Species diversity,susceptibility to disturbance and tree population dynamics in tropical rain forest

Abstract. In 1964 a census of all trees > 9.7 cm diameter at breast height (DBH) was conducted on 22 plots totalling 13.2 ha in lowland tropical evergreen rain forest on Kolombangara, Solomon Islands. Over the following 30 yr (1964–1994), populations of all individuals > 4.85 cm DBH of the 12 most common tree species and amounts of disturbance have been monitored on a declining number of these plots (in 1994, nine plots totalling 5.4 ha were still being recorded). Between November 1967 and April 1970, Kolombangara was struck by four cyclones, although only two of these caused substantial amounts of damage to the canopy structure. Multivariate analysis has identified six forest types on Kolombangara (Greig-Smith et al. 1967). The species richness and diversity of trees in the 1964 census, turnover rates of the populations monitored over 1964–1975, and the amount of disturbance sustained during a cyclone in 1970, were all positively correlated across five of the forest types. The sixth forest type was a consistent outlier in these analyses and is believed to have been seriously disturbed by humans about a century ago. The floristics, turnover and disturbance data support Connell's intermediate disturbance hypothesis. The most species-rich forest types contained a higher proportion of fast-growing individuals and species that are early-successional and which have low density timber. Properties of these species rendered them more susceptible to damage when struck by the 1970 cyclone. They showed higher turnover rates because disturbance-dependent species are also characterised by higher mortality and recruitment rates. Thus, periodic cyclones appear to favour the maintenance of differences in species diversity and composition between forest types.     

Tree species richness and turnover throughout New Zealand forests

Abstract. Patterns of mortality, recruitment, and forest turnover were investigated using permanent plot data from temperate forests in 14 localities throughout New Zealand. Tree mortality and recruitment rates were calculated from tagged trees ≥ 10 cm diameter at 1.4 m on individual 400 m² plots, and turnover rates were calculated as the mean of mortality and recruitment rates. Turnover rates (1.4% per year) were very similar to those recorded for tropical forests (i.e. 1.5% per year). As was shown in tropical forests, we also found significant relationships between forest turnover and species richness. In New Zealand forests there was also a decrease in species richness and turnover rates with increasing latitude. Although species richness is well known to decline with latitude, our study provides support for a possible link between seasonality and disturbance with tree turnover and species diversity. While tree mortality and recruitment rates were approximately in balance at some localities, in others there were imbalances between mortality and recruitment rates.     

Density‐dependence in common tree species in a tropical dry forest in Mudumalai,southern India

Abstract. Density‐dependence in tree population dynamics has seldom been examined in dry tropical forests. Using long‐term data from a large permanent plot, this study examined 16 common species in a dry tropical forest in southern India for density‐dependence. Employing quadrat‐based analyses, correlations of mortality, recruitment and population change with tree densities were examined. Mortality in 1–10 cm diameter trees was largely negatively correlated with conspecific density, whereas mortality in > 10 cm diameter trees was positively correlated. Mortality was, however, largely unaffected by the basal area and abundance of heterospecific trees. Recruitment was poor in most species, but in Lagerstroemia microcarpa (Lythraceae), Tectona grandis (Verbenaceae) and Cassia fistula (Fabaceae), species that recruited well, strong negative correlations of recruitment with conspecific basal area and abundance were found. In a few other species that could be tested, recruitment was again negatively correlated with conspecific density. In Lagerstroemia, recruitment was positively correlated with the basal area and abundance of heterospecific trees, but these correlations were non‐significant in other species. Similarly, although the rates of population change were negatively correlated with conspecific density they were positive when dry‐season ground fires occurred in the plot. Thus, the observed positive density‐dependence in large‐tree mortality and the negative density‐dependence in recruitment in many species were such that could potentially regulate tree populations. However, repeated fires influenced density‐dependence in the rates of population change in a way that could promote a few common species in the tree community.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Response of primary and secondary rainforest flowers and fruits to a cyclone,and implications for plant‐servicing bats

