Tree phenology and carbon dioxide fluxes - use of digital photography for process-based interpretation at the ecosystem scale

Vegetation phenology is an important indicator of climate change and climate variability and it is strongly connected to biospheric–atmospheric gas exchange. We aimed to evaluate the applicability of phenological information derived from digital imagery for the interpretation of CO2 exchange measurements. For the years 2005–2007 we analyzed seasonal phenological development of 2 temperate mixed forests using tower-based imagery from standard RGB cameras. Phenological information was jointly analyzed with gross primary productivity (GPP) derived from net ecosystem exchange data. Automated image analysis provided reliable information on vegetation developmental stages of beech and ash trees covering all seasons. A phenological index derived from image color values was strongly correlated with GPP, with a significant mean time lag of several days for ash trees and several weeks for beech trees in early summer (May to mid-July). Leaf emergence dates for the dominant tree species partly explained temporal behaviour of spring GPP but were also masked by local meteorological conditions. We conclude that digital cameras at flux measurement sites not only provide an objective measure of the physiological state of a forest canopy at high temporal and spatial resolutions, but also complement CO2 and water exchange measurements, improving our knowledge of ecosystem processes.

Richness of Lichen Species, Especially of Threatened Ones, Is Promoted by Management Methods Furthering Stand Continuity

Lichens are a key component of forest biodiversity. However, a comprehensive study analyzing lichen species richness in relation to several management types, extending over different regions and forest stages and including information on site conditions is missing for temperate European forests. In three German regions (Schwäbische Alb, Hainich-Dün, Schorfheide-Chorin), the so-called Biodiversity Exploratories, we studied lichen species richness in 631 forest plots of 400 m2 comprising different management types (unmanaged, selection cutting, deciduous and coniferous age-class forests resulting from clear cutting or shelterwood logging), various stand ages, and site conditions, typical for large parts of temperate Europe. We analyzed how lichen species richness responds to management and habitat variables (standing biomass, cover of deadwood, cover of rocks). We found strong regional differences with highest lichen species richness in the Schwäbische Alb, probably driven by regional differences in former air pollution, and in precipitation and habitat variables. Overall, unmanaged forests harbored 22% more threatened lichen species than managed age-class forests. In general, total, corticolous, and threatened lichen species richness did not differ among management types of deciduous forests. However, in the Schwäbische-Alb region, deciduous forests had 61% more lichen species than coniferous forests and they had 279% more threatened and 76% more corticolous lichen species. Old deciduous age classes were richer in corticolous lichen species than young ones, while old coniferous age-classes were poorer than young ones. Overall, our findings highlight the importance of stand continuity for conservation. To increase total and threatened lichen species richness we suggest (1) conserving unmanaged forests, (2) promoting silvicultural methods assuring stand continuity, (3) conserving old trees in managed forests, (4) promoting stands of native deciduous tree species instead of coniferous plantations, and (5) increasing the amount of deadwood in forests.

Rolling stones and tree rings: A state of research on dendrogeomorphic reconstructions of rockfall

This progress report focuses on the contribution of tree-ring series to rockfall research and on recent development and challenges in the field. Dendrogeomorphic techniques have been used extensively since the early 2000s and several approaches have been developed to extract rockfall signals from tree-ring records of conifer trees. The reconstruction of rockfall chronologies has been hampered in the past by sample sizes that decrease as one goes back in time, as well as by a paucity of studies that include broadleaved tree species, which are in fact quite common in rockfall-prone environments. In this report, we propose a new approach considering impact probability and quantification of uncertainty in the reconstruction of rockfall time series as well as a quantitative estimate of presumably missed events. In addition, we outline new approaches and future perspectives for the inclusion of woody vegetation in hazard assessment procedures, and end with future thematic perspectives.

