Low vertical transfer rates of carbon inferred from radiocarbon analysis in an Amazon Podzol

Hydromorphic Podzol soils in the Amazon Basin generally support low-stature forests with some of the lowest amounts of aboveground net primary production (NPP) in the region. However, they can also exhibit large values of belowground NPP that can contribute significantly to the total annual inputs of organic matter into the soil. These hydromorphic Podzol soils also exhibit a horizon rich in organic matter at around 1?2m depth, presumably as a result of eluviation of dissolved organic matter and sesquioxides of Fe and Al. Therefore, it is likely that these ecosystems store large quantities of carbon by (1) large amounts of C inputs to soils dominated by their high levels of fine-root production, (2) stabilization of organic matter in an illuviation horizon due to significant vertical transfers of C. To assess these ideas we studied soil carbon dynamics using radiocarbon in two adjacent Amazon forests growing on contrasting soils: a hydromorphic Podzol and a well-drained Alisol supporting a high-stature terra firme forest. Our measurements showed similar concentrations of C and radiocarbon in the litter layer and the first 5 cm of the mineral soil for both sites. This result is consistent with the idea that the hydromorphic Podzol soil has similar soil C storage and cycling rates compared to the well-drained Alisol that supports a more opulent vegetation. However, we found important differences in carbon dynamics and transfers along the vertical profile. At both soils, we found similar radiocarbon concentrations in the subsoil, but the carbon released after incubating soil samples presented radiocarbon concentrations of recent origin in the Alisol, but not in the Podzol. There were no indications of incorporation of C fixed after 1950 in the illuvial horizon of the Podzol. With the aid of a simulation model, we predicted that only a minor fraction (1.7 %) of the labile carbon decomposed in the topsoil is transferred to the subsoil of the Podzol, while this proportional transfer is about 30% in the Alisol. Furthermore, our estimates were 8 times lower than previous estimations of vertical C transfers in Amazon Podzols, and question the validity of these previous estimations for all Podzols within the Amazon Basin. Our results also challenge our previous ideas about the genesis of these particular soils and suggest that either they are not true Podzols or the podzolization processes had already stopped.

Tree height integrated into pantropical forest biomass estimates

Aboveground tropical tree biomass and carbon storage estimates commonly ignore tree height (H). We estimate the effect of incorporating H on tropics-wide forest biomass estimates in 327 plots across four continents using 42 656 H and diameter measurements and harvested trees from 20 sites to answer the following questions: 1. What is the best H-model form and geographic unit to include in biomass models to minimise site-level uncertainty in estimates of destructive biomass? 2. To what extent does including H estimates derived in (1) reduce uncertainty in biomass estimates across all 327 plots? 3. What effect does accounting for H have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of destructively harvested trees when including H (mean 0.06), was half that when excluding H (mean 0.13). Power- andWeibull-H models provided the greatest reduction in uncertainty, with regional Weibull-H models preferred because they reduce uncertainty in smaller-diameter classes (?40 cm D) that store about one-third of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows that including H reduces errors from 41.8Mgha?1 (range 6.6 to 112.4) to 8.0Mgha?1 (?2.5 to 23.0). For all plots, aboveground live biomass was ?52.2 Mgha?1 (?82.0 to ?20.3 bootstrapped 95%CI), or 13%, lower when including H estimates, with the greatest relative reductions in estimated biomass in forests of the Brazilian Shield, east Africa, and Australia, and relatively little change in the Guiana Shield, central Africa and southeast Asia. Appreciably different stand structure was observed among regions across the tropical continents, with some storing significantly more biomass in small diameter stems, which affects selection of the best height models to reduce uncertainty and biomass reductions due to H. After accounting for variation in H, total biomass per hectare is greatest in Australia, the Guiana Shield, Asia, central and east Africa, and lowest in eastcentral Amazonia, W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if tropical forests span 1668 million km2 and store 285 Pg C (estimate including H), then applying our regional relationships implies that carbon storage is overestimated by 35 PgC (31?39 bootstrapped 95%CI) if H is ignored, assuming that the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree H is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of tropical carbon stocks and emissions due to deforestation.

