Effect of nest composition, experience and nest quality on nest-building behaviour in the Bonelli's Eagle.

In bi-parentally built nests, there is evidence to suggest that nests are extended phenotypic signals that accurately indicate the quality of the building parent/s. Raptors often use a variety of materials to build their nests (natural, such as branches, but also non-natural objects), presumably due to their insulating properties, their suitability to advertise occupancy of the nest, and to decrease pathogen and parasite loads. However, in raptors where both sexes collaborate in nest construction, it is unclear whether nest building (taking the amount of material carried to the nest as the potential predictor) is an indicator of parental quality, and whether the effort expended by both sexes could constitute an honest signal of parental quality to their partners. Between 2011 and 2016, we monitored 16 nests of Bonelli's Eagles (Aquila fasciata), and we examined data on sex, type of material brought to the nest, breeding experience, nest quality, timing, and nest-building investment prior to egg-laying from 32 identifiable Bonelli's Eagles during the pre-laying period to investigate the relative contribution of the sexes to the amount of nest material gathered. Our results indicate that sex is not a determining factor in nest-building effort, and that females did not increase their parental effort in response to the male's contribution, and supply of materials did not increase during the pre-laying period. In contrast, our models showed that: (1) the type of material supplied to the nest by both sexes varied significantly throughout the pre-laying period and (2) nest-building effort was determined by individual experience and nest quality. Therefore, our study suggests that male nest-building behaviour and investment by Bonelli's Eagles cannot be considered as an extended phenotypic signal. The differential use of hard and green material by both sexes in the early and late stages of nest-building period, and the fact that the more experienced individuals contributed a larger amount of material on low quality nests, are discussed in the contexts of signaling nest occupancy to conspecifics and competitors and the decrease of ectoparasite loads during the pre-laying period.

Ecological values of intermittent rivers for terrestrial vertebrate fauna.

Rivers are generally considered critical habitats for biodiversity; however, this often ignores the fact that many rivers may run dry and support terrestrial as well as aquatic fauna. Here, we investigated the ecological value of intermittent rivers for terrestrial vertebrates by installing camera traps along rivers subject to varying dry periods in two contrasting European climatic zones. We then analysed i) species presence and behaviours (as a proxy of ecological functions) on perennial and intermittent streams; ii) environmental (hydrological and geomorphological) and anthropogenic factors affecting the frequency of occurrence and number of species recorded; and iii) the importance of hydrological factors as regards ecological functioning. In both study areas, we recorded a higher number of species and individuals along intermittent streams than perennial streams, with highest values in intermittent reaches exhibiting shorter dry periods. Both abundance and species richness were strongly affected by hydrological factors in both study areas, including not only the occurrence but also the duration of the dry period. Dry channels played a key role as migration corridors and as a source of food, being used more frequently than riparian habitats when the river ran dry. Our findings indicate that terrestrial vertebrate fauna benefit from dry phases in rivers. Intermittent rivers, supporting a high abundance and diversity of fauna, should be considered as target ecosystems for wildlife conservation. Not doing so will jeopardise urgently needed conservation strategies in the face of accelerating global climate change.

Territorial occupancy dynamics in a forest raptor community.

A Markovian modeling approach was used to explore territorial interactions among three forest raptors coexisting in a forested natural area in southeast Spain: the booted eagle (Hieraaetus pennatus), the common buzzard (Buteo buteo) and the northern goshawk (Accipiter gentilis). Using field data collected over a period of 12 years, 11 annual transition matrices were built, considering four occupancy states for each territory. The model describes transitional processes (colonization, abandonment, replacement and persistence), permits temporal variations in the transition matrix to be tested, and simulates territorial occupation for a few subsequent years. Parameters for the species and community dynamics were described in terms of turnover times and damping ratio. A perturbation analysis was performed to simulate the effects of changes in the transition probabilities on the stable state distribution. Our results indicate the existence of a stable community, largely dominated by the booted eagles, and described by a time-invariant transition matrix. Despite the stability observed, the territorial system is highly dynamic, with frequent abandonment and colonization events, although interspecific territorial interactions (the replacement of one species by another) are uncommon. Consequently, the three species appear to follow relatively independent occupancy dynamics. Simulation of potential management actions showed that substantial increases in the number of territories occupied by the less common species (goshawk and buzzard) can only be attained if relatively large increases in their reoccupation and colonization rates are considered.

