The composition and relationships of the marine molluscan fauna of the Pitcairn Islands

The recent expedition has doubled the number of marine molluscs known from the Pitcairn Islands. Over 80 taxa are recorded from Ducie and Pitcairn (both still poorly known), 240 from Oeno and 320 from Henderson. A total exceeding 400 taxa is now known from the group as a whole. Most of this increase results from a vastly improved knowledge of the small taxa that had been neglected in earlier surveys. Important faunal differences exist between the four islands in the group which are related to the different character of each island. These totals still underestimate the true diversities, but they provide data that are comparable in quality to those from adjacent regions. In this context the molluscan fauna from the Pitcairn Islands is seen to be impoverished when compared to those from islands further west. Most of the fauna is composed of widespread Indo-West Pacific species, but there are several taxa that have more restricted ranges centred on S.E. Polynesia. A few, including some undescribed taxa discovered on the expedition, appear to be endemic. Significantly, many of these are characterized by non-planktotrophic larval development. Comparison of the modern fauna from Henderson with the fossil fauna from the Pleistocene reef on its plateau reveals important differences. About 25% of the fossil molluscs are currently unknown from the group and about 5% appear to be undescribed. These high rates of turnover demonstrate that the faunas are temporally unstable.     

Survey of the terrestrial and freshwater molluscan fauna of the Pilliga forest area in northern inland New South Wales,Australia

The Pilliga forest in northern inland New South Wales, Australia, is one of the largest surviving remnants of native forest on the western slopes of the Great Dividing Range. The Pilliga landscape is a challenging environment for molluscs, dominated by dry sclerophyll forest and with limited and largely ephemeral aquatic habitats. A field survey of the area in 2006–2012 identified a surprisingly rich and relatively intact aquatic native molluscan fauna with five species of bivalves in three families and nine species of freshwater gastropods (four families), including some rare species and range extensions. The native land snail fauna comprised 18 species (six families), including an unusually rich pupillid fauna with nine species. Some range extensions are recorded and some species are narrow-range endemics. The distributions of many aquatic and terrestrial species were correlated with geology or soil type. Introduced molluscs were predominantly found in anthropogenic habitats and include two freshwater gastropods (two families) and nine terrestrial snails and slugs (eight families). This study provides insight into the original molluscan fauna of the western slopes prior to landscape-scale agricultural development and provides a benchmark for future reference.     

Henderson Island prehistory: colonization and extinction on a remote Polynesian island

Situated at the extreme margin of the Indo-West Pacific biotic province, the four islands of the isolated Pitcairn Group hold interest for biogeographers and archaeologists alike. Human settlement may have been as early as the 8th century AD for the uplifted limestone island of Henderson, the most pristine island of its kind. An archaeological survey of the Pitcairn Islands is provided, while Henderson is examined in detail. Recent extensive excavations provide a record of change during 600 years of human occupation. Adaptation to the ecologically-marginal conditions is documented by artefacts, more than 150000 vertebrate bones, molluscs and subfossil plant remains recovered from stratigraphic contexts. The effects of prehistoric human occupation on the pristine environment are revealed by Polynesian plant and animal introductions, bird extinctions and range reductions, possible over-predation of marine molluscs, exploitation of sea turtles, and large-scale burning for swidden agriculture. The origin of human colonists is documented by analysing imported artefacts by geochemical characterization (x-ray fluorescence analysis). The human abandonment of Henderson, by the seventeenth century, is viewed in the context of prehistoric regional dynamics.     