The response of primary (PF) and secondary (SF) rainforests to cyclones has broad implications for servicing fauna and the resilience of forest functions. We collected fine‐scale data on the reproductive phenology of plant communities in Fijian PF and SF in 12 monthly surveys before and after Cyclone Tomas (2010). We generated a resource index from the reproductive loads of 2218 trees and 1150 non‐trees (>190 species) and trunk and stem diameter to assess patterns in resource abundance for nectarivores and frugivores (hereafter NF resources). We aimed to determine (i) whether species richness of NF resources differed between forests; (ii) the patterns of resilience of NF resources at community level in both forests after a cyclone; and (iii) the effect of response on NF resources for plant‐servicing bats (Pteropodidae). In 12 months preceding the cyclone, NF resources were greater in PF trees; non‐tree resources fluctuated and were greater in SF. Lower species richness of NF resources in SF indicated that fewer opportunities exist there for exploitation by a diverse fauna. More resources were available for bats in PF. In 12 months following the cyclone, PF flowers and fruits, and SF fruits specifically used by pteropodid bats decreased for trees. Non‐tree resources were especially susceptible to the cyclone. No universal pattern of decline was associated with the cyclone; instead, some NF resources declined and others were resilient or responded rapidly to a post‐cyclone environment. Both PF and SF demonstrated resilience at the community level via increased flower survival (PF) and rapid flower production (SF). Reduced species richness of NF resources in SF will compromise future resilience and response to disturbance, including for threatened pteropodid bat species. These findings are critical for long‐term management of forests, given predicted increases in cyclone frequency and intensity associated with anthropogenic climate change.     

Differential temporal beta‐diversity patterns of native and non‐native arthropod species in a fragmented native forest landscape

An important factor that hinders the management of non‐native species is a general lack of information regarding the biogeography of non‐natives, and, in particular, their rates of turnover. Here, we address this research gap by analysing differences in temporal beta‐diversity (using both pairwise and multiple‐time dissimilarity metrics) between native and non‐native species, using a novel time‐series dataset of arthropods sampled in native forest fragments in the Azores. We use a null model approach to determine whether temporal beta‐diversity was due to deterministic processes or stochastic colonisation and extinction events, and linear modelling selection to assess the factors driving variation in temporal beta‐diversity between plots. In accordance with our predictions, we found that the temporal beta‐diversity was much greater for non‐native species than for native species, and the null model analyses indicated that the turnover of non‐native species was due to stochastic events. No predictor variables were found to explain the turnover of native or non‐native species. We attribute the greater turnover of non‐native species to source‐sink processes and the close proximity of anthropogenic habitats to the fragmented native forest plots sampled in our study. Thus, our findings point to ways in which the study of turnover can be adapted for future applications in habitat island systems. The implications of this for biodiversity conservation and management are significant. The high rate of stochastic turnover of non‐native species indicates that attempts to simply reduce the populations of non‐native species in situ within native habitats may not be successful. A more efficient management strategy would be to interrupt source‐sink dynamics by improving the harsh boundaries between native and adjacent anthropogenic habitats.     

Long‐term tree fern dynamics linked to disturbance and shade tolerance

Question: Do New Zealand tree ferns have recognizable shade tolerance niches? Location: Lowland temperate rain forest of New Zealand (41°20′S, 174°58′E). Methods: Growth, death and recruitment of five tree fern species were estimated from a 38‐year record of stem heights, collected within a 2.25‐ha block of forest, and electron transport rates (ETR) of photosystem II of fronds were measured. Results: Two species of Cyathea were comparatively common (603 and 351 stems in total) and two were comparatively rare (155 and 17 stems in total) on the site. The common species had lower rates of growth, recruitment and mortality than the rare species, had skewed age distributions typical of shade‐tolerant species and were probably recruited soon after a catastrophic earthquake in 1855. The two rare species were failing to recruit under closed forests; their age distributions indicated that all had regenerated long after the earthquake. ETR were higher for faster‐growing than for the shade‐tolerant species. A tree fern that regenerates vegetatively from aerial buds, Dicksonia squarrosa, was common on the site (361 stems in total). Its age distribution suggested it was relatively shade tolerant, but its mortality and recruitment rates were much higher than those of the two shade‐tolerating Cyathea species, suggesting that this multi‐stemmed species functions differently from the monopodial Cyathea species. Conclusions: New Zealand Cyathea tree ferns occupy distinct niches along a shade tolerance spectrum and their relative abundances are strongly influenced by disturbance history. The study provides evidence that tree fern species differ strongly in their responses to canopy disturbance and are not ecologically equivalent.     

