Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Demographic bottlenecks in tropical plant regeneration: A comparative analysis of causal influences

Mortality factors that act sequentially through the demographic transitions from seed to sapling may have critical effects on recruitment success. Understanding how habitat heterogeneity influences the causal factors that limit propagule establishment in natural populations is central to assess these demographic bottlenecks and their consequences. Bamboos often influence forest structure and dynamics and are a major factor in generating landscape complexity and habitat heterogeneity in tropical forests. To understand how patch heterogeneity influences plant recruitment we studied critical establishment stages during early recruitment of Euterpe edulis, Sloanea guianensis and Virola bicuhyba in bamboo and non-bamboo stands in the Brazilian Atlantic forest. We combined observational studies of seed rain and seedling emergence with seed addition experiments to evaluate the transition probabilities among regeneration stages within bamboo and non-bamboo stands. The relative importance of each mortality factor was evaluated by determining how the loss of propagules affected stage-specific recruitment success. Our results revealed that the seed addition treatment significantly increased seedling survivorship for all three species. E. edulis seedling survival probability increased in the addition treatment in the two stand types. However, for S. guianensis and V. bicuhyba this effect depended strongly on artificially protecting the seeds, as both species experienced increased seed and seedling losses due to post-dispersal seed predators and herbivores. Propagules of all three species had a greater probability of reaching subsequent recruitment stages when protected. The recruitment of large-seeded V. bicuhyba and E. edulis appears to be much more limited by post-dispersal factors than by dispersal limitation, whereas the small-seeded S. guianensis showed an even stronger effect of post-dispersal factors causing recruitment collapse in some situations. We demonstrated that E. edulis, S. guianensis and V. bicuhyba are especially susceptible to predation during early compared with later establishment stages and this early stage mortality can be more crucial than stand differences as determinants of successful regeneration. Among-species differences in the relative importance of dispersal vs. establishment limitation are mediated by variability in species responses to patch heterogeneity. Thus, bamboo effects on the early recruitment of non-bamboo species are patchy and species-specific, with successional bamboo patches exerting a far-reaching influence on the heterogeneity of plant species composition and abundance.     

Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

Land crabs as key drivers in tropical coastal forest recruitment

Plant populations are regulated by a diverse assortment of abiotic and biotic factors that influence seed dispersal and viability, and seedling establishment and growth at the microsite. Rarely does one animal guild exert as significant an influence on different plant assemblages as land crabs. We review three tropical coastal ecosystems–mangroves, island maritime forests, and mainland coastal terrestrial forests–where land crabs directly influence forest composition by limiting tree establishment and recruitment. Land crabs differentially prey on seeds, propagules and seedlings along nutrient, chemical and physical environmental gradients. In all of these ecosystems, but especially mangroves, abiotic gradients are well studied, strong and influence plant species distributions. However, we suggest that crab predation has primacy over many of these environmental factors by acting as the first limiting factor of tropical tree recruitment to drive the potential structural and compositional organisation of coastal forests. We show that the influence of crabs varies relative to tidal gradient, shoreline distance, canopy position, time, season, tree species and fruiting periodicity. Crabs also facilitate forest growth and development through such activities as excavation of burrows, creation of soil mounds, aeration of soils, removal of leaf litter into burrows and creation of carbon-rich soil microhabitats. For all three systems, land crabs influence the distribution, density and size-class structure of tree populations. Indeed, crabs are among the major drivers of tree recruitment in tropical coastal forest ecosystems, and their conservation should be included in management plans of these forests.     

Factors limiting plant recruitment in a tropical Afromontane Forest

Predicting how forest species composition may change in response to global change is essential for meaningful management. Which species are most likely to successfully recruit depends on a multitude of factors, but processes operating at the seed-to-seedling transition being especially important. Here, we explore how insufficient seed dispersal (dispersal limitation) and ecological filtering of seedlings (establishment limitation) influence species recruitment in an Afromontane Forest. We combined census data from seeds in seed traps and seed-seedling transition plots in a Nigerian montane forest to measure dispersal and establishment limitation across species, life history strategies, and seed sizes. From 106 seed traps and 318 seedling plots, we recorded 11,278 seeds from 38 species and 2688 seedlings of 61 species. All species except for one liana showed strong dispersal limitation while 23 out of the 30 species showed establishment limitation. The two lianas sampled had significantly lower dispersal limitation compared to the trees investigated in the study. Our results suggest that both dispersal and establishment limitation are markedly high in this Afromontane forest than would be expected in a lowland counterpart.     

