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West Nile virus (WNV) is a globally widespread arthropod-borne virus that poses a significant public health concern. Mosquitoes transmit the virus in an enzootic cycle among birds, which act as reservoirs. Climate plays a crucial role in these outbreaks as mosquitoes are highly influenced by climatic conditions, and bird migrations are also affected by weather patterns. Consequently, changes in climate can potentially impact the occurrence of WNV outbreaks. We used biogeographic modelling based on machine learning algorithms and fuzzy logic to analyse and evaluate separately the risk of WNV outbreaks in two different biogeographic regions, the Afrotropical and the Western Palaearctic region. By employing fuzzy logic tools, we constructed a comprehensive risk model that integrates the Afro-Palaearctic system as a unified operational unit for WNV spread. This innovative approach recognizes the Afro-Palaearctic region as a pathogeographic system, characterized by biannual connections facilitated by billions of migratory bird reservoirs carrying the disease. Subsequently, we forecasted the effects of different climate change scenarios on the spread of WNV in the Afro-Palaearctic system for the years 2040 and 2070. Our findings revealed an increasing epidemic and epizootic risk south of the Sahara. However, the area where an upsurge in risk was forecasted the most lies within Europe, with the anticipation of risk expansion into regions presently situated beyond the virus' distribution range, including central and northern Europe. Gaining insight into the risk within the Afro-Palaearctic system is crucial for establishing coordinated and international One Health surveillance efforts. This becomes particularly relevant in the face of ongoing climate change, which disrupts the ecological equilibrium among vectors, reservoirs, and human populations. We show that the application of biogeographical tools to assess risk of infectious disease, i.e. pathogeography, is a promising

West Nile virus (WNV) is a globally significant vector-borne disease that is primarily transmitted between birds and mosquitoes. Recently, there has been an increase in WNV in southern Europe, with new cases reported in more northern regions. Bird migration plays a crucial role in the introduction of WNV in distant areas. To better understand and address this complex issue, we adopted a One Health approach, integrating clinical, zoological, and ecological data. We analyzed the role of migratory birds in the Palaearctic-African region in the spread of WNV across Africa and Europe. We categorized bird species into breeding and wintering chorotypes based on their distribution during the breeding season in the Western Palaearctic and the wintering season in the Afrotropical region, respectively. By linking these chorotypes to the occurrence of WNV outbreaks in both continents throughout the annual bird migration cycle, we investigated the relationship between migratory patterns and virus spread. We demonstrate that WNV-risk areas are interconnected through the migration of birds. We identified a total of 61 species that potentially contribute to the intercontinental spread of the virus or its variants, as well as pinpointed high-risk areas for future outbreaks. This interdisciplinary approach, which considers the interconnectedness of animals, humans, and ecosystems, represents a pioneering effort to establish connections between zoonotic diseases across continents. The findings of our study can aid in anticipating the arrival of new WNV strains and predicting the occurrence of other re-emerging diseases. By incorporating various disciplines, we can enhance our understanding of these complex dynamics and provide valuable insights for proactive and comprehensive disease management strategies.

A classic issue in ecology is to understand how similar species coexist in a given area (i.e., sympatry). The situation of dolphins in the Western Mediterranean Sea may represent a special case of sympatry in that three similar species (the short-beaked common dolphin [Delphinus delphis], the striped dolphin [Stenella coeruleoalba], and the bottlenose dolphin [Tursiops truncatus) are under strong human impacts in the same area. From the viewpoint of ecology and conservation biology, it is challenging to determine how these three dolphin species live together and avoid competitive exclusion in the setting of such impacts. The Spanish Institute of Oceanography has a dataset of dolphin species opportunistic sightings. Using these data, we constructed three binary variables, comprising the sighting of one species versus the sighting of either of the other two species. We obtained three significant probability models after performing logistic regression of these binary variables on a set of spatio-temporal explanatory variables. We analysed these models from the perspective of fuzzy set theory by applying the favourability function to the probability models, fuzzy operations overlap, and entropy. The results show that common dolphins are differentially favoured in the eastern part of the study area and far away from main shipping routes. The striped dolphin was differentially favoured in the western part of the study area, above deep waters, near main shipping routes, and in summer and spring. Finally, bottlenose dolphins were differentially favoured in the mid-western part of the study area, in winter, and over shallow waters.

Understanding the biogeography of species in space and time is essential for the development of evidence-based conservation and management plans. In this paper we propose a biogeographical spatial modelling approach based on the favourability function, and developed under a fuzzy logic framework, to unravel the historical biogeography of the two European wild rabbit subspecies, Oryctolagus cuniculus algirus (Oca) and Oryctolagus cuniculus cuniculus (Occ), in the Iberian Peninsula (IP). We first reviewed published and unpublished information (PhD theses, scientific papers, technical reports, etc.) on the occurrence of each rabbit subspecies throughout the IP. We compiled data from 201 Iberian rabbit populations and from genetic information of 4348 rabbits that was used to identify subspecies. Only populations in which all rabbits surveyed belonged to one subspecies were considered in the modelling procedure. We modelled rabbit subspecies' distribution separately for populations in which nuclear and mitochondrial DNA sequences were available. We employed a trend surface analysis developed by logistic regressions, which applied the favourability function and fuzzy logic operations. Using our approach we indentify the expansion cores from which both rabbit subspecies would have expanded after isolation during the last glaciations. Furthermore, we reveal the possible existence of a competitive exclusion zone between both rabbit subspecies that may have prevented their further expansion. Finally, our study shows that the Oca subspecies is distributed in north-western areas previously attributed to Occ. This assessment of the actual and historical distribution of each rabbit subspecies may allow more specific conservation interventions, as the two subspecies are not just genetically distinct but also ecologically and behaviourally different. Our methodological approach could be useful in unravelling the historical biogeography of other lesser-known species.