Modelling Pinus pinea forest management to attain natural regeneration under present and future climatic scenarios

Natural regeneration-based silviculture has been increasingly regarded as a reliable option in sustainable forest management. However, successful natural regeneration is not always easy to achieve. Recently, new concerns have arisen because of changing future climate. To date, regeneration models have proved helpful in decision-making concerning natural regeneration. The implementation of such models into optimization routines is a promising approach in providing forest managers with accurate tools for forest planning. In the present study, we present a stochastic multistage regeneration model for Pinus pinea L. managed woodlands in Central Spain, where regeneration has been historically unsuccessful. The model is able to quantify recruitment under different silviculture alternatives and varying climatic scenarios, with further application to optimize management scheduling. The regeneration process in the species showed high between-year variation, with all subprocesses (seed production, dispersal, germination, predation, and seedling survival) having the potential to become bottlenecks. However, model simulations demonstrate that current intensive management is responsible for regeneration failure in the long term. Specifically, stand densities at rotation age are too low to guarantee adequate dispersal, the optimal density of seed-producing trees being around 150 stems·ha−1. In addition, rotation length needs to be extended up to 120 years to benefit from the higher seed production of older trees. Stochastic optimization confirms these results. Regeneration does not appear to worsen under climate change conditions; the species exhibiting resilience worthy of broader consideration in Mediterranean silviculture.

Climatic factors control rodent seed predation in Pinus pinea L. stands in Central Spain

- Context: Pinus pinea L. presents serious problems of natural regeneration in managed forest of Central Spain. The species exhibits specific traits linked to frugivore activity. Therefore, information on plant–animal interactions may be crucial to understand regeneration failure. - Aims: Determining the spatio-temporal pattern of P. pinea seed predation by Apodemus sylvaticus L. and the factors involved. Exploring the importance of A. sylvaticus L. as a disperser of P. pinea. Identifying other frugivores and their seasonal patterns. - Methods: An intensive 24-month seed predation trial was carried out. The probability of seeds escaping predation was modelled through a zero-inflated binomial mixed model. Experiments on seed dispersal by A. sylvaticus were conducted. Cameras were set up to identify other potential frugivores. - Results: Decreasing rodent population in summer and masting enhances seed survival. Seeds were exploited more rapidly nearby parent trees and shelters. A. sylvaticus dispersal activity was found to be scarce. Corvids marginally preyed upon P. pinea seeds. - Conclusions: Survival of P. pinea seeds is climate-controlled through the timing of the dry period together with masting occurrence. Should germination not take place during the survival period, establishment may be limited. A. sylvaticus mediated dispersal does not modify the seed shadow. Seasonality of corvid activity points to a role of corvids in dispersal.

Gestión de la fertilidad de suelos y la nutrición de plantaciones de teca (Tectona grandis L.f.) en América Central

