Avoiding misleading messages: Population assessment using camera trapping is not a simple task.

Population assessment is indispensable for appropriate and socially acceptable conservation and management of wildlife populations. This article critiques the paper by Campos-Candela et al. 2018 and highlights issues that could lead to inappropriate management and conservation policies.

Kleptoparasitism and Scavenging Can Stabilize Ecosystem Dynamics.

Scavenging is ubiquitous in nature, but its implications have rarely been investigated. We used camera traps on wolf kills to investigate the role of scavenging on predator and multiprey dynamics in a northern Apennine system in Italy. In contrast to North American systems, the omnivorous wild boar successfully competes with wolves for the meat of their kills. We developed a deterministic, multitrophic web model (wolf, vegetation, deer, and wild boar), tunable through a parameter that governs the impact of prey sharing between wolves and wild boar. When prey sharing is scarce, populations oscillate, but above a threshold value the trophic web is stabilized, with the regime solution becoming a fixed, stable point. Both deer and wild boar then increase as a function of prey sharing, and the impact of herbivores on the vegetation increases. When prey sharing exceeds another threshold, the system collapses due to the extinction of both wolves and wild boar. Our analysis shows that scavenging is crucial for the dynamics of this ecosystem, and thus it should not be overlooked in food web modeling. The exploitation of wolf kills by wild boar may allow juveniles and yearlings to obtain high-quality resources that are not usually available, helping the wild boar to compensate for losses caused by hunting. This is likely to make them even more invasive and difficult to control.

Reproductive allocation in pulsed-resource environments: a comparative study in two populations of wild boar.

Pulsed resources influence the demography and evolution of consumer populations and, by cascading effect, the dynamics of the entire community. Mast seeding provides a case study for exploring the evolution of life history traits of consumers in fluctuating environments. Wild boar (Sus scrofa) population dynamics is related to seed availability (acorns/beechnuts). From a long-term monitoring of two populations subjected to markedly different environmental contexts (i.e., both low vs. high frequency of pulsed resources and low vs. high hunting pressure in Italy and in France, respectively), we assessed how pulsed resources shape the reproductive output of females. Using path analyses, we showed that in both populations, abundant seed availability increases body mass and both the absolute and the relative (to body mass) allocation to reproduction through higher fertility. In the Italian population, females equally relied on past and current resources for reproduction and ranked at an intermediate position along the capital-income continuum of breeding tactics. In contrast, in the French population, females relied on current more than past resources and ranked closer to the income end of the continuum. In the French population, one-year old females born in acorn-mast years were heavier and had larger litter size than females born in beechnut-mast years. In addition to the quantity, the type of resources (acorns/beechnuts) has to be accounted for to assess reliably how females allocate resources to reproduction. Our findings highlight a high plasticity in breeding tactics in wild boar females and provide new insight on allocation strategies in fluctuating environments.

Lévy patterns in seabirds are multifaceted describing both spatial and temporal patterning.

BACKGROUND: The flight patterns of albatrosses and shearwaters have become a touchstone for much of Lévy flight research, spawning an extensive field of enquiry. There is now compelling evidence that the flight patterns of these seabirds would have been appreciated by Paul Lévy, the mathematician after whom Lévy flights are named. Here we show that Lévy patterns (here taken to mean spatial or temporal patterns characterized by distributions with power-law tails) are, in fact, multifaceted in shearwaters being evident in both spatial and temporal patterns of activity. RESULTS: We tested for Lévy patterns in the at-sea behaviours of two species of shearwater breeding in the North Atlantic Ocean (Calonectris borealis) and the Mediterranean sea (C. diomedea) during their incubating and chick-provisioning periods. We found that distributions of flight durations, on/in water durations and inter-dive time-intervals have power-law tails and so bear the hallmarks of Lévy patterns. CONCLUSIONS: The occurrence of these statistical laws is remarkable given that bird behaviours are strongly shaped by an individual's motivational state and by complex environmental interactions. Our observations could take Lévy patterns as models of animal behaviour to a new level by going beyond the characterisation of spatial movements to characterise how different behaviours are interwoven throughout daily animal life.

How many routes lead to migration? Comparison of methods to assess and characterize migratory movements.