In the Northern Alboran Sea, artisanal small-scale fisheries using trammel nets suffer economic losses, and local fishermen see their way-of-life endangered, due to interactions with wildlife species such as alien species and dolphins. On the one hand, the alien seaweed Rugulopteryx okamurae, which was first recorded in the Alboran Sea in 2015, has undergone an intensive expansion in the sub-region, monopolizing the available seabed, causing radical changes in the underwater seascape and clogging the trammel nets. On the other hand, the damage caused to the fishing nets by dolphin fish predation is an ancient problem worldwide, but it is intensifying in the last years. The main objective of this study is to understand the main environmental and technical conditions that favor damages of fishing trammel nets in the Alboran Sea, which entails an important loss of catchability, due to (i) the clogging of the artisanal fishing trammel nets by invasive seaweed, and (ii) the breaking of the nets by dolphin predation. Through close monitoring of fishermen in port, we obtained direct information of 548 sets. Our results indicate that approximately 30% of trammel sets suffered a damage due to unwanted interaction with alien seaweeds and dolphins. As seaweeds invasion is a global problem while dolphin-fishing gear interaction is more local, we concluded that only a large-scale management of exotic algae, together with the involvement of local fishermen, could solve the economic problems of this activity.

In biogeography the competitive exclusion principle (CEP) has been confirmed in some cases but not in others. This has fueled an unresolved debate between those advocating niche theory or the neutral theory in biodiversity and biogeography. We suggest that this situation mainly arises from the use of crisp logic, where the CEP is defined as either completely true or false. We propose the application of the fuzzy concepts of favorability (the degree to which environmental conditions are propitious for the occurrence of individual species) and favorableness (the degree to which environmental conditions are simultaneously favorable for competing species) to operationalize a fuzzy version of the CEP. Favorability was obtained by performing species distribution models applying favorability functions, while favorableness was derived from the application of the fuzzy intersection between the favorability for competing species. Then we plotted individual favorability values along the gradient of favorableness. Two potentially competing species would coexist in high-favorableness locations, as the demands of both species would be well fulfilled. In locations of low favorableness, the result would be either autecological exclusion of both species or autecological segregation, as abiotic conditions are unfavorable for at least one of the species. Competitive exclusion would occur at the intermediate stretch of the favorableness gradient, as the conditions would be good enough for persistence of each species separately but not enough for permanent coexistence. According to this theoretical framework, the observed probability that a location belongs to the intermediate favorableness area given that the two species co-occur in this location should be lower than expected according to the environmental probability models for the two species. We tested this prediction on published data about the distribution of pairs of native and introduced deer species in Great Britain, using a Ba

In biogeography the competitive exclusion principle (CEP) has been confirmed in some cases but not in others. This has fueled an unresolved debate between those advocating niche theory or the neutral theory in biodiversity and biogeography. We suggest that this situation mainly arises from the use of crisp logic, where the CEP is defined as either completely true or false. We propose the application of the fuzzy concepts of favorability (the degree to which environmental conditions are propitious for the occurrence of individual species) and favorableness (the degree to which environmental conditions are simultaneously favorable for competing species) to operationalize a fuzzy version of the CEP. Favorability was obtained by performing species distribution models applying favorability functions, while favorableness was derived from the application of the fuzzy intersection between the favorability for competing species. Then we plotted individual favorability values along the gradient of favorableness. Two potentially competing species would coexist in high-favorableness locations, as the demands of both species would be well fulfilled. In locations of low favorableness, the result would be either autecological exclusion of both species or autecological segregation, as abiotic conditions are unfavorable for at least one of the species. Competitive exclusion would occur at the intermediate stretch of the favorableness gradient, as the conditions would be good enough for persistence of each species separately but not enough for permanent coexistence. According to this theoretical framework, the observed probability that a location belongs to the intermediate favorableness area given that the two species co-occur in this location should be lower than expected according to the environmental probability models for the two species. We tested this prediction on published data about the distribution of pairs of native and introduced deer species in Great Britain, using a Ba

Dengue is a viral disease transmitted by mosquitoes. The rapid spread of dengue could lead to a global pandemic, and so the geographical extent of this spread needs to be assessed and predicted. There are also reasons to suggest that transmission of dengue from non-human primates in tropical forest cycles is being underestimated. We investigate the fine-scale geographic changes in transmission risk since the late 20th century, and take into account for the first time the potential role that primate biogeography and sylvatic vectors play in increasing the disease transmission risk. We apply a biogeographic framework to the most recent global dataset of dengue cases. Temporally stratified models describing favorable areas for vector presence and for disease transmission are combined. Our models were validated for predictive capacity, and point to a significant broadening of vector presence in tropical and non-tropical areas globally. We show that dengue transmission is likely to spread to affected areas in China, Papua New Guinea, Australia, USA, Colombia, Venezuela, Madagascar, as well as to cities in Europe and Japan. These models also suggest that dengue transmission is likely to spread to regions where there are presently no or very few reports of occurrence. According to our results, sylvatic dengue cycles account for a small percentage of the global extent of the human case record, but could be increasing in relevance in Asia, Africa, and South America. The spatial distribution of factors favoring transmission risk in different regions of the world allows for distinct management strategies to be prepared.

Dengue is a viral disease transmitted by mosquitoes. The rapid spread of dengue could lead to a global pandemic, and so the geographical extent of this spread needs to be assessed and predicted. There are also reasons to suggest that transmission of dengue from non-human primates in tropical forest cycles is being underestimated. We investigate the fine-scale geographic changes in transmission risk since the late 20th century, and take into account for the first time the potential role that primate biogeography and sylvatic vectors play in increasing the disease transmission risk. We apply a biogeographic framework to the most recent global dataset of dengue cases. Temporally stratified models describing favorable areas for vector presence and for disease transmission are combined. Our models were validated for predictive capacity, and point to a significant broadening of vector presence in tropical and non-tropical areas globally. We show that dengue transmission is likely to spread to affected areas in China, Papua New Guinea, Australia, USA, Colombia, Venezuela, Madagascar, as well as to cities in Europe and Japan. These models also suggest that dengue transmission is likely to spread to regions where there are presently no or very few reports of occurrence. According to our results, sylvatic dengue cycles account for a small percentage of the global extent of the human case record, but could be increasing in relevance in Asia, Africa, and South America. The spatial distribution of factors favoring transmission risk in different regions of the world allows for distinct management strategies to be prepared.

