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2021

Sacred groves in Greece are usually forest remnants with large trees around chapels, protected through centuries by Orthodox religion. We examined the comparative ecological value of 20 oak-dominated sacred groves vs managed oakwoods, in terms of their habitat characteristics and avian communities (passerines and woodpeckers). Sacred groves have maintained a more pronounced old-growth character than managed oakwoods in terms of average Diameter at Breast Height (DBH) and tree height. Besides holding significantly greater bird species richness and abundance, they supported greater functional richness, phylogenetic diversity, and phylogenetic bird species variability. Bird communities in sacred groves were more heterogeneous and showed greater avian specialization levels than in managed woods. Generalized Linear Models showed that the main factor positively affecting all aspects of bird diversity was DBH, while the abundance of dead trees increased bird abundance. Our results underline the importance of maintaining large-sized trees in forest management practices to support bird diversity and decrease biotic homogenization. Since the new European Biodiversity Strategy explicitly requires all remaining European primary and old-growth forests to be strictly protected by 2030, we argue that sacred groves, despite their small size, meet the criteria to be considered in the strict protection and restoration targets of the strategy, as primary old growth woods of high biodiversity value.
Sacred groves in Greece are usually forest remnants with large trees around chapels, protected through centuries by Orthodox religion. We examined the comparative ecological value of 20 oak-dominated sacred groves vs managed oakwoods, in terms of their habitat characteristics and avian communities (passerines and woodpeckers). Sacred groves have maintained a more pronounced old-growth character than managed oakwoods in terms of average Diameter at Breast Height (DBH) and tree height. Besides holding significantly greater bird species richness and abundance, they supported greater functional richness, phylogenetic diversity, and phylogenetic bird species variability. Bird communities in sacred groves were more heterogeneous and showed greater avian specialization levels than in managed woods. Generalized Linear Models showed that the main factor positively affecting all aspects of bird diversity was DBH, while the abundance of dead trees increased bird abundance. Our results underline the importance of maintaining large-sized trees in forest management practices to support bird diversity and decrease biotic homogenization. Since the new European Biodiversity Strategy explicitly requires all remaining European primary and old-growth forests to be strictly protected by 2030, we argue that sacred groves, despite their small size, meet the criteria to be considered in the strict protection and restoration targets of the strategy, as primary old growth woods of high biodiversity value.
Wind energy is the leading renewable technology towards achieving climate goals, yet biodiversity trade-offs via land take are emerging. Thus, we are facing the paradox of impacting on biodiversity to combat climate change. We suggest a novel method of spatial planning that enhances windfarm sustainability: investments are prioritized in the most fragmented zones that lie outside the Natura 2000 network of protected areas. We showcase it in Greece, a biodiversity hotspot with a strong climate policy and land conflict between conservation and wind energy schemes. The analysis indicates that the suggested investment zone supports wind harnessing 1.5 times higher than the 2030 national goal, having only marginally lower (4%) wind speed. It performs well for the conservation of the annexed habitats and species of the two Nature Directives and it greatly overlaps with the Important Bird Areas (93%) and the roadless areas (80%) of Greece. It also greatly overlaps (82%–91%) with the exclusion zones suggested according to three sensitivity maps for bird conservation. Since land use change triggers biodiversity decline, we underline the necessity of such approaches for meeting both climate and biodiversity goals and call for a greater environmental policy convergence towards biodiversity conservation and no net land take.
Wind energy is the leading renewable technology towards achieving climate goals, yet biodiversity trade-offs via land take are emerging. Thus, we are facing the paradox of impacting on biodiversity to combat climate change. We suggest a novel method of spatial planning that enhances windfarm sustainability: investments are prioritized in the most fragmented zones that lie outside the Natura 2000 network of protected areas. We showcase it in Greece, a biodiversity hotspot with a strong climate policy and land conflict between conservation and wind energy schemes. The analysis indicates that the suggested investment zone supports wind harnessing 1.5 times higher than the 2030 national goal, having only marginally lower (4%) wind speed. It performs well for the conservation of the annexed habitats and species of the two Nature Directives and it greatly overlaps with the Important Bird Areas (93%) and the roadless areas (80%) of Greece. It also greatly overlaps (82%–91%) with the exclusion zones suggested according to three sensitivity maps for bird conservation. Since land use change triggers biodiversity decline, we underline the necessity of such approaches for meeting both climate and biodiversity goals and call for a greater environmental policy convergence towards biodiversity conservation and no net land take.
Global environmental goals mandate the expansion of the protected area network to halt biodiversity loss. The European Union’s Natura 2000 network covers 27.3% of the terrestrial area of Greece, one of the highest percentages in Europe. However, the extent to which this network protects Europe’s biodiversity, especially in a biodiverse country like Greece, is unknown. Here, we overlap the country’s Natura 2000 network with the ranges of the 424 species assessed as threatened on the IUCN Red List and present in Greece. Natura 2000 overlaps on average 47.6% of the mapped range of threatened species; this overlap far exceeds that expected by random networks (21.4%). Special Protection Areas and Special Areas of Conservation (non-exclusive subsets of Natura 2000 sites) overlap 33.4% and 38.1% respectively. Crete and Peloponnese are the two regions with the highest percentage of threatened species, with Natura 2000 sites overlapping on average 62.3% with the threatened species’ ranges for the former, but only 30.6% for the latter. The Greek ranges of all 62 threatened species listed in Annexes 1 and II to the Birds and Habitats Directives are at least partially overlapped by the network (52.0%), and 18.0% of these are fully overlapped. However, the ranges of 27 threatened species, all of which are endemic to Greece, are not overlapped at all. These results can inform national policies for the protection of biodiversity beyond current Natura 2000 sites.