The Pitcairn Islands, South Pacific Ocean: plate tectonic and climatic contexts

The remote Pitcairn Group in the South Pacific Ocean comprises a volcanic island (Pitcairn Island), two low coral atolls (Oeno, Ducie) and a raised coralline island (Henderson Island). The geological history of these islands, on anomalously thin oceanic lithosphere, is related to the development of two subparallel island chains (Oeno-Henderson-Ducie; Pitcairn) associated with intra-Pacific plate 'hotspot' activity; the surface manifestation of this activity has been partly determined by structural lineations in the plate inherited from past plate history. The climate of the Pitcairn Islands is determined by the position of the subtropical high pressure system and the South Pacific Convergence Zone. Variations in the strength of this atmospheric circulation system, measured by changes in the Southern Oscillation index of pressure difference, provide a partial explanation of the long-term variability of mean annual rainfall at Pitcairn Island. Knowledge of past climates in the Pitcairn Group remains speculative. Maps of the Pitcairn Islands and a report of climate at Henderson Island (2/91-1/92) are included in the paper.     

The flora of the Pitcairn Islands: a review

vascular plant flora of the Pitcairn Islands, south-central Pacific Ocean, is described based on extensive new collections made in 1991 and previously published records. Two vascular plants occur on Ducie Atoll; one (Pemphis acidula) is a new record. Sixty-three native vascular plants occur on Henderson, of which nine are endemic; Canavalia rosea, Operculina lurpethum, Psilotum nudum and Solanum americanum are new records for the island. Oeno Atoll has 16 native vascular plants; the single endemic (Bidens hendersonensis var. oenoensis) was not found in 1991 despite careful searches. Triumfetta procumbens was new for Oeno. Sixty-six native vascular plants have now been recorded from Pitcairn Island, there are two endemic ferns and seven endemic angiosperms in this number. A number of non-native taxa were new to Pitcairn. Some of the previously described taxa could not be found on Pitcairn, probably because they are very rare and only a small amount of time was spent collecting on Pitcairn. Many of the Pitcairn taxa are threatened by the spread of introduced species, especially Syzygium jambos.     

The phytogeographical affinities of the Pitcairn Islands – a model for south‐eastern Polynesia?

Aim To identify how the Pitcairn group relates biogeographically to the south‐eastern Polynesian region and if, as a subset of the regions flora, it can then be used as a model for biogeographical analyses. Location The Pitcairn group (25°4′ S, 130°06′ W) comprises four islands: Pitcairn, a relatively young, high volcanic Island; Henderson, an uplifted atoll, the uplift caused by the eruption of Pitcairn; and two atolls, Ducie and Oeno. The remote location, young age and range of island types found in the Pitcairn Island group makes the group ideal for the study of island biogeography and evolution. Methods A detailed literature survey was carried out and several data sets were compiled. Dispersal method, propagule number and range data were collected for each of the 114 species that occurs in the Pitcairn group, and environmental data was also gathered for islands in Polynesia. Analyses were carried out using non‐metric multidimensional scaling and clustering techniques. Results The flora of the Pitcairn Islands is derived from the flora of other island groups in the south‐eastern Polynesian region, notably those of the Austral, Society and Cook Islands. Species with a Pacific‐wide distribution dominate the overall Pitcairn group flora. However, each of the islands show different patterns; Pitcairn is dominated by species with Pacific, Polynesian and endemic distributions, with anemochory as the dominant dispersal method (39.5%); Henderson is also dominated by species with Pacific, Polynesian and endemic distributions, but zoochory is the dominant dispersal method (59.4); Oeno and Ducie are dominated by Pantropic species with hydrochory as the most common dispersal method (52.9% and 100%, respectively). Main conclusions ? Habitat availability is the most significant factor determining the composition and size of the flora. ? South‐east Polynesia is a valid biogeographical unit, and should include the Cook, Austral, Society, Marquesas, Gambier, Tuamotu and Pitcairn Islands with Rapa, but should exclude Easter Island, Tonga and Samoa. ? Regionalization schemes should take island type into consideration. ? The Pitcairn Island group can serve as a useful model for Pacific biogeographical analyses.     

