Extinction‐driven changes in frugivore communities on oceanic islands

Global change and human expansion have resulted in many species extinctions worldwide, but the geographic variation and determinants of extinction risk in particular guilds still remain little explored. Here, we quantified insular extinctions of frugivorous vertebrates (including birds, mammals and reptiles) across 74 tropical and subtropical oceanic islands within 20 archipelagos worldwide and investigated extinction in relation to island characteristics (island area, isolation, elevation and climate) and species’ functional traits (body mass, diet and ability to fly). Out of the 74 islands, 33 islands (45%) have records of frugivore extinctions, with one third (mean: 34%, range: 2–100%) of the pre‐extinction frugivore community being lost. Geographic areas with more than 50% loss of pre‐extinction species richness include islands in the Pacific (within Hawaii, Cook Islands and Tonga Islands) and the Indian Ocean (Mascarenes, Seychelles). The proportion of species richness lost from original pre‐extinction communities is highest on small and isolated islands, increases with island elevation, but is unrelated to temperature or precipitation. Large and flightless species had higher extinction probability than small or volant species. Across islands with extinction events, a pronounced downsizing of the frugivore community is observed, with a strong extinction‐driven reduction of mean body mass (mean: 37%, range: –18–100%) and maximum body mass (mean: 51%, range: 0–100%). The results document a substantial trophic downgrading of frugivore communities on oceanic islands worldwide, with a non‐random pattern in relation to geography, island characteristics and species’ functional traits. This implies severe consequences for ecosystem processes that depend on mutualistic plant–animal interactions, including ecosystem dynamics that result from the dispersal of large‐seeded plants by large‐bodied frugivores. We suggest that targeted conservation and rewilding efforts on islands are needed to halt the defaunation of large and non‐volant seed dispersers and to restore frugivore communities and key ecological interactions.     

Late Quaternary reptile extinctions: size matters,insularity dominates

Aim A major Late Quaternary vertebrate extinction event affected mostly large‐bodied ‘megafauna’. This is well documented in both mammals and birds, but evidence of a similar trend in reptiles is scant. We assess the relationship between body size and Late Quaternary extinction in reptiles at the global level. Location Global. Methods We compile a body size database for all 82 reptile species that are known to have gone extinct during the last 50,000 years and compare them with the sizes of 10,090 extant reptile species (97% of known extant diversity). We assess the body size distributions in the major reptile groups: crocodiles, lizards, snakes and turtles, while testing and correcting for a size bias in the fossil record. We examine geographical biases in extinction by contrasting mainland and insular reptile assemblages, and testing for biases within regions and then globally by using geographically weighted models. Results Extinct reptiles were larger than extant ones, but there was considerable variation in extinction size biases among groups. Extinct lizards and turtles were large, extinct crocodiles were small and there was no trend in snakes. Lizard lineages vary in the way their extinction is related to size. Extinctions were particularly prevalent on islands, with 73 of the 82 extinct species being island endemics. Four others occurred in Australia. The fossil record is biased towards large‐bodied reptiles, but extinct lizards were larger than extant ones even after we account for this. Main conclusions Body size played a complex role in the extinction of Late Quaternary reptiles. Larger lizard and turtle species were clearly more affected by extinction mechanisms such as over exploitation and invasive species, resulting in a prevalence of large‐bodied species among extinct taxa. Insularity was by far the strongest correlate of recent reptile extinctions, suggesting that size‐biased extinction mechanisms are amplified in insular environments.     