Site- and species-specific responses of forest growth to climate across the European continent

Aim To evaluate the climate sensitivity of model-based forest productivity estimates using a continental-scale tree-ring network. Location Europe and North Africa (30–70° N, 10° W–40° E). Methods We compiled close to 1000 annually resolved records of radial tree growth for all major European tree species and quantified changes in growth as a function of historical climatic variation. Sites were grouped using a neural network clustering technique to isolate spatiotemporal and species-specific climate response patterns. The resulting empirical climate sensitivities were compared with the sensitivities of net primary production (NPP) estimates derived from the ORCHIDEE-FM and LPJ-wsl dynamic global vegetation models (DGVMs). Results We found coherent biogeographic patterns in climate response that depend upon (1) phylogenetic controls and (2) ambient environmental conditions delineated by latitudinal/elevational location. Temperature controls dominate forest productivity in high-elevation and high-latitude areas whereas moisture sensitive sites are widespread at low elevation in central and southern Europe. DGVM simulations broadly reproduce the empirical patterns, but show less temperature sensitivity in the boreal zone and stronger precipitation sensitivity towards the mid-latitudes. Main conclusions Large-scale forest productivity is driven by monthly to seasonal climate controls, but our results emphasize species-specific growth patterns under comparable environmental conditions. Furthermore, we demonstrate that carry-over effects from the previous growing season can significantly influence tree growth, particularly in areas with harsh climatic conditions – an element not considered in most current-state DGVMs. Model–data discrepancies suggest that the simulated climate sensitivity of NPP will need refinement before carbon-cycle climate feedbacks can be accurately quantified.

Carbon stocks, tree diversity, and the role of organic certification in different cocoa production systems in Alto Beni, Bolivia

This study compares aboveground and belowground carbon stocks and tree diversity in different cocoa cultivation systems in Bolivia: monoculture, simple agroforestry, and successional agroforestry, as well as fallow as a control. Since diversified, agroforestry-based cultivation systems are often considered important for sustainable development, we also evaluated the links between carbon stocks and tree diversity, as well as the role of organic certification in transitioning from monoculture to agroforestry. Biomass, tree diversity, and soil physiochemical parameters were sampled in 15 plots measuring 48 × 48 m. Semi-structured interviews with 52 cocoa farmers were used to evaluate the role of organic certification and farmers’ organizations (e.g., cocoa cooperatives) in promoting tree diversity. Total carbon stocks in simple agroforestry systems (128.4 ± 20 Mg ha−1) were similar to those on fallow plots (125.2 ± 10 Mg ha−1). Successional agroforestry systems had the highest carbon stocks (143.7 ± 5.3 Mg ha−1). Monocultures stored significantly less carbon than all other systems (86.3 ± 4.0 Mg ha−1, posterior probability P(Diff > 0) of 0.000–0.006). Among shade tree species, Schizolobium amazonicum, Centrolobium ochroxylum, and Anadenanthera sp. accumulated the most biomass. High-value timber species (S. amazonicum, C. ochroxylum, Amburana cearensis, and Swietenia macrophylla) accounted for 22.0 % of shade tree biomass. The Shannon index and tree species richness were highest in successional agroforestry systems. Cocoa plots on certified organic farms displayed significantly higher tree species richness than plots on non-certified farms. Thus, expanding the coverage of organic farmers’ organizations may be an effective strategy for fostering transitions from monoculture to agroforestry systems.

Modeling tree water deficit from microclimate: an approach to quantifying drought stress

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaW(E)), with soil water potential (Psi(soil)) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth, We followed changes in DeltaW and DeltaW(E) in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaW(E). This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psi(soil) or VPD alone provided poorer estimates of AWthan a model incorporating both factors. As a first approximation of DeltaW(E), Psi(soil) can be weighted so that the negative mean Psi(soil) reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: similar to65%, P abies: similar to70%, P sylvestris: similar to75%). The differences in DeltaW among species can be partially explained by a different weighting of Psi(soil) against VPD. The DeltaW of P. sylvestris was more dependent on Psi(soil) than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.

