Predation on endemic Hawaiian land snails by the invasive snail Oxychilus alliarius

The native land snails of the Hawaiian Islands have suffered numerous extinctions and continue to decline rapidly. A leading cause of these extinctions has been the impact of non-native predators. Oxychilus alliarius is a European predatory snail that has been in Hawaii since at least 1937 and has become one of the most abundant invasive snails in the islands. It is a facultative carnivore and will also consume non-animal food. However, this species has been mostly ignored as a threat and the majority of conservation attention has been focused on Euglandina rosea, the Rosy Wolf Snail. Here we report predation by O. alliarius on two rare and endemic Hawaiian snail species: Kaala subrutila (Helicarionidae) and Pleuropoma cf. sandwichiensis (Helicinidae). The wide distribution of O. alliarius in Hawaii, its high abundance, and now compelling evidence that it preys on native (and probably endangered) snails, indicate that it could be a much greater threat than previously thought.     

The cause of mollusk decline on the Ogasawara Islands

Decline of land snails on the Ogasawara Islands was studied. In Hahajima, major alien predators such as Euglandina rosea and Platydemus manokwari are not present, but some small endemic snails, for example, Hirasea spp. and Ogasawarana spp., are already rare and more common endemic snails, for example, Mandarina spp., are also declining in the northern mountains. The decline cannot be directly explained by forest deforestation and by its subsequent regeneration. Three species of flatworms were found to eat small snails under captive conditions. The distribution of these flatworms is restricted to the northern mountains of Hahajima where Mandarina is declining and its survival is low. These predators are plausible candidates as a cause of the decline of the endemic snails.     

On the extinction of the Dune Shearwater (Puffinus holeae) from the Canary Islands

Insular ecosystems have been subjected to severe hardship during the last millennia. Large numbers of insular bird species have undergone local disappearances and full extinctions, and a high number of insular birds are currently categorised as endangered species. In most of these cases, extinction—or endangerment—is in direct relation to the arrival of ‘aboriginal’ and/or imperialist waves of human settlement. Insular bird extinction events have been documented to have occurred at times corresponding to aboriginal settlement at many archipelagos and isolated islands, such as the Hawaiian Islands, New Zealand, the West Indies or the tropical Pacific Islands. However, no bird extinctions could be attributed to the first settlers of the Canary Islands—until now. The first accelerator mass spectrometer radiocarbon (¹⁴C) dating of collagen from a bone of the Dune Shearwater Puffinus holeae (3395 ± 30 year BP), an extinct bird from the Canary Islands, indicates a late Holocene extinction event. This relatively recent date, together with some features of this bird (large body size, breeding areas situated at very accessible places) and the absence of its bones from the entire archaeological record suggests that the extinction occurred close to the time that the first human settlement occurred on the islands.     

Prey preference and gregarious attacks by the invasive flatworm Platydemus manokwari

The flatworm Platydemus manokwari (Tricladida: Rhynchodemidae) preys on various species of land snail, and its introduction to areas outside of its native range is thought to have caused the extinction of native land snails on several Pacific islands. Platydemus manokwari occurs in areas where land snails have been absent since its invasion, suggesting that the flatworm can prey on animals other than land snails. To identify the alternative prey and prey preferences of P. manokwari, I examined the feeding ecology of P. manokwari under field and laboratory conditions. Individuals were observed attacking live earthworms in a forest where land snails are already extinct, on Chichijima, Ogasawara (Bonin) Islands, Japan. I also observed them attacking earthworms and a species of isopod and land snail in the lab. To prey on the worm, similar to other earthworm-eating flatworms, P. manokwari wrapped itself around the prey and fed on it by inserting its pharynx into the earthworm’s body. Large earthworms were able to escape P. manokwari attacks by autotomy, but the autotomized body parts were eaten. Several P. manokwari individuals together attacked earthworms that were larger than themselves. The laboratory experiments showed that such gregarious attacks increased predation success on both large earthworms and land snails. The flatworms also attacked the isopods, although the predation rate was low. These results demonstrate that P. manokwari is a polyphagous predator of slow-moving soil invertebrates (land snails, slugs, earthworms, and isopods) and that invasion by P. manokwari may directly and indirectly impact native soil fauna.     

