Regional assessment of the impact of climatic change on the distribution of a tropical conifer in the lowlands of South America

For decades, palynologists working in tropical South America are using the genus Podocarpus as a climate indicator although without referring to any modern data concerning its distribution and limiting factors. With the aim to characterize the modern and past distribution of the southern conifer Podocarpus in Brazil and to obtain new information on the distribution of the Atlantic rainforest during the Quaternary, we examined herbarium data to locate the populations of three Brazilian endemic Podocarpus species: P. sellowii, P. lambertii , and P. brasiliensis , and extracted DNA from fresh leaves from 26 populations. Our conclusions are drawn in the light of the combination of these three disciplines: botany, palynology, and genetics. We find that the modern distribution of endemic Podocarpus populations shows that they are widely dispersed in eastern Brazil, from north to south and reveals that the expansion of Podocarpus recorded in single Amazonian pollen records may have come from either western or eastern populations. Genetic analysis enabled us to delimit regional expansion: between 5° and 15° S grouping northern and central populations of P. sellowii expanded c . 16,000 years ago; between 15° and 23° S populations of either P. lambertii or sellowii expanded at different times since at least the last glaciation; and between 23° and 30° S, P. lambertii appeared during the recent expansion of the Araucaria forest. The combination of botany, pollen, and molecular analysis proved to be a rapid tool for inferring distribution borders for sparse populations and their regional evolution within tropical ecosystems. Today the refugia of rainforest communities we identified are crucial hotspots to allow the Atlantic forest to survive under unfavourable climatic conditions and, as such, offer the only possible opportunity for this type of forest to expand in the event of a future climate change.     

The Holocene history of low altitude Mediterranean Fagus sylvatica forests in southern France

Questions

As the dominant tree in many European forests, Fagus sylvatica functions as an ecosystem engineer, yet its istory remains little understood. Here we ask: (a) are there indications for its presence in southeast France during the last Glacial period; (b) what was the timing of the expansion and decline of F. sylvatica dominated forests; (c) which factors influenced their dynamics and in particular to what extent did past precipitation changes impact upon them; and (d) at which altitudes did these beech forests occur within the region?

Location

Languedoc, the French Mediterranean area.

Method

This article presents a well dated and high‐resolution pollen sequence covering the last 7,800 years from the Palavas Lagoon in the Languedoc together with a review of Fagus charcoal occurrences in the Languedoc and the lower Rhône Valley, and a review of pollen data from a compilation of 69 sites in southeast France.

Results

The Palavas pollen sequence provides a regional summary of F. sylvatica abundance changes near the Mediterranean coast. Around 6,000 years cal BP , an abrupt transition from small beech populations to well‐developed forests is recorded. The maximum development of beech forests occurred between 4,000 and 3,000 years cal BP , while F. sylvatica started to regress after 3,000 years cal BP .

Conclusion

Scattered F. sylvatica populations probably survived throughout southern France during the last Glacial period. F. sylvatica started to spread around 8,000 years cal BP while beech forests never expanded before 6,000 years cal BP . The complex patterns of F. sylvatica expansion in southern France after 6,000 years cal BP suggests that a combination of global (climate change) and local (human impact) factors were responsible for this major change. Recurrent abrupt climate changes, the aridity trend and human deforestation caused beech forests to decline after 3,000 years cal BP .

     

Mammalian responses to Middle Holocene climatic change in the Great Basin of the western United States

In spite of decades of intense research directed toward understanding the climates and ecology of the Great Basin (western United States) during the past 10,000 years, the responses of mammals to the extreme aridity of the Middle Holocene (c. 8000–5000 years ago) in this region have been poorly understood. Using a well‐dated small mammal sequence from Homestead Cave, north‐central Utah, I show that the Middle Holocene small mammal faunas of this area underwent a decrease in species richness and evenness, driven largely by a series of local extinctions and near‐extinctions coupled with a dramatic increase in the abundance of taxa well‐adapted to xeric conditions. At the end of this period, some taxa that require relatively mesic habitats began to increase in abundance immediately, others did not rebound in abundance until several thousand years later, while still others have not returned at all. This suite of responses has been difficult to detect because climatic change at the beginning of the Middle Holocene was so much more substantial than that which occurred toward its end.     