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

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

Demography of the long‐lived conifer Agathis ovata in maquis and rainforest,New Caledonia

Abstract. The endemic New Caledonian conifer Agathis ovata occurs as an emergent tree in fire‐prone shrublands (maquis), and fire‐sensitive rainforest. Growth, survivorship and recruitment over 5 yr were compared for populations from forest and maquis on ultramafic substrates in New Caledonia to investigate whether demographic behaviour varied in response to the strongly contrasting forest and shrubland environments. Growth of seedlings and of small (30–100 cm height) and large (100 cm height; 5 cm DBH) saplings was slow, but varied significantly among stages, site types and years. The greatest difference in growth rates was among stages, seedlings growing 0.34 cm.yr^?1, small saplings 1.06 cm.yr^?1 and large saplings 2.13 cm.yr^?1. Tree DBH increased by only 0.05 cm.yr^?1 and, based on these rates, individuals with DBH of 30 cm are estimated to be more than 700 yr old. Few trees (3.5%) produced cones in any year and seedling recruitment was low, but some recruitment was recorded each year in both maquis and forest. Rates of recruitment per parent were highest in forest (1.28.yr^?1, cf 0.78.yr^?1), but the higher density of trees in maquis meant that overall recruitment was greater there (92 ha^?1.yr^?1, cf 56 ha^?1.yr^?1). Seedling mortality ranged from 0.9 to 2.9% among years with no significant difference between maquis and forest. No sapling mortality was recorded, but annual tree mortality ranged from 0 to 1.4%. Evidence from a recently burned site indicated that while trees may survive fire, seedlings and saplings do not. Post‐fire seedling recruitment per ha from surviving trees was four times lower than in unburned sites, but growth rates were four times higher. Similar demographic attributes, including high survivorship, low growth rate and low rates of recruitment over a long reproductive life, characterize Agathis ovata populations in both maquis and rainforest in New Caledonia and are indicative of a broad tolerance of light environments that is unusual among tree species. These demographic attributes help to explain the long‐term persistence of the species in these strongly contrasting habitats.     

Effects of past disturbance and edges on tree community structure and dynamics within a fragment of tropical semideciduous forest in south-eastern Brazil over a five-year period (1987–1992)

Tree community structure and dynamics of a 5.8 ha fragment of montane semideciduous forest in south-eastern Brazil are described based on two successive surveys of trees with dbh 5 cm carried out in 1987 and 1992 in 126 20 × 20 m quadrats. The main purpose was to assess whether the spatial variation of dynamic and structural variables were related to edge-effects and past disturbance regimes. The totals for the two surveys were, respectively, 6528 and 6770 trees, and 94.89 and 108.53 m² of basal area. The forest fragment was at an aggrading sylvigenetic phase, indicated by an overall net increase in density and basal area, and by a declining proportion of trees of smaller size. The overall annual mortality and recruitment rates were 2.6% and 3.0%, respectively. The dynamic process, however, was strongly affected by spatial heterogeneity. The more severely disturbed sectors had lower average canopy heights and basal areas per hectare, and higher tree densities, proportions of smaller trees, and relative area of canopy gaps. They also had higher mortality and recruitment rates, resulting in higher turnover rates compared to the sectors which suffered only moderate disturbance regimes. Edge sectors were also affected by different past disturbance regimes. However, they stood out by having a particularly high frequency of liana tangles and the highest rates of net increase both in density and basal area. A canonical correspondence analysis indicated that the effects of past disturbance regimes and edges were highly related to the species distribution in the area. Pioneer species were concentrated on the edges and more severely disturbed sectors, shade-tolerant species were particularly more abundant on the moderately disturbed sites, while light-demanding ones were more widely distributed. Tree community structure was also undergoing important changes, with less common species enjoying higher recruitment rates and density increase compared to the more common ones. Shade-tolerant species were increasing in relative abundance and had comparatively lower mortality rates and higher recruitment rates. On the other hand, pioneer species were under retreat, with a decreasing relative abundance, higher mortality rates and lower recruitment rates.Nomenclature: Oliveira-Filho et al. 1994a,d.     

