Distribution,species diversity and life-form spectra of plant communities along an altitudinal gradient in the northern slopes of Qilianshan Mountains,Gansu, China

We studied the distribution pattern, species diversity and life-formspectra of plant communities along an altitudinal gradient in the mid-sectionofthe northern slopes of Qilianshan Mountains by means of multivariate analyses.Two data sets (167 species × 75 plots, 10 environmental variables ×75 plots), originated from the fieldworks in 1998–1999, were subjected toTWINSPAN and DCCA, resulting in 8 major plant communities: 1)Asterothamnus centraliasiaticus–Halogetonarachnoideus desert grassland on azonal substrates from 1450 to 1600m and 2) zonal Reaumuria soogorica desertgrassland on gravels from 1470 to 1900 m; 3) Stipaprzewalskii–Stipa purpurea montane grassland from 2200 to 2900m; 4) Polygonum viviparum alpine grasslandfrom 2900 to 3700 m; 5) Caraganastenophylla–Ajaniafruticulosa dry-warm shrubland from 2350 to 2800 m; 6)Sabina przewalskii mid-wet warm forest from 2700 to 3300m; 7) Picea crassifolia cold coniferousforestfrom 2450 to 3200 m; 8) Caragana jubatawet-cold alpine shrubland from 3100 to 3700 m. Species diversityand species richness of both grasslands and forests peaked at the intermediateportion of the elevational gradient. Evenness might be strongly influenced byeither biotic or abiotic factors at a local scale, while seems quiteindependentof an elevational gradient at landscape scales. Beta-diversity decreased from1500 to 3700 m, indicating that species turnover declined withincreased elevation. Both richness of life-form and total species richness in agiven altitudinal belt (gamma-diversity) peaked at intermediate elevations,while relative species richness of different life-form varied differently alongthe altitudinal gradient.     

The effect of fire interval on post‐fire understorey communities in Yellowstone National Park

Questions: How does the time interval between subsequent stand‐replacing fire events affect post‐fire understorey cover and composition following the recent event? How important is fire interval relative to broad‐ or local‐scale environmental variability in structuring post‐fire understorey communities? Location: Subalpine plateaus of Yellowstone National Park (USA) that burned in 1988. Methods: In 2000, we sampled understorey cover and Pinus contorta density in pairs of 12–yr old stands at 25 locations. In each pair, the previous fire interval was either short (7–100 yr) or long (100–395 yr). We analysed variation in understorey species richness, total cover, and cover of functional groups both between site pairs (using paired t‐tests) and across sites that experienced the short fire intervals (using regression and ordination). We regressed three principal components to assess the relative importance of disturbance and broad or local environmental variability on post‐fire understorey cover and richness. Results: Between paired plots, annuals were less abundant and fire‐intolerant species (mostly slow‐growing shrubs) were more abundant following long intervals between prior fires. However, mean total cover and richness did not vary between paired interval classes. Across a gradient of fire intervals ranging from 7–100 yr, total cover, species richness, and the cover of annuals and nitrogen‐fixing species all declined while the abundance of shrubs and fire‐intolerant species increased. The few exotics showed no response to fire interval. Across all sites, broad‐scale variability related to elevation influenced total cover and richness more than fire interval. Conclusions: Significant variation in fire intervals had only minor effects on post‐fire understorey communities following the 1988 fires in Yellowstone National Park.     

Factors affecting species richness of tree regeneration in mixed‐wood stands of central Maine

