Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Direct and indirect effects of understorey bamboo shape tree regeneration niches in a mixed temperate forest

Plant cover plays a major role in shaping the nature of recruitment microsites through direct (resource mediated) and indirect (consumer mediated) interactions. Understorey plants may differentially affect seedling establishment, thus contributing to regeneration-niche separation among canopy tree species. We examined patterns of early tree seedling survival resulting from interactive effects of understorey bamboo (Chusquea culeou) and resident consumers in a mixed temperate Patagonian forest, Argentina. Newly germinated seedlings of Nothofagus dombeyi and Austrocedrus chilensis were planted in bamboo thickets and non-bamboo patches, with or without small-vertebrate exclosures. We found species-specific patterns of seedling survival in relation to bamboo cover. Nothofagus survival was generally low but increased under bamboo, irrespective of cage treatment. Desiccation stress accounted for most Nothofagus mortality in open, non-bamboo areas. In contrast, Austrocedrus survival was highest in non-bamboo microsites, as most seedlings beneath bamboo were killed by small vertebrates through direct consumption or non-trophic physical damage. There was little evidence for a negative impact of bamboo on tree seedling survival attributable to resource competition. The balance of simultaneous positive and negative interactions implied that bamboo presence facilitated Nothofagus early establishment but inhibited Austrocedrus recruitment via apparent competition. These results illustrate the potential for dominant understorey plants to promote microsite segregation during early stages of recruitment between tree seedlings having different susceptibilities to water stress and herbivory. We recognise, however, that patterns of bamboo–seedling interactions may be conditional on moisture levels and consumer activity during establishment. Hence, both biotic and abiotic heterogeneity in understorey environments should be incorporated into conceptual models of regeneration dynamics and tree coexistence in forest communities.     

Woody debris and terrestrial invertebrates – effects on prey resources for brown trout (Salmo trutta) in a boreal stream

Intensive forestry and other activities that alter riparian vegetation may disrupt the connectivity and the flux of energy between terrestrial and aquatic habitats and have large effects on biota, especially in small streams. We manipulated the amount of in-stream wood and the flux of terrestrial invertebrate subsidies to determine how these factors affected potential food resources for drift-feeding brown trout (Salmo trutta ) in a boreal Swedish forest stream. Specifically, we followed the effects on the abundance of aquatic and terrestrial invertebrate fauna from June to August 2007. The treatments were 1) addition of wood, unmanipulated terrestrial invertebrate inputs, 2) reduction of terrestrial invertebrate inputs (using canopy covers), no addition of wood, 3) unmanipulated ambient conditions, 4) simultaneous addition of wood and reduction of terrestrial invertebrate inputs. Added wood resulted in greater biomass of aquatic invertebrate biomass, and both input and drift of terrestrial invertebrates were reduced by canopy covers. In terms of total potential prey biomass, the addition of wood with ambient levels of terrestrial invertebrate inputs had the highest standing crop of benthic, wood-living and terrestrial invertebrates combined, whereas the treatment with reduced terrestrial input and no wood added had the lowest standing crop. Our study indicates that forest practices that both reduce the recruitment of wood and the input of terrestrial invertebrates to small streams have negative effects on prey availability for drift-feeding brown trout. The positive effects of wood addition on biomass of aquatic macroinvertebrates may partly compensate for the negative effects of reduced terrestrial invertebrate subsidies.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

The effects of selective logging on the distribution of moths in a Bornean rainforest

The effects of selective logging on the diversity and species composition of moths were investigated by sampling from multiple sites in primary forest, both understorey and canopy, and logged forest at Danum Valley, Sabah, Malaysia. The diversity of individual sites was similar, although rarefied species richness of logged forest was 17% lower than for primary forest (understorey and canopy combined). There was significant heterogeneity in faunal composition and measures of similarity (NESS index) among primary forest understorey sites which may be as great as those between primary understorey and logged forest. The lowest similarity values were between primary forest understorey and canopy, indicating a distinct canopy fauna. A number of species encountered in the logged forest were confined to, or more abundant in, the canopy of primary forest. Approximately 10% of species were confined to primary forest across a range of species' abundances, suggesting this is a minimum estimate for the number of species lost following logging. The importance of accounting for heterogeneity within primary forest and sampling in the canopy when measuring the effects of disturbance on tropical forest communities are emphasized.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

