Are predictions of bovine tuberculosis-infected herds unbiased and precise?

Bovine tuberculosis (bTB) is prevalent among livestock and wildlife in many countries including New Zealand (NZ), a country which aims to eradicate bTB by 2055. This study evaluates predictions related to the numbers of livestock herds with bTB in NZ from 2012 to 2021 inclusive using both statistical and mechanistic (causal) modelling. Additionally, this study made predictions for the numbers of infected herds between 2022 and 2059. This study introduces a new graphical method representing the causal criteria of strength of association, such as R2, and the consistency of predictions, such as mean squared error. Mechanistic modelling predictions were, on average, more frequently (3 of 4) unbiased than statistical modelling predictions (1 of 4). Additionally, power model predictions were, on average, more frequently (3 of 4) unbiased than exponential model predictions (1 of 4). The mechanistic power model, along with annual updating, had the highest R2 and the lowest mean squared error of predictions. It also exhibited the closest approximation to unbiased predictions. Notably, significantly biased predictions were all underestimates. Based on the mechanistic power model, the biological eradication of bTB from New Zealand is predicted to occur after 2055. Disease eradication planning will benefit from annual updating of future predictions.

Unrecognized threat to global soil carbon by a widespread invasive species.

Most of Earth's terrestrial carbon is stored in the soil and can be released as carbon dioxide (CO2 ) when disturbed. Although humans are known to exacerbate soil CO2 emissions through land-use change, we know little about the global carbon footprint of invasive species. We predict the soil area disturbed and resulting CO2 emissions from wild pigs (Sus scrofa), a pervasive human-spread vertebrate that uproots soil. We do this using models of wild pig population density, soil damage, and their effect on soil carbon emissions. Our models suggest that wild pigs are uprooting a median area of 36,214 km2 (mean of 123,517 km2 ) in their non-native range, with a 95% prediction interval (PI) of 14,208 km2 -634,238 km2 . This soil disturbance results in median emissions of 4.9 million metric tonnes (MMT) CO2 per year (equivalent to 1.1 million passenger vehicles or 0.4% of annual emissions from land use, land-use change, and forestry; mean of 16.7 MMT) but that it is highly uncertain (95% PI, 0.3-94 MMT CO2 ) due to variability in wild pig density and soil dynamics. This uncertainty points to an urgent need for more research on the contribution of wild pigs to soil damage, not only for the reduction of anthropogenically related carbon emissions, but also for co-benefits to biodiversity and food security that are crucial for sustainable development.

Invasive wild pigs (Sus scrofa) as a human-mediated source of soil carbon emissions: Uncertainties and future directions.

Invasive wild pigs (Sus scrofa) have been spread by humans outside of their native range and are now established on every continent except Antarctica. Through their uprooting of soil, they affect societal and environmental values. Our recent article explored another threat from their soil disturbance: greenhouse gas emissions (O'Bryan et al., Global Change Biology, 2021). In response to our paper, Don (Global Change Biology, 2021) claims there is no threat to global soil carbon stocks by wild pigs. While we did not investigate soil carbon stocks, we examine uncertainties regarding soil carbon emissions from wild pig uprooting and their implications for management and future research.

Eruptive dynamics are common in managed mammal populations.

Successful conservation management is often based on the principle that small or declining populations can recover if we identify and remove the factors that caused them to decline in the first place. But what form will that recovery take? Theory tells us that when a strong limiting factor is removed, a population should increase in size to where it becomes limited by some other factor. However, if the subsequent limitation involves feedbacks between the density of a consumer and its resource, there is potential for the consumer population to undergo substantial fluctuations in size that we would characterize as boom-bust or eruptive dynamics. We analysed long-term (7.6-29 yr) data documenting changes in the abundance of 169 populations of 20 mammal species released from a strong limiting factor (fox predation) in Australia. We show that many populations (44) exhibited eruptive dynamics, with exponential increase to a peak and subsequent population decline. Of 51 populations showing eruptive dynamics (the Australian populations plus seven translocated ungulate populations), the time taken for erupting populations to reach a peak before declining was related negatively to the intrinsic rate of population growth and positively to body mass, such that larger-bodied species with slow rates of population growth had a longer period of population increase before declining. Our results suggest that a substantial proportion of populations recovering after removal of a threatening process are likely to exhibit eruptive dynamics, and that managers of recovering or translocated populations should anticipate this outcome in conservation planning.

Under what conditions do climate-driven sex ratios enhance versus diminish population persistence?

