Rapid responses of high-mountain vegetation to early Holocene environmental changes in the Swiss Alps

1 The Early Holocene sediment of a lake at tree line (Gouillé Rion, 2343 m a.s.l.) in the Swiss Central Alps was sampled for plant macrofossils. Thin (0.5 cm) slices, representing time intervals of c. 50 years each from 11 800 to 7800 cal. year bp, were analysed and the data compared with independent palaeoclimatic proxies to study vegetational responses to environmental change. 2 Alpine plant communities (e.g. with Salix herbacea) were established at 11 600–11 500 cal. year bp, when oxygen-isotope records showed that temperatures increased by c. 3–4 °C within decades. Larix decidua trees reached the site at c. 11 350 cal. year bp, probably in response to further warming by 1–2 °C. Forests dominated by L. decidua persisted until 9600 cal. year bp, when Pinus cembra became more important. 3 The dominance of Larix decidua for two millennia is explained by dry summer conditions, and possibly low winter temperatures, which favoured it over the late-successional Pinus cembra. Environmental conditions were a result of variations in the earth's orbit, leading to a maximum of summer and a minimum of winter solar radiation. Other heliophilous and drought-adapted species, such as Dryas octopetala and Juniperus nana, could persist in the open L. decidua forests, but were out-competed when the shade-tolerant P. cembra expanded. 4 The relative importance of Larix decidua decreased during periods of diminished solar radiation at 11 100, 10 100 and 9400 cal. year bp. Stable concentrations of L. decidua indicate that these percentage oscillations were caused by temporary increases of Pinus cembra, Dryas octopetala and Juniperus nana that can be explained by increases in moisture and/or decreases in summer temperature. 5 The final collapse of Larix decidua at 8400 cal. year bp was possibly related to abrupt climatic cooling as a consequence of a large meltwater input to the North Atlantic. Similarly, the temporary exclusion of Pinus cembra from tree line at 10 600–10 200 cal. year bp may be related to slowing down of thermohaline circulation at 10 700–10 300 cal. year bp. 6 Our results show that tree line vegetation was in dynamic equilibrium with climate, even during periods of extraordinarily rapid climatic change. They also imply that forecasted global warming may trigger rapid upslope movements of the tree line of up to 800 m within a few decades or centuries at most, probably inducing large-scale displacements of plant species as well as irrecoverable biodiversity losses.

Biotic and abiotic controls of nitrogen and phosphorus cycling in Central European forests

The functioning and services of Central European forests are threatened by global change and a loss of biodiversity. Nutrient cycling as a key forest function is affected by biotic drivers (e.g., dominant tree species, understory plants, soil organisms) that interact with abiotic conditions (e.g., climate, soil properties). In contrast to grassland ecosystems, evidence for the relationship of nutrient cycles and biodiversity in forests is scarce because the structural complexity of forests limits experimental control of driving factors. Alternatively, observational studies along gradients in abiotic conditions and biotic properties may elucidate the role of biodiversity for forest nutrient cycles. This thesis aims to improve the understanding of the functional importance of biodiversity for nutrient cycles in forests by analyzing water-bound fluxes of nitrogen (N) and phosphorus (P) along gradients in biodiversity in three regions of Germany. The tested hypotheses included: (1) temperate forest canopies retain atmospheric N and retention increases with increasing plant diversity, (2) N release from organic layers increases with resource availability and population size of decomposers but N leaching decreases along a gradient in plant diversity, (3) P leaching from forest canopies increases with improved P supply from recalcitrant P fractions by a more diverse ectomycorrhizal fungal community. In the canopies of 27 forest stands from three regions, 16 % to 51 % of atmospheric N inputs were retained. Regional differences in N retention likely resulted from different in N availability in the soil. Canopy N retention was greater in coniferous than in beech forests, but this was not the case on loessderived soils. Nitrogen retention increased with increasing tree and shrub diversity which suggested complementary aboveground N uptake. The strength of the diversity effect on canopy N uptake differed among regions and between coniferous and deciduous forests. The N processing in the canopy directly coupled back to N leaching from organic layers in beech forests because throughfall-derived N flushed almost completely through the mull-type organic layers at the 12 studied beech sites. The N release from organic layers increased with stand basal area but was rather low (< 10 % of annual aboveground litterfall) because of a potentially high microbial N immobilization and intensive incorporation of litter into the mineral soil by bioturbation. Soil fauna biomass stimulated N mineralization through trophic interactions with primary producers and soil microorganisms. Both gross and net leaching from organic layers decreased with increasing plant diversity. Especially the diversity but not the cover of herbs increased N uptake. In contrast to N, P was leached from the canopy. Throughfall-derived P was also flushed quickly through the mull-type organic layers and leached P was predominantly immobilized in non directly plant-available P fractions in the mineral soil. Concentrations of plant-available phosphate in mineral soil solution were low and P leaching from the canopy increased with increasing concentrations of the moderately labile P fraction in soil and increasing ectomycorrhiza diversity while leaf C:P ratios decreased. This suggested that tree P supply benefited from complementary mining of diverse mycorrhizal communities for recalcitrant P. Canopy P leaching increased in years with pronounced spring drought which could lead to a deterioration of P supply by an increasing frequency of drought events. This thesis showed that N and P cycling in Central European forests is controlled by a complex interplay of abiotic site conditions with biological processes mediated by various groups of organisms, and that diverse plant communities contribute to tightening the N cycle in Central European forests and that diverse mycorrhizal communities improve the limited P availability. Maintaining forest biodiversity seems essential to ensure forest services in the light of environmental change.