Direct litter interference and indirect soil competitive effects of two contrasting phenotypes of a spiny legume shrub drive the forb composition of an oromediterranean community

Contrasting phenotypes of foundation species are known to differentially affect understorey plants. However, there is little knowledge on both the mechanisms of competition (resource competition versus interference) of stress‐tolerant phenotypes and the importance of indirect interactions. In an oromediterranean community from Mount Lebanon we assessed the effects on understorey forbs of two contrasting phenotypes, a tight competitive from stressful habitat and a loose facilitative from more benign habitat. In a dry south and a wet north exposure we assessed short‐term resource effects removing shrub canopy and long‐term soil effects (including litter interference) with the comparison of forb performances in adjacent naturally open areas vs no shrub. Indirect effects were quantified through the removal of grasses. Abundance, richness and biomass of forbs were measured in all treatments after one year of experiment, together with litter depth and soil moisture. We found strong direct negative soil effects of the tight phenotype on all forb performances and in particular in south exposure. These effects were due to litter interference on water availability, but not to resource competition. They were likely explained by the high hydrophobicity of organic matter accumulating in the stressful habitat of the tight phenotype. We also found an indirect competition of the loose phenotype for forb richness, due to its direct positive soil effect on competitive grasses, and in particular in south exposure. Our results improve our knowledge on the importance of litter interference in dry nutrient‐poor habitats and the role of indirect interactions in phenotypic effects on understorey species.     

Crown illumination limits the population growth rate of a neotropical understorey palm (Geonoma macrostachys,Arecaceae)

The importance of light as a limiting factor for growth and reproductionin tropical understorey plants is well known, but the population-levelconsequences of light limitation remain little explored. Here, I usedemographicmodelling to examine if the effect of canopy gaps on individual-levelperformance in a tropical understorey palm translates into an effect on thepopulation growth rate. The demographic effects of heterogeneity in crownillumination (index of hemispherical canopy openness), forest-phase, and othermicroenvironmental parameters were quantified using data on 2592 individuals ofall sizes in permanent plots followed over a 1.5 year period. Supplementaryexperiments investigating the effects of canopy conditions on seedlingrecruitment were also conducted. Among the microenvironmental parameters onlycrown illumination and forest-phase had strong and consistent demographiceffects. Growth and fecundity generally increased with increasing crownillumination. Under open forest-phase conditions growth tended to increase andsurvival to decrease (due to increased physical damage). Seedlings transplantedinto gap centres were damaged by solarization. The population-level effect ofheterogeneity in crown illumination and forest-phase was investigated usingtransition matrix models. The overall population growth rate was 0.999, and notsignificantly different from 1. Illumination-specific models showed thatG. macrostachys would be unable to persist at permanentlylow illumination, but would increase in density under higher illumination.Forest-phase-specific models gave similar, but weaker results. Thus, inG. macrostachys effects of canopy openness onindividual-level performance strongly limit the population growth rate. Thepresent study thereby shows that fine-scale spatiotemporal variation in canopyopenness in the tropical rain forest understorey can play a central role in thepopulation ecology of shade-tolerant understorey plants.     

Primary forest dynamics in lowland dipterocarp forest at Danum Valley, Sabah, Malaysia, and the role of the understorey