For many species of reptile, crucial demographic parameters such as embryonic survival and individual sex (male or female) depend on ambient temperature during incubation. While much has been made of the role of climate on offspring sex ratios in species with temperature-dependent sex determination (TSD), the impact of variable sex ratio on populations is likely to depend on how limiting male numbers are to female fecundity in female-biased populations, and whether a climatic effect on embryonic survival overwhelms or interacts with sex ratio. To examine the sensitivity of populations to these interacting factors, we developed a generalized model to explore the effects of embryonic survival, hatchling sex ratio, and the interaction between these, on population size and persistence while varying the levels of male limitation. Populations with TSD reached a greater maximum number of females compared to populations with GSD, although this was often associated with a narrower range of persistence. When survival depended on temperature, TSD populations persisted over a greater range of temperatures than GSD populations. This benefit of TSD was greatly reduced by even modest male limitation, indicating very strong importance of this largely unmeasured biologic factor. Finally, when males were not limiting, a steep relationship between sex ratio and temperature favoured population persistence across a wider range of climates compared to the shallower relationships. The opposite was true when males were limiting - shallow relationships between sex ratio and temperature allowed greater persistence. The results highlight that, if we are to predict the response of populations with TSD to climate change, it is imperative to 1) accurately quantify the extent to which male abundance limits female fecundity, and 2) measure how sex ratios and peak survival coincide over climate.

How do climate-linked sex ratios and dispersal limit range boundaries?

BACKGROUND: Geographic ranges of ectotherms such as reptiles may be determined strongly by abiotic factors owing to causal links between ambient temperature, juvenile survival and individual sex (male or female). Unfortunately, we know little of how these factors interact with dispersal among populations across a species range. We used a simulation model to examine the effects of dispersal, temperature-dependent juvenile survival and sex determining mechanism (temperature-dependent sex determination (TSD) and genotypic sex determination (GSD)) and their interactions, on range limits in populations extending across a continuous range of air temperatures. In particular, we examined the relative importance of these parameters for population persistence to recommend targets for future empirical research. RESULTS: Dispersal influenced the range limits of species with TSD to a greater extent than in GSD species. Whereas male dispersal led to expanded species ranges across warm (female-producing) climates, female dispersal led to expanded ranges across cool (male-producing) climates. Two-sex dispersal eliminated the influence of biased sex ratios on ranges. CONCLUSION: The results highlight the importance of the demographic parameter of sex ratio in determining population persistence and species range limits.

Climatic amplification of the numerical response of a predator population to its prey.

We evaluated evidence of an effect of climate on the numerical response of a coyote (Canis latrans) population to their keystone prey, snowshoe hares (Lepus americanus), in a Canadian boreal forest. Six a priori hypotheses of the coyote numerical response were developed that postulated linear, nonlinear, additive, and interactive effects of prey and climate. Model selection procedures showed the North Atlantic Oscillation (NAO) had the strongest effect on the coyote numerical response via its interaction with snowshoe hare density, while other large-scale climate indices had very weak effects. For a given snowshoe hare density, a negative value of the NAO amplified the abundance of coyote and a positive NAO decreased coyote abundance. We hypothesize that the coyote numerical response is ultimately determined by the coyote functional response influenced by winter conditions determined by the NAO.

On the stability of populations of mammals, birds, fish and insects.

A key concern for conservation biologists is whether populations of plants and animals are likely to fluctuate widely in number or remain relatively stable around some steady-state value. In our study of 634 populations of mammals, birds, fish and insects, we find that most can be expected to remain stable despite year to year fluctuations caused by environmental factors. Mean return rates were generally around one but were higher in insects (1.09 +/- 0.02 SE) and declined with body size in mammals. In general, this is good news for conservation, as stable populations are less likely to go extinct. However, the lower return rates of the large mammals may make them more vulnerable to extinction. Our estimates of return rates were generally well below the threshold for chaos, which makes it unlikely that chaotic dynamics occur in natural populations--one of ecology's key unanswered questions.

Testing the metabolic theory of ecology: allometric scaling exponents in mammals.

Many fundamental traits of species measured at different levels of biological organization appear to scale as a power law to body mass (M) with exponents that are multiples of 1/4. Recent work has united these relationships in a "metabolic theory of ecology" (MTE) that explains the pervasiveness of quarter-power scaling by its dependence on basal metabolic rate (B), which scales as M(0.75). Central to the MTE is theory linking the observed -0.25 scaling of maximum population growth rate (rm) and body mass to the 0.75 scaling of metabolic rate and body mass via relationships with age at first reproduction (alpha) derived from a general growth model and demographic theory. We used this theory to derive two further predictions: that age at first reproduction should scale inversely to mass-corrected basal metabolic rate alpha infinity (B/M)(-l) such that rm infinity (B/M)1. We then used phylogenetic generalized least squares and model selection methods to test the predicted scaling relationships using data from 1197 mammalian species. There was a strong phylogenetic signal in these data, highlighting the need to account for phylogeny in allometric studies. The 95% confidence intervals included, or almost included, the scaling exponent predicted by MTE for B infinity M(0.75), rm infinity M(-0.25), and rm infinity alpha(-1), but not for alpha infinity M(0.25) or the two predictions that we generated. Our results highlight a mismatch between theory and observation and imply that the observed -0.25 scaling of maximum population growth rate and body mass does not arise via the mechanism proposed in the MTE.