La Alcarria is one of the most significant regions for honey production in Spain. Times series data of the annual Alcarria honey production from 2000 to 2013 revealed that the total production was highly variable ranging from ca. 60,000 kg in 2000 to almost three times as much in 2011, but without a remarkable trend through time. We hypothesized that these local fluctuations in honey production may be explained by broad-scale climatic patterns. Based on the multiple linear models, we found a significant positive relationship between honey production as a function of the overall means of the North Atlantic Oscillation (NAO) index and precipitation in January, which accounts for 80% of the variability in the Alcarria honey production. We propose that the effect of a negative phase of the NAO and precipitation in January could predict the Alcarria honey production a few months in advance.

The European rabbit Oryctolagus cuniculus is native to the lberian Península (IP), where the two known rabbit subspecies occur. Previous studies have suggested that O.c. cuniculus distribution extends towards the north-east of the IP, while O.c. algirus is restricted to south-westem regions; a transition area crosses the IP from the north-west to the south-east. Here, we revisited rabbit subspecies distribution to define with greater precision historical distribution areas, identifying the potential expansion cores and evaluating the spatial interaction between both subspecies expansion waves. We used different diagnostic loci located on mitochondrial and nuclear DNA to classify in any of the subspecies 4500 rabbits from 197 populations distributed throughout the IP. Using populations where all the rabbits were either O.c. cuniculus or O.c. algirus, we determined the spatial structure of the two molecules independently, by using a trend surface analysis based on favourability functions. We then combined the different spatial structures using fuzzy logic operations. Two areas in the Guadalquivir and Ebro valleys were identified as respective expansion cores. In comparison to previous studies, we found an increase in the distribution range of O.c. algirus to the northwest anda shift of the transition zone eastwards in its northern area. Our results suggest that competitive exclusion may ha ve a greater role than expected in regulating the expansion of both subspecies. The spatial structure of these genes reflects, and thus may be used to unravel, the historical biogeography of these subspecies, provided that we possess appropriate analytical tools, such as fuzzy logic.

Currently, there are naturalized populations of feral pigeon (Columba livia var. domestica) worldwide with an estimated global population between 165 and 330 million individuals. Feral pigeons are adapted to urban environments, creating problems related to human health, damage to architectural structures, unpleasant soiling, and foul odors. Thus, local governments usually implement measures to reduce feral pigeon populations in cities to a suitable size. We monitored a feral pigeon trapping campaign to investigate the use of cage traps as control method in residential zones. We analyzed efficiency (daily capture rate), effectiveness (defined as the number of feral pigeons captured relative to the total number of potential pigeon captures), animal welfare, selectivity (the proportion of captured feral pigeons in relation to the total number of captured animals), and the effect on feral pigeon abundance. All procedures were conducted in compliance with current European and Spanish legislation. Overall efficiency was 0.25 feral pigeons/trap-day, effectiveness was 22.5%, and selectivity was 99.7%. Over the study period, there was a decreasing trend in abundances. A significant decrease was found in the relative abundances of feral pigeons after 2 months of extractions, and population abundances remained low up to the end of the extraction period. The use of cage traps was a useful and effective method to reduce local feral pigeon populations in isolated urban areas with low building density in which the feral pigeon population has not yet reached high densities.

The evaluation of protected area networks on their capacity to preserve species distributions is a key topic in conservation biology. There are different types of protected areas, with National Parks those with highest level of protection. National Parks can be declared attending to many ecological features that include the presence of certain animal species. Here, we selected 37 vertebrate species that were highlighted as having relevant natural value for at least one of the 10 National Parks of mainland Spain. We modelled species distributions with the favourability function, and applied the Insecurity Index to detect the degree of protection of favourable areas for each species. Two metrics of Insecurity Index were defined for each species: the Insecurity Index in each of the cells, and the Overall Insecurity Index of a species. The former allows the identification of insecure areas for each species that can be used to establish spatial conservation priorities. The latter gives a value of Insecurity for each species, which we used to calculate the Representativeness of favourable areas for the species in the network. As expected, due to the limited extension of the National Park network, all species have high values of Insecurity; i.e., just a narrow proportion of their favourable areas are covered by a National Park. However, the majority of species favourable areas are well represented in the network, i.e., the percentage of favourable areas covered by the National Park network is higher than the percentage of mainland Spain covered by the network (result also supported by a randomization approach). Even if a reserve network only covers a low percentage of a country, the Overall Insecurity Index allows an objective assessment of its capacity to represent species. Beyond the results presented here, the Insecurity Index has the potential to be extrapolated to other areas and to cover a wide range of species.

Distribution of 37 vertebrate species and values of environmental variables in each 10 km x 10 km UTM cell of mainland Spain.

Chondrichthyes, which include Elasmobranchii (sharks and batoids) and Holocephali (chimaeras), are a relatively small group in the Mediterranean Sea (89 species) playing a key role in the ecosystems where they are found. At present, many species of this group are threatened as a result of anthropogenic effects, including fishing activity. Knowledge of the spatial distribution of these species is of great importance to understand their ecological role and for the efficient management of their populations, particularly if affected by fisheries. This study aims to analyze the spatial patterns of the distribution of Chondrichthyes species richness in the Mediterranean Sea. Information provided by the studied countries was used to model geographical and ecological variables affecting the Chondrichthyes species richness. The species were distributed in 16 Operational Geographical Units (OGUs), derived from the Geographical Sub-Areas (GSA) adopted by the General Fisheries Commission of the Mediterranean Sea (GFCM). Regression analyses with the species richness as a target variable were adjusted with a set of environmental and geographical variables, being the model that links richness of Chondrichthyes species with distance to the Strait of Gibraltar and number of taxonomic families of bony fishes the one that best explains it. This suggests that both historical and ecological factors affect the current distribution of Chondrichthyes within the Mediterranean Sea.