Are plant populations seed limited? A critique and meta-analysis of seed addition experiments

We examine the relative importance of processes that underlie plant population abundance and distribution. Two opposing views dominate the field. One posits that the ability to establish at a site is determined by the availability of suitable microsites (establishment limitation), while the second asserts that recruitment is limited by the availability of seeds (seed limitation). An underlying problem is that establishment and seed limitation are typically viewed as mutually exclusive. We conducted a meta-analysis of seed addition experiments to assess the relative strength of establishment and seed limitation to seedling recruitment. We asked (1) To what degree are populations seed and establishment limited? (2) Under what conditions (e.g., habitats and life-history traits) are species more or less limited by each? (3) How can seed addition studies be better designed to enhance our understanding of plant recruitment? We found that, in keeping with previous studies, most species are seed limited. However, the effects of seed addition are typically small, and most added seeds fail to recruit to the seedling stage. As a result, establishment limitation is stronger than seed limitation. Seed limitation was greater for large-seeded species, species in disturbed microsites, and species with relatively short-lived seed banks. Most seed addition experiments cannot assess the relationship between number of seeds added and number of subsequent recruits. This shortcoming can be overcome by increasing the number and range of seed addition treatments.     

Bamboo dominance reduces tree regeneration in a disturbed tropical forest

Human disturbance may change dominance hierarchies of plant communities, and may cause substantial changes in biotic environmental conditions if the new dominant species have properties that differ from the previous dominant species. We examined the effects of bamboos (Bambusa tulda and Cephalostachyum pergracile) and their litter on the overall woody seedling abundance, species richness and diversity in a mixed deciduous forest in northeastern Thailand. These bamboo species are gaining dominance after human disturbance. Our results show that seedling abundance and species richness were reduced by bamboo canopies. Seedling abundance and species diversity under bamboo canopies were affected by bamboo litter, whereas seedling abundance and species diversity outside bamboo canopies did not respond to the mixed-tree litter manipulation. Removal of bamboo litter increased seedling abundance and species diversity. However, bamboo litter addition did not affect seedling abundance or species diversity compared to either control or litter removal. This may indicate that the effect of natural amount of bamboo litter is as high as for litter addition in preventing seedling establishment by woody species and hence in minimizing resource competition. We conclude that undergrowth bamboos and their litter affect tree seedling regeneration differently from mixed-tree litter, causing changes in plant community composition and species diversity. Increased human disturbance, causing a shift in dominance structure of these forests, may result in a concomitant reduction in their overall woody species abundance, richness and diversity. Thus, management of bamboos by controlling their distribution in areas of high bamboo density can be an important forest restoration method.     

Experimental Seed Predator Removal Reveals Shifting Importance of Predation and Dispersal Limitation in Early Life History Stages of Tropical Forest Trees

Recruitment limitation of trees in tropical forests can occur because of high rates of seed predation or low rates of seed dispersal, but the degree to which limitation is influenced by variation in seed predator abundance, and hence variation in seed predation and dispersal, is not well understood. We experimentally reduced the density of a granivorous small mammal (Heteromys desmarestianus) by 90 % to assess the degree to which its rates of seed predation and dispersal limit seed to seedling survival of nine species of trees in a Neotropical lowland forest. Overall, the proportion of seeds that germinated was influenced more by high rates of predation than by limited dispersal. Reduction in density of H. desmarestianus resulted in an order of magnitude decrease in fruit removal rates and an order of magnitude increase in both the absolute and relative numbers of seeds that germinated. However, the proportion of seeds that were cached remained relatively constant across all periods and between control grids and removal plots. In removal plots, H. desmarestianus dispersed and cached about 10 % of the fruits they handled, of which approximately 25 % germinated. This was 2 to 3 times greater than the germination rates of undispersed seeds, and for two species dispersed seeds were the only ones that germinated. The results suggest the simultaneous occurrence of both seed predation and dispersal limitation for trees with animal-dispersed seeds, but there may also be a hierarchy of importance in the relative strength of these two mechanisms that is determined by the dynamics of seed predator populations.     