Global Patterns and Drivers of Avian Extinctions at the Species and Subspecies Level

Birds have long fascinated scientists and travellers, so their distribution and abundance through time have been better documented than those of other organisms. Many bird species are known to have gone extinct, but information on subspecies extinctions has never been synthesised comprehensively. We reviewed the timing, spatial patterns, trends and causes of avian extinctions on a global scale, identifying 279 ultrataxa (141 monotypic species and 138 subspecies of polytypic species) that have gone extinct since 1500. Species extinctions peaked in the early 20^th century, then fell until the mid 20^th century, and have subsequently accelerated. However, extinctions of ultrataxa peaked in the second half of the 20^th century. This trend reflects a consistent decline in the rate of extinctions on islands since the beginning of the 20^th century, but an acceleration in the extinction rate on continents. Most losses (78.7% of species and 63.0% of subspecies) occurred on oceanic islands. Geographic foci of extinctions include the Hawaiian Islands (36 taxa), mainland Australia and islands (29 taxa), the Mascarene Islands (27 taxa), New Zealand (22 taxa) and French Polynesia (19 taxa). The major proximate drivers of extinction for both species and subspecies are invasive alien species (58.2% and 50.7% of species and subspecies, respectively), hunting (52.4% and 18.8%) and agriculture, including non-timber crops and livestock farming (14.9% and 31.9%). In general, the distribution and drivers of subspecific extinctions are similar to those for species extinctions. However, our finding that, when subspecies are considered, the extinction rate has accelerated in recent decades is both novel and alarming.     

Extinction or survival: partulid tree snails in American Samoa

Four partulid tree snail species are known from American Samoa. In 1998, we surveyed the recently established National Park (units on three islands: Tutuila, Tau, Ofu) and neighboring areas for partulids. On Tutuila, Samoana abbreviata, previously considered probably extinct, was extremely rare (15 snails seen); Samoana conica was more common (288 snails) but still rare; Eua zebrina was the most common (1102 snails), at one locality perhaps near its natural abundance. The species have similar distributions within the Park. All three have declined dramatically since the 1920s. Before 1980, when the predatory snail Euglandina rosea was introduced, habitat destruction, and perhaps rat predation and shell collecting, probably caused the decline. The Park provides protection to the Tutuila partulids by protecting habitat, although development is still a potential problem. But predation by E. rosea may yet cause their extinction. On Ofu (only outside the Park), 12 live Samoana thurstoni were found; 31 have now been recorded by western science. Also, an apparently robust population of E. zebrina, previously considered a Tutuila endemic, was found. Euglandina rosea is not on Ofu, so these populations are important remnants of the fauna. No partulids were found on Tau; none has ever been recorded there.     

Tahitian tree snail mitochondrial clades survived recent mass extirpation

Oceanic islands frequently support endemic faunal radiations that are highly vulnerable to introduced predators [1]. This vulnerability is epitomized by the rapid extinction in the wild of all but five of 61 described Society Islands partulid tree snails [2], following the deliberate introduction of an alien biological control agent: the carnivorous snail Euglandina rosea[3]. Tahiti's tree snail populations have been almost completely extirpated and three of the island's eight endemic Partula species are officially extinct, a fourth persisting only in captivity [2]. We report a molecular phylogenetic estimate of Tahitian Partula mitochondrial lineage survival calibrated with a 1970 reference museum collection that pre-dates the predator's 1974 introduction to the island [4]. Although severe winnowing of lineage diversity has occurred, none of the five primary Tahitian Partula clades present in the museum samples is extinct. Targeted conservation measures, especially of montane refuge populations, may yet preserve a representative sub-sample of Tahiti's endemic tree snail genetic diversity in the wild.     