Global environmental goals mandate the expansion of the protected area network to halt biodiversity loss. The European Union’s Natura 2000 network covers 27.3% of the terrestrial area of Greece, one of the highest percentages in Europe. However, the extent to which this network protects Europe’s biodiversity, especially in a biodiverse country like Greece, is unknown. Here, we overlap the country’s Natura 2000 network with the ranges of the 424 species assessed as threatened on the IUCN Red List and present in Greece. Natura 2000 overlaps on average 47.6% of the mapped range of threatened species; this overlap far exceeds that expected by random networks (21.4%). Special Protection Areas and Special Areas of Conservation (non-exclusive subsets of Natura 2000 sites) overlap 33.4% and 38.1% respectively. Crete and Peloponnese are the two regions with the highest percentage of threatened species, with Natura 2000 sites overlapping on average 62.3% with the threatened species’ ranges for the former, but only 30.6% for the latter. The Greek ranges of all 62 threatened species listed in Annexes 1 and II to the Birds and Habitats Directives are at least partially overlapped by the network (52.0%), and 18.0% of these are fully overlapped. However, the ranges of 27 threatened species, all of which are endemic to Greece, are not overlapped at all. These results can inform national policies for the protection of biodiversity beyond current Natura 2000 sites.

2020

Land use change poses as the top threat for biodiversity decline, and road sprawl as a key driver behind it globally. According to the recent Landscape Fragmentation Indicator (LFI), Greece is less fragmented than the rest of Europe but presents higher rates of fragmentation increase. We developed the Roadless Fragmentation Indicator (RFI) to monitor fragmentation in more natural ecosystems. The RFI calculates the percentage of land that is covered by roadless areas (RAs), defined as land patches over 1 km2 that are over 1 km away from the nearest road. We produced the roadless map of Greece, concluding to 1115 RAs ranked by size (1–256 km2) and to a national RFI of less than 5%. The RFI reflected naturalness, was significantly higher in the Natura 2000 network, and was more sensitive in less fragmented zones. Six mountains (0.51% of Greek land) have remained largely roadless (RAs ≥ 50 km2) and should be protected as such. We call for a straightforward roadlessness policy under a “European Roadless Rule” that would legally protect at least 2% of European land as road-free area. We also call for no further unjustified road sprawl in more natural and least fragmented ecosystems, as a measure to be integrated in all sectors of EU policy and particularly in the spatial planning of development projects. We recommend a five-step roadlessness guideline to be implemented in the European Union, including Greece, as a measure to effectively address biodiversity decline.
Land use change poses as the top threat for biodiversity decline, and road sprawl as a key driver behind it globally. According to the recent Landscape Fragmentation Indicator (LFI), Greece is less fragmented than the rest of Europe but presents higher rates of fragmentation increase. We developed the Roadless Fragmentation Indicator (RFI) to monitor fragmentation in more natural ecosystems. The RFI calculates the percentage of land that is covered by roadless areas (RAs), defined as land patches over 1 km2 that are over 1 km away from the nearest road. We produced the roadless map of Greece, concluding to 1115 RAs ranked by size (1–256 km2) and to a national RFI of less than 5%. The RFI reflected naturalness, was significantly higher in the Natura 2000 network, and was more sensitive in less fragmented zones. Six mountains (0.51% of Greek land) have remained largely roadless (RAs ≥ 50 km2) and should be protected as such. We call for a straightforward roadlessness policy under a “European Roadless Rule” that would legally protect at least 2% of European land as road-free area. We also call for no further unjustified road sprawl in more natural and least fragmented ecosystems, as a measure to be integrated in all sectors of EU policy and particularly in the spatial planning of development projects. We recommend a five-step roadlessness guideline to be implemented in the European Union, including Greece, as a measure to effectively address biodiversity decline.
Balkan chamois (Rupicapra rupicapra balcanica) is a protected species with an Inadequate-Bad (U2) conservation status in Greece. Our study explores its seasonal range use pattern, demography and habitat selection in a site of the Natura 2000 network, Timfi Mountain. To this aim, we examined 1168 observations obtained from six seasonal surveys (2002: four seasons, 2014 and 2017: autumn) and performed an ecological-niche factor analysis (ENFA), using 16 environmental and human-disturbance variables. The species had an annual range of 6491 ha (25% of the study area), followed the typical range-use pattern, and presented the minimum core area during the rutting season (autumn). Timfi Mt hosted 469 individuals in 2017 (the largest population in Greece), increasing by 3.55 times since 2002. The species selected higher altitudes during summer and autumn, pinewoods over broad-leaved woods as winter grounds, and it avoided south-facing slopes. Our results supported the anthropogenic risk avoidance hypothesis; the species always selected remote areas away from roads, human settlements, and hunting grounds. In Greece, 40% of its distribution area falls within hunting ban areas (16.5% of the country). A national conservation policy is needed towards maintaining and increasing roadless areas and hunting-ban areas within Balkan chamois range nationwide.
Balkan chamois (Rupicapra rupicapra balcanica) is a protected species with an Inadequate-Bad (U2) conservation status in Greece. Our study explores its seasonal range use pattern, demography and habitat selection in a site of the Natura 2000 network, Timfi Mountain. To this aim, we examined 1168 observations obtained from six seasonal surveys (2002: four seasons, 2014 and 2017: autumn) and performed an ecological-niche factor analysis (ENFA), using 16 environmental and human-disturbance variables. The species had an annual range of 6491 ha (25% of the study area), followed the typical range-use pattern, and presented the minimum core area during the rutting season (autumn). Timfi Mt hosted 469 individuals in 2017 (the largest population in Greece), increasing by 3.55 times since 2002. The species selected higher altitudes during summer and autumn, pinewoods over broad-leaved woods as winter grounds, and it avoided south-facing slopes. Our results supported the anthropogenic risk avoidance hypothesis; the species always selected remote areas away from roads, human settlements, and hunting grounds. In Greece, 40% of its distribution area falls within hunting ban areas (16.5% of the country). A national conservation policy is needed towards maintaining and increasing roadless areas and hunting-ban areas within Balkan chamois range nationwide.
