Understanding decision making in a food-caching predator using hidden Markov models

Financial support was provided by the People’s Trust for Endangered Species (PTES), Zoologische Gesellschaft für Arten- und Populationsschutz (ZGAP), Quagga Conservation Fund and IdeaWild.Background Tackling behavioural questions often requires identifying points in space and time where animals make decisions and linking these to environmental variables. State-space modeling is useful for analysing movement trajectories, particularly with hidden Markov models (HMM). Yet importantly, the ontogeny of underlying (unobservable) behavioural states revealed by the HMMs has rarely been verified in the field. Methods Using hidden Markov models of individual movement from animal location, biotelemetry, and environmental data, we explored multistate behaviour and the effect of associated intrinsic and extrinsic drivers across life stages. We also decomposed the activity budgets of different movement states at two general and caching phases. The latter - defined as the period following a kill which likely involves the caching of uneaten prey - was subsequently confirmed by field inspections. We applied this method to GPS relocation data of a caching predator, Persian leopard Panthera pardus saxicolor in northeastern Iran. Results Multistate modeling provided strong evidence for an effect of life stage on the behavioural states and their associated time budget. Although environmental covariates (ambient temperature and diel period) and ecological outcomes (predation) affected behavioural states in non-resident leopards, the response in resident leopards was not clear, except that temporal patterns were consistent with a crepuscular and nocturnal movement pattern. Resident leopards adopt an energetically more costly mobile behaviour for most of their time while non-residents shift their behavioural states from high energetic expenditure states to energetically less costly encamped behaviour for most of their time, which is likely to be a risk avoidance strategy against conspecifics or humans. Conclusions This study demonstrates that plasticity in predator behaviour depending on life stage may tackle a trade-off between successful predation and avoiding the risks associated with conspecifics, human presence and maintaining home range. Range residency in territorial predators is energetically demanding and can outweigh the predator’s response to intrinsic and extrinsic variables such as thermoregulation or foraging needs. Our approach provides an insight into spatial behavior and decision making of leopards, and other large felids in rugged landscapes through the application of the HMMs in movement ecology.Publisher PDFPeer reviewe

A review of the ecological and socioeconomic characteristics of trophy hunting across Asia

The continuing debates about trophy hunting should be underpinned by an understanding of at least the basic characteristics of the practice (e.g. species, quotas, areas, prices). Whilst many countries in Asia have established trophy hunting programmes of considerable importance to conservation and local livelihoods, there remains some ambiguity over the extent of trophy hunting in Asia as its basic characteristics in each country have not been compiled. In this study, we compile information on various ecological and socioeconomic characteristics of trophy hunting of mammals for countries across Asia by reviewing published and unpublished literature, analysing trade data, and obtaining contributions from in-country contacts. Across Asia, established trophy hunting programmes exist in at least 11 countries and target at least 30 species and one hybrid (incl., five Vulnerable and one Endangered species). Trophy hunting in these countries varies markedly in areas (e.g. >1 million km2 in Kazakhstan, 37% of country, vs. 1325 km2 in Nepal, <1% of country) and annual offtakes (e.g. Kazakhstan: 4500 individuals from 4 of 5 trophy species; Pakistan: 229 from 4 of 7; Mongolia: 155 from 6 of 9; Tajikistan: 126 from 3 of 6; Nepal: 22 from 3 of the 4 that are trophy hunted in practice). Permit prices also vary across species and countries, with domestic and international hunters sometimes charged different rates. Hunters from the USA appear overwhelmingly prominent among international clients. National legislations typically mandate a proportion of trophy hunting revenue to accrue locally (range: 40–100%). We provide five key recommendations for research to inform trophy hunting policy in Asia: (1) Ecological impact assessments; (2) Socioeconomic impact assessments; (3) Evaluations of the contributions of trophy hunting to conservation spending; (4) Evaluations of the contributions of trophy hunting to the post-2020 Global Biodiversity Framework; (5) Further examinations of perceptions of trophy hunting

Optimal feedback control for dynamic systems with state constraints: An exact penalty approach

In this paper, we consider a class of nonlinear dynamic systems with terminal state and continuous inequality constraints. Our aim is to design an optimal feedback controller that minimizes total system cost and ensures satisfaction of all constraints. We first formulate this problem as a semi-infinite optimization problem. We then show that by using a new exact penalty approach, this semi-infinite optimization problem can be converted into a sequence of nonlinear programming problems, each of which can be solved using standard gradient-based optimization methods.We conclude the paper by discussing applications of our work to glider control

Coat Polymorphism in Eurasian Lynx: Adaptation to Environment or Phylogeographic Legacy?