Drivers of extinction: the case of Azorean beetles

Oceanic islands host a disproportionately high fraction of endangered or recently extinct endemic species. We report on species extinctions among endemic Azorean beetles following 97% habitat loss since AD 1440. We infer extinctions from historical and contemporary records and examine the influence of three predictors: geographical range, habitat specialization and body size. Of 55 endemic beetle species investigated (out of 63), seven can be considered extinct. Single-island endemics (SIEs) were more prone to extinction than multi-island endemics. Within SIEs restricted to native habitat, larger species were more extinction-prone. We thus show a hierarchical path to extinction in Azorean beetles: species with small geographical range face extinction first, with the larger bodied ones being the most threatened. Our study provides a clear warning of the impact of habitat loss on island endemic biotas.     

Characteristics of Pleistocene megafauna extinctions in Southeast Asia

The extinction of large-bodied taxa from the Pleistocene in Southeast Asia is examined. Although the chronological resolution of these extinctions is poor, and number of excavations in the region relatively few, broad characteristics of these extinctions can be described. Many taxa which became extinct appear to have been endemic to regions within Southeast Asia, while some taxa which experienced extinction or severe range reduction occurred in several regions. Members of the latter group include proboscideans (Stegodon and Palaeloxodon), the pygmy hippopotamus (Hexaprotodon), the orangutan (Pongo), hyenas (Crocuta and Hyaena), the giant panda (Ailuropoda), tapirs (Tapirus and Megatapirus), rhinoceroses (Rhinoceros), and the giant Asian ape Gigantopithecus. The loss of these species cannot be assigned to a single cause. Rather their disappearance is likely tied to both climatic and human agents. Unlike other regions which experienced megafauna extinctions, eustatic changes in sea level in Southeast Asia seems to have been an important factor.     

Micro‐evolutionary diversification among Indian Ocean parrots: temporal and spatial changes in phylogenetic diversity as a consequence of extinction and invasion

Almost 90% of global bird extinctions have occurred on islands. The loss of endemic species from island systems can dramatically alter evolutionary trajectories of insular species biodiversity, resulting in a loss of evolutionary diversity important for species adaptation to changing environments. The Western Indian Ocean islands have been the scene of evolution for a large number of endemic parrots. Since their discovery in the 16th century, many of these parrots have become extinct or have declined in numbers. Alongside the extinction of species, a number of the Indian Ocean islands have experienced colonization by highly invasive parrots, such as the Ring‐necked Parakeet Psittacula krameri. Such extinctions and invasions can, on an evolutionary timescale, drive changes in species composition, genetic diversity and turnover in phylogenetic diversity, all of which can have important impacts on species potential for adaptation to changing environmental and climatic conditions. Using mtDNA cytochrome b data, we resolve the taxonomic placement of three extinct Indian Ocean parrots: the Rodrigues Psittacula exsul, Seychelles Psittacula wardi and Reunion Parakeets Psittacula eques. This case study quantifies how the extinction of these species has resulted in lost historical endemic phylogenetic diversity and reduced levels of species richness, and illustrates how it is being replaced by non‐endemic invasive forms such as the Ring‐necked Parakeet. Finally, we use our phylogenetic framework to identify and recommend a number of phylogenetically appropriate ecological replacements for the extinct parrots. Such replacements may be introduced once invasive forms have been cleared, to rejuvenate ecosystem function and restore lost phylogenetic diversity.     