The importance of hornbeam (Carpinus sp.) pollen hypersensitivity in spring allergies

Hornbeam (Carpinus sp.) flowering occurs generallysimultaneously with birch anthesis. While birch pollenis known as the major tree-allergen in central andnorthern Europe, the quantitative and qualitative roleof hornbeam pollen in allergy has not been evaluated.In some regions of Switzerland, the annual ratiohornbeam SPI / birch SPI reaches 20–30% on average,and 50–180% some years. The relative importance ofhornbeam pollen hypersensitivity has been evaluated inmore than 200 patients suffering from spring allergiesto pollen during the last five years (1993–1997) inthe region of Geneva. Among symptomatic patientsduring the March-April period, 73.5% werehypersensitive to Betulaceae pollen. Overall, 71.7%of the patients allergic to Betulaceae pollen werefound to be sensitive to hornbeam, and 12.5% of thesame group of patients were sensitive to hornbeampollen only. Among this last group, half of thepatients, besides allergic rhino-conjunctivitis, werealso suffering from asthma. Hornbeam pollen isabundant and appears to have a significativeimportance in spring allergies. In some cases,specific immunotherapy to hornbeam pollen shouldperhaps be considered.     

Unexpected early extinction of the European pond turtle (Emys orbicularis) in Sweden and climatic impact on its Holocene range

Using ancient DNA sequences of subfossil European pond turtles ( Emys orbicularis ) from Britain, Central and North Europe and accelerator mass spectrometry radiocarbon dating for turtle remains from most Swedish sites, we provide evidence for a Holocene range expansion of the pond turtle from the southeastern Balkans into Britain, Central Europe and Scandinavia, according to the 'grasshopper pattern' of Hewitt. Northeastern Europe and adjacent Asia were colonized from another refuge located further east. With increasing annual mean temperatures, pond turtles reached southern Sweden approximately 9800 years ago. Until approximately 5500 years ago, rising temperatures facilitated a further range expansion up to Östergötland, Sweden (approximately 58°30'N). However, around 5500 years ago pond turtle records suddenly terminate in Sweden, some 1500 years before the Holocene thermal maximum ended in Scandinavia and distinctly earlier than previously thought. This extinction coincides with a temporary cooling oscillation during the Holocene thermal maximum and is likely related to lower summer temperatures deteriorating reproductive success. Although climatic conditions improved later again, recolonization of Sweden from southern source populations was prevented by the Holocene submergence of the previous land connection via the Danish Straits that occurred approximately 8500 years ago.     

Pleistocene glaciation leaves deep signature on the freshwater crab Aegla alacalufi in Chilean Patagonia

Quaternary glacial cycles have played an important role in shaping the biodiversity in temperate regions. This is well documented in Northern Hemisphere, but much less understood for Southern Hemisphere. We used mitochondrial DNA and nuclear elongation factor 1α intron sequences to examine the Pleistocene glacial impacts on the phylogeographical pattern of the freshwater crab Aegla alacalufi in Chilean Patagonia. Phylogenetic analyses, which separated the glaciated populations on eastern continent into a north group (seven populations) and a south group (one population), revealed a shallow phylogenetic structure in the north group but a deep one in the non-glaciated populations on western islands, indicating the significant influence of glaciation on these populations. Phylogenies also identified the Yaldad population on Chiloé Island as a potentially unrecognized new species. The non-glaciated populations showed higher among population genetic divergence than the glaciated ones, but lower population genetic diversity was not detected in the latter. The two glaciated groups, which diverged from the non-glaciated populations at ~96 800–29 500 years ago and ~104 200–73 800 years ago, respectively, seem to have different glacial refugia. Unexpectedly, the non-glaciated islands did not serve as refugia for them. Demographic expansion was detected in the glaciated north group, with a constant population increase after the last glacial maximum. Nested clade analyses suggest a possible colonization from western islands to eastern continent. After arriving on the continent and surviving the last glacial period there, populations likely have expanded from high to low altitude, following the flood of melting ice. Aegla alacalufi genetic diversity has been primarily affected by Pleistocene glaciation and minimally by drainage isolation.     