La teca (Tectona grandis L.f.) ha sido tradicionalmente considerada como una madera preciosa en los países del SE Asiático, de donde es originaria, pero durante las últimas décadas ha alcanzado especial relevancia en el sector internacional de las maderas tropicales duras de buena calidad. La especie ha sido ampliamente establecida en América Central, donde tiene una gran importancia socioeconómica, tanto por el impacto de las grandes empresas multinacionales que gestionan grandes plantaciones en la región, como por el gran número de pequeños y medianos propietarios que han elegido esta especie para reforestar sus tierras. Pese a la gran importancia de esta especie, se ha desarrollado relativamente poca investigación acerca de su nutrición y de la gestión del suelo necesaria para su establecimiento y mantenimiento en condiciones sostenibles y productivas. En la presente Tesis Doctoral, tras realizar una amplia revisión bibliográfica, se caracterizan los suelos y la nutrición de las plantaciones de teca en América Central y se proponen varias herramientas para la mejora de su gestión. Las plantaciones de teca de América Central presentan habitualmente deficiencias de K y P, además de algunos problemas de acidez ocasionales. Estos se originan, principalmente, por la mala selección de sitio que se realizó en las últimas dos décadas del siglo XX y por el establecimiento de plantaciones de teca por pequeños propietarios en terrenos que no tienen características propicias para la especie. Además, estos problemas comunes relativos a la baja disponibilidad de P y de K en el suelo son causantes de las relativamente bajas concentraciones foliares de estos elementos (0,88±0,07% K y 0,16±0,04% P) encontradas en plantaciones de teca características de la región. Se presentan varios modelos estadísticos que permiten a los gestores: (a) usarlos como referencia para la interpretación de análisis foliares, ya que ofrecen valores que se consideran característicos de plantaciones de teca con un buen estado nutricional; (b) estimar la cantidad de nutrientes acumulados en la biomasa aérea de sus plantaciones y, sobre todo, su extracción a través de la madera en un aprovechamiento forestal, bien sea una clara o la corta final. La gran acumulación de N, P y K en plantaciones de teca ha de ser considerada como un factor fundamental en su gestión. Además, P y K adquieren mayor relevancia aún ya que su extracción del sistema a través de la madera y su escasa disponibilidad en los suelos hacen que se presente un importante desequilibrio que pone en riesgo la sostenibilidad del sistema. En ese sentido, cambiar la época de cosecha, de la actual (en Enero-Mayo) a Septiembre o Diciembre, puede reducir entre un 24 y un 28% la salida de N asociada a la extracción de madera, un 29% la de P y entre un 14 y un 43% la de K. Se estima que la concentración foliar de P es un factor limitante de la productividad de plantaciones de teca en América Central, proponiéndose un nivel crítico de 0,125%. Además, la teca presenta una tolerancia muy baja a suelos salinos, tendencia que no había sido señalada hasta el momento, siendo muy alta la probabilidad de que la plantación tenga un crecimiento lento o muy lento cuando la Saturación de Na es mayor de 1,1%. Por otro lado, se confirma que K es uno de los elementos clave en la nutrición de las plantaciones de teca en la región centroamericana, proponiéndose un nivel crítico provisional de 3,09% para la Saturación de K, por encima del cual es muy probable que la plantación tenga un crecimiento muy alto. Se ha comprobado que las técnicas estadísticas de análisis multivariante pueden ser usadas como herramientas para agrupar los rodales en base a sus similitudes en cuanto a la fertilidad del suelo y mejorar así el diseño de planes de fertilización en plantaciones con una superficie relativamente grande. De esta manera, se pueden ajustar planes de fertilización más eficientes a escala de grupos de rodales, como un primer paso hacia la selvicultura de precisión, intensificando y diversificando la gestión en función de las diferencias edáficas. Finalmente, aunque los análisis foliares y de suelos indiquen la existencia de deficiencias nutricionales, la fertilización de las plantaciones no siempre va a producir efectos positivos sobre su crecimiento si no se diseña adecuadamente teniendo en cuenta varios factores que pueden estar influyendo negativamente en dicha respuesta, como la densidad de las plantaciones (sinergias con la programación de los clareos y claras) y la elección de la dosis y del producto a aplicar (habitualmente dosis bajas de N-P-K en lugar de incluir otros nutrientes como Mg, B y Zn o usar otros productos como micorrizas, biofertilizantes etc…). ABSTRACT Teak (Tectona grandis L.f.) has been traditionally considered as a precious wood in SE Asia, where it is indigenous. However, during recent decades the species has reached worldwide relevance in the tropical high quality hardwood sector. Teak has been widely established in Central America, where it has become a key species in the forest sector due to its socioeconomic impact, either because of the big-scale plantations of transnational companies and the abundant small-scale plantations established by many farmers. Despite the relevance of the species, little research has been carried out regarding its soil fertility and nutrition management, a key issue both for sustainability and productivity. The present Thesis performs a literature review to this respect, characterize the soil fertility and the nutrition of teak plantations of Central America and propose several management tools. Soil deficiencies of K and P are usually found in teak plantations in Central America, in addition to occasional acidity problems. These problems are mainly derived of (a) a poor site selection performed during 80s and 90s; and (b) small-scale plantations by farmers in sites which are not adequate for the species. These common soil fertility problems related with P and K soil availability are probably the cause of the relatively low P and K foliar concentration (0,88±0,07% K y 0,16±0,04% P) found in representative teak plantations of the region. Several statistical models are proposed, which allow forest managers to: (a) use them as a reference for foliar analysis interpretation, as they show values considered as representative for teak plantations with an adequate nutritional status in the region; (b) estimate the amount of nutrients accumulated in the aerial biomass of the plantations and, especially, the amount of them which are extracted from the systems as wood is harvested in thinning or final clearcuts. The accumulation of N, P and K result in a key factor for teak management in the region. This turns out to be especially relevant for the P and K because their high output rate by timber extraction and the low soil availability result in an important unbalance which constitutes a risk regarding the sustainability of the system. To this respect, modifying the harvesting time from the usual right now (January-May, business as usual scenario) to September or December (proposed alternatives) can reduce between 24 and 28% the N output associated to timber extraction, 29% the P output and between 14 and 43% the K. Foliar P concentration is a main limiting factor for teak plantations productivity in Central America and a 0.125% critical level is proposed. In addition, the results show a very low tolerance for soil salinity, tendency which was not previously reported. Hence, the probability of teak plantations to have low or very low Site Index is high when Na Saturation is higher than 1.1%. On the other hand, K is confirmed as one of the key nutrients regarding teak nutrition in Central America and a 3.09% provisional critical level is proposed for K Saturation; when values are above this level the probability of having very high Site Index is high. Multivariate statistical analyses have been successfully tested to be used as tools to group forest stands according to their soil fertility similarities. Hence, more efficient fertilization plans can be designed for each group of stands, intensifying and diversifying nutritional management according to soil fertility differences. This methodology, which is considered as a first step towards precision forestry, is regarded as helpful tool to design fertilization plans in big scale plantations. Finally, even though foliar and soil analysis would point out some nutritional deficiencies in a forest stand, the results show how the fertilization is not always going to have a positive effect over forest growth if it is not adequately designed. Some factors have been identified as determinants of tree response to fertilization: density (synergisms between fertilization and thinning scheduling) and the appropriate selection of dosages and product (usually low dosages are applied and N-P-K is preferred instead of applying other nutrients such as Mg, B or Zn or using other alternatives such as mycorrhizas or biofertilizers).

