Holocene forest development in the central Scandes Mountains, Sweden

Pollen analyses of sediment cores from two small lakes within the boreal forest in the central Scandes Mountains help to elucidate the Holocene forest dynamics of the region. Analyses of pore/pollen grain diameter ratios of Alnus grains indicate the early Holocene presence of Alnus glutinosa in the study area. The results are discussed in conjunction with available pollen records to evaluate the importance of thermophilous trees during the early Holocene and to deduce the regional spread of Picea abies. Corylus avellana, Alnus glutinosa and Ulmus glabra were probably common constituents of the early Holocene forest. Tilia cordata may have occurred there as a rare tree. Pollen stratigraphies from the region do not indicate the occurrence of Quercus robur. The regional spread of Picea abies can be separated into two phases: a mid-Holocene establishment or first expansion of small outpost populations and a late-Holocene population expansion. The mid-Holocene shift in vegetation composition may have been caused by changes in the westerly airflow.     

Complex effects of mammalian grazing on extramatrical mycelial biomass in the Scandes forest‐tundra ecotone

Mycorrhizal associations are widespread in high‐latitude ecosystems and are potentially of great importance for global carbon dynamics. Although large herbivores play a key part in shaping subarctic plant communities, their impact on mycorrhizal dynamics is largely unknown. We measured extramatrical mycelial (EMM) biomass during one growing season in 16‐year‐old herbivore exclosures and unenclosed control plots (ambient), at three mountain birch forests and two shrub heath sites, in the Scandes forest‐tundra ecotone. We also used high‐throughput amplicon sequencing for taxonomic identification to investigate differences in fungal species composition. At the birch forest sites, EMM biomass was significantly higher in exclosures (1.36 ± 0.43 g C/m²) than in ambient conditions (0.66 ± 0.17 g C/m²) and was positively influenced by soil thawing degree‐days. At the shrub heath sites, there was no significant effect on EMM biomass (exclosures: 0.72 ± 0.09 g C/m²; ambient plots: 1.43 ± 0.94). However, EMM biomass was negatively related to Betula nana abundance, which was greater in exclosures, suggesting that grazing affected EMM biomass positively. We found no significant treatment effects on fungal diversity but the most abundant ectomycorrhizal lineage/cortinarius, showed a near‐significant positive effect of herbivore exclusion (p = .08), indicating that herbivory also affects fungal community composition. These results suggest that herbivory can influence fungal biomass in highly context‐dependent ways in subarctic ecosystems. Considering the importance of root‐associated fungi for ecosystem carbon balance, these findings could have far‐reaching implications.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

Recent cooling and recession of Norway spruce (Picea abies (L.) Karst.) in the forest—alpine tundra ecotone of the Swedish Scandes

Abstract. Repeated crown condition surveys, 1974–94, of subalpine clonal groups of Norway spruce ( Picea abies (L.) Karst.) were carried out in the Swedish Scandes. Complementary analyses concerned radial and vertical growth, sexual regeneration and range limit responses of other plant species. Significant defoliation of spruce progressed linearly over the period of study, reaching cumulative values of about 85%. It is inferred that defoliation was preconditioned by decreasing radial growth since the thermal climax in 1937 and was proximately initiated by the extremely cold winter of 1965/66 and paralleled by consistently declining radial growth and staggering vertical increase. It appears that severe and prolonged ground freezing invoked winter desiccation (xylem cavitation), extensive needle loss and reduced radial growth. Hypothetically, from circumstantial evidence, these processes are interrelated in a positive feedback system, implying increasing sensitivity to climatic stress and decreasing ability to take advantage of positive climatic anomalies. Thus, the total demise of the supranival stems is cautiously predicted, by linear regression of the 20-yr defoliation pattern, to be less than a decade ahead. The recession of P. abies , clearly relevant in a landscape perspective, conforms with analogous responses of Pinus sylvestris L. and Betula pubescens Ehrh. ssp. tortuosa (Ledeb.) Nyman and a significant altitudinal range-limit retraction of certain silvine field-layer species. The structural development examined in this study concurs with long-term climate cooling and cold events and strongly contrasts with simulations of the performance of this system in response to a putative enhanced'greenhouse'effect.     

