E. Penelope Holland

Of mast and mean: differential-temperature cue makes mast seeding insensitive to climate change.

Mast-seeding plants often produce high seed crops the year after a warm spring or summer, but the warm-temperature model has inconsistent predictive ability. Here, we show for 26 long-term data sets from five plant families that the temperature difference between the two previous summers (ΔT) better predicts seed crops. This discovery explains how masting species tailor their flowering patterns to sites across altitudinal temperature gradients; predicts that masting will be unaffected by increasing mean temperatures under climate change; improves prediction of impacts on seed consumers; demonstrates that strongly masting species are hypersensitive to climate; explains the rarity of consecutive high-seed years without invoking resource constraints; and generates hypotheses about physiological mechanisms in plants and insect seed predators. For plants, ΔT has many attributes of an ideal cue. This temperature-difference model clarifies our understanding of mast seeding under environmental change, and could also be applied to other cues, such as rainfall.

Least-cost path length versus accumulated-cost as connectivity measures

Least-cost modelling has become a popular method for measuring connectivity. By representing the landscape as a cost-surface, least-cost paths can be calculated that represent the route of maximum efficiency between two locations as a function of the distance travelled and the costs traversed. Both the length and the accumulated-cost of a least-cost path have been used as measures of connectivity between pairs of locations. However, we are concerned that in some situations the length of a least-cost path may provide a misleading measure of connectivity as it only accounts for the distance travelled while ignoring the costs traversed, and results in a measure that may be little better than Euclidean distance. Through simulations using fractal landscapes we demonstrate that least-cost path length is often highly correlated with Euclidean distance. This indicates that least-cost path length provides a poor measure of connectivity in many situations, as it does not capture sufficient information about the ecological costs to movement represented by the cost-surface. We recommend that in most situations the accumulated-cost of a least-cost path provides a more appropriate measure of connectivity between locations as it accounts for both the distance travelled and costs traversed, and that the generation of vector least-cost paths should be reserved for visualisation purposes.

NLMpy: a python software package for the creation of neutral landscape models within a general numerical framework

Summary Neutral landscape models (NLMs) are widely used to model ecological patterns and processes across landscapes. However, the ability to generate NLMs is often made available through standalone bespoke software packages that have platform limitations. We have developed a python package that brings together some of the more popular NLM algorithms using a general numerical framework. The resulting NLMpy package: (i) allows for the creation of NLMs directly within a python modelling workflow or by other modelling software capable executing a python script, (ii) enables the first opportunity to create a NLM that combines different algorithms, (iii) provides easy integration with geographic information system data and (iv) creates a framework for developing other NLMs.

Thresholds in plant–herbivore interactions: predicting plant mortality due to herbivore browse damage

Patterns of herbivore browse at small scales, such as the rate of leaf consumption or plant preferences, drive the impact of herbivores on whole-plant processes, such as growth or survival, and subsequent changes in plant population structure. However, herbivore impacts are often non-linear, highly variable and context-dependent. Understanding the effect of herbivores on plant populations therefore requires a detailed understanding of the relationships that drive small-scale processes, and how these interact to generate dynamics at larger scales. We derive a mathematical model to predict annual rates of browse-induced tree mortality. We model individual plant mortality as a result of rates of foliage production, turnover and herbivore intake, and extend the model to the population scale by allowing for between-tree variation in levels of herbivore browse. The model is configurable for any broadleaved tree species subject to vertebrate or invertebrate browse, and is designed to be parameterized from field data typically collected as part of browse damage assessments. We parameterized and tested the model using data on foliage cover and browse damage recorded on kamahi trees (Weinmannia racemosa) browsed by possums (Trichosurus vulpecula) in New Zealand forests. The model replicated observed patterns of tree mortality at 12 independent validation sites with a wide range of herbivore densities and browse damage. The model reveals two key thresholds; in plant foliar cover, indicating when individual trees may be at high risk from browse-induced mortality, and in herbivore intake, leading to high rates of mortality across the whole population.

On the growth of locally interacting plants: differential equations for the dynamics of spatial moments

Ecologists are faced with the challenge of how to scale up from the activities of individual plants and animals to the macroscopic dynamics of populations and communities. It is especially difficult to do this in communities of plants where the fate of individuals depends on their immediate neighbors rather than an average over a larger region. This has meant that algorithmic, agent-based models are typically used to understand their dynamics, although certain macroscopic models have been developed for neighbor-dependent, birth death processes. Here we present a macroscopic model that, for the first time, incorporates explicit, gradual, neighbor-dependent plant growth, as a third fundamental process of plant communities. The model is derived from a stochastic, agent-based model, and describes the dynamics of the first and second spatial moments of a multispecies, spatially structured plant community with neighbor-dependent growth, births, and deaths. A simple example shows that strong neighborhood space-filling during tree growth in an even-aged stand of Scots pine is well captured by the spatial-moment model. The space-filling has a spatial signature consistent with that observed in several field studies of forests. Small neighborhoods of interaction, nonuniform spacing of trees, and asymmetric competition all contribute to the buildup of a wide range of tree sizes with some large dominant individuals and many smaller ones.

Assessing the efficacy of population-level models of mast seeding

We compare five models of mast seeding: the well-established resource-based model of Isagi et al. (J Theor Biol 187:231–239, 1997) and Satake and Iwasa (J Theor Biol 203:63–84, 2000), the regression based model of Masaka and Maguchi (Ann Bot 88:1049–1055, 2001), the more recent Delta temperature (ΔT) model of Kelly et al. (Ecol Lett 16:90–98, 2013), the resource model of Pearse et al. (Oikos 123:179–184, 2014) and the long-used temperature model. We discuss fitting methods for each model and compare the theoretical overlap between the models. Population level data from 28 seed sets and corresponding mean summer temperature are used and the error of each model is compared. On average, the ΔT model provides a better fit than the other models tested.