Litter mixture effects on tropical tree seedling growth--a greenhouse experiment

Decomposing litter provides critical nutrients for plants, particularly in nutrient-poor ecosystems such as tropical forests. We hypothesised that decomposing litter improves the performance of a variety of tropical tree seedlings, and that this litter effect varies depending on the species of litter present in litter mixtures. We addressed these hypotheses with a large pot experiment manipulating a range of different litter mixtures of contrasting quality and using seedlings of four tree species from the Amazonian forest of French Guiana. In contrast to our initial hypothesis, decomposing litter had either neutral or negative impacts on seedling growth, despite strongly different growth rates, biomass allocation patterns and leaf and root traits among tree species. Tree species varied in their responses to litter additions, which were further modified by species identity of the added litter. Our data show litter species-specific effects on growth, biomass allocation and leaf and root traits of tropical tree seedlings. These results suggest that a net nutrient release from decomposing litter does not necessarily improve tree seedling growth, even under nutrient-limiting conditions. In conclusion, litter layer composition may affect seedling establishment and recruitment success beyond litter-derived plant nutrient availability, which may contribute to tree species composition and dynamics in the studied tropical forest.     

Seed-deposition and recruitment patterns of Clusia species in a disturbed tropical montane forest in Bolivia

Spatial patterns of seed dispersal and recruitment of fleshy-fruited plants in tropical forests are supposed to be driven by the activity of animal seed dispersers, but the spatial patterns of seed dispersal, seedlings and saplings have rarely been analyzed simultaneously. We studied seed deposition and recruitment patterns of three Clusia species in a tropical montane forest of the Bolivian Andes and tested whether these patterns changed between habitat types (forest edge vs. forest interior), distance to the fruiting tree and consecutive recruitment stages of the seedlings. We recorded the number of seeds deposited in seed traps to assess the local seed-deposition pattern and the abundance and distribution of seedlings and saplings to evaluate the spatial pattern of recruitment. More seeds were removed and deposited at the forest edge than in the interior. The number of deposited seeds decreased with distance from the fruiting tree and was spatially clustered in both habitat types. The density of 1-yr-old seedlings and saplings was higher at forest edges, whereas the density of 2-yr-old seedlings was similar in both habitat types. While seedlings were almost randomly distributed, seeds and saplings were spatially clustered in both habitat types. Our findings demonstrate systematic changes in spatial patterns of recruits across the plant regeneration cycle and suggest that the differential effects of biotic and abiotic factors determine plant recruitment at the edges and in the interior of tropical montane forests. These differences in the spatial distribution of individuals across recruitment stages may have strong effects on plant community dynamics and influence plant species coexistence in disturbed tropical forests.     

Legacies of the past in the present-day forest biodiversity: a review of past land-use effects on forest plant species composition and diversity

Particularly in the temperate climate zone many forests have, at some moment in their history, been used as agriculture land. Forest cover is therefore often not as stable as it might look. How forest plant communities recovered after agriculture was abandoned allows us to explore some universal questions on how dispersal and environment limit plant species abundance and distribution. All studies looking at the effects of historical land use rely on adequate land use reconstruction. A variety of tools from maps, archival studies, and interviews to field evidence and soil analyses contribute to that. They allow us to distinguish ancient from recent forests and many studies found pronounced differences in forest plant species composition between them. A considerable percentage of our forest flora is associated with ancient forests. These ancient forest plant species (AFS) all have a low colonization capacity, suggesting that dispersal in space (distance related) and time (seed bank related) limit their distribution and abundance. However recent forests generally are suitable for the recruitment of AFS. There is clear evidence that dispersal limitation is more important than recruitment limitation in the distribution of AFS. Dispersal in time, through persistent seed banks, does not play a significant role. Ancient forests are not necessary more species-rich than recent forest, but if diversity is limited to typical forest plant species then ancient forests do have the highest number of plant species, making them highly important for nature conservation. The use of molecular markers, integrated approaches and modelling are all part of the way forward in this field of historical ecology.     