Prehistoric inter-archipelago trading of Polynesian tree snails leaves a conservation legacy

Inter-archipelago exchange networks were an important aspect of prehistoric Polynesian societies. We report here a novel genetic characterization of a prehistoric exchange network involving an endemic Pacific island tree snail, Partula hyalina. It occurs in the Society (Tahiti only), Austral and Southern Cook Islands. Our genetic data, based on museum, captive and wild-caught samples, establish Tahiti as the source island. The source lineage is polymorphic in shell coloration and contains a second nominal species, the dark-shelled Partula clara, in addition to the white-shelled P. hyalina. Prehistoric inter-island introductions were non-random: they involved white-shelled snails only and were exclusively inter-archipelago in scope. Partulid shells were commonly used in regional Polynesian jewellery, and we propose that the white-shelled P. hyalina, originally restricted to Tahiti, had aesthetic value throughout these archipelagoes. Demand within the Society Islands could be best met by trading dead shells, but a low rate of inter-archipelago exchange may have prompted the establishment of multiple founder populations in the Australs and Southern Cooks. The alien carnivorous land snail Euglandina rosea has recently devastated populations of all 61 endemic species of Society Island partulid snails. Southern Cooks and Australs P. hyalina now represent the only unscathed wild populations remaining of this once spectacular land snail radiation.     

Patterns of endemic extinctions among island bird species

The relationship between island biogeography and the vulnerability of island biota to extinction as a result of human activities was examined. In particular, this study analyzed whether island area, maximum elevation of an island, isolation from the nearest continental landmass, or date of human colonization had statistically significant relationships with the proportion of endemic island bird species that have become endangered or extinct. The study examined islands or island groups with endemic bird species, and which have never been connected to a continental landmass. Both modern and fossil bird species were incorporated into the analysis. Islands that were colonized by humans earliest had the lowest proportion of modern species alone, and modern and fossil species combined, that have gone extinct. However, date of human arrival was not correlated with the proportion of modern species that are endangered. Maximum elevation of an island was negatively correlated with the proportion of modern species that are extinct, and was positively correlated with the proportion of modern species that are endangered. Area was negatively correlated with the proportion of modern species that are endangered. Isolation of islands was not significantly correlated with the proportion of modern species extinct or endangered, but was positively correlated with the proportion of modern and fossil species combined that have gone extinct. These results indicate that the initial spasm of island bird extinctions due to human contact may have, in part, passed. They also indicate that bird species on islands colonized earliest by humans may have had more time to adapt to the presence of man and his commensal species, resulting in reduced extinction rates.     

Extinction of the autochthonous small mammals of Mallorca (Gymnesic Islands, Western Mediterranean) and its ecological consequences

Aim To investigate the chronology, causes and consequences of the extinction of the autochthonous Pleistocene small mammals of Mallorca. Location Mallorca (Gymnesic Islands, Balearics, Western Mediterranean). Methods We have obtained the first direct ¹⁴C ages from the bone collagen of selected samples of two extinct endemic small mammals from Mallorca: the Balearic dormouse, Eliomys morpheus (Rodentia: Myoxidae) and the Balearic shrew, Asoriculus hidalgoi (Soricomorpha: Soricidae). We also present evidence for the absence of both endemics from the earliest Mallorcan archaeological sites and for the introduction of the garden dormouse, Eliomys quercinus, and the wood mouse, Apodemus sylvaticus. Combined information from direct dating of bone collagen of E. quercinus and A. sylvaticus and from cultural associations provides an adequate framework to establish the chronology of the faunal change and to compare it with the chronological information available on climatic change and the first arrival of humans on the islands. Results The chronological record includes the latest evidence available for the survival of endemic species and the earliest introduction of small mammals into Mallorca. We present ‘uncertainty periods for extinction’ (UPEs) of both endemic mammals based on the chronology of their last occurrence and on the inferred timing of their extinction (restricted UPEs). Main conclusions Possible causes for the extinction of autochthonous small mammals on Mallorca are discussed. Once we have discarded climatic causes, predation by invasive species, competition with newcomers and habitat deterioration, the introduction of diseases emerges as the most reasonable explanation for these extinctions. Based on the identification of changes in keystone species in Mallorcan ecosystems, we propose a tentative schedule of key ecological changes that have taken place over the past 5 millennia.     

