The population bottleneck of the Iberian wolf impacted genetic diversity but not admixture with domestic dogs: A temporal genomic approach.

After decades of intense persecution, the Iberian wolf subspecies faced a severe bottleneck in the 1970s that considerably reduced its range and population size, nearly leading to its extinction in central and southern Iberian Peninsula. Such population decline could have impacted the genetic diversity of Iberian wolves through different processes, namely genetic drift and dynamics of hybridization with domestic dogs. By contrasting the genomes of 68 contemporary with 54 historical samples spanning the periods before and immediately after the 1970s bottleneck, we found evidence of its impact on genetic diversity and dynamics of wolf-dog hybridization. Our genome-wide assessment revealed that wolves and dogs form two well-differentiated genetic groups in Iberia and that hybridization rates did not increase during the bottleneck. However, an increased number of hybrid individuals was found over time during the population re-expansion, particularly at the edge of the wolf range. We estimated a low percentage of dog ancestry (~1.4%) in historical samples, suggesting that dog introgression was not a key driver for wolf extinction in central and southern Iberia. Our findings also unveil a significant decline in genetic diversity in contemporary samples, with the highest proportion of homozygous segments in the genome being recently inherited. Overall, our study provides unprecedented insight into the impact of a sharp decline on the Iberian wolf genome and refines our understanding of the ecological and evolutionary drivers of wolf-dog hybridization in the wild.

Carnivore conservation needs evidence-based livestock protection.

Carnivore predation on livestock often leads people to retaliate. Persecution by humans has contributed strongly to global endangerment of carnivores. Preventing livestock losses would help to achieve three goals common to many human societies: preserve nature, protect animal welfare, and safeguard human livelihoods. Between 2016 and 2018, four independent reviews evaluated >40 years of research on lethal and nonlethal interventions for reducing predation on livestock. From 114 studies, we find a striking conclusion: scarce quantitative comparisons of interventions and scarce comparisons against experimental controls preclude strong inference about the effectiveness of methods. For wise investment of public resources in protecting livestock and carnivores, evidence of effectiveness should be a prerequisite to policy making or large-scale funding of any method or, at a minimum, should be measured during implementation. An appropriate evidence base is needed, and we recommend a coalition of scientists and managers be formed to establish and encourage use of consistent standards in future experimental evaluations.

The place of nature in conservation conflicts.

Conservation conflicts are gaining importance in contemporary conservation scholarship such that conservation may have entered a conflict hype. We attempted to uncover and deconstruct the normative assumptions behind such studies by raising several questions: what are conservation conflicts, what justifies the attention they receive, do conservation-conflict studies limit wildlife conservation, is scientific knowledge stacked against wildlife in conservation conflicts, do conservation-conflict studies adopt a specific view of democracy, can laws be used to force conservation outcomes, why is flexibility needed in managing conservation conflicts, can conservation conflicts be managed by promoting tolerance, and who needs to compromise in conservation conflicts? We suggest that many of the intellectual premises in the field may defang conservation and prevent it from truly addressing the current conservation crisis as it accelerates. By framing conservation conflicts as conflicts between people about wildlife or nature, the field insidiously transfers guilt, whereby human activities are no longer blamed for causing species decline and extinctions but conservation is instead blamed for causing social conflicts. When the focus is on mitigating social conflicts without limiting in any powerful way human activities damaging to nature, conservation-conflict studies risk keeping conservation within the limits of human activities, instead of keeping human activities within the limits of nature. For conservation to successfully stop the biodiversity crisis, we suggest the alternative goal of recognizing nature's right to existence to maintenance of ecological functions and evolutionary processes. Nature being a rights bearer or legal person would imply its needs must be explicitly taken into account in conflict adjudication. If, even in conservation, nature's interests come second to human interests, it may be no surprise that conservation cannot succeed.

