Boom-bust population dynamics drive rapid genetic change.

Increasing environmental threats and more extreme environmental perturbations place species at risk of population declines, with associated loss of genetic diversity and evolutionary potential. While theory shows that rapid population declines can cause loss of genetic diversity, populations in some environments, like Australia's arid zone, are repeatedly subject to major population fluctuations yet persist and appear able to maintain genetic diversity. Here, we use repeated population sampling over 13 y and genotype-by-sequencing of 1903 individuals to investigate the genetic consequences of repeated population fluctuations in two small mammals in the Australian arid zone. The sandy inland mouse (Pseudomys hermannsburgensis) experiences marked boom-bust population dynamics in response to the highly variable desert environment. We show that heterozygosity levels declined, and population differentiation (FST) increased, during bust periods when populations became small and isolated, but that heterozygosity was rapidly restored during episodic population booms. In contrast, the lesser hairy-footed dunnart (Sminthopsis youngsoni), a desert marsupial that maintains relatively stable population sizes, showed no linear declines in heterozygosity. These results reveal two contrasting ways in which genetic diversity is maintained in highly variable environments. In one species, diversity is conserved through the maintenance of stable population sizes across time. In the other species, diversity is conserved through rapid genetic mixing during population booms that restores heterozygosity lost during population busts.

slimr: An R package for tailor-made integrations of data in population genomic simulations over space and time.

Software for realistically simulating complex population genomic processes is revolutionizing our understanding of evolutionary processes, and providing novel opportunities for integrating empirical data with simulations. However, the integration between standalone simulation software and R is currently not well developed. Here, we present slimr, an R package designed to create a seamless link between standalone software SLiM >3.0, one of the most powerful population genomic simulation frameworks, and the R development environment, with its powerful data manipulation and analysis tools. We show how slimr facilitates smooth integration between genetic data, ecological data and simulation in a single environment. The package enables pipelines that begin with data reading, cleaning and manipulation, proceed to constructing empirically based parameters and initial conditions for simulations, then to running numerical simulations and finally to retrieving simulation results in a format suitable for comparisons with empirical data - aided by advanced analysis and visualization tools provided by R. We demonstrate the use of slimr with an example from our own work on the landscape population genomics of desert mammals, highlighting the advantage of having a single integrated tool for both data analysis and simulation. slimr makes the powerful simulation ability of SLiM directly accessible to R users, allowing integrated simulation projects that incorporate empirical data without the need to switch between software environments. This should provide more opportunities for evolutionary biologists and ecologists to use realistic simulations to better understand the interplay between ecological and evolutionary processes.

Episodic population fragmentation and gene flow reveal a trade-off between heterozygosity and allelic richness.

In episodic environments like deserts, populations of some animal species exhibit irregular fluctuations such that populations are alternately large and connected or small and isolated. Such dynamics are typically driven by periodic resource pulses due, for example, to large but infrequent rainfall events. The repeated population bottlenecks resulting from fragmentation should lower genetic diversity over time, yet species undergoing these fluctuations appear to maintain high levels of genetic diversity. To resolve this apparent paradox, we simulated a metapopulation of constant size undergoing repeat episodes of fragmentation and change in gene flow to mimic outcomes experienced by mammals in an Australian desert. We show that episodic fragmentation and gene flow have contrasting effects on two measures of genetic diversity: heterozygosity and allelic richness. Specifically, fragmentation into many, small subpopulations, coupled with periods of infrequent gene flow, preserves allelic richness at the expense of heterozygosity. In contrast, fragmentation into a few, large subpopulations maintains heterozygosity at the expense of allelic richness. The strength of the trade-off between heterozygosity and allelic richness depends on the amount of gene flow and the frequency of gene flow events. Our results imply that the type of genetic diversity maintained among species living in strongly fluctuating environments will depend on the way populations fragment, with our results highlighting different mechanisms for maintaining allelic richness and heterozygosity in small, fragmented populations.

Long-term livestock exclusion increases plant richness and reproductive capacity in arid woodlands.