Efficiency of a protected-area network in a Mediterranean region: a multispecies assessment with raptors.

Three different systems of designating protected areas in a Mediterranean region in southeastern Spain were studied, referring to their effectiveness and efficiency for protecting both the breeding territories and the suitable habitat of a set of ten raptor species. Taking into consideration the varying degrees of endangerment of these species, a map of multispecies conservation values was also drawn up and superimposed on the three protected-area systems studied. In order to compare the levels of protection afforded by the three systems, we considered two indices that measured their relative effectiveness and efficiency. The effectiveness estimated the proportion of territories or optimal habitat protected by the networks while efficiency implicitly considered the area of each system (percentage of breeding territories or optimal habitat protected per 1% of land protected). Overall, our results showed that the most efficient system was that formed by the set of regional parks and reserves (17 protected breeding territories per 100 km²), although, given its small total area, it was by far the least effective (only protecting the 21% of the breeding territories of all species and 17% of the area of high conservation value). The systems formed by the Special Protection Areas (designated under the EU "Birds Directive") and by the Special Conservation Areas (designated under the EU "Habitats Directive") notably increased the percentages of protected territories of all species (61%) and area of high conservation value (57%), but their efficiency was not as high as expected in most cases. The overall level of protection was high for all species except for the Lesser Kestrel (Falco naumanni), an endangered falcon that inhabits pseudo-steppe and traditional agricultural habitats, which are clearly underrepresented in the protected-area network of the study region.

Comparing detectability patterns of bird species using multi-method occupancy modelling.

A robust knowledge of biodiversity distribution is essential for designing and developing effective conservation actions. The choice of a suitable sampling method is key to obtaining sufficiently accurate information of species distribution and consequently to improve biodiversity conservation. This study applies multi-method occupancy models to 36 common bird species associated with small ponds in the province of Murcia (south-eastern Spain), one of the most arid regions of Europe, in order to compare their effectiveness for detecting different bird species: direct observation, combined observation and video monitoring and mist netting captures. The results showed that the combined method and direct observation were similar and most effective than mist netting for detecting species occupancy, although detection rates ranged widely among bird groups, while some large species were poorly detected by all the methods used. Average detectability did not increase during the breeding period. The chosen approach is particularly applicable to both single- and multi-species bird monitoring programmes. However, we recommend evaluating the cost-effectiveness of all the available methods in order to reduce costs and improve the success of sampling designs.

Long-term population monitoring of a territorial forest raptor species.

We provide field monitoring data of a territorial raptor (the booted eagle, Hieraaetus pennatus), that was intensively monitored over a period of 18 years (1998-2015) in a Mediterranean forested area of south-eastern Spain designated as a Special Protection Area (Natura 2000 Network) for this species. The data set compiles all the relevant information about the occupation of territories and nests, reproductive ecology, long-term monitoring of marked individuals and influence of parent's colour morph on brood size. Several questions concerning the population ecology of forest-dwelling raptors and factors conditioning territorial occupancy, such as location cues or site fidelity, are addressed. This type of long-term population monitoring has high potential for replication, reuse and comparison purposes, providing insights for monitoring other long-lived, territorial species.

Blood concentrations of 50 elements in Eagle owl (Bubo bubo) at different contamination scenarios and related effects on plasma vitamin levels.

