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.     

Can Establishment Characteristics Explain the Poor Colonization Success of Late Successional Grassland Species on Ex‐Arable Land?

Abstract Many grassland restoration projects on former arable land face problems because early successional grassland species establish vigorously and persistently but late successional grassland species fail to establish. Differences in establishment characteristics of early and late successional species might provide an explanation for the failure of many late successional species to colonize grasslands on ex‐arable land. I examined whether early and late successional species had different establishment rates in the initial years of a grassland succession, whether a specific establishment stage (seedling emergence, mortality or growth) could be identified as the key process controlling establishment, and what management would enhance the establishment of late successional grassland species. Seeds of three early and three late successional species were sown separately in ex‐arable plots with bare soil, 1‐year‐old vegetation, and 2‐year‐old vegetation. Emergence, mortality, and seedling growth were monitored for 1 year. Early successional species established successfully in the bare soil plots but failed to establish in plots with 1‐ and 2‐year‐old vegetation. Late successional species showed either lower establishment rates in the younger succession stages or decreased establishment with succession that nevertheless resulted in significant establishment in the oldest plots. Seedling emergence proved to be the key factor determining the establishment pattern of early and late successional species. In absolute numbers, emergence of late successional species was, however, similar or higher than that of early successional species, even in the earliest succession stage. The poor establishment of late successional species on former arable land could therefore not be explained solely by differences in establishment characteristics between early and late successional grassland species. Competitive processes between early and late successional species later in the life cycle probably play an important role. The results do point out that establishment of late successional species can be promoted by creating vegetative cover from the start of the restoration effort.     

Phyllostomid bat assemblages in different successional stages of tropical rain forest in Chiapas,Mexico

Due to their role in seed dispersal, changes in the community of phyllostomid bats have direct consequences on ecological succession. The objective of this work was to document changes in the structure of bat assemblages among secondary successional stages of tropical rain forest in Chiapas, Mexico. Bats were mist-netted at ground level during 18 months in 10 sites belonging to 3 successional stages: four sites represented early succession (2–8 years of abandonment), four intermediate succession (10–20 years of abandonment), and two late succession (mature old-growth forest).We captured 1,179 phyllostomids comprising 29 species. Phyllostomid species richness was 17 (58% of all species) in the early stage, 18 (62%) in the intermediate stage and 24 (83%) in the late stage. The late successional mature forest possessed nine species that were exclusively found there, whereas early and intermediate successional stages contained only one exclusive species. Sturnira lilium, Artibeus lituratus, Carollia perpicillata, Artibeus jamaicensis and Glossophaga soricina represented 88% of all captured phyllostomid bats. Frugivores made up more than 90% of the species captured in early and intermediate successional stages and 84% in late successional forest. The Bray–Curtis index of dissimilarity showed a replacement of species through successional stages with the largest dissimilarity between early and late stages, followed by intermediate and late, and the lowest dissimilarity between early and intermediate stages. The number of gleaning insectivore species increased during succession. The carnivorous guild was exclusively found in the late stage (three species). We conclude that the late successional mature forest was the main reservoir for the gleaning insectivore and carnivore guilds; however, early and intermediate successional stages possessed a great diversity of species including many frugivores.     

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.     

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.     

Cropland edge,forest succession,and landscape affect shrubland bird nest predation

The effects of habitat edges on nest survival of shrubland birds, many of which have experienced significant declines in the eastern United States, have not been thoroughly studied. In 2007 and 2008, we collected data on nests of 5 shrubland passerine species in 12 early successional forest patches in North Carolina, USA. We used model selection methods to assess the effect of distance to cropland and mature forest edge on nest predation rates and additionally accounted for temporal trends, nest stage, vegetation structure, and landscape context. For nests of all species combined, nest predation decreased with increasing distance to cropland edge, by nearly 50% at 250 m from the cropland edge. Nest predation of all species combined also was higher in patches with taller saplings and less understory vegetation, especially in the second year of our study when trees were 4–6 m tall. Predation of field sparrow (Spizella pusilla) nests was lower in landscapes with higher agricultural landcover. Nest predation risk for shrubland birds appears to be greater near agricultural edges than mature forest edges, and natural forest succession may drive patterns of local extirpation of shrubland birds in early successional forest patches. Thus, we suggest that habitat patches managed for shrubland bird populations should be considerably large or wide (>250 m) when adjacent to crop fields and maintained in structurally diverse early seral stages. © 2011 The Wildlife Society.     

