Biomes of western North America at 18,000, 6000 and 0 14C yr bp reconstructed from pollen and packrat midden data

A new compilation of pollen and packrat midden data from western North America provides a refined reconstruction of the composition and distribution of biomes in western North America for today and for 6000 and 18,000 radiocarbon years before present (¹⁴C yr bp ). Modern biomes in western North America are adequately portrayed by pollen assemblages from lakes and bogs. Forest biomes in western North America share many taxa in their pollen spectra and it can be difficult to discriminate among these biomes. Plant macrofossils from packrat middens provide reliable identification of modern biomes from arid and semiarid regions, and this may also be true in similar environments in other parts of the world. However, a weighting factor for trees and shrubs must be used to reliably reconstruct modern biomes from plant macrofossils. A new biome, open conifer woodland, which includes eurythermic conifers and steppe plants, was defined to categorize much of the current and past vegetation of the semiarid interior of western North America. At 6000 ¹⁴C yr bp , the forest biomes of the coastal Pacific North‐west and the desert biomes of the South‐west were in near‐modern positions. Biomes in the interior Pacific North‐west differed from those of today in that taiga prevailed in modern cool/cold mixed forests. Steppe was present in areas occupied today by open conifer woodland in the northern Great Basin, while in the central and southern Rocky Mountains forests grew where steppe grows today. During the mid‐Holocene, cool conifer forests were expanded in the Rocky Mountains (relative to today) but contracted in the Sierra Nevada. These differences from the forests of today imply different climatic histories in these two regions between 6000 ¹⁴C yr bp and today. At 18,000 ¹⁴C yr bp , deserts were absent from the South‐west and the coverage of open conifer woodland was greatly expanded relative to today. Steppe and tundra were present in much of the region now covered by forests in the Pacific North‐west.     

Palaeovegetation of China: a pollen data-based synthesis for the mid-Holocene and last glacial maximum

Pollen data from China for 6000 and 18,000 ¹⁴C yr bp were compiled and used to reconstruct palaeovegetation patterns, using complete taxon lists where possible and a biomization procedure that entailed the assignment of 645 pollen taxa to plant functional types. A set of 658 modern pollen samples spanning all biomes and regions provided a comprehensive test for this procedure and showed convincing agreement between reconstructed biomes and present natural vegetation types, both geographically and in terms of the elevation gradients in mountain regions of north‐eastern and south‐western China. The 6000 ¹⁴C yr bp map confirms earlier studies in showing that the forest biomes in eastern China were systematically shifted northwards and extended westwards during the mid‐Holocene. Tropical rain forest occurred on mainland China at sites characterized today by either tropical seasonal or broadleaved evergreen/warm mixed forest. Broadleaved evergreen/warm mixed forest occurred further north than today, and at higher elevation sites within the modern latitudinal range of this biome. The northern limit of temperate deciduous forest was shifted c. 800 km north relative to today. The 18,000 ¹⁴C yr bp map shows that steppe and even desert vegetation extended to the modern coast of eastern China at the last glacial maximum, replacing today’s temperate deciduous forest. Tropical forests were excluded from China and broadleaved evergreen/warm mixed forest had retreated to tropical latitudes, while taiga extended southwards to c. 43°N.     

Pollen-based reconstructions of Japanese biomes at 0, 6000 and 18,000 14C yr bp

A biomization method, which objectively assigns individual pollen assemblages to biomes ( Prentice et al., 1996 ), was tested using modern pollen data from Japan and applied to fossil pollen data to reconstruct palaeovegetation patterns 6000 and 18,000 ¹⁴C yr bp Biomization started with the assignment of 135 pollen taxa to plant functional types (PFTs), and nine possible biomes were defined by specific combinations of PFTs. Biomes were correctly assigned to 54% of the 94 modern sites. Incorrect assignments occur near the altitudinal limits of individual biomes, where pollen transport from lower altitudes blurs the local pollen signals or continuous changes in species composition characterizes the range limits of biomes. As a result, the reconstructed changes in the altitudinal limits of biomes at 6000 and 18,000 ¹⁴C yr bp are likely to be conservative estimates of the actual changes. The biome distribution at 6000 ¹⁴C yr bp was rather similar to today, suggesting that changes in the bioclimate of Japan have been small since the mid‐Holocene. At 18,000 ¹⁴C yr bp the Japanese lowlands were covered by taiga and cool mixed forests. The southward expansion of these forests and the absence of broadleaved evergreen/warm mixed forests reflect a pronounced year‐round cooling.     