Some metals and metalloids (e.g. Pb, Hg, Cd and As) are well-known for their bioaccumulation capacity and their toxic effects on birds, but concerns on other minor elements and rare earth elements (ME and REE) are growing due to their intensive use in modern technology and potential toxicity. Vitamins and carotenoids play essential roles in nestling growth and proper development, and are known to be affected by the metals classically considered as toxic. However, we are unaware of any attempts to evaluate the exposure to 50 elements and related effects in plasma vitamins and carotenoids in raptor species. The main goals of this study are: (i) to assess the exposure to 50 elements (i.e. classic toxic elements, trace elements, REE and ME) in nestling Eagle owls (Bubo bubo) inhabiting three differently polluted environments (mining, industrial and control areas) in southeastern Spain, and (ii) to evaluate how element exposure affects plasma vitamin and carotenoid levels, hematocrit and body measurements (mass and wing length) of the individuals. Our results show that local contamination in the mining area contributes to increased blood concentrations of Pb, As and Tl in nestlings, while diet differences between control and mining/industrial areas may account for the different levels of Mn, Zn, and Sr in blood, and lutein in plasma. Plasma tocopherol levels were increased in the mining-impacted environment, which may be a mechanism of protection to prevent toxic element-related oxidative stress. Plasma α-tocopherol was enhanced by 20% at blood Pb concentrations ≥8 ng/ml, and nestlings exhibited up to 56% increase in α-tocopherol levels when blood Pb concentrations reached 170 ng/ml. Tocopherol seems to be a sensitive biomarker under an exposure to certain toxic elements (e.g. Pb, As, Tl).

Changes in blood pesticide levels in booted eagle (Hieraaetus pennatus) associated with agricultural land practices.

We estimated the degree of exposure to lindane and endosulfan in the blood of booted eagle nestlings (Hieraaetus pennatus) (1999-2003), in order to assess the usefulness of these samples as a unit for monitoring changes in exposure as a result of shifts in agricultural practices and the implementation of legal measures. The highest blood lindane concentrations were obtained 1 year prior to its prohibition by the European Union. Subsequent to that year, the drop in blood concentrations was dramatic. Furthermore, endosulfan blood concentrations follow a progression coinciding with an increase in olive, grape and plum-tree crops. We conclude that concentrations of organochlorine pesticides in the blood of booted eagle nestlings may be used to monitor the use of those pesticides over a particular agricultural region and alert the authorities of possible environmental or health risks.

Copulatory behaviour in the Bonelli´s Eagle: Assessing the paternity assurance hypothesis.

We examined copulatory behaviour in the Bonelli´s Eagle (Aquila fasciata) at nesting sites in the eastern zone of the Baetic Cordillera, southern Spain, between 2010 and 2012. We observed the copulatory behaviour of 15 pairs during the pre-laying period. Bonelli´s Eagles commenced sexual activity ca. 69 days before egg-laying. Ninety-six percent of mounting attempts were successful. Bonelli´s Eagle pairs averaged 99.8 copulation attempts per clutch, with an average copulation frequency of 0.86 copulation attempts per day. Pairs displayed a daily bimodal pattern of copulation activity, with copulations occurring most frequently in the evening. We used our data to test three predictions with regard to the paternity assurance hypothesis. Prediction 1, that within-pair copulations increase with local breeding density, was rejected because our models showed no evidence for it. Prediction 2, that within-pair copulations increase during the female fertile period, was marginally supported. Finally, Prediction 3, that mate attendance increases during the female fertile period, was also rejected because mate-guarding did not increase as the fertile period approached. However, mate-guarding was positively correlated with within-pair copulation frequency. Moderate copulation rates compared to other raptors and the absence of mate-guarding suggest that, in the study area, Bonelli´s Eagles exhibit only partially adaptive behaviour to assure their paternity. A possible explanation could be related to the low number of extra-pair encounters observed (opportunities for which appear to be rare), although the gradual increase in within-pair copulations during the female fertile period is consistent with the sperm competition hypothesis. The results are discussed based on the signalling hypothesis, which proposes that raptors signal territory ownership to conspecifics, and possibly to other raptor species, by copulating frequently and conspicuously in the defended nesting area.

Use of classical bird census transects as spatial replicates for hierarchical modeling of an avian community.