Biodiversity and biogeography of Henderson Island's insects

Henderson Island is an example of a raised coral atoll, and as such supports a much more diverse biota than geographically similar low atolls. It is also, due to its remote location, largely undisturbed by the activities of humans. It is therefore unique in the Pacific Ocean, and comparable to Aldabra in the Indian Ocean. Biogeographically it is also interesting as it is amongst the most easterly islands of the Indo-West Pacific region, and has been largely colonized by species dispersing from island to island across the ocean from the west. New and extensive collections of insects were made in 1991. Although not fully identified yet, it is clear that the fauna is depauperate, with only about 180 species known to date. Despite its easterly location almost all species are derived from the west, and much of the fauna is likely to be indigenous. The most diverse orders are the Lepidoptera ( c. 53 spp.), Coleoptera ( c. 38 spp.), Diptera ( c. 37 spp.), Hymenoptera ( c. 21 spp.), Homoptera ( c. 14 spp.) and Heteroptera (4 spp.). Of particular interest are the weevils, Heteroptera and Homoptera which exhibit considerable endemism with perhaps some examples of intra-island radiation. Although depauperate, Henderson supports a much larger insect fauna than that of nearby Ducie Atoll (total known insect fauna =15 spp).     

Geomorphology of the uplifted Pleistocene atoll at Henderson Island, Pitcairn Group

Henderson Island, in the Pitcairn Group, preserves a Pleistocene atoll physiography with the rim of the raised reef structure, supporting spur and groove topography, enclosing a central lagoon. Excellent preservation of coral reef communities occurs along the ancient atoll rim and within the central lagoon. The previously interpreted depositional nature of the fossil atoll structure is herein corroborated with geomorphologic and stratigraphic evidence from previously un-visited portions of the island. Stratigraphic and lateral facies relationships indicate a physiographic zonation which includes spur and grooves, outer reef flat, lagoon margin, and an interior lagoon with patch reefs. The in situ occurrence and zonation of reef coral communities around the periphery and within the interior of the island appear to reflect the original physiography of the atoll lagoon, with the most pronounced reef development on the SE side of the original atoll. Stratigraphic units which comprise the raised atoll lagoon structure represent different time intervals, so the atoll lagoon structure formed during various sea level fluctuations. The modern atolls of the Pitcairn Group, Oeno and Ducie, provide some comparisons (similarities and differences) with the fossil lagoon on top of Henderson Island.     

Biodiversity and origin of the non-flying terrestrial arthropods of Henderson Island

Henderson Island, a raised coral island on the extreme south-easterly edge of the Indo-Pacific plate, is of great importance as one of the few examples of a Pacific island with intact lowland forest. It is also of biogeographic interest as it is practically the final island in a series of island chains, along which the fauna of Polynesia has colonized, reaching back to New Guinea. New collections of the non-flying terrestrial arthropod fauna were made on Henderson Island in 1991. In excess of 100 taxa are now known. There is a rich mite fauna (especially oribatids), with many apparently endemic, some 26 species of spider, and nine species of isopod (including three endemic to Henderson or nearby Ducie Atoll). In addition Diplopoda, Chilopoda, Amphipoda, Pseudoscorpiones, Diplura, Protura and Collembola are represented. The majority of the fauna is derived from the west, as expected, though many taxa appear to be introduced, some of them from the neotropics (e.g. Frigga crocuta and perhaps Hoplophorella stilifer).     

The breeding biology of the gadfly petrels Pterodroma spp. of the Pitcairn Islands: characteristics, population sizes and controls