Herbivore exclusion is implemented globally to recover ecosystems from grazing by introduced and native herbivores, but evidence for large-scale biodiversity benefits is inconsistent in arid ecosystems. We examined the effects of livestock exclusion on dryland plant richness and reproductive capacity. We collected data on plant species richness and seeding (reproductive capacity), rainfall, vegetation productivity and cover, soil strength and herbivore grazing intensity from 68 sites across 6500 km2 of arid Georgina gidgee (Acacia georginae) woodlands in central Australia between 2018 and 2020. Sites were on an actively grazed cattle station and two destocked conservation reserves. We used structural equation modeling to examine indirect (via soil or vegetation modification) versus direct (herbivory) effects of grazing intensity by two introduced herbivores (cattle, camels) and a native herbivore (red kangaroo), on seasonal plant species richness and seeding of all plants, and the richness and seeding of four plant groups (native grasses, forbs, annual chenopod shrubs, and palatable perennial shrubs). Non-native herbivores had a strong indirect effect on plant richness and seeding by reducing vegetative ground cover, resulting in decreased richness and seeding of native grasses and forbs. Herbivores also had small but negative direct impacts on plant richness and seeding. This direct effect was explained by reductions in annual chenopod and palatable perennial shrub richness under grazing activity. Responses to grazing were herbivore-dependent; introduced herbivore grazing reduced native plant richness and seeding, while native herbivore grazing had no significant effect on richness or seeding of different plant functional groups. Soil strength decreased under grazing by cattle but not camels or kangaroos. Cattle had direct effects on palatable perennial shrub richness and seeding, whereas camels had indirect effects, reducing richness and seeding by reducing the abundance of shrubs. We show that considering indirect pathways improves evaluations of the effects of disturbances on biodiversity, as focusing only on direct effects can mask critical mechanisms of change. Our results indicate substantial biodiversity benefits from excluding livestock and controlling camels in drylands. Reducing introduced herbivore impacts will improve soil and vegetation condition, ensure reproduction and seasonal persistence of species, and protect native plant diversity.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Top-down response to spatial variation in productivity and bottom-up response to temporal variation in productivity in a long-term study of desert ants.

Under the Ecosystem Exploitation Hypothesis ecosystem productivity predicts trophic complexity, but it is unclear if spatial and temporal drivers of productivity have similar impacts. Long-term studies are necessary to capture temporal impacts on trophic structure in variable ecosystems such as deserts. We sampled ants and measured plant resources in the Simpson Desert, central Australia over a 22-year period, during which rainfall varied 10-fold. We sampled dune swales (higher nutrient) and crests (lower nutrient) to account for spatial variation in productivity. We asked how temporal and spatial variation in productivity affects the abundance of ant trophic guilds. Precipitation increased vegetation cover, with the difference more pronounced on dune crests; seeding and flowering also increased with precipitation. Generalist activity increased over time, irrespective of productivity. Predators were more active in more productive (swale) habitat, i.e. spatial impacts of productivity were greatest at the highest trophic level. By contrast, herbivores (seed harvesters and sugar feeders) increased with long-term rainfall; seed harvesters also increased as seeding increased. Temporal impacts of productivity were therefore greatest for low trophic levels. Whether productivity variation leads to top-down or bottom-up structured ecosystems thus depends on the scale and dimension (spatial or temporal) of productivity.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Characterising the spatiotemporal dynamics of drought and wet events in Australia.