Climate-based models for pulsed resources improve predictability of consumer population dynamics: outbreaks of house mice in forest ecosystems.

Accurate predictions of the timing and magnitude of consumer responses to episodic seeding events (masts) are important for understanding ecosystem dynamics and for managing outbreaks of invasive species generated by masts. While models relating consumer populations to resource fluctuations have been developed successfully for a range of natural and modified ecosystems, a critical gap that needs addressing is better prediction of resource pulses. A recent model used change in summer temperature from one year to the next (ΔT) for predicting masts for forest and grassland plants in New Zealand. We extend this climate-based method in the framework of a model for consumer–resource dynamics to predict invasive house mouse (Mus musculus) outbreaks in forest ecosystems. Compared with previous mast models based on absolute temperature, the ΔT method for predicting masts resulted in an improved model for mouse population dynamics. There was also a threshold effect of ΔT on the likelihood of an outbreak occurring. The improved climate-based method for predicting resource pulses and consumer responses provides a straightforward rule of thumb for determining, with one year’s advance warning, whether management intervention might be required in invaded ecosystems. The approach could be applied to consumer–resource systems worldwide where climatic variables are used to model the size and duration of resource pulses, and may have particular relevance for ecosystems where global change scenarios predict increased variability in climatic events.

A modelling framework for predicting the optimal balance between control and surveillance effort in the local eradication of tuberculosis in New Zealand wildlife.

Bovine tuberculosis (TB) impacts livestock farming in New Zealand, where the introduced marsupial brushtail possum (Trichosurus vulpecula) is the wildlife maintenance host for Mycobacterium bovis. New Zealand has implemented a campaign to control TB using a co-ordinated programme of livestock diagnostic testing and large-scale culling of possums, with the long-term aim of TB eradication. For management of the disease in wildlife, methods that can optimise the balance between control and surveillance effort will facilitate the objective of eradication on a fixed or limited budget. We modelled and compared management options to optimise the balance between the two activities necessary to achieve and verify eradication of TB from New Zealand wildlife: the number of lethal population control operations required to halt the M. bovis infection cycle in possums, and the subsequent surveillance effort needed to confidently declare TB freedom post-control. The approach considered the costs of control and surveillance, as well as the potential costs of re-control resulting from false declaration of TB freedom. The required years of surveillance decreased with increasing numbers of possum lethal control operations but the overall time to declare TB freedom depended on additional factors, such as the probability of freedom from disease after control and the probability of success of mop-up control, i.e. retroactive culling following detection of persistent disease in the residual possum population. The total expected cost was also dependent on a number of factors, many of which had wide cost ranges, suggesting that an optimal strategy is unlikely to be singular and fixed, but will likely vary for each different area being considered. Our approach provides a simple framework that considers the known and potential costs of possum control and TB surveillance, enabling managers to optimise the balance between these two activities to achieve and prove eradication of a wildlife disease, or the pest species that transmits it, in the most expedient and economic way. This cost- and risk-evaluation approach may be applicable to other wildlife disease problems where limited management funds exist.

Foliar Nutritional Quality Explains Patchy Browsing Damage Caused by an Invasive Mammal.

Introduced herbivores frequently inflict significant, yet patchy damage on native ecosystems through selective browsing. However, there are few instances where the underlying cause of this patchy damage has been revealed. We aimed to determine if the nutritional quality of foliage could predict the browsing preferences of an invasive mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), in a temperate forest in New Zealand. We quantified the spatial and temporal variation in four key aspects of the foliar chemistry (total nitrogen, available nitrogen, in vitro dry matter digestibility and tannin effect) of 275 trees representing five native tree species. Simultaneously, we assessed the severity of browsing damage caused by possums on those trees in order to relate selective browsing to foliar nutritional quality. We found significant spatial and temporal variation in nutritional quality among individuals of each tree species examined, as well as among tree species. There was a positive relationship between the available nitrogen concentration of foliage (a measure of in vitro digestible protein) and the severity of damage caused by browsing by possums. This study highlights the importance of nutritional quality, specifically, the foliar available nitrogen concentration of individual trees, in predicting the impact of an invasive mammal. Revealing the underlying cause of patchy browsing by an invasive mammal provides new insights for conservation of native forests and targeted control of invasive herbivores in forest ecosystems.

How Herbivore Browsing Strategy Affects Whole-Plant Photosynthetic Capacity

If a browse damage index indicates that a tree has been 50% browsed by herbivores, does this mean half the leaves are entirely eaten or are all the leaves half eaten? Were the affected leaves old or young? Large or small? In sunshine or shade? Understanding what effect browsing will have on the photosynthetic capacity and the plant’s survival ability clearly requires a greater understanding of browsing strategy across the canopy than can be given by a single index value. We developed stochastic models of leaf production, growth and consumption using data from kamahi (Weinmannia racemosa) trees in New Zealand which have been browsed by possums (Trichosurus vulpecula), to ascertain which of six feasible browsing strategies possums are most likely to be employing. We compared the area distribution of real fallen leaves to model output in order to select the best model, and used the model to predict the age distribution of leaves on the tree and thus infer its photosynthetic capability. The most likely browsing strategy that possums employ on kamahi trees is a preference for virgin (i.e. previously unbrowsed) leaves, consistent with the idea that browsing increases the production of chemical plant defences. More generally, our results show that herbivore browsing strategy can significantly change the whole-plant photosynthetic capability of any plant and hence its ability to survive, and therefore, herbivore damage indices should be used in conjunction with more detailed information about herbivore browsing strategy.