Question: Two questions about within‐stand spatial variability are addressed in this paper. How does species richness of tree regeneration respond to small‐scale ecological gradients, and what effect does natural Abies balsamea abundance have on the species richness of other tree regeneration? Location: A long‐term, gap‐silviculture experiment, Acadian mixed‐wood forest, Maine, USA. Methods: Eight stands treated with and without gap harvesting were sampled to capture sub‐stand heterogeneity of understorey tree regeneration concurrently with patterning of local stand conditions. Spatial and non‐spatial models were developed to test the relationships between two response variables [species richness of small (height ≥0.1 m, but <0.75 m) and large (height ≥0.75 m, but <1.4 m) regeneration] and five explanatory variables (depth to water table, percentage canopy transmittance, A. balsamea regeneration density, and overstorey basal area and species richness). Results: Despite high unexplained variance for all models, consistent associations among variables were found. Negative associations were found between: (1) the species richness of small regeneration and A. balsamea regeneration density and (2) the species richness of large regeneration and overstorey basal area. Positive associations were found between: (1) the species richness of small regeneration and both overstorey basal area and species richness and (2) the species richness of small and large regeneration and canopy transmittance. Conclusions: Promoting tree species diversity in Acadian mixed‐wood stands may not be achievable through the use of gap‐harvesting alone if the density of understorey Abies balsamea is not reduced either naturally or through silvicultural intervention.     

Formation and maintenance of community boundaries in a sub‐alpine forest landscape in northern Japan

Abstract. We focused on community boundaries in a sub‐alpine forest landscape in the Shiretoko Peninsula, northern Japan. Gradient‐directed transects were conducted on the northwestern slope (ranging 500–600 m a.s.l.) of Mount On'nebetsu (1331 m), where complex topography was formed by past landslides. Pioneer Picea glehnii made up a mosaic of pure stands related to landslides. Structural and compositional changes from P. glehnii pure stands to P. glehnii and Abies sachalinensis mixed stands were characterized by ca. 20 m transitional zones over the landscape. Stand density of the species changed across boundaries. A. sachalinensis preferred less undulated slopes with deep soil and P. glehnii preferred undulated rocky sites. Positive spatial associations between overstorey‐understorey P. glehnii were found at undulated core parts of P. glehnii pure stands. Short‐lived A. sachalinensis grew faster to the smaller maximum size than long‐lived P. glehnii. Undulated topography controlled the increase of A. sachalinensis and provided regeneration sites for P. glehnii, which prevented the general trend of canopy replacement from P. glehnii to late‐successional A. sachalinensis. However, the locations of current boundaries were not accordant with the topographic changes in the meso‐scale landscape. Initial P. glehnii pure stands would extend to larger areas if current patterns reflect vegetation recovery since the last landslide. P. glehnii pure stands with accurate boundaries were not maintained by topographic complexity, but were dynamically arranged by the one‐sided canopy replacements from P. glehnii to A. sachalinensis at less undulated slopes in the sub‐alpine forest landscape.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

A functional analysis of vegetation dynamics in abandoned and restored limestone grasslands

Question: How do various plant functional groups and types are related to regeneration of limestone grasslands and to temporal changes in cover of trees and shrubs? Which mechanisms are responsible for changes in differently treated grasslands? Location: Southern Poland, 50° 01'N, 19° 50'E. Methods: A 12‐year restoration experiment after the clearing of a 35‐yr‐old secondary pine wood developed on limestone grassland, and a parallel observation of succession in neighbouring abandoned grassland were carried out. Changes in cover of species with different functional traits and established strategies in relation to time and cover of shrub layer were analysed using multiple linear regression. Results: Over 12 years the composition and cover of species changed in the studied grasslands, but significant differences still occurred between the old grassland and the grassland restored in former wood gaps and that developed in former closed wood. Despite the very close proximity of the old grassland, the composition of restored grasslands was still much determined by initial conditions. In these grasslands, the cover of Festuco‐Brometea species was significantly lower than in the old grassland. In all sites, the cover of species with large leaves and with leaves distributed regularly along the stem, and the cover of CS strategists increased. With increasing shrub cover in restored grasslands, the cover of annuals and biennials, tall species, and R and CS strategists decreased. Conclusions: Limited availability of seeds seems the principal reason for the weak regeneration of xerothermic limestone grasslands. Trees and shrubs inhibit the developed of both early successional species and perennials occurring in established communities. They significantly hinder regeneration of grasslands and should be cut every five or six years.     