Decreasing rate of migration in several species as a consequence of climate change and anthropic pressure, together with increasing evidence of space-use strategies intermediate between residency and complete migration, are very strong motivations to evaluate migration occurrence and features in animal populations. The main goal of this paper was to perform a relative comparison between methods for identifying and characterizing migration at the individual and population level on the basis of animal location data. We classified 104 yearly individual trajectories from five populations of three deer species as migratory or non-migratory, by means of three methods: seasonal home range overlap, spatio-temporal separation of seasonal clusters and the Net Squared Displacement (NSD) method. For migratory cases, we also measured timing and distance of migration and residence time on the summer range. Finally, we compared the classification in migration cases across methods and populations. All methods consistently identified migration at the population level, that is, they coherently distinguished between complete or almost complete migratory populations and partially migratory populations. However, in the latter case, methods coherently classified only about 50% of the single cases, that is they classified differently at the individual-animal level. We therefore infer that the comparison of methods may help point to 'less-stereotyped' cases in the residency-to-migration continuum. For cases consistently classified by all methods, no significant differences were found in migration distance, or residence time on summer ranges. Timing of migration estimated by NSD was earlier than by the other two methods, both for spring and autumn migrations. We suggest three steps to identify improper inferences from migration data and to enhance understanding of intermediate space-use strategies. We recommend (i) classifying migration behaviours using more than one method, (ii) performing sensitivity analysis on method parameters to identify the extent of the differences and (iii) investigating inconsistently classified cases as these may often be ecologically interesting (i.e. less-stereotyped migratory behaviours).

Pelagic seabird flight patterns are consistent with a reliance on olfactory maps for oceanic navigation.

Homing studies have provided tantalizing evidence that the remarkable ability of shearwaters (Procellariiformes) to pinpoint their breeding colony after crossing vast expanses of featureless open ocean can be attributed to their assembling cognitive maps of wind-borne odours but crucially, it has not been tested whether olfactory cues are actually used as a system for navigation. Obtaining statistically important samples of wild birds for use in experimental approaches is, however, impossible because of invasive sensory manipulation. Using an innovative non-invasive approach, we provide strong evidence that shearwaters rely on olfactory cues for oceanic navigation. We tested for compliance with olfactory-cued navigation in the flight patterns of 210 shearwaters of three species (Cory's shearwaters, Calonectris borealis, North Atlantic Ocean, Scopoli's shearwaters, C. diomedea Mediterranean Sea, and Cape Verde shearwaters, C. edwardsii, Central Atlantic Ocean) tagged with high-resolution GPS loggers during both incubation and chick rearing.We found that most (69%) birds displayed exponentially truncated scale-free(Lévy-flight like) displacements, which we show are consistent with olfactory-cued navigation in the presence of atmospheric turbulence. Our analysis provides the strongest evidence yet for cognitive odour map navigation in wild birds. Thus, we may reconcile two highly disputed questions in movement ecology, by mechanistically connecting Lévy displacements and olfactory navigation. Our approach can be applied to any species which can be tracked at sufficient spatial resolution, using a GPS logger.

The Lévy flight foraging hypothesis in a pelagic seabird.

Lévy flight foraging represents an innovative paradigm for the analysis of animal random search by including models of heavy-tailed distribution of move length, which complements the correlated random walk paradigm that is founded on Brownian walks. Theory shows that the efficiency of the different foraging tactics is a function of prey abundance and dynamics with Lévy flight being especially efficient in poor prey fields. Lévy flights have been controversial in some quarters, because they previously have been wrongly ascribed to many species through the employment of inappropriate statistical techniques and by misunderstanding movement pattern data. More recent studies using state-of-the-art statistical tools have, however, provided seemingly compelling evidence for Lévy flights. In this study, we employ these maximum-likelihood methods and their Bayesian equivalents by analysing both turning angles and move length distributions. We tested, for compliance with Lévy flight foraging, a set of 77 independent foraging trajectories of Cory's shearwaters Calonectris diomedea diomedea. Birds were tagged with high-resolution GPS loggers in two Mediterranean colonies (Linosa and Tremiti) during both incubation and chick rearing. We found that the behaviour of six birds was fitted by a correlated random walk; the movement of 32 birds was better represented by adaptive correlated random walks by switching from intensive to extensive searches; and the trajectories of 36 birds were fitted by a Lévy flight pattern of movement. The probability of performing Lévy flights was higher for trips during chick provisioning when shearwaters were forced to forage in suboptimal areas. This study supports Lévy flight foraging as an appropriate framework to analyse search tactics in this pelagic bird species and highlights that the adoption of a given search strategy is a function of biological and ecological constraints.