Effects of Small Rodent and Large Mammal Exclusion on Seedling Recruitment in Costa Rica1

We examined whether the experimental exclusion of large mammalian and small rodent seed predators had differing effects on seedling recruitment under natural seed rain conditions. In both primary and late‐successional secondary forested areas, exclosure experiments using natural seed densities were designed to assess seedling recruitment. To assess the differences in seedling recruitment, we monitored three exclosure treatments (1.2 m radius/1.5 m height) in two forest types (primary vs. late‐successional secondary forest): (1) fenced exclosures that excluded large mammals; (2) fenced exclosures that excluded both large and small mammals; and (3) open controls. Within each exclosure treatment, we marked and identified all seedlings at the beginning of the experiment (February 2001), followed the marked seedlings' fate for a year, and then marked and identified all new seedlings after a year. Two preliminary findings were generated from these data: for some tree species, small rodents and large mammals have differential effects on seedling recruitment, and the effect of excluding mammals did not differ with habitat type (primary vs. late‐successional secondary forest). These preliminary results highlight the need to examine further how the effects of small rodent and large mammal exclusion may affect species‐specific seed predation and seedling recruitment in a variety of habitat/land use types (e.g., primary forest, late‐successional forest, and early‐successional forest).     

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.     

Arctiid moth ensembles along a successional gradient in the Ecuadorian montane rain forest zone: how different are subfamilies and tribes?

Aim We examined changes in the species diversity and faunal composition of arctiid moths along a successional gradient at a fine spatial scale in one of the world's hot spots for moths, the Andean montane rain forest zone. We specifically aimed to discover whether moth groups with divergent life histories respond differentially to forest recovery. Location Southern Ecuador (province Zamora‐Chinchipe) along a gradient from early successional stages to mature forest understorey at elevations of 1800–2005 m a.s.l. Methods Moths were sampled with weak light traps at 21 sites representing three habitat categories (early and late succession, mature forest understorey), and were analysed at species level. Relative proportions were calculated from species numbers as well as from specimen numbers. Fisher's α was used as a measure of local diversity, and for ordination analyses non‐metric multidimensional scaling (NMDS) was carried out. Results Proportions of higher arctiid taxa changed distinctly along the successional gradient. Ctenuchini (wasp moths) contributed more strongly to ensembles in natural forest, whereas Lithosiinae (lichen moths) decreased numerically with forest recovery. Arctiid species diversity (measured as Fisher's α) was high in all habitats sampled. The three larger subordinated taxa contributed differentially to richness: Phaegopterini (tiger moths) were always the most diverse clade, followed by Ctenuchini and Lithosiinae. Local species diversity was higher in successional habitats than in forest understorey, and this was most pronounced for the Phaegopterini. Dominance of a few common species was higher, and the proportion of species represented as singletons was lower, than reported for many other tropical arthropod communities. NMDS revealed a significant segregation between ensembles from successional sites and from forest understorey for all larger subordinated taxa (Phaegopterini, Ctenuchini, Lithosiinae). Abandoned pastures held an impoverished, distinct fauna. Faunal segregation was more pronounced for rare species. Ordination axes reflected primarily the degree of habitat disturbance (openness of vegetation, distance of sites from mature forest) and, to a lesser extent, altitude, but not distance between sampling sites. Main conclusions Despite the geographical proximity of the 21 sites and the pronounced dispersal abilities of adult arctiid moths, local ecological processes were strong enough to allow differentiation between ensembles from mature forest and disturbed sites, even at the level of subfamilies and tribes. Differences in morphology and life‐history characteristics of higher arctiid taxa were reflected in their differential representation (proportions of species and individuals) at the sites, whereas patterns of alpha and beta diversity were concordant. However, concordance was too low to allow for reliable extrapolation, in terms of biodiversity indication, from one tribe or subfamily to the entire family Arctiidae. Phaegopterini (comprising more putative generalist feeders during the larval stages) benefited from habitat disturbance, whereas Ctenuchini (with host‐specialist larvae) were more strongly affiliated with forest habitats.     

