taxa, in large part through the analysis of polymorphic molecular markers. We emphasize the profound evolutionary insights that oceanic island biodiversity has provided, and the fact that, although small in area, oceanic islands harbour disproportionately high biodiversity and numbers of endemic taxa. We further stress the importance of continuing research on mechanisms generating oceanic island biodiversity, especially detection of general, non-random patterns of dispersal, and hence the need to acknowledge oceanic dispersal as significant and worthy of research.     

Panbiogeography of New Caledonia,south‐west Pacific: basal angiosperms on basement terranes,ultramafic endemics inherited from volcanic island arcs and old taxa endemic to young islands

Aim To investigate areas of endemism in New Caledonia and their relationship with tectonic history. Location New Caledonia, south‐west Pacific. Methods Panbiogeographical analysis. Results Biogeographical patterns within New Caledonia are described and illustrated with reference to eight terranes and ten centres of endemism. The basement terranes make up a centre of endemism for taxa including Amborella, the basal angiosperm. Three of the terranes that accreted to the basement in the Eocene (high‐pressure metamorphic terrane, ultramafic nappe and Loyalty Ridge) have their own endemics. Main conclusions New Caledonia is not simply a fragment of Gondwana but, like New Zealand and New Guinea, is a complex mosaic of allochthonous terranes. The four New Caledonian basement terranes were all formed from island arc‐derived and arc‐associated material (including ophiolites) 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 break‐up of Gondwana. An Eocene collision of the basement terranes with an island arc to the north‐east – possibly the Loyalty Ridge – is of special biogeographical interest in connection with New Caledonia–central Pacific affinities. The Loyalty–Three Kings Ridge has had a separate history from that of the Norfolk Ridge/New Caledonia, although both now run in parallel between Vanuatu and New Zealand. The South Loyalty Basin opened between Grande Terre and the Loyalty Ridge in the Cretaceous and attained a width of 750 km. However, it was almost completely destroyed by subduction in the Eocene which brought the Loyalty Ridge and Grande Terre together again, after 30 Myr of separation. The tectonic history is reflected in the strong biogeographical differences between Grande Terre and the Loyalty Islands. Many Loyalty Islands taxa are widespread in the Pacific but do not occur on Grande Terre, and many Grande Terre/Australian groups are not on the Loyalty Islands. The Loyalty Islands are young (2 Myr old) but they are merely the currently emergent parts of the Loyalty Ridge whose ancestor arcs have a history of volcanism dating back to the Cretaceous. Old taxa endemic to the young Loyalty Ridge islands persist over geological time as a dynamic metapopulation surviving in situ on the individually ephemeral islands and atolls found around subduction zones. The current Loyalty Islands, like the Grande Terre terranes, have inherited their biota from previous islands. On Grande Terre, the ultramafic terrane was emplaced on Grande Terre in the Eocene (about the same time as the collision with the island arc). The very diverse endemic flora on the ultramafics may have been inherited by the obducting nappe from prior base‐rich habitat in the region, including the mafic Poya terrane and the limestones typical of arc and intraplate volcanic islands.     

Is a new paradigm emerging for oceanic island biogeography?

Following several decades during which two dissimilar and incompatible models (equilibrium and vicariance) dominated island biogeography, recent publications have documented patterns that point the way towards a new paradigm that includes elements of both models, as well as some novel aspects. Many of these seminal contributions have been made possible by the recent development of robust, temporally calibrated phylogenies used in concert with increasingly precise and reliable geological reconstructions of oceanic regions. Although a new general model of oceanic island biogeography has not yet been proposed, in this brief overview I present six hypotheses that summarize aspects of the emerging paradigm. These hypotheses deal with: the frequency of dispersal over oceanic water barriers by terrestrial organisms; the existence of substantial variation in the amount of dispersal (and gene flow) within a given set of related species within a given archipelago; the frequency, extent and impact on species richness of diversification within archipelagos; the frequent correlation of island age and the age of the species that live on the island; the long-term persistence of species on oceanic islands; and the occasional recolonization of continents by species from clades that diversified on islands. Identifying, testing, and seeking means of synthesizing these and other emerging hypotheses may allow a new conceptual paradigm to emerge.     

Mitochondrial DNA evolution and population history of the Tenerife skink Chalcides viridanus

Recent studies of island lizards have suggested that historical vicariance as a result of volcanism may have played an important role in shaping patterns of within-island genetic diversity. The skink, Chalcides viridanus, shows variation in morphology within the volcanic island of Tenerife. Two mitochondrial DNA (mtDNA) fragments (from the 12S and 16S rRNA regions) were sequenced in individuals from 17 sites to evaluate the relationship between current phylogeography and the geological history of the island. Three main clades were detected. The two most basal clades were restricted to areas representing the ancient precursor islands of Teno and Anaga in the northwest and northeast of Tenerife, respectively. The third clade showed a widespread geographical distribution and provided evidence of a recent rapid expansion after a bottleneck. Within-island cladogenesis appears to have taken place during a recent period of volcanic activity and long after the ancient islands had been united by the eruptions that led to the formation of the Canadas edifice. Evidence of similar biogeographical histories are found in other species in the Canary archipelago, supporting the volcanism scenario as a potentially widespread cause of within-island differentiation in reptiles.     

