Interspecific variation in extraction of buried seeds within an assemblage of sparrows

Most North American sparrows forage almost exclusively on herbaceous seeds during the winter months. Limited availability of surface seeds forces some birds to scratch for seeds buried beneath soil, snow, or litter. Artificial seed trays were used to test the ability of five different sparrow species to extract seeds buried at different depths in soil. The results suggest three functional groups based on relative scratching ability. Strong scratchers, which included eastern towhee and song and white-throated sparrows, met or exceeded their energetic requirements when foraging on seeds buried at all depths (down to a maximum depth of 1.50–2.25 cm). A weak scratching species, Savannah sparrow, scratched with the same frequency as the strong scratchers, but experienced negative energy budgets when forced to forage on sub-surface seeds. Finally, a non-scratching species, field sparrow, failed to extract any buried seeds. Level of scratching ability may influence foraging efficiency at low resource densities. As a result, interspecific differences in scratching ability may contribute to habitat selection. Strong scratchers may be adapted to foraging near woody vegetation where intense resource competition and abundant litter limit the availability of surface seeds. Weak scratchers, on the other hand, may be forced to feed away from areas where surface seeds are limited. Since woody vegetation serves as a primary source of protective cover in early successional habitats, a tradeoff between foraging efficiency and the risk of predation may promote the local coexistence of species that differ in relative scratching ability and adaptations to evading predators.     

Experimental evidence of density dependent activity pattern of a large herbivore in an alpine ecosystem

Density dependent processes affecting foraging strategies may in turn influence vital rates and population regulation in large herbivores. Increased competition may lower both forage availability and quality, but whether the main activity constraint at high density is increased searching time or increased digestion time is poorly investigated. In a fully replicated landscape‐scale experiment, we used long‐term data (2003–2009) from domestic sheep grazing at high and low density (80 and 25 sheep km^–2, respectively) on alpine summer ranges to test density dependence in allocation of time to feeding (moving) vs digestion (resting) activities and how this in turn affected body growth. Sheep at high density spent more time actively feeding than sheep at low density, but sheep moved shorter distances while foraging at high density. Increased activity levels at high density suggest that the main activity constraint at high density was availability of high‐quality food increasing searching time and possibly reducing intake rates. Increased movement distances at low density is consistent with a higher selection for more productive vegetation types since high‐quality patches are dispersed in the landscape. The alternative hypothesis, that food processing time increased at high density was not supported as it would have reduced overall activity levels. Individual activity levels increased body growth, but this was not sufficient to fully compensate for lower habitat quality leading to an overall reduced body growth at high density. Our experiment clearly documents changes in activity budgets and movement distances of a large herbivore at high population density, providing one potential behavioural mechanism of density dependent responses observed in vital rates.     

Costs of reproduction in a population of European adders

Eleven years of data on a small population of adders (Vipera berus) in southern Sweden provide quantitative information on the nature and degree of costs faced by reproducing animals. Reproduction imposes both an energy cost (measured by loss in body mass) and a mortality cost on adders of both sexes. The extent of the energy cost is broadly independent of levels of reproductive activity in males, but mortality costs are highest for large males, perhaps because they are more obvious to predators. In females, energy costs include a high ‘fixed’ (fecundity-independent) component, such that a large litter may cost little more to produce than would a small litter. Energy costs and mortality costs are separate in males, but inter-related in females. Mortality of reproducing females is high (40% per year), primarily because post-parturient females are emaciated and must forage actively, hence increasing their vulnerability to predators. Females producing relatively large litters (high Relative Clutch Mass) lose more body mass, and are less likely to survive after reproducing. The observed low reproductive frequencies of female adders may result from the presence of high fecundity-independent costs of reproduction.     

