Demographic models predict end-Pleistocene arrival and rapid expansion of pre-agropastoralist humans in Cyprus.

The antiquity of human dispersal into Mediterranean islands and ensuing coastal adaptation have remained largely unexplored due to the prevailing assumption that the sea was a barrier to movement and that islands were hostile environments to early hunter-gatherers [J. F. Cherry, T. P. Leppard, J. Isl. Coast. Archaeol. 13, 191-205 (2018), 10.1080/15564894.2016.1276489]. Using the latest archaeological data, hindcasted climate projections, and age-structured demographic models, we demonstrate evidence for early arrival (14,257 to 13,182 calendar years ago) to Cyprus and predicted that large groups of people (~1,000 to 1,375) arrived in 2 to 3 main events occurring within <100 y to ensure low extinction risk. These results indicate that the postglacial settlement of Cyprus involved only a few large-scale, organized events requiring advanced watercraft technology. Our spatially debiased and Signor-Lipps-corrected estimates indicate rapid settlement of the island within <200 y, and expansion to a median of 4,000 to 5,000 people (0.36 to 0.46 km-2) in <11 human generations (<300 y). Our results do not support the hypothesis of inaccessible and inhospitable islands in the Mediterranean for pre-agropastoralists, agreeing with analogous conclusions for other parts of the world [M. I. Bird et al., Sci. Rep. 9, 8220 (2019), 10.1038/s41598-019-42946-9]. Our results also highlight the need to revisit these questions in the Mediterranean and test their validity with new technologies, field methods, and data. By applying stochastic models to the Mediterranean region, we can place Cyprus and large islands in general as attractive and favorable destinations for paleolithic peoples.

Forest mosaics, not savanna corridors, dominated in Southeast Asia during the Last Glacial Maximum.

The dominant paradigm is that large tracts of Southeast Asia's lowland rainforests were replaced with a "savanna corridor" during the cooler, more seasonal climates of the Last Glacial Maximum (LGM) (23,000 to 19,000 y ago). This interpretation has implications for understanding the resilience of Asia's tropical forests to projected climate change, implying a vulnerability to "savannization". A savanna corridor is also an important foundation for archaeological interpretations of how humans moved through and settled insular Southeast Asia and Australia. Yet an up-to-date, multiproxy, and empirical examination of the palaeoecological evidence for this corridor is lacking. We conducted qualitative and statistical analyses of 59 palaeoecological records across Southeast Asia to test the evidence for LGM savannization and clarify the relationships between methods, biogeography, and ecological change in the region from the start of Late Glacial Period (119,000 y ago) to the present. The pollen records typically show montane forest persistence during the LGM, while δ13C biomarker proxies indicate the expansion of C4-rich grasslands. We reconcile this discrepancy by hypothesizing the expansion of montane forest in the uplands and replacement of rainforest with seasonally dry tropical forest in the lowlands. We also find that smooth forest transitions between 34,000 and 2,000 y ago point to the capacity of Southeast Asia's ecosystems both to resist and recover from climate stressors, suggesting resilience to savannization. Finally, the timing of ecological change observed in our combined datasets indicates an 'early' onset of the LGM in Southeast Asia from ~30,000 y ago.

Differential developmental rates and demographics in Red Kangaroo (Osphranter rufus) populations separated by the dingo barrier fence.

