Effects of snow accumulation on survival of beech (Fagus crenata) seed

The escape of beech seeds from seed predators and winter desiccation due to snow accumulation was studied by comparing two sites in Japan: one site that experiences much snow and another site that experiences less snow cover. At the site with greater snow cover, about 70% of the beech seeds escaped seed predation by rodents during winter and about 70% of surviving seeds germinated successfully in spring. At the site with less snow cover, however, all of the beech seeds were eaten by rodents, and all seeds that were protected from feeding were killed by winter desiccation. We confirmed that snow prevents beech seeds from predation by rodents because it conceals their sight and scent. These effects are one of the main reasons why beeches in snowy areas regenerate constantly and those in less snowy areas do not.     

Early reduction of post-fire recruitment of Pinus nigra by post-dispersal seed predation in different time-since-fire habitats

This study analyses the effects of post-dispersal predation of Pinus nigra seeds on the initial recruitment of this species in areas burned by large wildfires, where P. nigra shows very low regeneration. In three different habitats obtained in a gradient of time since fire in Catalonia (NE Spain), we have evaluated the effects of seed predators (ants, rodents and birds) on post-dispersal seed removal and early seedling establishment of P. nigra by using selective exclosures limiting their access to seeds. Ants were the most efficient seed predator group, followed by rodents and birds. The contribution of each group to overall predation showed large seasonal variations. The first seeds dispersed in winter were mainly predated by rodents, which also registered their highest abundance in this season of the year. In spring, at the end of the natural dissemination period of P. nigra seeds, ants became the major predators, this fact coinciding with their increased abundance. Birds showed the lowest predation values. In the seedling establishment experiment, only in the exclusion treatment of the three predator groups was there initial establishment in all habitats, especially in the recently burned area, where there was seedling establishment in all exclusion treatments. The post-dispersal seed predation by different animal groups and low seedling emergence in the different habitats obtained in this study, together with the low seed availability of P. nigra seeds in burned areas, do not predict a favourable outlook for the natural post-fire recolonization of this species, which might even affect its overall distribution area in the region.     

Factors affecting seed germination and seedling establishment of a long-lived desert shrub (<Emphasis Type="Italic">Coleogyne ramosissima</Emphasis>: Rosaceae)

Long-lived desert shrubs exhibit infrequent, episodic recruitment from seed. In spite of this long time scale, selection on life history attributes that affect seedling recruitment should be strong. We studied factors affecting germination phenology and seedling establishment for Coleogyne ramosissima, a dominant shrub species in the ecotone between warm and cold deserts in western North America. We also examined ecotypic differentiation in establishment strategy in response to selection regimes in two contrasting habitats. We followed patterns of dormancy loss, germination, emergence, and survival in reciprocal field experiments at warm winter Mojave Desert and cold winter Colorado Plateau study sites. Seed germination took place in late winter, under winter rain conditions at the warm desert site and under snow at the cold desert site. Distinctive germination phenologies for the two seed populations at contrasting field sites followed predictions based on laboratory germination experiments. There was no seed bank carryover across years. Seedling survival at the end of three growing seasons was remarkably high (mean survival 54%). Most seedling mortality was due to sprout predation by rodents early the first spring in unprotected caches. Emergence and establishment at each site were significantly higher for seeds from the local population, supporting the idea of ecotypic differentiation in establishment strategy. Establishment success was an order of magnitude greater overall at the Colorado Plateau site, which represents the leading edge of an upward elevational shift in distribution for this species under the current climatic regime. The Mojave Desert site is on the trailing edge of this shift, and recruitment there is apparently a much less frequent occurrence.     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Snow cover manipulations alter survival of early life stages of cold‐temperate tree species