Changes in species composition in two 4-ha plots of lowland dipterocarp rainforest at Danum, Sabah, were measured over ten years (1986-1996) for trees > or = 10 cm girth at breast height (gbh). Each included a lower-slope to ridge gradient. The period lay between two drought events of moderate intensity but the forest showed no large lasting responses, suggesting that its species were well adapted to this regime. Mortality and recruitment rates were not unusual in global or regional comparisons. The forest continued to aggrade from its relatively (for Sabah) low basal area in 1986 and, together with the very open upper canopy structure and an abundance of lianas, this suggests a forest in a late stage of recovery from a major disturbance, yet one continually affected by smaller recent setbacks. Mortality and recruitment rates were not related to population size in 1986, but across subplots recruitment was positively correlated with the density and basal area of small trees (10-< 50cm gbh) forming the dense understorey. Neither rate was related to topography. While species with larger mean gbh had greater relative growth rates (rgr) than smaller ones, subplot mean recruitment rates were correlated with rgr among small trees. Separating understorey species (typically the Euphorbiaceae) from the overstorey (Dipterocarpaceae) showed marked differences in change in mortality with increasing gbh: in the former it increased, in the latter it decreased. Forest processes are centred on this understorey quasi-stratum. The two replicate plots showed a high correspondence in the mortality, recruitment, population changes and growth rates of small trees for the 49 most abundant species in common to both. Overstorey species had higher rgrs than understorey ones, but both showed considerable ranges in mortality and recruitment rates. The supposed trade-off in traits, viz slower rgr, shade tolerance and lower population turnover in the understorey group versus faster potential growth rate, high light responsiveness and high turnover in the overstorey group, was only partly met, as some understorey species were also very dynamic. The forest at Danum, under such a disturbance-recovery regime, can be viewed as having a dynamic equilibrium in functional and structural terms. A second trade-off in shade-tolerance versus drought-tolerance is suggested for among the understorey species. A two-storey (or vertical component) model is proposed where the understorcy-overstorey species' ratio of small stems (currently 2:1) is maintained by a major feedback process. The understorey appears to be an important part of this forest, giving resilience against drought and protecting the overstorey saplings in the long term. This view could be valuable for understanding forest responses to climate change where drought frequency in Borneo is predicted to intensify in the coming decades.     

Patterns in the use of rainforest vertical space by Neotropical aerial insectivorous bats: all the action is up in the canopy

In tropical rainforests environmental conditions vary dramatically from the ground to the canopy, resulting in a marked stratification in the way vertical space is used by organisms, but research work is often limited to the understorey layer. Aerial insectivorous bats are a highly diverse group that plays key roles in the ecology of rainforests, but their use of vertical space remains elusive. Using automatic ultrasound recording stations placed in the canopy, subcanopy and understorey we tested if bat activity and species diversity are vertically stratified, both in the forest interior and near the edges of water bodies. These patterns were tested separately for individual species, and for two functional groups – open space and edge space bats. Insectivorous bat activity increased by roughly seven fold, and species diversity doubled, from the understorey to the canopy. Both edge space and open space bats were more active in the upper strata, but this tendency was much more accentuated in the latter. Myotis riparius was the only species with greater activity near the understorey. These patterns were altered at the edges of water bodies, where vertical stratification was much less marked. The observed patterns are parsimoniously explained by constraints imposed by vegetation clutter that change across strata, which affect bat species differently. Only bats better adapted to closed spaces are usually capable of foraging within the understorey, whereas the majority of species can exploit the free spaces immediately below the canopy; open space bats seem to concentrate their activity above the canopy. This importance of the inter strata open spaces for bat foraging highlights the need to preserve pristine stratified rainforests, as even selective logging usually disrupts vertical stratification. Moreover, the concentration of insectivorous bats at the upper strata of rainforests underlines the need to include canopy level sampling in ecological studies.     

Overstorey–Understorey Interactions Intensify After Drought-Induced Forest Die-Off: Long-Term Effects for Forest Structure and Composition

