The relationships between habitat topology, critical scales of connectivity and tick abundance Ixodes ricinus in a heterogeneous landscape in northern Spain

The habitat mosaic was used to quantify connectivity between patches of different tick density of the notorious tick species Ixodes ricinus in an attempt to determine the cause of variations in tick abundance among apparently homogeneous sites in northern Spain. The analysis revealed that patches with high tick abundance are "stepping-stone" territories that, when removed from the landscape, cause large changes in connectivity. Sites with medium tick abundance do not cause such a critical transition in connectivity. Patches with low tick abundance, but optimal abiotic conditions for survival, are located within the minimum cost corridors network joining the patches, while those sites where the tick has been intermittently collected are located at variable distances from this network. Sites where the tick is consistently absent, but where the habitat is predicted to be suitable (old, heterogeneous forests of Quercus spp.) for the tick, are very separated from this main network of connections. These results suggest that tick distribution in a zone is highly affected not only by abiotic variables (vegetation and weather) but also by host movements. Dispersal of the tick is a function of how the hosts perceive the habitat, and the habitat's permeability to host movement. Permanent tick populations seem to be supported by the existence of these critical, high density patches, located at significant places within the habitat network.     

Patch occupancy in the endangered butterfly Lycaena helle in a fragmented landscape: effects of habitat quality, patch size and isolation

While there is agreement that both habitat quality and habitat network characteristics (such as patch size and isolation) contribute to the occupancy of patches by any given species, the relative importance of these factors is under debate. This issue is of fundamental ecological importance, and moreover of special concern for conservation biologists aiming at preserving endangered species. Against this background we investigated patch occupancy in the violet copper Lycaena helle, one of the rarest butterfly species in Central Europe, in the Westerwald area (Rhineland-Palatinate, Western Germany). Occupied (n = 102) differed from vacant (n = 128) patches in altitude, size, connectivity, availability of wind shelter, in the abundance of the larval host-plant, in the abundance of a grass species indicating favorable habitat conditions and in the abundance of nitrophilous plants. Overall, patch occupancy was primarily determined by patch size, connectivity and the abundance of the larval host plant, while all other parameters of habitat quality were of subordinate importance. Therefore, our findings suggest that even for extremely sedentary species such as L. helle habitat networks are decisive and—next to the preservation of habitat quality—need to be an integral part of any conservation management for this species.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

Bird assemblages in fragmented agricultural landscapes: the role of small brigalow remnants and adjoining land uses

Agricultural intensification typically leads to changes in bird diversity and community composition, with fewer species and foraging guilds present in more intensively managed parts of the landscape. In this study, we compare bird communities in small (2–32 ha) brigalow (Acacia harpophylla) remnants with those in adjacent uncultivated grassland, previously cultivated grassland and current cropland, to determine the contribution of different land uses to bird diversity in the agricultural landscape. Twenty remnant brigalow patches and adjacent agricultural (‘matrix’) areas in southern inland Queensland, Australia were sampled for bird composition and habitat characteristics. The richness, abundance and diversity of birds were all significantly higher in brigalow remnants than in the adjacent matrix of cropping and grassland. Within the matrix, species richness and diversity were higher in uncultivated grasslands than in current cultivation or previously cultivated grasslands. Forty-four percent of bird species were recorded only in brigalow remnants and 78% of species were recorded in brigalow and at least one other land management category. Despite high levels of landscape fragmentation and modification, small patches of remnant brigalow vegetation provide important habitat for a unique and diverse assemblage of native birds. The less intensively managed components of the agricultural matrix also support diverse bird assemblages and thus, may be important for local and regional biodiversity conservation.     

Stochastic models for mainland-island metapopulations in static and dynamic landscapes

This paper has three primary aims: to establish an effective means for modelling mainland-island metapopulations inhabiting a dynamic landscape; to investigate the effect of immigration and dynamic changes in habitat on metapopulation patch occupancy dynamics; and to illustrate the implications of our results for decision-making and population management. We first extend the mainland-island metapopulation model of Alonso and McKane [Bull. Math. Biol. 64:913–958, 2002] to incorporate a dynamic landscape. It is shown, for both the static and the dynamic landscape models, that a suitably scaled version of the process converges to a unique deterministic model as the size of the system becomes large. We also establish that, under quite general conditions, the density of occupied patches, and the densities of suitable and occupied patches, for the respective models, have approximate normal distributions. Our results not only provide us with estimates for the means and variances that are valid at all stages in the evolution of the population, but also provide a tool for fitting the models to real metapopulations. We discuss the effect of immigration and habitat dynamics on metapopulations, showing thatmainland-like patches heavily influence metapopulation persistence, and we argue for adopting measures to increase connectivity between this large patch and the other island-like patches. We illustrate our results with specific reference to examples of populations of butterfly and the grasshopperBryodema tuberculata.     