A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic.

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8% (vs. 18.1% of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male's territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.

On the mean path length invariance property for random walks of animals in open environment.

Random walks are common in nature and are at the basis of many different phenomena that span from neutrons and light scattering to the behaviour of animals. Despite the evident differences among all these phenomena, theory predicts that they all share a common fascinating feature known as Invariance Property (IP). In a nutshell, IP means that the mean length of the total path of a random walker inside a closed domain is fixed by the geometry and size of the medium. Such a property has been demonstrated to hold not only in optics, but recently also in the field of biology, by studying the movement of bacteria. However, the range of validity of such a universal property, strictly linked to the fulfilment of equilibrium conditions and to the statistical distributions of the steps of the random walkers, is not trivial and needs to be studied in different contexts, such as in the case of biological entities occupied in random foraging in an open environment. Hence, in this paper the IP in a virtual medium inside an open environment has been studied by using actual movements of animals recorded in nature. In particular, we analysed the behaviour of a grazer mollusc, the chiton Acanthopleura granulata. The results depart from those predicted by the IP when the dimension of the medium increases. Such findings are framed in both the condition of nonequilibrium of the walkers, which is typical of animals in nature, and the characteristics of actual animal movements.

Steering complex networks toward desired dynamics.

We consider networks of dynamical units that evolve in time according to different laws, and are coupled to each other in highly irregular ways. Studying how to steer the dynamics of such systems towards a desired evolution is of great practical interest in many areas of science, as well as providing insight into the interplay between network structure and dynamical behavior. We propose a pinning protocol for imposing specific dynamic evolutions compatible with the equations of motion on a networked system. The method does not impose any restrictions on the local dynamics, which may vary from node to node, nor on the interactions between nodes, which may adopt in principle any nonlinear mathematical form and be represented by weighted, directed or undirected links. We first explore our method on small synthetic networks of chaotic oscillators, which allows us to unveil a correlation between the ordered sequence of pinned nodes and their topological influence in the network. We then consider a 12-species trophic web network, which is a model of a mammalian food web. By pinning a relatively small number of species, one can make the system abandon its spontaneous evolution from its (typically uncontrolled) initial state towards a target dynamics, or periodically control it so as to make the populations evolve within stipulated bounds. The relevance of these findings for environment management and conservation is discussed.

Female mating tactics in lekking fallow deer (Dama dama): experience explains inter-individual variability more than costs.

Most studies on ungulate reproduction have focused on the covariates of male reproductive success, while there is much less information on female tactics of mate choice. The aim of this work is to fill this gap and to assess condition-dependent variations in female tactics in a lekking fallow deer (Dama dama) population. In particular, we investigated three indirect selection mechanisms: i) aggregation: when females join an already formed female group; ii) copying: when females copy the mate choice of other females and iii) territory choice: when females select a territory where many copulations had previously occurred. Our results show that female fallow deer, which are less experienced (younger) and/or incur higher travel costs (home range far from the lek), adopt indirect forms of mate selection more often than older females or females residing near the lek, respectively. Compared to adults, younger females remained longer in the lek (almost three times) and in male territories, returning to the lek after copulation. However, despite the time spent at the lek, younger females were not able to select the highest-rank males, and relied on territory choice more often than older females. Farther does visited the lek less frequently (farthest females only once) and arrived on average 5 days later than closer females (which performed up to 7 visits), but they were seen more often within female groups (aggregation). We did not find a different amount of copying in younger or in farther females. Our results contribute to advance our understanding of female behaviours in ungulate leks.

Olfactory-cued navigation in shearwaters: linking movement patterns to mechanisms.

