Dynorphin / kappa-opioid receptor regulation of excitation-inhibition balance toggles afferent control of prefrontal cortical circuits in a pathway-specific manner.

The medial prefrontal cortex (mPFC) controls behavior via connections with limbic excitatory afferents that engage various inhibitory motifs to shape mPFC circuit function. The dynorphin (Dyn) / kappa-opioid receptor (KOR) system is highly enriched in the mPFC, and its dysregulation is implicated in neuropsychiatric disorders. However, it is unclear how the Dyn / KOR system modulates excitatory and inhibitory circuits that are integral for mPFC information processing and behavioral control. Here, we provide a circuit-based framework wherein mPFC Dyn / KOR signaling regulates excitation-inhibition balance by toggling which afferents drive mPFC neurons. Dyn / KOR regulation of afferent inputs is pathway-specific. Dyn acting on presynaptic KORs inhibits glutamate release from afferent inputs to the mPFC, including the basolateral amygdala (BLA), paraventricular nucleus of the thalamus, and contralateral cortex. The majority of excitatory synapses to mPFC neurons, including those from the ventral hippocampus (VH), do not express presynaptic KOR, rendering them insensitive to Dyn / KOR modulation. Dyn / KOR signaling also suppresses afferent-driven recruitment of specific inhibitory sub-networks, providing a basis for Dyn to disinhibit mPFC circuits. Specifically, Dyn / KOR signaling preferentially suppresses SST interneuron- relative to PV interneuron-mediated inhibition. Selective KOR action on afferents or within mPFC microcircuits gates how distinct limbic inputs drive spiking in mPFC neurons. Presynaptic Dyn / KOR signaling decreases KOR-positive input-driven (e.g. BLA) spiking of mPFC neurons. In contrast, KOR-negative input recruitment of mPFC neurons is enhanced by Dyn / KOR signaling via suppression of mPFC inhibitory microcircuits. Thus, by acting on distinct circuit elements, Dyn / KOR signaling shifts KOR-positive and negative afferent control of mPFC circuits, providing mechanistic insights into the role of neuropeptides in shaping mPFC function. Together, these findings highlight the utility of targeting the mPFC Dyn / KOR system as a means to treat neuropsychiatric disorders characterized by dysregulation in mPFC integration of long-range afferents with local inhibitory microcircuits.

Increased density of conspecifics caused niche contraction in a multispecific passerine assemblage.

Competition is a prominent mechanism driving population dynamics and structuring community assemblage, which can be investigated by linking shifts in species' ecological niche and the densities of sympatric species because the ecological release from competitive constraints is a density-dependent process. In this work we determine how a steppe passerine community segregates their ecological niches and evaluate the role of competition in inducing changes in the ecological niche of species. We built multidimensional ecological niches (with Gaussian kernel density estimators) using data on the habitat features used by 10 bird species collected from seven sites in the natural steppes of Central Spain over 2 consecutive years. We computed distance and niche similarity metrics to explore the ecological niche partitioning of the bird community. Next, we ran multivariate linear regression models to evaluate the effects of conspecific and heterospecific density (as proxies of intraspecific and interspecific competition, respectively) on niche breadth and/or position of the three most abundant species. We found low niche overlap in the community assemblage but varying levels of niche similarity among pairs of species, which could increase the likelihood of current competition operating in the community. However, we found no effect of heterospecific density on niche breadth or position, although conspecific density was negatively related to niche breadth. Contrary to predictions of competition theory, increased density of conspecifics caused niche contraction. Our results from a multispecies system contribute to advanced knowledge of the biotic mechanisms structuring wildlife communities within the framework of ecological niche theory.

Landscape features influencing gene flow and connectivity of an endangered passerine.

Dispersal of individuals and gene flow are crucial aspects to maintain genetic diversity and viability of populations, especially in the case of threatened species. Landscape composition and structure may facilitate or limit individual movement within and among populations. We used a landscape genetics approach to assess the connectivity patterns of the threatened Dupont's lark (Chersophilus duponti subsp. duponti), considering their genetic patterns and the landscape features associated with its gene flow in Spain. We analysed the genetic relatedness based on 11 species-specific polymorphic microsatellites on 416 Dupont's lark individuals sampled across peninsular Spain between 2017 and 2019, covering most of the European distribution of the species. To assess the relationship between the landscape composition and the species gene flow, we estimated genetic distance at the individual level (Dps). Next, we built a set of environmental surfaces from two time periods (years 1990 and 2018), based on factors such as land use and topography, influencing individuals' movement. We then obtained resistance surfaces from an optimization process on landscape variables. Landscape genetics analyses were done for single and composite surface models for each year separately. Our findings from both time points show that scatter or mosaic-structured vegetation composed by low agricultural and tree cover and high presence of sclerophyllous shrubs favoured Dupont's lark dispersal, while dense and continuous tree cover, as well as areas of intensive agriculture, were limiting factors. Our results suggest the importance of steppe habitat patches for the species' establishment and dispersal. In addition, our results provide key information to develop conservation measures, including conserving and restoring steppe habitats as scattered and/or mosaic-structured vegetation that could warrant the connectivity and persistence of Dupont's lark populations.

Prefrontal cortical dynorphin peptidergic transmission constrains threat-driven behavioral and network states.

Prefrontal cortical (PFC) circuits provide top-down control of threat reactivity. This includes ventromedial PFC (vmPFC) circuitry, which plays a role in suppressing fear-related behavioral states. Dynorphin (Dyn) has been implicated in mediating negative affect and mal-adaptive behaviors induced by severe threats and is expressed in limbic circuits, including the vmPFC. However, there is a critical knowledge gap in our understanding of how vmPFC Dyn-expressing neurons and Dyn transmission detect threats and regulate expression of defensive behaviors. Here, we demonstrate that Dyn cells are broadly activated by threats and release Dyn locally in the vmPFC to limit passive defensive behaviors. We further demonstrate that vmPFC Dyn-mediated signaling promotes a switch of vmPFC networks to a fear-related state. In conclusion, we reveal a previously unknown role of vmPFC Dyn neurons and Dyn neuropeptidergic transmission in suppressing defensive behaviors in response to threats via state-driven changes in vmPFC networks.

Prefrontal cortical dynorphin peptidergic transmission constrains threat-driven behavioral and network states.

Prefrontal cortical (PFC) circuits provide top-down control of threat reactivity. This includes ventromedial PFC (vmPFC) circuitry, which plays a role in suppressing fear-related behavioral states. Dynorphin (Dyn) has been implicated in mediating negative affect and maladaptive behaviors induced by severe threats and is expressed in limbic circuits, including the vmPFC. However, there is a critical knowledge gap in our understanding of how vmPFC Dyn-expressing neurons and Dyn transmission detect threats and regulate expression of defensive behaviors. Here, we demonstrate that Dyn cells are broadly activated by threats and release Dyn locally in the vmPFC to limit passive defensive behaviors. We further demonstrate that vmPFC Dyn-mediated signaling promotes a switch of vmPFC networks to a fear-related state. In conclusion, we reveal a previously unknown role of vmPFC Dyn neurons and Dyn neuropeptidergic transmission in suppressing defensive behaviors in response to threats via state-driven changes in vmPFC networks.

Dissociable control of motivation and reinforcement by distinct ventral striatal dopamine receptors.

Dopamine release in striatal circuits, including the nucleus accumbens (NAc), tracks separable features of reward such as motivation and reinforcement. However, the cellular and circuit mechanisms by which dopamine receptors transform dopamine release into distinct constructs of reward remain unclear. Here, we show that dopamine D3 receptor (D3R) signaling in the NAc drives motivated behavior by regulating local NAc microcircuits. Furthermore, D3Rs co-express with dopamine D1 receptors (D1Rs), which regulate reinforcement, but not motivation. Paralleling dissociable roles in reward function, we report non-overlapping physiological actions of D3R and D1R signaling in NAc neurons. Our results establish a novel cellular framework wherein dopamine signaling within the same NAc cell type is physiologically compartmentalized via actions on distinct dopamine receptors. This structural and functional organization provides neurons in a limbic circuit with the unique ability to orchestrate dissociable aspects of reward-related behaviors that are relevant to the etiology of neuropsychiatric disorders.

Extensive sheep grazing is associated with trends in steppe birds in Spain: recommendations for the Common Agricultural Policy.

Iberian natural steppes have traditionally been used for extensive sheep grazing, which has been noted to be positively associated with steppe bird abundance and diversity. Sheep numbers in Spain, which harbors the largest European populations of many steppe bird species, decreased by 9.2 million (37.3%) between 1992 and 2020. Steppe birds in Spain have faced dramatic declines during the same period, but there is a lack of knowledge about the potential association between sheep and open-habitat bird declines. We used sheep data from the Spanish Ministry of Agriculture and bird data (1998-2018) from the Spanish Common Bird Monitoring Program to assess the association at the Spanish scale between sheep decline and the Farmland Bird Index (FBI) and the Natural Shrub-steppe Bird Index (SBI). We also used an independent dataset on population trends of the Dupont's Lark (Chersophilus duponti) to assess the relationship between sheep numbers and the decline of this threatened steppe specialist passerine in Spain, whose European population is restricted to Iberian natural steppes. To test for a spurious relationship between temporal series, variables were tested for cointegration. After confirming cointegration, we found a strong positive relationship between sheep abundance and the trends of the FBI and SBI indices during the period 1998-2018. The association between sheep abundance and trends of the Dupont's Lark (2004-2015) was positive although it was not statistically significant. Although the main causes of decline of farmland and steppe birds are mainly related to agricultural intensification and land use changes, the correlation found, using two independent cointegrated datasets, between the reduction in farmland and shrub-steppe birds and sheep numbers at the country scale suggests that the decline of steppe birds in Spain may be also associated with the decline in sheep numbers. This agrees with previous studies that found a positive relationship between intermediate levels of sheep grazing and steppe bird abundance in Iberian steppes. Further research (e.g. experimental studies) is needed to corroborate our study and identify the most appropriate level of grazing intensity for protecting the most farmland and shrub-steppe birds. Our results suggest that the promotion of extensive grazing should be considered as a key factor in future Common Agricultural Policy reforms and conservation programmes to protect steppe birds.

Wind farm noise shifts vocalizations of a threatened shrub-steppe passerine.

Wind energy has experienced a notable development during the last decades, driving new challenges for animal communities. Although bird collisions with wind turbines and spatial displacement due to disturbance have been widely described in the literature, other potential impacts remain unclear. In this study, we addressed the effect of turbine noise on the vocal behaviour of a threatened shrub-steppe passerine highly dependent on acoustic communication, the Dupont's lark Chersophilus duponti. Based on directional recordings of 49 calling and singing males exposed to a gradient of turbine noise level (from 15 up to 51 dBA), we tested for differences in signal diversity, redundancy, and complexity, as well as temporal and spectral characteristics of their vocalizations (particularly the characteristic whistle). Our results unveiled that Dupont's lark males varied the vocal structure when subject to turbine noise, by increasing the probability of emitting more complex whistles (with increased number of notes) and shifting the dominant note (emphasizing the longest and higher-pitched note). In addition, males increased duration and minimum frequency of specific notes of the whistle, while repertoire size and signal redundancy remain constant. To our knowledge, this is the first study reporting multiple and complex responses on the vocal repertoire of animals exposed to turbine noise and unveiling a shift of the dominant note in response to anthropogenic noise in general. These findings suggest that the Dupont's lark exhibits some level of phenotypic plasticity, which might enable the species to cope with noisy environments, although the vocal adjustments observed might have associated costs or alter the functionality of the signal. Future wind energy projects must include fine-scale noise assessments to quantify the consequences of chronic noise exposure.

Protect European green agricultural policies for future food security.

European green agricultural policies have been relaxed to allow cultivation of fallow land to produce animal feed and meet shortfalls in exports from Ukraine and Russia. However, conversion of semi-natural habitats will disproportionately impact long term biodiversity and food security.

Population viability analysis of the endangered Dupont's Lark Chersophilus duponti in Spain.

Steppe lands in Europe are critically affected by habitat loss and fragmentation, and hold over 50% of IUCN Red List bird species in Europe. Dupont's Lark is a threatened steppe-specialist passerine whose European geographic range is restricted to Spain, with less than 2000 pairs and an annual population decline of - 3.9%. Its strongly fragmented habitat leads to a metapopulation structure in the Iberian Peninsula that includes 24 populations and 100 subpopulations. We present an updated Population Viability Analysis based on the latest scientific knowledge regarding distribution, population trends, breeding biology and connectivity. Our results predict metapopulation extinction in 2-3 decades, through a centripetal contraction process from the periphery to the core. The probability of extinction in 20 years was 84.2%, which supports its relisting to Endangered in Spain following IUCN criteria. We carried out a sensitivity analysis showing that some parameters, especially productivity and survival of adults and juveniles, help to increase metapopulation viability. Simulation of management scenarios showed that habitat restoration in a subset of key subpopulations had a positive effect on the overall metapopulation persistence. Translocations of a limited number of individuals from source to recipient locations may help to rescue the most endangered subpopulations without reducing the global time to extinction of the metapopulation. In addition, we identified the most critical areas for action, where local populations of the species are prone to extinction. This work suggests that the viability of the Dupont's Lark metapopulation could be improved and its risk of extinction reduced if urgent and localized conservation measures are applied. In the short-term, habitat loss and fragmentation due to ploughing, reforestation and infrastructures implementation in Dupont's Lark habitat must be avoided. Habitat restoration and translocations could help to avoid imminent extinction of critical subpopulations. Restoration of extensive grazing is recommended as the most effective way to achieve the long-term conservation of Dupont's Lark in Spain.

Connectivity in Spanish metapopulation of Dupont's lark may be maintained by dispersal over medium-distance range and stepping stones.

BACKGROUND: Dupont's Lark is an endangered bird, whose fragmented distribution in Europe is entirely restricted to Spain. This lark, suffering pronounced population decline and range contraction, inhabits steppes that have long been used for grazing sheep and are now threatened by rural abandonment and land use changes. Thus, for conservation of the lark, increasing knowledge about the connectivity of the Spanish metapopulation and identifying the most important connectivity nodes are crucial. METHODS: The study was carried out in Spain, using over 16,000 Dupont's Lark georeferenced observations. We used distance buffers to define populations and subpopulations, based on the available scientific information. We identified potential stepping stones using a MaxEnt probability of presence model. Connectivity was assessed using Conefor software, using the centroid of each subpopulation and stepping stone as nodes. Each node was assigned a quantitative attribute based on total habitat area, within-node habitat quality and internal fragmentation. We evaluated different connectivity scenarios by potential movement thresholds (5-20-100 km) and presence or absence of stepping stones in the network. RESULTS: Dupont's Lark Iberian metapopulation comprises 24 populations and 100 subpopulations, plus 294 potential stepping stones. Movement thresholds and stepping stones had a strong influence in the potential network connectivity. The most important nodes are located in the core of the metapopulation, which shows connectivity among subpopulations in the different indices and scenarios evaluated. Peripheral populations are more isolated and require stepping stones or medium (20 km) or long (100 km) potential movement thresholds to join the network. DISCUSSION: Metapopulation connectivity may be greater than expected, thanks to stepping stones and potential medium-distance movements. Connectivity is crucial for conservation and can be increased by preserving or improving adequate habitat in the most important nodes. Given the current species decline, steppe habitat should be urgently protected from land use changes and agriculture intensification, at least in the critical subpopulations and stepping stones. Long-term conservation of steppe lands and Dupont's Lark in Spain requires the recovery of traditional grazing and more research on juvenile dispersion. Meanwhile, the conservation of potentially critical stepping stones should be incorporated to management plans.

The decline of farmland birds in Spain is strongly associated to the loss of fallowland.

Farmland bird populations have strongly declined across Europe over the last decades due to agriculture intensification, despite successive reforms of EU's Common Agricultural Policy (CAP). In parallel, CAP has led to a reduction of fallow land, a critical habitat for biodiversity in agroecosystems. Fallow land in Spain, a country harboring the largest European populations of many endangered farmland birds, has decreased by 1.1 million ha in 15 years. The significant positive relationship between yearly change rates of the Spanish Farmland and Cereal Bird Indices (FBI and CBI) and fallow surface change highlights the adequacy of fallow land cover as an indicator of the state of farmland bird communities at country level. Moreover, the strong and positive association between the reduction in abundance of the fallow specialist little bustard and fallow surface suggests a potential causal link between these two factors. These results highlight the need for a new CAP that guarantees the maintenance of fallow land in European agroecosystems if farmland bird populations are to be conserved.

Hierarchical habitat-use by an endangered steppe bird in fragmented landscapes is associated with large connected patches and high food availability.

Multidimensional approaches must be employed when addressing habitat use patterns. In this study, we aim to elucidate the hierarchical nature of space use by species inhabiting fragmented landscapes, using the threatened Dupont's lark (Chersophilus duponti). The intensity of space use by Dupont's lark was estimated using the Kernel Density Function on territory locations in 2015. We measured descriptors of habitat quality at metapopulation (connectivity and patch size), landscape (land-use types and anthropogenic disturbance) and microhabitat-scale (plant structure and composition, herbivore abundance and food availability) at 37 sampling stations. We fitted a Partial Least Squares Regression (PLSR) which yielded two components, accounting for 81% of total variance. Metapopulation-scale factors had the greatest explanatory power (32%), followed by microhabitat (17%) landscape (10%) and spatial predictors (3.6%). Connectivity and patch size were key factors explaining habitat use, and wind farms had a negative effect. At microhabitat-scale, space use was positively associated with Coleoptera, Orthoptera, Araneae and Diptera biomass, but negatively with Formicidae and Blattodea biomass, the cover of Stipa spp, Koeleria vallesiana and moss. This research highlights the hierarchical nature of habitat use in fragmented landscapes. Therefore, conservation measures should ensure connectivity, guarantee a minimum patch size, and improve habitat quality within patches.

Factors affecting Dupont´s lark distribution and range regression in Spain.

In this work, we analyse factors explaining the distribution and range regression of Dupont's lark in Spain, the only European country in which this threatened alaudid is present. Dupont's lark is an extremely elusive and scarce species, distributed across a reduced and strongly fragmented range, showing a metapopulational structure with unknown dispersive and connective mechanisms. We used maximum entropy modelling (Maxent) on nearly 15,000 Dupont's lark observations (1985-2015) to assess the probability of presence at a 1 km resolution across its European range. Moreover, we tested the probability of extinction by comparing pre- and post-2000 observations by means of a GLM over a subset of cells with presence-absence data. We obtained strong model fitting (AUC = 0.919), in which species occurrence was explained by low values of plant productivity (NDVI), climate (high temperature range and medium annual precipitation), land use (increasing with sclerophyllous scrubland), flat topography and human disturbance (associated with low human population density). The species also tolerates dry farming, but not other farm types or forest cover. The probability map identified two main regions known as the species' core areas: the steppes of the Iberian System and the Ebro Valley. The North Plateau is characterised by a dispersed structure of small and very fragmented patches of suitable habitat, while a succession of discontinuous probability patches form an Eastern Corridor connecting the central core areas to the southernmost populations. Finally, the model identified small and isolated patches of high probability of presence along the eastern coastline. The species tends to occur in the best available areas but, at the same time, the model revealed a large area of suitable but unoccupied habitat. Our results correct the previous estimation of occupation area from 1,480 to 1,010.78 km2, a reduction of 26.22%. The current distribution of Dupont's lark is almost completely covered by Important Bird Areas (IBAs), highlighting their importance for bird conservation, but only 44.89% is included in Natura 2000 Special Protection Areas (SPAs). A comparison of pre- and post-2000 periods revealed a range contraction of 44%. Probability of extinction increased with higher temperature range and lower annual precipitation, and with decreases in population density, which suggests that this species is extremely vulnerable to both climate change and rural abandonment, due to its dependence on traditional grazing. These results suggest the need for a re-evaluation of the conservation status of Dupont's lark in Spain. They urge the preservation of not only current extant populations, but also the unoccupied suitable areas that could be critical for metapopulation structure, and the development of policies addressing the preservation of traditional grazing.

Correction: Factors affecting Dupont's lark distribution and range regression in Spain.

[This corrects the article DOI: 10.1371/journal.pone.0211549.].

Genetic Knockdown of mGluR5 in Striatal D1R-Containing Neurons Attenuates L-DOPA-Induced Dyskinesia in Aphakia Mice.

L-DOPA is the main pharmacological therapy for Parkinson's disease. However, long-term exposure to L-DOPA induces involuntary movements termed dyskinesia. Clinical trials show that dyskinesia is attenuated by metabotropic glutamate receptor type 5 (mGluR5) antagonists. Further, the onset of dyskinesia is delayed by nicotine and mGluR5 expression is lower in smokers than in non-smokers. However, the mechanisms by which mGluR5 modulates dyskinesia and how mGluR5 and nicotine interact have not been established. To address these issues, we studied the role of mGluR5 in D1R-containing neurons in dyskinesia and examined whether nicotine reduces dyskinesia via mGluR5. In the aphakia mouse model of Parkinson's disease, we selectively knocked down mGluR5 in D1R-containing neurons (aphakia-mGluR5KD-D1). We found that genetic downregulation of mGluR5 decreased dyskinesia in aphakia mice. Although chronic nicotine increased the therapeutic effect of L-DOPA in both aphakia and aphakia-mGluR5KD-D1 mice, it caused a robust reduction in dyskinesia only in aphakia, and not in aphakia-mGluR5KD-D1 mice. Downregulating mGluR5 or nicotine treatment after L-DOPA decreased ERK and histone 3 activation, and FosB expression. Combining nicotine and mGluR5 knockdown did not have an added antidyskinetic effect, indicating that the effect of nicotine might be mediated by downregulation of mGluR5 expression. Treatment of aphakia-mGluR5KD-D1 mice with a negative allosteric modulator did not further modify dyskinesia, suggesting that mGluR5 in non-D1R-containing neurons does not play a role in its development. In conclusion, this work suggests that mGluR5 antagonists reduce dyskinesia by mainly affecting D1R-containing neurons and that the effect of nicotine on dyskinetic signs in aphakia mice is likely via mGluR5.

European population trends and current conservation status of an endangered steppe-bird species: the Dupont's lark Chersophilus duponti.