Species distributions are typically represented by records of their observed occurrence at a given spatial and temporal scale. Such records are inevitably incomplete and contingent on the spatial–temporal circumstances under which the observations were made. Moreover, organisms may respond differently to similar environmental conditions at different places or moments, so their distribution is, in principle, not completely predictable. We argue that this uncertainty exists, and warrants considering species distributions as analogous to coherent quantum objects, whose distributions are better described by a wavefunction rather than by a set of locations. We use this to extend the existing concept of “dark diversity”, which incorporates into biodiversity metrics those species that could, but which have not yet been observed to, inhabit a region—thereby developing the idea of “potential biodiversity”. We show how conceptualizing species’ distributions in this way could help overcome important weaknesses in current biodiversity metrics, both in theory and by using a worked case study of mammal distributions in Spain over the last decade. We propose that considerable theoretical advances could eventually be gained through interdisciplinary collaboration between biogeographers and quantum physicists. [Biogeography; favorability; physics; predictability; probability; species occurrence; uncertainty; wavefunction., Species distributions are typically represented by records of their observed occurrence at a given spatial and temporal scale. Such records are inevitably incomplete and contingent on the spatial–temporal circumstances under which the observations were made. Moreover, organisms may respond differently to similar environmental conditions at different places or moments, so their distribution is, in principle, not completely predictable. We argue that this uncertainty exists, and warrants considering species distributions as analogous to coherent quantum objects, whose distributions are better described by a wavefunction rather than by a set of locations. We use this to extend the existing concept of “dark diversity”, which incorporates into biodiversity metrics those species that could, but which have not yet been observed to, inhabit a region—thereby developing the idea of “potential biodiversity”. We show how conceptualizing species’ distributions in this way could help overcome important weaknesses in current biodiversity metrics, both in theory and by using a worked case study of mammal distributions in Spain over the last decade. We propose that considerable theoretical advances could eventually be gained through interdisciplinary collaboration between biogeographers and quantum physicists. [Biogeography; favorability; physics; predictability; probability; species occurrence; uncertainty; wavefunction.

An accurate estimation of the expected consequences of climate change requires the proper quantification of the effect of climate on current species distributions. Several interrelated sources of uncertainty may affect the likelihood of species distribution models (SDMs) to determine the relative importance of climate. Our aim was to assess the relationship between the influence of geographical background (GB) delimitation and that of subtracting the non-climate effects from the weight of climatic predictors to estimate the combined influence of these two factors on predictions in climate change scenarios. The distribution of 40 endemic mammals in Western Europe have been modeled by (i) using the whole territory of Western Europe as the GB and also specifically delimiting the GB with a geographical criterion and (ii) considering climatic predictors in addition to other non-climatic variables in order to extract the pure effect of climate. The models were used to quantify species’ sensitivity to new climate scenarios. Results showed discrepancies among the analytical approaches. Changes in distribution obtained by considering the pure effect of climate were lower than those obtained by considering the apparent effect, and GB-delimited models yielded higher changes than those trained in Western Europe. We evidence that climate weighting and GB delimitation have dramatic influences on the projections of models when transferred to new scenarios. We emphasize that scientific studies and derived adaptation policies based on SDMs without an explicit consideration of the GB and the weighting of the climate-related variables may be misleading and in need of revision.

Species distribution models (SDMs) are basic tools in ecology, biogeography, and biodiversity. The usefulness of SDMs has expanded beyond the realm of ecological sciences, and their application in other research areas is currently frequent, e.g., spatial epidemiology. In any research area, the principal interest in these models resides in their capacity to predict species response in new scenarios, i.e., the models' transferability. Although the transferability of SDMs has been the subject of interest for many years, only in the 2000s did this topic gain particular attention. This article reviews the concept of the transferability of SDMs to new spatial scenarios, temporal periods, and (or) spatial resolutions, along with the potential constraints of the model's transferability, and more specifically: (i) the type of predictors and multicollinearity, (ii) the model complexity, and (iii) the species' intrinsic traits. Finally, we describe a practicable analytical protocol to be assessed before transferring a model to a new scenario. This protocol is based on three fundamental pillars: the environmental equilibrium of the species with the environment, the environmental similarity between the new scenario, and the areas used to model parametrisation and the correlation structure among predictors.

Leatherback sea turtle (Dermochelys coriacea, Vandelli, 1761)) is a large migratory species reproducing in Atlantic areas and abundant in the Mediterranean Sea, where nesting beaches do not exist. Leatherback stranding occasionally occurs in the Andalusia coasts (183 turtles during 1990- 2011 period). This communication study a possible inter and intra annual temporal patterns in leatherback strandings in the area and analyze the effects of atmospheric oscillations on their arrivals in the Mediterranean. The North Atlantic Oscillation (NAO) plays an essential role in climate fluctuations affecting the North Atlantic and associated biological components. Recent studies showed that the Arctic Oscillation (AO) is highly correlated with NAO, explaining more accurately variations found in ecological time series. The summer months concentrate significatively a higher number of leatherback turtle strandings. Moreover, the results show a significant negative correlation between the stranding frequency and the average rate of the AO.