Effects of topography on rainforest tree community structure and diversity in American Samoa, and implications for frugivore and nectarivore populations

Aim We investigated the spatial variation of rainforest tree community structure and composition to determine if forest structure and diversity varied as a function of topography; and in turn if this could influence patterns of habitat use by native forest birds and pteropodid bats. Location The study was undertaken on the island of Tutuila, American Samoa, located in the South Pacific Ocean. Methods All trees ≥10 cm diameter were censused in sixty 200 m² plots in ridge, slope and valley forest across the island of Tutuila. Results Forest structure varied significantly across topographical space. Ridge forest was shortest and had the highest stem densities, and valley forest was tallest with the fewest stems per unit area. Species richness was highest on ridges, and slope and valley forest were more similar in composition with each other than they were with ridge forest. Of the fifty-two tree species encountered in the plots, nine showed a statistical affiliation to one of the three topographical positions. Main conclusions We explain patterns of forest structure and diversity in the context of chronic and catastrophic disturbances. Higher stem densities in ridge forest suggested a higher degree of disturbance on ridges, and this was supported by the fact that the height/diameter ratio of the forest was lowest on ridges, which indicated wind-cropping. We hypothesize the potential effects of topographical variation and known phenological patterns on wildlife abundances. We predict that flowering episodes of ridge-affiliated, bird-visited species (particularly Syzygium inophylloides (A. Gray) C. Muell.; Myrtaceae) will concentrate honeyeater densities on ridges, and that fruiting of the tree Canarium vitiense A. Gray (Burseraceae) could localize populations of the Pacific pigeon (Ducula pacifica). Overall (i.e. net) bat foraging patterns are unlikely to be affected by either flowering or fruiting events. Most of the tree species on Tutuila are generalist in their demographic patterns, and the island is depauperate in wildlife fauna; the evolutionary and conservation implications are discussed. We conclude with the argument that conservation of vertebrate species is essential to maintain the current generalist demographic patterns of Samoan trees.     

A climate change‐induced threat to the ecological resilience of a subtropical monsoon evergreen broad‐leaved forest in Southern China

Recent studies have suggested that tropical forests may not be resilient against climate change in the long term, primarily owing to predicted reductions in rainfall and forest productivity, increased tree mortality, and declining forest biomass carbon sinks. These changes will be caused by drought‐induced water stress and ecosystem disturbances. Several recent studies have reported that climate change has increased tree mortality in temperate and boreal forests, or both mortality and recruitment rates in tropical forests. However, no study has yet examined these changes in the subtropical forests that account for the majority of China's forested land. In this study, we describe how the monsoon evergreen broad‐leaved forest has responded to global warming and drought stress using 32 years of data from forest observation plots. Due to an imbalance in mortality and recruitment, and changes in diameter growth rates between larger and smaller trees and among different functional groups, the average DBH of trees and forest biomass have decreased. Sap flow measurements also showed that larger trees were more stressed than smaller trees by the warming and drying environment. As a result, the monsoon evergreen broad‐leaved forest community is undergoing a transition from a forest dominated by a cohort of fewer and larger individuals to a forest dominated by a cohort of more and smaller individuals, with a different species composition, suggesting that subtropical forests are threatened by their lack of resilience against long‐term climate change.     

Edge disturbance drives liana abundance increase and alteration of liana–host tree interactions in tropical forest fragments