Projections of future climate suggest increases in global temperatures that are especially pronounced in winter in cold‐temperate regions. Thermal insulation provided by snow cover to litter, soil, and overwintering plants will likely be affected by changing winter temperatures and might influence future species composition and ranges. We investigated effects of changing snow cover on seed germination and sapling survival of several cold‐temperate tree species using a snow manipulation approach. Post‐winter seed germination increased or decreased with increasing snow cover, depending on species; decreased seed germination was found in species that characteristically disperse seed in summer or fall months prior to snowfall. Post‐winter sapling survival increased with increasing snow cover for all species, though some species benefitted more from increased snow cover than others. Sapling mortality was associated with root exposure, suggesting the possibility that soil frost heaving could be an important mechanism for observed effects. Our results suggest that altered snow regimes may cause re‐assembly of current species habitat relationships and may drive changes in species’ biogeographic range. However, local snow regimes also vary with associated vegetation cover and topography, suggesting that species distribution patterns may be strongly influenced by spatial heterogeneity in snow regimes and complicating future projections.     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Modelling oldfield species richness in a mountain area

Seed predation by rodents affects plant population dynamics and it mayrespond to changes in vegetation structure at forest edges. This studyinvestigates the magnitude and direction of a potential edge effect intemperatedeciduous forests, and it seeks possible explanations based on predatorabundance and vegetation structure. The study was conducted at twelve forestedges in northern Switzerland; in six sites all shrubs at the forest edge wereremoved, whereas the other sites maintained a structurally rich shrub layer. Ineach site three transects were laid out from the edge towards the forestinterior. In six plots along these transects (at 1, 3, 6, 12, 25, 50m) we studied seed removal from experimental dishes, rodentabundance in live-traps and four characteristics of the vegetation structure.For both woody seed species (Prunus avium,Viburnum lantana) predation was significantly higher nearthe forest edge in the structurally rich sites; in the other sites no suchgradient was found. Selectively accessible dishes revealed that rodents werethemain predators, whereas predation by insects or molluscs was not observed.Abundance of rodents (Apodemus flavicollis, A.sylvaticus, Clethrionomys glareolus) washighestunder dense shrubs close to the forest edge. In the structurally rich sitesthere was a clear gradient of decreasing shrub cover from the edge towards theforest interior; a weaker gradient was observed in the structurally poor sites.We conclude that high shrub cover near the forest edge is the main determinantfor edge effects in seed predation, and edges without a shrub belt show no sucheffect.     

Assessing context-dependent survival of <Emphasis Type="Italic">Quercus glauca</Emphasis> seeds in a humid maritime woodland

Whilst consumption by rodents is often invoked as a major mortality factor for large-seed species of trees, its relative importance compared with other mortality factors is poorly known. We investigated experimentally the fate of post-dispersal seeds of Quercus glauca under different understorey environments (areas covered by (i) a pteridophyte Pyrrosia lingua, (ii) a ground-vine, Trachelospermum asiaticum and (iii) no vegetation) from the germination stage to seedling emergence and establishment stages in humid maritime woodland. We employed a pair of caged and uncaged treatments to evaluate the impact of seed removal/predation by rodents, which allowed us to separate seed removal/predation mortality from mortality due to other factors. Effects by rodents were greater in the no-understorey habitat than in the Pyrrosia and Trachelospermum habitats at early stages of development, whilst non-rodent-associated mortality was relatively more important towards the seedling establishment stages in all habitats. In the absence of predation/removal by rodents (i.e. the caged treatment), more seedlings survived in the no-understorey habitat whilst seedlings were significantly taller in the Pyrrosia habitat. In contrast, no significant difference was observed in either seed/seedling survivorship or seedling height amongst habitats where seeds/seedlings were exposed to rodent predation/removal. Overall, this study in a humid maritime woodland has revealed the temporally variable influence of mortality factors and the context-dependent survival of oak seeds/seedlings, making a contrast to observations in drier woodlands; in the no-understorey environment predation/removal effect was heavier but later survivorship was higher, whilst in vegetated environments, predation/removal was reduced but survivorship was not high.     

Vertebrate herbivores are the main cause of seedling mortality in a logged African rainforest—implications for forest restoration