Significance of new records of Tridacna squamosa Lamarck, 1819, in the Tuamotu and Gambier Archipelagos (French Polynesia)

The giant clam subfamily Tridacninae (family Cardiidae) is an important group of bivalve molluscs found throughout the Red Sea and Indo-Pacific, from East Africa to the Eastern Pacific biogeographic region. The Tridacna genus is currently revised with numerous cryptic species identified with molecular markers. New Tridacna records from the fringe of the known distribution areas are extremely useful to identify genetically unique species, geographic ranges, and to examine processes associated with species differentiation. While Tridacna maxima is abundant in French Polynesia (Central South Pacific Ocean) the larger fluted giant clam Tridacna squamosa was formerly reported only in the Austral Islands in the south. Following a recent survey that spanned 23 islands and atolls of the Society, Tuamotu and Gambier Archipelagos, the presence of T. squamosa between the Cook Islands and Pitcairn Islands is confirmed using both morphological and molecular information, suggesting a relic distribution across the Central Pacific Ocean. Tridacna squamosa is rare, but present throughout Tuamotu and Gambier. However, it remained undetected from the Society Islands, probably due to historical over-fishing. This species is valued by local inhabitants, and is sought after mainly as gifts and also for a limited local shell trade. The rarity of T. squamosa may call for conservation measures in the near future.     

Human‐mediated extirpation of the unique Chatham Islands sea lion and implications for the conservation management of remaining New Zealand sea lion populations

While terrestrial megafaunal extinctions have been well characterized worldwide, our understanding of declines in marine megafauna remains limited. Here, we use ancient DNA analyses of prehistoric (<1450–1650 AD) sea lion specimens from New Zealand's isolated Chatham Islands to assess the demographic impacts of human settlement. These data suggest there was a large population of sea lions, unique to the Chatham Islands, at the time of Polynesian settlement. This distinct mitochondrial lineage became rapidly extinct within 200 years due to overhunting, paralleling the extirpation of a similarly large endemic mainland population. Whole mitogenomic analyses confirm substantial intraspecific diversity among prehistoric lineages. Demographic models suggest that even low harvest rates would likely have driven rapid extinction of these lineages. This study indicates that surviving Phocarctos populations are remnants of a once diverse and widespread sea lion assemblage, highlighting dramatic human impacts on endemic marine biodiversity. Our findings also suggest that Phocarctos bycatch in commercial fisheries may contribute to the ongoing population decline.     

Rediscovery of Leptoxis compacta (Anthony, 1854) (Gastropoda: Cerithioidea: Pleuroceridae)

The Mobile River Basin is a hotspot of molluscan endemism, but anthropogenic activities have caused at least 47 molluscan extinctions, 37 of which were gastropods, in the last century. Nine of these suspected extinctions were in the freshwater gastropod genus Leptoxis (Cerithioidea: Pleuroceridae). Leptoxis compacta, a Cahaba River endemic, has not been collected for >70 years and was formally declared extinct in 2000. Such gastropod extinctions underscore the imperilment of freshwater resources and the current biodiversity crisis in the Mobile River Basin. During a May 2011 gastropod survey of the Cahaba River in central Alabama, USA, L. compacta was rediscovered. The identification of snails collected was confirmed through conchological comparisons to the L. compacta lectotype, museum records, and radulae morphology of historically collected L. compacta. Through observations of L. compacta in captivity, we document for the first time that the species lays eggs in short, single lines. Leptoxis compacta is restricted to a single location in the Cahaba River, and is highly susceptible to a single catastrophic extinction event. As such, the species deserves immediate conservation attention. Artificial propagation and reintroduction of L. compacta into its native range may be a viable recovery strategy to prevent extinction from a single perturbation event.     

