Causes of arrested succession in coastal dune forest

Dispersal or recruitment limitation may arrest succession after disturbance. In north-eastern South Africa the Acacia karroo successional pathway is used to facilitate coastal forest recovery after strip-mining. However, although A. karroo establishes naturally, it forms monospecific stands, arresting forest succession for decades. This casts doubt on the efficacy of this restoration pathway. We investigated the causes of arrested succession. The seed and seedling banks of A. karroo stands and of forest at Cape Vidal, and three A. karroo stands (7–27 years old) on rehabilitated strip-mined dunes at nearby Richards Bay were examined. The establishment and growth of seedlings at Cape Vidal were also considered. The seed bank was larger and more diverse in forest, but the seedling bank was larger in Acacia stands. At Richards Bay, the size of the seed bank increased and the seedling bank decreased with Acacia stand age. Excluding mammalian herbivores in Acacia stands at Cape Vidal resulted in greater species richness and survival of naturally established seedlings, as well as two experimentally planted species. Neither seed dispersal nor seedling establishment limited recruitment of tree species in Acacia stands. Herbivory arrested forest succession by causing the differential mortality of seedlings. In contrast, at Richards Bay where there were few mammalian herbivores, the advanced regeneration in A. karroo stands converged on the diversity of nearby forests 29 years after restoration. Controlling herbivore access and seeding Acacia stands with forest species are site-specific options for preventing arrested succession when using the A. karroo successional pathway.     

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.     

Succession and Micro-elevation Effects on Seedling Establishment of Calophyllum brasiliense Camb. (Clusiaceae) in an Amazonian River Meander Forest1

I investigated the effects of successional stage and micro‐elevation on seedling establishment of Calophyllum brasiliense (Clusiaceae), a common canopy tree of seasonally flooded lowland forest along the Manú River meander zone in southeastern Peru. To compare seedling establishment between microhabitat types, I planted C. brasiliense seeds in a fully crossed experimental design of three successional stages (early, mid, and mature) and two micro‐elevations (levees and backwaters). Seedling establishment success in this study was affected by both successional stage and micro‐elevation, but micro‐elevation was most important in mid‐successional habitats. In general, seedlings in early succession experienced better conditions than in mature forest; light levels were higher, herbivory lower, and seedling growth higher. In mid‐successional forest, micro‐elevation determined habitat quality; backwaters had higher light levels, lower herbivory, and higher seedling growth and survival than levees. Mid‐successional backwaters were similar in quality to early successional forest for seedling establishment, while levees in that same successional stage were the poorest microhabitats for establishment. Although mid‐successional backwaters are similar to early succession for seedling establishment, in the long run, seedlings that establish in mid‐succession have a lower chance of reaching reproductive size before their habitat ages to mature forest than members of their cohort that established in early succession. I hypothesize that successful recruitment for C. brasiliense in the Manú River meander system requires dispersal to early successional habitat.     

Differences in seed rain composition in small and large fragments in the northeast Brazilian Atlantic Forest

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8–388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1‐year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal‐dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small‐sized seeds (<0.3 cm) and less large‐seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small‐sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large‐seeded tree species may facilitate the maintenance of species diversity.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

Functional traits associated with plant colonizing and competitive ability influence species abundance during secondary succession: Evidence from subalpine meadows of the Qinghai–Tibetan Plateau