Assessing forest response-type diversity to natural disturbances in Peninsular Spain and the Balearic Islands = Evaluación de la diversidad de tipos de respuesta a perturbaciones naturales en la España peninsular e Islas Baleares

Disturbances shape forest ecosystems by influencing their composition, structure, and processes. In the Mediterranean Basin, changes in the disturbance regimes have been predicted to occur in the next future with a higher occurrence of extreme events of drought, wildfire, and – to a lesser extent – windstorm. Woody species are the main elements defining the structure and functioning of forest ecosystems. Recently, response-type diversity has been pointed out as an appropriate indicator of ecosystems resilience. For this, we have elaborated a complete response-trait database for the tree and shrubby species considered in the Third Spanish National Forest Inventory (3SNFI). In the database, the presence or absence of nine response traits associated to drought, fire, and wind were assigned to each species. The database reflected the lack of information about some important traits (in particular for shrubby species) and allowed to determine those traits most widely distributed. The information contained in the database was then used to assess a relative index of forest resilience to these disturbances calculated from the abundance of response traits and the species redundancy for each plot of the 3SNFI; considering both tree and shrubby species. In general, few plots showed high values of the resilience index, probably because some traits were scarcely presented in the species and also because most plots presented very few species. The cartographic representation of the index showed low values for the stands located in mountainous ranges, which are mostly composed by species typical from central Europe. In the other side, Eucalyptus plantations in Galicia appeared as one thee the most resilient ecosystems, due to its higher adaptive capacity to persist after the occurrence of drought, fire, and windstorm events. We conclude that the response traits database can constitute a useful tool for forest management and planning and for future research to enhance the forest resilience.