Decommissioning the dingo barrier fence has been suggested to reduce destructive dingo control and encourage a free transfer of biota between environments in Australia. Yet the potential impacts that over a century of predator exclusion might have had on the population dynamics and developmental biology of prey populations has not been assessed. We here combine demographic data and both linear and geometric morphometrics to assess differences in populations among 166 red kangaroos (Osphranter rufus)-a primary prey species of the dingo-from two isolated populations on either side of the fence. We also quantified the differences in aboveground vegetation biomass for the last 10 years on either side of the fence. We found that the age structure and growth patterns, but not cranial shape, differed between the two kangaroo populations. In the population living with a higher density of dingoes, there were relatively fewer females and juveniles. These individuals were larger for a given age, despite what seems to be lower vegetation biomass. However, how much of this biomass represented kangaroo forage is uncertain and requires further on-site assessments. We also identified unexpected differences in the ontogenetic trajectories in relative pes length between the sexes for the whole sample, possibly associated with male competition or differential weight-bearing mechanics. We discuss potential mechanisms behind our findings and suggest that the impacts of contrasting predation pressures across the fence, for red kangaroos and other species, merit further investigation.

Estimating co-extinction threats in terrestrial ecosystems.

The biosphere is changing rapidly due to human endeavour. Because ecological communities underlie networks of interacting species, changes that directly affect some species can have indirect effects on others. Accurate tools to predict these direct and indirect effects are therefore required to guide conservation strategies. However, most extinction-risk studies only consider the direct effects of global change-such as predicting which species will breach their thermal limits under different warming scenarios-with predictions of trophic cascades and co-extinction risks remaining mostly speculative. To predict the potential indirect effects of primary extinctions, data describing community interactions and network modelling can estimate how extinctions cascade through communities. While theoretical studies have demonstrated the usefulness of models in predicting how communities react to threats like climate change, few have applied such methods to real-world communities. This gap partly reflects challenges in constructing trophic network models of real-world food webs, highlighting the need to develop approaches for quantifying co-extinction risk more accurately. We propose a framework for constructing ecological network models representing real-world food webs in terrestrial ecosystems and subjecting these models to co-extinction scenarios triggered by probable future environmental perturbations. Adopting our framework will improve estimates of how environmental perturbations affect whole ecological communities. Identifying species at risk of co-extinction (or those that might trigger co-extinctions) will also guide conservation interventions aiming to reduce the probability of co-extinction cascades and additional species losses.

Dredging activity in a highly urbanised estuary did not affect the long-term occurrence of Indo-Pacific bottlenose dolphins and long-nosed fur seals.

Dredging is an excavation activity used worldwide in marine and freshwater environments to create, deepen, and maintain waterways, harbours, channels, locks, docks, berths, river entrances, and approaches to ports and boat ramps. However, dredging impacts on marine life, including marine mammals (cetaceans, pinnipeds, and sirenians), remain largely unknown. Here we quantified the effect of dredging operations in 2005 and 2019 on the occurrence of Indo-Pacific bottlenose dolphins (Tursiops aduncus) and long-nosed fur seals (Arctocephalus forsteri) in the Port River estuary, a highly urbanized estuary in Adelaide, South Australia. We applied generalised linear models to two long-term sighting datasets (dolphins: 1992-2020, fur seals: 2010-2020), to analyse changes in sighting rates as a function of dredging operations, season, rainfall, and sea surface temperature. We showed that the fluctuations in both dolphin and fur seal occurrences were not correlated with dredging operations, whereas sea surface temperature and season were stronger predictors of both species sighting rates (with seals more prevalent during the colder months, and dolphins in summer). Given the highly industrial environment of the Port River estuary, it is possible that animals in this area are habituated to high noise levels and therefore were not disturbed by dredging operations. Future research would benefit from analysing short-term effects of dredging operations on behaviour, movement patterns and habitat use to determine effects of possible habitat alteration caused by dredging.

Landscape rules predict optimal superhighways for the first peopling of Sahul.

Archaeological data and demographic modelling suggest that the peopling of Sahul required substantial populations, occurred rapidly within a few thousand years and encompassed environments ranging from hyper-arid deserts to temperate uplands and tropical rainforests. How this migration occurred and how humans responded to the physical environments they encountered have, however, remained largely speculative. By constructing a high-resolution digital elevation model for Sahul and coupling it with fine-scale viewshed analysis of landscape prominence, least-cost pedestrian travel modelling and high-performance computing, we create over 125 billion potential migratory pathways, whereby the most parsimonious routes traversed emerge. Our analysis revealed several major pathways-superhighways-transecting the continent, that we evaluated using archaeological data. These results suggest that the earliest Australian ancestors adopted a set of fundamental rules shaped by physiological capacity, attraction to visually prominent landscape features and freshwater distribution to maximize survival, even without previous experience of the landscapes they encountered.

