Long‐term dynamics in a planted conifer forest with spontaneous ingrowth of broad‐leaved trees

Question: What are the age structure and growth trends in a 160‐year old not‐managed Pinus sylvestris plantation with spontaneous development of Quercus robur and can recruitment of Q. robur be related to the radial growth pattern of the P. sylvestris overstorey? Location: Mattemburgh forest reserve, The Netherlands. Methods: Throughout the forest, we sampled 103 oaks and 102 pines with an increment corer. Tree ring widths were measured and cross‐dated to produce mean ring width series. With these data we determined tree ages, investigated growth trends and identified growth releases and suppressions. Results: Q. robur is uneven‐aged: some individuals recruited around 1925, but most reached coring height in the 1940s. The latter recruitment period related to a transition from stressed to released growth of the overstorey pines, growth releases of the oldest Q. robur and occurrence of P. sylvestris regeneration. No further recruitment has taken place since 1950. Conclusions: This study demonstrates that an old pine plantation can develop spontaneously into well‐structured pine forest with an understorey of oak and pine. However, understorey recruitment in these forest types is not a continuous process and in this case a single allogenic canopy disturbance triggered its establishment.     

Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.     

Tree competition and species coexistence in a cool-temperate old-growth forest in southwestern Japan

Abstract. The growth dynamics and mode of competition between adult trees ≥ 4 cm in DBH (stem diameter at breast height 1.3 m) of eight abundant species occupying ca. 90 % of the total basal area were investigated in a 4-ha study plot (200 m × 200 m) of a cool-temperate, old-growth forest on Mount Daisen, southwestern Japan. In the study plot, 30 tree species with individuals ≥ 4.0 cm DBH co-occurred. A bimodal DBH distribution showing upper and lower-canopy layers was found for the most dominant and largest species, Fagus crenata (ca. 78 % of the total basal area), whilst other tree species showed unimodal DBH distributions corresponding mostly to the lower-canopy layer. We developed a model for individual growth incorporating both intra and interspecific competition and the degree of competitive asymmetry. Onesided interspecific competition was detected only from Fagus crenata (upper-canopy species) to Acer japonicum and Acanthopanax sciadophylloides (lower-canopy species) on the scale of the 4-ha study plot. Only Acanthopanax sciadophylloides showed symmetric intraspecific competition. However, a positive (non-competitive) interspecific relationship between adult trees prevailed over a competitive relationship; for example, individual DBH growth rate of Fagus crenata (especially lower-canopy trees) was correlated with the abundance of Acer mono. The positive relationship represented a group of species with similar habitat preference [soil type (mature or immature) caused by landslide disturbance and the presence/absence of Sasa dwarf bamboos in the understorey], where tree densities were not so high as to bring about competition. Competitive interactions between adult trees ≥ 4 cm in DBH occurred only locally between a few specific species and were suggested to be almost irrelevant to the variation in species coexistence on the 4-ha scale of cool-temperate forest. Rather, the coexistence of 30 tree species (species diversity) on this large scale was suggested to be governed by the regeneration pattern of each component species (habitat preference, seedling establishment, sapling competition) with respect to landslide disturbance.     

Density-dependence and co-existence of conifer and broad-leaved trees in a Japanese northern mixed forest

Abstract. The growth and survival of coniferous trees (particularly Abies sachalinensis) and broad-leaved trees (particularly Quercus crispula) were followed over a 15-yr period in a 15.5-ha area in a northern mixed forest in Japan, and the coexistence of the two groups was simulated by a density-dependent projection matrix model. The density-dependent model assumes that the density effect of mother trees due to one-sided competition for light on smaller-sized tree regulates the demographic functions. The mother tree densities of conifers and broad-leaved trees have stronger negative effects on the recruitment and survival of seedlings of their own group than of the other group. These results support the idea of reciprocal replacement for conifer and broad-leaved trees. Simulations using the density-dependent model showed that the two groups will co-exist within a particular range of recruitment rates. However, the density of both groups did not affect the growth rate of any tree, and equilibrium DBH-distributions from density-dependent matrices were quite different from present distributions both for conifers and broad-leaved trees. On the other hand, equilibrium DBH-distributions of conifer and broad-leaved trees from density-independent matrices were quite distinct from each other, reflecting different survivorship curves of the two dominants. These results suggest that density-dependent processes are not so important for shaping population structures in this northern mixed forest.     