It is widely recognized that colonists and competitors dominate early and late succession, respectively, with selected species having different colonizing and competitive abilities. However, it remains unknown whether colonizing and competitive ability can determine species abundance directly over succession. The data for five key functional traits were collected (photosynthesis rate, leaf turgor loss point, leaf proline content, seed mass, and seed germination rate), which are direct indicators of plant competitive and colonizing abilities including growth, drought and cold stress resistance, dispersal, and seed dormancy. Here, we tested the effects of colonizing and competitive abilities on species abundance, by employing a linear mixed‐effects model to examine the shifts in the relationship between species abundance and these five colonization and competition‐related traits in species‐rich subalpine secondary successional meadows (at 4, 6, 10, 13 years of age, and undisturbed, respectively) of the Qinghai–Tibetan Plateau. The abundant species at the early‐successional meadows tend to have high photosynthetic rate, high leaf proline content, low seed mass, and seed germination rate for having high colonizing ability, but low competitive ability. By contrast, late‐successional communities tend to be dominated by species with high competitive ability, but low colonizing ability, indicated by large seeds, high seed germination rate, low photosynthetic rate, and leaf proline content. The observed directional shifts in the relationships between traits (photosynthetic rate, leaf proline content, seed mass, and seed germination rate) and abundance with successional age, bring two new understandings of community assembly during succession of subalpine meadows in the Qinghai–Tibetan Plateau. First, it discloses that the differences in species abundance over succession can be directly attributed to differences in colonizing and competitive abilities of different species. Second, it expands the effects of multiple life historical differences including growth, resource competitive ability, cold stress resistance, dispersal, and seed germination strategy, represented by functional traits on community assembly along succession, that is, from the species to the community level.     

Diversity and ecology of lianas in tropical dry evergreen forests on the Coromandel Coast of India under various disturbance regimes

In this study we attempted to explore patterns of diversity, abundance, climbing and dispersal mode of lianas in relation to disturbance in 40 Indian subtropical dry forests. The sites were selected to represent four disturbance categories: relatively undisturbed, moderately disturbed, much disturbed and heavily disturbed. All lianas ≥1 cm dbh were counted, which resulted in a total amount of 5689 individuals of lianas, representing 77 species in 62 genera and 32 families. Liana species richness and abundance increased with forest disturbance, but the liana basal area values showed an opposite trend, with high scores in undisturbed sites. Twining was the main climbing mechanism (61.3%) and zoochory (59.6%) was the main dispersal mode in all the four forest categories. Application of Bray–Curtis cluster analysis produced three distinct clusters in which the much disturbed category was more distant from the others. High abundance of large lianas in undisturbed sites and that of the invasive Lantana camara in heavily disturbed site signals the conservation significance of the less disturbed study sites. The predominance of zoochorous dispersal indicates the faunal dependence of lianas, besides of host trees, thus underlining the need for a holistic approach in biodiversity conservation of this and similar tropical forests.     

Variations and Trade‐offs in Functional Traits of Tree Seedlings during Secondary Succession in a Tropical Lowland Rain Forest

The seedling stage is generally the most important bottleneck for the successful regeneration of trees in forests. The traits of seedlings, particularly biomass allocation and root traits, are more easily quantified than the traits of adults. In this study, we tested the hypothesis that seedling traits vary and trade‐off tracking the changing environment during secondary succession. We measured the major morphological traits of 27 dominant species and the major environmental factors in a chronosequence (30‐yr‐old fallow, 60‐yr‐old fallow, and old growth forest) after shifting cultivation in a tropical lowland rain forest on Hainan Island, China. The 30‐yr‐old fallow had higher light and nutrient availability, and the older forests had higher soil water content. Redundancy analysis based on species abundance and environmental factors revealed groups of seedlings that dominate in different stages of succession. Seedlings in different stages of succession had different strategies of biomass allocation for harvesting resources that varied in availability. Species characteristic of younger forest had higher allocation to roots and higher specific leaf area, while species characteristic of older forest had higher allocation to leaves. Our study suggests that the variations and trade‐offs in the major functional traits of tree seedlings among successional classes may reflect changes in environmental conditions during succession.     

Restoration of Araucaria Forest: The Role of Perches, Pioneer Vegetation, and Soil Fertility

The Araucaria forest is Brazil's highly threatened subtropical forest ecosystem that has been disappearing in recent decades. Restoration programs involving this forest type are scarce, and there is a lack of scientific information on how ecological processes such as competition, facilitation, and seed dispersal influence natural forest restoration. This work aims to investigate how use of perches to attract seed dispersers and the influence of pioneer vegetation and soil fertilization could affect the colonization of woody species in a degraded area. An experiment was conducted in an abandoned field where the natural establishment of seeds and seedlings of woody species was monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control); and (3) perches (presence and absence). Seed and seedling abundance, seed and seedling species richness, and seedling mortality were recorded monthly during 12 months. Seed abundance and species richness were significantly greater in places with perches than in control plots. These results were consistent over the year and more pronounced when the surrounding forest produced a higher amount of fruit. Species richness and abundance of seedlings were significantly greater in places with perches than in control plots, and in places with vegetation than without. Soil fertility did not influence seedling establishment. Facilitation and seed dispersal are important factors affecting the colonization of woody species in this subtropical area. Nutrient availability neither regulates the facilitation process nor influences species replacement during the early stages of Araucaria forest succession.     