Holocene environmental history of Lake Vuolep Njakajaure (Abisko National Park, northern Sweden) reconstructed using biological proxy indicators

Holocene environmental and climatic changes are reconstructed using analyses of biological proxies in lake sediments from Vuolep Njakajaure, a lake located near the altitudinal treeline in northern Sweden (68°20′ N, 18°47′ E). We analysed biological proxy indicators from both aquatic and terrestrial ecosystems, including diatoms, pollen and chironomid head capsules, in order to reconstruct regional Holocene climate and the development of the lake and its catchment. During the early Holocene and after 2500 cal b.p., Fragilaria taxa dominated the diatom assemblages, whereas planktonic Cyclotella taxa prevailed during the major part of the Holocene (7800–2300 cal b.p.), indicating the importance of the pelagic habitat for diatom assemblage composition. The planktonic diatoms appeared at the same time as Alnus became established in the catchment, probably altering nutrient availability and catchment stability. The pollen record is dominated by mountain birch (Betula pubescens ssp. tortuosa) pollen throughout the Holocene, but high percentage abundances of Scots pine (Pinus sylvestris) pollen suggest the presence of a mixed pine-birch forest during the mid-Holocene (6800–2300 cal b.p.). Head capsules of Tanytarsini and Psectrocladius dominated the chironomid assemblage composition throughout the Holocene, in combination with Corynocera ambigua after 2300 cal b.p. A quantitative, diatom-based reconstruction of mean July air temperature indicated a relatively cold temperature during the early Holocene (9000–8000 cal b.p.) and after ca. 2300 cal b.p., whereas the mid-Holocene period is characterised by stable and warm temperatures. The overall patterns of Holocene climate and environmental conditions are similarly described by all biological proxy-indicators, suggesting relatively warm conditions during the mid-Holocene (ca. 7800–2300 cal b.p.), with a subsequent colder climate after 2300 cal b.p. However, the onset and magnitude of the inferred changes differ slightly among the proxies, illustrating different responses to lake development phases, land-uplift, and climate forcing (e.g., insolation patterns) during the Holocene in northern Sweden.     

Tree dynamics and co‐existence in the montane–sub‐alpine ecotone: the role of different light‐induced strategies

Questions: Is light availability the main factor driving forest dynamics in Pyrenean sub‐alpine forests? Do pines and firs differ in growth, mortality and morphological response to low light availability? Can differences in shade tolerance affect predictions of future biome changes in Pyrenean sub‐alpine forests in the absence of thermal limitation? Location: Montane–sub‐alpine ecotones of the Eastern Pyrenees (NE Spain). Methods: We evaluated morphological plasticity, survival and growth response of saplings of Scots pine, mountain pine and silver fir to light availability in a mixed forest ecotone. For each species, we selected 100 living and 50 dead saplings and measured size, crown morphology and light availability. A wood disk at root collar was then removed for every sapling, and models relating growth and mortality to light were obtained. Results: Fir had the lowest mortality rate (<0.1) for any given light condition. Pines had comparable responses to light availability, although in deep shade Scots pine risked higher mortality (0.35) than mountain pine (0.19). Pines and fir developed opposing strategies to light deprivation: fir employed a conservative strategy based on sacrificing height growth, whereas pines enhanced height growth to escape from shade, but at the expense of higher mortality risk. Scots pine showed higher plasticity than mountain pine for all architectural and morphological traits analysed, having higher adaptive capacity to a changing environment. Conclusions: Our results support the prediction of future biome changes in Pyrenean sub‐alpine forests as silver fir and Scots pine may find appropriate conditions for colonizing mountain pine‐dominated stands due to land‐use change‐related forest densification and climate warming‐related temperature increases, respectively.     

Reconstruction of Holocene lake-level changes in Lake Kalvsjön, southern Sweden, with a contribution to the local palaeohydrology at the Elm Decline

Past lake-level changes in Lake Kalvsjön are reconstructed and compared with the changes recorded in nearby Lake Bysjön. The two major lowerings of lake level noted in Lake Bysjön are also recorded in Lake Kalvsjön, the older lowering taking place between ca. 9500–9200 B.P. and the younger occurring between ca. 6500–3000 B.P. A distinct decrease in the frequency ofUlmus, corresponding to the classical Elm Decline, is recorded in the pollen diagram from Lake Kalvsjön. The high rapidity of the decline strongly suggests that a pathogenic attack was primarily responsible. However, both human interference and palaeohydrological change may have interacted by disturbing the surrounding forests and increasing the susceptibility of elm to pathogenic attack. In the Lake Kalvsjön area, the disturbance resulting from palaeohydrological change is assumed to have been more influential in pre-disposing the forest to an outbreak of elm disease than any human interference.     