High seedling mortality can limit tropical forest regeneration after disturbance, such as selective logging. However, causes of seedling mortality are often poorly understood although this information is critical for designing appropriate restoration methods. We studied seedling mortality in a rainforest in Uganda where natural recovery has failed after selective logging carried out 40 years earlier and the sites are dominated by herbaceous vegetation. We established an experiment where seeds of 12 indigenous tree species were sown into different‐sized logging gaps in order to capture factors that cause early seedling mortality. We examined whether seedling mortality was influenced by vegetation cover (VC), vertebrate herbivores (VHs), gap size, or forest compartment and recorded causes of mortality. We found that seedling mortality was in general high but showed high tree species‐specific variation (ranging from 19 to 92% six months after emergence). VHs, particularly rodents, were the most important cause of mortality. Rotting caused mortality particularly in smaller gaps, but elephant trampling, drying, being washed away by rain and falling debris were only minor mortality causes. Dense VC was not a considerable barrier to seedling survival, but by providing shelter for rodents, it indirectly increased seedling herbivory. Our results show that high seedling mortality on logged sites can limit forest regeneration and rodents in particular can be a significant factor causing high seedling mortality.     

Critical stages in the recruitment process of Rhamnus alaternus L

BACKGROUND AND AIMS: Rhamnus alaternus is a Mediterranean shrub commonly used in reforestation programs. Although several aspects of its reproductive biology have been studied, little is known about the importance of the different recruitment stages in the overall regeneration process of this species, which limits its proper use in Mediterranean forests and shrubland management. The aim of the present work was to quantify the importance of the different recruitment stages in the regeneration process of R. alaternus. METHODS: Two populations of Rhamnus alaternus on the island of Mallorca that differ in climatic conditions, type of habitat and sex ratio were studied. The importance of seed production, seed dispersal and predation, seedling emergence and seedling survival for the regeneration of this species were quantified. KEY RESULTS: In both populations, fruit set and fruit removal by animals were not critical stages, since almost half of the flowers became mature fruits and 90% of those were dispersed. Most seeds were deposited under female conspecifics (86 and 47%, at Lloret and Esporles, respectively), and very few were found in open inter-spaces (1 and 5%). Post-dispersal seed predation (mostly by ants and rodents) was very high in both populations. Seedling emergence took place during autumn and early winter and it ranged from 31 to 68% depending upon year and microhabitat. The majority of emerged seedlings died during the first year, mainly due to desiccation; such mortality was influenced by rainfall and differed among microhabitats (varying from 67 to 100%). The general spatial distribution of seed rain was concordant with the seedling emergence and survival pattern in both populations. CONCLUSIONS: The recruitment of Rhamnus alaternus appeared to be mainly limited by seed and seedling survival, regardless of the type of habitat in which the species is found.     

Poachers and Forest Fragmentation Alter Seed Dispersal,Seed Survival,and Seedling Recruitment in the Palm Attalea butyraceae,with Implications for Tropical Tree Diversity1

We examined the interaction between a palm and two bruchid beetles along with several mammal species to explore how poachers and habitat fragmentation may indirectly alter the spatial pattern of seed dispersal, seed predation, and seedling recruitment in central Panama. The large, stony endocarps of Attalea butyraceae decay slowly and bear distinctive scars when opened by rodents or beetles. We determined the final distance between endocarps and reproductive trees (which we call an ecologically effective dispersal distance), the predation status of each endocarp, and the distance between seedlings and reproductive trees. The 68 focal trees were divided among 14 sites and four levels of anthropogenic disturbance. Levels of disturbance included full protection from poachers, light and heavy pressure from poachers, and small island habitat fragments. Ecologically effective seed dispersal distances were greatest for protected sites, intermediate for lightly poached sites, and shortest for heavily poached sites and habitat fragments. Seed predation by rodents increased with distance to the nearest reproductive Attalea and was greatest for fully protected sites, intermediate for lightly poached sites, and least for heavily poached sites and habitat fragments. Seed predation by beetles reversed the patterns described for seed predation by rodents. Total seed predation by beetles and rodents combined was independent of distance, greatest for fully protected sites, and lower for poached sites and habitat fragments. Seedling densities were always greatest close to reproductive trees; however, the increase in seedling densities close to reproductive trees was minimal for fully protected sites, clearly evident for poached sites, and pronounced for habitat fragments. Increased seedling recruitment near conspecific trees may in time reduce tree diversity where humans disrupt mammal communities.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.     