This paper reports the breeding biology and nesting seasons of the gadfly petrels which nest on the four islands of the Pitcairn group, Pitcairn, Henderson, Oeno and Ducie. The species currently breeding are Murphy's petrel Pterodroma ultima , Kermadec petrel P. neglecta , Herald petrel P. heraldica and Henderson petrel P. atrata. Of these, Murphy's petrel is the most numerous; an estimated 250000 pairs bred on Ducie, which is probably the major breeding station of the species. Novel basic breeding data for Murphy's petrel are presented. Incubation spells, averaging 19.3 days, are exceptionally long for a petrel. Phoenix petrel P. alba appears to have ceased to breed on the Pitcairn Islands since the 1922 surveys of the Whitney Expedition. Nesting success was low on Henderson Island during the study. For all four breeding species, less than 20% of eggs laid yielded fledglings. Failure occurred at the early chick stage and observations indicated that it was due to predation by Pacific rats Rattus exulans. Although rats are present on Ducie, predation was apparently less severe there. The situation on Oeno may be intermediate. I consider how the populations of Henderson are maintained in the face of this intense predation. The Murphy's petrel population may be sustained by immigration from Ducie while the Herald and Henderson petrel populations could be undergoing a long-term decline on Henderson. It is not clear how the Kermadec petrel population is maintained. The conservation implications of these findings are discussed.     

Local completion of the pelagic larval stage of coastal fishes in coral-reef lagoons of the Society and Tuamotu Islands

In four lagoons at two atolls and one high island in the Tuamotu and Society Islands, French Polynesia, plankton samples were taken weekly during 4 weeks in January/February 1989. A third atoll lagoon was sampled once. The lagoons varied in size and physical openness. We also sampled in the ocean near two atolls and the high island. All locations were sampled during the day, and three lagoons (two atolls and one high island) were also sampled at night. Pelagic fish eggs were more abundant in the ocean than in the lagoons at the atolls, but not at the high island. Larvae of coastal fishes were abundant in all lagoons. In the atoll lagoons, larvae of oceanic fishes were very rare to absent, but in the high-island lagoons and in the ocean, they were commonly encountered. In the ocean, larvae of many typical reef-fish taxa were abundant (58 taxa were represented by at least 10 individuals), but in the lagoons, most of these were rare or absent, and we conclude that these rare and absent taxa normally do not complete their larval phase in lagoons.Taxa were considered to be able to complete their pelagic phase in a lagoon (i.e., were 'completers') if they were present in the lagoon plankton samples from across a full larval size range. In the high-island barrier-reef lagoon, young, preflexion larvae were abundant, but only two taxa (of 56 captured) were present over a wide size range and were considered completers in this lagoon. In the high-island lagoonal bay, 11 taxa (of 67 captured) were considered completers. The numbers of taxa captured in the three atoll lagoons ranged from 39–44, and the number of taxa considered to be completers increased with increasing lagoon size and physical openness. The 17 completer taxa in the smallest, most enclosed atoll lagoon were, with one exception, a subset of those (18) in the second lagoon which, in turn, with one exception, were a subset of those in the largest, most open lagoon (26). Completer taxa were of the families Apogonidae, Blenniidae, Bothidae, Callionymidae, Carangidae, Gobiidae, Microdesmidae, Mullidae, Pomacentridae, Schindleriidae, and Tetraodontidae. The species that can complete their pelagic periods in coral-reef lagoons are a highly predictable group, and not simply a random selection of the potential species pool. Most of these species hatch from non-pelagic eggs. Water renewal times in the atoll lagoons, unlike the high-island barrier-reef lagoon, were much longer than expected pelagic larval durations of completer taxa. Demographically, lagoon populations of completer taxa apparently self-recruit and are probably near the closed end of the open/closed population continuum. The lagoonal bay on the high island differs from the other lagoons in containing larvae of species not found elsewhere, including some completers, and lacking some species that are abundant completers in other lagoons. In French Polynesia, lagoon size is a strong predictor of the number of lagoon completer taxa. The number of completer taxa apparently peaks at intermediate lagoon water-exchange times.     