Inferring species’ responses to climate change in the absence of long-term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14-day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23-day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8-day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space-for-time substitutions, understanding species-specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.
Inferring species’ responses to climate change in the absence of long-term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14-day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23-day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8-day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space-for-time substitutions, understanding species-specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.

2019

Understanding the feeding habits of wolves is essential for designing and implementing fundamental management processes across the range of the species. This is even more important within human-dominated areas, such as southern Europe, and more especially Greece. In this context, we analyzed 123 scat samples, collected between 2010 and 2012, from a mixed agricultural, forested and human-dominated area, centered on the municipality of Domokos in central continental Greece. We used standard laboratory procedures for scat analysis and calculated percentages of frequency of occurrence (FO%), average volume (AV%) and biomass index (BM%) to assess diet composition, and estimated prey selectivity. Domestic prey composed the bulk of wolf diet (FO%=73.5, AV%=84.8, BM%=97.2), wild ungulates were almost absent (FO%=0.5, AV%=0.8, BM%=1.2), whereas grass consumption was high in our area (FO%=19.5, AV%=11.0). The high dependence on livestock corroborates previous studies from Greece and other countries in southern Europe. Goat (FO%=46.0, AV%=61.2, BM%=64.9) was the main prey and was strongly selected, with sheep (FO%=11.5, AV%=9.0, BM%=11.2), pig carrion and cattle ranking behind (FO%=11.5, AV%=10.1, BM%=8.7 and FO%=4.5, AV%=4.5, BM%=12.4, respectively). No differences across seasons were detected, except from pig carrion, which increased during winter. The preference for goats is probably associated with its grazing behavior. High livestock consumption generally results in increased human-wolf conflict. Thus, substantial improvement of husbandry practices and restoration of wild ungulate populations are recommended to facilitate wolf-human coexistence in Greece.
Understanding the feeding habits of wolves is essential for designing and implementing fundamental management processes across the range of the species. This is even more important within human-dominated areas, such as southern Europe, and more especially Greece. In this context, we analyzed 123 scat samples, collected between 2010 and 2012, from a mixed agricultural, forested and human-dominated area, centered on the municipality of Domokos in central continental Greece. We used standard laboratory procedures for scat analysis and calculated percentages of frequency of occurrence (FO%), average volume (AV%) and biomass index (BM%) to assess diet composition, and estimated prey selectivity. Domestic prey composed the bulk of wolf diet (FO%=73.5, AV%=84.8, BM%=97.2), wild ungulates were almost absent (FO%=0.5, AV%=0.8, BM%=1.2), whereas grass consumption was high in our area (FO%=19.5, AV%=11.0). The high dependence on livestock corroborates previous studies from Greece and other countries in southern Europe. Goat (FO%=46.0, AV%=61.2, BM%=64.9) was the main prey and was strongly selected, with sheep (FO%=11.5, AV%=9.0, BM%=11.2), pig carrion and cattle ranking behind (FO%=11.5, AV%=10.1, BM%=8.7 and FO%=4.5, AV%=4.5, BM%=12.4, respectively). No differences across seasons were detected, except from pig carrion, which increased during winter. The preference for goats is probably associated with its grazing behavior. High livestock consumption generally results in increased human-wolf conflict. Thus, substantial improvement of husbandry practices and restoration of wild ungulate populations are recommended to facilitate wolf-human coexistence in Greece.
Understanding the feeding habits of wolves is essential for designing and implementing fundamental management processes across the range of the species. This is even more important within human-dominated areas, such as southern Europe, and more especially Greece. In this context, we analyzed 123 scat samples, collected between 2010 and 2012, from a mixed agricultural, forested and human-dominated area, centered on the municipality of Domokos in central continental Greece. We used standard laboratory procedures for scat analysis and calculated percentages of frequency of occurrence (FO%), average volume (AV%) and biomass index (BM%) to assess diet composition, and estimated prey selectivity. Domestic prey composed the bulk of wolf diet (FO%=73.5, AV%=84.8, BM%=97.2), wild ungulates were almost absent (FO%=0.5, AV%=0.8, BM%=1.2), whereas grass consumption was high in our area (FO%=19.5, AV%=11.0). The high dependence on livestock corroborates previous studies from Greece and other countries in southern Europe. Goat (FO%=46.0, AV%=61.2, BM%=64.9) was the main prey and was strongly selected, with sheep (FO%=11.5, AV%=9.0, BM%=11.2), pig carrion and cattle ranking behind (FO%=11.5, AV%=10.1, BM%=8.7 and FO%=4.5, AV%=4.5, BM%=12.4, respectively). No differences across seasons were detected, except from pig carrion, which increased during winter. The preference for goats is probably associated with its grazing behavior. High livestock consumption generally results in increased human-wolf conflict. Thus, substantial improvement of husbandry practices and restoration of wild ungulate populations are recommended to facilitate wolf-human coexistence in Greece.
