Reconstructing colonization dynamics to establish how human activities transformed island biodiversity.

Drivers and dynamics of initial human migrations across individual islands and archipelagos are poorly understood, hampering assessments of subsequent modification of island biodiversity. We developed and tested a new statistical-simulation approach for reconstructing the pattern and pace of human migration across islands at high spatiotemporal resolutions. Using Polynesian colonisation of New Zealand as an example, we show that process-explicit models, informed by archaeological records and spatiotemporal reconstructions of past climates and environments, can provide new and important insights into the patterns and mechanisms of arrival and establishment of people on islands. We find that colonisation of New Zealand required there to have been a single founding population of approximately 500 people, arriving between 1233 and 1257 AD, settling multiple areas, and expanding rapidly over both North and South Islands. These verified spatiotemporal reconstructions of colonisation dynamics provide new opportunities to explore more extensively the potential ecological impacts of human colonisation on New Zealand's native biota and ecosystems.

Biomonitoring tool for New Zealand peatlands: Testate amoebae and vascular plants as promising bioindicators.

In the last two centuries, a high proportion of peatlands have been lost or severely degraded across the world. The value of peatlands is now well-recognised for biodiversity conservation, flood management, and carbon mitigation, with peatland restoration now central to many government policies for climate action. A challenge, however, is to determine 'natural' and 'disturbed' conditions of peatlands to establish realistic baselines for assessing degradation and setting restoration targets. This requires a tool or set of tools that can rapidly and reliably capture peatland condition across space and time. Our aim was to develop such a tool based on combined analysis of plant and testate amoebae; a group of shelled protists commonly used as indicators of ecological change in peatlands. The value of testate amoebae is well established in Northern Hemisphere Sphagnum-dominated peatlands; however, relatively little work has been undertaken for Southern Hemisphere peat forming systems. Here we provide the first assessment and comparison of the bioindicator value of testate amoebae and vascular plants in the context of Southern Hemisphere peatlands. Our results further demonstrate the unique ecohydrological dynamics at play in New Zealand peat forming systems that set them apart from Northern Hemisphere peatlands. Our results show that plant and testate amoeba communities provided valuable information on peatland condition at different scales, we found that testate amoebae tracked changes in the abiotic variables (depth to water table, pH, and conductivity) more closely than vascular plants. Our results further demonstrate that functional traits of testate amoebae showed promising relationships with disturbance. Amoeba test compression, aperture position and test size were linked to changes in hydrology driven by fluctuations in ground water tables; however, trait responses manifested differently in ombrotrophic and minerotrophic peatlands. Overall, testate amoebae provide a promising bioindicator for tracking degradation in New Zealand peatlands and a potential additional tool to assess peatland condition.

Floristic homogenization of South Pacific islands commenced with human arrival.

The increasing similarity of plant species composition among distinct areas is leading to the homogenization of ecosystems globally. Human actions such as ecosystem modification, the introduction of non-native plant species and the extinction or extirpation of endemic and native plant species are considered the main drivers of this trend. However, little is known about when floristic homogenization began or about pre-human patterns of floristic similarity. Here we investigate vegetation trends during the past 5,000 years across the tropical, sub-tropical and warm temperate South Pacific using fossil pollen records from 15 sites on 13 islands within the biogeographical realm of Oceania. The site comparisons show that floristic homogenization has increased over the past 5,000 years. Pairwise Bray-Curtis similarity results also show that when two islands were settled by people in a given time interval, their floristic similarity is greater than when one or neither of the islands were settled. Importantly, higher elevation sites, which are less likely to have experienced human impacts, tended to show less floristic homogenization. While biotic homogenization is often referred to as a contemporary issue, we have identified a much earlier trend, likely driven by human colonization of the islands and subsequent impacts.

Long-term trajectories of non-native vegetation on islands globally.

Human-mediated changes in island vegetation are, among others, largely caused by the introduction and establishment of non-native species. However, data on past changes in non-native plant species abundance that predate historical documentation and censuses are scarce. Islands are among the few places where we can track human arrival in natural systems allowing us to reveal changes in vegetation dynamics with the arrival of non-native species. We matched fossil pollen data with botanical status information (native, non-native), and quantified the timing, trajectories and magnitude of non-native plant vegetational change on 29 islands over the past 5000 years. We recorded a proportional increase in pollen of non-native plant taxa within the last 1000 years. Individual island trajectories are context-dependent and linked to island settlement histories. Our data show that non-native plant introductions have a longer and more dynamic history than is generally recognized, with critical implications for biodiversity baselines and invasion biology.

