Earthquake impacts in old‐growth Nothofagus forests in New Zealand

Abstract. Six stands located on different land forms in mixed old‐growth Nothofagus forests in the Matiri Valley (northwest of South Island, New Zealand) were sampled to examine the effects of two recent large earthquakes on tree establishment and tree‐ring growth, and how these varied across land forms. 50 trees were cored in each stand to determine age structure and the cores were cross‐dated to precisely date unusual periods of radial growth. The 1968 earthquake (M = 7.1, epicentre 35 km from the study area) had no discernible impact on the sampled stands. The impact of the 1929 earthquake (M = 7.7, epicentre 20 km from the study area) varied between stands, depending on whether or not they had been damaged by soil or rock movement. In all stands, the age structures showed a pulse of N. fusca establishment following the 1929 earthquake, with this species dominating establishment in large gaps created by landslides. Smaller gaps, created by branch or tree death, were closed by both N. fusca and N. menziesii. The long period of releases (1929–1945) indicates that direct earthquake damage was not the only cause of tree death, and that many trees died subsequently most likely of pathogen attack or a drought in the early 1930s. The impacts of the 1929 earthquake are compared to a storm in 1905 and a drought in 1974–1978 which also affected forests in the region. Our results confirm that earthquakes are an important factor driving forest dynamics in this tectonically active region, and that the diversity of earthquake impacts is a major source of heterogeneity in forest structure and regeneration.     

Advances in spatial,individual‐based modelling of forest dynamics

Many individual‐based models of forest dynamics lack spatial complexity. Although, in certain cases, spatially simple models may not be substantially inferior to spatially complex models, advances in vegetation science indicate potential weaknesses, particularly the lack of consideration of propagule availability in horizontal space, and varying patch (or canopy gap) dimensions. Models with vertical and horizontal spatial complexity can address these issues, but, thus far, evidence that they outperform patch (or gap) models is limited. Comparison of projections from models that differ only in their spatial complexity is needed to address the effects of propagule availability in space, spatial pattern of canopy tree mortality, and spatial resolution.     

Integrating ecological features of species in spatial pattern analysis of a plant community

Objective: This empirical study was designed to explore the role of ecological features of species in the spatial patterning of a grassland community. Location: Banks of the river Rhône in France. Material and Methods: First, we explored the spatial pattern of 29 species recorded in the community using spatial autocorrelation analysis of species cover values. Second, we then explored the relationship between the patterns found and a set of life attributes that characterized the ecological features of species for resource foraging or dispersion. Finally, we explored the spatial relationship of groups of species that shared the same ecological features using cross‐correlation analysis. Results: We found a significant relationship between the spatial pattern and life attributes of the species highlighting three groups of species: (1) species characterized as competitors, reproducing by runner clonal organs and forming large, dense patches; (2) species characterized as competitive‐rud‐erals, dispersing exclusively by seed production and forming small periodic patches; and (3) species classified as CSR, characterized by rosette morphology and short rhizomes as clonal organs without any significant spatial autocorrelation. Spatial segregation was found between group 1 and group 2 up to 14 m; no significant cross‐correlation between groups 1 and 3 between 0 and 3.5 m, and association between groups 2 and 3 up to 14 m. Conclusions: These results helped to understand how species attributes (relative to stature or dispersion abilities); external factors (such as disturbance) and biotic processes (competition) interplay in structuring the plant community under study in space.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

Availability of coarse woody debris in a boreal old‐growth Picea abies forest

Abstract. This study reports temporal (based on cross‐dated dead trees) and spatial patterns of availability of coarse woody debris (CWD) from Picea abies in a Swedish boreal landscape with discrete old‐growth forest patches in a wetland matrix. Data were collected from 29 patches ranging in size from 0.3 to 28 ha. A total of 897 dead trees with a minimum diameter of > 15 cm occurred on the 7.2 ha area analysed. The year of death was established for 50% of these trees. CWD volume ranged from 17 to 65 m³/ha for downed logs and from 0.5 to 13 m³/ha for standing snags. CWD of all decay stages and diameter classes occurred abundantly and the probability of finding logs of all decay stages and sizes was very high at the scale of single hectares. Tree mortality differed among 5 yr periods. However, during the last 50 yr no 5 yr period produced less than 3 logs/ha. Decay rates were highly variable among different logs. Logs with soft wood and some wood pieces lost (decay stage 5) died ca. 34 years ago. This suggests a fairly rapid decay in this northern forest. The data indicate a high and continuous availability of CWD of all types. It is likely, therefore, that selection pressures for efficient dispersal among CWD dependent species may not be very high. Consequently, species with narrow habitat demands and/or low dispersal ability may have evolved and this may contribute to the decrease of certain species in the managed landscape.     