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa

BIOME 6000 is an international project to map vegetation globally at mid‐Holocene (6000 ¹⁴C yr bp ) and last glacial maximum (LGM, 18,000 ¹⁴C yr bp ), with a view to evaluating coupled climate‐biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site‐based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method’s skill in reconstructing present‐day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south‐western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 ¹⁴C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial‐interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now‐arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land‐surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere‐biosphere models. The data could also be objectively generalized to yield realistic gridded land‐surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation‐climate feedbacks have focused on the hypothesized positive feedback effects of climate‐induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid‐Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 ¹⁴C yr bp and for the LGM.     

Pollen-based biome reconstruction for southern Europe and Africa 18,000 yr bp

Pollen data from 18,000 ¹⁴C yr bp were compiled in order to reconstruct biome distributions at the last glacial maximum in southern Europe and Africa. Biome reconstructions were made using the objective biomization method applied to pollen counts using a complete list of dryland taxa wherever possible. Consistent and major differences from present‐day biomes are shown. Forest and xerophytic woods/scrub were replaced by steppe, both in the Mediterranean region and in southern Africa, except in south‐western Cape Province where fynbos (xerophytic scrub) persisted. Sites in the tropical highlands, characterized today by evergreen forest, were dominated by steppe and/or xerophytic vegetation (cf. today’s Ericaceous belt and Afroalpine grassland) at the last glacial maximum. Available data from the tropical lowlands are sparse but suggest that the modern tropical rain forest was largely replaced by tropical seasonal forest while the modern seasonal or dry forests were encroached on by savanna or steppe. Montane forest elements descended to lower elevations than today.     

Last glacial maximum biomes reconstructed from pollen and plant macrofossil data from northern Eurasia

Pollen and plant macrofossil data from northern Eurasia were used to reconstruct the vegetation of the last glacial maximum (LGM: 18,000 ± 2000 ¹⁴C yr bp ) using an objective quantitative method for interpreting pollen data in terms of the biomes they represent ( Prentice et al., 1996 ). The results confirm previous qualitative vegetation reconstructions at the LGM but provide a more comprehensive analysis of the data. Tundra dominated a large area of northern Eurasia (north of 57°N) to the west, south and east of the Scandinavian ice sheet at the LGM. Steppe‐like vegetation was reconstructed in the latitudinal band from western Ukraine, where temperate deciduous forests grow today, to western Siberia, where taiga and cold deciduous forests grow today. The reconstruction shows that steppe graded into tundra in Siberia, which is not the case today. Taiga grew on the northern coast of the Sea of Azov, about 1500 km south of its present limit in European Russia. In contrast, taiga was reconstructed only slightly south of its southern limit today in south‐western Siberia. Broadleaved trees were confined to small refuges, e.g. on the eastern coast of the Black Sea, where cool mixed forest was reconstructed from the LGM data. Cool conifer forests in western Georgia were reconstructed as growing more than 1000 m lower than they grow today. The few scattered sites with LGM data from the Tien‐Shan Mountains and from northern Mongolia yielded biome reconstructions of steppe and taiga, which are the biomes growing there today.     

Pollen‐based biomes for Beringia 18,000, 6000 and 0 14C yr bp†

The objective biomization method developed by Prentice et al. (1996) for Europe was extended using modern pollen samples from Beringia and then applied to fossil pollen data to reconstruct palaeovegetation patterns at 6000 and 18,000 ¹⁴C yr bp . The predicted modern distribution of tundra, taiga and cool conifer forests in Alaska and north‐western Canada generally corresponds well to actual vegetation patterns, although sites in regions characterized today by a mosaic of forest and tundra vegetation tend to be preferentially assigned to tundra. Siberian larch forests are delimited less well, probably due to the extreme under‐representation of Larix in pollen spectra. The biome distribution across Beringia at 6000 ¹⁴C yr bp was broadly similar to today, with little change in the northern forest limit, except for a possible northward advance in the Mackenzie delta region. The western forest limit in Alaska was probably east of its modern position. At 18,000 ¹⁴C yr bp the whole of Beringia was covered by tundra. However, the importance of the various plant functional types varied from site to site, supporting the idea that the vegetation cover was a mosaic of different tundra types.     

Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0, 6000 and 18,000 14C yr BP

Aim This paper documents reconstructions of the vegetation patterns in Australia, Southeast Asia and the Pacific (SEAPAC region) in the mid‐Holocene and at the last glacial maximum (LGM). Methods Vegetation patterns were reconstructed from pollen data using an objective biomization scheme based on plant functional types. The biomization scheme was first tested using 535 modern pollen samples from 377 sites, and then applied unchanged to fossil pollen samples dating to 6000 ± 500 or 18,000 ± 1000 ¹⁴C yr bp . Results 1. Tests using surface pollen sample sites showed that the biomization scheme is capable of reproducing the modern broad‐scale patterns of vegetation distribution. The north–south gradient in temperature, reflected in transitions from cool evergreen needleleaf forest in the extreme south through temperate rain forest or wet sclerophyll forest (WSFW) and into tropical forests, is well reconstructed. The transitions from xerophytic through sclerophyll woodlands and open forests to closed‐canopy forests, which reflect the gradient in plant available moisture from the continental interior towards the coast, are reconstructed with less geographical precision but nevertheless the broad‐scale pattern emerges. 2. Differences between the modern and mid‐Holocene vegetation patterns in mainland Australia are comparatively small and reflect changes in moisture availability rather than temperature. In south‐eastern Australia some sites show a shift towards more moisture‐stressed vegetation in the mid‐Holocene with xerophytic woods/scrub and temperate sclerophyll woodland and shrubland at sites characterized today by WSFW or warm‐temperate rain forest (WTRF). However, sites in the Snowy Mountains, on the Southern Tablelands and east of the Great Dividing Range have more moisture‐demanding vegetation in the mid‐Holocene than today. South‐western Australia was slightly drier than today. The single site in north‐western Australia also shows conditions drier than today in the mid‐Holocene. Changes in the tropics are also comparatively small, but the presence of WTRF and tropical deciduous broadleaf forest and woodland in the mid‐Holocene, in sites occupied today by cool‐temperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic vegetation in the south and tropical deciduous broadleaf forest and woodland in the north indicate drier conditions across mainland Australia at the LGM. None of these changes are informative about the degree of cooling. However the evidence from the tropics, showing lowering of the treeline and forest belts, indicates that conditions were between 1 and 9 °C (depending on elevation) colder. The encroachment of tropical deciduous broadleaf forest and woodland into lowland evergreen broadleaf forest implies greater aridity. Main conclusions This study provides the first continental‐scale reconstruction of mid‐Holocene and LGM vegetation patterns from Australia, Southeast Asia and the Pacific (SEAPAC region) using an objective biomization scheme. These data will provide a benchmark for evaluation of palaeoclimate simulations within the framework of the Palaeoclimate Modelling Intercomparison Project.     

Ploidy race distributions since the Last Glacial Maximum in the North American desert shrub, Larrea tridentata

• 1 A classic biogeographic pattern is the alignment of diploid, tetraploid and hexaploid races of creosote bush (Larrea tridentata) across the Chihuahuan, Sonoran and Mohave Deserts of western North America. We used statistically robust differences in guard cell size of modern plants and fossil leaves from packrat middens to map current and past distributions of these ploidy races since the Last Glacial Maximum (LGM).
• 2 Glacial/early Holocene (26–10 ¹⁴C kyr bp or thousands of radiocarbon years before present) populations included diploids along the lower Rio Grande of west Texas, 650 km removed from sympatric diploids and tetraploids in the lower Colorado River Basin of south‐eastern California/south‐western Arizona. Diploids migrated slowly from lower Rio Grande refugia with expansion into the northern Chihuahuan Desert sites forestalled until after ~4.0 ¹⁴C kyr bp . Tetraploids expanded from the lower Colorado River Basin into the northern limits of the Sonoran Desert in central Arizona by 6.4 ¹⁴C kyr bp . Hexaploids appeared by 8.5 ¹⁴C kyr bp in the lower Colorado River Basin, reaching their northernmost limits (~37°N) in the Mohave Desert between 5.6 and 3.9 ¹⁴C kyr bp .
• 3 Modern diploid isolates may have resulted from both vicariant and dispersal events. In central Baja California and the lower Colorado River Basin, modern diploids probably originated from relict populations near glacial refugia. Founder events in the middle and late Holocene established diploid outposts on isolated limestone outcrops in areas of central and southern Arizona dominated by tetraploid populations.
• 4 Geographic alignment of the three ploidy races along the modern gradient of increasingly drier and hotter summers is clearly a postglacial phenomenon, but evolution of both higher ploidy races must have happened before the Holocene. The exact timing and mechanism of polyploidy evolution in creosote bush remains a matter of conjecture.

