How do subordinate and dominant species in semi-natural mountain grasslands relate to productivity and land-use change?

Changes in agricultural practices of semi-natural mountain grasslands are expected to modify plant community structure and shift dominance patterns. Using vegetation surveys of 11 sites in semi-natural grasslands of the Swiss Jura and Swiss and French Alps, we determined the relative contribution of dominant, subordinate and transient plant species in grazed and abandoned communities and observed their changes along a gradient of productivity and in response to abandonment of pasturing. The results confirm the humpbacked diversity–productivity relationship in semi-natural grassland, which is due to the increase of subordinate species number at intermediate productivity levels. Grazed communities, at the lower or higher end of the species diversity gradient, suffered higher species loss after grazing abandonment. Species loss after abandonment of pasturing was mainly due to a higher reduction in the number of subordinate species, as a consequence of the increasing proportion of dominant species. When plant biodiversity maintenance is the aim, our results have direct implications for the way grasslands should be managed. Indeed, while intensification and abandonment have been accelerated since few decades, our findings in this multi-site analysis confirm the importance of maintaining intermediate levels of pasturing to preserve biodiversity.

Climatic and human impacts on mountain vegetation at Lauenensee (Bernese Alps, Switzerland) during the last 14,000 years

Lake sediments from Lauenensee (1381 m a.s.l.), a small lake in the Bernese Alps, were analysed to reconstruct the vegetation and fire history. The chronology is based on 11 calibrated radiocarbon dates on terrestrial plant macrofossils suggesting a basal age of 14,200 cal. BP. Pollen and macrofossil data imply that treeline never reached the lake catchment during the Bølling–Allerød interstadial. Treeline north of the Alps was depressed by c. 300 altitudinal meters, if compared with southern locations. We attribute this difference to colder temperatures and to unbuffered cold air excursions from the ice masses in northern Europe. Afforestation started after the Younger Dryas at 11,600 cal. BP. Early-Holocene tree-Betula and Pinus sylvestris forests were replaced by Abies alba forests around 7500 cal. BP. Continuous high-resolution pollen and macrofossil series allow quantitative assessments of vegetation dynamics at 5900–5200 cal. BP (first expansion of Picea abies, decline of Abies alba) and 4100–2900 cal. BP (first collapse of Abies alba). The first signs of human activity became noticeable during the late Neolithic c. 5700–5200 cal. BP. Cross-correlation analysis shows that the expansion of Alnus viridis and the replacement of Abies alba by Picea abies after c. 5500 cal. BP was most likely a consequence of human disturbance. Abies alba responded very sensitively to a combination of fire and grazing disturbance. Our results imply that the current dominance of Picea abies in the upper montane and subalpine belts is a consequence of anthropogenic activities through the millennia.

Analysis of coprolites from the extinct mountain goat Myotragus balearicus

Humans colonized the Balearic Islands 5-4 ka ago. They arrived in a uniquely adapted ecosystem with the Balearic mountain goat Myotragus balearicus (Bovidae, Antilopinae, Caprini) as the only large mammal. This mammal went extinct rapidly after human arrival. Several hypotheses have been proposed to explain the extinction of M. balearicus. For the present study ancient DNA analysis (Sanger sequencing, Roche-454, Ion Torrent), and pollen and macrofossil analyses were performed on preserved coprolites from M. balearicus, providing information on its diet and paleo-environment. The information retrieved shows that M. balearicus was heavily dependent on the Balearic box species Buxus balearica during at least part of the year, and that it was most probably a browser. Hindcast ecological niche modelling of B. balearica shows that local distribution of this plant species was affected by climate changes. This suggests that the extinction of M. balearicus can be related to the decline and regional extinction of a plant species that formed a major component of its diet. The vegetation change is thought to be caused by increased aridity occurring throughout the Mediterranean. Previous hypotheses relating the extinction of M. balearicus directly to the arrival of humans on the islands must therefore be adjusted. (C) 2013 University of Washington. Published by Elsevier Inn All rights reserved.

Ecosystem services, biodiversity and environmental change in a tropical mountain ecosystem of south Ecuador

An interdisciplinary research unit consisting of 30 teams in the natural, economic and social sciences analyzed biodiversity and ecosystem services of a mountain rainforest ecosystem in the hotspot of the tropical Andes, with special reference to past, current and future environmental changes. The group assessed ecosystem services using data from ecological field and scenario-driven model experiments, and with the help of comparative field surveys of the natural forest and its anthropogenic replacement system for agriculture. The book offers insights into the impacts of environmental change on various service categories mentioned in the Millennium Ecosystem Assessment (2005): cultural, regulating, supporting and provisioning ecosystem services. Examples focus on biodiversity of plants and animals including trophic networks, and abiotic/biotic parameters such as soils, regional climate, water, nutrient and sediment cycles. The types of threats considered include land use and climate changes, as well as atmospheric fertilization. In terms of regulating and provisioning services, the emphasis is primarily on water regulation and supply as well as climate regulation and carbon sequestration. With regard to provisioning services, the synthesis of the book provides science-based recommendations for a sustainable land use portfolio including several options such as forestry, pasture management and the practices of indigenous peoples. In closing, the authors show how they integrated the local society by pursuing capacity building in compliance with the CBD-ABS (Convention on Biological Diversity - Access and Benefit Sharing), in the form of education and knowledge transfer for application.

A model-data comparison of Holocene timberline changes in the Swiss Alps reveals past and future drivers of mountain forest dynamics

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11700years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid-inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim, a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid-inferred July-temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model-data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models (SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500m and treeline positions of ca. 2500m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low-impact pastoralism to preserve species-rich alpine meadows.