Edaphic controls on ecosystem-level carbon allocation in two contrasting Amazon forests

Three components of carbon allocation, biomass, flux, and partitioning, were measured in two contrasting Amazon forests growing under similar climatic conditions. Allocation to aboveground compartments was highest in a high-stature forest growing on clay soils, while allocation to fine roots was higher in a short-stature forest growing on white sands. Differences in carbon allocation components where not proportional between the two forests, with soils controlling a trade-off between allocation to fine roots versus aboveground parts.

Natural production of Tuber aestivum in central Spain: Pinus spp. versus Quercus spp. brûles

Aim of study: Tuber aestivum is the most widespread edible truffle, with increasing commercial interest. This species can produce carpophores with conifer hosts, in contrast with the inability of Pinus spp. to induce fruiting in other truffle species such as Tuber melanosporum. Therefore the objective is to compare the characteristics and carpophore production of T. aestivum brûlés associated with Pinus spp. versus Quercus spp. Area of study: We studied the natural habitats of T. aestivum in the Alto Tajo Nature Reserve in central Spain. Material and methods: During 5 years, we monitored the production of carpophores and brûlé size of 145 T. aestivum brûlés associated with Pinus nigra subsp. salzmanni and P. sylvestris and Quercus ilex subsp. ballota and Q. faginea hosts. Statistical treatment was performed using the Statistica Program v. 6. Main results: The size of brûlés associated with Pinus was significantly smaller than that of brûlés associated with Quercus. However, carpophore production per brûlé, and especially for brûlés of similar size, was greater when the host plant was a pine. After accounting for brûlé size, the production of brûlés associated with Pinus spp. was 2.23 (95% CI, between 1.35 and 3.69) and 1.61 (95% CI, between 1.02 and 2.54) times greater than the production of brûlés associated with Quercus faginea and Q. ilex subsp. ballota, respectively. Research highlights: The considerable ability of Pinus nigra subsp. salzmanni and P. sylvestris to form effective brûlés and to produce carpophores of Tuber aestivum in natural conditions was clearly demonstrated, and suggest that those species can be of use in the culture of T. aestivum.

Unravelling past flash flood activity in a forested mountain catchment of the Spanish Central System

Flash floods represent one of the most common natural hazards in mountain catchments, and are frequent in Mediterranean environments. As a result of the widespread lack of reliable data on past events, the understanding of their spatio-temporal occurrence and their climatic triggers remains rather limited. Here, we present a dendrogeomorphic reconstruction of past flash flood activity in the Arroyo de los Puentes stream (Sierra de Guadarrama, Spanish Central System). We analyze a total of 287 increment cores from 178 disturbed Scots pine trees (Pinus sylvestris L.) which yielded indications on 212 growth disturbances related to past flash flood impact. In combination with local archives, meteorological data, annual forest management records and highly-resolved terrestrial data (i.e., LiDAR data and aerial imagery), the dendrogeomorphic time series allowed dating 25 flash floods over the last three centuries, with a major event leaving an intense geomorphic footprint throughout the catchment in 1936. The analysis of meteorological records suggests that the rainfall thresholds of flash floods vary with the seasonality of events. Dated flash floods in the 20th century were primarily related with synoptic troughs owing to the arrival of air masses from north and west on the Iberian Peninsula during negative indices of the North Atlantic Oscillation. The results of this study contribute considerably to a better understanding of hazards related with hydrogeomorphic processes in central Spain in general and in the Sierra de Guadarrama National Park in particular.