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

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

A regional‐scale consideration of the effects of species richness on above‐ground biomass in temperate natural grasslands of China

Question: Can species richness be a predictor for above‐ground biomass in natural grasslands at a regional scale? Location: A total of 647 sites across temperate natural grasslands of northern China. Methods: Structural equation modelling (SEM) was used to examine the effect of species richness on above‐ground biomass. Asymptotically distribution‐free estimation was selected for parameter estimation. The SEM process was performed at five sample sizes (n=50, 100, 200, 300 and 647). Spatial structure in the original data was examined by calculating Moran's I. Results: SEM run at n=647 revealed a positive effect of species richness on above‐ground biomass after controlling for the influences of bioclimatic factors and grazing. At the four reduced sample size levels, the positive effect held true for most cases in 400 observations (>92%). Conclusions: Contrary to observations in previous studies in natural grasslands, our data showed a positive effect of species richness on above‐ground biomass. This suggests that, as a short‐term effect of diversity on productivity, niche complementarity among coexisting species tends to be an important process in arid and semi‐arid natural grasslands. We hold that biodiversity conservation is among the fundamental approaches required to maintain productivity of grasslands in arid and semi‐arid areas.     

Influence of rocky landscape features and fire regime on vegetation dynamics in Appalachian Quercus forests

Question : How do interactions between rocky landscape features and fire regime influence vegetation dynamics? Location : Continental Eastern USA. Methods : We measured vegetation, disturbance and site characteristics in 40 pairs of rocky and non‐rocky plots: 20 in recently burned stands, and 20 in stands with no evidence of recent fire (‘unburned’ stands). Two‐way analysis of variance (ANOVA) was used to assess the main and interaction effects of fire and rock cover on plant community composition. Results : In burned stands, rock cover had a strong influence on vegetation. Non‐rocky ‘matrix’ forests were dominated by Quercus, and had abundant ground cover and advance regeneration of early and mid‐successional tree species. Burned rocky patches supported greater density of fire‐sensitive species such as Acer rubrum, Sassafras albidum and Nyssa sylvatica and had little advance regeneration or ground cover. Quercus had fewer fire scars and catfaces (open, basal wounds) on rocky patches, suggesting that rocky features mitigate fire severity. In unburned stands, differences between rocky and non‐rocky patches were less distinct, with both patch types having sparse ground cover, little tree regeneration, and high understorey densities of relatively shade tolerant A. rubrum, N. sylvatica and Betula lenta. Conclusion : Under a sustained fire regime, heterogeneity in rock cover created a mosaic where fire‐adapted species such as Quercus dominate the landscape, but where fire‐sensitive species persisted in isolated pockets of lower fire severity. Without fire, species and landscape richness may decline as early‐mid successional species are replaced by more shade tolerant competitors.     

Snowbeds are more affected than other subalpine–alpine plant communities by climate change in the Swiss Alps

While the upward shift of plant species has been observed on many alpine and nival summits, the reaction of the subalpine and lower alpine plant communities to the current warming and lower snow precipitation has been little investigated so far. To this aim, 63 old, exhaustive plant inventories, distributed along a subalpine–alpine elevation gradient of the Swiss Alps and covering different plant community types (acidic and calcareous grasslands; windy ridges; snowbeds), were revisited after 25–50 years. Old and recent inventories were compared in terms of species diversity with Simpson diversity and Bray–Curtis dissimilarity indices, and in terms of community composition with principal component analysis. Changes in ecological conditions were inferred from the ecological indicator values. The alpha‐diversity increased in every plant community, likely because of the arrival of new species. As observed on mountain summits, the new species led to a homogenization of community compositions. The grasslands were quite stable in terms of species composition, whatever the bedrock type. Indeed, the newly arrived species were part of the typical species pool of the colonized community. In contrast, snowbed communities showed pronounced vegetation changes and a clear shift toward dryer conditions and shorter snow cover, evidenced by their colonization by species from surrounding grasslands. Longer growing seasons allow alpine grassland species, which are taller and hence more competitive, to colonize the snowbeds. This study showed that subalpine–alpine plant communities reacted differently to the ongoing climate changes. Lower snow/rain ratio and longer growing seasons seem to have a higher impact than warming, at least on plant communities dependent on long snow cover. Consequently, they are the most vulnerable to climate change and their persistence in the near future is seriously threatened. Subalpine and alpine grasslands are more stable, and, until now, they do not seem to be affected by a warmer climate.     