Does fragmentation of Urtica habitats affect phytophagous and predatory insects differentially?

Effects of habitat fragmentation on the insect community of stinging nettle (Urtica dioica L.) were studied, using 32 natural nettle patches of different area and degree of isolation in an agricultural landscape. Habitat fragmentation reduced the species richness of Heteroptera, Auchenorrhyncha, and Coleoptera, and the abundance of populations. Habitat isolation and area reduction did not affect all insect species equally. Monophagous herbivores had a higher probability of absence from small patches than all (monophagous and polyphagous) herbivore species, and the percentage of monophagous herbivores increased with habitat area. Abundance and population variability of species were negatively correlated and could both be used as a predictor of the percentage of occupied habitats. Species richness of herbivores correlated (positively) with habitat area, while species richness of predators correlated (negatively) with habitat isolation. In logistic regressions, the probability of absence of monophagous herbivores from habitat patches could only be explained by habitat area (in 4 out of 10 species) and predator absence probability only by habitat isolation (in 3 out of 14 species). Presumably because of the instability of higher-trophic-level populations and dispersal limitation, predators were more affected by habitat isolation than herbivores, while they did not differ from herbivore populations with respect to abundance or variability. Thus increasing habitat connectivity in the agricultural landscape should primarily promote predator populations. Received: 4 February 1998 / Accepted: 4 May 1998     

Differential effects of spatial network structure and scale on population size and genetic diversity of the ninespine stickleback in a remnant wetland system

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Rapid recovery of an insect–plant interaction following habitat loss and experimental wetland restoration

This study examined the impact of wetland habitat loss and isolation on an insect–plant interaction, and the subsequent rate of recovery of the interaction following experimental habitat restoration. We compared herbivore colonisation rates and herbivory damage by ‘Batrachedra’ sp. (Lepidoptera: Coleophoridae) on experimentally placed potted Sporadanthus ferrugineus (Restionaceae) plants at increasing distances (up to 800 m) from an intact habitat (the source population). These tests showed that even a moderate degree of isolation (i.e. greater than 400 m) from the intact wetland habitat caused an almost complete collapse of the insect–plant interaction, at least in the short term. The number of eggs and larvae of colonising ‘Batrachedra’ sp., as well as average larval size and the proportion of S. ferrugineus stems damaged, all decreased logarithmically with increasing distance from the intact habitat, presumably due to dispersal limitation of the herbivore. Subsequently, to test whether the interaction can recover following habitat restoration, we surveyed herbivore colonisation rates and herbivory damage on naturally regenerated S. ferrugineus plants on experimentally restored ‘islands’ at increasing distances (up to 800 m) from an intact habitat. The rate of recovery of the interaction was surprisingly rapid (i.e. between 196 and 308 weeks). The degree of difference in the density of eggs and larvae, and in the proportion of stems damaged with increasing isolation from the intact wetland, gradually diminished over 196 weeks. After 308 weeks there was no significant difference in the insect–plant interaction between the intact wetland sites and any of the experimentally restored sites up to 800 m away. These results suggest that some insect–plant interactions can recover rapidly from habitat loss with restoration management.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Boundary nature induces greater group size and group density in habitat edges: an agent-based model revealed

Apart from the amount and distribution of food resources and predation pressure, we sought to clarify habitat spatial features that affect group size and group density. To simulate animal groups interacting with one another, we made an agent-based model in a computational space. Assuming no predation pressure, we change the resource conditions between the habitat (comprising ‘interior’ and ‘edges’) and the ‘exterior’. The results are as follows: when the group density is high throughout the habitat, large groups tend to frequent the edges (i.e., the boundaries between the interior and exterior) even if food resources are scarce in the exterior. Additionally, when the group density is high and the group size variation is small, both group size and group density increase at the edges. These findings are discussed with reference to primates, particularly the Japanese macaque (Macaca fuscata) as an example of group-living animal.     