Holocene palaeoclimate and palaeovegetation on the islands of Flores and Pico

Palaeoclimatic stability is regarded as an important factor in explaining patterns of endemism in the Azorean flora. However, modelling simulations and quantitative reconstructions for the last 6000 years suggest considerable palaeoclimatic variability. Here we explore the link between Holocene palaeoclimate and palaeovegetation on the islands of Flores and Pico. Modern pollen assemblages indicate that most major plant communities can be detected using pollen analysis and that, in some cases, the pre-colonisation vegetation was quite similar to present-day relict vegetation. A 200–500-year pollen record from Alagoinha, a low-elevation mire in western Flores, shows that Juniperus brevifolia-dominated communities were widespread at lower elevations prior to large-scale deforestation. Today these communities are generally restricted to higher elevations. While our results are preliminary, there appears to be a weak link between palaeovegetation (which was primarily influenced by volcanism, soil formation and human impact) and palaeoclimatic changes detected through geochemical proxies. Even if the Azorean palaeoclimate varied substantially, its impact on the pristine vegetation, at least in terms of pollen production, was relatively small.

Impact of above- and below-ground invertebrates on temporal and spatial stability of grassland of different diversity

1. Recent theoretical studies suggest that the stability of ecosystem processes is not governed by diversity per se, but by multitrophic interactions in complex communities. However, experimental evidence supporting this assumption is scarce.2. We investigated the impact of plant diversity and the presence of above- and below-ground invertebrates on the stability of plant community productivity in space and time, as well as the interrelationship between both stability measures in experimental grassland communities.3. We sampled above-ground plant biomass on subplots with manipulated above- and below-ground invertebrate densities of a grassland biodiversity experiment (Jena Experiment) 1, 4 and 6 years after the establishment of the treatments to investigate temporal stability. Moreover, we harvested spatial replicates at the last sampling date to explore spatial stability.4. The coefficient of variation of spatial and temporal replicates served as a proxy for ecosystem stability. Both spatial and temporal stability increased to a similar extent with plant diversity. Moreover, there was a positive correlation between spatial and temporal stability, and elevated plant density might be a crucial factor governing the stability of diverse plant communities.5. Above-ground insects generally increased temporal stability, whereas impacts of both earthworms and above-ground insects depended on plant species richness and the presence of grasses. These results suggest that inconsistent results of previous studies on the diversity–stability relationship have in part been due to neglecting higher trophic-level interactions governing ecosystem stability.6. Changes in plant species diversity in one trophic level are thus unlikely to mirror changes in multitrophic interrelationships. Our results suggest that both above- and below-ground invertebrates decouple the relationship between spatial and temporal stability of plant community productivity by differently affecting the homogenizing mechanisms of plants in diverse plant communities.7.Synthesis. Species extinctions and accompanying changes in multitrophic interactions are likely to result not only in alterations in the magnitude of ecosystem functions but also in its variability complicating the assessment and prediction of consequences of current biodiversity loss.