The testate amoebae of New Zealand: A checklist, identification key and assessment of biogeographic patterns.

New Zealand (NZ) is a well-known hotspot of biodiversity and endemism for macroscopic organisms, but its microbial diversity is comparatively poorly documented. We assembled all records on NZ testate amoebae published since the early 20th century and present a comprehensive taxonomic checklist for NZ. Testate amoebae are reported from six major habitat types across both the North and South Islands of NZ, but the sampling effort is ecologically and geographically biased in favour of wetlands and the South Island. As a result, 93% of all 128 morphotypes recorded in NZ occur in wetlands, 28% are restricted to the South Island, and diversity is greater at higher latitudes. Around 50% of morphotypes have a broad latitudinal distribution across the NZ mainland, whereas 15% have narrow latitudinal ranges. Future research should aim to broaden the geographical and ecological ranges. We predict that our list of NZ testate amoebae will expand substantially with future work, and that the latitudinal diversity gradient will be inverted. We also introduce an interactive, fully illustrated, online Lucid key for the rapid identification of NZ testate amoebae. As many morphospecies are cosmopolitan, this key provides a useful tool for testate amoebae identification in other parts of the world.

The human dimension of biodiversity changes on islands.

Islands are among the last regions on Earth settled and transformed by human activities, and they provide replicated model systems for analysis of how people affect ecological functions. By analyzing 27 representative fossil pollen sequences encompassing the past 5000 years from islands globally, we quantified the rates of vegetation compositional change before and after human arrival. After human arrival, rates of turnover accelerate by a median factor of 11, with faster rates on islands colonized in the past 1500 years than for those colonized earlier. This global anthropogenic acceleration in turnover suggests that islands are on trajectories of continuing change. Strategies for biodiversity conservation and ecosystem restoration must acknowledge the long duration of human impacts and the degree to which ecological changes today differ from prehuman dynamics.

Vertical distribution of prokaryotes communities and predicted metabolic pathways in New Zealand wetlands, and potential for environmental DNA indicators of wetland condition.

Globally, wetlands are in decline due to anthropogenic modification and climate change. Knowledge about the spatial distribution of biodiversity and biological processes within wetlands provides essential baseline data for predicting and mitigating the effects of present and future environmental change on these critical ecosystems. To explore the potential for environmental DNA (eDNA) to provide such insights, we used 16S rRNA metabarcoding to characterise prokaryote communities and predict the distribution of prokaryote metabolic pathways in peats and sediments up to 4m below the surface across seven New Zealand wetlands. Our results reveal distinct vertical structuring of prokaryote communities and metabolic pathways in these wetlands. We also find evidence for differences in the relative abundance of certain metabolic pathways that may correspond to the degree of anthropogenic modification the wetlands have experienced. These patterns, specifically those for pathways related to aerobic respiration and the carbon cycle, can be explained predominantly by the expected effects of wetland drainage. Our study demonstrates that eDNA has the potential to be an important new tool for the assessment and monitoring of wetland health.

Using paleo-archives to safeguard biodiversity under climate change.

Strategies for 21st-century environmental management and conservation under global change require a strong understanding of the biological mechanisms that mediate responses to climate- and human-driven change to successfully mitigate range contractions, extinctions, and the degradation of ecosystem services. Biodiversity responses to past rapid warming events can be followed in situ and over extended periods, using cross-disciplinary approaches that provide cost-effective and scalable information for species' conservation and the maintenance of resilient ecosystems in many bioregions. Beyond the intrinsic knowledge gain such integrative research will increasingly provide the context, tools, and relevant case studies to assist in mitigating climate-driven biodiversity losses in the 21st century and beyond.

Comparing the effects of asynchronous herbivores on New Zealand montane vegetation communities.

Large herbivores facilitate a range of important ecological processes yet globally have experienced high rates of decline and extinction over the past 50,000 years. To some extent this lost function may be replaced through the introduction of ecological surrogate taxa, either by active management or via historic introductions. However, comparing the ecological effects of herbivores that existed in the same location, but at different times, can be a challenging proposition. Here we provide an example from New Zealand that demonstrates an approach for making such comparisons. In New Zealand it has been suggested that post-19th Century mammal introductions (e.g. deer and hare) may have filled ecological niches left vacant after the 15th Century AD extinction of large avian herbivores (moa). We quantified pollen assemblages from fecal samples deposited by these two asynchronous herbivore communities to see whether they were comparable. The fecal samples were collected at the same location, and in a native-dominated vegetation community that has experience little anthropogenic disturbance and their contents reflect both the local habitat and diet preferences of the depositing herbivore. The results reveal that the current forest understory is relatively sparse and species depauperate compared to the prehistoric state, indicating that deer and moa had quite different impacts on the local vegetation community. The study provides an example of how combining coprolite and fecal analyses of prehistoric and modern herbivores may clarify the degree of ecological overlap between asynchronous herbivore communities and provide insights into the extent of ecological surrogacy provided by introduced taxa.