Is there a link between the type of habitat and the patterns of abundance of holothurians in shallow rocky reefs?

The presence of a mosaic of habitats, largely determined by sea urchin grazing, across shallow rocky reefs may potentially influence in differences in the distribution patterns of invertebrates. The aim of this paper was to assess, using a correlative approach, whether the type of habitat influences the abundance patterns of holothurians in the eastern Atlantic. We hypothesized that abundances of large (> 10 cm) holothurians varied among four types of habitat (3 vegetated habitats with low abundances of the sea urchin D. antillarum vs. ‘barrens’ with hyperabundances of sea urchins), and that these differences were consistent at a hierarchy of spatial scales, including two islands and several replicated sites within each type of habitat and island. Three species of large holothurians were found, accounting for a total of 300 specimens. We found remarkable differences in abundances of holothurians between the ‘barrens’ and the three vegetated habitats. This pattern was strongest for the numerically dominant species, Holothuria sanctorii. Total abundances of holothurians were between 5 and 46 times more abundant in ‘barrens’ compared with the vegetated habitats. Inter-habitat differences were species-specific with some inconsistent patterns from one island to the other. The total abundances of holothurians tended to increase with the abundance of sea urchins within ‘barrens’. Our study suggests that there may be a link, at least for the dominant species Holothuria sanctorii, between the distribution and abundances of large holothurians and the habitat across shallow-waters of the eastern Atlantic.     

Spatial Graphs: Principles and Applications for Habitat Connectivity

ABSTRACT Well-founded methods to assess habitat connectivity are essential to inform land management decisions that include conservation and restoration goals. Indeed, to be able to develop a conservation plan that maintains animal movement through a fragmented landscape, spatial locations of habitat and paths among them need to be represented. Graph-based approaches have been proposed to determine paths among habitats at various scales and dispersal movement distances, and balance data requirements with information content. Conventional graphs, however, do not explicitly maintain geographic reference, reducing communication capacity and utility of other geo-spatial information. We present spatial graphs as a unifying theory for applying graph-based methods in a geographic context. Spatial graphs integrate a geometric reference system that ties patches and paths to specific spatial locations and spatial dimensions. Arguably, the complete graph, with paths between every pair of patches, may be one of the most relevant graphs from an ecosystem perspective, but it poses challenges to compute, process and visualize. We developed Minimum Planar Graphs as a spatial generalization of Delaunay triangulations to provide a reasonable approximation of complete graphs that facilitates visualization and comprehension of the network of connections across landscapes. If, as some authors have suggested, the minimum spanning tree identifies the connectivity “backbone” of a landscape, then the Minimum Planar Graph identifies the connectivity “network”. We applied spatial graphs, and in particular the Minimum Planar Graph, to analyze woodland caribou habitat in Manitoba, Canada to support the establishment of a national park.     

Determinants of distribution and abundance in the clouded apollo butterfly: a landscape ecological approach

Recent studies on the determinants of distribution and abundance of animals at landscape level have emphasized the usefulness of the metapopulation approach, in which patch area and habitat connectivity have often proved to explain satisfactorily existing patch occupancy patterns. A different approach is needed to study the common situation in which suitable habitat is difficult to determine or does not occur in well‐defined habitat patches. We applied a landscape ecological approach to study the determinants of distribution and abundance of the threatened clouded apollo Parnassius mnemosyne butterfly within an area of 6 km² of agricultural landscape in south‐western Finland. The relative role of 24 environmental variables potentially affecting the distribution and abundance of the butterfly was studied using a spatial grid system with 2408 grid squares of 0.25 ha, of which 349 were occupied by the clouded apollo. Both the probability of butterfly presence and abundance in a 0.25 ha square increased with the presence of the larval host plant Corydalis solida the cover of semi‐natural grassland, the amount of solar radiation and spalial autocorrelation in butterfly occurrence. Additionally, butterfly abundance increased with overall mean patch size and decreased with maximum slope angle and wind speed. Two advantages of the employment of a spatial grid system included the avoidance of a subjective definition of suitable habitat patches and an evaluation of the relative significance of different components of habitat quality at the same time with habitat availability and connectivity. The large variation in habitat quality was influenced by the abundance of the larval host plant and adult nectar sources but also by climatological. topographical and structural factors. The application of a spatial grid system as used here has potential for a wide use in studies on landscape‐level distribution and abundance patterns in species with complex habitat requirements and habitat availability patterns.     