Population Genetic Study of Mitochondrial DNA in Roseate Spoonbill (Aves; Platalea ajaja) Breeding Colonies from the Pantanal Wetlands, Brazil

Five breeding colonies of the Roseate spoonbill (Aves: Platalea ajaja) from two Brazilian wetland areas (Pantanal and Taim marshes) were sampled, and domain I of the mitochondrial DNA control region (483 bp) was sequenced in 50 birds. The average haplotype diversity (h = 0.75, s = 0.071) and average nucleotide diversity (pi = 0.004, s = 0.003) were evaluated, and nonsignificant differences were found among the colonies studied. The lack of differentiation among breeding colonies revealed by AMOVA analysis was explained either as a consequence of high gene flow or recent expansion. The significantly negative results of the neutrality tests (Fu's F ( s ) = -23.271, P < 0.01; Tajima's D = -1.941, P < 0.01) associated with the star shape of the haplotype tree and mismatch distribution data are evidence supporting the idea that these populations underwent a recent demographic expansion in the Pantanal region. The average time since the expansion is estimated to be 25,773 years, and this agrees with a period of increased moisture that occurred during the last glacial period.     

Compositional vegetation changes and increased red spruce abundance during the Little Ice Age in a sugar maple forest of north-eastern North America

In north-eastern North America, the recent red spruce decline has been linked to atmospheric pollution, notably acid rain, although climate was also advocated as a potential factor. A high resolution lake sediment pollen stratigraphy was obtained to elucidate long-term trends in tree-species abundance in a sugar maple??yellow birch forest. The reconstructed history (~250?C1996?A.D.) showed a steady increase of red spruce after 1300?A.D., with a peak between 1600 and 1900?A.D. followed by a strong decline in the last century, while sugar maple and yellow birch experienced an opposite trend. Red spruce abundance reached its apogee during the cool Little Ice Age (LIA) and decreased abruptly when annual temperature in the region increased by 2?°C in the last 125?years. American Beech was much more abundant in the forest before the LIA, typifying a sugar maple??American beech forest as the dominant forest type during the Late Holocene. Our results suggest that climate warming has played an important role in the current red spruce decline, the latter having been initiated well before acidic depositions reached deleterious potential effects on red spruce. Climate warming probably acted as a long-term predisposing factor that was aggravated by atmospheric pollution, in the last decades.     

Spatial extent of winter thaw events in eastern North America: historical weather records in relation to yellow birch decline

An algorithm (Weather Reader) was developed and used to analyze daily weather records from all existing Canadian and American weather stations of eastern North America (in excess of 2100 stations), from 1930 through 2000. Specifically, the Weather Reader was used to compile daily minimum, mean, and maximum air temperatures for weather stations with at least 30 years of data, and was used to calculate accumulated degree days for winter thaw–freeze events relevant to yellow birch (Betula alleghaniensis Britt.) from beginning to end. A thaw–freeze event relevant to yellow birch was considered to take place when (i) the station daily maximum temperature reached or exceeded +4°C after being below freezing for at least 2 months of the winter, (ii) sufficient growing degree days accumulated (>50 growing degree days) to cause the affected yellow birch trees to prematurely deharden, and (iii) the daily minimum temperature dropped below ?4°C causing roots and/or shoots of dehardened trees to experience freeze‐induced injury and possibly dieback. The threshold temperature of +4°C represents the daily temperature above which biological activity occurs in yellow birch. The station growing degree day summaries were subsequently spatially interpolated with the Kriging function in GS+^? and mapped in ArcView^? GIS in order to display the geographic extent of the most severe thaw–freeze events. The ArcView^? maps were then compared with the extent of historically observed yellow birch decline. It was found that the years 1936, 1944, and 1945 were particularly uncharacteristic in terms of region‐wide winter thaw–freeze extremes, and also in terms of observed birch decline events during 1930–1960. An overlay of suspected accumulated birch decline based on thaw–freeze mapping and observed decline maps prepared by Braathe (1995), Auclair (1987) , and Auclair et al. (1997) for 1930–1960 demonstrated similar geographic patterns. The thaw–freeze projection for 1930–1960 was shown to coincide with 83% of the birch decline map appearing in Braathe (1995) and 55% of the geographic range of yellow birch in eastern North America. Thaw–freeze mapping was also applied to two significant events in 1981. Greatest impact was recorded to occur mostly in southern Quebec and Ontario, and several American Great Lake States, specifically in northern Michigan and New York, where the greatest growing degree day accumulation prior to refreeze in late February (February 28th) was projected to have occurred; and in southern Quebec, most of Atlantic Canada, and Maine, prior to a late spring frost in mid‐April (April 17).     