Understanding the feeding habits of wolves is essential for designing and implementing fundamental management processes across the range of the species. This is even more important within human-dominated areas, such as southern Europe, and more especially Greece. In this context, we analyzed 123 scat samples, collected between 2010 and 2012, from a mixed agricultural, forested and human-dominated area, centered on the municipality of Domokos in central continental Greece. We used standard laboratory procedures for scat analysis and calculated percentages of frequency of occurrence (FO%), average volume (AV%) and biomass index (BM%) to assess diet composition, and estimated prey selectivity. Domestic prey composed the bulk of wolf diet (FO%=73.5, AV%=84.8, BM%=97.2), wild ungulates were almost absent (FO%=0.5, AV%=0.8, BM%=1.2), whereas grass consumption was high in our area (FO%=19.5, AV%=11.0). The high dependence on livestock corroborates previous studies from Greece and other countries in southern Europe. Goat (FO%=46.0, AV%=61.2, BM%=64.9) was the main prey and was strongly selected, with sheep (FO%=11.5, AV%=9.0, BM%=11.2), pig carrion and cattle ranking behind (FO%=11.5, AV%=10.1, BM%=8.7 and FO%=4.5, AV%=4.5, BM%=12.4, respectively). No differences across seasons were detected, except from pig carrion, which increased during winter. The preference for goats is probably associated with its grazing behavior. High livestock consumption generally results in increased human-wolf conflict. Thus, substantial improvement of husbandry practices and restoration of wild ungulate populations are recommended to facilitate wolf-human coexistence in Greece.
Increasing urbanisation is reported to have significant effects on bat communities, due to habitat modifications, light and noise pollution and reduced prey availability. Recent studies have indicated that species show varying responses to urbanisation, with a few able to exploit man-made structures and adjust to the new environmental conditions. This study aimed to identify how landscape composition influences bat diversity and community structure along the urbanisation gradient in a coastal Mediterranean city (Patras, Greece) and whether particular species benefit from the novel conditions. We conducted acoustic surveys along 45 transects during the post-breeding season for two years. The effect of land cover, the number of streetlamps (a proxy of artificial illumination), the presence of water bodies and weather conditions on bat activity, and community structure were investigated using Generalized Linear Mixed Models, and multivariate statistics respectively. Eight bat species and five species groups were identified. Bat communities were affected by urbanisation in general and diversity was low in the entire study area. The community was dominated by the synurbic species Pipistrellus kuhlii, which comprised more than 70% of the total bat activity recorded. A positive relationship between built-up areas and bat activity was found, probably because P. kuhlii usually forages around streetlamps in urban areas. In contrast, vegetation cover did not affect bat activity, even in the less urbanised areas. The remainder of the bat species were not frequently recorded and were mostly detected close to water bodies, highlighting their value for foraging bats and the need for their conservation.
Increasing urbanisation is reported to have significant effects on bat communities, due to habitat modifications, light and noise pollution and reduced prey availability. Recent studies have indicated that species show varying responses to urbanisation, with a few able to exploit man-made structures and adjust to the new environmental conditions. This study aimed to identify how landscape composition influences bat diversity and community structure along the urbanisation gradient in a coastal Mediterranean city (Patras, Greece) and whether particular species benefit from the novel conditions. We conducted acoustic surveys along 45 transects during the post-breeding season for two years. The effect of land cover, the number of streetlamps (a proxy of artificial illumination), the presence of water bodies and weather conditions on bat activity, and community structure were investigated using Generalized Linear Mixed Models, and multivariate statistics respectively. Eight bat species and five species groups were identified. Bat communities were affected by urbanisation in general and diversity was low in the entire study area. The community was dominated by the synurbic species Pipistrellus kuhlii, which comprised more than 70% of the total bat activity recorded. A positive relationship between built-up areas and bat activity was found, probably because P. kuhlii usually forages around streetlamps in urban areas. In contrast, vegetation cover did not affect bat activity, even in the less urbanised areas. The remainder of the bat species were not frequently recorded and were mostly detected close to water bodies, highlighting their value for foraging bats and the need for their conservation.
Urbanization induces rapid landscape and habitat modifications leading to alterations in species distribution patterns and biodiversity loss. As pollinating insects such as butterflies are particularly susceptible to urbanization, it is important to pinpoint the factors that could enhance their diversity in the urban areas in order to design adequate management and conservation actions. Our study aims to investigate the influence of land cover and local habitat characteristics on the butterfly diversity patterns and community structure in a densely built city in the eastern Mediterranean region. We carried out butterfly surveys (line transects) in 45 randomly selected sites, distributed along an urbanization gradient. In each site, we assessed the surrounding landscape by measuring the land cover in a 200-m buffer zone, and the local habitat by estimating the available plant resources along each transect. Overall, 1805 individuals belonging to 41 butterfly species were recorded. Land cover was found to have the strongest influence on butterfly species richness, abundance and community structure. Although plant resources were sufficiently available within the whole study area, the butterfly community was significantly poorer in the more urbanized areas, indicating the potential role of habitat fragmentation and patch isolation. In contrast, butterfly diversity was significantly higher in the peri-urban area, underlying its conservation value for butterflies in the urban landscape. We attribute these findings to the degradation of the more urbanized areas due to long-term inadequate planning and the disorganized expansion of the city.
Urbanization induces rapid landscape and habitat modifications leading to alterations in species distribution patterns and biodiversity loss. As pollinating insects such as butterflies are particularly susceptible to urbanization, it is important to pinpoint the factors that could enhance their diversity in the urban areas in order to design adequate management and conservation actions. Our study aims to investigate the influence of land cover and local habitat characteristics on the butterfly diversity patterns and community structure in a densely built city in the eastern Mediterranean region. We carried out butterfly surveys (line transects) in 45 randomly selected sites, distributed along an urbanization gradient. In each site, we assessed the surrounding landscape by measuring the land cover in a 200-m buffer zone, and the local habitat by estimating the available plant resources along each transect. Overall, 1805 individuals belonging to 41 butterfly species were recorded. Land cover was found to have the strongest influence on butterfly species richness, abundance and community structure. Although plant resources were sufficiently available within the whole study area, the butterfly community was significantly poorer in the more urbanized areas, indicating the potential role of habitat fragmentation and patch isolation. In contrast, butterfly diversity was significantly higher in the peri-urban area, underlying its conservation value for butterflies in the urban landscape. We attribute these findings to the degradation of the more urbanized areas due to long-term inadequate planning and the disorganized expansion of the city.