     

Carbon stock loss from deforestation through 2013 in Brazilian Amazonia

The largest carbon stock in tropical vegetation is in Brazilian Amazonia. In this ~5 million km² area, over 750 000 km² of forest and ~240 000 km² of nonforest vegetation types had been cleared through 2013. We estimate current carbon stocks and cumulative gross carbon loss from clearing of premodern vegetation in Brazil's ‘Legal Amazonia’ and ‘Amazonia biome’ regions. Biomass of ‘premodern’ vegetation (prior to major increases in disturbance beginning in the 1970s) was estimated by matching vegetation classes mapped at a scale of 1 : 250 000 and 29 biomass means from 41 published studies for vegetation types classified as forest (2317 1‐ha plots) and as either nonforest or contact zones (1830 plots and subplots of varied size). Total biomass (above and below‐ground, dry weight) underwent a gross reduction of 18.3% in Legal Amazonia (13.1 Pg C) and 16.7% in the Amazonia biome (11.2 Pg C) through 2013, excluding carbon loss from the effects of fragmentation, selective logging, fires, mortality induced by recent droughts and clearing of forest regrowth. In spite of the loss of carbon from clearing, large amounts of carbon were stored in stands of remaining vegetation in 2013, equivalent to 149 Mg C ha^?1 when weighted by the total area covered by each vegetation type in Legal Amazonia. Native vegetation in Legal Amazonia in 2013 originally contained 58.6 Pg C, while that in the Amazonia biome contained 56 Pg C. Emissions per unit area from clearing could potentially be larger in the future because previously cleared areas were mainly covered by vegetation with lower mean biomass than the remaining vegetation. Estimates of original biomass are essential for estimating losses to forest degradation. This study offers estimates of cumulative biomass loss, as well as estimates of premodern carbon stocks that have not been represented in recent estimates of deforestation impacts.     

A high‐resolution approach to estimating ecosystem respiration at continental scales using operational satellite data

A better understanding of the local variability in land‐atmosphere carbon fluxes is crucial to improving the accuracy of global carbon budgets. Operational satellite data backed by ground measurements at Fluxnet sites proved valuable in monitoring local variability of gross primary production at highly resolved spatio‐temporal resolutions. Yet, we lack similar operational estimates of ecosystem respiration (Re) to calculate net carbon fluxes. If successful, carbon fluxes from such a remote sensing approach would form an independent and sought after measure to complement widely used dynamic global vegetation models (DGVMs). Here, we establish an operational semi‐empirical Re model, based only on data from the Moderate Resolution Imaging Spectroradiometer (MODIS) with a resolution of 1 km and 8 days. Fluxnet measurements between 2000 and 2009 from 100 sites across North America and Europe are used for parameterization and validation. Our analysis shows that Re is closely tied to temperature and plant productivity. By separating temporal and intersite variation, we find that MODIS land surface temperature (LST) and enhanced vegetation index (EVI) are sufficient to explain observed Re across most major biomes with a negligible bias [R² = 0.62, RMSE = 1.32 (g C m^?2 d^?1), MBE = 0.05 (g C m^?2 d^?1)]. A comparison of such satellite‐derived Re with those simulated by the DGVM LPJmL reveals similar spatial patterns. However, LPJmL shows higher temperature sensitivities and consistently simulates higher Re values, in high‐latitude and subtropical regions. These differences remain difficult to explain and they are likely associated either with LPJmL parameterization or with systematic errors in the Fluxnet sampling technique. While uncertainties remain with Re estimates, the model formulated in this study provides an operational, cross‐validated and unbiased approach to scale Fluxnet Re to the continental scale and advances knowledge of spatio‐temporal Re variability.     