Severe drought events increasingly affect forests worldwide, but little is known about their long-term effects at the ecosystem level. Competition between trees and herbs (‘overstorey–understorey competition’) for soil water can reduce tree growth and regeneration success and may thereby alter forest structure and composition. However, these effects are typically ignored in modelling studies. To test the long-term impact of water competition by the herbaceous understorey on forest dynamics, we incorporated this process in the dynamic forest landscape model LandClim. Simulations were performed both with and without understorey under current and future climate scenarios (RCP4.5 and RCP8.5) in a drought-prone inner-Alpine valley in Switzerland. Under current climate, herbaceous understorey reduced tree regeneration biomass by up to 51%, particularly in drought-prone landscape positions (i.e., south-facing, low-elevation slopes), where it also caused a shift in forest composition towards drought-tolerant tree species (for example, Quercus pubescens). For adult trees, the understorey had a minor effect on growth. Under future climate change scenarios, increasing drought frequency and intensity resulted in large-scale mortality of canopy trees, which intensified the competitive interaction between the understorey and tree regeneration. At the driest landscape positions, a complete exclusion of tree regeneration and a shift towards an open, savannah-like vegetation occurred. Overall, our results demonstrate that water competition by the herbaceous understorey can cause long-lasting legacy effects on forest structure and composition across drought-prone landscapes, by affecting the vulnerable recruitment phase. Ignoring herbaceous vegetation may thus lead to a strong underestimation of future drought impacts on forests.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

Mechanisms underlying the interaction between Pinus halepensis and the native late-successional shrub Pistacia lentiscus in a semi-arid plantation

Pinus halepensis has been extensively planted in semi-arid areas throughout the world. This has often led to slow-growth stands that: a) suffer from insect plagues, b) promote nutrient depletion and c) fail to promote the recovery of native vegetation. The introduction of native late-successional shrubs in these stands could stimulate successional processes, improve soil conditions and enhance their resilience against disturbances. The main objective of this study was to evaluate the mechanisms underlying the interaction between Pinus and the native late-successional shrub Pistacia lentiscus in a semi-arid plantation. By using manipulative field and laboratory experiments, we evaluated direct (competition for soil resources and allelopathic effects) and indirect (competition with herbaceous understorey) interactions between Pinus and Pistacia . We found no effect of Pinus litter and root exudates on Pistacia growth. In the field, Pistacia seedlings planted under the canopy of Pinus showed higher survival than those planted in open areas with sparse vegetation. Girdling of Pinus trees did not affect the performance of planted Pistacia seedlings, but suppression of the herbaceous understorey significantly enhanced both survival and physiological status of Pistacia seedlings planted under the canopy of Pinus . The magnitude of the interference by herbaceous understorey was considerably higher than that by Pinus . Our results provide evidence that a negative indirect interaction between Pinus and Pistacia , mediated by the herbaceous understorey, is taking place in the afforestation studied, and can help to explain the low rates of colonisation of late-successional woody shrubs typically observed in semi-arid Pinus halepensis plantations.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

Effects of fire and water availability on the emergence and recruitment of grasses,forbs and woody species in a semiarid Chaco savanna

In the semiarid Chaco savannas, the balance between herbaceous and woody species affects landscape configuration and ecosystem services such as forage production. Therefore, understanding the factors that control this balance is crucial for management. In a factorial experiment, we assessed the effects of fire and water availability on the emergence and recruitment of grasses, forbs and woody species. The results showed that the effects of fire and water availability on early regeneration stages differed among the three functional groups. For grasses, fire and water addition enhanced emergence when acted solely, but only the positive effects of fire were translated into a greater recruitment, irrespective of the water availability. For forbs, only water addition, when acted solely, promoted the emergence, although this effect was not translated into a greater recruitment. Lastly, in the case of woody species, both fire and water addition reduced emergence and recruitment. Our results support the notion that both fire and greater water availability would contribute to the maintenance and recovery of a grass‐dominated state of vegetation through a positive effect of fire on grasses recruitment and a general negative effect of fire and water on woody species emergence and recruitment.     