Closed‐canopy forests are being rapidly fragmented across much of the tropical world. Determining the impacts of fragmentation on ecological processes enables better forest management and improves species‐conservation outcomes. Lianas are an integral part of tropical forests but can have detrimental and potentially complex interactions with their host trees. These effects can include reduced tree growth and fecundity, elevated tree mortality, alterations in tree‐species composition, degradation of forest succession, and a substantial decline in forest carbon storage. We examined the individual impacts of fragmentation and edge effects (0–100‐m transect from edge to forest interior) on the liana community and liana–host tree interactions in rainforests of the Atherton Tableland in north Queensland, Australia. We compared the liana and tree community, the traits of liana‐infested trees, and determinants of the rates of tree infestation within five forest fragments (23–58 ha in area) and five nearby intact‐forest sites. Fragmented forests experienced considerable disturbance‐induced degradation at their edges, resulting in a significant increase in liana abundance. This effect penetrated to significantly greater depths in forest fragments than in intact forests. The composition of the liana community in terms of climbing guilds was significantly different between fragmented and intact forests, likely because forest edges had more small‐sized trees favoring particular liana guilds which preferentially use these for climbing trellises. Sites that had higher liana abundances also exhibited higher infestation rates of trees, as did sites with the largest lianas. However, large lianas were associated with low‐disturbance forest sites. Our study shows that edge disturbance of forest fragments significantly altered the abundance and community composition of lianas and their ecological relationships with trees, with liana impacts on trees being elevated in fragments relative to intact forests. Consequently, effective control of lianas in forest fragments requires management practices which directly focus on minimizing forest edge disturbance.     

Without management interventions,endemic wet‐sclerophyll forest is transitioning to rainforest in World Heritage listed K’gari (Fraser Island), Australia

Wet‐sclerophyll forests are unique ecosystems that can transition to dry‐sclerophyll forests or to rainforests. Understanding of the dynamics of these forests for conservation is limited. We evaluated the long‐term succession of wet‐sclerophyll forest on World Heritage listed K'gari (Fraser Island)—the world's largest sand island. We recorded the presence and growth of tree species in three 0.4 hectare plots that had been subjected to selective logging, fire, and cyclone disturbance over 65 years, from 1952 to 2017. Irrespective of disturbance regimes, which varied between plots, rainforest trees recruited at much faster rates than the dominant wet‐sclerophyll forest trees, narrowly endemic species Syncarpia hillii and more common Lophostemon confertus. Syncarpia hillii did not recruit at the plot with the least disturbance and recruited only in low numbers at plots with more prominent disturbance regimes in the ≥10 cm at breast height size. Lophostemon confertus recruited at all plots but in much lower numbers than rainforest trees. Only five L. confertus were detected in the smallest size class (<10 cm diameter) in the 2017 survey. Overall, we find evidence that more pronounced disturbance regimes than those that have occurred over the past 65 years may be required to conserve this wet‐sclerophyll forest, as without intervention, transition to rainforest is a likely trajectory. Fire and other management tools should therefore be explored, in collaboration with Indigenous landowners, to ensure conservation of this wet‐sclerophyll forest.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

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.     

Forest regeneration following ungulate removal in a montane Hawaiian wet forest

Most Hawaiian forests lack resiliency following disturbance due to the presence of non‐native and invasive plant and animal species. The montane wet forest within Hakalau Forest National Wildlife Refuge on Hawai'i island has a long history of ungulate disturbance but portions of the refuge were fenced and most ungulates excluded by the early 1990s. We examined patterns of regeneration within two 100 ha study sites in this forest following the removal of ungulates and in the absence of invasive woody tree species to determine, in part, if passive restoration techniques can be successful under these conditions. We characterized growth, mortality, and basal area (BA) changes for approximately 7,100 marked individuals of all native tree species present in two surveys over a 17–18‐year period within two hundred 30 m diameter forest plots. Considerable recruitment within plots of new trees of all species significantly changed size class distributions and erased deficits in small‐sized trees observed during the first survey, particularly for the codominant canopy tree, koa (Acacia koa). Overall, growth of established dominant 'ōhi'a trees (Metrosideros polymorpha) and recruitment of mid‐canopy trees contributed to increases in BA while high levels of mortality for large A. koa trees contributed to decreased BA. This resulted in a slight increase in BA between the two surveys (+1.9%). This study demonstrates that fencing and ungulate removal may have rescued the A. koa population by facilitating the first real pulse in recruitment in over a century, and that passive restoration can be a successful management strategy in this forest.     