New monitoring programs are often designed with some form of temporal replication to deal with imperfect detection by means of occupancy models. However, classical bird census data from earlier times often lack temporal replication, precluding detection-corrected inferences about occupancy. Historical data have a key role in many ecological studies intended to document range shifts, and so need to be made comparable with present-day data by accounting for detection probability. We analyze a classical bird census conducted in the region of Murcia (SE Spain) in 1991 and 1992 and propose a solution to estimating detection probability for such historical data when used in a community occupancy model: the spatial replication of subplots nested within larger plots allows estimation of detection probability. In our study, the basic sample units were 1-km transects, which were considered spatial replicates in two aggregation schemes. We fit two Bayesian multispecies occupancy models, one for each aggregation scheme, and evaluated the linear and quadratic effect of forest cover and temperature, and a linear effect of precipitation on species occupancy probabilities. Using spatial rather than temporal replicates allowed us to obtain individual species occupancy probabilities and species richness accounting for imperfect detection. Species-specific occupancy and community size decreased with increasing annual mean temperature. Both aggregation schemes yielded estimates of occupancy and detectability that were highly correlated for each species, so in the design of future surveys ecological reasons and cost-effective sampling designs should be considered to select the most suitable aggregation scheme. In conclusion, the use of spatial replication may often allow historical survey data to be applied formally hierarchical occupancy models and be compared with modern-day data of the species community to analyze global change process.

Raptor ecotoxicology in Spain: a review on persistent environmental contaminants.

Initial studies on the pressure from environmental contaminants on raptor populations in Spain date back to the 1980s, and they have been carried out from a range of viewpoints using a range of sentinel raptor species. However, there is no national monitoring scheme, and therefore the research carried out has been sporadic both spatially and temporally. The exposure to metals has not varied over time, except in the case of lead, whose concentration in eggs and tissues has diminished. In general, the concentrations of metals detected in raptor samples from Spain are generally low and not sufficient to produce toxic effects. Excepting DDT and DDE, most organochlorine-based pesticides in raptors from Spain have diminished over the last 2 decades. The concentrations of DDE found in the eggs of various species could in part explain problems in the reproductive success of raptors in Spain.

Nest sites as a key resource for population persistence: A case study modelling nest occupancy under forestry practices.

Natural nest sites are important breeding resource in terms of population dynamics, especially in forest systems where nest trees limit populations or timber harvesting destroys nests. Nest structures usually have a long life and can be reused by breeding pairs across multiple breeding seasons, so studying their dynamics is of relevance for biodiversity conservation. In this study, we develop a dynamic model to evaluate nest site availability and its influence on the breeding settlement of a forest raptor community composed of booted eagle (Hieraaetus pennatus), common buzzard (Buteo buteo) and northern goshawk (Accipiter gentilis) in a Mediterranean forest ecosystem in southeast Spain. This model approach is also applied to analyse the influence of forestry practices on the dynamics of occupied nests for a simulated period (2010-2050). The simulated scenarios include unmanaged forest and timber harvesting practices of clearcuttings every ten years considering two factors: the age class of trees for clearcutting (40, 50, 60 and 70 years old) and the type of forest management (with or without nest protection). Our simulated results show that the number of breeding pairs is constant during the period without timber harvest, whereas breeding pairs gradually decrease in the scenario of clearcutting trees aged from 70 to 50-years without nest protection, and populations become extinct with the clearcutting of 40-year old trees. Considering the practice of clearcutting and nest protection, nest occupancy can reach the maximum number of occupied nests for the scenarios of cutting 70 and 60-year old trees, and maintain populations without extinction for the scenarios of cutting 40-year old trees. We conclude that nest sites (whether occupied or not) are key resources for increasing the occupancy of the forest raptor community and that nest protection measures buffer the effects of clearcuttings, thus preventing population extinction.

Organochlorine residues in booted eagle (Hieraaetus pennatus) and goshawk (Accipiter gentilis) eggs from southeastern Spain.