Airborne birch and oak pollen grains and birch pollen allergens at a common sampling station in Stockholm

In the present study, the airborne concentrations of birch and oak pollen grains and birch pollen allergens have been recorded in samples from a common sampling station in Stockholm. The sampling period was between April 22^nd and May 31^st 2003. The objectives were to evaluate if analysis of allergen particles in parallel with pollen grains would be relevant to aid subjects suffering from pollinosis. Days with low birch pollen counts had relatively high nominal allergen concentrations in the beginning of the sampling period. The birch pollen grain concentration peaks, during the dry pollination culmination interval in the middle of the period, were associated with correspondingly lower nominal concentrations of allergens than grains. At the end of the sampling period very high nominal amounts of allergen appeared, as reflected by high concentrations of oak pollen grains. The high allergen concentrations were obtained as a result of inherent cross‐reactivity of anti‐ Bet v 1 antibodies with Que a antigens, which are immunologically analogous with Bet v 1. Allergen concentrations increased and decreased after light and heavy rain, respectively. Results obtained indicate that adding a pollen count forecast with allergen concentration data should aid pollen allergic subjects to avoid particulate allergens which might be expected to penetrate more easily than pollen grains into indoor environments.     

Potential refugium on the Qinghai–Tibet Plateau revealed by the chloroplast DNA phylogeography of the alpine species Metagentiana striata (Gentianaceae)

Metagentiana striata is an alpine annual herbaceous plant endemic to the east of the Qinghai–Tibet (Q–T) Plateau and adjacent areas. The phylogeography of M. striata was studied by sequencing the chloroplast DNA (cpDNA) trn S– trn G intergenic spacer. Ten haplotypes were identified from an investigation of 232 individuals of M. striata from 14 populations covering the entire geographical range of this species. The level of differentiation amongst populations was very high ( G _ST = 0.746; N _ST = 0.774) and a significant phylogeographical structure was observed ( P  < 0.05). An analysis of molecular variance found a high variation amongst populations (76%), with F _ST = 0.762 (highly significant, P  < 0.001), indicating that little gene flow occurred amongst the different regions; this was explained by the isolation of populations by high mountains along the Q–T Plateau and adjacent areas ( N _m = 0.156). Only one ancestral haplotype (A) was common and widespread throughout the distributional range of M. striata. The populations of the Hengduan Mountains region of the south-eastern Q–T Plateau showed high diversity and uniqueness of haplotypes. It is suggested that this region was the potential refugium of M. striata during the Quaternary glaciation, and that interglacial and postglacial range expansion occurred from this refugium. This scenario was in good agreement with the results of nested clade analysis, which inferred that the current spatial distribution of cpDNA haplotypes and populations resulted from range expansion, together with past allopatric fragmentation events.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 125–140.     