Stochastic models support rapid peopling of Late Pleistocene Sahul.

The peopling of Sahul (the combined continent of Australia and New Guinea) represents the earliest continental migration and settlement event of solely anatomically modern humans, but its patterns and ecological drivers remain largely conceptual in the current literature. We present an advanced stochastic-ecological model to test the relative support for scenarios describing where and when the first humans entered Sahul, and their most probable routes of early settlement. The model supports a dominant entry via the northwest Sahul Shelf first, potentially followed by a second entry through New Guinea, with initial entry most consistent with 50,000 or 75,000 years ago based on comparison with bias-corrected archaeological map layers. The model's emergent properties predict that peopling of the entire continent occurred rapidly across all ecological environments within 156-208 human generations (4368-5599 years) and at a plausible rate of 0.71-0.92 km year-1. More broadly, our methods and approaches can readily inform other global migration debates, with results supporting an exit of anatomically modern humans from Africa 63,000-90,000 years ago, and the peopling of Eurasia in as little as 12,000-15,000 years via inland routes.

Relative demographic susceptibility does not explain the extinction chronology of Sahul's megafauna.

The causes of Sahul's megafauna extinctions remain uncertain, although several interacting factors were likely responsible. To examine the relative support for hypotheses regarding plausible ecological mechanisms underlying these extinctions, we constructed the first stochastic, age-structured models for 13 extinct megafauna species from five functional/taxonomic groups, as well as 8 extant species within these groups for comparison. Perturbing specific demographic rates individually, we tested which species were more demographically susceptible to extinction, and then compared these relative sensitivities to the fossil-derived extinction chronology. Our models show that the macropodiformes were the least demographically susceptible to extinction, followed by carnivores, monotremes, vombatiform herbivores, and large birds. Five of the eight extant species were as or more susceptible than the extinct species. There was no clear relationship between extinction susceptibility and the extinction chronology for any perturbation scenario, while body mass and generation length explained much of the variation in relative risk. Our results reveal that the actual mechanisms leading to the observed extinction chronology were unlikely related to variation in demographic susceptibility per se, but were possibly driven instead by finer-scale variation in climate change and/or human prey choice and relative hunting success.

FosSahul 2.0, an updated database for the Late Quaternary fossil records of Sahul.

The 2016 version of the FosSahul database compiled non-human vertebrate megafauna fossil ages from Sahul published up to 2013 in a standardized format. Its purpose was to create a publicly available, centralized, and comprehensive database for palaeoecological investigations of the continent. Such databases require regular updates and improvements to reflect recent scientific findings. Here we present an updated FosSahul (2.0) containing 11,871 dated non-human vertebrate fossil records from the Late Quaternary published up to 2018. Furthermore, we have extended the information captured in the database to include methodological details and have developed an algorithm to automate the quality-rating process. The algorithm makes the quality-rating more transparent and easier to reproduce, facilitating future database extensions and dissemination. FosSahul has already enabled several palaeoecological analyses, and its updated version will continue to provide a centralized organisation of Sahul's fossil records. As an example of an application of the database, we present the temporal pattern in megafauna genus richness inferred from available data in relation to palaeoclimate indices over the past 180,000 years.

Climate-human interaction associated with southeast Australian megafauna extinction patterns.