Most organochlorine (OC) use has been banned in Spain, but these compounds are persistent and may still adversely affect predatory birds. Data generally are lacking, however. Residues of hexachlorobenzene, alpha-hexachlorocyclohexane, lindane, hexachloro-octahydro-epoxy-dimethanonaphthalene, DDT, dichlorodiphenyldichloroethane, dichlorodiphenyldichloroethylene (DDE), and polychlorinated biphenyls (PCBs) were measured in 22 failed eggs of booted eagles (Hieraaetus pennatus) and goshawks (Accipiter gentilis) from southeastern Spain; both species are declining in this region. Hexachloro-octahydro-epoxy-dimethanonaphthalene, DDE, and sum PCB congener concentrations were significantly higher in booted eagle than in goshawk eggs, and an inverse relationship was found between shell thickness and DDE concentrations in booted eagles. Organochlorides may have been associated with the failure of some booted eagle eggs, but concentrations in booted eagle and goshawk eggs decreased over the period during which populations have dwindled. Thus, although OCs may be a contributory factor, they are unlikely to be the primary cause of the recent population declines in southeastern Spain.

Assessment of mercury exposure and maternal-foetal transfer in Miniopterus schreibersii (Chiroptera: Miniopteridae) from southeastern Iberian Peninsula.

Mercury (Hg) is a highly toxic and widely distributed metal that is bioaccumulated in insectivorous mammals and may cause adverse effects on the reproductive system. Bats are considered excellent Hg bioindicators due to their wide distribution, life span, trophic position, metabolic rate and food intake. However, few studies have analysed Hg residues in bats, and to the best of our knowledge, no studies have been made in the Iberian Peninsula. The main aim of this study was to undertake the first ever assessment of Hg exposure in Schreiber's bent-winged bats inhabiting a natural cave in the southeast of Spain. The findings suggest that Schreiber's bent-winged bats in the sampling area are chronically exposed to low levels of Hg. The Hg concentrations found in different tissues (fur, kidney, liver, muscle and brain) were below the threshold levels associated with toxic effects in mammals. Non-gestating females showed Hg concentrations in the brain and muscle that doubled those found in gestating females. This could be due to Hg mobilization from the mother to the foetus in gestating females, although other factors could contribute to explain this result such as variations in hunting areas and the insect-prey consumed and/or different energetic needs and average food consumption during the breeding season. Hg levels were 1.7 times higher, although not significant, in foetus' brains than in the maternal brains, and Hg concentration in foetus' brain was significantly correlated with levels in the corresponding mothers' kidney. These results suggest that there could be an active mother-to-foetus transfer of Hg in bats, which would be of special relevance in a scenario of higher Hg exposure than that found in this study. However, further research is needed to support this view due to the limited number of samples analysed. Given the scarce ecotoxicological data available for bats and their protected status, we encourage further opportunistic studies using carcasses found in the field, the validation of non-destructive samples such as fur and guano for Hg monitoring, and new modelling approaches that will increase the data needed for proper ecological risk assessment in bat populations.

Factors influencing territorial occupancy and reproductive success in a Eurasian Eagle-owl (Bubo bubo) population.

Modelling territorial occupancy and reproductive success is a key issue for better understanding the population dynamics of territorial species. This study aimed to investigate these ecological processes in a Eurasian Eagle-owl (Bubo bubo) population in south-eastern Spain during a seven-year period. A multi-season, multi-state modelling approach was followed to estimate the probabilities of occupancy and reproductive success in relation to previous state, time and habitat covariates, and accounting for imperfect detection. The best estimated models showed past breeding success in the territories to be the most important factor determining a high probability of reoccupation and reproductive success in the following year. In addition, alternative occupancy models suggested the positive influence of crops on the probability of territory occupation. By contrast, the best reproductive model revealed strong interannual variations in the rates of breeding success, which may be related to changes in the abundance of the European Rabbit, the main prey of the Eurasian Eagle-owl. Our models also estimated the probabilities of detecting the presence of owls in a given territory and the probability of detecting evidence of successful reproduction. Estimated detection probabilities were high throughout the breeding season, decreasing in time for unsuccessful breeders but increasing for successful breeders. The probability of detecting reproductive success increased with time, being close to one in the last survey. These results suggest that reproduction failure in the early stages of the breeding season is a determinant factor in the probability of detecting occupancy and reproductive success.

Integrating occurrence and detectability patterns based on interview data: a case study for threatened mammals in Equatorial Guinea.