Late Quaternary dynamics of pinewoods in the Iberian Mountains

The role of pinewoods as native forests in the Iberian Mountains, and their long-term dynamics during the last interglacial, has been intensely debated among geobotanists. This debate has resulted in several different hypotheses regarding the interpretation of the present day landscape.Recent research on Late Quaternary palaeoecological records has enabled long-term patterns of forest dynamics in the main ranges of the Iberian Peninsula to be tested, and the parallel analysis of micro- (mainly pollen) and macroremains (woods, charcoals, fruits, seeds and leaves) has enhanced the resolution of these palaeobiological records. Additional historical information has allowed a complete overview of the past vegetal landscapes to be obtained.This work summarises the available data for Iberian mountain pines during the Lateglacial and the Holocene, focussing on the mountain regions in scenarios of absence or dominance. Based on this overview, life history traits and patterns of plant dynamics during the Holocene are discussed and compared with present landscapes.In the long term, Pinus does not always respond as a serial element in succession dynamics, not even in very favourable environments for hardwoods. Processes such as incumbency or resilience, and features such as frugality and their ability to disperse, affect the persistence of pines to the present day. This is exemplified for continental mountain areas, in which the absence of anthropogenic activities could have determined the wider coverage of pinewoods seen today.     

The Late Holocene extinction of Pinus sylvestris in the western Cantabrian Range (Spain)

Aim To reconstruct the historical biogeography of Pinus sylvestris in the Cantabrian Range (Iberian Peninsula) during the Holocene, and to consider the interactions between vegetation dynamics, climate change and the role of man in the present‐day distribution of the species. Location The study site is a mire (1300 m a.s.l.) at Vega de Viejos, on a south‐facing slope of the western Cantabrian Range, Spain. The region’s present‐day landscape is almost treeless, with the exception of some patches of Quercus pyrenaica and a few copses of Salix and Betula along stream banks. Methods Tree macrofossils from Vega de Viejos were studied by transmitted light and dark‐field reflection microscopy; strobili were subjected to comparative morphological analyses. Two Pinus macrofossils were dated by conventional ¹⁴C methods. Results The taxonomic accuracy achieved in the identification of the macrofossils provided new information regarding the Holocene history of Pinus sylvestris in this territory. Ninety‐five cones of this species were identified; in fact, more than 80% of the 36 identified wood remains were of Pinus gr. sylvestris. Radiocarbon dating revealed that the forest to which the fossils belonged was present until at least 2170 ± 50 yr bp – its disappearance was therefore relatively recent. Main conclusions Pinus sylvestris suffered long‐term isolation, and after the Würm glacial period tended to migrate towards the east. In western Iberia, a temperate climate and autogenic succession favoured broadleaved taxa at the expense of Pinus. Late Holocene human disturbances may have further accelerated the decline of P. sylvestris; in the Cantabrian Range, only a few stands on southern slopes have persisted until the present day. The history of the capercaillie (Tetrao urogallus), a bird characteristic of pure or mixed Palaeartic coniferous forests, was almost certainly affected by the demise of these forests in this area. Cantabrian capercaillies are the only members of this species that live in purely deciduous forests, perhaps a recent adaptation to the regional extinction of pines. Today’s P. sylvestris and capercaillie populations are now highly fragmented and their future, given the predictions of global climate change, is uncertain.     

Warming‐induced upward migration of the alpine treeline in the Changbai Mountains,northeast China

Treeline responses to environmental changes describe an important phenomenon in global change research. Often conflicting results and generally too short observations are, however, still challenging our understanding of climate‐induced treeline dynamics. Here, we use a state‐of‐the‐art dendroecological approach to reconstruct long‐term changes in the position of the alpine treeline in relation to air temperature at two sides in the Changbai Mountains in northeast China. Over the past 160 years, the treeline increased by around 80 m, a process that can be divided into three phases of different rates and drives. The first phase was mainly influenced by vegetation recovery after an eruption of the Tianchi volcano in 1702. The slowly upward shift in the second phase was consistent with the slowly increasing temperature. The last phase coincided with rapid warming since 1985, and shows with 33 m per 1°C, the most intense upward shift. The spatial distribution and age structure of trees beyond the current treeline confirm the latest, warming‐induced upward shift. Our results suggest that the alpine treeline will continue to rise, and that the alpine tundra may disappear if temperatures will increase further. This study not only enhances mechanistic understanding of long‐term treeline dynamics, but also highlights the effects of rising temperatures on high‐elevation vegetation dynamics.     

Demography of Betula pubescens ssp. tortuosa sown in contrasting habitats close to the birch tree-limit in Central Sweden

Seeds of mountain birch (Betula pubescens Ehrh. ssp. tortuosa (Ledeb.) Nyman) were sown on permanent plots in three contrasting habitats close to the birch tree-limit in Central Sweden. The habitats differed mainly in regard to the length of the growth period, as determined by the pattern of local snow-accumulation. Germination success, seedling demography (1981–1983) and related aspects of the early stages were studied. Germination was found to start shortly after snowmelt in the year after ripening, in all three habitats. The conclusion is that germination is not the crucial phase of the life-cycle. In the habitat with the longest growth period, and where spontaneous birch stands are present, mortality mainly occurred in summer, evidently caused by soil drought. With shortening of the growth period, however, mortality during the winter became increasingly important. Survival was clearly correlated with the length of the growth period and the autumn height of seedlings. Factors which affect ecesis determine the altitude of the tree-limit and the long-term stability of the mountain birch forest ecosystem.     

