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.

Gene drives for invasive wasp control: Extinction is unlikely, with suppression dependent on dispersal and growth rates.

Gene drives offer a potentially revolutionary method for pest control over large spatial extents. These genetic modifications spread deleterious variants through a population and have been proposed as methods for pest suppression or even eradication. We examined the influence of local dispersal, long-distance and/or human-mediated dispersal, and variation in population growth on the success of a gene drive for the control of invasive social wasps (Vespula vulgaris). Our simulations incorporated a spatially realistic environment containing variable habitat quality in New Zealand. Pest eradication was not observed, except in extreme and unrealistic scenarios of constant, widespread, and spatially intense releases of genetically modified individuals every year for decades. Instead, the regional persistence of genetically modified and wild-type wasps was predicted. Simulations using spatially homogeneous versus realistic landscapes (incorporating uninhabitable areas and dispersal barriers) showed little difference in overall population dynamics. Overall, little impact on wasp abundance was observed in the first 15 years after introduction. After 25 years, populations were suppressed to levels <95% of starting populations. Populations exhibited "chase dynamics" with population cycles in space, with local extinction occurring in some areas while wasps became abundant in others. Increasing the wasps' local dispersal distance increased the spatial and temporal variability of the occupied area and population suppression. Varying levels of human-associated long-distance dispersal had little effect on population dynamics. Increasing intrinsic population growth rates interacted with local dispersal to cause higher mean populations and substantially higher levels of variation in population suppression and the total amount of landscape occupied. Gene drives appear unlikely to cause a rapid and widespread extinction of this and probably other pests but could offer long-term and cost-effective methods of pest suppression. The predicted level of <95% pest suppression would substantially reduce the predation pressure and competitive interactions of this invasive wasp on native species. However, the predicted long-term persistence of genetically modified pests will influence the ethics and likelihood of using gene drives for pest control, especially given concerns that modified wasps would eventually be transported back to their home range.

Climatic conditions affect shoot flammability by influencing flammability-related functional traits in nonfire-prone habitats.

Plant flammability is an important driver of wildfires, and flammability itself is determined by several plant functional traits. While many plant traits are influenced by climatic conditions, the interaction between climatic conditions and plant flammability has rarely been investigated. Here, we explored the relationships among climatic conditions, shoot-level flammability components, and flammability-related functional traits for 186 plant species from fire-prone and nonfire-prone habitats. For species originating from nonfire-prone habitats, those from warmer areas tended to have lower shoot moisture content and larger leaves, and had higher shoot flammability with higher ignitibility, combustibility, and sustainability. Plants in wetter areas tended to have lower shoot flammability with lower combustibility and sustainability due to higher shoot moisture contents. In fire-prone habitats, shoot flammability was not significantly related to any climatic factor. Our study suggests that for species originating in nonfire-prone habitats, climatic conditions have influenced plant flammability by shifting flammability-related functional traits, including leaf size and shoot moisture content. Climate does not predict shoot flammability in species from fire-prone habitats; here, fire regimes may have an important role in shaping plant flammability. Understanding these nuances in the determinants of plant flammability is important in an increasingly fire-prone world.

How far might plant-eating dinosaurs have moved seeds?

Fossilized gut contents suggest that seeds consumed by dinosaurs may have remained intact in their stomachs, and since seed dispersal distance increases with body-mass in extant vertebrates, dinosaurs may have moved seeds long distances. I simulated seed dispersal by dinosaurs across body-masses from 1 × 101 to 8 × 104 kg using allometric random walk models, informed by relationships between (i) body-mass and movement speed, and (ii) body-mass and seed retention time. Seed dispersal distances showed a hump-shaped relationship with body-mass, reflecting the allometric relationship between maximum movement speed and body-mass. Across a range of assumptions and parameterizations, the simulations suggest that plant-eating dinosaurs could have dispersed seeds long distances.

Strength of niche processes for species interactions is lower for generalists and exotic species.