Small plot mating disruption trials for tufted apple bud moth,Platynota idaeusalis, in Pennsylvania apple orchards

Mating disruption treatments for the tufted apple bud moth (TABM),Platynota idaeusalis (Walker), were tested in small plot trials in apple orchards in Pennsylvania. Treatments were evaluated by fruit injury and by capture of male TABM in traps baited with synthetic pheromone sources or virgin females. The TABM pheromone is a two component isomeric blend ofE-11-tetradecen-1-ol (E11-14:OH) andE-11-tetradecenyl acetate (E11-14:Ac). A 50∶50 ratio of these two components was used in standard monitoring septa and in mating disruption treatments released from either hollow fibers (‘fiber’) or PVC tubes (‘PVC’). Other pheromone blends tested included a 90∶10 ratio of E11-14: Ac and E11-14:OH (‘EAc’) and its reverse (‘EOH’), mixture ‘EAc’ with 30% of the Z-isomers (‘low AEc’), and a blend similar to the preceding with 2% Z9-12:Ac (‘generic’). These other blends were released from multi tube tape (‘tape’) or Shin-Etsu type rope (‘rope’) dispensers. Seasonal dispenser release rate in mg ha⁻¹ h⁻¹ was ca. 30 for the ‘rope’ dispensers, 14 for ‘PVC’ and 6 for ‘fiber’. ‘EAc-tape’ and ‘EOH-tape’ were equally effective in reducing catches of males in traps baited with synthetic lures and in traps baited with virgin females. Both treatments also reduced fruit injury. ‘EAc-rope’, ‘fiber’ and ‘PVC’ also were generally effective; whereas, the ‘low EAc’ and ‘generic’ treatments reduced trap capture less than 90% and did not reduce fruit injury. Dispenser density was positively correlated with reduction in trap capture for the ‘low EAc-rope’ and ‘genericrope’ treatments. Traps loaded with ‘fiber’ dispensers captured more male TABM than the other treatments in non-pheromone permeated environments. Trap capture of other tortricids was reduced in pheromone treatments. ‘EAc-rope’ and the ‘TABM’ treatments provided mean (s.e.) percent reduction in trap catch of 99.5 (0.4) and 42.9 (10.1), respectively, for the redbanded leafroller,Argyrotaenia velutinana (Walker), and 90.4 (6.8) and 90.4 (1.3), respectively, for the obliquebanded leafroller,Choristoneura rosaceana (Harris).     

气候变化对孑遗植物银杉的潜在分布及生境破碎度的影响

以孑遗植物银杉(Cathaya argyrophylla Chun et Kuang)为研究对象,选取65个地理分布记录和19个生物气候因子(bio1—bio19),利用MaxEnt模型预测四种不同浓度路径下(RCP 2.6、RCP 4.5、RCP 6.0和RCP 8.5),银杉在2050s和2070s两个年代的潜在分布变化,并利用景观指数对气候变化情景下银杉适宜生境空间格局特征转变及生境破碎度变化进行分析。结果表明:在当前气候情景下,银杉适宜生境面积约占研究区面积的14.32%,主要分布于北纬24°—32°、东经105°—114°之间,位于四川盆地东南地区、云贵高原东北地区、南岭西段地区以及浙闽丘陵的北部地区。在未来不同气候情景下,银杉适宜生境变化特征显著,面积呈增加趋势,形状上整体呈四周向中间聚集。气候变化对银杉适宜生境的景观指数影响主要表现在斑块数量增多、斑块密度增加、面积加权平均形状指数变大,对分离度与聚散性影响较小;气候变化对银杉生境破碎化程度的影响表现在破碎化两极现象减弱,总体破碎化程度加剧。研究选取7个景观指数并结合PCA法得到综合的破碎度指数来定量分析银杉适宜生境破碎化程度变化,相比单一指标的定量评价和多个指标的定性分析,更能代表银杉生境的实际破碎化程度。     

Colonisation of ephemeral forest habitats by specialised species: beetles and bugs associated with recently dead aspen wood