The history and control of the pine beauty moth, Panolis flammea (D. & S.) (Lepidoptera: Noctuidae), in Scotland from 1976 to 2000

1 The pine beauty moth, Panolis flammea, has been a serious pest of lodgepole pine plantations in Scotland since 1976. It historically feeds on native Scots pine throughout Europe but population levels of P. flammea on this host have never been high enough to cause tree mortality in the U.K. 2 This paper reviews recent advances in the biology of the pest and documents control programmes from 1976 to 1999. 3 There has been practically uninterrupted population monitoring of P. flammea from 1977 to the present day in Scottish lodgepole pine plantations. Intervention with chemical spraying has often been necessary. 4 The population data suggest that populations of P. flammea may have had a cyclic pattern over the monitoring period, with outbreaks occurring at regular intervals of between 6 and 7 years. 5 The amplitude of population cycles was large during the 1970s and 1980s, but has dampened in recent years. Natural enemies are believed to contribute to this trend. Fungal disease, specifically, appears to have had a greater effect on pest populations in recent years than in the past and is suggested to have contributed significantly to the population dynamics observed since 1990.     

Did tree‐Betula,Pinus and Picea survive the last glaciation along the west coast of Norway? A review of the evidence,in light of Kullman (2002)

Aim We discuss the hypotheses proposed by Kullman [Geo‐Öko 21 (2000) 141; Nordic Journal of Botany 21 (2001) 39; Journal of Biogeography 29 (2002) 1117] on the basis of radiocarbon‐dated megafossils of late‐glacial age from the central Swedish mountains that boreal trees survived the glaciation along the south‐west coast of Norway and subsequently migrated eastward early in the late‐glacial to early deglaciated parts of the central Swedish Scandes mountains. Methods We assess these hypotheses on the basis of glacial geological evidence and four lines of palaeoecological evidence, namely macrofossil records of the tree species, vegetation and climate reconstructions from plant evidence, independent climate reconstructions from other proxies for the late‐glacial environment of south‐west Norway, and the patterns of post‐glacial spread of the tree species. Location South and west Norway, central Swedish Scandes mountains (Jämtland). Results and conclusions South‐west Norway and the adjacent continental shelf were under ice at the last‐glacial maximum (LGM). The late‐glacial vegetation of south‐west Norway was treeless and summer temperatures were below the thermal limits for Betula pubescens Ehrh., Pinus sylvestris L. and Picea abies (L.) Karst. Instead of spreading immediately after the onset of Holocene warming, as might have been expected if local populations were surviving, B. pubescens showed a lag of local arrival of 600 to > 1000 years, Pinus lagged by 1500 to > 2000 years, and Picea only reached southern Norway c. 1500 years ago and has not colonized most of south‐west Norway west of the watershed. Glacial geological evidence shows the presence of an ice sheet in the Scandes at the LGM and in the Younger Dryas, which was cold‐based near or at the area where the late‐glacial‐dated megafossils were recovered by Kullman. We conclude that the samples dated by Kullman (2002) should be evaluated carefully for possible sources of contamination. All the available evidence shows that the biogeographical hypotheses, based on these radiocarbon dates taken at face value, of late‐glacial tree survival at the Norwegian coast and subsequent eastwards spread to the mountains, are unsupportable.     

Vegetation history of Lago Battaglia (eastern Gargano coast,Apulia, Italy) during the middle-late Holocene

A pollen and charcoal record from Gargano (southern Italy) provides new information on the vegetation history and environmental change in southern Italy during the middle to late Holocene. The chronological framework is provided by six AMS radiocarbon dates carried out on plant macroremains. Pollen diagrams record a Mediterranean vegetation along the coastland and a stable mixed oak forest at higher elevations between ca. 5900 and 4200 cal b.p. A sharp and dramatic fall of tree pollen concentrations and a change in fire frequencies occurred from approximately 4200 cal b.p. suggesting a climate change towards drier conditions. This event is coherent with regional and extra-regional palaeoclimatic records that suggest that a mid-Holocene dry period was experienced in southern Italy, southern Spain, and perhaps further afield. Human impact on vegetation seems to have occurred since about 2700 cal b.p.