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.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.     

Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.     

Drivers of seedling survival in a temperate forest and their relative importance at three stages of succession

Negative density dependence (NDD) and niche partitioning have been perceived as important mechanisms for the maintenance of species diversity. However, little is known about their relative contributions to seedling survival. We examined the effects of biotic and abiotic neighborhoods and the variations of biotic neighborhoods among species using survival data for 7503 seedlings belonging to 22 woody species over a period of 2 years in three different forest types, a half‐mature forest (HF), a mature forest (MF), and an old‐growth forest (OGF), each of these representing a specific successional stage in a temperate forest ecosystem in northeastern China. We found a convincing evidence for the existence of NDD in temperate forest ecosystems. The biotic and abiotic variables affecting seedlings survival change with successional stage, seedling size, and age. The strength of NDD for the smaller (<20 cm in height) and younger seedlings (1–2 years) as well as all seedlings combined varies significantly among species. We found no evidence that a community compensatory trend (CCT) existed in our study area. The results of this study demonstrate that the relative importance of NDD and habitat niche partitioning in driving seedling survival varies with seedling size and age and that the biotic and abiotic factors affecting seedlings survival change with successional stage.     

Seed rain and advanced regeneration in a tropical rain forest

By comparing seed rain, seedling and sapling community structures we assessed the possible role played by vertebrate seed dispersal as a structuring factor in advanced regeneration of closed-canopied sites in the tropical rain forest of Los Tuxtlas, Mexico. Seed weight, initial morphology of seedlings and species abundance were also analyzed to determine if these traits influenced the probability of establishment in the shade.About half of the seed species falling in five closed forest sites (25×25 m) during one year came from fruiting trees growing within the sites (local seeds) and half from fruiting trees found outside the sites (immigrant seeds). Seeds of liana and upper-canopy species were over-represented among immigrant seeds compared with seeds of understory tree species. This probably reflects the activity of frugivorous arboreal mammals, bats, and birds. Species with immigrant seeds had both a lower abundance and a narrower spatial distribution than locally produced seeds. Therefore, immigrant seeds showed higher diversity values than locally produced seeds.Average seed size and the proportion of epigeous seedlings were similar in local and immigrant species. Under closed-canopied sites, factors affecting community organization seem to operate selectively, favoring the establishment of large-seeded, local abundant species in the advanced regeneration. However, the fact that some saplings of immigrant species were found in the plots suggest that a slow species infiltration may be occurring leading to a slow shift in the advanced regeneration species composition. We propose that the influence of seed dispersal on advanced regeneration structure depends on the disturbance history of the patches where seeds land.     

Effect of the grass Leersia hexandra on the dispersal,seed germination,and establishment of Pachira aquatica seedlings