The mechanisms leading to megafauna (>44 kg) extinctions in Late Pleistocene (126,000-12,000 years ago) Australia are highly contested because standard chronological analyses rely on scarce data of varying quality and ignore spatial complexity. Relevant archaeological and palaeontological records are most often also biased by differential preservation resulting in under-representated older events. Chronological analyses have attributed megafaunal extinctions to climate change, humans, or a combination of the two, but rarely consider spatial variation in extinction patterns, initial human appearance trajectories, and palaeoclimate change together. Here we develop a statistical approach to infer spatio-temporal trajectories of megafauna extirpations (local extinctions) and initial human appearance in south-eastern Australia. We identify a combined climate-human effect on regional extirpation patterns suggesting that small, mobile Aboriginal populations potentially needed access to drinkable water to survive arid ecosystems, but were simultaneously constrained by climate-dependent net landscape primary productivity. Thus, the co-drivers of megafauna extirpations were themselves constrained by the spatial distribution of climate-dependent water sources.

Modeling the distribution of a wide-ranging invasive species using the sampling efforts of expert and citizen scientists.

In its invasive range in Australia, the European rabbit threatens the persistence of native flora and fauna and damages agricultural production. Understanding its distribution and ecological niche is critical for developing management plans to reduce populations and avoid further biodiversity and economic losses.We developed an ensemble of species distribution models (SDMs) to determine the geographic range limits and habitat suitability of the rabbit in Australia. We examined the advantage of incorporating data collected by citizens (separately and jointly with expert data) and explored issues of spatial biases in occurrence data by implementing different approaches to generate pseudo-absences. We evaluated the skill of our model using three approaches: cross-validation, out-of-region validation, and evaluation of the covariate response curves according to expert knowledge of rabbit ecology.Combining citizen and expert occurrence data improved model skill based on cross-validation, spatially reproduced important aspects of rabbit ecology, and reduced the need to extrapolate results beyond the studied areas.Our ensemble model projects that rabbits are distributed across approximately two thirds of Australia. Annual maximum temperatures >25°C and annual minimum temperatures >10°C define, respectively, the southern and northern most range limits of its distribution. In the arid and central regions, close access to permanent water (≤~ 0.4 km) and reduced clay soil composition (~20%-50%) were the major factors influencing the probability of occurrence of rabbits. Synthesis and applications. Our results show that citizen science data can play an important role in managing invasive species by providing missing information on occurrences in regions not surveyed by experts because of logistics or financial constraints. The additional sampling effort provided by citizens can improve the capacity of SDMs to capture important elements of a species ecological niche, improving the capacity of statistical models to accurately predict the geographic range of invasive species.

Early human settlement of Sahul was not an accident.

The first peopling of Sahul (Australia, New Guinea and the Aru Islands joined at lower sea levels) by anatomically modern humans required multiple maritime crossings through Wallacea, with at least one approaching 100 km. Whether these crossings were accidental or intentional is unknown. Using coastal-viewshed analysis and ocean drift modelling combined with population projections, we show that the probability of randomly reaching Sahul by any route is <5% until ≥40 adults are 'washed off' an island at least once every 20 years. We then demonstrate that choosing a time of departure and making minimal headway (0.5 knots) toward a destination greatly increases the likelihood of arrival. While drift modelling demonstrates the existence of 'bottleneck' crossings on all routes, arrival via New Guinea is more likely than via northwestern Australia. We conclude that anatomically modern humans had the capacity to plan and make open-sea voyages lasting several days by at least 50,000 years ago.

Minimum founding populations for the first peopling of Sahul.

The timing, context and nature of the first people to enter Sahul is still poorly understood owing to a fragmented archaeological record. However, quantifying the plausible demographic context of this founding population is essential to determine how and why the initial peopling of Sahul occurred. We developed a stochastic, age-structured model using demographic rates from hunter-gatherer societies, and relative carrying capacity hindcasted with LOVECLIM's net primary productivity for northern Sahul. We projected these populations to determine the resilience and minimum sizes required to avoid extinction. A census founding population of between 1,300 and 1,550 individuals was necessary to maintain a quasi-extinction threshold of ≲0.1. This minimum founding population could have arrived at a single point in time, or through multiple voyages of ≥130 people over ~700-900 years. This result shows that substantial population amalgamation in Sunda and Wallacea in Marine Isotope Stages 3-4 provided the conditions for the successful, large-scale and probably planned peopling of Sahul.