Red Lists are very valuable tools in nature conservation at global, continental and (sub-) national scales. In an attempt to prioritise conservation actions for European butterflies, we compiled a database with species lists and Red Lists of all European countries, including the Macaronesian archipelagos (Azores, Madeira and Canary Islands). In total, we compiled national species lists for 42 countries and national Red Lists for 34 of these. The most species-rich countries in Europe are Italy, Russia and France with more than 250 species each. Endemic species are mainly found on the Macaronesian archipelagos and on the Mediterranean islands. By attributing numerical values proportionate to the threat statuses in the different national Red List categories, we calculated a mean Red List value for every country (cRLV) and a weighted Red List value for every species (wsRLV) using the square root of the country’s area as a weighting factor. Countries with the highest cRLV were industrialised (NW) European countries such as the Netherlands, Belgium, the Czech Republic and Denmark, whereas large Mediterranean countries such as Spain and Italy had the lowest cRLV. Species for which a Red List assessment was available in at least two European countries and with a relatively high wsRLV (≥ 50) are Colias myrmidone, Pseudochazara orestes, Tomares nogelii, Colias chrysotheme and Coenonympha oedippus. We compared these wsRLVs with the species statuses on the European Red List to identify possible mismatches. We discuss how this complementary method can help to prioritise butterfly conservation on the continental and/or the (sub-)national scale.
Red Lists are very valuable tools in nature conservation at global, continental and (sub-) national scales. In an attempt to prioritise conservation actions for European butterflies, we compiled a database with species lists and Red Lists of all European countries, including the Macaronesian archipelagos (Azores, Madeira and Canary Islands). In total, we compiled national species lists for 42 countries and national Red Lists for 34 of these. The most species-rich countries in Europe are Italy, Russia and France with more than 250 species each. Endemic species are mainly found on the Macaronesian archipelagos and on the Mediterranean islands. By attributing numerical values proportionate to the threat statuses in the different national Red List categories, we calculated a mean Red List value for every country (cRLV) and a weighted Red List value for every species (wsRLV) using the square root of the country’s area as a weighting factor. Countries with the highest cRLV were industrialised (NW) European countries such as the Netherlands, Belgium, the Czech Republic and Denmark, whereas large Mediterranean countries such as Spain and Italy had the lowest cRLV. Species for which a Red List assessment was available in at least two European countries and with a relatively high wsRLV (≥ 50) are Colias myrmidone, Pseudochazara orestes, Tomares nogelii, Colias chrysotheme and Coenonympha oedippus. We compared these wsRLVs with the species statuses on the European Red List to identify possible mismatches. We discuss how this complementary method can help to prioritise butterfly conservation on the continental and/or the (sub-)national scale.
In this study we investigate the environmental factors influencing butterfly communities and evaluate the Natura 2000 network’s effectiveness in representing butterfly species richness and abundance, taking as a case study the island of Cyprus. We sampled butterflies and 11 environmental factors in 60 randomly selected sites across four 500-m elevation zones, representing seven habitat types. Rural mosaics and riverine vegetation were the habitats with the highest diversity of butterflies. Within habitats, the number of flower heads was the most important factor favouring butterfly species richness and abundance and endemic butterfly richness, while soil humidity had a positive effect on species richness and abundance. Although the Natura 2000 network succeeds in including the majority of butterfly species and all Cyprian endemics, the transects sampled within the network did not support more butterfly species than those outside it, and were significantly poorer in terms of butterfly abundance and endemic butterfly species richness and abundance. We found a similar pattern for the Habitats Directive priority habitats, which held poorer overall and endemic butterfly communities than the other habitats. The effectiveness of existing protected area networks may need to be reassessed in regions such as the South East Mediterranean, to ensure that regionally important components of biological diversity are adequately protected. To this aim, our results suggest that new European and national policies as well as further inclusion of rural mosaics and riverine habitats in protected area networks are needed for effective butterfly conservation in Cyprus.
In this study we investigate the environmental factors influencing butterfly communities and evaluate the Natura 2000 network’s effectiveness in representing butterfly species richness and abundance, taking as a case study the island of Cyprus. We sampled butterflies and 11 environmental factors in 60 randomly selected sites across four 500-m elevation zones, representing seven habitat types. Rural mosaics and riverine vegetation were the habitats with the highest diversity of butterflies. Within habitats, the number of flower heads was the most important factor favouring butterfly species richness and abundance and endemic butterfly richness, while soil humidity had a positive effect on species richness and abundance. Although the Natura 2000 network succeeds in including the majority of butterfly species and all Cyprian endemics, the transects sampled within the network did not support more butterfly species than those outside it, and were significantly poorer in terms of butterfly abundance and endemic butterfly species richness and abundance. We found a similar pattern for the Habitats Directive priority habitats, which held poorer overall and endemic butterfly communities than the other habitats. The effectiveness of existing protected area networks may need to be reassessed in regions such as the South East Mediterranean, to ensure that regionally important components of biological diversity are adequately protected. To this aim, our results suggest that new European and national policies as well as further inclusion of rural mosaics and riverine habitats in protected area networks are needed for effective butterfly conservation in Cyprus.