Niche and neutral processes jointly influence species interactions. Predictions of interactions based on these processes assume that they operate similarly across all species. However, species characteristics could systematically create differences in the strength of niche or neutral processes for each interspecific interaction. We used national-level records of plant-frugivore interactions, species traits, biogeographic status (native vs. exotic), phylogenies and species range sizes to test the hypothesis that the strength of niche processes in species interactions changes in predictable ways depending on trophic generalism and biogeographic status of the interacting species. The strength of niche processes (measured as trait matching) decreased when the generalism of the interacting partners increased. Furthermore, the slope of this negative relationship between trait matching and generalism of the interacting partners was steeper (more negative) for interactions between exotic species than those between native species. These results remained significant after accounting for the potential effects of neutral processes (estimated by species range size). These observed changes in the strength of niche processes in generating species interactions, after accounting for effects of neutral processes, could improve predictions of ecological networks from species trait data. Specifically, due to their shorter co-evolutionary history, exotic species tend to interact with native species even when lower trait matching occurs than in interactions among native species. Likewise, interactions between generalist bird species and generalist plant species should be expected to occur despite low trait matching between species, whereas interactions between specialist species involve higher trait matching.

Trait matching and phenological overlap increase the spatio-temporal stability and functionality of plant-pollinator interactions.

Morphology and phenology influence plant-pollinator network structure, but whether they generate more stable pairwise interactions with higher pollination success remains unknown. Here we evaluate the importance of morphological trait matching, phenological overlap and specialisation for the spatio-temporal stability (measured as variability) of plant-pollinator interactions and for pollination success, while controlling for species' abundance. To this end, we combined a 6-year plant-pollinator interaction dataset, with information on species traits, phenologies, specialisation, abundance and pollination success, into structural equation models. Interactions among abundant plants and pollinators with well-matched traits and phenologies formed the stable and functional backbone of the pollination network, whereas poorly matched interactions were variable in time and had lower pollination success. We conclude that phenological overlap could be more useful for predicting changes in species interactions than species abundances, and that non-random extinction of species with well-matched traits could decrease the stability of interactions within communities and reduce their functioning.

Shoot-level flammability across the Dracophyllum (Ericaceae) phylogeny: evidence for flammability being an emergent property in a land with little fire.

Plant flammability varies across species, but the evolutionary basis for this variation is not well understood. Phylogenetic analysis of interspecific variation in flammability can provide insights into the evolution of plant flammability. We measured four components of flammability (ignitability, sustainability, combustibility and consumability) to assess the shoot-level flammability of 21 species of Dracophyllum (Ericaceae). Using a macroevolutionary approach, we explored phylogenetic patterns of variation in shoot-level flammability. Shoot-level flammability varied widely in Dracophyllum. Species in the subgenus Oreothamnus had higher flammability and smaller leaves than those in the subgenus Dracophyllum. Shoot flammability (ignitability, combustibility and consumability) and leaf length showed phylogenetic conservatism across genus Dracophyllum, but exhibited lability among some closely related species, such as D. menziesii and D. fiordense. Shoot flammability of Dracophyllum species was negatively correlated with leaf length and shoot moisture content, but had no relationship with the geographic distribution of Dracophyllum species. Shoot-level flammability varied widely in the genus Dracophyllum, but showed phylogenetic conservatism. The higher flammability of the subgenus Oreothamnus may be an incidental or emergent property as a result of the evolution of flammability-related traits, such as smaller leaves, which were selected for other functions and incidentally changed flammability.

Shoot flammability of vascular plants is phylogenetically conserved and related to habitat fire-proneness and growth form.

Terrestrial plants and fire have interacted for at least 420 million years1. Whether recurrent fire drives plants to evolve higher flammability and what the evolutionary pattern of plant flammability is remain unclear2-7. Here, we show that phylogeny, the susceptibility of a habitat to have recurrent fires (that is, fire-proneness) and growth form are important predictors of the shoot flammability of 194 indigenous and introduced vascular plant species (Tracheophyta) from New Zealand. The phylogenetic signal of the flammability components and the variation in flammability among phylogenetic groups (families and higher taxonomic level clades) demonstrate that shoot flammability is phylogenetically conserved. Some closely related species, such as in Dracophyllum (Ericaceae), vary in flammability, indicating that flammability exhibits evolutionary flexibility. Species in fire-prone ecosystems tend to be more flammable than species from non-fire-prone ecosystems, suggesting that fire may have an important role in the evolution of plant flammability. Growth form also influenced flammability-forbs were less flammable than grasses, trees and shrubs; by contrast, grasses had higher biomass consumption by fire than other groups. The results show that shoot flammability of plants is largely correlated with phylogenetic relatedness, and high flammability may result in parallel evolution driven by environmental factors, such as fire regime.

Will Peri-Urban Cydia pomonella (Lepidoptera: Tortricidae) Challenge Local Eradication?

Codling moth, Cydia pomonella (Lepidoptera: Tortricidae), is a phytosanitary pest of New Zealand's export apples. The sterile insect technique supplements other controls in an eradication attempt at an isolated group of orchards in Hawke's Bay, New Zealand. There has been no attempt in New Zealand to characterize potential sources of uncontrolled peri-urban populations, which we predicted to be larger than in managed orchards. We installed 200 pheromone traps across Hastings city, which averaged 0.32 moths/trap/week. We also mapped host trees around the pilot eradication orchards and installed 28 traps in rural Ongaonga, which averaged 0.59 moths/trap/week. In Hastings, traps in host trees caught significantly more males than traps in non-host trees, and spatial interpolation showed evidence of spatial clustering. Traps in orchards operating the most stringent codling moth management averaged half the catch rate of Hastings peri-urban traps. Orchards with less rigorous moth control had a 5-fold higher trap catch rate. We conclude that peri-urban populations are significant and ubiquitous, and that special measures to reduce pest prevalence are needed to achieve area-wide suppression and reduce the risk of immigration into export orchards. Because the location of all host trees in Hastings is not known, it could be more cost-effectively assumed that hosts are ubiquitous across the city and the area treated accordingly.

Peri-Urban Community Attitudes towards Codling Moth Trapping and Suppression Using the Sterile Insect Technique in New Zealand.

New, more socially-acceptable technologies are being developed to suppress horticultural pests, because suppression is technically difficult with current technologies, especially in urban areas. One technique involves the release of sterile insects to prevent offspring in the next generation. This technology involves aerial or ground release systems, but this could also create issues for the public. This study investigated community perceptions of a recently-introduced response to codling moth control in New Zealand-Sterile Insect Technique (SIT). Community attitudes to SIT were examined in Hastings, New Zealand, in April, 2018. Eighty-six detailed interviews were undertaken with a random sample of households. This community was very willing (98% agreement) to host a sex pheromone trap in their gardens, and condoned regular visits to monitor traps. Attitudes to SIT were very positive (98% in favor). Once explained, the concept of using unmanned aerial vehicles to deliver sterile insects was also acceptable (98%) to the community. Use of unmanned aerial vehicles to release sterile insects during a hypothetical incursion response of an exotic fruit fly was also supported at 98% by respondent householders. Investigation of community attitudes can be valuable to guide practitioners in determining suitable technologies before an area-wide programme is launched.

Gametophyte niche differences among sympatric tree ferns.

Forest community assembly is usually framed in terms of sporophyte dynamics; however, the recruitment and maintenance of fern populations, frequently influential in forest composition and structure, are initially determined by gametophytes. Sporophytes of three Cyathea tree fern species show habitat partitioning along gradients of phosphorus and light; we asked whether gametophyte niche differences parallel this pattern. To compare niche characteristics among taxa we compared growth rates to a size threshold (≥3 mm) of gametophytes under controlled conditions using a multi-factorial, multi-level (3 × 4) experiment, varying irradiance (5.4 ± 4.4; 59.1 ± 44.3; 107 ± 74.1 µmol m-2 s-1) and orthophosphate concentrations (5, 10, 20, 40 mg kg-1). Gametophytes of the pioneer species C. medullaris developed to the size threshold across a broad range of phosphate and irradiance treatments (more than 20% of gametophytes in ≥ 7 of the 12 treatments), peaking at 20 mg kg-1 P and 60 µmol m-2 s-1 irradiance. The growth rates of the forest understorey species C. dealbata and C. smithii also peaked at 60 µmol m-2 s-1 but varied across treatments, suggesting niche differentiation along irradiance and orthophosphate gradients. Our analysis suggests that gametophyte development is strategically aligned to the ecological habits of sporophytes and that forest community assembly is likely strongly influenced by the independent gametophyte life-stage.

Effects of mis-alignment between dispersal traits and landscape structure on dispersal success in fragmented landscapes.