Climate, food, density and wildlife population growth rate.

1. The aim of this study was to derive and evaluate a priori models of the relationship between annual instantaneous population growth rate (r) and climate. These were derived from the numerical response of annual r and food, and the effect of climate on a parameter in the numerical response. The goodness of fit of a range of such deductive models to data on annual r of Soay sheep and red deer were evaluated using information-theoretic (AICc-based) analyses. 2. The analysis for sheep annual r showed negative effects of abundance and negative effects of the interaction of abundance and climate, measured as March rainfall (and winter NAO) in the best fitting models. The analysis for deer annual r showed a negative effect of deer abundance and a positive effect of climate measured as March rainfall (but a negative effect of winter NAO), but no interaction of abundance and climate in the best fitting models. 3. There was most support in the analysis of sheep dynamics for the ratio numerical response and the assumption that parameter J (equilibrium food per animal) was influenced by climate. In the analysis of deer dynamics there was most support for the numerical responses assuming effects of food and density (Ivlev and density, food and density, and additive responses) and slightly less support for the ratio numerical response. The evaluation of such models would be aided by the collection of and incorporation of food data into the analyses.

On the regulation of populations of mammals, birds, fish, and insects.

A key unresolved question in population ecology concerns the relationship between a population's size and its growth rate. We estimated this relationship for 1780 time series of mammals, birds, fish, and insects. We found that rates of population growth are high at low population densities but, contrary to previous predictions, decline rapidly with increasing population size and then flatten out, for all four taxa. This produces a strongly concave relationship between a population's growth rate and its size. These findings have fundamental implications for our understanding of animals' lives, suggesting in particular that many animals in these taxa will be found living at densities above the carrying capacity of their environments.

Population growth rate and its determinants: an overview.

We argue that population growth rate is the key unifying variable linking the various facets of population ecology. The importance of population growth rate lies partly in its central role in forecasting future population trends; indeed if the form of density dependence were constant and known, then the future population dynamics could to some degree be predicted. We argue that population growth rate is also central to our understanding of environmental stress: environmental stressors should be defined as factors which when first applied to a population reduce population growth rate. The joint action of such stressors determines an organism's ecological niche, which should be defined as the set of environmental conditions where population growth rate is greater than zero (where population growth rate = r = log(e)(N(t+1)/N(t))). While environmental stressors have negative effects on population growth rate, the same is true of population density, the case of negative linear effects corresponding to the well-known logistic equation. Following Sinclair, we recognize population regulation as occurring when population growth rate is negatively density dependent. Surprisingly, given its fundamental importance in population ecology, only 25 studies were discovered in the literature in which population growth rate has been plotted against population density. In 12 of these the effects of density were linear; in all but two of the remainder the relationship was concave viewed from above. Alternative approaches to establishing the determinants of population growth rate are reviewed, paying special attention to the demographic and mechanistic approaches. The effects of population density on population growth rate may act through their effects on food availability and associated effects on somatic growth, fecundity and survival, according to a 'numerical response', the evidence for which is briefly reviewed. Alternatively, there may be effects on population growth rate of population density in addition to those that arise through the partitioning of food between competitors; this is 'interference competition'. The distinction is illustrated using a replicated laboratory experiment on a marine copepod, Tisbe battagliae. Application of these approaches in conservation biology, ecotoxicology and human demography is briefly considered. We conclude that population regulation, density dependence, resource and interference competition, the effects of environmental stress and the form of the ecological niche, are all best defined and analysed in terms of population growth rate.

Demographic, mechanistic and density-dependent determinants of population growth rate: a case study in an avian predator.

Identifying the determinants of population growth rate is a central topic in population ecology. Three approaches (demographic, mechanistic and density-dependent) used historically to describe the determinants of population growth rate are here compared and combined for an avian predator, the barn owl (Tyto alba). The owl population remained approximately stable (r approximately 0) throughout the period from 1979 to 1991. There was no evidence of density dependence as assessed by goodness of fit to logistic population growth. The finite (lambda) and instantaneous (r) population growth rates were significantly positively related to food (field vole) availability. The demographic rates, annual adult mortality, juvenile mortality and annual fecundity were reported to be correlated with vole abundance. The best fit (R(2) = 0.82) numerical response of the owl population described a positive effect of food (field voles) and a negative additive effect of owl abundance on r. The numerical response of the barn owl population to food availability was estimated from both census and demographic data, with very similar results. Our analysis shows how the demographic and mechanistic determinants of population growth rate are linked; food availability determines demographic rates, and demographic rates determine population growth rate. The effects of food availability on population growth rate are modified by predator abundance.