2018

This study examined the effects of pastoralism, including cattle grazing, on populations of three species of locally endemic and rare Peripodisma grasshoppers in calcareous grassland mountain habitats of northwestern Greece and southern Albania. The three Peripodisma species are on the IUCN Red List as near threatened, endangered, and critically endangered species, and cattle grazing had been identified as a key threat to the species. The study sites represented 70% of the known locations of Peripodisma genus. The region was historically grazed by local breeds of nomadic sheep and goats, but grazing practices had recently shifted to cattle grazing from non-local cattle breeds. We found a clear relationship between local abundance of Peripodisma and overall richness of Orthoptera communities. Orthoptera richness decreased at sites with medium to high impacts of livestock grazing. Cattle grazing had significant adverse effects on overall Orthoptera species richness and on Peripodisma abundance. Further studies are urgently needed to gather more data and information to guide grazing management and conservation planning that will provide a more balanced coexistence between livestock and Orthoptera, especially for the rare Peripodisma species that are in dire need of conservation management.
This study examined the effects of pastoralism, including cattle grazing, on populations of three species of locally endemic and rare Peripodisma grasshoppers in calcareous grassland mountain habitats of northwestern Greece and southern Albania. The three Peripodisma species are on the IUCN Red List as near threatened, endangered, and critically endangered species, and cattle grazing had been identified as a key threat to the species. The study sites represented 70% of the known locations of Peripodisma genus. The region was historically grazed by local breeds of nomadic sheep and goats, but grazing practices had recently shifted to cattle grazing from non-local cattle breeds. We found a clear relationship between local abundance of Peripodisma and overall richness of Orthoptera communities. Orthoptera richness decreased at sites with medium to high impacts of livestock grazing. Cattle grazing had significant adverse effects on overall Orthoptera species richness and on Peripodisma abundance. Further studies are urgently needed to gather more data and information to guide grazing management and conservation planning that will provide a more balanced coexistence between livestock and Orthoptera, especially for the rare Peripodisma species that are in dire need of conservation management.
Question: What are the woody vegetation encroachment patterns after agricultural land abandonment? Focusing on two parameters, woody plant species richness and vertical vegetation heterogeneity (number of different vegetation strata and their relative cover) we investigated: (a) the effect of forest encroachment following land abandonment; (b) the comparative importance of forest encroachment vis-a-vis topographic and climatic parameters, and finally; (c) the ecological importance of eight land-cover types encountered in abandoned agricultural landscapes. Location: The Balkan Peninsula (Albania, Bulgaria, Croatia, Greece). Methods: We set up a standard methodology for 1 × 1 km site selection (70 sites) and data collection within 497 plots, along a well-defined forest encroachment gradient that reflects land abandonment in terms of woody vegetation cover. Results: The pattern that emerged was neither clear nor common for the Balkan region, regarding the effect of forest encroachment on the woody plant species richness in young forests, formed from 20 to 50 years after land abandonment. However, at national level, species diversity was significantly affected by elevation (Bulgaria and Croatia) and temperature (Croatia), with lower and cooler areas being richer. Elevation was of great importance in determining vertical vegetation heterogeneity. Woodlots, broad-leaved forests, hedges and shrublands held the highest woody species richness in comparison to the more open land-cover types, and vertical vegetation heterogeneity was higher in open forests and woodlots. Conclusions: We expect the expansion of broad-leaved forests following land abandonment to enhance woody species richness. However, other land-cover types that were found to be important for woody plants should be maintained. As woody plants play a key role in supporting overall biodiversity, by providing suitable habitat for many species, we consider the preservation of a mosaic of patches of different land-cover types essential for the conservation of both plant and animal species diversity. Young forests should be preserved at intermediate stages of succession, through intermediate disturbance activities, including medium intensity grazing and the enhancement of wild ungulates.
Question: What are the woody vegetation encroachment patterns after agricultural land abandonment? Focusing on two parameters, woody plant species richness and vertical vegetation heterogeneity (number of different vegetation strata and their relative cover) we investigated: (a) the effect of forest encroachment following land abandonment; (b) the comparative importance of forest encroachment vis-a-vis topographic and climatic parameters, and finally; (c) the ecological importance of eight land-cover types encountered in abandoned agricultural landscapes. Location: The Balkan Peninsula (Albania, Bulgaria, Croatia, Greece). Methods: We set up a standard methodology for 1 × 1 km site selection (70 sites) and data collection within 497 plots, along a well-defined forest encroachment gradient that reflects land abandonment in terms of woody vegetation cover. Results: The pattern that emerged was neither clear nor common for the Balkan region, regarding the effect of forest encroachment on the woody plant species richness in young forests, formed from 20 to 50 years after land abandonment. However, at national level, species diversity was significantly affected by elevation (Bulgaria and Croatia) and temperature (Croatia), with lower and cooler areas being richer. Elevation was of great importance in determining vertical vegetation heterogeneity. Woodlots, broad-leaved forests, hedges and shrublands held the highest woody species richness in comparison to the more open land-cover types, and vertical vegetation heterogeneity was higher in open forests and woodlots. Conclusions: We expect the expansion of broad-leaved forests following land abandonment to enhance woody species richness. However, other land-cover types that were found to be important for woody plants should be maintained. As woody plants play a key role in supporting overall biodiversity, by providing suitable habitat for many species, we consider the preservation of a mosaic of patches of different land-cover types essential for the conservation of both plant and animal species diversity. Young forests should be preserved at intermediate stages of succession, through intermediate disturbance activities, including medium intensity grazing and the enhancement of wild ungulates.