Spatial distribution of Eurya japonica in an old-growth evergreen broad-leaved forest,SW Japan

Abstract. The spatial distribution of Eurya japonica, an evergreen understorey species, was studied on a 4-ha permanent plot in an old-growth, temperate, evergreen broad-leaved forest in southwestern Japan. The pattern of spatial dispersion varied with size class, the environmental heterogeneity produced by tree-fall gaps and microtopography and the scale of observation. Seedlings (0.05 to < 0.5 m high) had the most clumped distributions, followed by saplings (0.5 to 1.3 m high) and adults (> 1.3 m). Individuals were more clumped at the smallest grid size, 1.56m × 1.56 m, and became less clumped as grid size increased for all size classes. On grids of less than 6.25 m × 6.25 m, no spatial correlations were found between seedlings and larger size classes; positive spatial correlations were found between saplings and adults at this scale. Abundant seedlings appeared in tree-fall gaps formed by uprooted trees, but seedlings were frequent on ridge tops and upper slopes. Saplings and adults also occurred on upper slopes, but their spatial distribution had little relation to current tree-fall gaps. The spatial distribution of E. japonica individuals was largely attributed to the distribution of tree-fall gaps and the differential responses to microtopographic variation by individuals at different size classes.     

Sapling survival and growth of coniferous and broad‐leaved trees in successional highland habitats in Mexico

Abstract. We assessed survival and growth of transplanted saplings of understory broad‐leaved trees (Oreopanax xalapensis, Rapaneajuergensenii, Rhamnus sharpii and Tern‐stroemia lineata) and canopy conifers (Abies guatemalensis, Pinus ayacahuite and P. pseudostrobus) into Grassland (GR), Mid‐Successional Oak Forest (MS), and Old‐Growth Oak Forest (OF) stands in the central highlands of Chiapas, Mexico. A total of 727 plants were monitored over eight years. The results suggest habitat preferences of the studied species that should be considered for their conservation oriented management. Conifers had highest survival and growth in GR, while broad‐leaved trees survived better under sparse canopy in MS, but grew tallest in full light. Saplings of all species in dense canopy sites in OF were smaller, and the survival of conifers was lower. An integrated response index (IRI) calculated for each species with survival and relative growth rates showed similar trends. Re‐introduction of the endangered A. guatemalensis is possible in open habitats if fire, trampling and grazing are suppressed. Open habitats in current man‐made landscapes in the highlands of Chiapas may limit the establishment of understory trees requiring partial shade. Inclusion of such species in restoration programs has not been usual so far; yet, local and global interest in native species for such programs is increasing. Information on the response of these kind of species in contrasting habitats may help to incorporate them in the management of high‐diversity forests that may follow pine plantations.     

Scale‐dependent effects of neighborhood biodiversity on individual tree productivity in a coniferous and broad‐leaved mixed forest in China

The relationship between biodiversity and productivity has stimulated an increasing body of research over the past decades, and this topic still occupies a central place in ecology. While most studies have focused on biomass production in quadrats or plots, few have investigated the scale‐dependent relationship from an individual plant perspective. We present an analysis of the effects of biodiversity (species diversity and functional diversity) on individual tree growth with a data set of 16,060 growth records from a 30‐ha temperate forest plot using spatially explicit individual tree‐based methods. A significant relationship between species diversity and tree growth was found at the individual tree level in our study. The magnitude and direction of biodiversity effects varies with the spatial scale. We found positive effects of species diversity on tree growth at scales exceeding 9 m. Individual tree growth rates increased when there was a greater diversity of species in the neighborhood of the focal tree, which provides evidence of a niche complementarity effect. At small scales (3–5 m), species diversity had negative effects on tree growth, suggesting that competition is more prevalent than complementarity or facilitation in these close neighborhoods. The results also revealed many confounding factors which influence tree growth, such as elevation and available sun light. We conclude that the use of individual tree‐based methods may lead to a better understanding of the biodiversity‐productivity relationship in forest communities.     

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.     