Occurrence models that account for imperfect detection of species are increasingly used for estimating geographical range, for determining species-landscape relations and to prioritize conservation actions worldwide. In 2010, we conducted a large-scale survey in Río Muni, the mainland territory of Equatorial Guinea, which aimed to estimate the probabilities of occurrence and detection of threatened mammals based on environmental covariates, and to identify priority areas for conservation. Interviews with hunters were designed to record presence/absence data of seven species (golden cat, leopard, forest elephant, forest buffalo, western gorilla, chimpanzee and mandrill) in 225 sites throughout the region. We fitted single season occupancy models and recently developed models which also include false positive errors (i.e. species detected in places where it actually does not occur), which should provide more accurate estimates for most species, which are susceptible to mis-identification. Golden cat and leopard had the lowest occurrence rates in the region, whereas primates had the highest rates. All species, except gorilla, were affected negatively by human settlements. The southern half of Río Muni showed the highest occurrence of the species studied, and conservation strategies for ensuring the persistence of threatened mammals should be focused on this area.

Lifespan analyses of forest raptor nests: patterns of creation, persistence and reuse.

Structural elements for breeding such as nests are key resources for the conservation of bird populations. This is especially true when structural elements require a specific and restricted habitat, or if the construction of nests is costly in time and energy. The availability of nesting-platforms is influenced by nest creation and persistence. In a Mediterranean forest in southeastern Spain, nesting-platforms are the only structural element for three forest-dwelling raptor species: booted eagle Aquila pennata, common buzzard Buteo buteo and northern goshawk Accipiter gentilis. From 1998 to 2013, we tracked the fate of 157 nesting-platforms built and reused by these species with the aim of determining the rates of creation and destruction of nesting-platforms, estimating nest persistence by applying two survival analyses, describing the pattern of nest reuse and testing the effects of nest use on breeding success. Nest creation and destruction rates were low (0.14 and 0.05, respectively). Using Kaplan Meier survival estimates and Cox proportional-hazards regression models we found that median nest longevity was 12 years and that this was not significantly affected by nest characteristics, nest-tree dimensions, nest-builder species, or frequency of use of the platform. We also estimated a transition matrix, considering the different stages of nest occupation (vacant or occupied by one of the focal species), to obtain the fundamental matrix and the average life expectancies of nests, which varied from 17.9 to 19.7 years. Eighty six percent of nests were used in at least one breeding attempt, 67.5% were reused and 17.8% were successively occupied by at least two of the study species. The frequency of nest use had no significant effects on the breeding success of any species. We conclude that nesting-platforms constitute an important resource for forest raptors and that their longevity is sufficiently high to allow their reuse in multiple breeding attempts.

Factors that influence mercury concentrations in nestling Eagle Owls (Bubo bubo).

Mercury (Hg) is a global pollutant that bioaccumulates and biomagnifies in food chains, and is associated with adverse effects in both humans and wildlife. The Hg levels detected in blood obtained from Eagle Owl (Bubo bubo) chicks in Southeast Spain (Murcia) can be considered low (mean Hg concentration in blood from 2006 to 2012 was 36.83 ± 145.58 µg/l wet weigh, n=600), and it is therefore unlikely that Hg pollution can negatively affect their breeding. Positive correlation (r=0.339, p<0.001, n=229) was found to exist between the Hg concentrations in the blood and back feathers of the chicks. We provide a regression equation that could be helpful to estimate blood Hg levels when analyzing Hg concentrations in back feathers. Blood Hg concentrations in Eagle Owls have shown positive correlations with Hg levels in rabbit muscles, more evident in nests from the Northern area (r=0.600, p=0.014, n=16), where rabbits are the main prey of Eagle Owls. The best Linear Mixed Model to explain variations in blood Hg concentrations in nestling Eagle Owls includes year and location within the mining area as variables. The variable year is assigned the largest value of relative importance, followed by the location in the ancient mining sites and then the zone. Rainfalls may have an effect on the temporal differences in the blood Hg concentrations of nestling Eagle Owls. Although the studied region is not considered Hg polluted, the Hg levels were higher in Eagle Owls and European Rabbits from areas within the ancient mining sites as compared to those in the entire region. This result shows that spatial differences in Hg concentrations in Eagle Owls may be affected by local contamination, and that the role of diet composition may be less significant.