Aluminum toxicity to tropical montane forest tree seedlings in southern Ecuador: response of biomass and plant morphology to elevated Al concentrations

In acid tropical forest soils (pH < 5.5) increased mobility of aluminum might limit aboveground productivity. Therefore, we evaluated Al phytotoxicity of three native tree species of tropical montane forests in southern Ecuador. An hydroponic dose-response experiment was conducted. Seedlings of Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson were treated with 0, 300, 600, 1200, and 2400 mu M Al and an organic layer leachate. Dose-response curves were generated for root and shoot morphologic properties to determine effective concentrations (EC). Shoot biomass and healthy leaf area decreased by 44 % to 83 % at 2400 mu M Al, root biomass did not respond (C. odorata), declined by 51 % (H. americanus), or was stimulated at low Al concentrations of 300 mu M (T. chrysantha). EC10 (i.e. reduction by 10 %) values of Al for total biomass were 315 mu M (C. odorata), 219 mu M (H. americanus), and 368 mu M (T. chrysantha). Helicarpus americanus, a fast growing pioneer tree species, was most sensitive to Al toxicity. Negative effects were strongest if plants grew in organic layer leachate, indicating limitation of plant growth by nutrient scarcity rather than Al toxicity. Al toxicity occurred at Al concentrations far above those in native organic layer leachate.

Density-dependent dynamics of a dominant rain forest tree change with juvenile stage and time of masting

Although negative density dependence (NDD) can facilitate tree species coexistence in forests, the underlying mechanisms can differ, and rarely are the dynamics of seedlings and saplings studied together. Herein we present and discuss a novel mechanism based on our investigation of NDD predictions for the large, grove-forming ectomycorrhizal mast fruiting tree, Microberlinia bisulcata (Caesalpiniaceae), in an 82.5-ha plot at Korup, Cameroon. We tested whether juvenile density, size, growth and survival decreases with increasing conspecific adult basal area for 3245 ‘new’ seedlings and 540 ‘old’ seedlings (< 75-cm tall) during an approximately 4-year study period (2008–2012) and for 234 ‘saplings’ (≥ 75-cm tall) during an approximately 6-year study period (2008–2014). We found that the respective densities of new seedlings, old seedlings and saplings were positively, not and negatively related to increasing BA. Maximum leaf numbers and heights of old seedlings were negatively correlated with increasing basal areas, as were sapling heights and stem diameters. Whereas survivorship of new seedlings decreased by more than one-half with increasing basal area over its range in 2010–2012, that of old seedlings decreased by almost two-thirds, but only in 2008–2010, and was generally unrelated to conspecific seedling density. In 2010–2012 relative growth rates in new seedlings’ heights decreased with increasing basal area, as well as with increasing seedling density, together with increasing leaf numbers, whereas old seedlings’ growth was unrelated to either conspecific density or basal area. Saplings of below-average height had reduced survivorship with increasing basal area (probability decreasing from approx. 0.4 to 0.05 over the basal area range tested), but only sapling growth in terms of leaf numbers decreased with increasing basal area. These static and dynamic results indicate that NDD is operating within this system, possibly stabilizing the M. bisulcata population. However, these NDD patterns are unlikely to be caused by symmetric competition or by consumers. Instead, an alternative mechanism for conspecific adult–juvenile negative feedback is proposed, one which involves the interaction between tree phenology and ectomycorrhizal linkages.

Tree architecture in a Bornean lowland rain forest: intraspecific and interspecific patterns

Intraspecific and interspecific architectural patterns were studied for eight tree species of a Bornean rain forest. Trees 5--19 m tall in two 4-ha permanent sample plots in primary forest were selected, and three light descriptors and seven architectural traits for each tree were measured. Two general predictions were made: (1) Slow growing individuals (or short ones) encounter lower light, and have flatter crowns, fewer leaf layers, and thinner stems, than do fast growing individuals (or tall ones). (2) Species with higher shade-tolerance receive less light and have flatter crowns, fewer leaf layers, and thinner stems, than do species with lower shade-tolerance. Shade-tolerance is assumed to decrease with maximum growth rate, mortality rate, and adult stature of a species.