Long‐term dynamics of the long‐lived conifer Libocedrus bidwillii after a volcanic eruption 2000 years ago

Abstract. Question: Following a volcanic eruption of ca. 232 AD, known as the Taupo eruption, the emergent conifer Libocedrus bidwillii expanded on Mt. Hauhungatahi, upwards above the current tree‐line, and downwards into the mixed montane forest. We ask: (1) if current age‐structures at different altitudes support the patterns predicted by the temporal stand replacement model, with cohort senescence and progressively depleting recruitment at ca. 600 year intervals (average cohort age) since the eruption: and (2) if the case history of the population sheds light on the persistence of mixed conifer‐hardwood forests in general. Location: Mt. Hauhungatahi, Tongariro National Park, New Zealand. Methods: The species composition and structure of seven stands covering the altitudinal range of Libocedrus bidwillii, were quantified. Libocedrus trees were cored, and regression equations used to predict ages. Cohorts were identified. Results: Libocedrus densities and basal areas, and the abundance of seedlings and saplings, peaked at different altitudes. At the species’lower limits there has been no recruitment for ca. 550 years, and the angiosperm Weinmannia racemosa has gained dominance. In the tree line and sub‐alpine forest stands, a low level of continuous regeneration has been boosted by periodic cohort recruitment following exogenous disturbances. Conclusions: In the montane zone, the Libocedrus age structure, and its replacement by Weinmannia, are consistent with a model of depleting cohorts separated by ca. 600 years since the Taupo eruption. At higher altitudes more frequent disturbances and reduced competition have allowed Libocedrus persistence. Comparison with other studies suggests long‐term relationships between gymnosperms and angiosperms are mediated by the scale and frequency of disturbance.     

Spatial analysis of structural and tree‐ring related parameters in a timberline forest in the Italian Alps

Abstract. We investigated the variability in spatial pattern of some structural, dendrochronological and dendroclimatological features of a mixed Larix decidua‐Pinus cembra forest at the timberline in the eastern Italian Alps at fine geographical and temporal scales. Forest structure variables such as stem diameter, tree height, age and tree‐ring related parameters (yearly growth index, mean sensitivity, first order autocorrelation and some dendroclimatic variables) have been compared at various scale levels. We observed that most of the variables show positive autocorrelated structures due to both forest dynamics and fine‐scale driving forces, probably related to microrelief. Spatial structure of yearly indexed radial growth appears sensitive to extreme climatic events. Secondary succession after past disturbances drives the forest towards a structure governed by a gap regeneration dynamics that seems to ensure the different requirements of the two main tree species present. Small spatial scale studies of forest structures, especially if integrated to dendro‐ecological data, seem an efficient tool to assess the disturbance regime and species sensitivity to environmental change.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Linking radial growth patterns and moderate‐severity disturbance dynamics in boreal old‐growth forests driven by recurrent insect outbreaks: A tale of opportunities,successes, and failures

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.     

Convergence and divergence in a long‐term old‐field succession: the importance of spatial scale and species abundance

Whether plant communities in a given region converge towards a particular stable state during succession has long been debated, but rarely tested at a sufficiently long time scale. By analysing a 50‐year continuous study of post‐agricultural secondary succession in New Jersey, USA, we show that the extent of community convergence varies with the spatial scale and species abundance classes. At the larger field scale, abundance‐based dissimilarities among communities decreased over time, indicating convergence of dominant species, whereas incidence‐based dissimilarities showed little temporal tend, indicating no sign of convergence. In contrast, plots within each field diverged in both species composition and abundance. Abundance‐based successional rates decreased over time, whereas rare species and herbaceous plants showed little change in temporal turnover rates. Initial abandonment conditions only influenced community structure early in succession. Overall, our findings provide strong evidence for scale and abundance dependence of stochastic and deterministic processes over old‐field succession.     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.

     

A tree‐ring reconstruction of wind disturbances in a forest of the Slovakian Tatra Mountains,Western Carpathians

Question: Have past windstorm events influenced the structure and composition of mountain forests in the Tatra Mountains? Can severe and infrequent wind disturbances lead to dynamic coexistence of two tree species with different ecological requirements? Location: Subalpine mixed spruce‐larch forest at 1200‐1300 m a.s.l. in the Slovakian Tatra Mountains. A forested site affected by catastrophic large‐scale windthrow on 19 November 2004. Methods: Sixty‐seven spruce and 30 larch cross‐sections from the oldest cohorts were collected in a regular pattern in a 100‐ha plot. Tree‐ring series were analysed to reconstruct growth releases associated with past windthrows. A boundary‐line release criterion was applied to detect disturbance year. Spatial patterns of release signals were statistically detected with Mantel's test. We compared reconstructed years of disturbance events with historical records. Results: Releases in both species showed three main pulses. More than 85% showed major or moderate releases in 1865‐1879, 48% in 1915‐1924, and 25% in 1940‐1949. All of these disturbance events affected the whole 100‐ha area. Releases were spatially patterned in the first disturbances, but distributed randomly in the last. Releases co‐occurred in time with enhanced production of compression wood, suggesting disturbances were of wind origin. Reconstructed dates of windthrows were confirmed using historical data on storms. Conclusions: At least three windthrows of major and moderate severity took place in the last 150 years on southern slopes of the Tatra Mountains. This disturbance regime may contribute to coexistence of spruce and larch through differences in vulnerability and response to heavy windstorms.