Environmental correlates of species rank − abundance distributions in global drylands

Theoretical models predict lognormal species abundance distributions (SADs) in stable and productive environments, with log-series SADs in less stable, dispersal driven communities. We studied patterns of relative species abundances of perennial vascular plants in global dryland communities to: (i) assess the influence of climatic and soil characteristics on the observed SADs, (ii) infer how environmental variability influences relative abundances, and (iii) evaluate how colonisation dynamics and environmental filters shape abundance distributions. We fitted lognormal and log-series SADs to 91 sites containing at least 15 species of perennial vascular plants. The dependence of species relative abundances on soil and climate variables was assessed using general linear models. Irrespective of habitat type and latitude, the majority of the SADs (70.3%) were best described by a lognormal distribution. Lognormal SADs were associated with low annual precipitation, higher aridity, high soil carbon content, and higher variability of climate variables and soil nitrate. Our results do not corroborate models predicting the prevalence of log-series SADs in dryland communities. As lognormal SADs were particularly associated with sites with drier conditions and a higher environmental variability, we reject models linking lognormality to environmental stability and high productivity conditions. Instead our results point to the prevalence of lognormal SADs in heterogeneous environments, allowing for more evenly distributed plant communities, or in stressful ecosystems, which are generally shaped by strong habitat filters and limited colonisation. This suggests that drylands may be resilient to environmental changes because the many species with intermediate relative abundances could take over ecosystem functioning if the environment becomes suboptimal for dominant species.

(Table 1) Species richness and diversity of tundra plants in herbivore exclosure and control plots near Barrow, Alaska

To determine the role lemmings play in structuring plant communities and their contribution to the 'greening of the Arctic', we measured plant cover and biomass in 50 + year old lemming exclosures and control plots in the coastal tundra near Barrow, Alaska. The response of plant functional types to herbivore exclusion varied among land cover types. In general, the abundance of lichens and bryophytes increased with the exclusion of lemmings, whereas graminoids decreased, although the magnitude of these responses varied among land cover types. These results suggest that sustained lemming activity promotes a higher biomass of vascular plant functional types than would be expected without their presence and highlights the importance of considering herbivory when interpreting patterns of greening in the Arctic. In light of the rapid environmental change ongoing in the Arctic and the potential regional to global implications of this change, further exploration regarding the long-term influence of arvicoline rodents on ecosystem function (e.g. carbon and energy balance) should be considered a research priority.

Parches de vegetación arvense en una toposecuencia de altos y bajos en cultivos de manzanos : Mendoza (Argentina)

En un cultivo de manzanos en la provincia de Mendoza, Argentina, se observó la presencia de las comunidades dominadas por Cynodon dactylon y Galinsoga parviflora localizadas entre y bajo las copas de los árboles frutales, en coincidencia con una toposecuencia de relieves de altos y bajos, respectivamente. Ante el hecho de estar frente a un interesante ejemplo agronómico de establecimiento de comunidades arvenses se las analizó fisonómica, florística y sociológicamente, relacionándolas con las condiciones ecológicas, físico-químicas de los suelos y bioclimáticas de los sitios que las contienen. Mientras la comunidad de Cynodon dactylon presente en los altos o entre plantas resultó más luminosa, cálida, seca y menos fértil, la de Galinsoga parviflora en los bajos fue más sombría, fría, húmeda y más fértil. En esta última, la alta concentración de materia orgánica, nutrientes minerales y humedad de los suelos parece estar relacionada con la presencia de plantas anuales bajo las copas. Los resultados obtenidos muestran la importancia de las toposecuencias de relieves de altos y bajos en la distribución, estructura y composición florística de las comunidades vegetales arvenses, las cuales pueden resultar potenciadas con las coberturas de las copas de los árboles frutales.

Flora y vegetación del centro urbano de Luján de Cuyo : Mendoza (Argentina)

Se analizó la flora y vegetación del centro urbano de Luján de Cuyo, Mendoza, Argentina. Florísticamente se reconocieron 20 familias, 54 géneros y 61 especies. Estas últimas son mayormente introducidas (68.8 %) y el resto, nativas y endémicas (31.2 %); son más numerosas las perennes (54.0 %) que las anuales (42.7 %) y bienales (3.3 %). Estructuralemente dominan las terófitas (44.3 %) sobre las hemicriptófitas (26.2 %), geófitas (16.4 %), caméfitas (9.8 %) y nanofanerófitas (3.3 %). Se reconocieron 15 comunidades vegetales y 2 clases fitosociológicas: Stellarietea mediae y Molinio-Arrhenatheretea. La primera de las clases contiene las comunidades dominadas por Sisymbrium irio, Malva parviflora, Bromus catharticus y Sonchus oleraceus, que son las de mayor cobertura y representación en el área.

Pérdidas de biodiversidad vegetal en ambientes de cerrilladas pedemontanas de Mendoza, Argentina.

Se analizaron las pérdidas de biodiversidad vegetal en un área desmontada de las cerrilladas pedemontanas de Mendoza, Argentina. El análisis de la vegetación y su flora reveló la pérdida total de las comunidades vegetales de Larrea cuneifolia Cav., bosques riparios de Acacia furcatispina Burkart y de álveos y de la flora compuesta de 30 familias de plantas, 72 géneros y 84 especies. Esta última incluyó la de 34,1 % de especies endémicas, 58,8 % nativas, 4,7 % adventicias y 2,4 % introducidas. Se sugiere tener en cuenta estos tipos de estudios antes de realizar planificaciones urbanas sobre la vegetación natural con el fin de reconocer las comunidades vegetales y su flora y rescatar sus bancos de germoplasma.

Propuesta metodológica para la evaluación de tierras periféricas al oasis irrigado de Lavalle, Mendoza (Argentina)

La factibilidad de obtener productos agrícolas de calidad, irrigados con agua salina, hace necesario evaluar áreas periféricas al oasis irrigado para determinar su aptitud para el cultivo. El objetivo es articular diversas metodologías para cartografiar y evaluar unidades ecológicamente homogéneas en tierras marginales de Mendoza utilizando las comunidades vegetales como indicador. Se relevó un área piloto mediante procesamiento de imagen LANDSAT/TM y se analizó la vegetación, suelo y freática. Las unidades evaluadas y sus respectivos rangos de salinidad, en dS m-1, hasta 50 cm fueron: estepa de Heterostachys ritteriana, 91-83; matorral de Allenrolfea vaginata con H. ritteriana, 83-48; matorral abierto de A.vaginata con Prosopis strombulifera, 62-55; estepa abierta de Suaeda divaricata con Atriplex argentina, 52-83; matorral de S. divaricata con Lycium tenuispinosum, 40-43; bosquecillo abierto de Prosopis flexuosa, 37-26; matorral denso de Prosopis alpataco, 7-6. La concentración de sales solubles sobrepasa los niveles críticos tolerados por las plantas cultivadas. No se detectaron niveles freáticos en los primeros 15 m de profundidad. Se considera que los suelos tienen buen drenaje interno, por lo que pueden ser recuperados sin inversiones elevadas. Este enfoque metodológico mostró una buena correspondencia entre las comunidades vegetales, su expresión espectral y las condiciones de salinidad y drenaje del suelo.

La vegetación de los médanos grandes, provincia de San Juan

El objetivo general de la tesis es contribuir al conocimiento de la vegetación sammófila del centro-oeste de la Argentina mediante el análisis geosinfitosociológico (florístico y sinecológico) (paisaje vegetal) de la vegetación de los Médanos Grandes–San Juan, uno de los sistemas eólicos más importantes de Argentina. El análisis realizado permite definir en el sistema dos subambientes, el dominado por procesos eólicos, con megadunas, y el dominado por procesos fluvio-eólicos, sin megadunas. El sistema, en general estabilizado y fijo, tiene actividad sólo en las crestas con escasa cobertura vegetal. La vegetación juega un papel determinante en la dinámica de la arena y en el modelado de este sistema, estando estrechamente relacionada con la disponibilidad de agua que se ajusta a tres modelos: -el de escurrimiento superficial y subsuperficial desde la bajada pedemontana de la sierra de Pie de Palo al norte, -el de la freática relacionada con los ríos San Juan, al oeste, y Bermejo, al este, y – el del agua de lluvia que en las megadunas determina un bulbo húmedo con contenidos de 1,21-2,4 g de agua/100 g de arena, entre los 15-35 m de profundidad, aprovechada por las raíces de los arbustos. Cuatro comunidades vegetales dominan en el sistema:-el pastizal de Panicum urvilleanum en las crestas, el -matorral de Tricomaria usillo-Bulnesia retama en las laderas de las dunas e intermédanos a más de 690 m, el -matorral de Atriplex lampa en los sectores marginales con suelos salinos y el -bosque de Prosopis flexuosa en los intermédanos bajos. Sintaxonómicamente la vegetación pertenece a tres Clases, la Panico urvilleani-Sporoboletea rigentis Esk., 1992 en ambientes sammófilos, la Suaedetea divaricatae Alonso et Conticello ex Martinez Carretero, 2001 en ambientes halófilos y la Larreetea divaricato-cuneifoliae Roig, 89 en la estepa arbustiva del Monte. La relación entre las comunidades vegetales y las asociaciones geomorfológicas permite establecer dos paisajes: el Paisaje I o de Sistema eólico que incluye el 55 % de la superficie y el Paisaje II o de Sistema fluvio-eólico. Palabras claves: eólico, megadunas, comunidades vegetales, dinamismo, bioclima, paisaje vegetal.