Context-dependent behavior and the benefits of communal nesting

We present a model for the behavior of communally nesting insects. Females may forage for food to provision offspring or may remain in the nest, with the option of eating and replacing nest mates' eggs. Orphaned brood are at risk of predation. The optimal behavior of solitary females is determined using stochastic dynamic programming; static and dynamic evolutionarily stable strategies (ESSs) are then calculated for colonies of various sizes. A solitary female should forage if her brood is smaller than a time-dependent threshold. Females in small colonies should forage. In colonies above some threshold size, the static ESS is for one female to forage and the rest to cheat. The dynamic ESS in large colonies is for no females to forage until some time close to the end of the foraging season and for all females to forage thereafter. Mixed dynamic ESSs, with some foragers and some cheats, may arise if individuals differ in their chances of surviving a foraging interval or if females with new offspring vary their guarding behavior, depending on the numbers of cheats and new cells in the nest. We discuss these predictions in the light of published observations and preliminary data on the halictine bee Lasioglossum (Chilalictus) hemichalceum.     

Reproductive phenology of a food-hoarding mast-seed consumer: resource- and density-dependent benefits of early breeding in red squirrels

The production of offspring by vertebrates is often timed to coincide with the annual peak in resource availability. However, capital breeders can extend the energetic benefits of a resource pulse by storing food or fat, thus relaxing the need for synchrony between energy supply and demand. Food-hoarding red squirrels (Tamiasciurus hudsonicus) breeding in the boreal forest are reliant on cones from a masting conifer for their nutrition, yet lactation is typically completed before the annual crop of cones is available for consumption such that peaks in energy supply and demand are not synchronized. We investigated the phenological response of red squirrels to annual variation in environmental conditions over a 20-year span and examined how intra- and inter-annual variation in the timing of reproduction affected offspring recruitment. Reproductive phenology was strongly affected by past resource availability with offspring born earlier in years following large cone crops, presumably because this affected the amount of capital available for reproduction. Early breeders had higher offspring survival and were more likely to renest following early litter loss when population density was high, perhaps because late-born offspring are less competitive in obtaining a territory when vacancies are limited. Early breeders were also more likely to renest after successfully weaning their first litter, but renesting predominantly occurred during mast years. Because of their increased propensity to renest and the higher survival rates of their offspring, early breeders contribute more recruits to the population but the advantage of early breeding depends on population density and resource availability.     

Changes in metabolism in response to fasting and food restriction in the Steller sea lion (Eumetopias jubatus)

Many animals lower their resting metabolism (metabolic depression) when fasting or consuming inadequate food. We sought to document this response by subjecting five Steller sea lions to periods of: (1) complete fasting; or (2) restricting them to 50% of their normal herring diet. The sea lions lost an average of 1.5% of their initial body mass per day (2.30 kg/d) during the 9-14-day fast, and their resting metabolic rates decreased 31%, which is typical of a "fasting response". However, metabolic depression did not occur during the 28-day food restriction trials, despite the loss of 0.30% of body mass per day (0.42 kg/d). This difference in response suggests that undernutrition caused by reduced food intake may stimulate a "hunger response", which in turn might lead to increased foraging effort. The progressive changes in metabolism we observed during the fasts were related to, but were not directly caused by, changes in body mass from control levels. Combining these results with data collected from experiments when Steller sea lions were losing mass on low energy squid and pollock diets reveals a strong relationship between relative changes in body mass and relative changes in resting metabolism across experimental conditions. While metabolic depression caused by fasting or consuming large amounts of low energy food reduced the direct costs from resting metabolism, it was insufficient to completely overcome the incurred energy deficit.     

Chemical composition and tissue energy density of the cuttlefish (Sepia apama) and its assimilation efficiency by Diomedea albatrosses

The cuttlefish Sepia apama Gray (Mollusca: Cephalopoda) is a seasonally abundant food resource exploited annually by moulting albatrosses throughout winter and early spring in the coastal waters of New South Wales, Australia. To assess its nutritional value as albatross forage, we analysed S. apama for water, lipid protein, ash contents, energy density and amino acid composition. Because albatrosses consistently consume S. apama parts preferentially in the order of head, viscera and mantle, we analysed these sections separately, but did not identify any nutritional basis for this selective feeding behaviour. The gross energy value of S. apama bodies was 20.9 kJ/g dry mass, but their high water content (>83%; cf <70% for fish) results in a relatively low energy density of 3.53 kJ/g. This may contribute to a need to take large meals, which subsequently degrade flight performance. Protein content was typically >75% dry mass, whereas fat content was only about 1%. Albatrosses feed on many species of cephalopods and teleost fish, and we found the amino acid composition of S. apama to be comparable to a range of species within these taxa. We used S. apama exclusively in feeding trials to estimate the energy assimilation efficiency for Diomedea albatrosses. We estimated their nitrogen-corrected apparent energy assimilation efficiency for consuming this prey to be 81.82 ± 0.72% and nitrogen retention as 2.90 ± 0.11 g N kg⁻¹ d⁻¹. Although S. apama has a high water content and relatively low energy density, its protein composition is otherwise comparable to other albatross prey species. Consequently, the large size and seasonal abundance of this prey should ensure that albatrosses remain replete and adequately nourished on this forage while undergoing moult.     