We studied the relationship between the variability and contemporary distribution of pelage phenotypes in one of most widely distributed felid species and an array of environmental and demographic conditions. We collected 672 photographic georeferenced records of the Eurasian lynx throughout Eurasia. We assigned each lynx coat to one of five phenotypes. Then we fitted the coat patterns to different environmental and anthropogenic variables, as well as the effective geographic distances from inferred glacial refugia. A majority of lynx were either of the large spotted (41.5%) or unspotted (uniform, 36.2%) phenotype. The remaining patterns (rosettes, small spots and pseudo-rosettes) were represented in 11.0%, 7.4%, and 3.9% of samples, respectively. Although various environmental variables greatly affected lynx distribution and habitat suitability, it was the effect of least-cost distances from locations of the inferred refugia during the Last Glacial Maximum that explained the distribution of lynx coat patterns the best. Whereas the occurrence of lynx phenotypes with large spots was explained by the proximity to refugia located in the Caucasus/Middle East, the uniform phenotype was associated with refugia in the Far East and Central Asia. Despite the widely accepted hypothesis of adaptive functionality of coat patterns in mammals and exceptionally high phenotypic polymorphism in Eurasian lynx, we did not find well-defined signs of habitat matching in the coat pattern of this species. Instead, we showed how the global patterns of morphological variability in this large mammal and its environmental adaptations may have been shaped by past climatic change.publishedVersio

The control parameterization method for nonlinear optimal control: A survey

The control parameterization method is a popular numerical technique for solving optimal control problems. The main idea of control parameterization is to discretize the control space by approximating the control function by a linear combination of basis functions. Under this approximation scheme, the optimal control problem is reduced to an approximate nonlinear optimization problem with a finite number of decision variables. This approximate problem can then be solved using nonlinear programming techniques. The aim of this paper is to introduce the fundamentals of the control parameterization method and survey its various applications to non-standard optimal control problems. Topics discussed include gradient computation, numerical convergence, variable switching times, and methods for handling state constraints. We conclude the paper with some suggestions for future research

Effects of body size on estimation of mammalian area requirements.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied blockcross validation to quantify bias in empirical home range estimates. Area requirements of mammals 1, meaning the scaling of the relationship changedsubstantially at the upper end of the mass spectrum

Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations

Background: Resolving the historical biogeography of the leopard (Panthera pardus) is a complex issue, because patterns inferred from fossils and from molecular data lack congruence. Fossil evidence supports an African origin, and suggests that leopards were already present in Eurasia during the Early Pleistocene. Analysis of DNA sequences however, suggests a more recent, Middle Pleistocene shared ancestry of Asian and African leopards. These contrasting patterns led researchers to propose a two-stage hypothesis of leopard dispersal out of Africa: an initial Early Pleistocene colonisation of Asia and a subsequent replacement by a second colonisation wave during the Middle Pleistocene. The status of Late Pleistocene European leopards within this scenario is unclear: were these populations remnants of the first dispersal, or do the last surviving European leopards share more recent ancestry with their African counterparts? Results: In this study, we generate and analyse mitogenome sequences from historical samples that span the entire modern leopard distribution, as well as from Late Pleistocene remains. We find a deep bifurcation between African and Eurasian mitochondrial lineages (~ 710 Ka), with the European ancient samples as sister to all Asian lineages (~ 483 Ka). The modern and historical mainland Asian lineages share a relatively recent common ancestor (~ 122 Ka), and we find one Javan sample nested within these. Conclusions: The phylogenetic placement of the ancient European leopard as sister group to Asian leopards suggests that these populations originate from the same out-of-Africa dispersal which founded the Asian lineages. The coalescence time found for the mitochondrial lineages aligns well with the earliest undisputed fossils in Eurasia, and thus encourages a re-evaluation of the identification of the much older putative leopard fossils from the region. The relatively recent ancestry of all mainland Asian leopard lineages suggests that these populations underwent a severe population bottleneck during the Pleistocene. Finally, although only based on a single sample, the unexpected phylogenetic placement of the Javan leopard could be interpreted as evidence for exchange of mitochondrial lineages between Java and mainland Asia, calling for further investigation into the evolutionary history of this subspecies

Effects of body size on estimation of mammalian area requirements

Accurately quantifying species’ area requirements is a prerequisite for effective area‐based conservation. This typically involves collecting tracking data on species of interest and then conducting home‐range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home‐range areas with GPS locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4,000 kg. We then applied block cross‐validation to quantify bias in empirical home‐range estimates. Area requirements of mammals 1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum