Trophic structure and habitat determinants of a Mediterranean carnivore guild: insights from the Sierra de la Culebra (NW Spain)

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Indeed, the Sierra de la Culebra (Zamora, NW Spain) hosts one of the densest populations of Iberian wolves ( Canis lupus signatus ) in Western Europe, together with a diverse guild of mesocarnivores. We investigated habitat determinants of carnivore diversity and trophic structure using a non-invasive, multi-method approach that combined camera trapping, faecal DNA identification, and interviews with local rangers across twelve 7.5 × 7.5 km grid cells (65,000 ha). Seven camera trap stations were positioned in the grid, with one camera trap per station, after the results were cross-referenced with 253 fecal samples and a survey conducted among Park staff. Finally, to determine the habitat, the relative abundance index (RAI) for each taxon was calculated using the formula suggested by Lira-Torres and Briones Salas. Twelve carnivore species were recorded, with wolves and red foxes ubiquitous and smaller mustelids showing patchy distributions although wolf presence exerted a weak negative influence on mesocarnivore richness, suggesting moderate top-down regulation within the guild. As regards the characterization of the habitat multiple regression models revealed that deciduous forests and shrublands were positively associated with total species richness, while pine plantations and croplands had negative effects therefore the carnivore community of the Sierra de la Culebra has shown a significant predilection for forest areas with deciduous or thicket forests, strongly reaching coniferous forests (generally repopulation) and cultivated areas. In the European context, where the wolf is a sensitive species, these results demonstrate that carnivore diversity in Mediterranean landscapes depends on both trophic and habitat complexity. Ecological restoration in the Sierra de la Culebra should therefore prioritize the recovery of native oak woodlands and mixed shrub mosaics to maintain a functionally rich and balanced predator community. carnivore guild trophic structure habitat heterogeneity top-down regulation community ecology Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 1. Introduction Habitat fragmentation and landscape transformation are among the most pervasive drivers of biodiversity loss worldwide. habitats that require large, continuous areas of natural habitat are particularly vulnerable to these changes, as fragmentation leads to isolation, reduced gene flow, and altered community dynamics (Beier et al., 1993; Mortelliti et al., 2010). Mammalian carnivores are especially sensitive to habitat reduction because of their large home ranges and high trophic position, which make them reliable indicators of ecosystem integrity (Crooks, 2002; Reed, 2004).In Mediterranean environments, the progressive replacement of native woodlands and shrublands by pine plantations, crops, and infrastructures has reshaped ecological networks and modified the structure of predator communities (Waldron et al., 2006; Debinski & Holt, 2000). While some species exhibit high tolerance to human-modified landscapes, others show pronounced declines, making comparative studies of habitat selection essential for conservation and restoration planning. The Sierra de la Culebra, located in northwestern Zamora (Castilla y León, Spain), represents one of the most emblematic Mediterranean mountain ranges in the Iberian Peninsula. Covering approximately 65,000 ha, this Regional Hunting Reserve and UNESCO Biosphere Reserve is internationally known for hosting one of the densest populations of Iberian wolves ( Canis lupus signatus ) in Western Europe (Vicente et al., 2000). The area comprises a mosaic of habitats (heathlands, pine forests ( Pinus pinaster, P. sylvestris ), oak woodlands ( Quercus pyrenaica, Q. ilex ), pastures, and croplands) shaped by centuries of silviculture, grazing, and hunting (Ramos Santos, 2006; Hidalgo, 2008). Despite its ecological importance, few studies have addressed the structure of the carnivore community as a whole or the relationships between top predators and mesocarnivores in this unique landscape. Understanding habitat preferences and interspecific interactions within the carnivore guild of the Sierra de la Culebra is essential for developing effective management and ecological restoration strategies. Habitat selection models can provide insights into how vegetation composition, land-use intensity, and predator presence influence species richness and distribution patterns. The objectives of this study were therefore to (i) assess the species composition and relative abundance of carnivores in the Sierra de la Culebra using complementary non-invasive methods (camera trapping, faecal DNA analysis, and interviews), (ii) identify habitat preferences and key environmental predictors of carnivore richness, and (iii) evaluate the influence of top predators (wolves) and exotic species (American mink) on community structure. We hypothesized that ( 1 ) carnivore richness would be higher in deciduous forests and shrublands than in pine plantations and croplands, due to the greater structural complexity and prey availability of natural habitats. ( 2 ) The presence of the top predator, the Iberian wolf ( Canis lupus signatus ), would moderately reduce mesocarnivore richness through interspecific competition and top-down regulation. ( 3 ) The integration of complementary non-invasive methods, camera trapping, faecal DNA analysis, and interviews, would provide a more comprehensive and accurate assessment of community composition than any single approach alone. 2. Material and Methods 2.1. Study area The study was conducted in the Sierra de la Culebra Regional Hunting Reserve (Zamora, Castilla y León, NW Spain), an area also designated as a UNESCO Biosphere Reserve. The reserve extends over approximately 65,000 ha, ranging from 800 to 1,243 m a.s.l., with Peña Mira as its highest peak (Vicente et al., 2000). The mountain range is composed of parallel quartzite ridges-oriented NNW-SSE and intersected by small fluvial valleys (Fig. 1 ). Its natural limits are the River Tera to the north and the River Aliste to the south, while it borders the region of Tábara in the east and the foothills of the Sierra Segundera in the west (Ramos Santos, 2006). Both the structure and composition of the vegetation have been substantially modified by human activities. The autochthonous deciduous forests, mainly Quercus pyrenaica , Q. ilex subsp. rotundifolia and Q. faginea , are now interspersed with cultivated lands and large pine plantations ( Pinus pinaster, P. sylvestris, P. nigra ) established for timber production. Heather ( Erica australis, E. arborea, E. umbellata ) and Cistus shrubs ( C. laurifolius, C. ladanifer ) cover approximately 55.7% of the study area, followed by pine forests (22.7%), deciduous woodlands (5.7%), pastures (7.9%) and croplands (7.7%) (Centro Nacional de Información Geográfica, 2018). The local fauna is highly diverse, including at least six fish, fourteen amphibians, fourteen reptiles, twenty-nine mammals, and over one hundred bird species (Vicente et al., 2000). 2.2. Sampling design Fieldwork was carried out between March and June 2019. The reserve was divided into twelve 7.5 × 7.5 km grid cells (Fig. 2 ), each covering approximately 5,625 ha. Within each grid, seven camera-trapping stations were established, giving a total of 84 stations across the study area. Cameras (Campark models equipped with 44 infrared LEDs and IR filters) were active 24 h per day for seven consecutive days at each station (Zielinski & Kucera, 1995; Torres et al., 2003; Barea-Azcón et al., 2007). To increase the detection probability of mesocarnivores, each station was baited with sardines in oil and lynx urine as olfactory attractants (Ferreras et al., 2016). Total sampling effort amounted to 588 camera-days. In parallel, faecal surveys were conducted along linear transects of approximately 3 km per grid cell (Rodríguez-Mazzini, 1996), each surveyed twice. A total of 253 faecal samples was collected and preserved in autoclaved tubes containing 96% ethanol, then stored at − 20°C until genetic analysis. During fieldwork, all tracks and visual records of carnivores were noted following the identification criteria of Sanz & Turón (2017). Geographic coordinates of every detection were recorded with a Garmin GPS. Finally, 21 structured interviews were conducted with forest agents and hunting wardens working in the reserve to obtain complementary information on carnivore presence and activity patterns. 2.3 Laboratory procedures DNA was extracted from faecal samples using the FavorPrep Mini Stool DNA Isolation Kit (Favorgen Biotech Corp.) according to the manufacturer’s protocol and the general molecular procedures of Sambrook et al. (1989). Species identification focused on mustelids and other small carnivores was achieved by a PCR-RFLP approach following Ruiz-González et al. (2008). Amplified mtDNA fragments were digested simultaneously with RsaI and BsuRI, generating species-specific restriction patterns that allowed unequivocal discrimination of sympatric species such as Martes martes , Martes foina , Neogale vison and Lutra lutra (Ruiz-González et al., 2008; Dettori et al., 2021). PCR reactions were run on a GeneAmp PCR System 2700 (Applied Biosystems). Amplicons were sequenced at Macrogen Europe (Amsterdam, Netherlands) using an ABI Prism 3730XL analyzer. The resulting sequences were aligned in GENEIOUS v7.1.7 (Biomatters Ltd.) and compared with homologous sequences from the GenBank database using the BLAST algorithm of the National Center for Biotechnology Information (NCBI). 2.4 Habitat classification Habitat information was derived from the CORINE Land Cover database (Centro Nacional de Información Geográfica, 2018) and processed in QGIS v3.14. Each 7.5 × 7.5 km grid was overlaid with the land-cover map, and the proportional area (%) of each of seven major habitat categories was calculated (Table 1 ): H1 = Crops, H2 = Pastures, H3 = Shrublands, H4 = Deciduous forests, H5 = Coniferous forests, H6 = Watercourses, H7 = Anthropic elements. Two additional categorical variables were included: H8 = presence of wolf home ranges (1 = absent, 2 = partially overlapping, 3 = fully overlapping) and H9 = presence/absence of the invasive American mink ( Neogale vison ) (0/1) (Table 2 ). These variables were later incorporated into multiple linear regression models to assess how habitat composition, predator presence, and alien species influenced carnivore richness across the grid cells. Table 1 Habitat variables, obtained by merging Corine Land Covers (Centro Nacional de Información Geográfica, 2018). Corine Land Cover Code Land 211 Dry farmland 221 Vineyards H1-Crops 242 Crop mosaic 243 Mainly agricultural land, but with important spaces of natural vegetation 333 Spaces with sparse vegetation H2-Pastures 231 Grasslands 321 Natural grasslands H3-Shrubs 322 Moors and scrub 323 Sclerophyll vegetation 324 Transitional scrub forest H4-Deciduous forest 311 Leafy woods 313 Mixed forest H5-Conifers 312 Coniferous forests H6-Water courses 511 Water courses 512 Water sheets 112 Discontinuous urban areas H7-Anthropic elements 131 Mining extraction areas 132 Dumps and landfills 133 Area under construction Table 2 H1 = Crops; H2 = Pastures; H3 = Scrub; H4 = Deciduous forest; H5 = Coniferous forest; H6 = Bodies of water; H7 = Anthropic elements; P/A H8 = Presence/Absence of Wolf and P/A H9 = Presence/Absence of American mink. In the P/A H8 column, the value 1 = No wolf vital domain present; 2 = Presence shared life domain between grids; 3 = Vital domain exclusive presence on grid. In the P/A H9 column, the value 0 = Absence of mink; 1 = presence of mink. Grids Richness % H1 % H2 % H3 % H4 % H5 % H6 % H7 P/A H8 P/A H9 1 7 7,91 11,87 62,91 1,12 14,27 0,00 1,92 3,00 0,00 2 5 1,09 2,45 42,00 2,94 50,17 0,03 1,33 3,00 0,00 3 11 3,47 22,09 48,99 15,91 1,78 2,93 4,83 1,00 1,00 4 5 0,09 6,85 56,99 3,06 31,74 1,09 0,18 3,00 0,00 5 8 2,84 2,53 88,54 2,09 3,94 0,00 0,07 3,00 1,00 6 10 4,43 4,48 75,21 2,72 11,18 0,00 1,99 3,00 1,00 7 9 3,36 6,40 57,65 12,81 18,45 0,36 0,95 3,00 1,00 8 9 0,50 5,53 61,42 7,85 20,70 0,00 4,00 3,00 0,00 9 5 9,63 10,66 50,40 2,31 25,72 0,00 1,27 2,00 0,00 10 6 24,89 11,02 35,46 0,78 27,79 0,00 0,05 2,00 1,00 11 4 12,65 6,90 42,36 6,19 28,38 0,00 3,52 3,00 0,00 12 7 21,77 4,20 47,34 10,90 15,42 0,00 0,37 2,00 0,00 Table 3 Summary of the 1st explanatory model of the carnivores richness in the Sierra de la Culebra. First model Coefficient Sig (p) R² R² (adjusted) Constant 9,27205 0,0000 0,7307 0,6297 Pine grove -0,10826 0,0124 Deciduous forest 0,12684 0,1873 Crops -0,07593 0,1651 Table 4 Summary of the 2nd model used to determine the richness of carnivores in the Sierra de la Culebra. Second model Coefficient Sig (p) R² R² (adjusted) Constant 2,68575 0,2923 0,7107 0,622 Shrub 0,10407 0,0105 Deciduous forest 0,23761 0,0361 Wolf presence -1,03685 0,2246 2.5 Data analysis Habitat selection: To compare habitat use at the transect scale, faecal samples were assigned to one of the seven habitat types defined above. Habitat selection was tested using Chi-squared (χ²) goodness-of-fit tests (White & Garrott, 1990; Balestrieri et al., 2015), with expected frequencies based on the relative length of transects in each habitat. Significant selection or avoidance was evaluated using Bonferroni confidence intervals (Neu et al., 1974; Byers et al., 1984; Manly et al., 1993; Di Luzio, 2010; Dettori et al., 2017). Community-level models: At the community level, species richness per grid cell was modeled through multiple linear regression to determine the effects of land-cover variables (H1-H7) and biotic predictors (H8-H9). Two models were tested: i) Model 1 included only habitat variables (percent cover of pine forest, deciduous forest, cropland). ii) Model 2 added shrub cover and wolf presence to evaluate biotic modulation of richness. Model performance was evaluated by R² and adjusted R², and predictor significance was assessed at p < 0.05. Normality of residuals and multicollinearity were examined before interpretation. All statistical analyses were performed in SPSS v13.0 (IBM Corp., Armonk, NY, USA). Graphical outputs and spatial visualizations were prepared in QGIS and Microsoft Excel. 2.6 Ethical statement All field procedures were non-invasive and complied with Spanish wildlife protection regulations (Law 4/1996, Castilla y León). Sampling and data collection were conducted with the authorization of the Regional Hunting Reserve authorities. 3. Results and discussion Carnivore community composition. A total of twelve carnivore species were detected in the Sierra de la Culebra during the 2019 survey: red fox ( Vulpes vulpes ), Iberian wolf ( Canis lupus signatus ), common genet ( Genetta genetta ), stone marten ( Martes foina ), pine marten ( Martes martes ), Eurasian badger ( Meles meles ), European polecat ( Mustela putorius ), least weasel ( Mustela nivalis ), wildcat ( Felis silvestris ), Egyptian mongoose ( Herpestes ichneumon ), Eurasian otter ( Lutra lutra ), and American mink ( Neogale vison ). This diverse assemblage includes both apex predators and mesocarnivores, confirming that the Sierra de la Culebra maintains a functionally complete predator guild (Vicente et al., 2000). Camera trapping, faecal DNA analysis, and interviews proved complementary. Wolves and red foxes were the most ubiquitous species, detected in all twelve grid cells, followed by pine and stone martens (11 grids), badger and genet (10), while the American mink and wildcat appeared in 5 grids. The polecat, otter, weasel, and mongoose were recorded only in 2-4 grids, mostly near riparian environments. The total camera-trapping effort was 588 camera-days, yielding a capture success of 37.4% (37.4 detections/100 camera-days). American mink, red fox, and genet showed the highest activity indices. These results highlight that species with generalist or semi-aquatic habits were the most detectable through camera and bait-based methods (Lira-Torres & Briones-Salas, 2012). 3.1 Habitat models and community patterns Habitat models and community patterns. Multiple linear regression models revealed that deciduous forests and shrublands were the primary positive predictors of carnivore species richness (Tables 2, 3). In Model 1, habitat variables alone explained 73% of the variance (R² = 0.73). Richness increased with deciduous forest cover and declined in areas dominated by pine plantations and croplands. A 10% increase in deciduous forest corresponded to an average gain of 1.2 species, whereas the same increase in pine or cropland covers reduced richness by approximately one species. This pattern underscores the ecological importance of native oak woodlands ( Quercus pyrenaica, Q. ilex ) as biodiversity refuges, while intensively managed pine forests offer limited structural diversity and prey availability due to shrub removal and silvicultural disturbance (Ramos Santos, 2006; Ferreras et al., 2016). Model 2, which included shrub cover and wolf presence, explained 71% of the variance (R² = 0.71). Both deciduous and shrub habitats were positively associated with richness, while wolf presence had a slight negative, though non-significant, effect (p = 0.22). This suggests that the Iberian wolf may exert moderate top-down pressure on mesocarnivores, but the high spatial overlap within the reserve prevents clear segregation patterns. Similar findings have been reported in Mediterranean landscapes where apex predators regulate smaller carnivores without completely excluding them (Soulé et al., 1988; Peterson, 1995; Blanco, 2017). Unexpectedly, anthropogenic land cover did not significantly affect species richness, likely because urban and infrastructure elements were scarce and evenly distributed. The high heterogeneity and low human density of the Sierra likely buffer these effects compared to more fragmented Iberian regions (Rosalino et al., 2008). 3.2 Habitat selection by species At the species level, significant selection patterns were detected for the pine marten, common genet, and Eurasian otter (Figures 3, 5). The pine marten used pastures (H2) more frequently than expected and avoided shrublands (H3) (χ² = 224.5, p < 0.001). This atypical pattern may reflect competitive displacement by the stone marten ( Martes foina ), which dominates forested habitats (Erlinge & Sandell, 1986; Pilot et al., 2007). The common genet preferred croplands (H1) and avoided coniferous forests (H5) (χ² = 17.8, p < 0.03). Genets exploit arthropods, small mammals, and dung beetles abundant in grazed fields, aligning with previous observations that link genet abundance to livestock-associated habitats (Virgós & Casanovas, 1997; Rosalino & Santos-Reis, 2002). The Eurasian otter selected deciduous forest (H4) corridors along small streams and avoided shrub formations (H3) (χ² = 10.9, p < 0.05). Riparian deciduous stands thus represent essential habitat elements for semi-aquatic predators (Clavero et al., 2006; Lanszki et al., 2008). Other species displayed no significant differences between used and available habitats, suggesting generalist or overlapping habitat use at the scale considered. 3.3 Predator hierarchy and interspecific coexistence Predator hierarchy and interspecific coexistence. The broad coexistence of wolves and foxes in almost all sampling cells, together with the spatial overlap of martens, badgers, and genets, reflects a hierarchical but tolerant carnivore community. Wolves likely influence the spatial or temporal activity of mesocarnivores rather than their absolute occurrence, consistent with the concept of trophic modulation rather than strict exclusion (Donadio & Buskirk, 2006; Mech & Peterson, 2003). The coexistence of sympatric mustelids appears to be mediated by niche partitioning based on body size and feeding ecology (Erlinge & Sandell, 1986). Pine martens tend to exploit open areas or forest edges, while stone martens remain within human-modified woodlands and settlements (Clevenger, 1994). Such micro-scale segregation minimizes competition and facilitates coexistence, as documented in other European systems (Rosellini et al., 2008). 3.4 Functional implications for management and restoration The positive relationships between carnivore richness, deciduous forests, and shrublands emphasize that habitat heterogeneity is fundamental for sustaining trophic complexity. In contrast, intensive pine forestry and agricultural expansion contribute to ecological simplification and lower species diversity. Therefore, ecological restoration within the Sierra de la Culebra should prioritize: i) Recovery of native deciduous forests and natural shrub mosaics. ii) Reduction of silvicultural intensity in pine stands to allow shrub and understorey regeneration. iii) Control of invasive species, particularly the American mink, which poses risks to native riparian fauna such as the Eurasian otter (Dettori et al., 2021). The integration of non-invasive monitoring techniques, camera trapping, faecal DNA analysis, and local ecological knowledge, proved highly effective for assessing carnivore assemblages without disturbing wildlife. This methodological framework can serve as a model for biodiversity assessment and adaptive management in other Mediterranean protected areas. 3.5 Functional interpretation and synthesis The carnivore guild of the Sierra de la Culebra represents a functionally structured community where trophic interactions, habitat heterogeneity, and human disturbance jointly determine species coexistence. The persistence of the Iberian wolf ( Canis lupus signatus ) as a top predator plays a pivotal ecological role in maintaining this structure. By exerting top-down control on ungulate and mesocarnivore populations, wolves indirectly influence vegetation dynamics and the availability of carrion resources, contributing to what can be described as a trophic cascade in motion (Ripple & Beschta, 2012; Newsome et al., 2015). The moderate negative relationship observed between wolf presence and mesocarnivore richness supports the predator-mediated coexistence hypothesis: instead of excluding smaller species, apex predators regulate their density and spatial behavior, reducing exploitative competition among mesocarnivores and stabilizing community composition (Ritchie & Johnson, 2009). This hierarchical yet tolerant structure typifies resilient carnivore assemblages in which competitive and facilitative interactions coexist. At the same time, habitat heterogeneity emerges as the key bottom-up driver of diversity. The positive associations between species richness and deciduous or shrub habitats underline that structurally complex landscapes promote resource partitioning and niche complementarity (Tews et al., 2004). Conversely, intensive pine plantations and croplands reduce vertical and horizontal habitat complexity, homogenizing prey availability and limiting refuge opportunities. Therefore, trophic stability in Mediterranean systems depends on the combined maintenance of top-down regulation by large predators and bottom-up diversity generated by natural vegetation mosaics. From a restoration perspective, the Sierra de la Culebra exemplifies the dual need to conserve apex predators as ecological engineers and restore native vegetation as functional habitat. Actions that facilitate oak ( Quercus spp. ) and heathland regeneration, decrease understorey clearance in pine stands, and preserve riparian deciduous corridors will enhance the carrying capacity for a wide range of carnivores. The functional recovery of these habitats is also expected to strengthen ecosystem services such as carcass removal, rodent control, and seed dispersal via scavenger pathways. Furthermore, the methodological framework developed here, integrating camera trapping, genetic verification, and local ecological knowledge, constitutes a valuable tool for adaptive management. It allows long-term monitoring of carnivore responses to habitat restoration, forestry policies, and human-wildlife interactions, aligning conservation practice with the One Health and rewilding paradigms emerging across Europe (Perino et al., 2019; Svenning et al., 2016). In summary, the carnivore community of the Sierra de la Culebra exemplifies a resilient Mediterranean predator network in which functional diversity is maintained by the interplay of trophic regulation and landscape heterogeneity. Safeguarding both the wolf population and the structural diversity of habitats should remain central goals of regional management plans. By promoting coexistence rather than control, and by restoring ecological complexity rather than uniform productivity, managers can ensure the long-term integrity of one of the most emblematic carnivore systems in the Iberian Peninsula. 4. Concluding remarks The carnivore community of the Sierra de la Culebra is shaped by a combination of habitat structure, land-use history, and predator interactions. Deciduous forests and shrublands sustain higher diversity, while pine monocultures and croplands are associated with lower richness. The presence of the Iberian wolf contributes to maintaining a balanced trophic hierarchy without displacing mesocarnivores. In ecological and management terms, the preservation and restoration of natural vegetation, particularly native oak and heath formations, represent the key to conserving a functionally diverse carnivore community and reinforcing the ecological resilience of this emblematic Mediterranean landscape. Declarations ETHICAL STATEMENT This work did not require ethical clearance. CONFLICT OF INTEREST : The authors declare that this work does not report any conflict of interest. ETHICAL APPROVAL : This work complies with all ethical standards. INFORMED CONSENT : The authors give their consent to the use of their data and to the publication of this work. FUNDING: This work has not received any type of funding. Author Contribution All authors contributed equally to the fieldwork, laboratory analyses, data analysis, and generation of the final manuscript. Acknowledgement To my students Juan Sánchez, Iván Rebollo y Román Abela. A Javier Talegón for their knowledge of the Sierra. DATA AVAILABILITY STATEMENT: Not Applicable. References Balestrieri, A., Remonti, L., & Prigioni, C. (2009). Exploitation of food resources by the Eurasian badger (Meles meles) at the altitudinal limit of its Alpine range (NW Italy). Zoological science , 26(12), 821-827. DOI:10.2108/zsj.26.821 Barea-Azcón, J. M., Virgós, E., Ballesteros-Duperon, E., Moleón, M., & Chirosa, M. (2007). Surveying carnivores at large spatial scales: a comparison of four broad-applied methods. Biodiversity and Conservation , 16(4), 1213-1230. DOI:10.1007/s10531-006-9114-x Barrientos, L. M. (1995). El lobo, distribución, tamaño de población y estudio de su alimentación en la Sierra de la Culebra (Zamora, 1995). Junta de Castilla y León. Inédito. Barrientos, R., and Virgós, E. (2006). Reduction of potential food interference in two sympatric carnivores by sequential use of shared resources. Acta Oecologica 30,107–116. DOI:10.1016/j.actao.2006.02.006 Beier, C., Hansen, K., & Gundersen, P. (1993). Spatial variability of throughfall fluxes in a spruce forest. Environmental Pollution , 81(3), 257-267. https://doi.org/10.1016/0269-7491(93)90208-6 Begon, M., Harper, J. L., and Townsend, C. R. (Eds) (1986). ‘Ecology. Individuals, Populations and Communities.’ (Blackwell Scientific Publications: Oxford.). Blanco, J. C., & Salvador Milla, A. (2017). Lobo–Canis lupus Linnaeus, 1758. Bright, P. W., & Morris, P. A. (1994). Animal translocation for conservation: performance of dormice in relation to release methods, origin and season. Journal of Applied Ecology , 699-708. DOI:10.2307/2404160 Byers, C. R., Steinhorst, R. K., & Krausman, P. R. (1984). Clarification of a technique for analysis of utilization-availability data. T he Journal of wildlife management , 48(3), 1050-1053. Bunnell, F. L. (1987). Reproductive tactics of cervidae and their relationships to habitat. In RES. SYMP. NATL. ZOOL. PARK. (pp. 145-167). Centro de Información Geográfica Nacional (2018). CORINE Land Cover. En: Centro de descargas. Clavero, M., Prenda, J., & Delibes, M. (2006). Seasonal use of coastal resources by otters: comparing sandy and rocky stretches. Estuarine, Coastal and Shelf Science , 66(3-4), 387-394. DOI:10.1016/j.ecss.2005.09.011 Clevenger, A. P. (1994). Habitat characteristics of Eurasian pine martens Martes martes in an insular Mediterranean environment. Ecography , 17(3), 257-263. DOI:10.1111/j.1600-0587.1994.tb00101.x Crooks, K. R. (2002). Relative sensitivities of mammalian carnivores to habitat fragmentation. Conservation biology , 16(2), 488-502. DOI:10.1046/j.1523-1739.2002.00386.x Debano, S. J. (2006). Effects of livestock grazing on aboveground insect communities in semi-arid grasslands of southeastern Arizona. Biodiversity & Conservation , 15, 2547-2564. DOI:10.1007/s10531-005-2786-9 Debinski, D. M., & Holt, R. D. (2000). A survey and overview of habitat fragmentation experiments. Conservation biology , 14(2), 342-355. DOI:10.1046/j.1523-1739.2000.98081.x Delibes, M. (1982). Notas sobre la distribución pasada y actual del meloncillo Herpestes ichneumon (L.) en la Península Ibérica. Doñana Acta Vertebrata , 9, 341-352. Dettori, E. E., Balestrieri, A., Román, L. G., Zapata-Perez, V. M., Ruiu, A., Carcangiu, L., & Robledano-Aymerich, F. (2021). A cost-effective PCR–RFLP method for monitoring invasive American mink: preliminary field test in control areas of Spain. Conservation Genetics Resources , 13, 143-149. DOI:10.1007/s12686-020-01183-3 Dettori, E. E., Zapata-Pérez, V. M., Collados, D. B., Martínez, H. I., Balestrieri, A., & Aymerich, F. R. (2017). A non-invasive genetic survey of sympatric carnivorous mammals in a riparian corridor along the intensively cultivated valley of the Segura River (SE Spain). In III Jornadas Doctorales Escuela Internacional de Doctorado de la Universidad de Murcia (EIDUM) (pp. 1122-1123). Servicio de Publicaciones. Di Luzio, V. B. (2010). Risultati preliminari sul comportamento spaziale della lepre italica (lepus corsicanus) nella rnr monterano (RM). In: Workshop nazionale sulla conservazione della lepre itálica (p. 111). Erlinge, S., & Sandell, M. (1986). Seasonal changes in the social organization of male stoats, Mustela erminea: an effect of shifts between two decisive resources. Oikos , 57-62. Ferreras, P., Díaz-Ruiz, F., Alves, P. C., & Monterroso, P. S. (2016). Factores de la coexistencia de mesocarnívoros en parques nacionales de ambiente mediterráneo. Gese, E. M. (2001). Territorial defense by coyotes (Canis latrans) in Yellowstone National Park, Wyoming: who, how, where, when, and why. Canadian Journal of Zoology , 79(6), 980-987. DOI:10.1139/cjz-79-6-980 Kauhala, K., & Salonen, L. (2012). Does a non-invasive method-latrine surveys-reveal habitat preferences of raccoon dogs and badgers?. Mammalian Biology , 77, 264-270. DOI:10.1016/j.mambio.2012.02.007 Kucera, T. E., & Barrett, R. H. (1993). In my experience: the Trailmaster® camera system for detecting wildlife. Wildlife Society Bulletin, 505-508. Kohn, M. H., & Wayne, R. K. (1997). Facts from feces revisited. Trends in ecology & evolution , 12(6), 223-227. Lanszki, J., & Nagy, L. (2003). A Látrányi Puszta Természetvédelmi Terület gerinces (Vertebrata) faunájának felmérése-Investigation on vertebrata fauna of the Látrányi Puszta Nature Conservation Area. Natura Somogyiensis, (5), 279-290. DOI:10.24394/NatSom.2003.5.279 Lanszki, J., Hidas, A., Szentes, K., Révay, T., Lehoczky, I., & Weiss, S. (2008). Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in Hungary. Mammalian biology , 73(1), 40-47. DOI:10.1016/j.mambio.2007.08.005 Leonard, J. A., Vila, C., & Wayne, R. K. (2005). FAST TRACK: Legacy lost: genetic variability and population size of extirpated US grey wolves (Canis lupus). Molecular Ecology, 14(1), 9-17. DOI:10.1111/j.1365-294X.2004.02389.x Libois, R., & Waechter, A. (1991). La fouine (Martes foina Erxleben, 1777. Société française pour l'étude et la protection des mammifères, Bourges, France. Manly, B. F. (1993). Comments on design and analysis of multiple-choice feeding-preference experiments. Oecologia , 149-152. Martínez, F. A., & Pastor-Alfonso, M. J. (2019). El turismo lobero en la Sierra de La Culebra (España). Revista LIDER , 21(35), 137-160. DOI:10.32735/S0719-526520193513 Mech, L. D. (1966). The Wolves of isle royale (No. 7). US Government Printing Office. Mech, L. D., & Peterson, R. O. (2003). Wolf-prey relations. Mendoza, M., Janis, C. M., & Palmqvist, P. (2005). Ecological patterns in the trophic-size structure of large mammal communities: a ‘taxon-free’characterization. Evolutionary Ecology Research , 7(4), 505-530. Núñez, I. B. (2005). Patrones de marcaje con heces por la marta europea (Martes martes) en el noroeste de España: importancia para su estudio. Galemys: Boletín informativo de la Sociedad Española para la conservación y estudio de los mamíferos, 17(1), 123-134. DOI:10.7325/Galemys.2005.NE.A10 Lira-Torres, I., & Briones-Salas, M. (2012). Abundancia relativa y patrones de actividad de los mamíferos de los Chimalapas, Oaxaca, México. Acta zoológica mexicana , 28(3), 566-585. DOI:10.21829/azm.2012.283859 Mortelliti, A., Amori, G., & Boitani, L. (2010). The role of habitat quality in fragmented landscapes: a conceptual overview and prospectus for future research. Oecologia , 163(2), 535-547. DOI:10.1007/s00442-010-1623-3 Pilot, M., Gralak, B., Goszczyński, J., & Posłuszny, M. (2007). A method of genetic identification of pine marten (Martes martes) and stone marten (Martes foina) and its application to faecal samples. Journal of Zoology , 271(2), 140-147. DOI:10.1111/j.1469-7998.2006.00179.x Neu, C. W., Byers, C. R., & Peek, J. M. (1974). A technique for analysis of utilization-availability data. The Journal of Wildlife Management, 541-545. Newsome, T. M., Dellinger, J. A., Pavey, C. R., Ripple, W. J., Shores, C. R., Wirsing, A. J., & Dickman, C. R. (2015). The ecological effects of providing resource subsidies to predators. Global Ecology and Biogeography, 24(1), 1-11. https://doi.org/10.1111/geb.12236 Palacios, J. y J. L. Vicente (1997). Gestión de Fauna en la Reserva Nacional de Caza “Sierra de la Culebra”. Pp. 18/1-18/15. En: Master Universitario en Veterinaria y Fauna Salvaje; Tomo IV: Gestión Cinegética y de Especies Protegidas. Colegio Oficial de Veterinarios de Zamora e I.C.E. Universidad de León. Zamora. Palomares, F., Ferreras, P., Fedriani, J. M., and Delibes, M. (1996). Spatial relationships between Iberian lynx and other carnivores in an area of south-west Spain. Journal of Applied Ecology 33, 5–13. Perino, A., Pereira, H. M., Navarro, L. M., Fernández, N., Bullock, J. M., Ceaușu, S., ... & Svenning, J. C. (2019). Rewilding complex ecosystems. Science , 364(6438), eaav5570. https://doi.org/10.1126/science.aav5570 Peterson, R. O. (1995). Wolves as interspecific competitors in canid ecology. Ecology and conservation of wolves in a changing world, 315-324. Putman, R. J. (1984). Facts from faeces. Mammal review , 14(2), 79-97. Ramos Santos, J. M. (2006). La transformación del paisaje vegetal y los aprovechamientos forestales tradicionales en la Sierra de la Culebra (Zamora). Investigaciones geográficas, nº 40, 55-72. Remonti, L., Balestrieri, A., Ruiz-González, A., Gómez-Moliner, B. J., Capelli, E., & Prigioni, C. (2012). Intraguild dietary overlap and its possible relationship to the coexistence of mesocarnivores in intensive agricultural habitats. Population Ecology , 54, 521-532. DOI:10.1007/s10144-012-0337-2 Ripple, W. J., & Beschta, R. L. (2012). Trophic cascades in Yellowstone: The first 15 years after wolf reintroduction. Biological Conservation , 145(1), 205–213. https://doi.org/10.1016/j.biocon.2011.11.005 Ritchie, E. G., & Johnson, C. N. (2009). Predator interactions, mesopredator release and biodiversity conservation. Ecology Letters , 12(9), 982–998. https://doi.org/10.1111/j.1461-0248.2009.01347.x Rodríguez-Mazzini, R. (1996). Uso de la técnica de estaciones olfativas (SCENT-STATION TECHNIQUE) en estudios de ecología de mamíferos (No. 599.74 ROD). Rosalino, L. M., & Santos-Reis, M. (2002). Feeding habits of the common genet Genetta genetta (Carnivora: Viverridae) in a semi-natural landscape of central Portugal. Rosalino, L. M., Santos, M. J., Beier, P., & Santos-Reis, M. (2008). Eurasian badger habitat selection in Mediterranean environments: does scale really matter?. Mammalian Biology , 73(3), 189-198. https://doi.org/10.1016/j.mambio.2007.02.004 Rosellini, S., Osorio, E., Ruiz-González, A., Piñeiro, A., & Barja, I. (2008). Monitoring the small-scale distribution of sympatric European pine martens (Martes martes) and stone martens (Martes foina): a multievidence approach using faecal DNA analysis and camera-traps. Wildlife research , 35(5), 434-440. DOI:10.1071/wr07030 Ruiz-González, A., Rubines, J., Berdión, O., & Gómez-Moliner, B. J. (2008). A non-invasive genetic method to identify the sympatric mustelids pine marten (Martes martes) and stone marten (Martes foina): preliminary distribution survey on the northern Iberian Peninsula. European Journal of Wildlife Research , 54, 253-261. DOI:10.1007/s10344-007-0138-7 Sambrook, J., Fritsch, E. F., & Maniatis, T. (1989). Molecular cloning: a laboratory manual (No. Ed. 2). Cold spring harbor laboratory press. Sanz, B. y Turón, J.V. (2017). Guía de mamíferos terrestres. Península ibérica y balear. Ed. Prames. Zaragoza. Soulé, M. E., Bolger, D. T., Alberts, A. C., Wrights, J., Sorice, M., & Hill, S. (1988). Reconstructed dynamics of rapid extinctions of chaparral‐requiring birds in urban habitat islands. Conservation Biology , 2(1), 75-92. https://doi.org/10.1111/j.1523-1739.1988.tb00337.x Suárez-Tangil, B. D., & Rodríguez, A. (2021). Estimates of species richness and composition depend on detection method in assemblages of terrestrial mammals. Animals , 11(1), 186. DOI:10.3390/ani11010186 Svenning, J. C., Pedersen, P. B., Donlan, C. J., Ejrnæs, R., Faurby, S., Galetti, M., ... & Vera, F. W. (2016). Science for a wilder Anthropocene: Synthesis and future directions for trophic rewilding research. Proceedings of the National Academy of Sciences , 113(4), 898-906. https://doi.org/10.1073/pnas.1502556112 Tews, J., Brose, U., Grimm, V., Tielbörger, K., Wichmann, M. C., Schwager, M., & Jeltsch, F. (2004). Animal species diversity driven by habitat heterogeneity/diversity: The importance of keystone structures. Journal of Biogeography , 31(1), 79–92. https://doi.org/10.1046/j.0305-0270.2003.00994 Torre, I., Arrizabalaga, A. y Flaquer, C. (2003). Estudio de la distribución y abundancia de carnívoros en el Parque Natural del Montnegre y El Corredor mediante trampeo fotográfico. Galemys , 15 (1), 15-28. DOI:10.7325/Galemys.2003.1.A2 Vicente, J. L., Rodríguez, M., & Palacios, J. (2000). Gestión del lobo ibérico (Canis lupus signatus Cabrera, 1907) en la Reserva Regional de Caza “Sierra de la Culebra”(Zamora). Galemys , 12(1), 181-199. Vilà, C., Amorim, I. R., Leonard, J. A., Posada, D., Castroviejo, J., Petrucci‐Fonseca, F., ... & Wayne, R. K. (1999). Mitochondrial DNA phylogeography and population history of the grey wolf Canis lupus. Molecular ecology , 8(12), 2089-2103. DOI:10.1046/j.1365-294x.1999.00825.x Virgós, E., & Casanovas, J. G. (1997). Habitat selection of genet Genetta genetta in the mountains of central Spain. Acta Theriologica , 42(2), 169-177. DOI:10.7325/Galemys.2007.NE.A6 Waldron, J. L., Bennett, S. H., Welch, S. M., Dorcas, M. E., Lanham, J. D., & Kalinowsky, W. (2006). Habitat specificity and home‐range size as attributes of species vulnerability to extinction: a case study using sympatric rattlesnakes. Animal conservation, 9(4), 414-420. DOI:10.1111/j.1469-1795.2006.00050.x White, G. C., & Garrott, R. A. (2012). Analysis of wildlife radio-tracking data. Elsevier. Zielinski, W. J., & Kucera, T. E. (1995). American marten, fisher, lynx, and wolverine: survey methods for their detection (Vol. 157). Pacific Southwest Research Station. https://doi.org/10.2737/PSW-GTR-157 Additional Declarations No competing interests reported. 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1","display":"","copyAsset":false,"role":"figure","size":229021,"visible":true,"origin":"","legend":"\u003cp\u003eStudy area (re-elaboration of available data from the Spanish National Geographic Institute).\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/40e4c21d08bc8b604ab244ec.jpg"},{"id":96611192,"identity":"f37bad6c-e9eb-4d6f-baa9-9a06c11cd484","added_by":"auto","created_at":"2025-11-24 09:33:02","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":183822,"visible":true,"origin":"","legend":"\u003cp\u003eSampling grid (7.5 x 7.5 km) in the Sierra de la Culebra.\u003c/p\u003e","description":"","filename":"2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/8a27581b8b6028b1a8663d1c.jpg"},{"id":96708163,"identity":"c6701f11-22c7-47db-9c51-2dddbb24ac1e","added_by":"auto","created_at":"2025-11-25 09:58:26","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":113785,"visible":true,"origin":"","legend":"\u003cp\u003eLinear adjustment of the variables that intervene in the model with the species’ richness.\u003c/p\u003e","description":"","filename":"3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/089b33d02b144b6b22cd494f.jpg"},{"id":96611244,"identity":"71b17a98-4c76-42c3-b5ab-416e2103d859","added_by":"auto","created_at":"2025-11-24 09:33:17","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":24220,"visible":true,"origin":"","legend":"\u003cp\u003eObserved (use) and expected (availability) proportion of use of the major habitats by the pine marten, showing Bonferroni’s confidence intervals.\u003c/p\u003e","description":"","filename":"4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/863805893474ec191995cbe6.jpg"},{"id":96611170,"identity":"4835b80d-1c8a-4ad0-8e9e-0c1f35ee30fe","added_by":"auto","created_at":"2025-11-24 09:32:59","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":23527,"visible":true,"origin":"","legend":"\u003cp\u003eObserved (use) and expected (availability) proportion of use of the major habitats by the common genet, showing Bonferroni’s confidence intervals.\u003c/p\u003e","description":"","filename":"5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/fc7b72272b10dddcca0eeaba.jpg"},{"id":96611158,"identity":"03144b90-6105-4497-9d04-6cb79f0bf0a3","added_by":"auto","created_at":"2025-11-24 09:32:53","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":24780,"visible":true,"origin":"","legend":"\u003cp\u003eObserved (use) and expected (availability) proportion of use of the major habitats by the Eurasian otter, showing Bonferroni’s confidence intervals.\u003c/p\u003e","description":"","filename":"6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/dee4a240613450069a61b57d.jpg"},{"id":102321099,"identity":"8f2f01b3-de27-4cc7-ba0c-3f84145ad8b5","added_by":"auto","created_at":"2026-02-10 13:43:17","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1374494,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8029414/v1/be5ed018-56da-4cac-a251-0c0a95368398.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Trophic structure and habitat determinants of a Mediterranean carnivore guild: insights from the Sierra de la Culebra (NW Spain)","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eHabitat fragmentation and landscape transformation are among the most pervasive drivers of biodiversity loss worldwide. habitats that require large, continuous areas of natural habitat are particularly vulnerable to these changes, as fragmentation leads to isolation, reduced gene flow, and altered community dynamics (Beier et al., 1993; Mortelliti et al., 2010). Mammalian carnivores are especially sensitive to habitat reduction because of their large home ranges and high trophic position, which make them reliable indicators of ecosystem integrity (Crooks, 2002; Reed, 2004).In Mediterranean environments, the progressive replacement of native woodlands and shrublands by pine plantations, crops, and infrastructures has reshaped ecological networks and modified the structure of predator communities (Waldron et al., 2006; Debinski \u0026amp; Holt, 2000). While some species exhibit high tolerance to human-modified landscapes, others show pronounced declines, making comparative studies of habitat selection essential for conservation and restoration planning. The Sierra de la Culebra, located in northwestern Zamora (Castilla y Le\u0026oacute;n, Spain), represents one of the most emblematic Mediterranean mountain ranges in the Iberian Peninsula. Covering approximately 65,000 ha, this Regional Hunting Reserve and UNESCO Biosphere Reserve is internationally known for hosting one of the densest populations of Iberian wolves (\u003cem\u003eCanis lupus signatus\u003c/em\u003e) in Western Europe (Vicente et al., 2000). The area comprises a mosaic of habitats (heathlands, pine forests (\u003cem\u003ePinus pinaster, P. sylvestris\u003c/em\u003e), oak woodlands (\u003cem\u003eQuercus pyrenaica, Q. ilex\u003c/em\u003e), pastures, and croplands) shaped by centuries of silviculture, grazing, and hunting (Ramos Santos, 2006; Hidalgo, 2008). Despite its ecological importance, few studies have addressed the structure of the carnivore community as a whole or the relationships between top predators and mesocarnivores in this unique landscape. Understanding habitat preferences and interspecific interactions within the carnivore guild of the Sierra de la Culebra is essential for developing effective management and ecological restoration strategies. Habitat selection models can provide insights into how vegetation composition, land-use intensity, and predator presence influence species richness and distribution patterns. The objectives of this study were therefore to (i) assess the species composition and relative abundance of carnivores in the Sierra de la Culebra using complementary non-invasive methods (camera trapping, faecal DNA analysis, and interviews), (ii) identify habitat preferences and key environmental predictors of carnivore richness, and (iii) evaluate the influence of top predators (wolves) and exotic species (American mink) on community structure. We hypothesized that (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e) carnivore richness would be higher in deciduous forests and shrublands than in pine plantations and croplands, due to the greater structural complexity and prey availability of natural habitats. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e) The presence of the top predator, the Iberian wolf (\u003cem\u003eCanis lupus signatus\u003c/em\u003e), would moderately reduce mesocarnivore richness through interspecific competition and top-down regulation. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e) The integration of complementary non-invasive methods, camera trapping, faecal DNA analysis, and interviews, would provide a more comprehensive and accurate assessment of community composition than any single approach alone.\u003c/p\u003e"},{"header":"2. Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1. Study area\u003c/h2\u003e\u003cp\u003eThe study was conducted in the Sierra de la Culebra Regional Hunting Reserve (Zamora, Castilla y Le\u0026oacute;n, NW Spain), an area also designated as a UNESCO Biosphere Reserve. The reserve extends over approximately 65,000 ha, ranging from 800 to 1,243 m a.s.l., with Pe\u0026ntilde;a Mira as its highest peak (Vicente et al., 2000). The mountain range is composed of parallel quartzite ridges-oriented NNW-SSE and intersected by small fluvial valleys (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Its natural limits are the River Tera to the north and the River Aliste to the south, while it borders the region of T\u0026aacute;bara in the east and the foothills of the Sierra Segundera in the west (Ramos Santos, 2006). Both the structure and composition of the vegetation have been substantially modified by human activities. The autochthonous deciduous forests, mainly \u003cem\u003eQuercus pyrenaica\u003c/em\u003e, Q. ilex subsp. rotundifolia and \u003cem\u003eQ. faginea\u003c/em\u003e, are now interspersed with cultivated lands and large pine plantations (\u003cem\u003ePinus pinaster, P. sylvestris, P. nigra\u003c/em\u003e) established for timber production. Heather (\u003cem\u003eErica australis, E. arborea, E. umbellata\u003c/em\u003e) and Cistus shrubs (\u003cem\u003eC. laurifolius, C. ladanifer\u003c/em\u003e) cover approximately 55.7% of the study area, followed by pine forests (22.7%), deciduous woodlands (5.7%), pastures (7.9%) and croplands (7.7%) (Centro Nacional de Informaci\u0026oacute;n Geogr\u0026aacute;fica, 2018). The local fauna is highly diverse, including at least six fish, fourteen amphibians, fourteen reptiles, twenty-nine mammals, and over one hundred bird species (Vicente et al., 2000).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2. Sampling design\u003c/h2\u003e\u003cp\u003eFieldwork was carried out between March and June 2019. The reserve was divided into twelve 7.5 \u0026times; 7.5 km grid cells (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), each covering approximately 5,625 ha. Within each grid, seven camera-trapping stations were established, giving a total of 84 stations across the study area. Cameras (Campark models equipped with 44 infrared LEDs and IR filters) were active 24 h per day for seven consecutive days at each station (Zielinski \u0026amp; Kucera, 1995; Torres et al., 2003; Barea-Azc\u0026oacute;n et al., 2007). To increase the detection probability of mesocarnivores, each station was baited with sardines in oil and lynx urine as olfactory attractants (Ferreras et al., 2016). Total sampling effort amounted to 588 camera-days. In parallel, faecal surveys were conducted along linear transects of approximately 3 km per grid cell (Rodr\u0026iacute;guez-Mazzini, 1996), each surveyed twice. A total of 253 faecal samples was collected and preserved in autoclaved tubes containing 96% ethanol, then stored at \u0026minus;\u0026thinsp;20\u0026deg;C until genetic analysis. During fieldwork, all tracks and visual records of carnivores were noted following the identification criteria of Sanz \u0026amp; Tur\u0026oacute;n (2017). Geographic coordinates of every detection were recorded with a Garmin GPS. Finally, 21 structured interviews were conducted with forest agents and hunting wardens working in the reserve to obtain complementary information on carnivore presence and activity patterns.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\u003ch2\u003e2.3 Laboratory procedures\u003c/h2\u003e\u003cp\u003eDNA was extracted from faecal samples using the FavorPrep Mini Stool DNA Isolation Kit (Favorgen Biotech Corp.) according to the manufacturer\u0026rsquo;s protocol and the general molecular procedures of Sambrook et al. (1989). Species identification focused on mustelids and other small carnivores was achieved by a PCR-RFLP approach following Ruiz-Gonz\u0026aacute;lez et al. (2008). Amplified mtDNA fragments were digested simultaneously with RsaI and BsuRI, generating species-specific restriction patterns that allowed unequivocal discrimination of sympatric species such as \u003cem\u003eMartes martes\u003c/em\u003e, \u003cem\u003eMartes foina\u003c/em\u003e, Neogale vison and Lutra lutra (Ruiz-Gonz\u0026aacute;lez et al., 2008; Dettori et al., 2021). PCR reactions were run on a GeneAmp PCR System 2700 (Applied Biosystems). Amplicons were sequenced at Macrogen Europe (Amsterdam, Netherlands) using an ABI Prism 3730XL analyzer. The resulting sequences were aligned in GENEIOUS v7.1.7 (Biomatters Ltd.) and compared with homologous sequences from the GenBank database using the BLAST algorithm of the National Center for Biotechnology Information (NCBI).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003e2.4 Habitat classification\u003c/h2\u003e\u003cp\u003eHabitat information was derived from the CORINE Land Cover database (Centro Nacional de Informaci\u0026oacute;n Geogr\u0026aacute;fica, 2018) and processed in QGIS v3.14. Each 7.5 \u0026times; 7.5 km grid was overlaid with the land-cover map, and the proportional area (%) of each of seven major habitat categories was calculated (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e): H1\u0026thinsp;=\u0026thinsp;Crops, H2\u0026thinsp;=\u0026thinsp;Pastures, H3\u0026thinsp;=\u0026thinsp;Shrublands, H4\u0026thinsp;=\u0026thinsp;Deciduous forests, H5\u0026thinsp;=\u0026thinsp;Coniferous forests, H6\u0026thinsp;=\u0026thinsp;Watercourses, H7\u0026thinsp;=\u0026thinsp;Anthropic elements. Two additional categorical variables were included: H8\u0026thinsp;=\u0026thinsp;presence of wolf home ranges (1\u0026thinsp;=\u0026thinsp;absent, 2\u0026thinsp;=\u0026thinsp;partially overlapping, 3\u0026thinsp;=\u0026thinsp;fully overlapping) and H9\u0026thinsp;=\u0026thinsp;presence/absence of the invasive American mink (\u003cem\u003eNeogale vison\u003c/em\u003e) (0/1) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These variables were later incorporated into multiple linear regression models to assess how habitat composition, predator presence, and alien species influenced carnivore richness across the grid cells.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eHabitat variables, obtained by merging Corine Land Covers (Centro Nacional de Informaci\u0026oacute;n Geogr\u0026aacute;fica, 2018).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"3\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCorine Land Cover\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCode\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLand\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e211\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDry farmland\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e221\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eVineyards\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eH1-Crops\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e242\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCrop mosaic\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e243\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMainly agricultural land, but with important spaces of natural vegetation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e333\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSpaces with sparse vegetation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e\u003cb\u003eH2-Pastures\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e231\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eGrasslands\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e321\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNatural grasslands\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003e\u003cb\u003eH3-Shrubs\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e322\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMoors and scrub\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e323\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSclerophyll vegetation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e324\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTransitional scrub forest\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e\u003cb\u003eH4-Deciduous forest\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e311\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLeafy woods\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e313\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMixed forest\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eH5-Conifers\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e312\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eConiferous forests\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003e\u003cb\u003eH6-Water courses\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e511\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eWater courses\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e512\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eWater sheets\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e112\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDiscontinuous urban areas\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eH7-Anthropic elements\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e131\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eMining extraction areas\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e132\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDumps and landfills\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e133\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eArea under construction\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eH1\u0026thinsp;=\u0026thinsp;Crops; H2\u0026thinsp;=\u0026thinsp;Pastures; H3\u0026thinsp;=\u0026thinsp;Scrub; H4\u0026thinsp;=\u0026thinsp;Deciduous forest; H5\u0026thinsp;=\u0026thinsp;Coniferous forest; H6\u0026thinsp;=\u0026thinsp;Bodies of water; H7\u0026thinsp;=\u0026thinsp;Anthropic elements; P/A H8\u0026thinsp;=\u0026thinsp;Presence/Absence of Wolf and P/A H9\u0026thinsp;=\u0026thinsp;Presence/Absence of American mink. In the P/A H8 column, the value 1\u0026thinsp;=\u0026thinsp;No wolf vital domain present; 2\u0026thinsp;=\u0026thinsp;Presence shared life domain between grids; 3\u0026thinsp;=\u0026thinsp;Vital domain exclusive presence on grid. In the P/A H9 column, the value 0\u0026thinsp;=\u0026thinsp;Absence of mink; 1\u0026thinsp;=\u0026thinsp;presence of mink.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"11\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGrids\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRichness\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e%\u003c/p\u003e\u003cp\u003eH1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e%\u003c/p\u003e\u003cp\u003eH2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e%\u003c/p\u003e\u003cp\u003eH3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e%\u003c/p\u003e\u003cp\u003eH4\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e%\u003c/p\u003e\u003cp\u003eH5\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e% H6\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e% H7\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003eP/A H8\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c11\"\u003e\u003cp\u003eP/A H9\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7,91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e11,87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e62,91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1,12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e14,27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e1,92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e2\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1,09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2,45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e42,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2,94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e50,17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e1,33\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e3\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3,47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e22,09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e48,99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e15,91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1,78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e2,93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e4,83\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e4\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0,09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6,85\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e56,99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e3,06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e31,74\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1,09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e0,18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e5\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2,84\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2,53\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e88,54\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2,09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e3,94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e0,07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e6\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e4,43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4,48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e75,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2,72\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e11,18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e1,99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e7\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3,36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6,40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e57,65\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e12,81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e18,45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e0,95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e8\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0,50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5,53\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e61,42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e7,85\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e20,70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e4,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e9\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9,63\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e10,66\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e50,40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2,31\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e25,72\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e1,27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e10\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e24,89\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e11,02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e35,46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0,78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e27,79\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e0,05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e1,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e11\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e12,65\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6,90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e42,36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e6,19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e28,38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e3,52\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e12\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e21,77\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4,20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e47,34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e10,90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e15,42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e0,37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2,00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e\u003cp\u003e0,00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSummary of the 1st explanatory model of the carnivores richness in the Sierra de la Culebra.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e\u003cp\u003eFirst model\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eCoefficient\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eSig (p)\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eR\u0026sup2;\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eR\u0026sup2; (adjusted)\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eConstant\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e9,27205\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,0000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e0,7307\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e0,6297\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePine grove\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0,10826\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,0124\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDeciduous forest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0,12684\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,1873\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCrops\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0,07593\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,1651\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSummary of the 2nd model used to determine the richness of carnivores in the Sierra de la Culebra.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e\u003cp\u003eSecond model\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eCoefficient\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eSig (p)\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eR\u0026sup2;\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eR\u0026sup2; (adjusted)\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eConstant\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2,68575\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,2923\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e0,7107\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003e0,622\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eShrub\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0,10407\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,0105\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDeciduous forest\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0,23761\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,0361\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eWolf presence\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-1,03685\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,2246\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\u003ch2\u003e2.5 Data analysis\u003c/h2\u003e\u003cp\u003eHabitat selection: To compare habitat use at the transect scale, faecal samples were assigned to one of the seven habitat types defined above. Habitat selection was tested using Chi-squared (χ\u0026sup2;) goodness-of-fit tests (White \u0026amp; Garrott, 1990; Balestrieri et al., 2015), with expected frequencies based on the relative length of transects in each habitat. Significant selection or avoidance was evaluated using Bonferroni confidence intervals (Neu et al., 1974; Byers et al., 1984; Manly et al., 1993; Di Luzio, 2010; Dettori et al., 2017). Community-level models: At the community level, species richness per grid cell was modeled through multiple linear regression to determine the effects of land-cover variables (H1-H7) and biotic predictors (H8-H9). Two models were tested: i) Model 1 included only habitat variables (percent cover of pine forest, deciduous forest, cropland). ii) Model 2 added shrub cover and wolf presence to evaluate biotic modulation of richness. Model performance was evaluated by R\u0026sup2; and adjusted R\u0026sup2;, and predictor significance was assessed at p\u0026thinsp;\u0026lt;\u0026thinsp;0.05. Normality of residuals and multicollinearity were examined before interpretation. All statistical analyses were performed in SPSS v13.0 (IBM Corp., Armonk, NY, USA). Graphical outputs and spatial visualizations were prepared in QGIS and Microsoft Excel.\u003c/p\u003e\u003c/div\u003e\n\u003cp\u003e\u003cem\u003e2.6 Ethical statement\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAll field procedures were non-invasive and complied with Spanish wildlife protection regulations (Law 4/1996, Castilla y Le\u0026oacute;n). Sampling and data collection were conducted with the authorization of the Regional Hunting Reserve authorities.\u003c/p\u003e"},{"header":"3. Results and discussion","content":"\u003cp\u003eCarnivore community composition. A total of twelve carnivore species were detected in the Sierra de la Culebra during the 2019 survey: red fox (\u003cem\u003eVulpes vulpes\u003c/em\u003e), Iberian wolf (\u003cem\u003eCanis lupus signatus\u003c/em\u003e), common genet (\u003cem\u003eGenetta genetta\u003c/em\u003e), stone marten (\u003cem\u003eMartes foina\u003c/em\u003e), pine marten (\u003cem\u003eMartes martes\u003c/em\u003e), Eurasian badger (\u003cem\u003eMeles meles\u003c/em\u003e), European polecat (\u003cem\u003eMustela putorius\u003c/em\u003e), least weasel (\u003cem\u003eMustela nivalis\u003c/em\u003e), wildcat (\u003cem\u003eFelis silvestris\u003c/em\u003e), Egyptian mongoose (\u003cem\u003eHerpestes ichneumon\u003c/em\u003e), Eurasian otter (\u003cem\u003eLutra lutra\u003c/em\u003e), and American mink (\u003cem\u003eNeogale vison\u003c/em\u003e). This diverse assemblage includes both apex predators and mesocarnivores, confirming that the Sierra de la Culebra maintains a functionally complete predator guild (Vicente et al., 2000). Camera trapping, faecal DNA analysis, and interviews proved complementary. Wolves and red foxes were the most ubiquitous species, detected in all twelve grid cells, followed by pine and stone martens (11 grids), badger and genet (10), while the American mink and wildcat appeared in 5 grids. The polecat, otter, weasel, and mongoose were recorded only in 2-4 grids, mostly near riparian environments. The total camera-trapping effort was 588 camera-days, yielding a capture success of 37.4% (37.4 detections/100 camera-days). American mink, red fox, and genet showed the highest activity indices. These results highlight that species with generalist or semi-aquatic habits were the most detectable through camera and bait-based methods (Lira-Torres \u0026amp; Briones-Salas, 2012).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.1 Habitat models and community patterns\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eHabitat models and community patterns. Multiple linear regression models revealed that deciduous forests and shrublands were the primary positive predictors of carnivore species richness (Tables 2, 3). In Model 1, habitat variables alone explained 73% of the variance (R\u0026sup2; = 0.73). Richness increased with deciduous forest cover and declined in areas dominated by pine plantations and croplands. A 10% increase in deciduous forest corresponded to an average gain of 1.2 species, whereas the same increase in pine or cropland covers reduced richness by approximately one species. This pattern underscores the ecological importance of native oak woodlands (\u003cem\u003eQuercus pyrenaica, Q. ilex\u003c/em\u003e) as biodiversity refuges, while intensively managed pine forests offer limited structural diversity and prey availability due to shrub removal and silvicultural disturbance (Ramos Santos, 2006; Ferreras et al., 2016). Model 2, which included shrub cover and wolf presence, explained 71% of the variance (R\u0026sup2; = 0.71). Both deciduous and shrub habitats were positively associated with richness, while wolf presence had a slight negative, though non-significant, effect (p = 0.22). This suggests that the Iberian wolf may exert moderate top-down pressure on mesocarnivores, but the high spatial overlap within the reserve prevents clear segregation patterns. Similar findings have been reported in Mediterranean landscapes where apex predators regulate smaller carnivores without completely excluding them (Soul\u0026eacute; et al., 1988; Peterson, 1995; Blanco, 2017). Unexpectedly, anthropogenic land cover did not significantly affect species richness, likely because urban and infrastructure elements were scarce and evenly distributed. The high heterogeneity and low human density of the Sierra likely buffer these effects compared to more fragmented Iberian regions (Rosalino et al., 2008).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.2 Habitat selection by species\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAt the species level, significant selection patterns were detected for the pine marten, common genet, and Eurasian otter (Figures 3, 5). The pine marten used pastures (H2) more frequently than expected and avoided shrublands (H3) (\u0026chi;\u0026sup2; = 224.5, p \u0026lt; 0.001). This atypical pattern may reflect competitive displacement by the stone marten (\u003cem\u003eMartes foina\u003c/em\u003e), which dominates forested habitats (Erlinge \u0026amp; Sandell, 1986; Pilot et al., 2007). The common genet preferred croplands (H1) and avoided coniferous forests (H5) (\u0026chi;\u0026sup2; = 17.8, p \u0026lt; 0.03). Genets exploit arthropods, small mammals, and dung beetles abundant in grazed fields, aligning with previous observations that link genet abundance to livestock-associated habitats (Virg\u0026oacute;s \u0026amp; Casanovas, 1997; Rosalino \u0026amp; Santos-Reis, 2002). The Eurasian otter selected deciduous forest (H4) corridors along small streams and avoided shrub formations (H3) (\u0026chi;\u0026sup2; = 10.9, p \u0026lt; 0.05). Riparian deciduous stands thus represent essential habitat elements for semi-aquatic predators (Clavero et al., 2006; Lanszki et al., 2008). Other species displayed no significant differences between used and available habitats, suggesting generalist or overlapping habitat use at the scale considered.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.3 Predator hierarchy and interspecific coexistence\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003ePredator hierarchy and interspecific coexistence. The broad coexistence of wolves and foxes in almost all sampling cells, together with the spatial overlap of martens, badgers, and genets, reflects a hierarchical but tolerant carnivore community. Wolves likely influence the spatial or temporal activity of mesocarnivores rather than their absolute occurrence, consistent with the concept of trophic modulation rather than strict exclusion (Donadio \u0026amp; Buskirk, 2006; Mech \u0026amp; Peterson, 2003). The coexistence of sympatric mustelids appears to be mediated by niche partitioning based on body size and feeding ecology (Erlinge \u0026amp; Sandell, 1986). Pine martens tend to exploit open areas or forest edges, while stone martens remain within human-modified woodlands and settlements (Clevenger, 1994). Such micro-scale segregation minimizes competition and facilitates coexistence, as documented in other European systems (Rosellini et al., 2008).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.4 Functional implications for management and restoration\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe positive relationships between carnivore richness, deciduous forests, and shrublands emphasize that habitat heterogeneity is fundamental for sustaining trophic complexity. In contrast, intensive pine forestry and agricultural expansion contribute to ecological simplification and lower species diversity. Therefore, ecological restoration within the Sierra de la Culebra should prioritize: i) Recovery of native deciduous forests and natural shrub mosaics. ii) Reduction of silvicultural intensity in pine stands to allow shrub and understorey regeneration. iii) Control of invasive species, particularly the American mink, which poses risks to native riparian fauna such as the Eurasian otter (Dettori et al., 2021). The integration of non-invasive monitoring techniques, camera trapping, faecal DNA analysis, and local ecological knowledge, proved highly effective for assessing carnivore assemblages without disturbing wildlife. This methodological framework can serve as a model for biodiversity assessment and adaptive management in other Mediterranean protected areas.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.5 Functional interpretation and synthesis\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe carnivore guild of the Sierra de la Culebra represents a functionally structured community where trophic interactions, habitat heterogeneity, and human disturbance jointly determine species coexistence. The persistence of the Iberian wolf (\u003cem\u003eCanis lupus signatus\u003c/em\u003e) as a top predator plays a pivotal ecological role in maintaining this structure. By exerting top-down control on ungulate and mesocarnivore populations, wolves indirectly influence vegetation dynamics and the availability of carrion resources, contributing to what can be described as a trophic cascade in motion (Ripple \u0026amp; Beschta, 2012; Newsome et al., 2015). The moderate negative relationship observed between wolf presence and mesocarnivore richness supports the predator-mediated coexistence hypothesis: instead of excluding smaller species, apex predators regulate their density and spatial behavior, reducing exploitative competition among mesocarnivores and stabilizing community composition (Ritchie \u0026amp; Johnson, 2009). This hierarchical yet tolerant structure typifies resilient carnivore assemblages in which competitive and facilitative interactions coexist. At the same time, habitat heterogeneity emerges as the key bottom-up driver of diversity. The positive associations between species richness and deciduous or shrub habitats underline that structurally complex landscapes promote resource partitioning and niche complementarity (Tews et al., 2004). Conversely, intensive pine plantations and croplands reduce vertical and horizontal habitat complexity, homogenizing prey availability and limiting refuge opportunities. Therefore, trophic stability in Mediterranean systems depends on the combined maintenance of top-down regulation by large predators and bottom-up diversity generated by natural vegetation mosaics. From a restoration perspective, the Sierra de la Culebra exemplifies the dual need to conserve apex predators as ecological engineers and restore native vegetation as functional habitat. Actions that facilitate oak (\u003cem\u003eQuercus spp.\u003c/em\u003e) and heathland regeneration, decrease understorey clearance in pine stands, and preserve riparian deciduous corridors will enhance the carrying capacity for a wide range of carnivores. The functional recovery of these habitats is also expected to strengthen ecosystem services such as carcass removal, rodent control, and seed dispersal via scavenger pathways. Furthermore, the methodological framework developed here, integrating camera trapping, genetic verification, and local ecological knowledge, constitutes a valuable tool for adaptive management. It allows long-term monitoring of carnivore responses to habitat restoration, forestry policies, and human-wildlife interactions, aligning conservation practice with the One Health and rewilding paradigms emerging across Europe (Perino et al., 2019; Svenning et al., 2016). In summary, the carnivore community of the Sierra de la Culebra exemplifies a resilient Mediterranean predator network in which functional diversity is maintained by the interplay of trophic regulation and landscape heterogeneity. Safeguarding both the wolf population and the structural diversity of habitats should remain central goals of regional management plans. By promoting coexistence rather than control, and by restoring ecological complexity rather than uniform productivity, managers can ensure the long-term integrity of one of the most emblematic carnivore systems in the Iberian Peninsula.\u003c/p\u003e"},{"header":"4. Concluding remarks","content":"\u003cp\u003eThe carnivore community of the Sierra de la Culebra is shaped by a combination of habitat structure, land-use history, and predator interactions. Deciduous forests and shrublands sustain higher diversity, while pine monocultures and croplands are associated with lower richness. The presence of the Iberian wolf contributes to maintaining a balanced trophic hierarchy without displacing mesocarnivores. In ecological and management terms, the preservation and restoration of natural vegetation, particularly native oak and heath formations, represent the key to conserving a functionally diverse carnivore community and reinforcing the ecological resilience of this emblematic Mediterranean landscape.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eETHICAL STATEMENT\u003c/h2\u003e\u003cp\u003eThis work did not require ethical clearance.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003ch2\u003e\u003cb\u003eCONFLICT OF INTEREST\u003c/b\u003e:\u003c/h2\u003e\u003cp\u003eThe authors declare that this work does not report any conflict of interest.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003e\u003cb\u003eETHICAL APPROVAL\u003c/b\u003e:\u003c/strong\u003e\u003cp\u003eThis work complies with all ethical standards.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003cstrong\u003e\u003cb\u003eINFORMED CONSENT\u003c/b\u003e:\u003c/strong\u003e\u003cp\u003eThe authors give their consent to the use of their data and to the publication of this work.\u003c/p\u003e\u003c/p\u003e\u003ch2\u003eFUNDING:\u003c/h2\u003e\u003cp\u003eThis work has not received any type of funding.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors contributed equally to the fieldwork, laboratory analyses, data analysis, and generation of the final manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eTo my students Juan S\u0026aacute;nchez, Iv\u0026aacute;n Rebollo y Rom\u0026aacute;n Abela. A Javier Taleg\u0026oacute;n for their knowledge of the Sierra.\u003c/p\u003e\u003ch2\u003eDATA AVAILABILITY STATEMENT:\u003c/h2\u003e\u003cp\u003eNot Applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBalestrieri, A., Remonti, L., \u0026amp; Prigioni, C. (2009). Exploitation of food resources by the Eurasian badger (Meles meles) at the altitudinal limit of its Alpine range (NW Italy). \u003cem\u003eZoological science\u003c/em\u003e, 26(12), 821-827. DOI:10.2108/zsj.26.821\u003c/li\u003e\n\u003cli\u003eBarea-Azc\u0026oacute;n, J. M., Virg\u0026oacute;s, E., Ballesteros-Duperon, E., Mole\u0026oacute;n, M., \u0026amp; Chirosa, M. (2007). Surveying carnivores at large spatial scales: a comparison of four broad-applied methods. \u003cem\u003eBiodiversity and Conservation\u003c/em\u003e, 16(4), 1213-1230. DOI:10.1007/s10531-006-9114-x\u003c/li\u003e\n\u003cli\u003eBarrientos, L. M. (1995). El lobo, distribuci\u0026oacute;n, tama\u0026ntilde;o de poblaci\u0026oacute;n y estudio de su alimentaci\u0026oacute;n en la Sierra de la Culebra (Zamora, 1995). Junta de Castilla y Le\u0026oacute;n. In\u0026eacute;dito.\u003c/li\u003e\n\u003cli\u003eBarrientos, R., and Virg\u0026oacute;s, E. (2006). Reduction of potential food interference in two sympatric carnivores by sequential use of shared resources. \u003cem\u003eActa Oecologica\u003c/em\u003e 30,107\u0026ndash;116. DOI:10.1016/j.actao.2006.02.006\u003c/li\u003e\n\u003cli\u003eBeier, C., Hansen, K., \u0026amp; Gundersen, P. (1993). Spatial variability of throughfall fluxes in a spruce forest. \u003cem\u003eEnvironmental Pollution\u003c/em\u003e, 81(3), 257-267. https://doi.org/10.1016/0269-7491(93)90208-6\u003c/li\u003e\n\u003cli\u003eBegon, M., Harper, J. L., and Townsend, C. R. (Eds) (1986). \u0026lsquo;Ecology. Individuals, Populations and Communities.\u0026rsquo; (Blackwell Scientific Publications: Oxford.).\u003c/li\u003e\n\u003cli\u003eBlanco, J. C., \u0026amp; Salvador Milla, A. (2017). Lobo\u0026ndash;Canis lupus Linnaeus, 1758.\u003c/li\u003e\n\u003cli\u003eBright, P. W., \u0026amp; Morris, P. A. (1994). Animal translocation for conservation: performance of dormice in relation to release methods, origin and season. Journal of \u003cem\u003eApplied Ecology\u003c/em\u003e, 699-708. DOI:10.2307/2404160\u003c/li\u003e\n\u003cli\u003eByers, C. R., Steinhorst, R. K., \u0026amp; Krausman, P. R. (1984). Clarification of a technique for analysis of utilization-availability data. T\u003cem\u003ehe Journal of wildlife management\u003c/em\u003e, 48(3), 1050-1053.\u003c/li\u003e\n\u003cli\u003eBunnell, F. L. (1987). Reproductive tactics of cervidae and their relationships to habitat. In RES. SYMP. NATL. ZOOL. PARK. (pp. 145-167).\u003c/li\u003e\n\u003cli\u003eCentro de Informaci\u0026oacute;n Geogr\u0026aacute;fica Nacional (2018). CORINE Land Cover. En: Centro de descargas.\u003c/li\u003e\n\u003cli\u003eClavero, M., Prenda, J., \u0026amp; Delibes, M. (2006). Seasonal use of coastal resources by otters: comparing sandy and rocky stretches. \u003cem\u003eEstuarine, Coastal and Shelf Science\u003c/em\u003e, 66(3-4), 387-394. DOI:10.1016/j.ecss.2005.09.011\u003c/li\u003e\n\u003cli\u003eClevenger, A. P. (1994). Habitat characteristics of Eurasian pine martens Martes martes in an insular Mediterranean environment. \u003cem\u003eEcography\u003c/em\u003e, 17(3), 257-263. DOI:10.1111/j.1600-0587.1994.tb00101.x\u003c/li\u003e\n\u003cli\u003eCrooks, K. R. (2002). Relative sensitivities of mammalian carnivores to habitat fragmentation. \u003cem\u003eConservation biology\u003c/em\u003e, 16(2), 488-502. DOI:10.1046/j.1523-1739.2002.00386.x\u003c/li\u003e\n\u003cli\u003eDebano, S. J. (2006). Effects of livestock grazing on aboveground insect communities in semi-arid grasslands of southeastern Arizona. \u003cem\u003eBiodiversity \u0026amp; Conservation\u003c/em\u003e, 15, 2547-2564. DOI:10.1007/s10531-005-2786-9\u003c/li\u003e\n\u003cli\u003eDebinski, D. M., \u0026amp; Holt, R. D. (2000). A survey and overview of habitat fragmentation experiments. \u003cem\u003eConservation biology\u003c/em\u003e, 14(2), 342-355. DOI:10.1046/j.1523-1739.2000.98081.x\u003c/li\u003e\n\u003cli\u003eDelibes, M. (1982). Notas sobre la distribuci\u0026oacute;n pasada y actual del meloncillo Herpestes ichneumon (L.) en la Pen\u0026iacute;nsula Ib\u0026eacute;rica. \u003cem\u003eDo\u0026ntilde;ana Acta Vertebrata\u003c/em\u003e, 9, 341-352.\u003c/li\u003e\n\u003cli\u003eDettori, E. E., Balestrieri, A., Rom\u0026aacute;n, L. G., Zapata-Perez, V. M., Ruiu, A., Carcangiu, L., \u0026amp; Robledano-Aymerich, F. (2021). A cost-effective PCR\u0026ndash;RFLP method for monitoring invasive American mink: preliminary field test in control areas of Spain. \u003cem\u003eConservation Genetics Resources\u003c/em\u003e, 13, 143-149. DOI:10.1007/s12686-020-01183-3\u003c/li\u003e\n\u003cli\u003eDettori, E. E., Zapata-P\u0026eacute;rez, V. M., Collados, D. B., Mart\u0026iacute;nez, H. I., Balestrieri, A., \u0026amp; Aymerich, F. R. (2017). A non-invasive genetic survey of sympatric carnivorous mammals in a riparian corridor along the intensively cultivated valley of the Segura River (SE Spain). In III Jornadas Doctorales Escuela Internacional de Doctorado de la Universidad de Murcia (EIDUM) (pp. 1122-1123). Servicio de Publicaciones.\u003c/li\u003e\n\u003cli\u003eDi Luzio, V. B. (2010). Risultati preliminari sul comportamento spaziale della lepre italica (lepus corsicanus) nella rnr monterano (RM). In: Workshop nazionale sulla conservazione della lepre it\u0026aacute;lica (p. 111).\u003c/li\u003e\n\u003cli\u003eErlinge, S., \u0026amp; Sandell, M. (1986). Seasonal changes in the social organization of male stoats, Mustela erminea: an effect of shifts between two decisive resources. \u003cem\u003eOikos\u003c/em\u003e, 57-62.\u003c/li\u003e\n\u003cli\u003eFerreras, P., D\u0026iacute;az-Ruiz, F., Alves, P. C., \u0026amp; Monterroso, P. S. (2016). Factores de la coexistencia de mesocarn\u0026iacute;voros en parques nacionales de ambiente mediterr\u0026aacute;neo.\u003c/li\u003e\n\u003cli\u003eGese, E. M. (2001). Territorial defense by coyotes (Canis latrans) in Yellowstone National Park, Wyoming: who, how, where, when, and why. \u003cem\u003eCanadian Journal of Zoology\u003c/em\u003e, 79(6), 980-987. DOI:10.1139/cjz-79-6-980\u003c/li\u003e\n\u003cli\u003eKauhala, K., \u0026amp; Salonen, L. (2012). Does a non-invasive method-latrine surveys-reveal habitat preferences of raccoon dogs and badgers?. \u003cem\u003eMammalian Biology\u003c/em\u003e, 77, 264-270. DOI:10.1016/j.mambio.2012.02.007\u003c/li\u003e\n\u003cli\u003eKucera, T. E., \u0026amp; Barrett, R. H. (1993). In my experience: the Trailmaster\u0026reg; camera system for detecting wildlife. Wildlife Society Bulletin, 505-508.\u003c/li\u003e\n\u003cli\u003eKohn, M. H., \u0026amp; Wayne, R. K. (1997). Facts from feces revisited. \u003cem\u003eTrends in ecology \u0026amp; evolution\u003c/em\u003e, 12(6), 223-227.\u003c/li\u003e\n\u003cli\u003eLanszki, J., \u0026amp; Nagy, L. (2003). A L\u0026aacute;tr\u0026aacute;nyi Puszta Term\u0026eacute;szetv\u0026eacute;delmi Ter\u0026uuml;let gerinces (Vertebrata) faun\u0026aacute;j\u0026aacute;nak felm\u0026eacute;r\u0026eacute;se-Investigation on vertebrata fauna of the L\u0026aacute;tr\u0026aacute;nyi Puszta Nature Conservation Area. Natura Somogyiensis, (5), 279-290. DOI:10.24394/NatSom.2003.5.279\u003c/li\u003e\n\u003cli\u003eLanszki, J., Hidas, A., Szentes, K., R\u0026eacute;vay, T., Lehoczky, I., \u0026amp; Weiss, S. (2008). Relative spraint density and genetic structure of otter (Lutra lutra) along the Drava River in Hungary. \u003cem\u003eMammalian biology\u003c/em\u003e, 73(1), 40-47. DOI:10.1016/j.mambio.2007.08.005\u003c/li\u003e\n\u003cli\u003eLeonard, J. A., Vila, C., \u0026amp; Wayne, R. K. (2005). FAST TRACK: Legacy lost: genetic variability and population size of extirpated US grey wolves (Canis lupus). Molecular Ecology, 14(1), 9-17. DOI:10.1111/j.1365-294X.2004.02389.x\u003c/li\u003e\n\u003cli\u003eLibois, R., \u0026amp; Waechter, A. (1991). La fouine (Martes foina Erxleben, 1777. Soci\u0026eacute;t\u0026eacute; fran\u0026ccedil;aise pour l\u0026apos;\u0026eacute;tude et la protection des mammif\u0026egrave;res, Bourges, France.\u003c/li\u003e\n\u003cli\u003eManly, B. F. (1993). Comments on design and analysis of multiple-choice feeding-preference experiments. \u003cem\u003eOecologia\u003c/em\u003e, 149-152.\u003c/li\u003e\n\u003cli\u003eMart\u0026iacute;nez, F. A., \u0026amp; Pastor-Alfonso, M. J. (2019). El turismo lobero en la Sierra de La Culebra (Espa\u0026ntilde;a). \u003cem\u003eRevista LIDER\u003c/em\u003e, 21(35), 137-160. DOI:10.32735/S0719-526520193513\u003c/li\u003e\n\u003cli\u003eMech, L. D. (1966). The Wolves of isle royale (No. 7). US Government Printing Office.\u003c/li\u003e\n\u003cli\u003eMech, L. D., \u0026amp; Peterson, R. O. (2003). Wolf-prey relations.\u003c/li\u003e\n\u003cli\u003eMendoza, M., Janis, C. M., \u0026amp; Palmqvist, P. (2005). Ecological patterns in the trophic-size structure of large mammal communities: a \u0026lsquo;taxon-free\u0026rsquo;characterization. \u003cem\u003eEvolutionary Ecology Research\u003c/em\u003e, 7(4), 505-530.\u003c/li\u003e\n\u003cli\u003eN\u0026uacute;\u0026ntilde;ez, I. B. (2005). Patrones de marcaje con heces por la marta europea (Martes martes) en el noroeste de Espa\u0026ntilde;a: importancia para su estudio. Galemys: Bolet\u0026iacute;n informativo de la Sociedad Espa\u0026ntilde;ola para la conservaci\u0026oacute;n y estudio de los mam\u0026iacute;feros, 17(1), 123-134. DOI:10.7325/Galemys.2005.NE.A10\u003c/li\u003e\n\u003cli\u003eLira-Torres, I., \u0026amp; Briones-Salas, M. (2012). Abundancia relativa y patrones de actividad de los mam\u0026iacute;feros de los Chimalapas, Oaxaca, M\u0026eacute;xico. \u003cem\u003eActa zool\u0026oacute;gica mexicana\u003c/em\u003e, 28(3), 566-585. DOI:10.21829/azm.2012.283859\u003c/li\u003e\n\u003cli\u003eMortelliti, A., Amori, G., \u0026amp; Boitani, L. (2010). The role of habitat quality in fragmented landscapes: a conceptual overview and prospectus for future research. \u003cem\u003eOecologia\u003c/em\u003e, 163(2), 535-547. DOI:10.1007/s00442-010-1623-3\u003c/li\u003e\n\u003cli\u003ePilot, M., Gralak, B., Goszczyński, J., \u0026amp; Posłuszny, M. (2007). A method of genetic identification of pine marten (Martes martes) and stone marten (Martes foina) and its application to faecal samples. \u003cem\u003eJournal of Zoology\u003c/em\u003e, 271(2), 140-147. DOI:10.1111/j.1469-7998.2006.00179.x\u003c/li\u003e\n\u003cli\u003eNeu, C. W., Byers, C. R., \u0026amp; Peek, J. M. (1974). A technique for analysis of utilization-availability data. The Journal of Wildlife Management, 541-545.\u003c/li\u003e\n\u003cli\u003eNewsome, T. M., Dellinger, J. A., Pavey, C. R., Ripple, W. J., Shores, C. R., Wirsing, A. J., \u0026amp; Dickman, C. R. (2015). The ecological effects of providing resource subsidies to predators. Global Ecology and Biogeography, 24(1), 1-11. https://doi.org/10.1111/geb.12236\u003c/li\u003e\n\u003cli\u003ePalacios, J. y J. L. Vicente (1997). Gesti\u0026oacute;n de Fauna en la Reserva Nacional de Caza \u0026ldquo;Sierra de la Culebra\u0026rdquo;. Pp. 18/1-18/15. En: Master Universitario en Veterinaria y Fauna Salvaje; Tomo IV: Gesti\u0026oacute;n Cineg\u0026eacute;tica y de Especies Protegidas. Colegio Oficial de Veterinarios de Zamora e I.C.E. Universidad de Le\u0026oacute;n. Zamora.\u003c/li\u003e\n\u003cli\u003ePalomares, F., Ferreras, P., Fedriani, J. M., and Delibes, M. (1996). Spatial relationships between Iberian lynx and other carnivores in an area of south-west Spain. \u003cem\u003eJournal of Applied Ecology\u003c/em\u003e 33, 5\u0026ndash;13.\u003c/li\u003e\n\u003cli\u003ePerino, A., Pereira, H. M., Navarro, L. M., Fern\u0026aacute;ndez, N., Bullock, J. M., Ceaușu, S., ... \u0026amp; Svenning, J. C. (2019). \u003cem\u003eRewilding complex ecosystems. Science\u003c/em\u003e, 364(6438), eaav5570. https://doi.org/10.1126/science.aav5570\u003c/li\u003e\n\u003cli\u003ePeterson, R. O. (1995). Wolves as interspecific competitors in canid ecology. Ecology and conservation of wolves in a changing world, 315-324.\u003c/li\u003e\n\u003cli\u003ePutman, R. J. (1984). Facts from faeces. \u003cem\u003eMammal review\u003c/em\u003e, 14(2), 79-97.\u003c/li\u003e\n\u003cli\u003eRamos Santos, J. M. (2006). La transformaci\u0026oacute;n del paisaje vegetal y los aprovechamientos forestales tradicionales en la Sierra de la Culebra (Zamora). Investigaciones geogr\u0026aacute;ficas, n\u0026ordm; 40, 55-72.\u003c/li\u003e\n\u003cli\u003eRemonti, L., Balestrieri, A., Ruiz-Gonz\u0026aacute;lez, A., G\u0026oacute;mez-Moliner, B. J., Capelli, E., \u0026amp; Prigioni, C. (2012). Intraguild dietary overlap and its possible relationship to the coexistence of mesocarnivores in intensive agricultural habitats. \u003cem\u003ePopulation Ecology\u003c/em\u003e, 54, 521-532. DOI:10.1007/s10144-012-0337-2\u003c/li\u003e\n\u003cli\u003eRipple, W. J., \u0026amp; Beschta, R. L. (2012). Trophic cascades in Yellowstone: The first 15 years after wolf reintroduction. \u003cem\u003eBiological Conservation\u003c/em\u003e, 145(1), 205\u0026ndash;213. https://doi.org/10.1016/j.biocon.2011.11.005 \u003c/li\u003e\n\u003cli\u003eRitchie, E. G., \u0026amp; Johnson, C. N. (2009). Predator interactions, mesopredator release and biodiversity conservation. \u003cem\u003eEcology Letters\u003c/em\u003e, 12(9), 982\u0026ndash;998. https://doi.org/10.1111/j.1461-0248.2009.01347.x\u003c/li\u003e\n\u003cli\u003eRodr\u0026iacute;guez-Mazzini, R. (1996). Uso de la t\u0026eacute;cnica de estaciones olfativas (SCENT-STATION TECHNIQUE) en estudios de ecolog\u0026iacute;a de mam\u0026iacute;feros (No. 599.74 ROD).\u003c/li\u003e\n\u003cli\u003eRosalino, L. M., \u0026amp; Santos-Reis, M. (2002). Feeding habits of the common genet Genetta genetta (Carnivora: Viverridae) in a semi-natural landscape of central Portugal.\u003c/li\u003e\n\u003cli\u003eRosalino, L. M., Santos, M. J., Beier, P., \u0026amp; Santos-Reis, M. (2008). Eurasian badger habitat selection in Mediterranean environments: does scale really matter?. \u003cem\u003eMammalian Biology\u003c/em\u003e, 73(3), 189-198. https://doi.org/10.1016/j.mambio.2007.02.004\u003c/li\u003e\n\u003cli\u003eRosellini, S., Osorio, E., Ruiz-Gonz\u0026aacute;lez, A., Pi\u0026ntilde;eiro, A., \u0026amp; Barja, I. (2008). Monitoring the small-scale distribution of sympatric European pine martens (Martes martes) and stone martens (Martes foina): a multievidence approach using faecal DNA analysis and camera-traps. \u003cem\u003eWildlife research\u003c/em\u003e, 35(5), 434-440. DOI:10.1071/wr07030\u003c/li\u003e\n\u003cli\u003eRuiz-Gonz\u0026aacute;lez, A., Rubines, J., Berdi\u0026oacute;n, O., \u0026amp; G\u0026oacute;mez-Moliner, B. J. (2008). A non-invasive genetic method to identify the sympatric mustelids pine marten (Martes martes) and stone marten (Martes foina): preliminary distribution survey on the northern Iberian Peninsula. \u003cem\u003eEuropean Journal of Wildlife Research\u003c/em\u003e, 54, 253-261. DOI:10.1007/s10344-007-0138-7\u003c/li\u003e\n\u003cli\u003eSambrook, J., Fritsch, E. F., \u0026amp; Maniatis, T. (1989). Molecular cloning: a laboratory manual (No. Ed. 2). Cold spring harbor laboratory press.\u003c/li\u003e\n\u003cli\u003eSanz, B. y Tur\u0026oacute;n, J.V. (2017). Gu\u0026iacute;a de mam\u0026iacute;feros terrestres. Pen\u0026iacute;nsula ib\u0026eacute;rica y balear. Ed. Prames. Zaragoza.\u003c/li\u003e\n\u003cli\u003eSoul\u0026eacute;, M. E., Bolger, D. T., Alberts, A. C., Wrights, J., Sorice, M., \u0026amp; Hill, S. (1988). Reconstructed dynamics of rapid extinctions of chaparral‐requiring birds in urban habitat islands. \u003cem\u003eConservation Biology\u003c/em\u003e, 2(1), 75-92. https://doi.org/10.1111/j.1523-1739.1988.tb00337.x\u003c/li\u003e\n\u003cli\u003eSu\u0026aacute;rez-Tangil, B. D., \u0026amp; Rodr\u0026iacute;guez, A. (2021). Estimates of species richness and composition depend on detection method in assemblages of terrestrial mammals. \u003cem\u003eAnimals\u003c/em\u003e, 11(1), 186. DOI:10.3390/ani11010186\u003c/li\u003e\n\u003cli\u003eSvenning, J. C., Pedersen, P. B., Donlan, C. J., Ejrn\u0026aelig;s, R., Faurby, S., Galetti, M., ... \u0026amp; Vera, F. W. (2016). Science for a wilder Anthropocene: Synthesis and future directions for trophic rewilding research. \u003cem\u003eProceedings of the National Academy of Sciences\u003c/em\u003e, 113(4), 898-906. https://doi.org/10.1073/pnas.1502556112\u003c/li\u003e\n\u003cli\u003eTews, J., Brose, U., Grimm, V., Tielb\u0026ouml;rger, K., Wichmann, M. C., Schwager, M., \u0026amp; Jeltsch, F. (2004). Animal species diversity driven by habitat heterogeneity/diversity: The importance of keystone structures. \u003cem\u003eJournal of Biogeography\u003c/em\u003e, 31(1), 79\u0026ndash;92. https://doi.org/10.1046/j.0305-0270.2003.00994\u003c/li\u003e\n\u003cli\u003eTorre, I., Arrizabalaga, A. y Flaquer, C. (2003). Estudio de la distribuci\u0026oacute;n y abundancia de carn\u0026iacute;voros en el Parque Natural del Montnegre y El Corredor mediante trampeo fotogr\u0026aacute;fico. \u003cem\u003eGalemys\u003c/em\u003e, 15 (1), 15-28. DOI:10.7325/Galemys.2003.1.A2\u003c/li\u003e\n\u003cli\u003eVicente, J. L., Rodr\u0026iacute;guez, M., \u0026amp; Palacios, J. (2000). Gesti\u0026oacute;n del lobo ib\u0026eacute;rico (Canis lupus signatus Cabrera, 1907) en la Reserva Regional de Caza \u0026ldquo;Sierra de la Culebra\u0026rdquo;(Zamora). \u003cem\u003eGalemys\u003c/em\u003e, 12(1), 181-199.\u003c/li\u003e\n\u003cli\u003eVil\u0026agrave;, C., Amorim, I. R., Leonard, J. A., Posada, D., Castroviejo, J., Petrucci‐Fonseca, F., ... \u0026amp; Wayne, R. K. (1999). Mitochondrial DNA phylogeography and population history of the grey wolf Canis lupus. \u003cem\u003eMolecular ecology\u003c/em\u003e, 8(12), 2089-2103. DOI:10.1046/j.1365-294x.1999.00825.x\u003c/li\u003e\n\u003cli\u003eVirg\u0026oacute;s, E., \u0026amp; Casanovas, J. G. (1997). Habitat selection of genet Genetta genetta in the mountains of central Spain. \u003cem\u003eActa Theriologica\u003c/em\u003e, 42(2), 169-177. DOI:10.7325/Galemys.2007.NE.A6\u003c/li\u003e\n\u003cli\u003eWaldron, J. L., Bennett, S. H., Welch, S. M., Dorcas, M. E., Lanham, J. D., \u0026amp; Kalinowsky, W. (2006). Habitat specificity and home‐range size as attributes of species vulnerability to extinction: a case study using sympatric rattlesnakes. Animal conservation, 9(4), 414-420. DOI:10.1111/j.1469-1795.2006.00050.x\u003c/li\u003e\n\u003cli\u003eWhite, G. C., \u0026amp; Garrott, R. A. (2012). Analysis of wildlife radio-tracking data. Elsevier.\u003c/li\u003e\n\u003cli\u003eZielinski, W. J., \u0026amp; Kucera, T. E. (1995). American marten, fisher, lynx, and wolverine: survey methods for their detection (Vol. 157). Pacific Southwest Research Station. https://doi.org/10.2737/PSW-GTR-157\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"carnivore guild, trophic structure, habitat heterogeneity, top-down regulation, community ecology","lastPublishedDoi":"10.21203/rs.3.rs-8029414/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8029414/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe aim of this study was to understand how mesocarnivores are structured as a function of habitat type and in the presence of a top predator such as the wolf. Indeed, the Sierra de la Culebra (Zamora, NW Spain) hosts one of the densest populations of Iberian wolves (\u003cem\u003eCanis lupus signatus\u003c/em\u003e) in Western Europe, together with a diverse guild of mesocarnivores. We investigated habitat determinants of carnivore diversity and trophic structure using a non-invasive, multi-method approach that combined camera trapping, faecal DNA identification, and interviews with local rangers across twelve 7.5 \u0026times; 7.5 km grid cells (65,000 ha). Seven camera trap stations were positioned in the grid, with one camera trap per station, after the results were cross-referenced with 253 fecal samples and a survey conducted among Park staff. Finally, to determine the habitat, the relative abundance index (RAI) for each taxon was calculated using the formula suggested by Lira-Torres and Briones Salas. Twelve carnivore species were recorded, with wolves and red foxes ubiquitous and smaller mustelids showing patchy distributions although wolf presence exerted a weak negative influence on mesocarnivore richness, suggesting moderate top-down regulation within the guild. As regards the characterization of the habitat multiple regression models revealed that deciduous forests and shrublands were positively associated with total species richness, while pine plantations and croplands had negative effects therefore the carnivore community of the Sierra de la Culebra has shown a significant predilection for forest areas with deciduous or thicket forests, strongly reaching coniferous forests (generally repopulation) and cultivated areas. In the European context, where the wolf is a sensitive species, these results demonstrate that carnivore diversity in Mediterranean landscapes depends on both trophic and habitat complexity. Ecological restoration in the Sierra de la Culebra should therefore prioritize the recovery of native oak woodlands and mixed shrub mosaics to maintain a functionally rich and balanced predator community.\u003c/p\u003e","manuscriptTitle":"Trophic structure and habitat determinants of a Mediterranean carnivore guild: insights from the Sierra de la Culebra (NW Spain)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-11-24 09:28:29","doi":"10.21203/rs.3.rs-8029414/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"e418a64e-53ae-498c-b393-f710d6849eff","owner":[],"postedDate":"November 24th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-02-10T13:41:42+00:00","versionOfRecord":[],"versionCreatedAt":"2025-11-24 09:28:29","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8029414","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8029414","identity":"rs-8029414","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}