Does foraging behaviour explain the poor breeding success of great tits Parus major in northern Europe?

Great tits Parus major have generally much poorer breeding success in northern Finland than in mid- and western Europe. The aim of this study is to find out whether the poor success is linked to foraging behaviour. This was studied by monitoring great tits' foraging behaviour and food abundance in different substrates during the breeding cycle in two populations, N Finland (65°N) and Latvia (56°N). It was shown that breeding success, from eggs to fledglings, was significantly poorer (56% vs. 93%) and caterpillar abundance considerably lower in the northern population. The general patterns in foraging behaviour were the same in N Finland and Latvia (especially the preference of birches Betula spp.) indicating that great tits used basically the same species-typical foraging strategy in both populations. However, thin branches of birch were preferred in the north but avoided in Latvia, which may suggest that northern great tits have changed their foraging niche towards the outer parts of canopy, a niche typically occupied by the blue tit. This shift is theoretically advantageous, since the outer parts of the canopy are the richest caterpillar source. In practice, however, the poorer success of great tits indicates that this is not a beneficial strategy. Primarily, great tits in the northern population seem to be food limited since they lay too large clutches in relation to accessible food resources. This may be because great tits are not adapted to lower caterpillar production in the northern margins of their distribution and cannot change their narrow diet (3/4 caterpillars), like e.g. willow tits can. In search for caterpillars, food accessibility is further limited because the great tits' normal foraging behaviour, with wide search radius, may not function properly in the denser, outer parts of the canopy. Great tits may also be too heavy to forage efficiently on leafed twigs.     

The ecology of asociality in Namibian leopards

Data on the ecology of leopards ( Panthera pardus ) from north-eastern Namibia are presented and discussed in terms of the possible costs and benefits of solitary behaviour. In an area of low leopard density, where individuals lived alone, both males and females occupied large home ranges,(♂= 210-1164km², ♀= 183-194km²). Despite resource and reproductive advantages in maintaining exclusive ranges, the degree of range overlap both between and within sexes was substantial. Average overlap between males was 46% and between females 35%. The cost of dispersal appeared high as all three marked sub-adults died, and most recorded mortalities were of sub-adults. Females with dependent young showed a significant increase in per capita food intake compared to single females and males. Higher foraging success by females with cubs was revealed through two energy expenditure parameters (kg/km travelled/day and kg/hunt/day). Differential food intake between females with cubs, single females and males can be explained partly by differences in day ranges, body size and costs of parental care. Females shared 27% of their food with cubs and the costs of sharing food does not appear as high as previously suggested. Inter-specific competition over food and the defence of carcasses is suggested as an important cost to group living. Leopard kills were visited by other large carnivores (12%) but food loss was minimal (2%). Leopards successfully avoided conflict with inter-specific competitors by dragging and hiding kills in thick vegetation. We argue that solitary and secretive behaviour enables leopards to avoid the costs of defending carcasses against larger and gregarious carnivores.     