Species-level phylogeographical history of Myricaria plants in the mountain ranges of western China and the origin of M. laxiflora in the Three Gorges mountain region

Myricaria species in China occur mostly in the major high-altitude mountain areas in and around the Qinghai-Tibetan Plateau. The one major exception to this is M. laxiflora which is restricted to the Three Gorges mountain region. In this study, we investigate species-level phylogeographical patterns of Myricaria species in western China and the origin of M. laxiflora . The results show that most chloroplast haplotypes are species-specific, except for one haplotype which is shared by three widespread species. Higher haplotype diversity within the Qinghai-Tibetan Plateau region supports the hypothesis that the Himalayas are the centre of origin for Myricaria . The phylogeny of Myricaria was geographically structured, and an estimated Bayesian chronology suggested the main divergence events occurred during the Late Pliocene and Early Pleistocene (~1.46–2.30 million years ago). The overall phylogeographical pattern was characterized by vicariance events and regional demographical expansion, reflecting a major influence of geological and climatic events on the evolution of Myricaria species. Our data suggest that M. laxiflora has an ancient origin, but has experienced recent population expansion through the Three Gorges Valley. The origin of M. laxiflora was estimated to be during the Early Pleistocene but its demographical expansion was more recent at about 0.015 million years ago. This highlights the unique phylogeographical history of the Three Gorges mountain region, and the deep imprint of the watercourse connections of the Yangtze River Valley on the phylogeographical structure of the species in this region.     

Genetic variation and phylogeography of free-living mouse species (genus Mus) in the Balkans and the Middle East

This work presents a study of the distribution and pattern of variation throughout the ranges of three free-living mouse species of the genus Mus-M. macedonicus, M. spicilegus, and a M. cypriacus - based on sequencing of two segments of the mitochondrial DNA (mtDNA) control region. The study shows a similar level of variability in the three species and suggests their recent population expansion. The highest proportion of variation is found within populations indicating low genetic structuring. Phylogenetic analysis confirms the significant divergence of a mitochondrial lineage of M. macedonicus from Israel, recently described as a new subspecies, M. macedonicus spretoides. Conversely, no genetic hiatus is revealed between European and Asian populations of M. macedonicus macedonicus. Although phylogenetic relationships among M. spicilegus populations could not be unravelled precisely, the results suggest a recent westward expansion of the species. The mtDNA divergence between M. macedonicus and M. spicilegus is 7.3%, suggesting their split between c. 700,000 and 1 million years ago. These dates correspond with a coalescent estimate about 720,000 years ago. On the other hand, M. cypriacus appeared almost twice as divergent from the former species (4.5%) as from the latter (8.8%) suggesting a divergence of c. 430,000-610,000 years ago (coalescent approximately 490,000 years ago) and 830,000-1.2 million years ago (coalescent approximately 780,000 years ago), respectively. Approximate times of population expansion have also been estimated for all taxa and groups of populations. Existence of several glacial refuges and various colonization scenarios are discussed; since all estimated divergence times fall within interglacial periods it seems that climatic oscillations did not play a crucial role in the evolution of the three species.     

Elevated CO2 influences the responses of two birch species to soil moisture: implications for forest community structure

Increased levels of atmospheric CO₂ may alter the structure and composition of plant communities by affecting how species respond to their physical and biological environment. We investigated how elevated CO₂ influenced the response of paper birch ( Betula papyrifera Marsh.) and yellow birch (Betula alleghaniensis Britt.) seedlings to variation in soil moisture. Seedlings were grown for four months on a soil moisture gradient, individually and in mixed species stands, in controlled environment facilities at ambient (375 μL L^–1) and elevated (700 μL L^–1) atmospheric CO₂. For both individually and competitively grown paper birch seedlings, there was a greater CO₂ growth enhancement for seedlings watered less frequently than for well-watered seedlings. This differential change in CO₂ responsiveness across the moisture gradient reduced the difference in seedling growth between high and low water levels and effectively broadened the regeneration niche of paper birch. In contrast, for yellow birch seedlings, elevated CO₂ only produced a significant growth enhancement at the wet end of the soil moisture gradient, and increased the size difference between seedlings at the two ends of the gradient. Gas exchange measurements showed that paper birch seedlings were more sensitive than yellow birch seedlings to declines in soil moisture, and that elevated CO₂ reduced this sensitivity. Additionally, elevated CO₂ improved survival of yellow birch seedlings growing in competition with paper birch in dry stands. Thus, elevated CO₂ may influence regeneration patterns of paper birch and yellow birch on sites of differing soil moisture. In the future, as atmospheric CO₂ levels rise, growth of paper birch seedlings and survival of yellow birch seedlings may be enhanced on xeric sites, while yellow birch may show improved growth on mesic sites.     