Spatial patterns of association at local and regional scales in coastal sand dune communities

Questions: Are positive understorey‐dominant associations important in physically severe dune communities and does the strength of positive associations vary with disturbance at the local scale and with stress at the regional scale? Do associational patterns observed at the neighbourhood scale predict diversity at higher scales? Location: Coastal sand dunes, Aquitaine (France). Methods: Associational patterns with five dominant species were recorded along a local gradient of disturbance and a 240‐km long regional gradient. Density, richness, cover and variance ratio of understorey species were recorded in quadrats located in dominant and in open areas. Spatial pattern of dominant plant species was recorded using a distance‐based method. Results: Positive understorey‐dominant associations were most frequent at both regional and local scale, although negative associations with understorey species were observed for one of the five dominants. At the regional scale, there was a shift in the magnitude of spatial associations, with higher positive associations in the most stressful sites, whereas spatial associations where not affected by the local disturbance gradient. Positive associations were not related to the size of the dominants but rather influenced by the identity of the dominant species. Conclusions: Our study highlights the potential crucial role of facilitation together with the importance of turnover of the dominants in explaining large‐scale variation in diversity. However, because positive associations may also be attributed to environmental heterogeneity or co‐occurrence of microhabitat preferences of species, experiments are needed to fully assess the relative importance of facilitation versus other drivers of community diversity.     

Positive effects of an extremely hot summer on propagule rain in upper alpine to subnival habitats of the Central Eastern Alps

Background: Propagule production and dispersal largely determine the distribution and potential migration ability of alpine plant species. Variation in reproductive success caused by year-to-year variation in climate may critically influence these processes.

Aims: To obtain estimates for the propagule rain in high-alpine plant communities and detect potential dispersal events from lower elevations.

Methods: The magnitude and composition of the propagule rain was studied in different plant communities along an elevation gradient from the upper alpine to subnival zone. Propagules were trapped at eight elevations from 2760 to 3070 m a.s.l. for three years from July to September 2003–2005. Vascular plant species and their cover were recorded in an area with a radius of 10 m surrounding the traps.

Results: A five- to 10-fold higher propagule rain was observed in 2003, a year with an exceptionally hot summer, compared to 2004 and 2005. Propagule and species numbers varied highly among years and community types. Few propagules of non-local origin were recorded in any year.

Conclusions: Extremely hot summers are likely to greatly magnify the propagule rain size of species in alpine habitats. Such ‘mast years’ may contribute to enhanced and accelerated vegetation changes in alpine habitats in the absence of limiting factors.     

Shrub invasion into subalpine vegetation: implications for restoration of the native ecosystem

The ability of plant communities to recover after non-native species invasion will depend upon the nature of their soil seed bank and seed rain characteristics. This study assessed changes in the soil seed bank and seed rain associated with the invasion of the non-native shrub Cytisus scoparius in subalpine vegetation. Soil seed bank and seed rain composition, density and richness were investigated at three areas of different stages of invasion: (i) recent (8–10 years), (ii) mature (15–16 years) and (iii) long-term (25 years). There were few changes in seed bank composition or richness regardless of invasion stage. By contrast, the seed rain composition, richness and density was substantially different within long-invaded areas. Very few seeds were able to colonise the dense barrier characteristic of larger, more mature C. scoparius stands. Some prominent herbs from the native vegetation were under-represented or absent from the seed bank, both in invaded and uninvaded areas. Laboratory germination experiments demonstrated that most native species germinate easily, which may imply a transient seed bank, rather than a persistent one. The majority of herbaceous and shrub species were capable of resprouting vegetatively. Therefore, regeneration appeared more reliant on the bud and tuber bank than a persistent soil seed bank. The dominance of graminoid species and C. scoparius rather than other herbaceous, shrub or tree species suggests that the regenerating vegetation will be dominated by grass species and/or C. scoparius. Hence, in areas where long-invaded C.␣scoparius stands are present the recovery of native subalpine vegetation maybe difficult. Recovery may only be possible through wind dispersal from the surrounding intact vegetation or through actively reseeding the area. This study highlights the importance of early intervention in invasive species management.     