Stratification and diel activity of arthropods in a lowland rainforest in Gabon

The abundance, activity and species richness of arthropods, particularly of insect herbivores, were investigated in the upper canopy and understorey of a lowland rainforest at La Makandé, Gabon. In total 14 161 arthropods were collected with beating, flight interception and sticky traps, from six canopy sites, during the day and at night, from mid-January to mid-March 1999. The effects of stratum were most important, representing between 40 and 70% of the explained variance in arthropod distribution. Site effects represented between 20 and 40% of the variance and emphasized the néed for replication of sampling among canopy sites. Time effects (diel activity) explained a much lower percentage of variance (6–9%). The density and abundance of many arthropod taxa and species were significantly higher in the upper canopy than in the understorey. Arthropod activity was also higher during the day than at night. In particular, insect herbivores were 2.5 times more abundant and twice as speciose in the upper canopy than in the understorey, a probable response to the greater and more diverse food resources in the former stratum. Faunal overlap between the upper canopy and understorey was low. The most dissimilar herbivore communities foraged in the understorey at night and the upper canopy during the day. Further, a taxonomic study of a species-rich genus of herbivore collected there (Agrilus , Coleoptera Buprestidae) confirmed that the fauna of the upper canopy was different, diverse and very poorly known in comparison to that of the understorey. Herbivore turnover between day and night was rather high in the upper canopy and no strong influx of insect herbivores from lower foliage to the upper canopy was detected at night. This suggests that insect herbivores of the upper canopy may be resident and well adapted to environmental conditions there.     

Competitive interactions between overstorey proteas and sprouting understorey species in South African mountain fynbos

Abstract. Previous studies in the mountain fynbos of South Africa have demonstrated that short fire cycles favour the establishment of dense covers of understorey sprouters while longer fire intervals enable the establishment from seed of overstorey proteas and the formation of a overstorey. One consequence of these differences between fire cycle lengths is the effect that understorey sprouters and an overstorey protea canopy have on species richness. In the case of short fire intervals, species richness is decreased while longer intervals between fires allow species richness to decrease or increase depending on the patchiness of the overstorey canopy. Such results are suggestive of competitive effects between understorey sprouters and overstorey canopy proteas. In this study, data were collected from several pyric successional stages in mountain fynbos to study the effect of overstorey proteas on the growth and flowering of understorey sprouters since the last fire. Data were also collected to determine the effect that understorey sprouters had on the establishment and fecundity of overstorey protea species. Competitive interactions between overstorey proteas and sprouting understorey species were evident at all the sites studied. The vegetative growth and seed production of understorey sprouters, which grew under a canopy of overstorey proteas during the current interfire period, were significantly lower than that for plants growing in the open. In addition, the postfire growth and seed production of understorey sprouters were significantly lower for individuals, which grew under an overstorey protea canopy during the previous fire cycle, than for those individuals which grew in the open. The fecundity of overstorey proteas, which grew near understorey sprouters, was lower than that of plants which grew in the open. This effect was evident for up to the first 15 years after a fire. However, not all understorey sprouters affected the overstorey proteas equally. Also, seedlings of overstorey proteas established significantly less successfully in close proximity to understorey sprouters after a fire than in the open or under proteas. Finally, the results demonstrate that complex species‐specific, understorey–overstorey interactions are important in mountain fynbos. For example, some overstorey species depend on trophically similar species to reduce potential competition from understorey sprouters for their successful establishment at a site.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Forest bird diversity in Mediterranean areas affected by wildfires: a multi-scale approach

Fire is a key mechanism creating and maintaining habitat heterogeneity in Mediterranean landscapes by turning continuous woody landscapes into mosaics of forests and shrublands. Due to the long historical role of fires in the Mediterranean, we hypothesised a moderate negative effect of this type of perturbation on forest bird distribution at a landscape level. We conducted point bird censuses in Aleppo pine forest patches surrounded by burnt shrublands and studied the relationships between three ecological groups of bird species (forest canopy species, forest understorey species, and ubiquitous species) and the features of local habitat, whole patch and surrounding landscape. We used a multi-scale approach to assess the effects of landscape variables at increasing spatial scales on point bird richness. Regarding local habitat components, canopy species were positively associated with tall pines while understorey species with the cover of shrubs and plants from holm-oak forests. Forest birds were positively related to patch size and irregular forest shapes, that is, with high perimeter/size ratios. Thus, these species did not seem to perceive edges as low quality but rather favourable microhabitats. We did not detect any negative effect of isolation or cover of woodlands in the landscape on the presence of forest species after local habitat factors had been accounted for. Finally, only local habitat factors entered the model for ubiquitous species. We suggest that mosaic-like landscapes shaped by fires in the Mediterranean basin are not strongly associated with negative effects fragmentation on forest birds other than those related with habitat loss.