Pleistocene population expansions of Antarctic seals

We sequenced a portion ( c . 475 bp) of the mitochondrial control region of three species of Antarctic phocid carnivores (Weddell seal, Leptonychotes weddellii , N  = 181; crabeater seal, Lobodon carcinophaga , N  = 143; and Ross seal, Ommatophoca rossii , N  = 41) that live seasonally or permanently in the fast ice and seasonal pack ice of the western Amundsen and Ross seas of western Antarctica. We resolved 251 haplotypes with a haplotype diversity of 0.98 to 0.99. Bayesian estimates of Θ from the program LAMARC ranged from 0.075 for Weddell seals to 0.576 for crabeater seals. We used the values of theta to estimate female effective population sizes ( N_EF ), which were 40 700 to 63 000 for Weddell seals, 44 400 to 97 800 for Ross seals, and 358 500 to 531 900 for crabeater seals. We used mismatch distributions to test for historical population size expansions. Weddell seals and crabeater seals had significant, unimodal mean pairwise difference distributions ( P  = 0.56 and 0.36, respectively), suggesting that their populations expanded suddenly around 731 000 years ago (Weddell seals) and around 1.6 million years ago (crabeater seals). Both of these expansions occurred during times of intensified glaciations and may have been fostered by expanding pack ice habitat.     

Pollen fates and the limits on male reproductive success in an orchid population

Male reproductive success in higher plants depends largely on the fates of pollen, but current methodologies have given only partial insights into this important aspect of plant mating. We present a detailed analysis of the proportions and absolute amounts of stained pollen involved in six key fates for the hawkmoth-pollinated African orchid Disa cooperi . Despite being packaged into pollinaria, high proportions (> 0.95) of the pollen removed from anthers were lost during transport by hawkmoths in both years. The proportion of pollen lost correlated positively with the number of pollinaria removed from a plant, so that pollen export did not vary with pollen removal. Most pollen was dispersed to neighbouring plants, with rare long-distance dispersal up to 65 m. Of the pollen that reached stigmas during both years, roughly equal amounts were involved in facilitated self-pollination vs. cross-pollination, but the relative proportions of these fates differed between years. Contrary to expectation, we found that self-pollination between flowers did not increase with the number of open flowers, even though moths probed significantly more flowers on larger plants. However, during both years the fraction of removed pollen exported to other plants declined significantly with increasing self-pollination on the source plant, indicating that once self-pollination occurred it reduced (discounted) subsequent pollen export opportunities. The packaging of pollen into pollinaria in orchids appears to increase overall transfer efficiency by at least an order of magnitude relative to plants with granular pollen. Nevertheless, considerable uncertainties remain in the male reproductive success of individual orchids. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 175–190.     

Phylogeny and historical demography of economically important rodents of the genus Arvicanthis (Mammalia: Muridae) from the Nile Valley: of mice and men

The present study clarified the taxonomy, phylogeny and historical demography of semicommensal Nile rats ( Arvicanthis ) from the Nile Valley in Sudan. Nile rats are important crop pests and zoonotic disease reservoirs and are closely associated with agricultural settlements in the Nile Valley. Phylogenetic analysis of the complete cytochrome b gene (1140 bp) of 23 individuals from six localities in Sudan (from two previously recognized species, Arvicanthis niloticus and Arvicanthis testicularis ), supported the existence of only a single species, A. niloticus , from the Nile Valley. Historical demography of the Sudanese Nile Valley population inferred from mismatch coefficients indicated that an exponential population expansion event occurred approximately 144 000–288 000 years ago, corresponding in time with early human expansion and colonization from Africa to the Middle East, Europe, and the world. The inferred high level of gene flow and large size of Sudanese Nile populations of A. niloticus is consistent with historically recent (300 years ago) exponential human population growth and intense agricultural activity inferred from archaeological and historical evidence. Two African Arvicanthis clades were well supported by the broader phylogenetic analysis: (1) A. niloticus , Arvicanthis abyssinicus and Arvicanthis neumanni and (2) Arvicanthis rufinus and Arvicanthis ansorgei from western Africa. Within the first clade, divergence between lineages of A. niloticus s.s. from west and north-east Africa (8.9%) suggests specific recognition, but sampling of geographically intermediate localities is required. Based on hypothesized palaeodrainage and palaeoclimatic patterns, we propose a simple model for speciation of Arvicanthis in Africa.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 641–655.     