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.

Late Glacial and Holocene vegetational changes on the Ulagan high-mountain plateau, Altai Mountains, southern Siberia

Three well-dated pollen diagrams from 1985 m, 2050 m, and at the tree line at 2150 m asl show the vegetational succession in the central Altai Mountains since 16 cal ka BP. Pioneer vegetation after deglaciation was recorded first at the lowest site. Subsequently, dense dry steppe vegetation developed coincident with the change from silt to organic sediments at the two lower sites, but silt lasted longer at the highest site, indicating the persistence of bare ground there. Forests of Pinus sibirica, Pinus sylvestris, Picea obovata, Larix sibirica, Abies sibirica, and Betula pendula started to develop about 12 cal ka BP with the change to a warmer and wetter climate at the beginning of the Holocene. Results indicate that the timberline did not rise above the highest site. Mesophilous dark-coniferous forests were fully developed by 9.5 cal ka BP. The role of Abies and Picea decreased by about 7.5 cal ka BP suggesting cooler climate, after which the forests changed little until today. The vegetational development in this portion of the central Altai Mountains is compatible with that described in neighbouring areas of the Altai, southern Siberia, Mongolia, and Kazakhstan.

Reconstruction of Holocene vegetation dynamics at Lac de Bretaye, a high-mountain lake in the Swiss Alps

A deeper understanding of past vegetation dynamics is required to better assess future vegetation responses to global warming in the Alps. Lake sediments from Lac de Bretaye, a small subalpine lake in the Northern Swiss Alps (1780 m a.s.l.), were analysed to reconstruct past vegetation dynamics for the entire Holocene, using pollen, macrofossil and charcoal analyses as main proxies. The results show that timberline reached the lake’s catchment area at around 10,300 cal. BP, supporting the hypothesis of a delayed postglacial afforestation in the Northern Alps. At the same time, thermophilous trees such as Ulmus, Tilia and Acer established in the lowlands and expanded to the altitude of the lake, forming distinctive boreo-nemoral forests with Betula, Pinus cembra and Larix decidua. From about 5000 to 3500 cal. BP, thermophilous trees declined because of increasing human land use, mainly driven by the mass expansion of Picea abies and severe anthropogenic fire activity. From the Bronze Age onwards (c. 4200–2800 cal. BP), grazing indicators and high values for charcoal concentration and influx attest an intensifying human impact, fostering the expansion of Alnus viridis and Picea abies. Hence, biodiversity in alpine meadows increased, whereas forest diversity declined, as can be seen in other regional records. We argue that the anticipated climate change and decreasing human impact in the Alps today will not only lead to an upward movement of timberline with consequent loss of area for grasslands, but also to a disruption of Picea abies forests, which may allow the re-expansion of thermophilous tree species.

Long-term Responses of Mountain Ecosystems to Environmental Changes: Resilience, Adjustment, and Vulnerability

The steep environmental gradients of mountain ecosystems over short distances reflect large gradients of several climatic parameters and hence provide excellent possibilities for ecological research on the effects of environmental change. To gain a better understanding of the dynamics of abiotic and biotic parameters of mountain ecosystems, long-term records are required since permanent plots in mountain regions cover in the best case about 50 - 70 years. In order to extend investigations of ecological dynamics beyond these temporal limitations of permanent plots, paleoecological approaches can be used if the sampling resolution can be adapted to ecological research questions, e.g. a sample every 10 years. Paleoecological studies in mountain ecosystems can provide new ecological insights through the combination of different spatial and temporal scales. [f we thus improve our understanding of processes across both steep environmental gradients and different time scales, we may be able to better estimate ecosystem responses to current and future environmental change (Ammann et al. 1993; Lotter et al. 1997). The complexity of ecological interactions in mountain regions forces us to concentrate on a number of sub-systems - without losing sight of the wider context. Here, we summarize a few case studies on the effects of Holocene climate change and disturbance on the vegetation of the Western Alps. To categorize the main response modes of vegetation to climatic change and disturbance in the Alps we use three classes of ecological behaviour: "resilience", "adjustment", and "vulnerability", We assume a resilient (or elastic) behaviour if vegetation is able to recover to its former state, regaining important ecosystem characteristics, such as floristic composition, biodiversity, species abundances, and biomass (e.g. Küttel 1990; Aber and Melillo 199 1). Conversely, vegetation displacements may occur in response to climatic change and/or disturbance. In some cases, this may culminate in irreversible large-scale processes such as species and/or community extinctions. Such drastic developments indicate high ecosystem vulnerability (or inelasticity or instability, for detailed definitions see Küttel 1990; Aber and Melillo 199 1) to climatic change and/or disturbance. In this sense, the "vulnerability" (or instability) of an ecosystem is expressed by the degree of failure to recover to the original state before disturbance and/or climatic change. Between these two extremes (resilience vs. vulnerability), ecosystem adjustments to climatic change and/or disturbance may occur, including the appearance of new and/or the disappearance of old species. The term "adjustment" is hence used to indicate the response of vegetational communities, which adapted to new environmental conditions without losing their main character. For forest ecosystems, we assume vegetational adjustments (rather than vulnerability) if the dominant (or co-dominant) tree species are not outnumbered or replaced by formerly unimportant plant species or new invaders. Adaptation as a genetic process is not discussed here and will require additional pbylogeographical studies (that incorporate the analysis of ancient DNA) in order to fully understand the distributions of ecotypes.