Plant β-diversity i in human-altered forest ecosystems: the importance of the structural, spatial, and topographical characteristics of stands in patterning plant species assemblages

An understanding of spatial patterns of plant species diversity and the factors that drive those patterns is critical for the development of appropriate biodiversity management in forest ecosystems. We studied the spatial organization of plants species in human- modified and managed oak forests (primarily, Quercus faginea) in the Central Pre- Pyrenees, Spain. To test whether plant community assemblages varied non-randomly across the spatial scales, we used multiplicative diversity partitioning based on a nested hierarchical design of three increasingly coarser spatial scales (transect, stand, region). To quantify the importance of the structural, spatial, and topographical characteristics of stands in patterning plant species assemblages and identify the determinants of plant diversity patterns, we used canonical ordination. We observed a high contribution of ˟-diversity to total -diversity and found ˟-diversity to be higher and ˞-diversity to be lower than expected by random distributions of individuals at different spatial scales. Results, however, partly depended on the weighting of rare and abundant species. Variables expressing the historical management intensities of the stand such as mean stand age, the abundance of the dominant tree species (Q. faginea), age structure of the stand, and stand size were the main factors that explained the compositional variation in plant communities. The results indicate that (1) the structural, spatial, and topographical characteristics of the forest stands have the greatest effect on diversity patterns, (2) forests in landscapes that have different land use histories are environmentally heterogeneous and, therefore, can experience high levels of compositional differentiation, even at local scales (e.g., within the same stand). Maintaining habitat heterogeneity at multiple spatial scales should be considered in the development of management plans for enhancing plant diversity and related functions in human-altered forests

Evaluating the quality of riparian forest vegetation: The Riparian Forest Evaluation (RFV) index

Aim of study: This paper presents a novel index, the Riparian Forest Evaluation (RFV) index, for assessing the ecological condition of riparian forests. The status of riparian ecosystems has global importance due to the ecological and social benefits and services they provide. The initiation of the European Water Framework Directive (2000/60/CE) requires the assessment of the hydromorphological quality of natural channels. The Directive describes riparian forests as one of the fundamental components that determine the structure of riverine areas. The RFV index was developed to meet the aim of the Directive and to complement the existing methodologies for the evaluation of riparian forests. Area of study: The RFV index was applied to a wide range of streams and rivers (170 water bodies) inSpain. Materials and methods: The calculation of the RFV index is based on the assessment of both the spatial continuity of the forest (in its three core dimensions: longitudinal, transversal and vertical) and the regeneration capacity of the forest, in a sampling area related to the river hydromorphological pattern. This index enables an evaluation of the quality and degree of alteration of riparian forests. In addition, it helps to determine the scenarios that are necessary to improve the status of riparian forests and to develop processes for restoring their structure and composition. Main results: The results were compared with some previous tools for the assessment of riparian vegetation. The RFV index got the highest average scores in the basins of northernSpain, which suffer lower human influence. The forests in central and southern rivers got worse scores. The bigger differences with other tools were found in complex and partially altered streams and rivers. Research highlights: The study showed the index’s applicability under diverse hydromorphological and ecological conditions and the main advantages of its application. The utilization of the index allows a better understanding of the status of riparian forests, and enhances improvements in the conservation and management of riparian areas.

An epidemic of tuberculosis with a high rate of tuberculin anergy among a population previously unexposed to tuberculosis, the Yanomami Indians of the Brazilian Amazon