Dispersal is fundamental to population dynamics and hence extinction risk. The dispersal success of animals depends on the biophysical structure of their environments and their biological traits; however, comparatively little is known about how evolutionary trade-offs among suites of biological traits affect dispersal potential. We developed a spatially explicit agent-based simulation model to evaluate the influence of trade-offs among a suite of biological traits on the dispersal success of vagile animals in fragmented landscapes. We specifically chose traits known to influence dispersal success: speed of movement, perceptual range, risk of predation, need to forage during dispersal, and amount of suitable habitat required for successful settlement in a patch. Using the metric of relative dispersal success rate, we assessed how the costs and benefits of evolutionary investment in these biological traits varied with landscape structure. In heterogeneous environments with low habitat availability and scattered habitat patches, individuals with more equal allocation across the trait spectrum dispersed most successfully. Our analyses suggest that the dispersal success of animals in heterogeneous environments is highly dependent on hierarchical interactions between trait trade-offs and the geometric configurations of the habitat patches in the landscapes through which they disperse. In an applied sense, our results indicate potential for ecological mis-alignment between species' evolved suites of dispersal-related traits and altered environmental conditions as a result of rapid global change. In many cases identifying the processes that shape patterns of animal dispersal, and the consequences of abiotic changes for these processes, will require consideration of complex relationships among a range of organism-specific and environmental factors.

Patterns and drivers of recent disturbances across the temperate forest biome.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001-2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes.

Increasing agricultural land use is associated with the spread of an invasive fish (Gambusia affinis).

Land-use change and invasive species pose major threats to ecosystems globally. These stressors can act together, with disturbance due to changes in land-use facilitating invasion. We examined the potential for agricultural land use to facilitate the establishment and population growth (abundance) of a globally invasive fish (Gambusia affinis). To achieve this we examined Gambusia presence, abundance, and life history traits in 31 streams spanning an agricultural land use gradient in the North Island of New Zealand. We used regression models to quantify the relationship between agricultural land use and in-stream physiochemical and habitat variables, and zero-inflated models to explore the relationship among physiochemical, habitat and catchment-scale variables and Gambusia's distribution and abundance. The percentage of the catchment in agricultural land use was associated with changes to physiochemical and habitat conditions. Increasing agricultural land use was associated with increasing macrophyte cover and water temperature and decreasing velocity in streams. Catchment-scale variables (land use and site position in the network) and water temperature were the most important determinants of whether Gambusia occurred at a site. Local in-stream habitat (macrophyte cover and water velocity) and nutrient conditions were the most influential predictors of Gambusia abundance given Gambusia were present. Gambusia life-history traits, sex ratio and body length varied among sites but were not predicted by physiochemical gradients. The distribution of Gambusia in streams in New Zealand is partially controlled by catchment-scale conditions via a combination of dispersal limitation and environmental filtering, both of which are affected by agricultural land use. Agricultural land use alters local in-stream conditions, resulting in systems that are similar to those in Gambusia's natural range; these altered systems have the potential to support an increased abundance of Gambusia. This study provides preliminary quantitative evidence that agricultural land use is related to the spread of a globally invasive freshwater fish.

Pyrodiversity is the coupling of biodiversity and fire regimes in food webs.

Fire positively and negatively affects food webs across all trophic levels and guilds and influences a range of ecological processes that reinforce fire regimes, such as nutrient cycling and soil development, plant regeneration and growth, plant community assembly and dynamics, herbivory and predation. Thus we argue that rather than merely describing spatio-temporal patterns of fire regimes, pyrodiversity must be understood in terms of feedbacks between fire regimes, biodiversity and ecological processes. Humans shape pyrodiversity both directly, by manipulating the intensity, severity, frequency and extent of fires, and indirectly, by influencing the abundance and distribution of various trophic guilds through hunting and husbandry of animals, and introduction and cultivation of plant species. Conceptualizing landscape fire as deeply embedded in food webs suggests that the restoration of degraded ecosystems requires the simultaneous careful management of fire regimes and native and invasive plants and animals, and may include introducing new vertebrates to compensate for extinctions that occurred in the recent and more distant past.This article is part of the themed issue 'The interaction of fire and mankind'.

Extending the Functionality of Behavioural Change-Point Analysis with k-Means Clustering: A Case Study with the Little Penguin (Eudyptula minor).