Positive indirect effect of tadpoles on a detritivore through nutrient regeneration

In aquatic food webs consumers can affect other members of the web by releasing nutrients as a result of their feeding activity. There is increasing evidence of these positive effects on primary producers, but such nutrient regeneration can also affect detritivores, by favoring the activities of detritus-associated microbes. We examined the effects of nutrient regeneration by tadpoles on leaf-eating detritivores under laboratory conditions. We fed four species of tadpoles three different food items (leaf litter, algae, and sludgeworms). We then conditioned terrestrial dead leaves with water from reared tadpoles (treatments) or food items alone (controls), and compared the C:N ratios of the conditioned leaves and the growth of the isopod Asellus hilgendorfii fed on the conditioned leaves. Tadpole feeding activity reduced the C:N ratio of conditioned leaves, and the effect was greatest when tadpoles were fed algae. Isopod growth rates were often higher when they were fed the litter conditioned with water from reared tadpoles. Thus, nutrient regeneration by tadpoles had a positive indirect effect on detritivores by enhancing leaf quality. Tadpoles often occur in nutrient-limited habitats where leaf litter is the major energy source, and their facilitative effects on leaf-eating detritivores may be of great significance in food webs by enhancing litter decomposition.     

Large litter size increases maternal energy intake but has no effect on UCP1 content and serum-leptin concentrations in lactating Brandt’s voles (<Emphasis Type="Italic">Lasiopodomys brandtii</Emphasis>)

Lactation is the most energetically demanding period in the female mammal's life. We measured maternal energy intake, uncoupling protein 1 (UCP1) content in brown adipose tissue (BAT), serum-leptin concentration, and litter growth in lactating Brandt's voles (Lasiopodomys brandtii) with different litter sizes. Litter mass was positively related to litter size but there was no difference in pup mass at birth. Maternal gross energy intake at peak lactation was positively correlated with litter size and litter mass. Maternal resting metabolic rate (RMR) was positively correlated with litter mass, but not with litter size. No significant differences were detected in body-fat mass, serum-leptin concentration, or UCP1 in lactating voles with different litter sizes. Serum-leptin concentration was negatively correlated with energy intake during lactation. Our data did not support the hypothesis that there is a trade-off in energy allocation between maternal maintenance and offspring growth in lactating Brandt's voles, but support the idea that if the mothers with ten pups should have less energy available for their maintenance than mothers raising fewer pups. Also, leptin is probably not the only factor that induces the high energy intake in mothers with large litter sizes, although it was involved in the regulation of energy intake during lactation.     

Perception of Human-Derived Risk Influences Choice at Top of the Food Chain

On human-used landscapes, animal behavior is a trade-off between maximizing fitness and minimizing human-derived risk. Understanding risk perception in wildlife can allow mitigation of anthropogenic risk, with benefits to long-term animal fitness. Areas where animals choose to rest should minimize risk from predators, which for large carnivores typically equate to humans. We hypothesize that high human activity leads to selection for habitat security, whereas low activity enables trading security for forage. We investigated selection of resting (bedding) sites by GPS radiocollared adult grizzly bears (n = 10) in a low density population on a multiple-use landscape in Canada. We compared security and foods at resting and random locations while accounting for land use, season, and time of day. On reclaimed mines with low human access, bears selected high horizontal cover far from trails, but did not avoid open (herbaceous) areas, resting primarily at night. In protected areas bears also bedded at night, in areas with berry shrubs and Hedysarum spp., with horizontal cover selected in the summer, during high human access. On public lands with substantial human recreation, bears bedded at day, selected resting sites with high horizontal cover in the summer and habitat edges, with bedding associated with herbaceous foods. These spatial and temporal patterns of selection suggest that bears perceive human-related risk differentially in relation to human activity level, season and time of day, and employ a security-food trade-off strategy. Although grizzly bears are presently not hunted in Alberta, their perceived risks associated with humans influence resting-site selection.     