The most appropriate strategy for preserving fragmented populations depends on a species’ ability to colonise distant habitat patches. Insects associated with early decay stages of dead wood are expected to have a high capacity to colonise new habitat patches. To study the dispersal ranges of beetles (Coleoptera) and flat bugs (Hemiptera: Aradidae) dependent on recently dead aspen (Populus tremula) wood in Finland, we set out 58 piles of recently cut aspen logs at various distances up to 1.6 km from forests that contained a high density of old aspen trees. We captured insects by trunk window-traps, and counted beetles’ exit holes. Habitat connectivity was measured in terms of the amount of suitable aspen-wood in the surrounding environment, with the closest dead wood items up-weighted by a negative-exponential function. The log-piles attracted many saproxylic insects including four red-listed aspen-specialist species. The exposure of log-piles to the sun, and high levels of habitat connectivity increased the species richness of aspen-specialists, whereas bark peeling by moose decreased richness. The spatial scale at which species richness had its strongest response to habitat was 93 m. Among individual species there was a wide variability in spatial scale of response. This study supports the view that conservation efforts in boreal forests should be concentrated on sites where colonisation by target species is most likely. Restoration of habitat by re-locating logs may be useful at localities with a rich and specialised fauna but which have too low rate of formation of dead wood by natural processes.     

Effects of landscape complexity and stand factors on arthropod communities in poplar forests

The arthropod communities are influenced by both local conditions and features of the surrounding landscape. Landscape complexity and stand factors may both influence arthropod communities in poplar forests, but the multiscale effects of these factors on poplar defoliators and natural enemies are still poorly understood. We collected poplar arthropods at 30 sampling sites within five forest landscapes in Xinjiang, China, and assessed whether landscape complexity and stand factors influence species abundance and diversity of poplar arthropods. Landscape complexity was quantified by several independent metrics of landscape composition, configuration, and connectivity at three spatial scales. We also determined the most powerful explanatory variables and the scale effect of each arthropod. Results found that landscape complexity and stand factors had different effects on different poplar arthropod communities. Landscape complexity promoted natural enemies at different spatial scales, but it inhibited the population of poplar defoliators at the scale of 200 m. Specifically, the abundance and diversity of all defoliators decreased with increasing proportion of nonhost plants. Landscape diversity only had a negative effect on defoliator abundance. The shape complexity of habitat patches increased the abundance of carabid beetles but reduced the abundance of green leafhoppers and migratory locusts. The abundance and diversity of predators increased with increasing structural connectivity of forest landscape. Additionally, both the abundance and diversity of all defoliators were positively correlated with the average height of herbaceous plants. Diversity of all defoliators increased with increasing size of host trees. The distance from sampling site to the nearest village positively influenced the abundance and diversity of all predators. Arthropod abundance and diversity in poplar forests were driven by stand factors and landscape complexity. Therefore, maintaining complex shape and structural connectivity of habitat patches and keeping poplar stands away from the village are crucial for management of forest landscape to enhance natural enemies. And in order to reduce the abundance of defoliators in poplar forest, the diversity of surrounding habitat types should be promoted within 200 m radii.     

基于景观格局和连接度评价的生态网络方法优化与应用

景观生态学中常凭借最小累积阻力模型构建目标种生态网络,以提升破碎栖息地间的景观连接度,缓解生境破碎化负面影响.但传统最小累积阻力生态网络方法缺乏对生态网络的效用验证,对研究地的景观结构变化与生态过程的影响认识不足.本研究运用景观格局指数与连接度概率指数,定量评价生态网络构建前后的研究地景观结构与连接度特征,并以崇左白头叶猴栖息地生态网络为例,详尽叙述此生态网络方法的优化与应用过程.通过对白头叶猴栖息地斑块进行辨认、踏脚石斑块识别,对研究地用地类型进行划分并进行阻力赋值,运用最小累积阻力模型生成了20条白头叶猴栖息地生态网络廊道;然后利用景观结构指数与连接度概率指数结合的方法,对生成的生态网络结构和功能连接度进行评价.结果表明: 凭借最小累积阻力模型构建的目标种生态网络,能有效提升栖息地生境的完整性和连续性,降低总体破碎化水平,并改善生境质量.同时,该生态网络构建能提升生境景观的结构连接度与功能连接度,且两方面的连接度变化在结果上具有极显著的一致性(R²=98.3%,P<0.01).生态网络带来的景观结构方面变化与功能连接度的关联性不强,两种指数间的相互关系不如结构与功能的内在关系显著.     