Integration of palaeontological, historical, and geographical data on the extinction of koa-finches

Identifying the root causes of extinction or endangerment requires long chronological records that begin before a population started to decline and extend until its extinction or functional extinction. We present a case study of the koa‐finches, genus Rhodacanthis, an extinct group of Hawaiian honeycreepers that was specialized to feed on green pods and seeds of the koa tree or other leguminous plants. Six island populations of koa‐finches are known; four in the Holocene fossil record and two that survived until the 1890s. We document the palaeoecological context of the fossils and identify constraints on the age span of the specimen record for each population using stratigraphic contexts, associated radiometric determinations, and museum specimen data. We estimate the potential geographical range of koa‐finches at the time of human arrival using two methods: assessment of their historical and palaeo‐habitats, and geographical information system mapping of the pre‐human distribution of the koa plant (Acacia koa) and its sister species, the koai‘a plant (Acacia koaia). After integrating the foregoing data with chronological records and distributional maps of the potential forcing agents of extinction, we conclude that at least two extinctions of island populations were due to ecological change in the lowlands in the prehistorical and perhaps the early historical periods. In the same time frame, the koa‐finch populations on Hawai‘i Island became rare and restricted to upland refugia, making them vulnerable to the upland forest harvesting and degradation that was accelerating in the 1890s. Neither climatic variation nor mosquito‐vectored diseases are likely to have caused the observed extinctions. This study illustrates an approach that can be applied to many other extinct and endangered island species to better understand the causes of high extinction rates in the human era.     

Reconstructing past species assemblages reveals the changing patterns and drivers of extinction through time

Predicting future species extinctions from patterns of past extinctions or current threat status relies on the assumption that the taxonomic and biological selectivity of extinction is consistent through time. If the driving forces of extinction change through time, this assumption may be unrealistic. Testing the consistency of extinction patterns between the past and the present has been difficult, because the phylogenetically explicit methods used to model present-day extinction risk typically cannot be applied to the data from the fossil record. However, the detailed historical and fossil records of the New Zealand avifauna provide a unique opportunity to reconstruct a complete, large faunal assemblage for different periods in the past. Using the first complete phylogeny of all known native New Zealand bird species, both extant and extinct, we show how the taxonomic and phylogenetic selectivity of extinction, and biological correlates of extinction, change from the pre-human period through Polynesian and European occupation, to the present. These changes can be explained both by changes in primary threatening processes, and by the operation of extinction filter effects. The variable patterns of extinction through time may confound attempts to identify risk factors that apply across time periods, and to infer future species declines from past extinction patterns and current threat status.     

The best sections method of studying mass extinctions

Phanerozoic mass extinctions have been studied primarily by analysing global diversity patterns compiled from the published literature. However, such compilations are beset by problems of incorrect correlation, imprecise age assignments and changing taxonomy. An alternative approach is to analyse mass extinctions by the ‘best sections’ method. This method identifies abundantly fossiliferous, well‐studied, stratigraphically dense and temporally extensive fossil records in strata that contain geochemical and other relevant non‐palaeontological data from a single depositional basin or geographically restricted outcrop area as the ‘best sections’ by which to analyse extinctions. A strength of the best sections method is that it allows the extinctions identified to be compared directly to changes in facies and other factors recorded in the best section. And, the hypothesis of a widespread extinction based on an extinction seen in a best section can be tested by its presence or absence in temporally equivalent sections. What we need are more field‐based studies of the best sections that encompass mass extinctions (real and hypothetical) and less of a reliance on literature‐based diversity compilations to produce a more reliable and comprehensive understanding of the history of extinctions.     

The fossil record of the sixth extinction

Comparing the magnitude of the current biodiversity crisis with those in the fossil record is difficult without an understanding of differential preservation. Integrating data from palaeontological databases with information on IUCN status, ecology and life history characteristics of contemporary mammals, we demonstrate that only a small and biased fraction of threatened species (< 9%) have a fossil record, compared with 20% of non‐threatened species. We find strong taphonomic biases related to body size and geographic range. Modern species with a fossil record tend to be large and widespread and were described in the 19^th century. The expected magnitude of the current extinction based only on species with a fossil record is about half of that of one based on all modern species; values for genera are similar. The record of ancient extinctions may be similarly biased, with many species having originated and gone extinct without leaving a tangible record.     