The Australian National Rabbit Database: 50 yr of population monitoring of an invasive species.

With ongoing introductions into Australia since the 1700s, the European rabbit (Oryctolagus cuniculus) has become one of the most widely distributed and abundant vertebrate pests, adversely impacting Australia's biodiversity and agroeconomy. To understand the population and range dynamics of the species and its impacts better, occurrence and abundance data have been collected by researchers and citizens from sites covering a broad spectrum of climatic and environmental conditions in Australia. The lack of a common and accessible repository for these data has, however, limited their use in determining important spatiotemporal drivers of the structure and dynamics of the geographical range of rabbits in Australia. To meet this need, we created the Australian National Rabbit Database, which combines more than 50 yr of historical and contemporary survey data collected from throughout the range of the species in Australia. The survey data, obtained from a suite of complementary monitoring methods, were combined with high-resolution weather, climate, and environmental information, and an assessment of data quality. The database provides records of rabbit occurrence (689,265 records) and abundance (51,241 records, >120 distinct sites) suitable for identifying the spatiotemporal drivers of the rabbit's distribution and for determining spatial patterns of variation in its key life-history traits, including maximum rates of population growth. Because all data are georeferenced and date stamped, they can be coupled with information from other databases and spatial layers to explore the potential effects of rabbit occurrence and abundance on Australia's native wildlife and agricultural production. The Australian National Rabbit Database is an important tool for understanding and managing the European rabbit in its invasive range and its effects on native biodiversity and agricultural production. It also provides a valuable resource for addressing questions related to the biology, success, and impacts of invasive species more generally. No copyright or proprietary restrictions are associated with the use of this data set other than citation of this Data Paper.

High-quality fossil dates support a synchronous, Late Holocene extinction of devils and thylacines in mainland Australia.

The last large marsupial carnivores-the Tasmanian devil (Sarcophilis harrisii) and thylacine (Thylacinus cynocephalus)-went extinct on mainland Australia during the mid-Holocene. Based on the youngest fossil dates (approx. 3500 years before present, BP), these extinctions are often considered synchronous and driven by a common cause. However, many published devil dates have recently been rejected as unreliable, shifting the youngest mainland fossil age to 25 500 years BP and challenging the synchronous-extinction hypothesis. Here we provide 24 and 20 new ages for devils and thylacines, respectively, and collate existing, reliable radiocarbon dates by quality-filtering available records. We use this new dataset to estimate an extinction time for both species by applying the Gaussian-resampled, inverse-weighted McInerney (GRIWM) method. Our new data and analysis definitively support the synchronous-extinction hypothesis, estimating that the mainland devil and thylacine extinctions occurred between 3179 and 3227 years BP.

Why decadal to century timescale palaeoclimate data are needed to explain present-day patterns of biological diversity and change.

The current distribution of species, environmental conditions and their interactions represent only one snapshot of a planet that is continuously changing, in part due to human influences. To distinguish human impacts from natural factors, the magnitude and pace of climate shifts, since the Last Glacial Maximum, are often used to determine whether patterns of diversity today are artefacts of past climate change. In the absence of high-temporal resolution palaeoclimate reconstructions, this is generally done by assuming that past climate change occurred at a linear pace between widely spaced (usually, ≥1,000 years) climate snapshots. We show here that this is a flawed assumption because regional climates have changed significantly across decades and centuries during glacial-interglacial cycles, likely causing rapid regional replacement of biota. We demonstrate how recent atmosphere-ocean general circulation model (AOGCM) simulations of the climate of the past 21,000 years can provide credible estimates of the details of climate change on decadal to centennial timescales, showing that these details differ radically from what might be inferred from longer timescale information. High-temporal resolution information can provide more meaningful estimates of the magnitude and pace of climate shifts, the location and timing of drivers of physiological stress, and the extent of novel climates. They also produce new opportunities to directly investigate whether short-term climate variability is more important in shaping biodiversity patterns rather than gradual changes in long-term climatic means. Together, these more accurate measures of past climate instability are likely to bring about a better understanding of the role of palaeoclimatic change and variability in shaping current macroecological patterns in many regions of the world.