Mitogenomes Uncover Extinct Penguin Taxa and Reveal Island Formation as a Key Driver of Speciation.

The emergence of islands has been linked to spectacular radiations of diverse organisms. Although penguins spend much of their lives at sea, they rely on land for nesting, and a high proportion of extant species are endemic to geologically young islands. Islands may thus have been crucial to the evolutionary diversification of penguins. We test this hypothesis using a fossil-calibrated phylogeny of mitochondrial genomes (mitogenomes) from all extant and recently extinct penguin taxa. Our temporal analysis demonstrates that numerous recent island-endemic penguin taxa diverged following the formation of their islands during the Plio-Pleistocene, including the Galápagos (Galápagos Islands), northern rockhopper (Gough Island), erect-crested (Antipodes Islands), Snares crested (Snares) and royal (Macquarie Island) penguins. Our analysis also reveals two new recently extinct island-endemic penguin taxa from New Zealand's Chatham Islands: Eudyptes warhami sp. nov. and a dwarf subspecies of the yellow-eyed penguin, Megadyptes antipodes richdalei ssp. nov. Eudyptes warhami diverged from the Antipodes Islands erect-crested penguin between 1.1 and 2.5 Ma, shortly after the emergence of the Chatham Islands (∼3 Ma). This new finding of recently evolved taxa on this young archipelago provides further evidence that the radiation of penguins over the last 5 Ma has been linked to island emergence. Mitogenomic analyses of all penguin species, and the discovery of two new extinct penguin taxa, highlight the importance of island formation in the diversification of penguins, as well as the extent to which anthropogenic extinctions have affected island-endemic taxa across the Southern Hemisphere's isolated archipelagos.

Multiscale climate change impacts on plant diversity in the Atacama Desert.

Comprehending ecological dynamics requires not only knowledge of modern communities but also detailed reconstructions of ecosystem history. Ancient DNA (aDNA) metabarcoding allows biodiversity responses to major climatic change to be explored at different spatial and temporal scales. We extracted aDNA preserved in fossil rodent middens to reconstruct late Quaternary vegetation dynamics in the hyperarid Atacama Desert. By comparing our paleo-informed millennial record with contemporary observations of interannual variations in diversity, we show local plant communities behave differentially at different timescales. In the interannual (years to decades) time frame, only annual herbaceous expand and contract their distributional ranges (emerging from persistent seed banks) in response to precipitation, whereas perennials distribution appears to be extraordinarily resilient. In contrast, at longer timescales (thousands of years) many perennial species were displaced up to 1,000 m downslope during pluvial events. Given ongoing and future natural and anthropogenically induced climate change, our results not only provide baselines for vegetation in the Atacama Desert, but also help to inform how these and other high mountain plant communities may respond to fluctuations of climate in the future.

Assessing the ecological value of small testate amoebae (<45 µm) in New Zealand peatlands.

Methodological advances are essential for robust ecological research. Quantitative reconstructions of environmental conditions using testate amoebae rely on sound taxonomy. While the taxonomy of large species is relatively well resolved, this is not the case for most small taxa (typically <45 µm long). In New Zealand, peatlands contain a diversity of both cosmopolitan and characteristic large southern endemic taxa, but also have a high abundance of small taxa. The latter are often lumped into morphotypes reducing their value as ecological indicators. In this study, we demonstrate how (a) lumping small taxa versus splitting them into unique types, and (b) including or excluding them from community analysis influenced their ecological inference. We assessed testate amoeba composition in six peat bogs from New Zealand, three that were moderately-to-highly impacted, and three that were non-impacted. Environmental variables were measured at each sampling site and the surface testate amoeba community patterns and community-environment relationships compared. We found a clear division between impacted and non-impacted sites. Several distinct small taxa were more strongly related to water-table depth and conductivity, while the larger taxa were more correlated to pH. These results show that improved taxonomic resolution of small taxa can provide more informed environmental assessment.

Plant pathogen responses to Late Pleistocene and Holocene climate change in the central Atacama Desert, Chile.