El “principio de exclusión competitiva” se ha visto confirmado en algunas situaciones pero no en otras. Sostenemos que esta situación equívoca surge del uso de la lógica clásica, que define las proposiciones como totalmente ciertas o completamente falsas. La alternativa es tomar una perspectiva desde la lógica difusa mediante el empleo del concepto asociado de favorabilidad. Favorabilidad se define como un conjunto borroso de condiciones ambientales que son favorables para la presencia de una especie a lo largo de un gradiente que va desde muy favorable a muy desfavorable. La favorabilidad compartida se define como el grado en que las condiciones autoecológicas son simultáneamente favorables para dos (o más) especies que interactúan. Desde esta perspectiva, se describe un nuevo paradigma para explicar la coexistencia o la exclusión competitiva de especies. Al contrastar favorabilidad individual y favorabilidad compartida es posible determinar y comprobar cómo la segregación autoecológica, la exclusión competitiva y la coexistencia actúan diferencialmente a lo largo de un gradiente de favorabilidad compartida entre especies competidoras. Se presentan algunos ejemplos de mamíferos (cérvidos y liebres) que aportan un apoyo empírico a este nuevo enfoque lógico.

[Aim]: A striking case of mitochondrial DNA (mtDNA) introgression occurs in the Iberian Peninsula: the mtDNA of Lepus timidus, a species that inhabited this territory during the Last Glacial Maximum (LGM), is almost fixed in the Iberian range of Lepus europaeus, is very frequent in the northern half of Iberia within Lepus granatensis, and is fixed in Lepus castroviejoi. Here, we used a transdisciplinary approach to understand the ecological drivers of the distribution of this introgressed mtDNA in the Iberian hare species. [Location]: Europe, with an emphasis on the Iberian Peninsula. [Methods]: First, we modelled the climatic niche of L. timidus to predict the favourability for the species during the LGM. Second, we gathered data regarding L. timidus mtDNA introgression for 1137 individuals (139 populations) from the Iberian Peninsula. We tested whether the climatic favourability for L. timidus during the LGM reflects the frequency of the introgressed mtDNA in the carrier hare populations. [Results]: The climatic favourability for the LGM is positively related to the presence of individuals with L. timidus mtDNA, and discriminates better than by chance between individuals with and without this lineage. At the population level, the favourability values relate to the observed frequencies of mtDNA introgression and discriminate between populations with and without L. timidus mtDNA. [Main conclusions]: The results suggest that the geographical distribution of L. timidus mtDNA introgression is linked to L. timidus distribution in Iberia during the LGM. Our study helps to improve our understanding of the role of climate in the reticulated evolutionary history of hares, and is a first step towards understanding the ecological impact of mtDNA introgression on Iberian hares.

Assessing the response of exploited marine populations in a context of rapid climate change: the case of blackspot seabream from the Strait of Gibraltar.— There is a growing concern over the decline of fisheries and the possibility of the decline becoming worse due to climate change. Studies on small–scale fisheries could help to improve our understanding of the effect of climate on the ecology of exploited stocks. The Strait of Gibraltar is an important fishery ground for artisanal fleets. In this area, b lackspot seabream ( Pagellus bogaraveo ) is the main species targeted by artisanal fisheries in view of its relevance in landed weight. The aims of this study were to explore the possible effects of two atmospheric oscillations, the North Atlantic Oscillation (NAO) and the Arctic Oscillation (AO), on the capture of blackspot seabream in the Strait of Gibraltar, to determine their association with oceanographic conditions, and to improve our knowledge about the possible effects of climate change on fisheries ecology so that fishery management can be improved. We used two types of data from different sources: (i) landings per unit of effort reported from a second working group between Morocco and Spain on Pagellus bogaraveo in the Gibraltar Strait area, for the period 1983–2011 , and (ii) the recorded blackspot seabream landings obtained from the annual fisheries statistics published by the Junta de Andalucía (Andalusian Regional Government). Our results indicate that the long–term landing of blackspot seabream in the Strait of Gibraltar is closely associated with atmospheric oscillations. Thus, prolonged periods of positive trends in the NAO and AO could favour high fishery yields. In contrast, negative trends in NAO and AO could drastically reduce yield.

Álvarez de Quevedo et al. (2013; Mar Ecol Prog Ser 489:225–234) tagged 26 by- caught loggerhead sea turtles, released them with the hook and 40 cm of line, tracked them for 90 d, and concluded that the Spanish longline fleet in the southwestern Mediterranean Sea has an annual by-catch of 10 656 loggerhead turtles, resulting in 3421 to 4028 turtle deaths. We consider the methodology and results in Álvarez de Quevedo et al. (2013) to be correct overall, but based on the loggerhead CPUE (per 1000 hooks) in the 2006−2007 period, we estimate that only 6060 ind. (SE: ± 40%) are captured annually; this would lead to a lower estimate of turtle mortality of 1945 to 2290 ind. (± 40%).

We assessed the vulnerability of the native Mediterranean pond turtle to encroachment by the invasive red-eared slider in southern Spain. We first obtained an ecogeographical favourability model for the Mediterranean pond turtle. We then modelled the presence/absence of the red-eared slider in the Mediterranean pond turtle range and obtained an encroachment favourability model. We also obtained a favourability model for the red-eared slider using the ecogeographical favourability for the Mediterranean pond turtle as a predictor. When favourability for the Mediterranean pond turtle was high, favourability for the red-eared slider was low, suggesting that in these areas the Mediterranean pond turtle may resist encroachment by the red-eared slider. We also calculated favourability overlap between the two species, which is their simultaneous favourability. Grids with low overlap had higher favourability values for the Mediterranean pond turtle and, consequently, were of lesser conservation concern. A few grids had high values for both species, being potentially suitable for coexistence. Grids with intermediate overlap had similar intermediate favourability values for both species and were therefore areas where the Mediterranean pond turtle was more vulnerable to encroachment by the red-eared slider. We mapped the favourability overlap to provide a map of vulnerability of the Mediterranean pond turtle to encroachment by the red-eared slider.

