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.     

A conceptual model of ecosystem restoration triage based on experiences from three remote oceanic islands

A conceptual model, that illustrates restoration, ecological landscaping, rehabilitation and regreening, is developed. It considers biocentric, historical, aesthetic and engineering aspects. The term ecosystem restoration triage is used because the first step is to decide whether to do nothing (because, on the one hand, the system is too degraded to warrant restoration, or, on the other, because biological integrity is relatively intact and therefore either none, or minimal, restoration is required) or to do something (because restoration is worthwhile, urgent and feasible). This approach hinges on the definition that restoration in the strictist sense is a biocentric activity that returns the original compositional, structural and functional diversity, along with its dynamics and natural evolutionary potential. Original is a difficult qualifier as it depends on just how far back in time we go. Where human values are involved, this is not restoration in the pure sense of restoring ecological integrity, but is ecological landscaping, rehabilitation or regreening. Experience from three remote oceanic islands [Easter Island, Cousine Island (Seychelles), Marion Island (Sub-Antarctic)] and which represent near extremes of this model are used to illustrate it.     

The ecological impact of oceanic island colonization – a palaeoecological perspective from the Azores

Aim In many cases, human colonization drastically modified the ecosystems of remote oceanic islands before scientists arrived to document the changes. Palaeoecological records before and after human colonization provide insights into the original ecosystems and an assessment of subsequent human impact. We used pollen analysis to compare the impact of 15th century colonization of the Azores with that of natural disturbances such as volcanic eruptions and climate changes. Location Azores archipelago, Atlantic Ocean. Methods Sediment records from three highland sites in the Azores (on the islands of Pico and Flores) were dated radiometrically and analysed palynologically. Pollen taxa were classified as native, endemic or introduced based on comparison with flora lists. Data were statistically zoned and temporal trends identified using detrended correspondence analysis. Results Human colonization of the Azores resulted in rapid, widespread, persistent vegetation changes on a scale unprecedented in the last 2700 years, detectable through the decline of dominant trees, the spread of grasses and fire‐tolerant species, the introduction of exotic plants, evidence for grazing and fire, and changes to soils and moisture availability. During the same period, volcanic eruptions appear to have had more localized impacts on the vegetation, lasting 500–1000 years and favouring endemic taxa. The effect of late Holocene climatic changes on the highland vegetation of the Azores seems to have been minor. Palaeoecological data indicate that at least two plant species went extinct on Pico after human colonization and that some plants regarded as introduced were almost certainly part of the original flora of the islands. Despite a consistent signal of human impact, compositional differences between Juniperus brevifolia communities on Pico and Flores remained after colonization. Main conclusions Human colonization had a greater impact on the pristine vegetation of Pico and Flores than climatic changes and volcanic activity during recent millennia. The similarity between post‐colonization changes on the Azores and other oceanic islands suggests a consistent pattern and scale to historical‐era human impact on otherwise pristine ecosystems. These characteristics could be used to further elaborate biogeographical theory and direct conservation efforts towards species that appear most susceptible to human activity.     

Palaeo‐islands as refugia and sources of genetic diversity within volcanic archipelagos: the case of the widespread endemic Canarina canariensis (Campanulaceae)

Geographical isolation by oceanic barriers and climatic stability has been postulated as some of the main factors driving diversification within volcanic archipelagos. However, few studies have focused on the effect that catastrophic volcanic events have had on patterns of within‐island differentiation in geological time. This study employed data from the chloroplast (cpDNA haplotypes) and the nuclear (AFLPs) genomes to examine the patterns of genetic variation in Canarina canariensis, an iconic plant species associated with the endemic laurel forest of the Canary Islands. We found a strong geographical population structure, with a first divergence around 0.8 Ma that has Tenerife as its central axis and divides Canarian populations into eastern and western clades. Genetic diversity was greatest in the geologically stable ‘palaeo‐islands’ of Anaga, Teno and Roque del Conde; these areas were also inferred as the ancestral location of migrant alleles towards other disturbed areas within Tenerife or the nearby islands using a Bayesian approach to phylogeographical clustering. Oceanic barriers, in contrast, appear to have played a lesser role in structuring genetic variation, with intra‐island levels of genetic diversity larger than those between‐islands. We argue that volcanic eruptions and landslides after the merging of the palaeo‐islands 3.5 Ma played key roles in generating genetic boundaries within Tenerife, with the palaeo‐islands acting as refugia against extinction, and as cradles and sources of genetic diversity to other areas within the archipelago.     