Prehistoric bird extinctions and human hunting

Holocene fossils document the extinction of hundreds of bird species on Pacific islands during prehistoric human occupation. Human hunting is implicated in these extinctions, but the impact of hunting is difficult to disentangle from the effects of other changes induced by humans, including habitat destruction and the introduction of other mammalian predators. Here, we use data from bones collected at a natural sand dune site and associated archaeological middens in New Zealand to show that, having controlled for differences in body mass and family membership (and hence for variation in life-history traits related to population growth rate), birds that were more intensively hunted by prehistoric humans had a higher probability of extinction. This result cannot be attributed to preservation biases and provides clear evidence that selective hunting contributed significantly to prehistoric bird extinctions at this site.     

The uncertain blitzkrieg of Pleistocene megafauna

We investigated, using meta‐analysis of empirical data and population modelling, plausible scenarios for the cause of late Pleistocene global mammal extinctions. We also considered the rate at which these extinctions may have occurred, providing a test of the so‐called ‘blitzkrieg’ hypothesis, which postulates a rapid, anthropogenically driven, extinction event. The empirical foundation for this work was a comprehensive data base of estimated body masses of mammals, comprising 198 extinct and 433 surviving species > 5 kg, which we compiled through an extensive literature search. We used mechanistic population modelling to simulate the role of human hunting efficiency, meat off‐take, relative naivety of prey to invading humans, variation in reproductive fitness of prey and deterioration of habitat quality (due to either anthropogenic landscape burning or climate change), and explored the capacity of different modelling scenarios to recover the observed empirical relationship between body mass and extinction proneness. For the best‐fitting scenarios, we calculated the rate at which the extinction event would have occurred. All of the modelling was based on sampling randomly from a plausible range of parameters (and their interactions), which affect human and animal population demographics. Our analyses of the empirical data base revealed that the relationship between body mass and extinction risk relationship increases continuously from small‐ to large‐sized animals, with no clear ‘megafaunal’ threshold. A logistic ancova model incorporating body mass and geography (continent) explains 92% of the variation in the observed extinctions. Population modelling demonstrates that there were many plausible mechanistic scenarios capable of reproducing the empirical body mass–extinction risk relationship, such as specific targeting of large animals by humans, or various combinations of habitat change and opportunistic hunting. Yet, given the current imperfect knowledge base, it is equally impossible to use modelling to isolate definitively any single scenario to explain the observed extinctions. However, one universal prediction, which applied in all scenarios in which the empirical distribution was correctly predicted, was for the extinctions to be rapid following human arrival and for surviving fauna to be suppressed below their pre‐‘blitzkrieg’ densities. In sum, human colonization in the late Pleistocene almost certainly triggered a ‘blitzkrieg’ of the ‘megafauna’, but the operational details remain elusive.     

Convergent evolution of morphology and habitat use in the explosive Hawaiian fancy case caterpillar radiation

Species occurring in unconnected, but similar habitats and under similar selection pressures often display strikingly comparable morphology, behaviour and life history. On island archipelagos where colonizations and extinctions are common, it is often difficult to separate whether similar traits are a result of in situ diversification or independent colonization without a phylogeny. Here, we use one of Hawaii's most ecologically diverse and explosive endemic species radiations, the Hawaiian fancy case caterpillar genus Hyposmocoma, to test whether in situ diversification resulted in convergence. Specifically, we examine whether similar species utilizing similar microhabitats independently developed largely congruent larval case phenotypes in lineages that are in comparable, but isolated environments. Larvae of these moths are found on all Hawaiian Islands and are characterized by an extraordinary array of ecomorphs and larval case morphology. We focus on the ‘purse cases’, a group that is largely specialized for living within rotting wood. Purse cases were considered a monophyletic group, because morphological, behavioural and ecological traits appeared to be shared among all members. We constructed a phylogeny based on nuclear and mitochondrial DNA sequences from 38 Hyposmocoma species, including all 14 purse case species and 24 of non‐purse case congeners. Divergence time estimation suggests that purse case lineages evolved independently within dead wood and developed nearly identical case morphology twice: once on the distant Northwest Hawaiian Islands between 15.5 and 9 Ma and once on the younger main Hawaiian Islands around 3.0 Ma. Multiple ecomorphs are usually found on each island, and the ancestral ecomorph of Hyposmocoma appears to have lived on tree bark. Unlike most endemic Hawaiian radiations that follow a clear stepwise progression of colonization, purse case Hyposmocoma do not follow a pattern of colonization from older to younger island. We postulate that the diversity of microhabitats and selection from parasitism/predation from endemic predators may have shaped case architecture in this extraordinary endemic radiation of Hawaiian insects.     