Recovery from clearing,cyclone and fire in rain forests of Tonga,South Pacific: Vegetation dynamics 1995–2005

Abstract In late 2001 a category 3 cyclone impacted forest plots that were established in Tonga in 1995, and additionally, one plot was accidentally burned by an escaped land‐clearing fire. Subsequent surveys provide observations of 10 years of forest dynamics in this poorly studied region, and the first reported observations of large interannual variation in juvenile (seedling and sapling) abundance in the western tropical Pacific. The severely disturbed (burned) plot was initially colonized by a non‐native early pioneer, Carica papaya L., but 3.5 years later a native pioneer, Macaranga harveyana (Muell. Arg.) Muell. Arg., was the most abundant tree species. The seedling layer included some long‐lived pioneers and shade‐tolerant species. Two mature forest plots affected only by the cyclone changed very little over a decade. Late‐successional shade‐tolerant species that dominated the overstory were also abundant as seedlings and saplings. This is in contrast with a 30‐ to 40‐year‐old, formerly cultivated, secondary forest plot that still shows no recruitment of late‐successional dominants, in spite of the proximity of remnant forest patches. This study suggests differing pathways of succession following shifting cultivation versus cyclone and fire disturbances in Tonga. Land use legacies appear to have a long‐lasting effect on community composition.     

An ericoid shrub plays a dual role in recruiting both pines and their fungal symbionts along primary succession gradients

Relative importance of positive and negative interactions between plant species may change along disturbance and resource gradients. Positive interactions are suggested to prevail in low resource, low productivity (high stress) conditions and negative interactions in high resource availability. A dwarf shrub, mountain crowberry Empetrum nigrum ssp. hermaphroditum, is known to have allelopathic impacts on both Scots pine Pinus sylvestris and its ectomycorrhizal symbionts. We aimed to study if the outcome of Empetrum impacts on Scots pine changes along primary succession gradients on the dune shores of Bothnian Bay, in Finland, where abiotic stress gradually changes to biotic stress along the succession. We found that Empetrum may act as a facilitator despite its allelopathic effects, since the proportion of Scots pine seedlings established in Empetrum patches was higher than in patches without the shrub in early and mid succession stages, whereas patches without Empetrum were preferred in late succession. The amount of mycelial fungal biomass (ergosterol) in the soil in the vicinity of the seedling roots was higher in Empetrum patches than in patches without Empetrum and it increased along the succession gradient. Proportion of pine root tips colonised by suilloid morphotypes with abundant external mycelia and the diversity of ectomycorrhizal morphotypes were higher in mid successional stage in Empetrum patches compared to patches without Empetrum. Our results suggest that in the harsh physical conditions of the dune shore Empetrum facilitates pine seedling establishment in the early and mid stages of succession by providing mechanical and physical shelter whereas in late succession negative interactions (competition and allelopathy) between the shrub and the pine are dominating. To our knowledge we present the first finding that an ericoid mycorrhizal shrub could enhance both the performance of the ectomycorrhizal host tree and the tree's fungal symbionts.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Limited by the host: Host age hampers establishment of holoparasite Cuscuta epithymum

A good understanding of the relationship between plant establishment and the ecosystem of which they are part of is needed to conserve rare plant species. Introduction experiments offer a direct test of recruitment limitation, but generally only the seed germination and seedling phases are monitored. Thus the relative importance of different establishment stages in the process of recruitment is not considered. This is particularly true for parasitic plants where empirical data are generally missing. During two consecutive growing seasons we examined the effect of heathland management applications, degree of heathland succession (pioneer, building and mature phase) and seed-density on the recruitment and establishment of the endangered holoparasite Cuscuta epithymum. In general, recruitment after two growing seasons was low with 4.79% of the sown seeds that successfully emerged to the seedling stage and a final establishment of 89 flowering adults (i.e. <1.5% of the sown seeds). Although a higher seed-density resulted in a higher number of seedlings, seed-density did not significantly affected relative germination percentages. The management type and subsequent heath succession had no significant effect on seedling emergence; whereas, seedling attachment to the host, establishment and growth to full-grown size were hampered in older heath vegetation (i.e. high, dense, and mature canopy). Establishment was most successful in turf-cut pioneer heathland, characterised by a relatively open and low vegetation of young Calluna vulgaris. The age of C. vulgaris, C. epithymum's main host, proved to be the most limiting factor. These results emphasise the importance of site quality (i.e. successional phase of its host) on recruitment success of C. epithymum, which is directly affected by the management applied to the vegetation. Lack of any heathland management will thus seriously restrict establishment of the endangered parasite.     