A comprehensive database of quality-rated fossil ages for Sahul's Quaternary vertebrates.

The study of palaeo-chronologies using fossil data provides evidence for past ecological and evolutionary processes, and is therefore useful for predicting patterns and impacts of future environmental change. However, the robustness of inferences made from fossil ages relies heavily on both the quantity and quality of available data. We compiled Quaternary non-human vertebrate fossil ages from Sahul published up to 2013. This, the FosSahul database, includes 9,302 fossil records from 363 deposits, for a total of 478 species within 215 genera, of which 27 are from extinct and extant megafaunal species (2,559 records). We also provide a rating of reliability of individual absolute age based on the dating protocols and association between the dated materials and the fossil remains. Our proposed rating system identified 2,422 records with high-quality ages (i.e., a reduction of 74%). There are many applications of the database, including disentangling the confounding influences of hypothetical extinction drivers, better spatial distribution estimates of species relative to palaeo-climates, and potentially identifying new areas for fossil discovery.

Where to Dig for Fossils: Combining Climate-Envelope, Taphonomy and Discovery Models.

Fossils represent invaluable data to reconstruct the past history of life, yet fossil-rich sites are often rare and difficult to find. The traditional fossil-hunting approach focuses on small areas and has not yet taken advantage of modelling techniques commonly used in ecology to account for an organism's past distributions. We propose a new method to assist finding fossils at continental scales based on modelling the past distribution of species, the geological suitability of fossil preservation and the likelihood of fossil discovery in the field, and apply it to several genera of Australian megafauna that went extinct in the Late Quaternary. Our models predicted higher fossil potentials for independent sites than for randomly selected locations (mean Kolmogorov-Smirnov statistic = 0.66). We demonstrate the utility of accounting for the distribution history of fossil taxa when trying to find the most suitable areas to look for fossils. For some genera, the probability of finding fossils based on simple climate-envelope models was higher than the probability based on models incorporating current conditions associated with fossil preservation and discovery as predictors. However, combining the outputs from climate-envelope, preservation, and discovery models resulted in the most accurate predictions of potential fossil sites at a continental scale. We proposed potential areas to discover new fossils of Diprotodon, Zygomaturus, Protemnodon, Thylacoleo, and Genyornis, and provide guidelines on how to apply our approach to assist fossil hunting in other continents and geological settings.

Climate change not to blame for late Quaternary megafauna extinctions in Australia.

Late Quaternary megafauna extinctions impoverished mammalian diversity worldwide. The causes of these extinctions in Australia are most controversial but essential to resolve, because this continent-wide event presaged similar losses that occurred thousands of years later on other continents. Here we apply a rigorous metadata analysis and new ensemble-hindcasting approach to 659 Australian megafauna fossil ages. When coupled with analysis of several high-resolution climate records, we show that megafaunal extinctions were broadly synchronous among genera and independent of climate aridity and variability in Australia over the last 120,000 years. Our results reject climate change as the primary driver of megafauna extinctions in the world's most controversial context, and instead estimate that the megafauna disappeared Australia-wide ∼13,500 years after human arrival, with shorter periods of coexistence in some regions. This is the first comprehensive approach to incorporate uncertainty in fossil ages, extinction timing and climatology, to quantify mechanisms of prehistorical extinctions.