The Foraminifera of the Pitcairn Islands

Foraminifera were recovered from 18 samples collected in the Pitcairn Islands, 12 from Henderson Island (including the best and most comprehensive collections) and three each from Oeno Atoll and Pitcairn Island itself. Although both algae and sediment samples were collected, the living Foraminifera came, almost exclusively, from phytal (attached or clinging) habitats. Foraminifera in the sediment samples are mainly thanatocoenoses. The fauna is an exclusively calcareous, relatively low diversity assemblage, dominated by large soritids [Marginopora, Amphisorus, Sorites) and Amphistegina , all of which are ubiquitous throughout the tropical Pacific. These larger Foraminifera are usually accompanied by small miliolids in particular, as well as by small attached Foraminifera (discorbids and the like). Typical reefal Foraminifera are generally under-represented. So far, no endemic species have been found. Of more significance, perhaps, is the apparent absence of Calcarina , small rotaliids, elphidiids and agglutinating species, so common in the western Pacific islands. One sample of fossil Foraminifera was analysed, from a shelly sand (c. 30 m above present sea-level) on Henderson Island. Though, for the most part, like the Recent assemblages, this was characterized by Archaias , a soritid which was not found in any of the modern collections made by the 1991– 92 Expedition. This could either be a sampling artifact or refer to a real environmental change since the Pleistocene.     

Three detailed,radiocarbon-dated,Holocene tufa and alluvial fan mollusc successions from southern Ontario: The first in northeastern North America

Fossil mollusc remains, particularly those of land snails, provide a wealth of detailed information concerning local Quaternary paleoenvironments and regional paleoclimatic conditions. However, this important resource is generally overlooked in North America and consequently quantitative, well-dated terrestrial mollusc successions are extremely rare here. In an attempt to redress this imbalance, a project examining the Holocene molluscan faunal history of northeastern North America has been initiated. Thus far, detailed mollusc successions and associated radiocarbon age determinations have been obtained from three sites in southern Ontario; two alluvial fans (Chine Drive Fan and Cudia Park Fan, both from the Scarborough Bluffs east of Toronto) and a tufa (West River Road, near Cambridge). The observed land snail successions to provide an initial view of the molluscan faunal history of this region during the period from the late glacial to the mid Holocene, between approximately 10,500 and 5000 radiocarbon years BP (yr BP). The molluscan data have also been used to reconstruct the paleoenvironmental history of southern Ontario during this time, documenting the progression from exposed, open habitats characterised by species such as Vertigo modesta, Vallonia albula, Columella columella and Vitrina angelicae limpida, to closed woodland biotopes indicated by the presence of taxa including Carychium nannodes, Nesovitrea binneyana, Paravitrea multidenta and Helicodiscus parallelus. This study has already demonstrated the potential of this technique in the study of Quaternary climatic and environmental change in eastern North America and represents a strong foundation for extending the analysis of fossil land snail successions to other sites throughout this region. It has also provided new and important insights into the establishment of the modern molluscan fauna, which are essential to the understanding of present day distributional patterns and environmental associations.     

Reassembling island ecosystems: the case of Lord Howe Island

Exotic species that invade remote islands, usually following human settlement, have had catastrophic effects on native biota. However, on islands it is increasingly feasible to eradicate key exotic species allowing extant native species to recover in situ or to return naturally. The practice of marooning threatened species on islands where the threat is absent, irrespective of whether the threatened species once occurred on the island, is well established. However, less focus has been given to the 'island' as the management unit on which to return extirpated species or related surrogates for extinct species. We use the example of Lord Howe Island as a case study to explore options for island restoration should the remaining critical exotic pests (rodents and perhaps owls in this case) be eradicated as planned. Lord Howe Island, in the south-west Pacific Ocean, is remote, biologically diverse, has a high degree of endemism, and was only discovered by humans in 1778. Consequently, the original and exotic biota and their interactions are all better known than for many islands with a more ancient human history. Two species of plants, nine terrestrial birds, one bat and at least four invertebrates have been lost from the island since 1778. One plant and two invertebrates could be returned as conspecifics. One plant and all the terrestrial birds that are extinct could be replaced by closely related species from elsewhere in the Pacific Ocean. Decisions on replacing extinct species with surrogates should be based on the taxonomic relatedness of the candidates for reintroduction: the same species before subspecies before genera, with functional replacement being a further filter on candidates that are not the same species. In our opinion, taxa with functional equivalence but without taxonomic relatedness would not be acceptable candidates for reintroduction.     