Effects of the duration of cold stratification on early life stages of the Mediterranean alpine plant Silene ciliata

Cold stratification provided by snow cover is essential to break seed dormancy in many alpine plant species. The forecast reduction in snow precipitation and snow cover duration in most temperate mountains as a result of global warming could threaten alpine plant populations, especially those at the edge of their species distribution, by altering the dynamics of early life stages. We simulated some effects of a reduction in the snow cover period by manipulating the duration of cold stratification in seeds of Silene ciliata, a Mediterranean alpine specialist. Seeds from three populations distributed along an altitudinal gradient were exposed to different periods of cold stratification (2, 4 and 6 months) in the laboratory and then moved to common garden conditions in a greenhouse. The duration of the cold stratification treatment and population origin significantly affected seed emergence percentage, emergence rate and seedling size, but not the number of seedling leaves. The 6‐month and 4‐month cold stratification treatments produced higher emergence percentages and faster emergence rates than seeds without cold stratification treatment. No significant cold stratification duration x seed population origin interactions were found, thus differential sensitivity to cold stratification along elevation is not supported.     

Effects of fire severity in a north Patagonian subalpine forest

Abstract. Question: What is the relative importance of fire‐induced canopy mortality, soil burning and post‐fire herbivory on tree seedling performance? Location: Subalpine Nothofagus pumilio forests at Challhuaco valley (41°13'S, 71°19'W), Nahuel Huapi National Park, Argentina. Methods: We fenced and transplanted soils of three burning severities along a fire severity gradient produced by a fire in 1996. Over two growing seasons we monitored soil water, direct incoming solar radiation, seedling survival, final seedling total biomass and root/shoot ratio. Additionally, we assessed severity‐related changes in soil properties. Results: Incoming radiation (an indicator of the amount of canopy cover left by the fire) was the primary factor influencing spring and summer top soil water availability, first and second‐year seedling survival and seedling growth. While seedling survival and soil water content were negatively affected by increased radiation, seedling final biomass was highest in very open microsites. Burned soils showed lower water holding capacity and soil carbon; however these changes did not affect topsoil water, and, contrary to expectation, there was a slight tendency toward higher seedling survival on more heavily burned soils. Herbivory significantly reduced seedling survival, but only under high‐radiation conditions. While the effect of radiation on final seedling biomass was not affected by herbivory, R/S ratios were significantly reduced by herbivory in high radiation micro sites. Conclusions: Despite inducing faster aerial growth, increased radiation and herbivory in severely burned sites may effectively prevent post‐fire regeneration in north Patagonian subalpine forest where seed sources are not limiting.     

Ant foraging strategies vary along a natural resource gradient

Food selection by foragers is sensitive to the availability of resources, which may vary along geographical gradients. Hence, selectivity of food types by foragers is expected to track these resource gradients. Here we addressed this hypothesis and asked if foraging decisions of seed-eating ants differ along a geographic gradient of habitat productivity. The study was carried out for two years at five sites along a natural climatic gradient, ranging from arid to Mediterranean, where plant productivity varies six-fold across a short geographic distance of 250 km. We found that in ant colonies of the genus Messor, collective foraging decisions differed along the gradient. Specifically, at the high-productivity sites, a stronger association was found between plant seed availability and selectivity, suggesting that colonies respond more accurately to within-patch variation in food amounts. In contrast, colonies in low-productivity sites foraged in patches with higher concentration of seeds, suggesting that they respond more accurately to among-patch variation in food amounts. Moreover, at the high-productivity sites, colonies were more discriminating in their choice of food and preferred bigger seeds, while in the low-productivity sites, where smaller seeds were relatively more abundant, food collection depended mostly on seed availability. An experiment with artificial seed patches performed along the same climatic gradient, revealed no difference in food selectivity across sites when food type and availability were similar, and a general preference for bigger over medium-sized seeds. Overall, our findings suggest that resource availability is an important factor explaining food choice along a climatic gradient and imply that in low-productivity regions small-seeded species incur high predation pressure, whereas in high-productivity regions, large-seeded species suffer higher predation. This could have important consequences for plant species composition, particularly at the face of climate change, which could dramatically alter the foraging decisions of granivores.     