Many have asserted that Sacred Natural Sites (SNS) play an important role in nature protection but few have assessed their conservation effectiveness for different taxa. We studied sacred groves in Epirus, NW Greece, where a large number of such SNS have been identified. Based on historical, ethnographic and ecological criteria, we selected eight of these groves and matching control sites and in them we studied fungi, lichens, herbaceous plants, woody plants, nematodes, insects, bats and passerine birds. Our results reveal that the contribution of SNS to species conservation is nuanced by taxon, vegetation type and management history. We found that the sacred groves have a small conservation advantage over the corresponding control sites. More specifically, there are more distinct sets of organisms amongst sacred groves than amongst control sites, and overall biodiversity, diversity per taxonomic group, and numbers of species from the European SCI list (Species of Community Interest) are all marginally higher in them. Conservationists regard the often small size of SNS as a factor limiting their conservation value. The sizes of SNS around the globe vary greatly, from a few square meters to millions of hectares. Given that those surveyed by us (ranging from 5 to 116 ha) are at the lower end of this spectrum, the small conservation advantage that we testified becomes important. Our results provide clear evidence that even small-size SNS have considerable conservation relevance; they would contribute most to species conservation if incorporated in networks.
Many have asserted that Sacred Natural Sites (SNS) play an important role in nature protection but few have assessed their conservation effectiveness for different taxa. We studied sacred groves in Epirus, NW Greece, where a large number of such SNS have been identified. Based on historical, ethnographic and ecological criteria, we selected eight of these groves and matching control sites and in them we studied fungi, lichens, herbaceous plants, woody plants, nematodes, insects, bats and passerine birds. Our results reveal that the contribution of SNS to species conservation is nuanced by taxon, vegetation type and management history. We found that the sacred groves have a small conservation advantage over the corresponding control sites. More specifically, there are more distinct sets of organisms amongst sacred groves than amongst control sites, and overall biodiversity, diversity per taxonomic group, and numbers of species from the European SCI list (Species of Community Interest) are all marginally higher in them. Conservationists regard the often small size of SNS as a factor limiting their conservation value. The sizes of SNS around the globe vary greatly, from a few square meters to millions of hectares. Given that those surveyed by us (ranging from 5 to 116 ha) are at the lower end of this spectrum, the small conservation advantage that we testified becomes important. Our results provide clear evidence that even small-size SNS have considerable conservation relevance; they would contribute most to species conservation if incorporated in networks.
The sacred groves in the mountains of Epirus in NW Greece have been established during the Ottoman period and consist of locally adapted systems set apart from the surrounding intensively managed, anthropogenic landscape. We inventoried eight sacred groves and compared them with nearby control (managed) forests. In total, 166 taxa of lichens and five of lichenicolous fungi were recorded. The most common lichen species were Anaptychia ciliaris, Phlyctis argena and Lecidella elaeochroma. Seven species are new for Greece: Calicium quercinum, Chaenotheca ferruginea, Chaenotheca trichialis, Chaenothecopsis nana, Leptogium hibernicum, Parvoplaca nigroblastidiata and Rinodina orculata. The sacred groves appeared not very different from the control forests; more pronounced differences were observed between deciduous oak evergreen oak and pine forests. Localities characterized by deciduous oak forest hosted the highest number of taxa belonging to the order Peltigerales, the most frequent were: Nephroma laevigatum, Collema subflaccidum, Leptogium lichenoides and Lobaria pulmonaria, but also rare species such as Polychidium muscicola, Koerberia biformis and Degelia atlantica were recorded.
The sacred groves in the mountains of Epirus in NW Greece have been established during the Ottoman period and consist of locally adapted systems set apart from the surrounding intensively managed, anthropogenic landscape. We inventoried eight sacred groves and compared them with nearby control (managed) forests. In total, 166 taxa of lichens and five of lichenicolous fungi were recorded. The most common lichen species were Anaptychia ciliaris, Phlyctis argena and Lecidella elaeochroma. Seven species are new for Greece: Calicium quercinum, Chaenotheca ferruginea, Chaenotheca trichialis, Chaenothecopsis nana, Leptogium hibernicum, Parvoplaca nigroblastidiata and Rinodina orculata. The sacred groves appeared not very different from the control forests; more pronounced differences were observed between deciduous oak evergreen oak and pine forests. Localities characterized by deciduous oak forest hosted the highest number of taxa belonging to the order Peltigerales, the most frequent were: Nephroma laevigatum, Collema subflaccidum, Leptogium lichenoides and Lobaria pulmonaria, but also rare species such as Polychidium muscicola, Koerberia biformis and Degelia atlantica were recorded.

2017

Our understanding of arthropod responses to environmental pressures is limited, especially for the poorly studied Mediterranean region. In the light of likely further environmental change and the need for protocols for rapid biodiversity assessment, we measured how the abundance and species richness of two taxa, ground spiders and Orthoptera, belonging to different functional groups, fluctuates intra- seasonally (early-mid-late summer) and across habitat types (grasslands, maquis, forests). We also tested their surrogate value. Spiders were found to have higher species richness and abundance almost throughout the investigation. Orthoptera had lower species richness and abundance in forests compared to grasslands and maquis, while no significant difference between habitats was revealed for spiders. Early-summer was the richest period for spiders while mid-summer was the richest for Orthoptera. Canopy cover was found to significantly influence community composition of both groups, while herb height and cover of stones was a determinant factor for Orthoptera only. There was a significant congruence between the two groups and Orthoptera provided the best complementary network. Our results show that diversity patterns of both spiders and Orthoptera are sensitive to environmental changes even over short time-scales (e.g. within the summer period) and space (e.g. across different habitat types), suggesting that small inexpensive experimental designs may still reveal community dynamics. For conservation purposes, we advise a focus on variables regulating habitat heterogeneity and microhabitat characteristics. We provide a list of the most influential species and propose the most effective network for obtaining information on the local fauna.