El Lugar de la Naturaleza en los Conflictos de Conservación Resumen Los conflictos de conservación están ganando importancia en los últimos años en la disciplina de la biología de la conservación de tal manera que puede ser que la conservación haya entrado en una exageración de conflictos. En este ensayo, hemos llevado a cabo un ejercicio de deconstrucción de las asunciones detrás de dichos estudios al formular varias preguntas: qué son los conflictos de conservación, qué justifica la atención que reciben, si los estudios sobre los conflictos de conservación pueden llegar a limitar la conservación de la fauna, si el conocimiento científico puede llegar a afectar la conservación cuando se involucra en los conflictos de conservación, si los estudios sobre los conflictos de conservación adoptan una visión específica de la democracia, si las leyes ambientales pueden usarse para forzar resultados positivos en conservación, por qué se requiere flexibilidad para gestionar los conflictos de conservación, si los conflictos de conservación pueden gestionarse con el objetivo de aumentar la tolerancia hacia las especies, y quién necesita ceder en los conflictos de conservación. Sugerimos que muchas de las premisas intelectuales detrás de este tipo de estudios pueden debilitar el objetivo de conservar la naturaleza de forma efectiva. Al enmarcar a los conflictos de conservación como conflictos entre personas en relación a la conservación de la naturaleza, se transfiere la culpa de manera más insidiosa, mientras que las actividades humanas dejan de ser culpables por originar el declive y extinción de las especies y, al mismo tiempo, la conservación de la naturaleza pasa a ser culpable de ocasionar conflictos sociales. Cuando el enfoque de los conflictos de conservación se centra en mitigar el conflicto social sin limitar de ninguna manera las actividades humanas que son dañinas para la naturaleza, los estudios sobre los conflictos de conservación corren el riesgo de enmarcar a la conservación de la naturaleza dentro de los límites de las actividades humanas, en lugar de mantener a las actividades humanas dentro de los límites de la naturaleza. Para que la conservación de la naturaleza se efectiva ante la actual crisis de la biodiversidad sugerimos el objetivo alternativo de reconocer el derecho de la naturaleza a su existencia, el mantenimiento de sus funciones ecológicas y los procesos evolutivos. Que la naturaleza tenga derechos o sea considerada como una persona legal implicaría que se deben considerar explícitamente sus necesidades durante la gestión de conflictos de conservación. Si por el contrario, dentro de la conservación los intereses de la naturaleza son subordinados por defecto a los intereses humanos, no debería sorprendernos que la conservación frecuentemente no sea exitosa.

Multimethod, multistate Bayesian hierarchical modeling approach for use in regional monitoring of wolves.

In many cases, the first step in large-carnivore management is to obtain objective, reliable, and cost-effective estimates of population parameters through procedures that are reproducible over time. However, monitoring predators over large areas is difficult, and the data have a high level of uncertainty. We devised a practical multimethod and multistate modeling approach based on Bayesian hierarchical-site-occupancy models that combined multiple survey methods to estimate different population states for use in monitoring large predators at a regional scale. We used wolves (Canis lupus) as our model species and generated reliable estimates of the number of sites with wolf reproduction (presence of pups). We used 2 wolf data sets from Spain (Western Galicia in 2013 and Asturias in 2004) to test the approach. Based on howling surveys, the naïve estimation (i.e., estimate based only on observations) of the number of sites with reproduction was 9 and 25 sites in Western Galicia and Asturias, respectively. Our model showed 33.4 (SD 9.6) and 34.4 (3.9) sites with wolf reproduction, respectively. The number of occupied sites with wolf reproduction was 0.67 (SD 0.19) and 0.76 (0.11), respectively. This approach can be used to design more cost-effective monitoring programs (i.e., to define the sampling effort needed per site). Our approach should inspire well-coordinated surveys across multiple administrative borders and populations and lead to improved decision making for management of large carnivores on a landscape level. The use of this Bayesian framework provides a simple way to visualize the degree of uncertainty around population-parameter estimates and thus provides managers and stakeholders an intuitive approach to interpreting monitoring results. Our approach can be widely applied to large spatial scales in wildlife monitoring where detection probabilities differ between population states and where several methods are being used to estimate different population parameters.

Using a top predator as a sentinel for environmental contamination with pathogenic bacteria: the Iberian wolf and leptospires.

The Iberian wolf (Canis lupus) is the top predator in the Iberian environments in which it lives, feeding on a wide range of species, thus encountering a wide range of disease agents. Therefore, the wolf can serve as sentinel of environmental contamination with pathogens. We investigated the exposure of free-living wolves to 14 serovars of Leptospira interrogans sensu lato. Kidney samples from 49 wolves collected from 2010-2013 in northwestern Spain were analysed by culture, direct immunofluorescence and polymerase chain reaction. Tissue fluids were analysed for antibodies by a microscopic agglutination test. Ten wolves (observed prevalence: 20%, 95% confidence interval = 11-33%) showed evidence of contact with leptospires, eight through direct detection and nine through serology (7 wolves were positive according to both techniques). Titres below the cut-off level were also detected in seven cases. Serovars confirmed were Canicola (n = 4), Icterohaemorrhagiae (n = 3) and Sejroë, Ballum and Grippotyphosa (n = 1 each), indicating that wolves were infected with serovars for which dogs, rodents and ungulates, are the natural hosts and supporting the utility of the wolf and other large predators as environmental sentinels for pathogens.