Introductions do not compensate for functional and phylogenetic losses following extinctions in insular bird assemblages

The ratio of species extinctions to introductions has been comparable for many insular assemblages, suggesting that introductions could have ‘compensated’ for extinctions. However, the capacity for introduced species to replace ecological roles and evolutionary history lost following extinction is unclear. We investigated changes in bird functional and phylogenetic diversity in the wake of extinctions and introductions across a sample of 32 islands worldwide. We found that extinct and introduced species have comparable functional and phylogenetic alpha diversity. However, this was distributed at different positions in functional space and in the phylogeny, indicating a ‘false compensation’. Introduced and extinct species did not have equivalent functional roles nor belong to similar lineages. This makes it unlikely that novel island biotas composed of introduced taxa will be able to maintain ecological roles and represent the evolutionary histories of pre‐disturbance assemblages and highlights the importance of evaluating changes in alpha and beta diversity concurrently.     

ORIGINAL ARTICLE: Mass extinctions in the Azores during the last glaciation: fact or myth?

Aim The influence of the last glaciation on the shallow‐water marine malacofauna of the Azores Islands is reviewed. We test, for this fauna, the ‘Pleistocene temperature theory’ of J.C. Briggs, which hypothesizes that a (supposed) lack of endemics in the older (Azorean endemic) fauna resulted from extinctions caused by a severe drop in sea surface temperatures during the Pleistocene. Location Santa Maria Island, Azores, Portugal. Methods We compare the fossil mollusc fauna of Prainha, Praia do Calhau and Lagoinhas Pleistocene outcrops with the recent mollusc fauna of the Azores Islands. We dated the fossil fauna, using shells of Patella aspera Röding, 1798, by standard U/Th methodology at the GEOTOP laboratory (Université du Québec à Montreal, Canada). Results Dating of the shells of P. aspera indicates that the deposition of the lower unit of the Prainha outcrop corresponded to Marine Oxygen Isotope Substage 5e (MISS 5e). Not a single endemic Azorean species of mollusc that is present in the Pleistocene fossil record has since become extinct, and we found no signs of ‘mass extinctions’ in the littoral marine molluscs of the Azores. The only species that were extirpated from these islands were thermophilic molluscs and littoral bivalves living in fine sand. Main conclusions Our results do not support Briggs’‘Pleistocene temperature theory’. Nor did we find evidence supporting the hypothesis that most of the marine organisms now present in the Azores recolonized the islands after the last glacial maximum.     

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.     

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.     

Competition and introduction regime shape exotic bird communities in Hawaii

Complex combinations of historical and local-regional processes determine the assembly of ecological communities. We investigated such processes in the Hawaiian introduced avifauna, comprising 140 years of historical records of invasions and extinctions of birds. Here the particular introduction regime (i.e., colonization attempts and number of introduced species) and priority effects constitute the historical (and regional) component, and competition is the local component. These processes are theoretically supported by means of a Lotka–Volterra model of species competition, finding that changes in the specific introduction regime might result in different extinction dynamics. Both field data and model outcomes support the biotic resistance hypothesis, so that the invasibility of new incomers decrease with species richness. Finally, we found that the resistance to new invaders depends on the particular introduction regime. Thus, community assembly models built to predict the success of exotic species should consider more scenarios than random introduction regimes.     

Characterising extinction debt following habitat fragmentation using neutral theory

Habitat loss leads to species extinctions, both immediately and over the long term as ‘extinction debt’ is repaid. The same quantity of habitat can be lost in different spatial patterns with varying habitat fragmentation. How this translates to species loss remains an open problem requiring an understanding of the interplay between community dynamics and habitat structure across temporal and spatial scales. Here we develop formulas that characterise extinction debt in a spatial neutral model after habitat loss and fragmentation. Central to our formulas are two new metrics, which depend on properties of the taxa and landscape: ‘effective area’, measuring the remaining number of individuals and ‘effective connectivity’, measuring individuals’ ability to disperse through fragmented habitat. This formalises the conventional wisdom that habitat area and habitat connectivity are the two critical requirements for long‐term preservation of biodiversity. Our approach suggests that mechanistic fragmentation metrics help resolve debates about fragmentation and species loss.     

Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis

Cascales‐Miñana, B., Muñoz‐Bertomeu, J., Ros, R., Segura, J. 2010: Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis. Lethaia, 10.1111/j.1502‐3931.2009.00212.x Studying the macroevolutionary patterns of vascular plants from the Silurian to the present‐day provides a global record of plant life history. Evolutionary rates (origination, extinction and diversification) for families, orders, classes and divisions were analysed, as was abundance and richness for 21 time intervals. An accumulative analysis, based on the total plant fossil record, the accumulated extinctions and relative diversity, was also carried out. The diversification rate shows a uniquely constant and progressive reduction from the end of the Carboniferous to the Permian when the lowest values are registered. Very small peaks seem to reflect Cretaceous extinction for families. At family level, only two time intervals present higher extinctions, than originations. Richness and accumulative analyses reveal that only 32% of the families analysed became extinct, and that approximately 90% of them disappeared at the end of the Palaeozoic. Our results indicate that plants did not undergo mass extinction events in the ‘big five’ sense, but rather, mass ecological reorganization the absence of important extinction events or evolutionary innovations producing diversification patterns without abrupt changes. □Diversification, evolutionary, extinction, fossil record, innovations, radiation, vascular plants.     

Was a ‘hyperdisease’ responsible for the late Pleistocene megafaunal extinction?

Numerous hypotheses have been proposed to explain the end Pleistocene extinction of large bodied mammals. The disease hypothesis attributes the extinction to the arrival of a novel ‘hyperdisease’ brought by immigrating aboriginal humans. However, until West Nile virus (WNV) invaded the United States, no known disease met the criteria of a hyperdisease. We evaluate the disease hypothesis using WNV in the United States as a model system. We show that WNV is size‐biased in its infection of North America birds, but is unlikely to result in an extinction similar to that of the end Pleistocene. WNV infects birds more uniformly across the body size spectrum than extinctions did across mammals and is not size‐biased within orders. Our study explores the potential impact of WNV on bird populations and provides no support for disease as a causal mechanism for the end Pleistocene megafaunal extinction.     

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.     

Was the Late Ordovician mass extinction truly exceptional?

The Late Ordovician mass extinction event is the oldest of the five great extinction events in the fossil record. It has long been regarded as an outlier among mass extinctions, primarily due to its association with a cooling climate. However, recent temporally better resolved fossil biodiversity estimates complicate this view, providing growing evidence for a prolonged but punctuated biodiversity decline modulated by changes in atmospheric composition, ocean chemistry, and viable habitat area. This evolving view invokes extinction drivers similar to those that occurred during other major extinctions; some are even factors in the current human-induced biodiversity crisis. Even this very ancient and, at first glance, exceptional event conveys important lessons about the intensifying ‘sixth mass extinction’.     

The loss of indirect interactions leads to cascading extinctions of carnivores

Species extinctions are biased towards higher trophic levels, and primary extinctions are often followed by unexpected secondary extinctions. Currently, predictions on the vulnerability of ecological communities to extinction cascades are based on models that focus on bottom‐up effects, which cannot capture the effects of extinctions at higher trophic levels. We show, in experimental insect communities, that harvesting of single carnivorous parasitoid species led to a significant increase in extinction rate of other parasitoid species, separated by four trophic links. Harvesting resulted in the release of prey from top‐down control, leading to increased interspecific competition at the herbivore trophic level. This resulted in increased extinction rates of non‐harvested parasitoid species when their host had become rare relative to other herbivores. The results demonstrate a mechanism for horizontal extinction cascades, and illustrate that altering the relationship between a predator and its prey can cause wide‐ranging ripple effects through ecosystems, including unexpected extinctions.