The age and origin of the Pacific islands: a geological overview

The Pacific Ocean evolved from the Panthalassic Ocean that was first formed ca 750 Ma with the rifting apart of Rodinia. By 160 Ma, the first ocean floor ascribed to the current Pacific plate was produced to the west of a spreading centre in the central Pacific, ultimately growing to become the largest oceanic plate on the Earth. The current Nazca, Cocos and Juan de Fuca (Gorda) plates were initially one plate, produced to the east of the original spreading centre before becoming split into three. The islands of the Pacific have originated as: linear chains of volcanic islands on the above plates either by mantle plume or propagating fracture origin, atolls, uplifted coralline reefs, fragments of continental crust, obducted portions of adjoining lithospheric plates and islands resulting from subduction along convergent plate margins. Out of the 11 linear volcanic chains identified, each is briefly described and its history summarized. The geology of 10 exemplar archipelagos (Japan, Izu-Bonin, Palau, Solomons, Fiji, New Caledonia, New Zealand, Society, Galápagos and Hawaii) is then discussed in detail.     

Minimizing energy expenditure facilitates vertebrate persistence on oceanic islands

The characteristics of terrestrial vertebrates on oceanic islands are examined. They often include a reduced body size, a tolerance of conspecifics, flightlessness, a reduced basal rate of metabolism, and a propensity to enter torpor. On oceanic islands ectotherms frequently replace endotherms. These changes reduce the energy expenditure and resource requirements of vertebrates. Such reductions are permitted by the absence of mammalian predators and facilitate the survival of island endemics in the face of a restricted resource base and a variable environment through an increase in population size. Some insular species increase body size, but this occurs only when the resource base is large, due either to a fortuitously abundant resource, or to the absence of other species that exploit normally abundant resources. Some questions are posed to guide future work. They examine of the characteristics that permit species to disperse over water barriers, the conditions that require a reduction in resource use, the rapidity of response by immigrants to island conditions, whether supertramps show physiological differentiation with respect to island distance or size, and whether island size is absolute or relative to the size of the immigrants.     

Biodiversity hotspots through time: an introduction

International targets set for reducing the rate of biodiversity loss--the 2010 target--and ensuring environmental stability (Millennium Development Goals) have helped to focus the efforts of the scientific community on providing the data necessary for their implementation. The urgency of these goals, coupled with the increased rate of habitat alteration worldwide, has meant that actions have largely not taken into account the increasing body of data about the biodiversity change in the past. We know a lot about how our planet has been altered and recovered in the past, both in deep time and through prehistory. Linking this knowledge to conservation action has not been widely practised, by either the palaeoecology or the conservation communities. Long-term data, however, have much to offer current conservation practice, and in the papers for this volume we have tried to bring together a variety of different perspectives as to how this might happen in the most effective way. We also identify areas for productive collaboration and some key synergies for work in the near future to enable our knowledge of the past to be used for conservation action in the here and now. Lateral thinking, across knowledge systems and with open-mindness about bridging data gaps, will be necessary for our accumulating knowledge about our planet's past to be brought to bear on our attempts to conserve it in the future.     

IUCN Red List assessment of the Cape Verde endemic flora: towards a global strategy for plant conservation in Macaronesia

We review the conservation status and threats to the endemic vascular flora of the Cape Verde islands, mostly based on the past two decades of collecting, literature review and herbarium specimens. The application of IUCN Red List criteria and categories using RAMAS software reveals that 78% of the endemic plants are threatened (29.3% Critically Endangered, 41.3% Endangered, 7.6% Vulnerable). Most of these endemics have a limited geographical range, and half of them have Areas of Occupancy and Extents of Occurrence of < 20 and 200 km², respectively. Our data show that, over the last two decades, the Cape Verde vascular plants have become more threatened and their conservation status has declined, mostly as a consequence of the increase in exotic species, habitat degradation and human disturbance. This paper presents the first comprehensive IUCN Red List data review for the plants endemic to Cape Verde, thus providing an important step towards the recognition and conservation of its threatened endemic flora at the national and global level. It also fills a knowledge gap, as it represents the first thorough assessment of the conservation status of the entire endemic flora of a Macaronesian archipelago.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

Evolution on oceanic islands: molecular phylogenetic approaches to understanding pattern and process

By their very nature oceanic island ecosystems offer great opportunities for the study of evolution and have for a long time been recognized as natural laboratories for studying evolution owing to their discrete geographical nature and diversity of species and habitats. The development of molecular genetic methods for phylogenetic reconstruction has been a significant advance for evolutionary biologists, providing a tool for answering questions about the diversity among the flora and fauna on such islands. These questions relate to both the origin and causes of species diversity both within an archipelago and on individual islands. Within a phylogenetic framework one can answer fundamental questions such as whether ecologically and/or morphologically similar species on different islands are the result of island colonization or convergent evolution. Testing hypotheses about ages of the individual species groups or entire community assemblages is also possible within a phylogenetic framework. Evolutionary biologists and ecologists are increasingly turning to molecular phylogenetics for studying oceanic island plant and animal communities and it is important to review what has been attempted and achieved so far, with some cautionary notes about interpreting phylogeographical pattern on oceanic islands.     