Responses of a top and a meso predator and their prey to moon phases.

We compared movement patterns and rhythms of activity of a top predator, the Iberian lynx Lynx pardinus, a mesopredator, the red fox Vulpes vulpes, and their shared principal prey, the rabbit Oryctolagus cuniculus, in relation to moon phases. Because the three species are mostly nocturnal and crepuscular, we hypothesized that the shared prey would reduce its activity at most risky moon phases (i.e. during the brightest nights), but that fox, an intraguild prey of lynx, would avoid lynx activity peaks at the same time. Rabbits generally moved further from their core areas on darkest nights (i.e. new moon), using direct movements which minimize predation risk. Though rabbits responded to the increased predation risk by reducing their activity during the full moon, this response may require several days, and the moon effect we observed on the rabbits had, therefore, a temporal gap. Lynx activity patterns may be at least partially mirroring rabbit activity: around new moons, when rabbits moved furthest and were more active, lynxes reduced their travelling distances and their movements were concentrated in the core areas of their home ranges, which generally correspond to areas of high density of rabbits. Red foxes were more active during the darkest nights, when both the conditions for rabbit hunting were the best and lynxes moved less. On the one hand, foxes increased their activity when rabbits were further from their core areas and moved with more discrete displacements; on the other hand, fox activity in relation to the moon seemed to reduce dangerous encounters with its intraguild predator.

Possible extinction vortex for a population of Iberian lynx on the verge of extirpation.

Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female-biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co-occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.

Linking landscape structure and vegetation productivity with nut consumption by the Cantabrian brown bear during hyperphagia.

In bears, reproduction is dependent on the body reserves accumulated during hyperphagia. The Cantabrian brown bear mainly feeds on nuts during the hyperphagia period. Understanding how landscape heterogeneity and vegetation productivity in human-dominated landscapes influence the feeding habits of bears may therefore be important for disentangling species-habitat relationships of conservation interest. We determined the spatial patterns of nut consumption by brown bears during the hyperphagia period in relation to landscape structure, characteristics of fruit-producing patches and vegetation productivity. For this purpose, we constructed foraging models based on nut consumption data (obtained by scat analysis), by combining vegetation productivity data, topographical variables and landscape metrics to identify nut foraging patterns during this critical period for bears. The average wooded area of patches where scats were collected and where the nuts that the bears had consumed were produced was larger than that of the corresponding patches where nuts were not produced. For scats collected outside of nut-producing patches, the distance between the scats and the patches was greatest for chestnut-producing patches. Elevation, Gross Primary Production (GPP) and the Aggregation Index (AI) were good predictors of acorn consumption in the models. Good model fits were not obtained for data on chestnut consumption in bears. The findings confirm that brown bears feeding on nuts show a preference for relatively large, highly aggregated patches with a high degree of diversity in the landscape pattern, which may help the bears to remain undetected. The nut prediction model highlights areas of particular importance for brown bears during hyperphagia. The human presence associated with sweet chestnut forest stands or orchards may make bears feel more vulnerable when feeding.

Hair cortisol concentration reflects the life cycle and management of grey wolves across four European populations.

The grey wolf (Canis lupus) persists in a variety of human-dominated landscapes and is subjected to various legal management regimes throughout Europe. Our aim was to assess the effects of intrinsic and methodological determinants on the hair cortisol concentration (HCC) of wolves from four European populations under different legal management. We determined HCC by an enzyme-linked immune assay in 259 hair samples of 133 wolves from the Iberian, Alpine, Dinaric-Balkan, and Scandinavian populations. The HCC showed significant differences between body regions. Mean HCC in lumbar guard hair was 11.6 ± 9.7 pg/mg (range 1.6-108.8 pg/mg). Wolves from the Dinaric-Balkan and Scandinavian populations showed significantly higher HCC than Iberian wolves, suggesting that harvest policies could reflected in the level of chronic stress. A significant negative relationship with body size was found. The seasonal, sex and age patterns are consistent with other studies, supporting HCC as a biomarker of chronic stress in wolves for a retrospective time frame of several weeks. Our results highlight the need for standardization of sampling and analytical techniques to ensure the value of HCC in informing management at a continental scale.