Early herbaceous succession along a topographical gradient on forest clear-felling sites in mountainous terrain,central Japan

Early successional patterns of herbaceous communities in forest clear-felling sites were investigated along a topographical gradient, which included ridge, slope and valley habitat types, in warm-temperate evergreen forest regions of central Japan for 5 years. Three dominant species with wind-dispersed seeds played a major role in the succession: an annual,Crassocephalum crepidioides, a biennial,Erigeron canadensis, and a perennial,Miscanthus sinensis. Pioneer herbs that have a seed-bank strategy, which are common in old field succession, were not found in the mountainous sites. The persistence of the annual or biennial dominants during the very early stages of secondary succession was different in the three topographical habitat types. In the ridge habitat,M. sinensis dominated from the first year state, andC. crepidioides andE. canadensis were less prominent.Crassocephalum crepidioides andE. canadensis became dominant as one moved down the slope. In the valley habitat,C. crepidioides dominated in the first year stage, was succeeded byE. canadensis in the second year, and thenM. sinensis gradually replaced it in later years. As all three wind-dispersed dominants simultaneously invaded in all the habitat types after clear-felling, the different successional patterns along the topographical gradient might have resulted from differences in the establishment ability and the growth rate of the three dominants depending on the three habitat types.     

Herbivory and competition slow down invasion of a tall grass along a productivity gradient

Competition models including competition for light predict that small plant species preferred by herbivores will be outshaded by taller unpreferred plant species with increasing productivity. When the tall plant species is little grazed by the herbivores, it can easily invade and dominate short vegetation. The tall-growing grass Elymus athericus dominates the highly productive stages of a salt-marsh succession in Schiermonnikoog and is not preferred by the herbivores which occur there, hares and geese. We studied how interspecific competition and herbivory affected performance during early establishment of this species with increasing productivity. Seedlings were planted in the field in a full factorial design, manipulating both interspecific competition and herbivory. The experiment was replicated along a natural productivity gradient. Competition reduced aboveground biomass production and decreased the number of ramets that were produced but did not affect survival of seedlings. The negative effects of competition on seedling performance increased with increasing productivity. In contrast to our expectations, herbivory strongly reduced seedling survival, especially at the unproductive sites and had only small effects on seedling growth. The present study shows that unpreferred tall-growing species cannot easily invade vegetation composed of short preferred species. Grazing by (intermediate-sized) herbivores can prevent establishment at unproductive sites, and increased competition can prevent a rapid invasion of highly productive sites. Herbivores can have a long-lasting impact on vegetation succession by preventing the establishment of tall-growing species, such as E. athericus, in a window of opportunity at young unproductive successional stages.Plant nomenclature follows Van der Meijden et al. (1990)     

Diversity and composition of Arctiidae moth ensembles along a successional gradient in the Ecuadorian Andes

Andean montane rain forests are among the most species‐rich terrestrial habitats. Little is known about their insect communities and how these respond to anthropogenic habitat alteration. We investigated exceptionally speciose ensembles of nocturnal tiger moths (Arctiidae) at 15 anthropogenically disturbed sites, which together depict a gradient of forest recovery and six closed‐forest understorey sites in southern Ecuador. At weak light traps we sampled 9211 arctiids, representing 287 species. Arctiid abundance and diversity were highest at advanced succession sites, where secondary scrub or young forest had re‐established, followed by early succession sites, and were lowest, but still high, in mature forest understorey. The proportion of rare species showed the reverse pattern. We ordinated moth samples by non‐metric multidimensional scaling using the chord‐normalized expected species shared index (CNESS) index at various levels of the sample size parameter m. A distinct segregation of arctiid ensembles at succession sites from those in mature forest consistently emerged only at high m‐values. Segregation between ensembles of early vs. late succession stages was also clear at high m values only, and was rather weak. Rare species were responsible for much of the faunal difference along the succession gradient, whereas many common arctiid species occurred in all sites. Matrix correlation tests as well as exploration of relationships between ordination axes and environmental variables revealed the degree of habitat openness, and to a lesser extent, elevation, as best predictors of faunal dissimilarity. Faunal differences were not related to geographical distances between sampling sites. Our results suggest that many of the more common tiger moths of Neotropical montane forests have a substantial recolonization potential at the small spatial scale of our study and accordingly occur also in landscape mosaics surrounding nature reserves. These species contribute to the unexpectedly high diversity of arctiid ensembles at disturbed sites, whereas the proportion of rare species declines outside mature forest.     