After foraging in the open ocean pelagic birds can pinpoint their breeding colonies, located on remote islands in visually featureless seascapes. This remarkable ability to navigate over vast distances has been attributed to the birds being able to learn an olfactory map on the basis of wind-borne odors. Odor-cued navigation has been linked mechanistically to displacements with exponentially-truncated power-law distributions. Such distributions were previously identified in three species of Atlantic and Mediterranean shearwaters but crucially it has not been demonstrated that these distributions are wind-speed dependent, as expected if navigation was olfactory-cued. Here we show that the distributions are wind-speed dependent, in accordance with theoretical expectations. We thereby link movement patterns to underlying generative mechanisms. Our novel analysis is consistent with the results of more traditional, non-mathematical, invasive methods and thereby provides independent evidence for olfactory-cued navigation in wild birds. Our non-invasive diagnostic tool can be applied across taxa, potentially allowing for the assessment of its pervasiveness.

Generalized structural equations improve sexual-selection analyses.

Sexual selection is an intense evolutionary force, which operates through competition for the access to breeding resources. There are many cases where male copulatory success is highly asymmetric, and few males are able to sire most females. Two main hypotheses were proposed to explain this asymmetry: "female choice" and "male dominance". The literature reports contrasting results. This variability may reflect actual differences among studied populations, but it may also be generated by methodological differences and statistical shortcomings in data analysis. A review of the statistical methods used so far in lek studies, shows a prevalence of Linear Models (LM) and Generalized Linear Models (GLM) which may be affected by problems in inferring cause-effect relationships; multi-collinearity among explanatory variables and erroneous handling of non-normal and non-continuous distributions of the response variable. In lek breeding, selective pressure is maximal, because large numbers of males and females congregate in small arenas. We used a dataset on lekking fallow deer (Dama dama), to contrast the methods and procedures employed so far, and we propose a novel approach based on Generalized Structural Equations Models (GSEMs). GSEMs combine the power and flexibility of both SEM and GLM in a unified modeling framework. We showed that LMs fail to identify several important predictors of male copulatory success and yields very imprecise parameter estimates. Minor variations in data transformation yield wide changes in results and the method appears unreliable. GLMs improved the analysis, but GSEMs provided better results, because the use of latent variables decreases the impact of measurement errors. Using GSEMs, we were able to test contrasting hypotheses and calculate both direct and indirect effects, and we reached a high precision of the estimates, which implies a high predictive ability. In synthesis, we recommend the use of GSEMs in studies on lekking behaviour, and we provide guidelines to implement these models.

The Weierstrassian movement patterns of snails.

Weierstrassian Lévy walks are the archetypical form of random walk that do not satisfy the central limit theorem and are instead characterized by scale invariance. They were originally regarded as a mathematical abstraction but subsequent theoretical studies showed that they can, in principle, at least, be generated by chaos. Recently, Weierstrassian Lévy walks have been found to provide accurate representations of the movement patterns of mussels (Mytilus edulis) and mud snails (Hydrobia ulvae) recorded in the laboratory under controlled conditions. Here, we tested whether Weierstrassian Lévy walks and chaos are present under natural conditions in intertidal limpets Patella vulgata and P. rustica, and found that both characteristics are pervasive. We thereby show that Weierstrassian Lévy walks may be fundamental to how molluscs experience and interact with the world across a wide range of ecological contexts. We also show in an easily accessible way how chaos can produce a wide variety of Weierstrassian Lévy walk movement patterns. Our findings support the Lévy flight foraging hypothesis that posits that because Lévy walks can optimize search efficiencies, natural selection should have led to adaptations for Lévy walks.

Spatial patterns and density dependence in the dynamics of a roe deer (Capreolus capreolus) population in central Italy.

The demography of roe deer living in a mountain area of central Italy was studied from 1995 to 1999 with 104 radio-tagged animals, including fawns and adults of both sexes. From spring surveys we estimated population density by mark-resighting (average: 53.8±4.8 individuals km-2) and found an average fawn/doe ratio of 0.75±0.4. The fawn/doe ratio was negatively correlated to density suggesting density-dependent regulation in this population. Using culled and net-trapped individuals we evaluated the dressed body weights of adult males (23.1±1.0 kg) and females (22.0±1.0 kg), which indicated a low level of sexual dimorphism. The potential litter size (1.44±0.1 embryos) depended on female body weight and a threshold of 20.9±1.4 kg separated adult females carrying one or two embryos. Both fawn (0.38±0.07) and adult survival (0.90±0.07) were evaluated from radiotagged individuals and no gender effect was observed in either age class. During the study period we recorded a population decline in one part of the study area and an increasing fawn mortality, which was attributed to the spreading of an enteropathogenic desease. The study revealed an unexpected spatial structure in population dynamics at a scale of few square kilometres. In the two studied subareas, which are very close and ecologically similar, we documented significant differences in several demographic parameters: females in the subarea with the highest deer density produced smaller litters and allocated their reproductive effort preferentially to males, which is consistent with the hypothesis that local resource competition determines sex allocation in roe deer. The importance of spatial variability with respect to roe deer demography was overlooked in previous studies and our results raise new interesting research questions relative to the study of population equilibria which are also relevant for the management of this important game species.