Global climate change has altered precipitation patterns and disrupted the characteristics of drought and rainfall events. Climate projections confirm that more frequent, intense, and extreme droughts and rainfall events will continue. However, knowledge around how drought and wet events move dynamically through space and time is limited, especially in the southern hemisphere. Australia is the driest inhabited continent, renowned as the land of droughts and flooding rains, but recent climate-driven changes to the severity of wildfires and floods have garnered global attention. Here we used S-TRACK, a novel method for spatial drought tracking, to build pathways for past drought and wet events in Australia to examine their spatiotemporal dynamics. Characteristics such as duration, severity, and intensity were obtained from these pathways, and modified Mann-Kendall tests and Sen's slope were used to detect significant trends in characteristics over time. Drought conditions in southern Australia have intensified, particularly in the southwest of Australia and Tasmania, while the north of the country is experiencing longer, more severe, and more intense wet conditions. We also found that the location of drought and wet hotspots has clearly shifted in response to precipitation changes since the 1970's. Finally, pathways for the most extreme events show peak severity is reached in the middle to late stages of pathways, and that the largest drought and wet areas of a pathway have moved further west in recent times. The findings in this study provide the necessary knowledge to improve preparedness for extreme precipitation events as they become more common and to inform predictions for agricultural output or the extent of other climate events such as wildfires and flooding.

Soil properties as key predictors of global grassland production: Have we overlooked micronutrients?

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

Phenotypic plasticity masks range-wide genetic differentiation for vegetative but not reproductive traits in a short-lived plant.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

Combating ecosystem collapse from the tropics to the Antarctic.

Globally, collapse of ecosystems-potentially irreversible change to ecosystem structure, composition and function-imperils biodiversity, human health and well-being. We examine the current state and recent trajectories of 19 ecosystems, spanning 58° of latitude across 7.7 M km2 , from Australia's coral reefs to terrestrial Antarctica. Pressures from global climate change and regional human impacts, occurring as chronic 'presses' and/or acute 'pulses', drive ecosystem collapse. Ecosystem responses to 5-17 pressures were categorised as four collapse profiles-abrupt, smooth, stepped and fluctuating. The manifestation of widespread ecosystem collapse is a stark warning of the necessity to take action. We present a three-step assessment and management framework (3As Pathway Awareness, Anticipation and Action) to aid strategic and effective mitigation to alleviate further degradation to help secure our future.

General destabilizing effects of eutrophication on grassland productivity at multiple spatial scales.

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities.

Effective ecosystem monitoring requires a multi-scaled approach.

Ecosystem monitoring is fundamental to our understanding of how ecosystem change is impacting our natural resources and is vital for developing evidence-based policy and management. However, the different types of ecosystem monitoring, along with their recommended applications, are often poorly understood and contentious. Varying definitions and strict adherence to a specific monitoring type can inhibit effective ecosystem monitoring, leading to poor program development, implementation and outcomes. In an effort to develop a more consistent and clear understanding of ecosystem monitoring programs, we here review the main types of monitoring and recommend the widespread adoption of three classifications of monitoring, namely, targeted, surveillance and landscape monitoring. Landscape monitoring is conducted over large areas, provides spatial data, and enables questions relating to where and when ecosystem change is occurring to be addressed. Surveillance monitoring uses standardised field methods to inform on what is changing in our environments and the direction and magnitude of that change, whilst targeted monitoring is designed around testable hypotheses over defined areas and is the best approach for determining the causes of ecosystem change. The classification system is flexible and can incorporate different interests, objectives, targets and characteristics as well as different spatial scales and temporal frequencies, while also providing valuable structure and consistency across distinct ecosystem monitoring programs. To support our argument, we examine the ability of each monitoring type to inform on six key types of questions that are routinely posed for ecosystem monitoring programs, such as where and when change is occurring, what is the magnitude of change, and how can the change be managed? As we demonstrate, each type of ecosystem monitoring has its own strengths and weaknesses, which should be carefully considered relative to the desired results. Using this scheme, scientists and land managers can design programs best suited to their needs. Finally, we assert that for our most serious environmental challenges, it is essential that we include information from each of these monitoring scales to inform on all facets of ecosystem change, and this is best achieved through close collaboration between the scales. With a renewed understanding of the importance of each monitoring type, along with greater commitment to monitor cooperatively, we will be well placed to address some of our greatest environmental challenges.

Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains.

Global gene flow releases invasive plants from environmental constraints on genetic diversity.

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area.

Long-term responses of desert ant assemblages to climate.