Factors influencing seasonal and monthly changes in the group size of chital or axis deer in southern India

Chital or axis deer (Axis axis) form fluid groups that change in size temporally and in relation to habitat. Predictions of hypotheses relating animal density, rainfall, habitat structure, and breeding seasonality, to changes in chital group size were assessed simultaneously using multiple regression models of monthly data collected over a 2 yr period in Guindy National Park, in southern India. Over 2,700 detections of chital groups were made during four seasons in three habitats (forest, scrubland and grassland). In scrubland and grassland, chital group size was positively related to animal density, which increased with rainfall. This suggests that in these habitats, chital density increases in relation to food availability, and group sizes increase due to higher encounter rate and fusion of groups. The density of chital in forest was inversely related to rainfall, but positively to the number of fruiting tree species and availability of fallen litter, their forage in this habitat. There was little change in mean group size in the forest, although chital density more than doubled during the dry season and summer. Dispersion of food items or the closed nature of the forest may preclude formation of larger groups. At low densities, group sizes in all three habitats were similar. Group sizes increased with chital density in scrubland and grassland, but more rapidly in the latter—leading to a positive relationship between openness and mean group size at higher densities. It is not clear, however, that this relationship is solely because of the influence of habitat structure. The rutting index (monthly percentage of adult males in hard antler) was positively related to mean group size in forest and scrubland, probably reflecting the increase in group size due to solitary males joining with females during the rut. The fission-fusion system of group formation in chital is thus interactively influenced by several factors. Aspects that need further study, such as interannual variability, are highlighted.     

Trade-offs in Energy Allocation During Lactation

SYNOPSIS. During lactation, mothers require energy to meet bothmaternal and offspring requirements. If a mother exports toomuch energy to dependent offspring (in milk), her weight lossmay be excessive and maternal risk may increase. Conversely,too little energy allocation to offspring may reduce the growthrate or induce mortality of dependent offspring. This paradigmwas evaluated in cotton rats (Sigmodon hispidus) supportingsmall (3 pup) and large (6 pup) litters from early to late lactation.Several types of evidence indicate that physiological constraintslimit the ability of mothers with large litters to provide resourcesto offspring. Mothers with large litters produced a dilute,energy-poor milk and their rates of food intake, weight lossand energy export per litter appeared to approach physiologicalmaxima. Whereas the energy exported to pups in small littersincreased from early to late lactation, the energy flow perpup in large litters was consistently low; consequently, offspringin large litters had low growth rates. An increase in eithermaternal food intake or weight loss (catabolism of maternaltissue) could have provided additional energy to offset thelow growth rate of pups in large litters. However, mothers withlarge litters did not substantially increase their food intakeor weight loss compared with mothers supporting small litters.These results indicate that the maternal support of offspringin large litters is limited. The pattern of energy allocationshown by cotton rats with large litters likely reflects a compromisebetween meeting maternal and offspring energy requirements (cf.,Parker and Macnair, 1979). The energy flow is greater than optimalfor the parent but less than optimal for the offspring. Lessmaternal-offspring conflict occurs in small than large littersbecause offspring in small litters maintain a high growth rateat a relatively low maternal cost. Yet, under favorable environmentalconditions, the reduction in maternal-offspring conflict hasno apparent fitness benefit.     

Aggregation in a Desert Tenebrionid Beetle: A Cost/Benefit Analysis

Parastizopus armaticeps (Coleoptera: Tenebrionidae), a nocturnal fossorial detritivore inhabiting southern Kalahari dunes, aggregates in burrows during the day. Group size increases during drought but 25% of beetles are still found alone or in pairs. During drought, beetles from large groups leave burrows after sunset synchronously and carlier than pairs and single animals and earlier than beetles of any group size after rain. Detritus from the beetles' major foodplant is scarce and food competition high. Beetles emerging early preferentially select and carry high-quality transportable items into burrows to eat (forage); late-emerging ones feed on the low-quality large twigs on the surface. Foraging is shown to be a strategy to secure food items against surface competitors, not one to reduce body water loss during surface exposure. The costs and benefits of group vs. solitary lifestyles and alternate hypotheses for early and synchronous emergence were tested experimentally. Grouped beetles had lower body water loss rates but, due to competition with burrow mates, higher feeding costs than single ones. It is hunger that advances and thus synchronizes emergence time, not social facilitation. Field data support a model predicting that, for maximal benefits, beetles should alternate between solitary and group life at optimal time intervals.     