Do age-specific survival patterns of wild boar fit current evolutionary theories of senescence?

Actuarial senescence is widespread in age-structured populations. In growing populations, the progressive decline of Hamiltonian forces of selection with age leads to decreasing survival. As actuarial senescence is overcompensated by a high fertility, actuarial senescence should be more intense in species with high reproductive effort, a theoretical prediction that has not been yet explicitly tested across species. Wild boar (Sus scrofa) females have an unusual life-history strategy among large mammals by associating both early and high reproductive effort with potentially long lifespan. Therefore, wild boar females should show stronger actuarial senescence than similar-sized related mammals. Moreover, being polygynous and much larger than females, males should display higher senescence rates than females. Using a long-term monitoring (18 years) of a wild boar population, we tested these predictions. We provided clear evidence of actuarial senescence in both sexes. Wild boar females had earlier but not stronger actuarial senescence than similar-sized ungulates. Both sexes displayed similar senescence rates. Our study indicates that the timing of senescence, not the rate, is associated with the magnitude of fertility in ungulates. This demonstrates the importance of including the timing of senescence in addition to its rate to understand variation in senescence patterns in wild populations.

Stochastic modelling of animal movement.

Modern animal movement modelling derives from two traditions. Lagrangian models, based on random walk behaviour, are useful for multi-step trajectories of single animals. Continuous Eulerian models describe expected behaviour, averaged over stochastic realizations, and are usefully applied to ensembles of individuals. We illustrate three modern research arenas. (i) Models of home-range formation describe the process of an animal 'settling down', accomplished by including one or more focal points that attract the animal's movements. (ii) Memory-based models are used to predict how accumulated experience translates into biased movement choices, employing reinforced random walk behaviour, with previous visitation increasing or decreasing the probability of repetition. (iii) Lévy movement involves a step-length distribution that is over-dispersed, relative to standard probability distributions, and adaptive in exploring new environments or searching for rare targets. Each of these modelling arenas implies more detail in the movement pattern than general models of movement can accommodate, but realistic empiric evaluation of their predictions requires dense locational data, both in time and space, only available with modern GPS telemetry.

Adaptive Lévy walks in foraging fallow deer.

BACKGROUND: Lévy flights are random walks, the step lengths of which come from probability distributions with heavy power-law tails, such that clusters of short steps are connected by rare long steps. Lévy walks maximise search efficiency of mobile foragers. Recently, several studies raised some concerns about the reliability of the statistical analysis used in previous analyses. Further, it is unclear whether Lévy walks represent adaptive strategies or emergent properties determined by the interaction between foragers and resource distribution. Thus two fundamental questions still need to be addressed: the presence of Lévy walks in the wild and whether or not they represent a form of adaptive behaviour. METHODOLOGY/PRINCIPAL FINDINGS: We studied 235 paths of solitary and clustered (i.e. foraging in group) fallow deer (Dama dama), exploiting the same pasture. We used maximum likelihood estimation for discriminating between a power-tailed distribution and the exponential alternative and rank/frequency plots to discriminate between Lévy walks and composite Brownian walks. We showed that solitary deer perform Lévy searches, while clustered animals did not adopt that strategy. CONCLUSION/SIGNIFICANCE: Our demonstration of the presence of Lévy walks is, at our knowledge, the first available which adopts up-to-date statistical methodologies in a terrestrial mammal. Comparing solitary and clustered deer, we concluded that the Lévy walks of solitary deer represent an adaptation maximising encounter rates with forage resources and not an epiphenomenon induced by a peculiar food distribution.