Future climate change has the potential to alter the distribution and prevalence of plant pathogens, which may have significant implications for both agricultural crops and natural plant communities. However, there are few long-term datasets against which modelled predictions of pathogen responses to climate change can be tested. Here, we use 18S metabarcoding of 28 rodent middens (solidified deposits of rodent coprolites and nesting material) from the Central Atacama, spanning the last ca. 49 ka, to provide the first long-term late Quaternary record of change in plant pathogen communities in response to changing climate. Plant pathogen richness was significantly greater in middens deposited during the Central Andean Pluvial Event (CAPE); a period of increased precipitation between 17.5-8.5 ka. Moreover, the occurrence frequency of Pucciniaceae (rust fungi) was significantly greater during the CAPE, and the highest relative abundances for five additional potentially pathogenic taxa also occurred during this period. The results demonstrate the promising potential for ancient DNA analysis of late Quaternary samples to reveal insights into how plant pathogens responded to past climatic and environmental change, which could help predict how pathogens may responded to future change.

An avian seed dispersal paradox: New Zealand's extinct megafaunal birds did not disperse large seeds.

Often the mutualistic roles of extinct species are inferred based on plausible assumptions, but sometimes palaeoecological evidence can overturn such inferences. We present an example from New Zealand, where it has been widely assumed that some of the largest-seeded plants were dispersed by the giant extinct herbivorous moa (Dinornithiformes). The presence of large seeds in preserved moa gizzard contents supported this hypothesis, and five slow-germinating plant species (Elaeocarpus dentatus, E. hookerianus, Prumnopitys ferruginea, P. taxifolia, Vitex lucens) with thick seedcoats prompted speculation about whether these plants were adapted for moa dispersal. However, we demonstrate that all these assumptions are incorrect. While large seeds were present in 48% of moa gizzards analysed, analysis of 152 moa coprolites (subfossil faeces) revealed a very fine-grained consistency unparalleled in extant herbivores, with no intact seeds larger than 3.3 mm diameter. Secondly, prolonged experimental mechanical scarification of E. dentatus and P. ferruginea seeds did not reduce time to germination, providing no experimental support for the hypothesis that present-day slow germination results from the loss of scarification in moa guts. Paradoxically, although moa were New Zealand's largest native herbivores, the only seeds to survive moa gut passage intact were those of small-seeded herbs and shrubs.

Global Peak in Atmospheric Radiocarbon Provides a Potential Definition for the Onset of the Anthropocene Epoch in 1965.

Anthropogenic activity is now recognised as having profoundly and permanently altered the Earth system, suggesting we have entered a human-dominated geological epoch, the 'Anthropocene'. To formally define the onset of the Anthropocene, a synchronous global signature within geological-forming materials is required. Here we report a series of precisely-dated tree-ring records from Campbell Island (Southern Ocean) that capture peak atmospheric radiocarbon (14C) resulting from Northern Hemisphere-dominated thermonuclear bomb tests during the 1950s and 1960s. The only alien tree on the island, a Sitka spruce (Picea sitchensis), allows us to seasonally-resolve Southern Hemisphere atmospheric 14C, demonstrating the 'bomb peak' in this remote and pristine location occurred in the last-quarter of 1965 (October-December), coincident with the broader changes associated with the post-World War II 'Great Acceleration' in industrial capacity and consumption. Our findings provide a precisely-resolved potential Global Stratotype Section and Point (GSSP) or 'golden spike', marking the onset of the Anthropocene Epoch.

Island biodiversity conservation needs palaeoecology.

The discovery and colonization of islands by humans has invariably resulted in their widespread ecological transformation. The small and isolated populations of many island taxa, and their evolution in the absence of humans and their introduced taxa, mean that they are particularly vulnerable to human activities. Consequently, even the most degraded islands are a focus for restoration, eradication, and monitoring programmes to protect the remaining endemic and/or relict populations. Here, we build a framework that incorporates an assessment of the degree of change from multiple baseline reference periods using long-term ecological data. The use of multiple reference points may provide information on both the variability of natural systems and responses to successive waves of cultural transformation of island ecosystems, involving, for example, the alteration of fire and grazing regimes and the introduction of non-native species. We provide exemplification of how such approaches can provide valuable information for biodiversity conservation managers of island ecosystems.

Biodiversity losses and conservation responses in the Anthropocene.