A test of the escape and colonization hypotheses for zoochorous tree species in a Western Amazonian forest

In order to assess the importance of seed dispersal (escape and colonization hypotheses), I used transplant experiments for seeds and seedlings of 5–11 plant species with fleshy fruits in a lowland tropical forest (Tinigua National Park, Colombia). I controlled seed density, distance to parental tree, and habitat type. I monitored seed removal, seedling survival, and seedling growth during the first year of development for an average of 554 seeds and 169 seedlings for each species. I supplemented the experimental results with measurements of natural recruitment. I found little support for the escape hypothesis during the seed and seedling stages. For six species that showed differences in seed removal associated with distance, five showed highest removal away from, than close to parent trees, suggesting predator satiation. Seedling survival during the first year was not consistently associated with low densities and long distances from parent trees. For the majority of species, seedlings did not survive flooding in low basins, and there was growth advantage for most plant species in canopy gaps. These differences imply advantages for seed dispersal to adequate habitats, as predicted by the colonization hypothesis. In contrast to experiments, strong negative distance-dependent effects were evident when analyzing natural recruitment patterns. The ratio between saplings and seedlings was higher away from parent trees for the species with enough recruitment to be analyzed and this suggests that a negative distance-dependent effect may also occur after seedling establishment. This pattern is suspected for several other species, but an analysis with some of the other most common trees showed a variety of negative, neutral, and positive distance dependent effects. This study emphasizes the importance of long-term studies to asses the role of seed dispersal.     

Establishment of Epiphytic Bromeliads in Successional Tropical Premontane Forests in Costa Rica

Plant community composition is the combined result of species-specific competitive abilities and the availability of propagules. For epiphytic plants, current hypotheses consider that dispersal-related factors are most important. By controlling seed dispersal constraints, we experimentally examined whether the community composition of epiphytic bromeliads in a tropical premontane area is determined during early phases of seedling recruitment. Also, we tested whether establishment success was related to eco-physiological traits of the species. A total of 7200 seeds were artificially affixed on several host trees in two secondary forest patches and in a mature forest stand. Four bromeliad species with differing physiological characteristics (CAM, C₃-CAM, and C₃) and habitat preference (secondary vs. primary forest) were selected. We found that differences in seed germination probability among habitats and species were not likely to influence community assembly. After 2 yr, seedling survival and plant development were relatively higher in the early-successional forest. Seedling establishment success was not associated with specific physiological and morphological adaptations or habitat preference of the studied species. Our results were not consistent with the described community composition and rates of population recruitment of the studied species in the same successional habitats. The results support the hypothesis that chance and historic events related to seed dispersal have an important influence on community assembly of epiphytic plants. In addition, differences in growth rates and reproductive turnover among species are expected to influence the relative abundance and recruitment rates in a particular habitat.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

Roles of non‐native species in large‐scale regeneration of moist tropical forests on anthropogenic grassland

This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.     

Epiphytic bromeliad communities in secondary and mature forest in a tropical premontane area

We analyzed the differences in species richness, community composition, population structure and within-tree location of epiphytic bromeliads in contiguous secondary and mature forests in a premontane area in Costa Rica. Diversity in the mature forest was highest, and the communities differed in their composition as well as in the recruitment rates of the dominant species. Guzmania monostachia and Catopsis nutans dominated the secondary forests, whereas Tillandsia fasciculata and T. tricolor were more abundant in the mature forest. The secondary forest species showed high rates of seedling recruitment while the opposite was found for the mature forest species. Species presence and abundance among and within habitats did not correlate with their physiological (i.e. CAM vs. C₃ photosynthesis) or morphological attributes. The spatial distribution patterns were similar among habitats; bromeliads tended to aggregate on a few relatively large phorophytes. The species shared a similar vertical stratification within habitats, except for the two dominant species in the early and mid-successional stages, although its ecological implication is not clear. With some exceptions, conspecifics of different ages were located on similar substrate types (i.e. stems, primary, secondary, or tertiary branches) within the tree-crowns, which suggests limited within-tree dispersion. Differences in species composition and rates of seedling recruitment among secondary and mature forest may arise from ecophysiological differences among species; however, the combined effect of seed availability and dispersal differences may have a larger influence. Thus, epiphyte community assembly can only be understood when the differences in habitat conditions, the availability of propagules, their dispersal characteristics and requirements for seedling establishment are known.