What saplings can tell us about forest expansion over natural grasslands

Questions: 1. Do the species composition, richness and diversity of sapling communities vary significantly in differently sized patches? 2. Do forest patches of different sizes differ in woody plant colonization patterns? Location: São Francisco de Paula, Rio Grande do Sul, Brazil, 29°28'S,50°13'W. Methods: Three woody vegetation types, differing in structural development (patch size) and recovering for 10 years from cattle and burning disturbances, were sampled on grassland. We analysed the composition and complexity of the woody sapling communities, through relative abundance, richness and diversity patterns. We also evaluated recruitment status (residents vs. colonizers) of species in communities occurring in different forest patch size classes. Results : 1. There is a compositional gradient in sapling communities strongly associated with forest patch area. 2. Richness and diversity are positively correlated to patch area, but only in poorly structured patches; large patches present richness and diversity values similar to small patches. 3. Resident to colonizer abundance ratio increases from nurse plants to large patches. The species number proportion between residents and colonizers is similar in small and large patches and did not differ between these patch types. 4. Large patches presented a high number of exclusive species, while nurse plants and small patches did not. Conclusions: Woody plant communities in Araucaria forest patches are associated with patch structure development. Richness and diversity patterns are linked to patch colonization patterns. Generalist species colonize the understorey of nurse plants and small patches; resident species cannot recruit many new individuals. In large patches, sapling recruitment by resident adults precludes the immigration of new species into the patches, limiting richness and diversity.     

Contribution of arbuscular mycorrhizal fungi of sedges to soil aggregation along an altitudinal alpine grassland gradient on the Tibetan Plateau

The diversity of arbuscular mycorrhizal fungi (AMF) in sedges on the Tibetan Plateau remains largely unexplored, and their contribution to soil aggregation can be important in understanding the ecological function of AMF in alpine ecosystems. Roots of Kobresia pygmaea C.B. Clarke and Carex pseudofoetida Kük. in alpine Kobresia pastures along an elevational transect (4149–5033 m) on Mount Mila were analysed for AMF diversity. A structural equation model was built to explore the contribution of biotic factors to soil aggregation. Sedges harboured abundant AMF communities covering seven families and some operational taxonomic units are habitat specific. The two plant species hosted similar AMF communities at most altitudes. The relative abundance of the two sedges contributed largely to soil macroaggregates, followed by extraradical mycorrhizal hyphae (EMH) and total glomalin‐related soil protein (T‐GRSP). The influence of plant richness was mainly due to its indirect influence on T‐GRSP and EMH. There was a strong positive correlation between GRSP and soil total carbon and nitrogen. Our results indicate that mycorrhization might not be a major trait leading to niche differentiation of the two co‐occurring sedge species. However, AMF contribute to soil aggregation and thus may have the potential to greatly influence C and N cycling in alpine grasslands.     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

Maintaining diversity through intermediate disturbances: evidence from rodents colonizing rehabilitating coastal dunes

Rodents inhabit the coastal dune forests of KwaZulu‐Natal, South Africa. Here habitat rehabilitation following mining of dunes has resulted in coastal dune forest succession similar to that recorded in nonmined forests. We investigated the colonization of rehabilitating stands and evaluate the role of disturbance in maintaining rodent diversity. A trapping programme was established between July 1993 and February 1995 during which rodent colonization, local extinction and species richness were recorded for rehabilitating stands of different ages. Trends in these variables were closely associated with one of three possible outcomes for a disturbed patch over time, with no intervening disturbances following the initial disturbance. Colonization was initially high which led to an increase in species richness. Extinction was lower than colonization, but became higher when the habitat was 3 years old, which led to a decline in richness. We extrapolate this result assuming negligibly small disturbances after the initiation of rehabilitation and suggest that intermediate levels of disturbance maintain rodent species richness in coastal dune forests. Furthermore, our results illustrated species turnover, a prediction of the recorded outcome, with young stands dominated by Mastomys natalensis and older stands by Saccostomus campestris or Aethomys chrysophilus.     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.