Seed plants of Fiji: an ecological analysis

An annotated list of indigenous Fijian seed plant genera is presented and comprises 484 genera and 1315 species in 137 families. The relative diversity of the largest families and genera in Fiji is indicated and compared with floras in New Caledonia and the Upper Watut Valley, Papua New Guinea. Differences and similarities appear to be due to biogeographical/phylogenetic factors rather than ecological differences or means of dispersal. Generic diversity for the seed plants as a whole is greatest between 0–100 m and decreases monotonically with altitude. However, in the largest family, Orchidaceae, maximum diversity occurs between 200–400 m. Fifty percent of the families are recorded from shore habitat. Twenty‐seven percent of the families and 80 species occur in or around mangrove, where the most diverse families are Orchidaceae, Rubiaceae, and the legumes. Some of the mangrove‐associate species are pantropical or Indo‐Pacific but most are locally or regionally endemic. Fifty‐six percent of the Fijian families are recorded on limestone. Twenty‐nine species are restricted to limestone and 12 species usually occur on limestone. The importance of calcium in reducing the effects of salinity is emphasized and 39 species are recorded from both mangrove and limestone. A plagiotropic habit occurs in 38 species which occur on limestone or around beaches, and 20 of these are Pacific endemics. Genera restricted to higher altitudes include many present elsewhere in Melanesia but absent from Australia despite suitable habitat there, again indicating the importance of biogeographical and historical factors. Altitudinal anomalies in Fiji taxa are cited and include 7 anomalously high records from northern Viti Levu, a site of major uplift, and 22 anomalously low altitudinal records in the Lau Group, a site of subsidence. It is suggested that the Fijian flora has not been derived from immigrants from Asia, but has evolved more or less in situ. Taxa would have survived as metapopulations on the individually ephemeral volcanic islands always found at oceanic subduction zones and hot spots, and the atolls which characterize areas of subsidence. The complex geology of Fiji is determined by its position between two subduction zones of opposite polarity, the Vanuatu and Tonga Trenches, in what is currently a region of transform faulting. The large islands comprise fragments of island arcs that have amalgamated and welded together. There has been considerable uplift as well as subsidence in the islands and it is suggested that both these processes have had drastic effects on the altitudinal range of the taxa. Limestone and mangrove floras could have provided a widespread, diverse ancestral species pool from which freshwater swamp forest, lowland rainforest, dry forest, secondary forest, thickets, and montane forest have been derived during phases of uplift. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 407–431.     

The resurrection of oceanic dispersal in historical biogeography

Geographical distributions of terrestrial or freshwater taxa that are broken up by oceans can be explained by either oceanic dispersal or vicariance in the form of fragmentation of a previously contiguous landmass. The validation of plate-tectonics theory provided a global vicariance mechanism and, along with cladistic arguments for the primacy of vicariance, helped create a view of oceanic dispersal as a rare phenomenon and an explanation of last resort. Here, I describe recent work that suggests that the importance of oceanic dispersal has been strongly underestimated. In particular, molecular dating of lineage divergences favors oceanic dispersal over tectonic vicariance as an explanation for disjunct distributions in a wide variety of taxa, from frogs to beetles to baobab trees. Other evidence, such as substantial gene flow among island populations of Anolis lizards, also indicates unexpectedly high frequencies of oceanic dispersal. The resurrection of oceanic dispersal is the most striking aspect of a major shift in historical biogeography toward a more even balance between vicariance and dispersal explanations. This new view implies that biotas are more dynamic and have more recent origins than had been thought previously. A high frequency of dispersal also suggests that a fundamental methodological assumption of many biogeographical studies--that vicariance is a priori a more probable explanation than dispersal--needs to be re-evaluated and perhaps discarded.     

Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago

We analysed the phylogenetic relationships of ten of the 13 known species of the genus Apomys using DNA sequences from cytochrome b . Apomys, endemic to oceanic portions of the Philippine archipelago, diversified during the Pliocene as these oceanic islands arose de novo . Several of the speciation events probably took place on Luzon or Mindanao, the two largest, oldest, and most topographically complex islands. Only one speciation event is associated with vicariance due to Pleistocene sea-level fluctuation, and a Pleistocene diversification model in which isolation is driven by sea-level changes is inconsistent with the data. Tectonic vicariance is nearly absent from the Philippines, in which tectonic coalescence plays a significant role. Most speciation events (about two-thirds) are associated with dispersal to newly developed oceanic islands. The data imply that the species have persisted for long periods, measured in millions of years after their origins; further implications therefore are that faunal turnover is very slow, and persistence over geological time spans is more prominent than repeated colonization and extinction. Neither the equilibrium nor the vicariance model of biogeography adequately encompasses these results; a model incorporating colonization, extinction, and speciation is necessary and must incorporate long-term persistence to accommodate our observations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 699–715.     

Biodiversity growth on the volcanic ocean islands and the roles of in situ cladogenesis and immigration: case with the reptiles

Models for biodiversity growth on the remote oceanic islands assume that in situ cladogenesis is a major contributor. To test this, we compiled occurrence data for 194 terrestrial reptile species on 53 volcanically‐constructed middle‐ to low‐latitude landmasses worldwide. Despite 273 native island‐species records, there are only 8–12 cases of the phenomenon, including just two radiations. Diversification frequencies are largely uncorrelated with island area, age, maximum altitude, and isolation. Furthermore, there is no indication that the presence of non‐sister congeners on an island stymies the process. Diversity on individual oceanic islands therefore results primarily from immigration and anageneis, but this is not a simple matter. Clusters that are difficult to reach (far or challenging to get to) or thrive upon (e.g. Canaries, Galápagos) have relatively few clades (3–8), some of which have many species (6–14), and all host at least one endemic genus. In these settings, diversity grows mainly by intra‐archipelago transfer followed by within‐island anagenetic speciation. In contrast, those island groups that are easier to disperse to (characterized by short distances and conducive transit conditions) and harbour more benign habitats (e.g. Comoros, Lesser Antilles) have been settled by many ancestor‐colonizers (≥ 14), but each clade has few derived species (≤ 4). These archipelagoes lack especially distinctive lineages. Models explaining the assembly and growth of terrestrial biotic suites on the volcanic ocean islands thus need to accommodate these new insights.