The Decimation of Endemic Hawai'ian Tree Snails by Alien Predators

Endemic terrestrial tree snails of the Hawai'ian Islands, likethose of other oceanic islands and even some continental areas,are extremely sensitive to disturbance because of their lowpopulation numbers and small geographic ranges. Like many otherplants and animals of oceanic islands, they have evolved nodefenses against introduced predators and competitors. The rangeof Achatinella mustelina, a tree-snail species found only ina short mountain range on the island of O'ahu, typifies thisproblem. Mark-recapture studies at two field locations revealthat the snails exhibit slow growth andlate maturity (3–5years). Fecundity is estimated at about 7 offspring per adultper year. The young are born live at about 4.6 mm. Populationgrowth typically depends on considerable longevity (> 10years). Demographic effects of the depredations by alien predators,rats and a North American predatory snail, Euglandina rosea,were documented in two long-term study sites. The predatorysnail eats all sizes of A. mustelina and can rapidly drive populationsto extinction (less than one year). Rats tend to select largersnails as prey and may leave an area before destroying all ofthe prey snails present; while reproductive output is temporarilydestroyed, populations may survive. Actions necessary to conserveHawai;ian tree snails, or indeed any group of relatively sedentaryinvertebrates with small species ranges, must include predatorabatement, but also preservation and restoration of sufficientlyarge and complex forest habitats that the invertebrates mayfind refuge from alien predators.     

The ghosts of mammals past: biological and geographical patterns of global mammalian extinction across the Holocene

Although the recent historical period is usually treated as a temporal base-line for understanding patterns of mammal extinction, mammalian biodiversity loss has also taken place throughout the Late Quaternary. We explore the spatial, taxonomic and phylogenetic patterns of 241 mammal species extinctions known to have occurred during the Holocene up to the present day. To assess whether our understanding of mammalian threat processes has been affected by excluding these taxa, we incorporate extinct species data into analyses of the impact of body mass on extinction risk. We find that Holocene extinctions have been phylogenetically and spatially concentrated in specific taxa and geographical regions, which are often not congruent with those disproportionately at risk today. Large-bodied mammals have also been more extinction-prone in most geographical regions across the Holocene. Our data support the extinction filter hypothesis, whereby regional faunas from which susceptible species have already become extinct now appear less threatened; they may also suggest that different processes are responsible for driving past and present extinctions. We also find overall incompleteness and inter-regional biases in extinction data from the recent fossil record. Although direct use of fossil data in future projections of extinction risk is therefore not straightforward, insights into extinction processes from the Holocene record are still useful in understanding mammalian threat.     

Distribution,movement, and microhabitat use of the introduced predatory snail Euglandina rosea in Hawaii: implications for management

The purposeful introduction of the land snail Euglandina rosea, which feeds exclusively on snails and slugs, has been implicated as a major factor in the decline of diverse Pacific island land snail faunas. We report on the distribution, movement patterns, and microhabitat preferences of E. rosea in a gulch in the Waianae Mountains, Oahu, Hawaii, because such data will help focus management actions at a local scale to protect native snail populations in areas where E. rosea is established. The Waianae Mountains harbor many endangered or threatened snails, most currently found in isolated habitat patches near the ridges. Conversely, most living individuals (28/29) and shells (46/56) of E. rosea were collected within the gulch, which supported higher densities of other native and non‐native snails, and was cooler and more moist than the ridges. Thirteen individuals of E. rosea were tracked (eight directly using a bobbin and thread method, and five indirectly by mark–recapture); most (10/13) moved on average <2.5 m per week (range 0.1–25.21 m), and all stayed within the gulch. Members of E. rosea preferred leaf litter over open, fern/shrub, or wood microhabitats. There were large differences in the population density of E. rosea over small spatial scales, indicating that there may be places where native snail populations could persist even in areas where populations of E. rosea are established. Identifying areas with differing population densities of E. rosea is critical for not only understanding why some native snail species may be more vulnerable to extinction, but also to locate areas where predation pressure is low and conservation efforts will be most likely to succeed.