Canopy manipulation as a tool for restoring mature forest conifers under an early‐successional angiosperm canopy

Low‐light environments in early‐successional forests that have established after abandonment of farming often restrict the establishment of later successional species resulting in an arrested succession. This 6‐year study tested the potential of different canopy manipulations to facilitate the establishment of a light‐demanding canopy tree species, tōtara (Podocarpus totara), within a regenerating kānuka (Kunzea robusta) stand. Results highlighted the effectiveness of artificial gaps over other methods (ring‐barking and edge‐planting) in accelerating the growth of planted tōtara. Seedlings under gaps grew consistently taller and faster over time indicative of an improved understorey light environment. Ring‐barking did not have a significant effect on tōtara growth because only a portion of the treated trees died, and after 6 years dead trees remained standing with intact branches resulting in insignificant increases in light transmission. At the forest edge sites, tōtara growth was highly variable. Although some seedlings grew as tall as in the gaps, others did not. Survival was also lower in the edge sites than in other treatments, which was likely due to enhanced herbivory from ungulates which impacted some plants at these sites. Gap creation is likely to be an important tool for restoring late‐successional canopy species in regenerating stands both through providing ideal sites for the growth of light‐demanding species such as tōtara and through natural establishment of other future canopy trees into the gaps.     

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.     

Germination and seedling survival in fens undergoing succession

We studied mechanisms of vegetation change in fens subject to succession from open water to floating mats and finally herbaceous rich-fens. Earlier research showed that these systems are characterized by transient seed banks. Our main question was whether seedlings of later successional fen stages are already present in earlier stages, remaining subordinate in the vegetation until conditions become suitable for them. If, however, conditions during succession change in a way that only a limited set of species can survive as seedlings during each of the successional stages, no seedling bank will exist. The transient character of the seeds would then imply that seeds will not germinate and will subsequently die and that seeds that have germinated in the “wrong” stage will not become established. We hypothesized that: (1) germination and seedling survival of fen species are significantly better in the successional fen stage for which these species are characteristic, (2) as a consequence no seedling banks occur in these fens. In a field experiment, seeds of five characteristic fen species in the standing vegetation of three successional fen stages i.e. raft fen, quaking fen and rich fen were sown in each of these stages in a turf pond in the Tienhoven area, The Netherlands. Germination and seedling survival were measured over two growing seasons together with environmental variables. Germination was higher in the “own” stage for all species groups as was survival for quaking fen species and rich fen species. For both these stages, percentage of germination and survival of four out of five characteristic species were significantly higher in the “own” stage. Germination and survival can be considered stage-dependent and it was concluded that seedling banks do not exist in these fens. Site-specific environmental variables act as a sieve and differentiate on species presence already during early life history stages. We found clues that the environmental sieve acts at the level of nutrient availability, tolerance for high sulphide concentration and light climate. Because of the transient seed bank and absence of a seedling bank in these fen wetlands, successful establishment of species necessitates a continuous dispersal of characteristic species until the environmental conditions permit establishment. This also implies that species of the whole successional sere should be present within dispersal distance.     

Population dynamics of Echinops gmelinii Turcz. at different successional stages of biological soil crusts in a temperate desert in China

• The effects of biological soil crusts (BSC) on vascular plant growth can be positive, neutral or negative, and little information is available on the impacts of different BSC successional stages on vascular plant population dynamics.
• We analysed seedling emergence, survival, plant growth and reproduction in response to different BSC successional stages (i.e. habitats: bare soil, cyanobacteria, lichen and moss crusts) in natural populations of Echinops gmelinii Turcz. in the Tengger Desert of northwest China. The winter annual E. gmelinii is a dominant pioneer herb after sand stabilisation.
• During the early stages of BSC succession, the studied populations of E. gmelinii were characterised by high density, plant growth and fecundity. As the BSC succession proceeded beyond moss crusts, the fecundity decreased sharply, which limited seedling recruitment. Differences in seedling survival among the successional stages were not evident, indicating that BSC have little effect on survival in arid desert regions. Moreover, E. gmelinii biomass allocation exhibited low plasticity, and only reproductive allocation was sensitive to the various habitats. Our results further suggest that the negative effects of BSC succession on population dynamics are primarily driven by increasing topsoil water‐holding capacity and decreasing rain water infiltration into deeper soil.
• We conclude that BSC succession drives population dynamics of E. gmelinii, primarily via its effect on soil moisture. The primary cause for E. gmelinii population decline during the moss‐dominated stage of BSC succession is decreased fecundity of individual plants, with declining seed mass possibly reducing the success of seedling establishment.