Biodiversity is essential to human well-being, but people have been reducing biodiversity throughout human history. Loss of species and degradation of ecosystems are likely to further accelerate in the coming years. Our understanding of this crisis is now clear, and world leaders have pledged to avert it. Nonetheless, global goals to reduce the rate of biodiversity loss have mostly not been achieved. However, many examples of conservation success show that losses can be halted and even reversed. Building on these lessons to turn the tide of biodiversity loss will require bold and innovative action to transform historical relationships between human populations and nature.

Origin and timing of New Zealand's earliest domestic chickens: Polynesian commensals or European introductions?

Human settlers transported chickens (Gallus gallus domesticus) to most East Polynesian archipelagos between AD 1000 and 1300; however, it has long been assumed that New Zealand was an exception. Despite the fact that chicken bones have been recovered from localities of early archaeological middens in New Zealand, their age and genetic relationships have never been critically assessed. Here, we test the assumption that chickens were not introduced to New Zealand during prehistory through ancient DNA and radiocarbon analyses of chicken bones from sites of Maori middens containing prehistoric material. The chickens belong to the widespread mitochondrial control region haplogroup E. Radiocarbon dating reveals that the bones are not prehistoric, but are still the earliest chicken remains known from New Zealand. Two of the bones pre-date permanent European settlement (ca 1803s onwards) but overlap with the arrival of James Cook's second voyage (1773-1774), and, therefore, they are likely to be chickens, or progeny thereof, liberated during that voyage. Our results support the idea that chickens were first introduced to New Zealand by Europeans, and provide new insights into Maori uptake and integration of resources introduced during the early post-European period.

Functional and environmental determinants of bark thickness in fire-free temperate rain forest communities.

PREMISE OF THE STUDY: In fire-prone ecosystems, variation in bark thickness among species and communities has been explained by fire frequency; thick bark is necessary to protect cambium from lethal temperatures. Elsewhere this investment is deemed unnecessary, and thin bark is thought to prevail. However, in rain forest ecosystems where fire is rare, bark thickness varies widely among species and communities, and the causes of this variation remain enigmatic. We tested for functional explanations of bark thickness variation in temperate rain forest species and communities. METHODS: We measured bark thickness in 82 tree species throughout New Zealand temperate rain forests that historically have experienced little fire and applied two complementary analyses. First, we examined correlations between bark traits and leaf habit, and leaf and stem traits. Second, we calculated community-weighted mean (CWM) bark thickness for 272 plots distributed throughout New Zealand to identify the environments in which thicker-barked communities occur. KEY RESULTS: Conifers had higher size-independent bark thickness than evergreen angiosperms. Species with thicker bark or higher bark allocation coefficients were not associated with "slow economic" plant traits. Across 272 forest plots, communities with thicker bark occurred on infertile soils, and communities with thicker bark and higher bark allocation coefficients occurred in cooler, drier climates. CONCLUSIONS: In non-fire-prone temperate rain forest ecosystems, investment in bark is driven by soil resources, cool minimum temperatures, and seasonal moisture stress. The role of these factors in fire-prone ecosystems warrants testing.

Long-term ecology resolves the timing, region of origin and process of establishment for a disputed alien tree.

Alien plants are a pervasive environmental problem, particularly on islands where they can rapidly transform unique indigenous ecosystems. However, often it is difficult to confidently determine whether a species is native or alien, especially if establishment occurred before historical records. This can present a management challenge: for example, should such taxa be eradicated or left alone until their region of origin and status are clarified? Here we show how combining palaeoecological and historical records can help resolve such dilemmas, using the tree daisy Olearia lyallii on the remote New Zealand subantarctic Auckland Islands as a case study. The status of this tree as native or introduced has remained uncertain for the 175 years since it was first discovered on the Auckland Islands, and its appropriate management is debated. Elsewhere, O. lyallii has a highly restricted distribution on small sea bird-rich islands within a 2° latitudinal band south of mainland New Zealand. Analysis of palaeoecological and historical records from the Auckland Islands suggest that O. lyallii established there c. 1807 when these islands were first exploited by European sealers. Establishment was facilitated by anthropogenic burning and clearing and its subsequent spread has been slow, limited in distribution and probably human-assisted. Olearia lyallii has succeeded mostly in highly disturbed sites which are also nutrient enriched from nesting sea birds, seals and sea spray. This marine subsidy has fuelled the rapid growth of O. lyallii and allowed this tree to be competitive against the maritime communities it has replaced. Although endemic to the New Zealand region, our evidence suggests that O. lyallii is alien to the Auckland Islands. Although such 'native' aliens can pose unique management challenges on islands, in this instance we suggest that ongoing monitoring with no control is an appropriate management action, as O. lyallii appears to pose minimal risk to ecological integrity.