[Aim]: Lepus castroviejoi and Lepus corsicanus are sister species with allopatric distributions that share extensive phenotypic and genetic variation. Under the framework of niche conservatism, we assessed whether these species have similar ecological niches, which could provide insights into their mode of divergence, conservation, and taxonomic status. [Location]: The distribution range of L. castroviejoi in the northern Iberian Peninsula, and that of L. corsicanus in mainland Italy and Sicily. [Methods]: We developed spatially explicit ecological models to characterize the niches of the two species by modelling them separately and together. Individual models were transferred to the territory of the sister species to explore their niche relationships. Predictions were assessed for discrimination and calibration in a cross-assessment procedure. [Results]: The model trained with L. castroviejoi was not able to predict the range of L. corsicanus, whereas the model trained with L. corsicanus was able to discriminate the L. castroviejoi distribution better than by chance alone (AUC = 0.814), although the reliability of the predictions was limited. The model trained with L. corsicanus in Italy's mainland (excluding the range in Sicily), however, discriminated L. castroviejoi presences/absences (AUC = 0.788) and accurately predicted its probability of occurrence. Furthermore, a well-calibrated model, which was able to discriminate the species distributions (L. castroviejoi, AUC = 0.828; L. corsicanus, AUC = 0.956), was obtained when the species were considered together. [Main conclusions]: Our results suggest that L. castroviejoi and L. corsicanus share extensive niche properties, which reinforces their possible conspecific status. The ecological niche of their ancestor may have resembled the present occupied niche of L. corsicanus in mainland Italy, given that this model was able to accurately predict the distribution range of both species. Finally, ecological evidence s

Analysis of the effect of atmospheric oscillations on physical condition of pre–reproductive bluefin tuna from the Strait of Gibraltar .— The aim of this study was to explore the possible effects of atmospheric oscillations: North Atlantic Oscillation (NAO) and Arctic Oscillation (AO), on the physical condition of bluefin tuna ( Thunnus thynnus ). We estimated a fitness ratio from 3,501 pairs of length–weight data based on bluefin tuna caught in bait–boat fisheries before the spawning season (January, February and March), for each length class and year. In order to obtain a single fitness ratio (K–mean) per year we determined the average for all length classes. We also evaluated Le Cren’s condition index (K LC ). We observed significant positive correlations between the atmospheric oscillations and both physical condition indexes. In the case of K–mean, the AO explained 75% of the observed variability. Regarding K LC , the NAO explained approximately 73% of the observed variability, while the AO explained 70% of the observed variability. The increase in physical conditions of bluefin tuna in association with positive atmospheric oscillations could be mediated by the increase in the prevalence of strong trade winds. We concluded that the increase in the prevalence of strong westerly winds, mediated by a positive AO or NAO, favours the trip from the Atlantic to the Mediterranean by reducing energy costs due to migration and by increasing the supply of nutrients at the surface by the mixing of deep water and surface water in local areas such as the Strait of Gibraltar.

Fuzzy logic is a form of many-valued logic whose variables have a truth value that varies in degree. Spatial favourability for species occurrence may be considered a fuzzy concept, as historical, geographical, human, and environmental conditions make locations more or less favourable for the occurrence of particular species. The favourability function was conceptually conceived to define spatial favourability in a fuzzy gradient from 0 to 1, so facilitating the application of fuzzy logic to spatial modelling. Favourability values derived from the favourability function have the same meaning and the same mathematical value regardless the prevalence of the species, so enabling direct comparison of models built for different species and their combination using fuzzy logic operators. This characteristics make the favourability function particularly useful in the spatial modelling of ecologically interacting species. In particular, the fuzzy intersection of favourability for different species is useful to model the biogeographical consequences of different degrees of competition between species. Fuzzy logic operations allow also to combine autoecological and sinecological responses in a way that may account for the existence of parapatric distributions in current and future environments, as exemplified by hare species in Europe. Fuzzy logic may provide biogeographical modellers with the necessary flexibility in concepts and operational tools to deal with a highly unstable and intertwined biogeographical world.

Models that predict species’ distributions or ecological niches, based on inferred relationships between species occurrence and environmental variables, need to be evaluated for their quality or predictive capacity. However, commonly used model evaluation measures, such as the area under the receiver operating characteristic curve (AUC), correct classification rates (including sensitivity and specificity), Cohen’s kappa, and the true skill statistic, are known to be strongly affected by the species’ prevalence or extent of occurrence in the study area: widespread species yield apparently poorer models than species with restricted distributions. Nevertheless, models for widespread species may actually be quite accurate and correlate with independent measures such as species abundance. The problem lies in that these evaluation measures assess only the discrimination capacity of models: they start by converting continuous model predictions into an overly simple binary output (or a series of possible binary outputs, in the case of the AUC), and then evaluate the performance of these binary outputs instead of what the models really predict. We argue that, when the modelling aim does not require binary predictions, discrimination measures should be replaced (or at least complemented) with calibration or reliability measures applied directly to the continuous predictions. We propose a new model evaluation method, the Area Between the Curves (ABC), which can be applied to presence probability models and provides, besides various quantitative metrics, a visually explicit plot showing how much, in which direction and where in the explanatory and predictor space each model departs from observed distribution patterns. It also includes several methods to divide the predicted probability values into bins that allow meaningful comparisons between observations and predictions. We illustrate this procedure on a set of generalised linear models built for European tree distributions (