Litterfall dynamics in a mixed conifer-angiosperm forest in northern New Zealand

Litterfall in a mixed conifer-angiosperm temperate forest in northern New Zealand was traced for 5 years to determine the patterns of litter production and turnover for conifer and angiosperm components of the forest. Basal area and above-ground biomass was shared approximately equally between conifer (mostly Agathis australis; New Zealand kauri) and angiosperm species (plus tree ferns). The five-year mean annual litterfall, excluding macro-litter, was 7.76± 0.39(SEM) t ha^?1 and ranged from 6.77±0.70 t ha^?1 in 1983–4 to 8.79±1.00 t ha^?1 in 1987–8. Mean monthly litterfall showed a strong seasonal pattern with low rates in winter and early spring, increasing to a peak in early autumn. There were major differences in the nature and timing of litterfall between the conifer and angiosperm fractions. Angiosperm leaf litter reached a maximum in early summer, while conifer litterfall showed highest rates for leaves, twigs and cone scales in late summer-autumn. Conifer reproductive structures (strobili and cone scales) contributed from 13 to 21% of total litterfall, a value high relative to other temperate forests. However, conifer leaf turnover was low relative to that for the angiosperms. Size of the microlitter store was 16.16±1.97 t ha^?1 prior to conifer cone fall, and 18.70±2.02 t ha^?1 following it, and conifer litter made up 76–78% of the total litter store. The estimated mean annual decomposition constant, k, was 0.39 overall, 0.33 for conifer leaf litter and 0.71 for angiosperm leaf litter, values which agree well with previously published rates for decomposition in this forest stand. Differences in the costs of biomass production and rates of turnover, as measured by litterfall and decomposition, may help to explain the functional coexistence of conifers and angiosperms in mixed forests.     

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.     

Nuclear and chloroplast microsatellite analysis of Abies sachalinensis regeneration on fallen logs in a subboreal forest in Hokkaido, Japan

Fallen logs are the main regeneration sites for Abies sachalinensis (Sachalin fir) in subboreal forests in Japan. We surveyed the spatial genetic structure of different demographic stages in a 4.29-ha natural population of A. sachalinensis . Genetic structure was significant at short distances throughout all stages in this wind-dispersed conifer. To evaluate the effects of fallen-log-dependent recruitment on demography and spatial genetic structure, we conducted parentage analysis of offspring with highly polymorphic nuclear simple sequence repeat and chloroplast simple sequence repeat markers, and developed a new hierarchical Bayesian model to estimate the effects of mother trees and fallen logs on seed dispersal and offspring recruitment. Combined application of nuclear and chloroplast simple sequence repeat markers allowed us to unambiguously identify mother trees of most offspring (> 90%). Female reproductive success was extremely skewed; a few adults produced most of the offspring on fallen logs. Limited distance of effective dispersal and recruitment was seen in the parentage analysis and modelling estimation. These recruitment characters of A. sachalinensis , with a fallen-log-dependent recruitment process, may result in significant genetic structure in early demographic stages.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Demographic traits of understory trees and population dynamics of aPicea-Abies forest in Taisetsuzan National Park, northern Japan

The size structure transition matrices ofPicea jezoensis, Picea glehnii andAbies sachalinensis of a sub-boreal forest in Hokkaido, northern Japan were constructed based on the demography of each species (Picea jezoensis andPicea glehnii were dealt with together asPicea) during a 4-year period. Two types of matrices, density-independent and density-dependent population dynamics models, were investigated for evaluating the ‘waiting pattern’ betweenPicea spp. andA. sachalinensis. For the density-dependent model, it was assumed that the demographic traits of understory trees, the recruitment rate, the understory mortality rate and the transition probability from the understory to canopy stages, were regulated by the one-sided competitive effect of canopy trees. The observed size structure ofPicea was almost consistent with the stationary size structure obtained in both the density-independent and the density-dependent models, whereas the observed size structure ofA. sachalinensis was not realized in the two models. The effects of both the transition probability from the understory to canopy stages and the recruitment rate on the dynamics of canopy trees were investigated. ForPicea, two parameters—recruitment rate (e _i ) and transition probability from the understory to canopy stages-exponentially affected the dynamics of canopy trees. In contrast, forAbies sachalinensis, the two parameters affected linearly the dynamics of canopy trees. In conclusion, the population dynamics ofPicea andA. sachalinensis was determined by the parameters of the recruitment rate and the transition probability from the understory to canopy stages, relating to waiting patterns of understory trees for future gap formation. InPicea, the demographic parameters of understory trees intensively regulated the dynamics of canopy trees if compared withA. sachalinensis, suggesting that the performance of understory trees plays a key role in the population dynamics ofPicea. This reflects the growth pattern of understory trees in the regeneration of the two species.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.