Host plant density and patch isolation drive occupancy and abundance at a butterfly's northern range margin

Marginal populations are usually small, fragmented, and vulnerable to extinction, which makes them particularly interesting from a conservation point of view. They are also the starting point of range shifts that result from climate change, through a process involving colonization of newly suitable sites at the cool margin of species distributions. Hence, understanding the processes that drive demography and distribution at high‐latitude populations is essential to forecast the response of species to global changes. We investigated the relative importance of solar irradiance (as a proxy for microclimate), habitat quality, and connectivity on occupancy, abundance, and population stability at the northern range margin of the Oberthür's grizzled skipper butterfly Pyrgus armoricanus. For this purpose, butterfly abundance was surveyed in a habitat network consisting of 50 habitat patches over 12 years. We found that occupancy and abundance (average and variability) were mostly influenced by the density of host plants and the spatial isolation of patches, while solar irradiance and grazing frequency had only an effect on patch occupancy. Knowing that the distribution of host plants extends further north, we hypothesize that the actual variable limiting the northern distribution of P. armoricanus might be its dispersal capacity that prevents it from reaching more northern habitat patches. The persistence of this metapopulation in the face of global changes will thus be fundamentally linked to the maintenance of an efficient network of habitats.     

Remnants fragments preserve genetic diversity of the old forest lichen <Emphasis Type="Italic">Lobaria pulmonaria</Emphasis> in a fragmented Mediterranean mountain forest

Fragmentation represents a serious threat to biodiversity worldwide, however its effects on epiphytic organisms is still poorly understood. We study the effect of habitat fragmentation on the genetic population structure and diversity of the red-listed epiphytic lichen, Lobaria pulmonaria, in a Mediterranean forest landscape. We tested the relative importance of forest patch quality, matrix surrounding fragments and connectivity on the genetic variation within populations and the differentiation among them. A total of 855 thalli were sampled in 44 plots (400 m²) of 31 suitable forest fragments (beeches and oaks), in the Sierra de Ayllón in central Spain. Variables related to landscape attributes of the remnant forest patches such as size and connectivity and also the nature of the matrix or tree species had no significant effects on the genetic diversity of L. pulmonaria. Values of genetic diversity (Nei’s) were only affected by habitat quality estimated as the age patches. Most of the variation (76%) in all populations was observed at the smallest sampled unit (plots). Using multiple regression analysis, we found that habitat quality is more important in explaining the genetic structure of the L. pulmonaria populations than spatial distance. The relatively high level of genetic diversity of the species in old forest patches regardless of patch size indicates that habitat quality in a highly structured forest stand determines the population size and distribution pattern of this species and its associated lichen community. Thus, conservation programmes of Mediterranean mountain forests have to prioritize area and habitat quality of old forest patches.     

Landscape connectivity and the role of small habitat patches as stepping stones: an assessment of the grassland biome in South America

Connectivity losses lead to a reduction of the amount of habitat resources that can be reached and used by species, and hence to a decline in the ranges and abundance of multiple taxa. Despite the recognized important role of small habitat patches for many species inhabiting fragmented landscapes, their potential contribution as stepping stones for maintaining overall landscape connectivity has received less attention. Using connectivity metrics based on a graph-theoretic approach we (i) quantified the connectivity of grassland patches in a sector of the Pampa region in Argentina, using a range of dispersal distances (from 100 to 10,000 m) representative of the scale of dispersal of different species; (ii) identified the most relevant patches for maintaining overall connectivity; and (iii) studied the importance of small patches (defined for different area thresholds of 5, 20, and 50 ha) as connectivity providers in the landscape. Although grassland patches were in general poorly connected at all distances, some of them were critical for overall connectivity and were found to play different crucial roles in the patch network. The location of small patches in the grassland network allowed them to function as stepping stones, yielding significant connectivity gains for species that move large distances (>5000 m) for the three area thresholds considered. Thus, under the spatial pattern of the studied landscape, species that move long distances would benefit from stepping stones, while less mobile organisms would benefit from, and mostly rely on the largest patches. We recommend that future management activities should (i) aim at preserving the grassland patches with the highest potential as stepping stones to promote landscape-level connectivity; and (ii) pay more attention to the conservation of key small patches, particularly given that usually they are those more vulnerable to land clearing for agriculture.