We present a simple framework for classifying mutually exclusive behavioural states within the geospatial lifelines of animals. This method involves use of three sequentially applied statistical procedures: (1) behavioural change point analysis to partition movement trajectories into discrete bouts of same-state behaviours, based on abrupt changes in the spatio-temporal autocorrelation structure of movement parameters; (2) hierarchical multivariate cluster analysis to determine the number of different behavioural states; and (3) k-means clustering to classify inferred bouts of same-state location observations into behavioural modes. We demonstrate application of the method by analysing synthetic trajectories of known 'artificial behaviours' comprised of different correlated random walks, as well as real foraging trajectories of little penguins (Eudyptula minor) obtained by global-positioning-system telemetry. Our results show that the modelling procedure correctly classified 92.5% of all individual location observations in the synthetic trajectories, demonstrating reasonable ability to successfully discriminate behavioural modes. Most individual little penguins were found to exhibit three unique behavioural states (resting, commuting/active searching, area-restricted foraging), with variation in the timing and locations of observations apparently related to ambient light, bathymetry, and proximity to coastlines and river mouths. Addition of k-means clustering extends the utility of behavioural change point analysis, by providing a simple means through which the behaviours inferred for the location observations comprising individual movement trajectories can be objectively classified.

Feedbacks and landscape-level vegetation dynamics.

Alternative stable-state theory (ASS) is widely accepted as explaining landscape-level vegetation dynamics, such as switches between forest and grassland. This theory argues that webs of feedbacks stabilise vegetation composition and structure, and that abrupt state shifts can occur if stabilising feedbacks are weakened. However, it is difficult to identify stabilising feedback loops and the disturbance thresholds beyond which state changes occur. Here, we argue that doing this requires a synthetic approach blending observation, experimentation, simulation, conceptual models, and narratives. Using forest boundaries and large mammal extinctions, we illustrate how a multifaceted research program can advance understanding of feedback-driven ecosystem change. Our integrative approach has applicability to other complex macroecological systems controlled by numerous feedbacks where controlled experimentation is impossible.

Quantifying the direct transfer costs of common brushtail possum dispersal using least-cost modelling: a combined cost-surface and accumulated-cost dispersal kernel approach.

Dispersal costs need to be quantified from empirical data and incorporated into dispersal models to improve our understanding of the dispersal process. We are interested in quantifying how landscape features affect the immediately incurred direct costs associated with the transfer of an organism from one location to another. We propose that least-cost modelling is one method that can be used to quantify direct transfer costs. By representing the landscape as a cost-surface, which describes the costs associated with traversing different landscape features, least-cost modelling is often applied to measure connectivity between locations in accumulated-cost units that are a combination of both the distance travelled and the costs traversed. However, we take an additional step by defining an accumulated-cost dispersal kernel, which describes the probability of dispersal in accumulated-cost units. This novel combination of cost-surface and accumulated-cost dispersal kernel enables the transfer stage of dispersal to incorporate the effects of landscape features by modifying the direction of dispersal based on the cost-surface and the distance of dispersal based on the accumulated-cost dispersal kernel. We apply this approach to the common brushtail possum (Trichosurus vulpecula) within the North Island of New Zealand, demonstrating how commonly collected empirical dispersal data can be used to calibrate a cost-surface and associated accumulated-cost dispersal kernel. Our results indicate that considerable improvements could be made to the modelling of the transfer stage of possum dispersal by using a cost-surface and associated accumulated-cost dispersal kernel instead of a more traditional straight-line distance based dispersal kernel. We envisage a variety of ways in which the information from this novel combination of a cost-surface and accumulated-cost dispersal kernel could be gainfully incorporated into existing dispersal models. This would enable more realistic modelling of the direct transfer costs associated with the dispersal process, without requiring existing dispersal models to be abandoned.

Species-specific traits plus stabilizing processes best explain coexistence in biodiverse fire-prone plant communities.

Coexistence in fire-prone Mediterranean-type shrublands has been explored in the past using both neutral and niche-based models. However, distinct differences between plant functional types (PFTs), such as fire-killed vs resprouting responses to fire, and the relative similarity of species within a PFT, suggest that coexistence models might benefit from combining both neutral and niche-based (stabilizing) approaches. We developed a multispecies metacommunity model where species are grouped into two PFTs (fire-killed vs resprouting) to investigate the roles of neutral and stabilizing processes on species richness and rank-abundance distributions. Our results show that species richness can be maintained in two ways: i) strictly neutral species within each PFT, or ii) species within PFTs differing in key demographic properties, provided that additional stabilizing processes, such as negative density regulation, also operate. However, only simulations including stabilizing processes resulted in structurally realistic rank-abundance distributions over plausible time scales. This result underscores the importance of including both key species traits and stabilizing (niche) processes in explaining species coexistence and community structure.