Rates of change in tree communities of secondary Neotropical forests following major disturbances

Rates of change in tree communities following major disturbances are determined by a complex set of interactions between local site factors, landscape history and structure, regional species pools and species life histories. Our analysis focuses on vegetation change following abandonment of agricultural fields or pastures, as this is the most extensive form of major disturbance in Neotropical forests. We consider five tree community attributes: stem density, basal area, species density, species richness and species composition. We describe two case studies, in northeastern Costa Rica and Chiapas, Mexico, where both chronosequence and annual tree dynamics studies are being applied. These case studies show that the rates of change in tree communities often deviate from chronosequence trends. With respect to tree species composition, sites of different ages differ more than a single site followed over time through the same age range. Dynamic changes in basal area within stands, on the other hand, generally followed chronosequence trends. Basal area accumulation was more linked with tree growth rates than with net changes in tree density due to recruitment and mortality. Stem turnover rates were poor predictors of species turnover rates, particularly at longer time-intervals. Effects of the surrounding landscape on tree community dynamics within individual plots are poorly understood, but are likely to be important determinants of species accumulation rates and relative abundance patterns.     

Intermediate disturbances are a key driver of long‐term tree demography across old‐growth temperate forests

Disentangling the relative influence of background versus disturbance related mortality on forest demography is crucial for understanding long‐term dynamics and predicting the influence of global change on forests. Quantifying the rates and drivers of tree demography requires direct observations of tree populations over multiple decades, yet such studies are rare in old‐growth forest, particularly in the temperate zone of Europe. We use multi‐decade (1980–2020) monitoring of permanent plots, including observations of mode of mortality and disturbance events, to quantify rates and drivers of tree demography across a network of old‐growth remnants in temperate mountain forests of Slovenia. Annual rates of mortality and recruitment varied markedly among sites and over time; census intervals that captured intermediate severity canopy disturbances caused subtle peaks in annual mortality (e.g., >2%/year), while rates of background mortality in non‐disturbed intervals averaged about 1%/year. Roughly half of the trees died from modes of mortality associated with disturbance (i.e., uprooting or snapped‐alive). Results of a Bayesian multilevel model indicate that beech (Fagus sylvatica) had a higher likelihood of disturbance related mortality compared to fir (Abies alba), which mainly died standing, and there was a notable increase in the odds of disturbance mortality with increasing diameter for all species. Annual recruitment rates were consistently low at sites (<0.5%) that lacked evidence of disturbance, but often exceeded 3% on sites with higher levels of past canopy mortality. Recruitment was dominated by beech on sites with more diffuse background mortality, while the less shade tolerant maple (Acer pseudoplatanus) recruited following known disturbance events. Our study highlights the important role of stand‐scale, partial canopy disturbance for long‐term forest demography. These results suggest that subtle climate‐driven changes in the regime of intermediate severity disturbances could have an important influence on future forest dynamics and warrant attention.     

Large‐scale assessment of regeneration and diversity in Mediterranean planted pine forests along ecological gradients

Aim There is increasing concern regarding sustainable management and restoration of planted forests, particularly in the Mediterranean Basin where pine species have been widely used. The aim of this study was to analyse the environmental and structural characteristics of Mediterranean planted pine forests in relation to natural pine forests. Specifically, we assessed recruitment and woody species richness along climatic, structural and perturbation gradients to aid in developing restoration guidelines. Location Continental Spain. Methods We conducted a multivariate comparison of ecological characteristics in planted and natural stands of main Iberian native pine species (Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus nigra and Pinus sylvestris). We fitted species‐specific statistical models of recruitment and woody species richness and analysed the response of natural and planted stands along ecological gradients. Results Planted pine forests occurred on average on poorer soils and experienced higher anthropic disturbance rates (fire frequency and anthropic mortality) than natural pine forests. Planted pine forests had lower regeneration and diversity levels than natural pine forests, and these differences were more pronounced in mountain pine stands. The largest differences in recruitment – chiefly oak seedling abundance – and species richness between planted and natural stands occurred at low‐medium values of annual precipitation, stand tree density, distance to Quercus forests and fire frequency, whereas differences usually disappeared in the upper part of the gradients. Main conclusions Structural characteristics and patterns of recruitment and species richness differ in pine planted forests compared to natural pine ecosystems in the Mediterranean, especially for mountain pines. However, management options exist that would reduce differences between these forest types, where restoration towards more natural conditions is feasible. To increase recruitment and diversity, vertical and horizontal heterogeneity could be promoted by thinning in high‐density and homogeneous stands, while enrichment planting would be desirable in mesic and medium‐density planted forests.