(Table 2) Survivorship of transplanted seedlings of Picea glauca inside and outside three tree islands in 1994, 1995, 2009 and after 15 years

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Juvenile–adult tree associations in a continental Mediterranean ecosystem: no evidence for sustained and general facilitation at increased aridity

Question: How do tree species identity, microhabitat and water availability affect inter- and intra-specific interactions between juvenile and adult woody plants? Location: Continental Mediterranean forests in Alto Tajo Natural Park, Guadalajara, Spain. Methods: A total of 2066 juveniles and adults of four co-occurring tree species were mapped in 17 plots. The frequency of juveniles at different microhabitats and water availability levels was analysed using log-linear models. We used nearest-neighbour contingency table analysis of spatial segregation and J-functions to describe the spatial patterns. Results: We found a complex spatial pattern that varied according to species identity and microhabitat. Recruitment was more frequent in gaps for Quercus ilex, while the other three species recruited preferentially under shrubs or trees depending on the water availability level. Juveniles were not spatially associated to conspecific adults, experiencing segregation from them inmany cases. Spatial associations, both positive and negative, were more common at higher water availability levels. Conclusions: Our results do not agree with expectations from the stressgradient hypothesis, suggesting that positive interactions do not increase in importance with increasing aridity in the study ecosystem. Regeneration patterns are species-specific and depend on microhabitat characteristics and dispersal strategies. In general, juveniles do not look for conspecific adult protection. This work contributes to the understanding of species co-existence, proving the importance of considering a multispecies approach at several plots to overcome limitations of simple pair-wise comparisons in a limited number of sites.

Forest connectivity restoration through reforestation: an integrated methodology for prioritizing agricultural lands and selecting reforestation species

Forest connectivity restoration is a major goal in natural resource planning. Given the high amount of abandoned cultivated lands, setting efficient methods for the reforestation of agricultural lands offers a good opportunity to face this issue. However, reforestations must be carefully planned, which poses two main challenges. In first place, to determine those agricultural lands that, once reforested, would meet more effectively the planning goals. As a further step, in order to grant the success of the activity, it is fairly advisable to select those tree species that are more adapted to each particular environment. Here we intend to give response to both requirements by proposing a sequential and integrated methodology that has been implemented in two Spanish forest districts, which are formed by several landscape types that were previously defined and characterized. Using the software Conefor Sensinode, a powerful tool for quantifying habitat availability that is based on graph theory concepts, we determined the landscapes where forest planning should have connectivity as a major concern and, afterwards, we detected the agricultural patches that would contribute most to enhance connectivity if they were reforested. The subsequent reforestation species assessment was performed within these priority patches. Using penalized logistic regressions we fitted ecological niche models for the Spanish native tree species. The models were trained with species distribution data from the Spanish Forest Map and used climatic and lithological variables as predictors. Model predictions were used to build ordered lists of suitable species for each priority patch. The lists include dominant and non dominant tree species and allow adding biodiversity goals to the reforestation planning. The result of this combined methodology is a map of agricultural patches that would contribute most to uphold forest connectivity if they were reforested and a list of suitable tree species for each patch ordered by occurrence probability. Therefore the proposed methodology may be useful for suitable and efficient forest planning and landscape designing.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

On the spatial distribution of woody plant species in a tropical montane cloud forest