1. In wetlands, hydrochory is one of the main mechanisms of seed dispersal and there is often synchrony between propagule production and the flood season. Different sources of disturbance can prevent seed dispersal to suitable sites, and if environmental conditions are not adequate for germination and seedling establishment, recruitment will be limited, affecting succession.
2. We worked in a disturbed tropical freshwater swamp where the native grass Leersia hexandra has dominated open areas, creating a grass matrix that surrounds patches of swamp forest. Leersia grows vigorously, forming cushions of dry matter that cover the soil, forming a potential obstacle to seed dispersal. We asked whether the vegetative growth of this grass prevents the entry and dispersal of seeds of the tropical swamp tree Pachira aquatica, thwarting seed germination and seedling establishment, and arresting succession. We set up transects in the grass matrix in two zones: close to the river (R) and bordering the tree patches or fragments (F). We quantified tree seed and seedling presence, survival and growth in situ and experimentally introduced seeds and seedlings in the field and monitored seed germination and the survival and growth of their seedlings, as well as that of transplanted seedlings.
3. There was a negative relationship between the number of seeds and established seedlings, and the distance to river or fragment (r = −0.86, p < 0.001 for zone R; and r = −0.77, p < 0.001 for zone F) and with the grass cushion (r = −0.68, p = 0.005 for zone R; and r = −0.66, p = 0.007 for zone F); the grass creating a barrier to dispersal. When seeds were sown after clearing the grass cushion, germination success was high, so this stage is not limited. The transplanted seedlings had better survival and a greater final height than the seedlings of the sown seeds. Grass cover had a negative effect on both types of seedlings. Seedling survival rates were inversely related to grass cover, showing that seedlings overgrown by grass had low survival rates. Flooding is a stress factor for seedlings and produced mortality, in addition to the effects of the grass.
4. Together, the field survey and the experiment show that succession is being arrested in two ways: (1) by limiting seed dispersal because the grass cushion slows the dispersal and penetration of seeds into the vegetation; and (2) by limiting seedling establishment because the grass competes for space and light. Our results show that even where the grass is native, slower growing, seed-dependent species may struggle to compete and establish. If grass cover is increasing, these swamps are very vulnerable to a decrease in area because it is very difficult for them to regenerate naturally.

     

Differential seedling regeneration patterns across forest–grassland ecotones in two tropical treeline species (Polylepis spp.)

Successful forest expansion into grassland can be limited by seed dispersal and adverse conditions for tree seedlings in the grassland environment. In the high‐elevation Andes, human‐induced fragmentation has exacerbated the patchy distribution of Polylepis forests, threatening their unique biological communities and spurring restoration interest. Studies of Polylepis forest extent in Peru suggest that forest borders have remained stable over the past century despite decreasing anthropogenic disturbance, suggesting that tree seedling recruitment is being limited in the open grassland habitat. We studied natural seedling dispersion patterns of Polylepis sericea and Polylepis weberbaueri (Rosaceae) at forest–grassland edges across a range of environmental conditions to examine seedling recruitment and colonization of grasslands in Huascaran National Park (Peru). Using data from 2367 seedlings found in 48 forest–grassland edge plots (15 m × 15 m) at forest patches between 3900–4500 masl, we employed generalized mixed modelling to identify the significant associations of seedling densities with environmental covariates. In addition, we compared these associations to patterns of adult presence on the landscape. Seedling densities were associated with a combination of variables varying within (distance to forest edge) and among (elevation and dry season solar irradiation) plots across the landscape. For both species, seedling densities decreased with increasing distance away from the forest in a manner consistent with short‐distance seed dispersal by wind. Our results suggest that such short‐distance dispersal may slow forest expansion, but that there also appear to be substantial post‐dispersal limitations to seedling establishment in the grassland. Polylepis sericea densities decreased with elevation, while P. weberbaueri increased with elevation and decreased with solar irradiation. Associations of adult presence with elevation and solar irradiation mirrored those of seedling densities. Management of areas with forest patches dominated by these species should consider these differences in their environmental tolerances, particularly during species selection and zonation for reforestation.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than environmental factors.     

The potential of small exclosures in assisting regeneration of coffee shade trees in South‐Western Ethiopian coffee forests

Ethiopian Afromontane moist forests where coffee grows as understorey shrub are traditionally managed by the local communities for coffee production through thinning of the shade tree canopy and slashing of competing undergrowth. This management practice has a negative impact on the coffee shrubs, because the removal of shade tree saplings and seedlings reduces the succession potential of the shade tree canopy, which threatens the very existence of the shade coffee production system. We assessed the functionality of small exclosures to initiate coffee shade tree canopy restoration through natural regeneration. Our results show that small exclosures have a strong restoration potential for the coffee shade trees preferred by farmers (Albizia schimperiana, A. gummifera and Millettia ferruginea), as evidenced from their seedling abundance, survival and growth. The regeneration of late‐successional tree species of the moist Afromontane forest was not successful in the small exclosures, most probably due to the low abundance or absence of adult trees as seed sources for regeneration. Therefore, temporary establishment of small exclosures in degraded coffee forest fragments where shade trees are getting old or dying is recommended for sustainable shade coffee production.