An interactive effect of simultaneous death of dwarf bamboo, canopy gap, and predatory rodents on beech regeneration

To clarify the interactive effect of the simultaneous death of dwarf bamboo (Sasa kurilensis), forest canopy gap formation, and seed predators on beech (Fagus crenata) regeneration, we analyzed beech demography from seed fall until the end of the first growing season of seedlings in an old-growth forest near Lake Towada, northern Japan. The simultaneous death of S. kurilensis took place in 1995. We established four types of sampling site differing in forest canopy conditions (closed or gap) and Sasa status (dead or alive). Beech seed survival and emergence ratio were both highest in gaps with dead Sasa (gap-dead), because rate of predation was lowest. Seedling survival during the first growing season was also highest in the gap-dead treatment, because of less predation and less damping off. As a result, even though density of seed fall was lowest in the gap-dead treatment, the living seedling density there was highest at the end of the first growing season. Predation, which caused the greatest mortality during the seed and seedling stages, was significantly lower at both sites in gaps and sites with dead Sasa. This was probably due to changes in the behavior of rodents in response to the structure of the forest canopy and undergrowth. Both the death of Sasa and canopy gap formation allowed seedlings to avoid damping off because of the high light availability. The indirect effect of the simultaneous death of Sasa and canopy gap formation in reducing predation contributed more to beech regeneration than their direct effect in increasing light for the seedlings.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

Snow evens fragmentation effects and food determines overwintering success in ground-dwelling voles

Climate instability strongly affects overwintering conditions in organisms living in a strongly seasonal environment and consequently their survival and population dynamics. Food, predation and density effects are also strong during winter, but the effect of fragmentation of ground vegetation on ground-dwelling small mammals is unknown. Here, we report the results of a winter experiment on the effects of habitat fragmentation and food on experimental overwintering populations of bank voles Myodes glareolus. The study was conducted in large outdoor enclosures containing one large, two medium-sized or four small habitat patches or the total enclosure area covered with protective tall-grass habitat. During the stable snow cover of midwinter, only food affected the overwintering success, body condition, trappability and earlier onset of breeding in bank voles. However, after the snow thaw in spring, habitat fragmentation gained importance again, and breeding activities increased the movements of voles in the most fragmented habitat. The use of an open, risky matrix area increased along the habitat fragmentation. Our experiment showed that long-lasting stable snow cover protects overwintering individuals in otherwise exposed and risky ground habitats. We conclude that a stable winter climate and snow cover should even out habitat fragmentation effects on small mammals. However, along prolonged snow-free early winter and in an earlier spring thaw, this means loss of protection by snow cover both in terms of thermoregulation and predation. Thus, habitat cover is important for the survival of small ground-dwelling boreal mammals also during the non-breeding season.     

The Interplay among Acorn Abundance and Rodent Behavior Drives the Spatial Pattern of Seedling Recruitment in Mature Mediterranean Oak Forests

The patterns of seedling recruitment in animal-dispersed plants result from the interactions among environmental and behavioral variables. However, we know little on the contribution and combined effect of both kinds of variables. We designed a field study to assess the interplay between environment (vegetation structure, seed abundance, rodent abundance) and behavior (seed dispersal and predation by rodents, and rooting by wild boars), and their contribution to the spatial patterns of seedling recruitment in a Mediterranean mixed-oak forest. In a spatially explicit design, we monitored intensively all environmental and behavioral variables in fixed points at a small spatial scale from autumn to spring, as well as seedling emergence and survival. Our results revealed that the spatial patterns of seedling emergence were strongly related to acorn availability on the ground, but not by a facilitationeffect of vegetation cover. Rodents changed seed shadows generated by mother trees by dispersing most seeds from shrubby to open areas, but the spatial patterns of acorn dispersal/predation had no direct effect on recruitment. By contrast, rodents had a strong impact on recruitment as pilferers of cached seeds. Rooting by wild boars also reduced recruitment by reducing seed abundance, but also by changing rodent’s behavior towards higher consumption of acorns in situ. Hence, seed abundance and the foraging behavior of scatter-hoarding rodents and wild boars are driving the spatial patterns of seedling recruitment in this mature oak forest, rather than vegetation features. The contribution of vegetation to seedling recruitment (e.g. facilitation by shrubs) may be context dependent, having a little role in closed forests, or being overridden by directed seed dispersal from shrubby to open areas. We warn about the need of using broad approaches that consider the combined action of environment and behavior to improve our knowledge on the dynamics of natural regeneration in forests.