Assessment of the conservation efforts to prevent extinction of the Iberian lynx.

The Iberian lynx (Lynx pardinus) may be the first charismatic felid to become extinct in a high-income country, despite decades of study and much data that show extinction is highly probable. The International Union for Conservation of Nature categorizes it as critically endangered; about 200 free-ranging individuals remain in two populations in southern Spain. Conservation measures aimed at averting extirpation have been extensively undertaken with 4 of the former 10 Iberian lynx populations recorded 25 years ago. Two of the four populations have been extirpated. The number of individuals in the third population have declined by 83%, and in the fourth the probability of extirpation has increased from 34% to 95%. Major drivers of the pending extinction are the small areas to which conservation measures have been applied; lack of incorporation of evidence-based conservation, scientific monitoring, and adaptive management into conservation efforts; a lack of continuity in recovery efforts, and distrust by conservation agencies of scientific information. In contrast to situations in which conservation and economic objectives conflict, in the case of the Iberian lynx all stakeholders desire the species to be conserved.

Reproductive traits in captive and free-ranging males of the critically endangered Iberian lynx (Lynx pardinus).

The Iberian lynx (Lynx pardinus) is the most endangered felid in the world. Adequate genetic management of in situ and ex situ populations, and linkage between both, require knowledge on male reproductive biology and factors influencing it. We examined the influence of age, free-ranging versus captive conditions and seasonality on phenotypic, endocrine and semen traits, and links between reproductive traits and male fertility. Males had relatively small testes, produced low sperm numbers, a low proportion of normal sperm, and a high proportion of motile sperm. Young (2-year-old) males had lower testosterone levels, fewer sperm, and a lower proportion of motile and normal sperm than > or =4-year-old males. No major differences were found in semen traits before and after the mating season or between free-ranging and captive males, although the latter had better sperm motility. Males with larger relative testes weight and more sperm copulated more frequently, whereas males that produced more sperm with higher motility produced more cubs per female. In conclusion, small relative testes size and low sperm quality could indicate either low levels of sperm competition or high levels of inbreeding. Young males are probably subfertile; there is a slight trend for males in the captive breeding programme to have better semen quality than wild males, and males with higher sperm production are sexually more active and more fertile. These findings have major implications for decisions regarding which males should breed, provide samples for the genetic resource bank, or participate in programmes involving the use of assisted reproductive techniques.

Wireless Sensor Network deployment for monitoring wildlife passages.

Wireless Sensor Networks (WSNs) are being deployed in very diverse application scenarios, including rural and forest environments. In these particular contexts, specimen protection and conservation is a challenge, especially in natural reserves, dangerous locations or hot spots of these reserves (i.e., roads, railways, and other civil infrastructures). This paper proposes and studies a WSN based system for generic target (animal) tracking in the surrounding area of wildlife passages built to establish safe ways for animals to cross transportation infrastructures. In addition, it allows target identification through the use of video sensors connected to strategically deployed nodes. This deployment is designed on the basis of the IEEE 802.15.4 standard, but it increases the lifetime of the nodes through an appropriate scheduling. The system has been evaluated for the particular scenario of wildlife monitoring in passages across roads. For this purpose, different schemes have been simulated in order to find the most appropriate network operational parameters. Moreover, a novel prototype, provided with motion detector sensors, has also been developed and its design feasibility demonstrated. Original software modules providing new functionalities have been implemented and included in this prototype. Finally, main performance evaluation results of the whole system are presented and discussed in depth.

The effectiveness of conditioned aversion in wolves: Insights from experimental tests.