[Aim]: When faced with dichotomous events, such as the presence or absence of a species, discrimination capacity (the ability to separate the instances of presence from the instances of absence) is usually the only characteristic that is assessed in the evaluation of the performance of predictive models. Although neglected, calibration or reliability (how well the estimated probability of presence represents the observed proportion of presences) is another aspect of the performance of predictive models that provides important information. In this study, we explore how changes in the distribution of the probability of presence make discrimination capacity a context-dependent characteristic of models. For the first time, we explain the implications that ignoring the context dependence of discrimination can have in the interpretation of species distribution models. [Innovation]: In this paper we corroborate that, under a uniform distribution of the estimated probability of presence, a well-calibrated model will not attain high discrimination power and the value of the area under the curve will be 0.83. Under non-uniform distributions of the probability of presence, simulations show that a well-calibrated model can attain a broad range of discrimination values. These results illustrate that discrimination is a context-dependent property, i.e. it gives information about the performance of a certain algorithm in a certain data population. [Main conclusions]: In species distribution modelling, the discrimination capacity of a model is only meaningful for a certain species in a given geographic area and temporal snapshot. This is because the representativeness of the environmental domain changes with the geographical and temporal context, which unavoidably entails changes in the distribution of the probability of presence. Comparative studies that intend to generalize their results only based on the discrimination capacity of models may not be broadly extrapolated. Assessme

We explored the possible effects of the North Atlantic Oscillation (NAO) and Arctic Oscillation (AO) on interannual sea surface temperature (SST) variations in the Alborán Sea, both separately and combined. The probability of observing mean annual SST values higher than average was related to NAO and AO values of the previous year. The effect of NAO on SST was negative, while that of AO was positive. The pure effects of NAO and AO on SST are obscuring each other, due to the positive correlation between them. When decomposing SST, NAO and AO in seasonal values, we found that variation in mean annual SST and mean winter SST was significantly related to the mean autumn NAO of the previous year, while mean summer SST was related to mean autumn AO of the previous year. The one year delay in the effect of the NAO and AO on the SST could be partially related to the amount of accumulated snow, as we found a significant correlation between the total snow in the North Alborán watershed for a year with the annual average SST of the subsequent year. A positive AO implies a colder atmosphere in the Polar Regions, which could favour occasional cold waves over the Iberian Peninsula which, when coupled with precipitations favoured by a negative NAO, may result in snow precipitation. This snow may be accumulated in the high peaks and melt down in spring-summer of the following year, which consequently increases the runoff of freshwater to the sea, which in turn causes a diminution of sea surface salinity and density, and blocks the local upwelling of colder water, resulting in a higher SST.

The Alborán Sea hosts the largest population of common dolphin in the Mediterranean Sea. This species of cetacean has the highest number of recorded strandings on the Andalusian coast. We analyzed the spatial and temporal patterns of 633 short-beaked common dolphins stranded on this coast during the period 1996-2009. The spatial distribution of the strandings differed between the Atlantic and Mediterranean coasts of the Andalusian littoral. Mean body size was 163.3 ± 35.6 cm, although body size was not homogeneously distributed along the coast. These cetaceans tended to be larger on the Atlantic coast than on the rest of the Andalusian coast. Although others factors must be taken into account, the stranding patterns may be explained by the density of short-beaked common dolphins in inshore waters. The number of strandings suggests intense use by this species of the continental shelf along the Andalusian coast, with the possibility of conflict between dolphins and human activities in the Alborán Sea.

Fisheries by-catch is considered to be a major threat to loggerhead turtles in the Mediterranean Sea. Technical differences in both gear configurations (e.g. hook and bait type) and fisheries operations carried out by the Spanish Mediterranean surface longline fleet could have an effect on by-catch rates and size selectivity. The aim of the present study was to test the differences in by-catch per unit effort and body size of loggerhead sea turtles caught using different gear types in the Mediterranean surface longline fishery. Our results suggest that differences in the gear type used have an effect on catch rates and size selec- tivity. Thus, surface longliners targeting albacore (LLALB) using smaller hooks tend to capture smaller loggerheads but have the highest by-catch per unit of effort (BPUE), whereas other longlines, such as surface longliners targeting bluefin tuna (LLJAP) and traditional surface longliners targeting swordfish (LLHB), using larger hooks tend to select the larger animals; moreover, LLHB had the lowest BPUE. Disproportionate rates of fisheries-induced mortality on certain size/age- classes can differentially affect sea turtle populations, as each sea turtle age-class contributes differently to current and future reproduction. Thus, fisheries management should not only be focusing on preserving the large juvenile and mature turtles, but also on reducing the total by-catch. Thus, we recommend encouraging the use of LLHB versus other surface gears. We suggest that it is very important to take into account the gear type (and its particular catch rates) when making inferences about the impact of longline fisheries on sea turtle populations.

Hyperpredation can be described as a restrictive case of apparent competition where an increased number of primary prey species indirectly induces the decrease of the secondary prey species through numerical response of predators to the primary prey dynamics. It has been proposed that rabbit hemorrhagic disease (RHD), which decimated populations of European wild rabbit (Oryctolagus cuniculus) in Spain, led to prey switching by raptors towards red-legged partridges (Alectoris rufa) causing declines in their populations as a peculiar case of hyperpredation. We have reviewed field studies that disagree with estimates (based on bag records) of recent increases of rabbit and partridge numbers in Spain. Because of an increase in releases of farm-reared animals, there is doubt about the use of bag records to accurately estimate population trends in recent decades. We also provide new data and discuss some analytical considerations related to the temporal and spatial scales that might affect the interpretation of data. Finally, we discuss why studies associated with raptor food habits, predator population dynamics and predator distribution suggest that diet data are not sufficient to link patterns and processes. Although we agree that the RHD outbreak has markedly affected rabbit populations, which has indirectly affected many other species in the Iberian Peninsula, we consider that hyperpredation mediated by raptors has not been clearly demonstrated. In contrast, endorsing the hypothesis of raptor-mediated hyperpredation without sufficient proof may have conflictive consequences if we consider the increased persecution of raptors in recent decades in Spain.