Abstract The cloud forest is a special type of forest ecosystem that depends on suitable conditions of humidity and temperature to exist; hence, it is a very fragile ecosystem. The cloud forest is also one of the richest ecosystems in terms of species diversity and rate of endemism. However, today, it is one of the most threatened ecosystems in the world. Little is known about tree species distribution and coexistence among cloud forest trees. Trees are essential to understanding ecosystem functioning and maintenance because they support the ecosystem in important ways. For this dissertation, an analysis of woody plant species distribution at a small scale in a north-Peruvian Andean cloud forest was performed, and some of the factors implicated in the observed patterns were identified. Towards that end, different natural factors acting on species distribution within the forest were investigated: (i) intra-specific arrangements, (ii) heterospecific spatial relationships and (iii) relationships with external environmental factors. These analyses were conducted first on standing woody plants and then on seedlings. The woody plants were found to be clumped in the forest, either considering all the species together or each species separately. However, each species presented a specific pattern and specific spatial relationship among different-age individuals. Dispersal mode, growth form and shade tolerance played roles in the final distribution of the species. Furthermore, spatial associations among species, either positive or negative, were observed. These associations were more numerous when considering individuals of the interacting species at different developmental stages, i.e., younger individuals from one species and older individuals from another. Accordingly, competition and facilitation are asymmetric processes and vary throughout the life of an individual. Moreover, some species appear to prefer certain habitat conditions and avoid other habitats. The habitat definition that best explains species distribution is that which includes both environmental and stand characteristics; thus, a combination of these factors is necessary to understanding species' niche preferences. Seedling distribution was also associated with habitat conditions, but these conditions explained less than the 30% of the spatial variation. The position of conspecific adult individuals also affected seedling distribution; although the seedlings of many tree species avoid the vicinity of conspecifics, a few species appeared to prefer the formation of cohorts around their parent trees. The importance of habitat conditions and distance dependence with conspecifics varied among regions within the forest as well as on the developmental stage of the stand. The results from this thesis suggest that different species can coexist within a given space, forming a “puzzle” of species as a result of the intra- and interspecific spatial relationships along with niche preferences and adaptations that operate at different scales. These factors not only affect each species in a different way, but specific preferences also vary throughout species' lifespans. Resumen Resumen El bosque de niebla es uno de los ecosistemas más amenazados del mundo además de ser uno de los más frágiles. Son formaciones azonales que dependen de la existencia de unas condiciones de humedad y temperatura que permitan la formación de nubes que cubran el bosque; lo que dificulta en gran medida su conservación. También es uno de los ecosistemas con mayor riqueza de especies además de tener uno de los mayores porcentajes de endemismos. Uno de los aspectos más importantes para entender el ecosistema, es identificar y entender los elementos que lo componen y los mecanismos que regulan las relaciones entre ellos. Los árboles son el soporte del ecosistema. Sin embargo, apenas hay información sobre la distribución y coexistencia de los árboles en los bosques de niebla. Esta tesis presenta un análisis de la distribución a pequeña escala de las plantas leñosas en un bosque de niebla situado en la cordillera andina del norte de Perú; así como el análisis de algunos de los factores que pueden estar implicados en que se origine la distribución observada. Para este propósito se estudia cómo influyen factores de diferente naturaleza en la distribución de las especies (i) organización intra-específica (ii) relaciones espaciales heterospecíficas y (iii) relación con factores ambientales externos. En estos análisis se estudiaron primero las plantas jóvenes y las adultas, y después las plántulas. Los árboles aparecieron agregados en el bosque, tanto considerando todos a la vez como cuando se estudió cada especie por separado. Sin embargo, cada especie mostró un patrón distinto así como una particular relación espacial entre individuos jóvenes y adultos. El modo de dispersión, la forma de vida y la tolerancia de la especies estuvieron relacionados con el patrón general observado. Se vio también que ciertas especies aparecían relacionadas con otras, tanto de forma positiva (compartiendo zonas) como negativa (apareciendo en áreas distintas). Las asociaciones fueron mucho más numerosas cuando se consideraron los pares de especies en diferente estado de desarrollo, es decir, individuos jóvenes de una especie e individuos mayores de la otra. Eso indicaría que los procesos de competencia y facilitación son asimétricos y además varían durante la vida de la planta. Por otro lado, algunas especies aparecen preferentemente bajo ciertas condiciones de hábitat y evitan otras. La definición de hábitat a la que mejor responden las especies es cuando se incluyen tanto variables ambientales como de masa; así que ambos tipos de variables son necesarias para entender la preferencia de las especies por ciertos nichos. La distribución de las plántulas también estuvo relacionada con condiciones de hábitat, pero eso sólo llegaba a explicar hasta un 30% de la variabilidad espacial. La posición de los adultos de la misma especie también afectó a la distribución de las plántulas. En bastantes especies las plántulas evitan la cercanía de adultos de su misma especie, padres potenciales, aunque algunas especies aisladas mostraron el patrón contrario y aparecieron preferentemente en las mismas áreas que sus padres. La importancia de las condiciones de hábitat y posición de los adultos en la disposición de las plántulas varía de una zona a otra del bosque y además también varía según el estado de desarrollo de la masa.