Productivity is a key driver of ecosystem structure and function, so long-term studies are critical to understanding ecosystems with high temporal variation in productivity. In some deserts, productivity, driven by moisture availability, varies immensely over time (rainfall) and space (landscape factors). At high productivity, species richness is expected to be driven in opposing directions by abundance (More Individuals Hypothesis - MIH) and competition. While studies investigating the impacts of spatial variation in productivity on community structure are common, the impacts of temporal variability on productivity are poorly understood. We tested how well rainfall predicted the activity, species numbers and assemblage composition of ants and if responses were moderated by landscape position. We also asked whether the number of species (richness per sampling unit and estimated species richness) responded directly to rainfall or was moderated by ant activity or competition from dominant ants. Over a 22-year period, when annual rainfall fluctuated between 79 mm and 570 mm, we sampled ants using pitfall traps in paired dune and swale habitats in the Simpson Desert, Australia. We used climate records over this period to model changes in ant assemblages. Activity of dominant ants responded primarily to long-term rainfall, increasing exponentially, while subordinate ants responded to short-term weather and time. Consistent with the MIH, the number of ant species was best predicted by activity, particularly that of subordinate ants. Activity of dominant ants had a declining positive effect on numbers of species. Landscape position strongly predicted species composition, while long-term rainfall determined composition at genus level but not species level. Over time, species composition fluctuated, but several genera consistently increased in activity. Productivity moderators such as long-term rainfall and landscape position are key drivers of ant activity and composition in the study ecosystem, acting indirectly on numbers of species. Numbers of species were explained largely by ant activity, making a strong case for the MIH, but not competition. Longer periods of low rainfall may indirectly reduce species richness in desert ecosystems. However, a trend to increasing richness over time may indicate that conservation management can ameliorate this impact.

A decision tree for assessing the risks and benefits of publishing biodiversity data.

Inadequate information on the geographical distribution of biodiversity hampers decision-making for conservation. Major efforts are underway to fill knowledge gaps, but there are increasing concerns that publishing the locations of species is dangerous, particularly for species at risk of exploitation. While we recognize that well-informed control of location data for highly sensitive taxa is necessary to avoid risks, such as poaching or habitat disturbance by recreational visitors, we argue that ignoring the benefits of sharing biodiversity data could unnecessarily obstruct conservation efforts for species and locations with low risks of exploitation. We provide a decision tree protocol for scientists that systematically considers both the risks of exploitation and potential benefits of increased conservation activities. Our protocol helps scientists assess the impacts of publishing biodiversity data and aims to enhance conservation opportunities, promote community engagement and reduce duplication of survey efforts.

Biodiversity responds to increasing climatic extremes in a biome-specific manner.

An unprecedented rate of global environmental change is predicted for the next century. The response to this change by ecosystems around the world is highly uncertain. To address this uncertainty, it is critical to understand the potential drivers and mechanisms of change in order to develop more reliable predictions. Australia's Long Term Ecological Research Network (LTERN) has brought together some of the longest running (10-60years) continuous environmental monitoring programs in the southern hemisphere. Here, we compare climatic variables recorded at five LTERN plot network sites during their period of operation and place them into the context of long-term climatic trends. Then, using our unique Australian long-term datasets (total 117 survey years across four biomes), we synthesize results from a series of case studies to test two hypotheses: 1) extreme weather events for each plot network have increased over the last decade, and; 2) trends in biodiversity will be associated with recent climate change, either directly or indirectly through climate-mediated disturbance (wildfire) responses. We examined the biodiversity responses to environmental change for evidence of non-linear behavior. In line with hypothesis 1), an increase in extreme climate events occurred within the last decade for each plot network. For hypothesis 2), climate, wildfire, or both were correlated with biodiversity responses at each plot network, but there was no evidence of non-linear change. However, the influence of climate or fire was context-specific. Biodiversity responded to recent climate change either directly or indirectly as a consequence of changes in fire regimes or climate-mediated fire responses. A national long-term monitoring framework allowed us to find contrasting species abundance or community responses to climate and disturbance across four of the major biomes of Australia, highlighting the need to establish and resource long-term monitoring programs across representative ecosystem types, which are likely to show context-specific responses.