(Table 1) n-alkane distribution, and stable carbon and deuterium isotope ratios of DSDP Hole 10-94

Hydrology, source region, and timing of precipitation are important controls on the climate of the Great Plains of North America and the composition of terrestrial ecosystems. Moisture delivered to the Great Plains varies seasonally and predominately derives from the Gulf of Mexico/Atlantic Ocean with minor contributions from the Pacific Ocean and Arctic region. For this work, we evaluate long-term relationships for the past ~ 35 million years between North American hydrology, climate, and floral change, using isotopic records and average carbon chain lengths of higher plant n-alkanes from Gulf of Mexico sediments (DSDP Site 94). We find that carbon isotope values (d13C) of n-alkanes, corrected for variations in the d13C value of atmospheric CO2, provide minor evidence for contributions of C4 plants prior to the Middle Miocene. A sharp spike in C4 input is identified during the Middle Miocene Climatic Optimum, and the influence of C4 plants steadily increased during the Late Miocene into the Pleistocene - consistent with other North American records. Chain-length distributions of n-alkanes, indicative of the composition of higher plant communities, remained remarkably constant from 33 to 4 Ma. However, a trend toward longer chain lengths occurred during the past 4 million years, concurrent with an increase in d13C values, indicating increased C4 plant influence and potentially aridity. The hydrogen isotope values (dD) of n-alkanes are relatively invariant between 33 and 9 Ma, and then become substantially more negative (75 per mil) from 9 to 2 Ma. Changes in the plant community and temperature of precipitation can solely account for the observed variations in dD from 33 to 5 Ma, but cannot account for Plio-Pleistocene dD variations and imply substantial changes in the source region of precipitation and seasonality of moisture delivery. We posit that hydrological changes were linked to tectonic and oceanographic processes including the shoaling and closure of the Panamanian Seaway, amplification of North Atlantic Deep Water Production and an associated increase of meridional winds. The southerly movement of the Intertropical Convergence Zone near 4 Ma allowed for the development of a near-modern pressure/storm track system, driving increased aridity and changes in seasonality within the North American interior.

Flower and seed biomass, and germination rate of arctic tundra plants in response to long term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long-term (12 years) experimental warming in the High Arctic. Open-top chambers (OTCs) were used to increase growing season temperatures by 1-2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long-term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers - via increased forage availability, and the global carbon budget - as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.

Contribution of woody habitat islands to the conservation of birds and their potential ecosystem services in an extensive Colombian rangeland

The conservation of birds and their habitats is essential to maintain well-functioning ecosystems including human-dominated habitats. In simplified or homogenized landscapes, patches of natural and semi-natural habitat are essential for the survival of plant and animal populations. We compared species composition and diversity of trees and birds between gallery forests, tree islands and hedges in a Colombian savanna landscape to assess how fragmented woody plant communities affect forest bird communities and how differences in habitat characteristics influenced bird species traits and their potential ecosystem function. Bird and tree diversity was higher in forests than in tree islands and hedges. Soil depth influenced woody species distribution, and canopy cover and tree height determined bird species distribution, resulting in plant and bird communities that mainly differed between forest and non-forest habitat. Bird and tree species and traits widely co-varied. Bird species in tree islands and hedges were on average smaller, less specialized to habitat and more tolerant to disturbance than in forest, but dietary differences did not emerge. Despite being less complex and diverse than forests, hedges and tree islands significantly contribute to the conservation of forest biodiversity in the savanna matrix. Forest fragments remain essential for the conservation of forest specialists, but hedges and tree islands facilitate spillover of more tolerant forest birds and their ecological functions such as seed dispersal from forest to the savanna matrix.

Environmental characteristics, and growth traits and leaf chemistry of tundra plants in a warming experiment at Alexandra Fiord

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate-vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open-top chambers (OTCs) to passively warm vegetation by 1-2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf d15N. Long-term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect d15N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.