When North and South don’t mix: genetic connectivity of a recently endangered oceanic cycad,Cycas micronesica,in Guam using EST‐microsatellites

Subject to environmental changes and recurrent isolation in the last ca. 250 Ma, cycads are often described as relicts of a previously common lineage, with populations characterized by low genetic variation and restricted gene flow. We found that on the island of Guam, the endemic Cycas micronesica has most of the genetic variation of 14 EST‐microsatellites distributed within each of 18 genetic populations, from 24 original sampling sites. There were high levels of genetic variation in terms of total number of alleles and private alleles, and moderate levels of inbreeding. Restricted but ongoing gene flow among populations within Guam reveals a genetic mosaic, probably more typical of cycads than previously assumed. Contiguous cycad populations in the north of Guam had higher self‐recruitment rates compared to fragmented populations in the south, with no substantial connection between them except for one population. Guam’s genetic mosaic may be explained by the influence of forest continuity, seed size, edaphic differences, and human transport of cycads. Also important are the extent of synchrony among flushes of reproductive female seed‐bearing sporophylls and restricted pollen movement by an obligate mutualist and generalist insects. An NADH EST‐locus under positive selection may reflect pressure from edaphic differences across Guam. This and three other loci are ideal candidates for ecological genomic studies. Given this species’ vulnerability due to the recent introduction of the cycad aulacaspis scale, we also identify priority populations for ex situ conservation, and provide a genetic baseline for understanding the effects of invasive species on cycads in the Western Pacific, and islands in general.     

Northern Islands,human error,and environmental degradation: A view of social and ecological change in the Medieval North Atlantic

Between ca. 790 and 1000 AD, Scandinavian settlers occupied the islands of the North Atlantic: Shetland, the Orkneys, the Hebrides, the Faroes, Iceland, and Greenland. These offshore islands initially supported stands of willow, alder, and birch, and a range of non-arboreal species suitable for pasture for the imported Norse domestic animals. Overstocking of domestic animals, fuel collection, ironworking, and construction activity seems to have rapidly depleted the dwarf trees, and several scholars argue that soil erosion and other forms of environmental degradation also resulted from Norse landuse practices in the region. Such degradation of pasture communities may have played a significant role in changing social relationships and late medieval economic decline in the western tier colonies of Iceland and Greenland. This paper presents simple quantified models for Scandinavian environmental impact in the region, and suggests some sociopolitical causes for ultimately maladaptive floral degradation.     

Immigration, species radiation and extinction in a highly diverse songbird lineage: white-eyes on Indian Ocean islands

Molecular phylogenetic hypotheses of species-rich lineages in regions where geological history can be reliably inferred may provide insights into the scale of processes driving diversification. Here we sample all extant or recently extinct white-eye (Zosterops) taxa of the southwest Indian Ocean, combined with samples from all principal continental lineages. Results support a high dispersal capability, with at least two independent continental sources for white-eyes of the region. An early (within 1.8 million years ago) expansion into the Indian Ocean may have originated either from Asia or Africa; the three resulting lineages show a disparate distribution consistent with considerable extinction following their arrival. Africa is supported as the origin of a later expansion into the region (within 1.2 million years ago). On two islands, a pair of Zosterops species derived from independent immigrations into the Indian Ocean co-occur or may have formerly co-occurred, providing strong support for their origin by double-island colonization rather than within-island (sympatric or microallopatric) speciation. On Mauritius and La Réunion, phylogenetic placement of sympatric white-eyes allow us to rule out a scenario in which independent within-island speciation occurred on both islands; one of the species pairs must have arisen by double colonization, while the other pair is likely to have arisen by the same mechanism. Long-distance immigration therefore appears to be responsible for much of the region's white-eye diversity. Independent immigrations into the region have resulted in lineages with mutually exclusive distributions and it seems likely that competition with congeneric species, rather than arrival frequency, may limit present-day diversity.     

First discovery of Stephanus Jurine (Hymenoptera: Stephanidae) in Japan,with description of a new species from Anijima Island of Ogasawara Islands

The genus Stephanus Jurine (in Panzer) is newly recorded from Japan and is represented by a new species. S. anijimensis sp. nov. from Anijima Island of the Ogasawara Islands. This species resembles S. tridentatus van Achterberg & Yang from the Oriental part of China, but can be distinguished by the combination of following characters: pronotum with a cavity under pronotal fold (cavity absent in S. tridentatus); hind femur comparatively slender, 4.8–5.1 times as long as maximum width (about 4.1 times in S. tridentatus); apical 0.5 of hind femur black or blackish‐brown (more or less yellowish in S. tridentatus). This new species was not found from other islands of the Ogasawara Islands, in which fauna is being severely endangered mainly due to predation by the introduced green anoles. Recently, it was introduced into Anijima Island in March 2013 and thus the extinction risk of this species is very high on this island.