Evaluating the efficacy of land snail survey techniques in Hawaii: implications for conservation throughout the Pacific

Terrestrial micromolluscs (snails with an adult maximum shell dimension <5 mm) constitute a considerable proportion of the land snail fauna of the Pacific. However, micromolluscs are often underestimated in biological surveys because of size bias. It has been argued that visual searches are preferable on Pacific islands because: (1) size biases are limited based on the understanding that most native Pacific island land snails are very small, and (2) amount of labor is less than other methods such as soil surveys and adequate for inventory purposes (though not for abundance assessments). To test whether visual surveys and soil surveys were accurately recording all taxa, land snail inventories were completed in three forest reserves (5 sampling sites in each) on the island of Oahu, Hawaii. Visual surveys involved 30-min visual search in a 10 m² site; soil surveys involved sieving leaf litter and topsoil from four 0.3 m² quadrats and extracting snails with the aid of a microscope. The data indicate a size and microhabitat bias associated with both techniques. Visual surveys consistently collected large arboreal and litter-dwelling species but missed a significant portion of micromolluscs, while soil surveys collected micromolluscs but missed larger snails. Because of such biases, employing both methods is critical for collecting all taxa at a survey location. As such, we recommend that future land snail surveys on Pacific Islands incorporate both survey techniques. Obtaining a complete inventory is critical if we are to understand species distributions and patterns of diversity and make well-informed conservation recommendations.     

Hawaiian biogeography and the islands' freshwater fish fauna

Aim This paper describes known patterns in the distributions and relationships of Hawaiian freshwater fishes, and compares these patterns with those exhibited by Hawaii's terrestrial biota. Location The study is based in Hawaii, and seeks patterns across the tropical and subtropical Indo‐west Pacific. Methods The study is based primarily on literature analysis. Results The Hawaiian freshwater fish fauna comprises five species of goby in five different genera (Gobiidae). Four species are Hawaiian endemics, the fifth shared with islands in the western tropical Pacific Ocean. All genera are represented widely across the Indo‐west Pacific. All five species are present on all of the major Hawaiian islands. All five species are amphidromous – their larval and early juvenile life being spent in the sea. Although there has been some local phyletic evolution to produce Hawaiian endemics, there has been no local radiation to produce single‐island endemics across the archipelago. Nor is there evidence for genetic structuring among populations in the various islands. Main conclusions In this regard, the freshwater fish fauna of Hawaii differs from the well‐known patterns of local evolution and radiation in Hawaiian Island terrestrial taxa. Amphidromy probably explains the biogeographical idiosyncrasies of the fish fauna – dispersal through the sea initially brought the fish species to Hawaii, and gene flow among populations, across the archipelago, has hitherto inhibited the evolution of local island endemics, apparently even retarding genetic structuring on individual islands.     

Using Mark-recapture to provide population census data for use in Red Listing of invertebrates: the rare terrestrial snail Prestonella bowkeri as a case study

Invertebrates in general, and terrestrial molluscs in particular, are greatly affected by the current biodiversity extinction crisis, and accurate assessment of these organisms for inclusion in Red Lists is essential for the conservation of these taxa. However, obtaining accurate population counts of molluscs for this purpose can be challenging. Using Prestonella bowkeri, a terrestrial snail up to 23 mm in length and 14 mm in width found in mesic rocky cliff face habitats on the Great Escarpment of southern Africa as an example, we show that the use of Mark-recapture techniques to obtain population census data is appropriate to terrestrial molluscs of this size and larger. While currently listed as vulnerable, our results indicate that there are as many as 4,108 individuals of this species in one population, which suggests that the population size (sensu the IUCN definition) is probably >20,000, and the species could be down-listed and classified as near threatened. However, when considering the problems associated with applying the IUCN Red List criteria to invertebrates, and the very small number of populations and hence area of occupancy of this species, down-listing would be premature.