Our understanding of arthropod responses to environmental pressures is limited, especially for the poorly studied Mediterranean region. In the light of likely further environmental change and the need for protocols for rapid biodiversity assessment, we measured how the abundance and species richness of two taxa, ground spiders and Orthoptera, belonging to different functional groups, fluctuates intra- seasonally (early-mid-late summer) and across habitat types (grasslands, maquis, forests). We also tested their surrogate value. Spiders were found to have higher species richness and abundance almost throughout the investigation. Orthoptera had lower species richness and abundance in forests compared to grasslands and maquis, while no significant difference between habitats was revealed for spiders. Early-summer was the richest period for spiders while mid-summer was the richest for Orthoptera. Canopy cover was found to significantly influence community composition of both groups, while herb height and cover of stones was a determinant factor for Orthoptera only. There was a significant congruence between the two groups and Orthoptera provided the best complementary network. Our results show that diversity patterns of both spiders and Orthoptera are sensitive to environmental changes even over short time-scales (e.g. within the summer period) and space (e.g. across different habitat types), suggesting that small inexpensive experimental designs may still reveal community dynamics. For conservation purposes, we advise a focus on variables regulating habitat heterogeneity and microhabitat characteristics. We provide a list of the most influential species and propose the most effective network for obtaining information on the local fauna.
Mountains are complex ecosystems supporting a great variety of taxa. Here, we explored the diversity patterns of arthropods in two mountains, pinpointing the spatial scale that accounts most for overall diversity variation, using an additive partitioning framework. Butterflies and Orthoptera were sampled in Rodopi (2012) and Grammos (2013) mountains. Diversity was partitioned into five hierarchical levels (mountain, elevational zone, habitat, transect and plot). We compared the estimated diversity values for each level to the respective permuted values expected by chance, for all species, as well as for species identified as “rare” or “common”. At broader spatial levels, the variation in total diversity was attributed to the beta diversity component: mountains accounted for 20.94 and 26.25% of butterfly and Orthoptera diversity, and elevational zones accounted for 28.94 and 35.87% respectively. At finer spatial scales, beta diversity was higher than expected by chance in terms of the Shannon index. The type of habitat was found to play a significant role only for rare orthopterans. Finally, common species were recognized for shaping overall species diversity. We highlight the importance of the spatial levels of elevation zone and then mountain position in conservation planning, due to the greater beta diversity recorded at this scale as compared to habitat or more finite scales. Monitoring programs might need to adapt different strategies with respect to the focal organisms, and consider patterns of common rather than rare species that found to drive the patterns of the entire community.
Mountains are complex ecosystems supporting a great variety of taxa. Here, we explored the diversity patterns of arthropods in two mountains, pinpointing the spatial scale that accounts most for overall diversity variation, using an additive partitioning framework. Butterflies and Orthoptera were sampled in Rodopi (2012) and Grammos (2013) mountains. Diversity was partitioned into five hierarchical levels (mountain, elevational zone, habitat, transect and plot). We compared the estimated diversity values for each level to the respective permuted values expected by chance, for all species, as well as for species identified as “rare” or “common”. At broader spatial levels, the variation in total diversity was attributed to the beta diversity component: mountains accounted for 20.94 and 26.25% of butterfly and Orthoptera diversity, and elevational zones accounted for 28.94 and 35.87% respectively. At finer spatial scales, beta diversity was higher than expected by chance in terms of the Shannon index. The type of habitat was found to play a significant role only for rare orthopterans. Finally, common species were recognized for shaping overall species diversity. We highlight the importance of the spatial levels of elevation zone and then mountain position in conservation planning, due to the greater beta diversity recorded at this scale as compared to habitat or more finite scales. Monitoring programs might need to adapt different strategies with respect to the focal organisms, and consider patterns of common rather than rare species that found to drive the patterns of the entire community.
Urbanization causes rapid changes in the landscape and land use, exerting a significant pressure on bird communities. The effect of urbanization on bird diversity has been widely investigated in many cities worldwide; however, our knowledge on urban bird communities from the eastern Mediterranean region is very scarce. In this context, we aimed to investigate the effect of the different land-cover types on bird species richness and abundance in a densely built coastal Mediterranean city (Patras, Greece) during the breeding and wintering seasons. We sampled the bird community in 90 randomly selected sites along an urbanization gradient. Open green spaces proved to be the most significant factor favouring bird diversity in both seasons. In winter, woody vegetation and impervious surfaces had a positive effect on species richness as well. The local bird community consisted of a large number of species associated with open and semi-open unmanaged green areas, 12 of which are Species of European Conservation Concern (SPECs) showing a declining trend in Europe. On the other hand, in winter the number of forest-dwellers increased significantly. Species richness was significantly higher in winter indicating that the urban environment provides important wintering grounds. Thus, management actions in cities with similar characteristics in the Mediterranean region should focus on the maintenance of open green spaces and woody vegetation patches to enhance bird diversity.
Urbanization causes rapid changes in the landscape and land use, exerting a significant pressure on bird communities. The effect of urbanization on bird diversity has been widely investigated in many cities worldwide; however, our knowledge on urban bird communities from the eastern Mediterranean region is very scarce. In this context, we aimed to investigate the effect of the different land-cover types on bird species richness and abundance in a densely built coastal Mediterranean city (Patras, Greece) during the breeding and wintering seasons. We sampled the bird community in 90 randomly selected sites along an urbanization gradient. Open green spaces proved to be the most significant factor favouring bird diversity in both seasons. In winter, woody vegetation and impervious surfaces had a positive effect on species richness as well. The local bird community consisted of a large number of species associated with open and semi-open unmanaged green areas, 12 of which are Species of European Conservation Concern (SPECs) showing a declining trend in Europe. On the other hand, in winter the number of forest-dwellers increased significantly. Species richness was significantly higher in winter indicating that the urban environment provides important wintering grounds. Thus, management actions in cities with similar characteristics in the Mediterranean region should focus on the maintenance of open green spaces and woody vegetation patches to enhance bird diversity.

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