A survey of an emerging tuberculosis epidemic among the Yanomami Indians of the Amazonian rain forest provided a unique opportunity to study the impact of tuberculosis on a population isolated from contact with the tubercle bacillus for millennia until the mid-1960s. Within the Yanomami population, an extraordinary high prevalence of active tuberculosis (6.4% of 625 individuals clinically examined) was observed, indicating a high susceptibility to disease, even among bacille Calmette–Guérin-vaccinated individuals. Observational studies on cell-mediated and humoral immune responses of the Yanomami Indians compared with contemporary residents of the region suggest profound differences in immunological responsiveness to Mycobacterium tuberculosis infection. Among the Yanomami, a very high prevalence of tuberculin skin test anergy was found. Of patients with active tuberculosis, 46% had purified protein derivative of tuberculosis reactions <10 mm; similarly 58% of recent bacillus Calmette–Guérin vaccines exhibited skin test reactions <5 mm. The Yanomami also had higher titers of antibodies against M. tuberculosis glycolipid antigens (>70%) than the control subjects comprised of Brazilians of European descent (14%). The antibodies were mostly of the IgM isotype. Among the tuberculosis patients who also produced IgG antibodies, the titers of IgG4 were significantly higher among the Yanomami than in the control population. Although it was not possible to analyze T-cell responses or patterns of lymphokine production in vitro because of the remoteness of the villages from laboratory facilities, the results suggest that the first encounter of the Yanomami Indian population with tuberculosis engenders a diminished cell-mediated immune response and an increased production antibody responses, relative to other populations with extensive previous contact with the pathogen. These findings suggest that tuberculosis may represent a powerful selective pressure on human evolution that over centuries has shaped the nature of human immune responses to infection.

Late Holocene human-induced modifications to a central Polynesian island ecosystem.

A 7000-year-long sequence of environmental change during the Holocene has been reconstructed for a central Pacific island (Mangaia, Cook Islands). The research design used geomorphological and palynological methods to reconstruct vegetation history, fire regime, and erosion and depositional rates, whereas archaeological methods were used to determine prehistoric Polynesian land use and resource exploitation. Certain mid-Holocene environmental changes are putatively linked with natural phenomena such as eustatic sea-level rise and periodic El Niño-Southern Oscillation events. However, the most significant changes were initiated between 2500 and 1800 years and were directly or indirectly associated with colonization by seafaring Polynesian peoples. These human-induced effects included major forest clearance, increased erosion of volcanic hillsides and alluvial deposition in valley bottoms, significant increases in charcoal influx, extinctions of endemic terrestrial species, and the introduction of exotic species.

Uppermost Limit, Extent, and Fluctuations of the Timberline and Treeline Ecocline in the Swiss Central Alps during the past 11,500 Years

Pollen and macrofossils were analyzed at two sites above today's treeline (or tree limit) in the Swiss Central Alps (Gouillé Loéré, 2503 m a.s.l., and Lengi Egga, 2557 m a.s.l.) to test two contrasting hypotheses about the natural formation of timberline (the upper limit of closed forest) in the Alps. Our results revealed that Pinus cembra--Larix decidua forests near timberline were rather closed between 9000 and 2500 B.C. (9600-4000 ¹⁴C yr BP), when timberline fluctuations occurred within a belt 100-150 m above today's tree limit. The treeline ecocline above timberline was characterized by the mixed occurrence of tree, shrub, dwarf-shrub, and herbaceous species, but it did not encompass more than 100-150 altitudinal meters. The uppermost limit reached by timberline and treeline during the Holocene was ca. 2420 and 2530 m, respectively, i.e., about 120 to 180 m higher than today. Between 3500 and 2500 B.C. (4700-4000 ¹⁴C yr BP) timberline progressively sank by about 300 m, while treeline was lowered only ca. 100 m. This change led to an enlargement of the treeline-ecocline belt (by ca. 300 m) after 2500 B.C. (4000 ¹⁴C yr BP). Above the treeline ecocline, natural meadows dominated by dwarf shrubs (e.g., Salix herbacea) and herbaceous species (e.g., Helianthemum, Taraxacum, Potentialla, Leontodon t., Cerastium alpinum t., Cirsium spinosissimum, Silene exscapa t., and Saxifraga stellaris) have been present since at least 11,000 cal yr ago. In these meadows tree and tall shrub species (>0.5 m) never played a major role. These results support the conventional hypothesis of a narrow ecocline with rather sharp upper timberline and treeline boundaries and imply that today's treeless alpine communities in the Alps are close to a natural stage. Pollen (percentages and influx), stomata, and charcoal data may be useful for determining whether or not a site was treeless. Nevertheless, a reliable and detailed record of past local vegetation near and above timberline is best achieved through the inclusion of macrofossil analysis.