It has been suggested that conditioned food aversion (CFA) could be a potential non-lethal intervention by which to deter attacks on livestock by large carnivores. CFA occurs when an animal associates the characteristics of a food with an illness, thus rejecting that food in subsequent encounters. CFA can be associated with an artificial odour during conditioning. Despite the debate surrounding the use of this intervention, more studies evaluating the effectiveness of CFA are necessary. We experimentally evaluated the potential of microgranulated levamisole + a vanilla odour cue to induce CFA in captive Iberian wolves (Canis lupus signatus). Four out of the five wolves treated showed an aversion to the meat for a minimum of one month after conditioning. The microgranulated presentation masked the flavour and smell of the levamisole but increased its volume, which may have facilitated its detection by the wolves. We also observed that the strength of the odour played an important role in the aversion extinction. The use of microgranulated levamisole + an odour cue has the potential to be used as an intervention by which to induce aversive conditioning in wolves in the wild, although rigorous field tests are required. We discuss the potential of CFA to deter attacks on livestock by large carnivores.

Not exodus, but population increase and gene flow restoration in Cantabrian brown bear (Ursus arctos) subpopulations. Comment on Gregório et al. 2020.

In a genetic study on brown bears (Ursus arctos) in the Cantabrian Mountains, Gregório et al. (2020) interpreted the asymmetrical gene flow they found from the eastern subpopulation towards the western one as an exodus of bears forced to flee from the eastern nucleus "with higher human disturbance and poaching", concluding that connectivity may be operating as a means for eastern Cantabrian bears to find more suitable territories. In this reply, we maintain that the explanations of Gregorio et al. contradict the source-sink theory and we also present demographic data not considered by these authors showing that the eastern subpopulation is not declining, but persistently increasing. After reviewing the demographic and genetic studies published during the last 20 years, we conclude that the connectivity between the two subpopulations is operating as a route which allows the regular movement of males and the restoration of the gene flow across the whole Cantabrian population.

Anthropogenic food resources sustain wolves in conflict scenarios of Western Iran.

The feeding ecology of gray wolves has been investigated extensively worldwide. Despite previous studies on food habits of wolves in Asia and Iran, none has focused on the diet of the species in a scenario of depleted of wild prey and with recent records of attacks on humans. Here, we combined telemetry methods and scat analysis to study the diet of wolves in areas of Hamadan province, Iran, where medium to large wild prey is almost absent. Between October 2015 and March 2017, we studied the feeding behavior (by identifying feeding sites through clusters of GPS locations) of three wolves fitted with GPS collars, belonging to different wolf packs. We also collected and analyzed 110 wolf scats during the same period within the same areas. Overall, we investigated 850 clusters of GPS locations in the field, and identified 312 feeding sites. Most feeding clusters were linked to dumpsites and poultry farms around villages. We found 142 and 170 events of predatory (kill sites) and scavenging behavior, respectively. Prey composition based on kill sites was comprised of 74.6% livestock, 19.7% lagomorphs, 3.5% dogs, 1.4% red fox, and 0.7% golden jackal. Similarly, prey composition based on scavenging clusters was comprised of 79.9% livestock, 10.6% red fox, and 9.4% golden jackal. Scat analysis, however, indicated that livestock (34.3%), garbage (23.7%), poultry (16.0%), and European hare (15.4%) were the most frequent food items. We discuss the role of anthropogenic food sources in a context where agonistic wolf-human encounters occur recurrently, and suggest management guidelines regarding illegal dumping of animal carcasses and garbage dumpsites, in order to minimize wolf-human negative interactions.

Eurasian lynx fitness shows little variation across Scandinavian human-dominated landscapes.

Despite extensive research on the ecology and behavioural adaptations of large carnivores in human-dominated landscapes, information about the fitness consequences of sharing landscapes is still limited. We assessed the variation in three consecutive components of female fitness: the probability of reproduction, litter size and juvenile survival in relation to environmental and human factors in a solitary carnivore, the Eurasian lynx (Lynx lynx), occurring in human-dominated landscapes in Scandinavia. We used demographic data from 57 radio-collared adult females between 1995-2011 (126 radio-years). Overall, the yearly probability of female reproduction was 0.80, mean litter size was 2.34 (range 1-4) and the probability to find a female that reproduced in the spring being accompanied by at least one offspring during the subsequent winter was 0.70. We did not find evidence that food availability was a key factor influencing female fitness. Female lynx may adapt to food availability when establishing their home ranges by adopting an obstinate strategy, ensuring a minimum amount of prey necessary for survival and reproduction even during periods of prey scarcity. In human-dominated landscapes, where sufficient prey are available for lynx, mortality risk may have a larger influence on lynx population dynamics compared to food availability. Our results suggest that lynx population dynamics in human-dominated landscapes may be mainly driven by human impacts on survival.