Long distance pollen transport cause problems for determining the timing of birch pollen season in Fennoscandia by using phenological observations

The male flowering and leaf bud burst of birch take place almost simultaneously, suggesting that the observations of leaf bud burst could be used to determine the timing of birch pollen release. However, long‐distance transport of birch pollen before the onset of local flowering may complicate the utilization of phenological observations in pollen forecasting.

We compared the timing of leaf bud burst of silver birch with the timing of the stages of birch pollen season during an eight year period (1997–2004) at five sites in Finland. The stages of the birch pollen season were defined using four different thresholds: 1) the first date of the earliest three‐day period with airborne birch pollen counts exceeding 10 grains m^?3 air; and the dates when the accumulated pollen sum reaches 2) 5%; 3) 50% and 4) 95% of the annual total. Atmospheric modelling was used to determine the source areas for the observed long‐distance transported pollen, and the exploitability of phenological observations in pollen forecasting was evaluated.

Pair‐wise comparisons of means indicate that the timing of leaf bud burst fell closest to the date when the accumulated pollen sum reached 5% of the annual total, and did not differ significantly from it at any site (p<0.05; Student‐Newman‐Keuls test). It was found that the timing of leaf bud burst of silver birch overlaps with the first half of the main birch pollen season. However, phenological observations alone do not suffice to determine the timing of the main birch pollen season because of long‐distance transport of birch pollen.     

Ecosystem nitrogen fixation throughout the snow‐free period in subarctic tundra: effects of willow and birch litter addition and warming

Nitrogen (N) fixation in moss‐associated cyanobacteria is one of the main sources of available N for N‐limited ecosystems such as subarctic tundra. Yet, N₂ fixation in mosses is strongly influenced by soil moisture and temperature. Thus, temporal scaling up of low‐frequency in situ measurements to several weeks, months or even the entire growing season without taking into account changes in abiotic conditions cannot capture the variation in moss‐associated N₂ fixation. We therefore aimed to estimate moss‐associated N₂ fixation throughout the snow‐free period in subarctic tundra in field experiments simulating climate change: willow (Salix myrsinifolia) and birch (Betula pubescens spp. tortuosa) litter addition, and warming. To achieve this, we established relationships between measured in situ N₂ fixation rates and soil moisture and soil temperature and used high‐resolution measurements of soil moisture and soil temperature (hourly from May to October) to model N₂ fixation. The modelled N₂ fixation rates were highest in the warmed (2.8 ± 0.3 kg N ha^?1) and birch litter addition plots (2.8 ± 0.2 kg N ha^?1), and lowest in the plots receiving willow litter (1.6 ± 0.2 kg N ha^?1). The control plots had intermediate rates (2.2 ± 0.2 kg N ha^?1). Further, N₂ fixation was highest during the summer in the warmed plots, but was lowest in the litter addition plots during the same period. The temperature and moisture dependence of N₂ fixation was different between the climate change treatments, indicating a shift in the N₂ fixer community. Our findings, using a combined empirical and modelling approach, suggest that a longer snow‐free period and increased temperatures in a future climate will likely lead to higher N₂ fixation rates in mosses. Yet, the consequences of increased litter fall on moss‐associated N₂ fixation due to shrub expansion in the Arctic will depend on the shrub species’ litter traits.