Some species are very difficult to observe in the wild, and some of these present an apparently incongruent distribution, as is the case of the spur-thighed tortoise Testudo graeca in the southern Iberian Peninsula. This species has a discontinuous distribution with two main reproductive areas: one in the contiguous provinces of Almeria and Murcia, and the other in Huelva province. Both populations are thought to maintain reproductive isolation. However, we present new records obtained from two areas (the provinces of Malaga and Cadiz) which are outside their previously known range. Malaga province and Cadiz province are located between the two traditional reproductive areas. The new records were based on 16 interviews with shepherds as well as information provided by three naturalists. The aim of this work was to analyze the spatial distribution of areas favourable to the spur-thighed tortoise in the southern Iberian Peninsula, using different models with the same group of variables but with a different number of records: known records, new records and a combination of them. We obtained a distribution model for the species in Andalusia consistent with all observations, which was related to climatic stability, climatic disturbances, and spatial situation. This model presented three main favourable areas: two are coincident with the two known nuclei, and the other is situated in the southern part of the region, in the contiguous provinces of Cadiz and Malaga. We conclude that it is important to consider all the observations in the distribution models because records outside the traditional reproductive areas can include other environmental characteristics appropriate for the species.

The idea of analysing the general favourability for the occurrence of an event was presented in 2006 through a mathematical function. However, even when favourability has been used in species distribution modelling, the conceptual framework of this function is not yet well perceived among many researchers. The present paper is conceived for providing a wider and more in-depth presentation of the idea of favourability; concretely, we aimed to clarify both the concept and the main distinctive characteristics of the favourability function, especially in relation to probability and suitability, the most common outputs in species distribution modelling. As the capabilities of the favourability function go beyond species distribution modelling, we also illustrate its usefulness for different research disciplines for which this function remains unknown. In particular, we stressed that the favourability function has potential to be applied in all the cases where the probability of occurrence of an event is analysed, such as, for example, habitat selection or epidemiological studies.

[Aim]: The extent of the study area (geographical background, GB) can strongly affect the results of species distribution models (SDMs), but as yet we lack objective and practicable criteria for delimiting the appropriate GB. We propose an approach to this problem using trend surface analysis (TSA) and provide an assessment of the effects of varying GB extent on the performance of SDMs for four species. [Location]: Mainland Spain. [Methods]: Using data for four well known wild ungulate species and different GBs delimited with a TSA, we assessed the effects of GB extent on the predictive performance of SDMs: specifically on model calibration (Miller's statistic) and discrimination (area under the curve of the receiver operating characteristic plot, AUC; sensitivity and specificity), and on the tendency of the models to predict environmental potential when they are projected beyond their training area. [Results]: In the training area, discrimination significantly increased and calibration decreased as the GB was enlarged. In contrast, as GB was enlarged, both discriminatory power and calibration decreased when assessed in the core area of the species distributions. When models trained using small GBs were projected beyond their training area, they showed a tendency to predict higher environmental potential for the species than those models trained using large GBs. [Main conclusions]: By restricting GB extent using a geographical criterion, model performance in the core area of the species distribution can be significantly improved. Large GBs make models demonstrate high discriminatory power but are barely informative. By delimiting GB using a geographical criterion, the effect of historical events on model parameterization may be reduced. Thus purely environmental models are obtained that, when projected onto a new scenario, depict the potential distribution of the species. We therefore recommend the use of TSA in geographically delimiting the GB for use in SDMs.

The application of species distribution models (SDMs) in ecology and conservation biology is increasing and assuming an important role, mainly because they can be used to hindcast past and predict current and future species distributions. However, the accuracy of SDMs depends on the quality of the data and on appropriate theoretical frameworks. In this study, comprehensive data on the current distribution of the Iberian hare (Lepus granatensis) were used to i) determine the species' ecogeographical constraints, ii) hindcast a climatic model for the last glacial maximum (LGM), relating it to inferences derived from molecular studies, and iii) calibrate a model to assess the species future distribution trends (up to 2080). Our results showed that the climatic factor (in its pure effect and when it is combined with the land-cover factor) is the most important descriptor of the current distribution of the Iberian hare. In addition, the model's output was a reliable index of the local probability of species occurrence, which is a valuable tool to guide species management decisions and conservation planning. Climatic potential obtained for the LGM was combined with molecular data and the results suggest that several glacial refugia may have existed for the species within the major Iberian refugium. Finally, a high probability of occurrence of the Iberian hare in the current species range and a northward expansion were predicted for future. Given its current environmental envelope and evolutionary history, we discuss the macroecology of the Iberian hare and its sensitivity to climate change.

Parapatry is a biogeographical term used to refer to organisms whose ranges do not overlap, but are immediately adjacent to each other; they only co-occur - if at all - in a narrow contact zone. Often there are no environmental barriers in the contact zones, hence competitive interaction is usually advocated as the factor that modulates species distribution ranges. Even though the effects of climate change on species distribution have been widely studied, few studies have explored these effects on the biogeographical relationships between closely related, parapatric, species. We modelled environmental favourability for three parapatric hare species in Europe - Lepus granatensis, L. europaeus and L. timidus - using ecogeographical variables and projected the models into the future according to the IPCC A2 emissions scenario. Favourabilities for present and future scenarios were combined using fuzzy logic with the following aims: (i) to determine the biogeographical relationships between hare species in parapatry, that is L. granatensis/L. europaeus and L. europaeus/L. timidus and (ii) to assess the effects of climate change on each species as well as on their interspecific interactions. In their contact area L. granatensis achieved higher favourability values than L. europaeus, suggesting that if both species have a similar population status, the former species may have some advantages over the latter if competitive relationships are established. Climate change had the most striking effect on the distribution of L. timidus, especially when interspecific interactions with L. europaeus were taken into account, which may compromise the co-existence of L. timidus. The results of this study are relevant not only for understanding the distribution patterns of the hares studied and the effects of climate change on these patterns, but also for improving the general application of species distribution models to the prediction of the effects of climate change on biodiversity.