BACKGROUND: Steppe-birds face drastic population declines throughout Europe. The Dupont's lark Chersophilus duponti is an endangered steppe-bird species whose European distribution is restricted to Spain. This scarce passerine bird could be considered an 'umbrella species', since its population trends may reveal the conservation status of shrub-steppes. However, trends for the Spanish, and thus European, population of Dupont's lark are unknown. In this work, we evaluated Dupont's lark population trends in Europe employing the most recent and largest compiled database to date (92 populations over 12 years). In addition, we assessed the species threat category according to current applicable criteria (approved in March 2017) in the Spanish catalogue of threatened species (SCTS), which have never been applied to the Dupont's lark nor to any other Spanish species. Finally, we compared the resulting threat categories with the current conservation status at European, national and regional levels. METHODS: We fitted switching linear trend models (software TRIM-Trends and Indices for Monitoring data) to evaluate population trends at national and regional scale (i.e. per Autonomous Community) during the period 2004-2015. In addition, the average finite annual rate of change ( λ ¯ ) obtained from the TRIM analysis was employed to estimate the percentage of population size change in a 10-year period. A threat category was assigned following A1 and A2 criteria applicable in the SCTS. RESULTS: Trends showed an overall 3.9% annual decline rate for the Spanish population (moderate decline, following TRIM). Regional analyses showed high inter-regional variability. We forecasted a 32.8% average decline over the next 10 years. According to these results, the species should be listed as 'Vulnerable' at a national scale (SCTS). At the regional level, the conservation status of the species is of particular concern in Andalusia and Castile-Leon, where the species qualifies for listing as 'Endangered'. DISCUSSION: Our results highlight the concerning conservation status of the European Dupont's lark population, undergoing a 3.9% annual decline rate. Under this scenario, the implementation of a wide-ranging conservation plan is urgently needed and is vital to ensuring the conservation of this steppe-bird species. The role of administrations in matters of nature protection and the cataloguing of endangered species is crucial to reverse declining population trends of this and other endangered taxa.

Intraspecific and interspecific competition induces density-dependent habitat niche shifts in an endangered steppe bird.

Interspecific competition is a dominant force in animal communities that induces niche shifts in ecological and evolutionary time. If competition occurs, niche expansion can be expected when the competitor disappears because resources previously inaccessible due to competitive constraints can then be exploited (i.e., ecological release). Here, we aimed to determine the potential effects of interspecific competition between the little bustard (Tetrax tetrax) and the great bustard (Otis tarda) using a multidimensional niche approach with habitat distribution data. We explored whether the degree of niche overlap between the species was a density-dependent function of interspecific competition. We then looked for evidences of ecological release by comparing measures of niche breadth and position of the little bustard between allopatric and sympatric situations. Furthermore, we evaluated whether niche shifts could depend not only on the presence of great bustard but also on the density of little and great bustards. The habitat niches of these bustard species partially overlapped when co-occurring, but we found no relationship between degree of overlap and great bustard density. In the presence of the competitor, little bustard's niche was displaced toward increased use of the species' primary habitat. Little bustard's niche breadth decreased proportionally with great bustard density in sympatric sites, in consistence with theory. Overall, our results suggest that density-dependent variation in little bustard's niche is the outcome of interspecific competition with the great bustard. The use of computational tools like kernel density estimators to obtain multidimensional niches should bring novel insights on how species' ecological niches behave under the effects of interspecific competition in ecological communities.

Forecasting Large-Scale Habitat Suitability of European Bustards under Climate Change: The Role of Environmental and Geographic Variables.

We modelled the distribution of two vulnerable steppe birds, Otis tarda and Tetrax tetrax, in the Western Palearctic and projected their suitability up to the year 2080. We performed two types of models for each species: one that included environmental and geographic variables (space-included model) and a second one that only included environmental variables (space-excluded model). Our assumption was that ignoring geographic variables in the modelling procedure may result in inaccurate forecasting of species distributions. On the other hand, the inclusion of geographic variables may generate an artificial constraint on future projections. Our results show that space-included models performed better than space-excluded models. While distribution of suitable areas for T. tetrax in the future was approximately the same as at present in the space-included model, the space-excluded model predicted a pronounced geographic change of suitable areas for this species. In the case of O. tarda, the space-included model showed that many areas of current presence shifted to low or medium suitability in the future, whereas a northward expansion of intermediate suitable areas was predicted by the space-excluded one. According to the best models, current distribution of these species can restrict future distribution, probably due to dispersal constraints and site fidelity. Species ranges would be expected to shift gradually over the studied time period and, therefore, we consider it unlikely that most of the current distribution of these species in southern Europe will disappear in less than one hundred years. Therefore, populations currently occupying suitable areas should be a priority for conservation policies. Our results also show that climate-only models may have low explanatory power, and could benefit from adjustments using information on other environmental variables and biological traits; if the latter are not available, including the geographic predictor may improve the reliability of predicted results.