Density-dependent foraging behaviour of sheep on alpine pastures: effects of scale

Foraging patterns of large herbivores may give important clues as to why their life history varies depending on population density. In this landscape-scale experiment, domestic sheep Ovis aries were kept at high (80 sheep km⁻²) and low (25 sheep km⁻²) population densities during summer in high mountain pastures in Hol, Norway. We predicted an increasing use of less preferred plant species or habitat types with increasing sheep population density. Foraging behaviour was investigated by direct observations of individually marked sheep on different spatial scales, and diet composition was also assessed with microhistological analysis of faecal samples from known individuals. We found that the effects of density on foraging behaviour depended on scale and were only detected at the scale of diet choice. Use of the common grass species Deschampsia flexuosa , which provided the bulk forage (10–65% of the diet), remained constant throughout the season at low densities, but increased significantly over time at high densities. On a coarser spatial scale, neither within vegetation type nor between vegetation types, selection was affected by density, but vegetation type selection differed depending on whether the sheep were grazing or resting. Our study provides evidence of density dependence in foraging behaviour, but only at the finest spatial scale (diet choice).     

Habitat selection and diet of Great Snipe Gallinago media during breeding

The Great Snipe Gallinago media is considered to be an endangered species. This paper examines its food and habitat use on a sub-alpine/low-alpine breeding area in central Norway. It was estimated that earthworms constitute more than 90% of Great Snipe food (by weight). Feeding birds selected the low herb willow scrub vegetation community and to a lesser degree eutrophic fen. Birds did not selectively feed on eutrophic dwarf birch/juniper heath despite it being the third most used vegetation community by virtue of its extent. Although soil penetrability, vegetation cover and earthworm density varied across vegetation communities, Great Snipe selected sites with similar habitat characteristics in different vegetation communities. Great Snipe seemed to select for an optimal combination of soil penetrability and earthworm density, and for medium scrub cover. Nests were found in a broad range of vegetation communities, but only low herb willow scrub was selectively used. However, an equal number of nests was found in eutrophic fen due to its larger extent. Dense vegetation cover around nests and short flushing distance of incubating females indicate low sensitivity to disturbance. We consider the Great Snipe to be a food and habitat specialist, requiring habitats rich in sub-surface invertebrates to breed. This may explain its scattered distribution in Scandinavia, and may render it vulnerable to habitat modification and loss.     

The social organization of non-breeding Magpies Pica pica

The social behaviour of non-breeding individuals in a colour-marked population of Magpies was studied. In early autumn most non-breeders began to forage in a common area, the 'Non-breeding Flock Area'. A few individuals remained on their natal territories away from the flock area as solitary non-breeders. Most non-breeders were first-year birds, but some were second years or adults which no longer held a territory. Birds foraged in groups, their food intake rate differing with both group size and location. It appeared that the area in which birds foraged had a significant effect on food intake rate; foraging groups tended to form at sites rich in food. There was a dominance hierarchy amongst non-breeders; an individual's foraging behaviour, survivorship and chances of breeding were status dependent, with subordinates feeding less in groups and being less likely to survive and breed. Solitary individuals' chances of breeding were similar to those of high status birds, although their survivorship to breeding age may have been lower. Non-breeding Magpies are compared with non-breeders of other species and the factors which may influence their social behaviour are discussed. It is suggested that remaining as a solitary non-breeder is a viable alternative to becoming a low-status flock member for some birds.     

Foraging Rules for Group Feeders: Area Copying Depends upon Food Density in Shoaling Goldfish

The decision rules governing forage area copying behaviour were investigated in shoaling fish. Shoaling goldfish were offered two equal food patches, one of which was adjacent to an equal-sized shoal feeding behind a transparent barrier. When food was low, goldfish foraged according to an area copying rule, but under high and zero food area copying disappeared. Only under high food density did equal numbers of fish feed at both sites as predicted by foraging theory. Under zero food the fish were less certain about where to forage. Precise visual cues from feeding fish were required: non-feeders did not attract area copiers. Furthermore, area copying was task-dependent since it reappeared strongly if fish were not able to forage on patches like their fellows. Control experiments eliminated an increase in group size for anti-predator advantage as an explanation. Two sequential decisions: to stay or move, and to join or leave may explain the results, which are not accommodated by simple optimality models. These decisions may be based on a comparison of current food intake with the anticipation of a higher reward by foraging socially.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.