Sensitivity of African biomes to changes in the precipitation regime

Aim Africa is identified by the Inter‐governmental Panel on Climate Change (IPCC) as the least studied continent in terms of ecosystem dynamics and climate variability. The aim of this study was (1) to adapt the Lund‐Postdam‐Jena‐GUESS (LPJ‐GUESS) ecological modelling framework to Africa by providing new parameter values for tropical plant functional types (PFT), and (2) to assess the sensitivity of some African biomes to changes in precipitation regime. Location The study area was a representative transect (0–22° N and 7–18° E) through the transition from equatorial evergreen forests to savannas, steppes and desert northwards. The transect showed large latitudinal variation in precipitation (mean rainfall ranged from 50 to 2300 mm year^?1). Methods New PFT parameters used to calibrate LPJ‐GUESS were based on modern pollen PFTs and remote sensed leaf area index (LAI). The model was validated using independent modern pollen assemblages, LAI and through comparison with White's modern potential vegetation map. Several scenarios were developed by combining changes in total rainfall amount with variation in the length of the dry season in order to test the sensitivity of African biomes. Results Simulated vegetation compared well to observed data at local and regional scales, in terms of ecosystem functioning (LAI), and composition (pollen and White's vegetation map). The assessment of the sensitivity of biomes to changes in precipitation showed that none of the ecosystems would shift towards a new type under the range of precipitation increases suggested by the IPCC (increases from 5 to 20%). However, deciduous and semi‐deciduous forests may be very sensitive to small reductions in both the amount and seasonality of precipitation. Main conclusions This version of LPJ‐GUESS parameterized for Africa simulated correctly the vegetation present over a wide precipitation gradient. The biome sensitivity assessment showed that, compared with savannas and grasslands, closed canopy forests may be more sensitive to change in precipitation regime due to the synergetic effects of changed rainfall amounts and seasonality on vegetation functioning.     

Modern pollen-based biome reconstructions in East Africa expanded to southern Tanzania

New detailed biome reconstructions are proposed in East Africa from modern pollen data derived from 150 sites located in northern Kenya (40 sites), north-western Uganda (51 sites) and southern Tanzania (59 new sites presented as pollen diagram), which are representative of the major vegetation associations occurring in seven phytogeographical regions, mosaics or centres of endemism. We use the standard biomisation method previously published for the African continent, but we reconsider the taxa assignment to plant functional types. We include in this approach all identified taxa (408) except aquatics, ferns and exotic taxa. The method is validated by comparison with local vegetation data and we show that 124 (82.6%) sites are assigned to the correct biome and that for all the biomes under investigation, the number of correct assignments always exceeds the number of incorrect ones. When an incorrect biome reconstruction occurs, mainly toward drier biomes, this is generally linked to the local open/degraded structure of the original vegetation or to the occurrence of a mosaic of open/closed vegetation. In turn, most of the reconstructions of more humid/closed biomes than the corresponding local vegetation (8.6%) remain unexplained. A comparison of our reconstructed biomes with the main East African vegetation types of White's map indicates that 121 (80.6%) sites are assigned to the correct biomes. However, the majority of sites are incorrectly reconstructed compared to Olson and IGBP maps from satellite data, mainly due to incorrect allocation of the land cover classes compared to the potential vegetation. The application of this method to our pollen data set demonstrates that modern pollen assemblages can successfully reconstruct the main modern East African vegetation types.     

Post-Hypsithermal plant disjunctions in western Alberta, Canada

Aim Evaluate the hypothesis that nine disjunct vascular plant species along the eastern slopes of the Rocky Mountains and in the Peace River District of west‐central Alberta represent remnants of more southerly vegetation that occupied these areas during the Holocene Hypsithermal (9000–6000 yr bp ). Alternatively, these plants represent populations that became established because of independent chance dispersal events. Location This study focuses on the area east of the Rocky Mountain Continental Divide in the Province of Alberta and the State of Montana in western Canada and USA, respectively. Methods Disjunct species were identified and their distributions mapped based on a review of occurrence maps and records, botanical floras and checklists, herbaria specimens, ecological and botanical studies, and field surveys of selected species. A disjunct species was defined as a plant population separated from its next nearest occurrence by a distance of > 300 km. Evaluation of the hypothesis was based on a review of published and unpublished pollen stratigraphy and palaeoecological studies. The potential geographical distribution of Hypsithermal vegetation was based on modern regional‐based ecosystem mapping and associated monthly temperature summaries as well as future climatic warming models. Results The hypothesis was compatible with Holocene pollen stratigraphy, Hypsithermal permafrost and fen occurrence, and palaeosol phytolith analyses; and future global climatic warming models. Modelled regional Hypsithermal vegetation based on a 1 °C increase in July temperatures relative to current conditions, indicated that much of the boreal forest zone in Alberta could have been grassland, which would explain the occurrence of Prairie species in the Peace River District. This amount of latitudinal vegetation shift (6.5°) was similar to an earlier Hypsithermal permafrost zone location study. An equivalent shift in vegetation along the eastern Cordillera would have placed south‐western Montana‐like vegetation and species such as Boykinia heucheriformis (Rydb.) Rosend. and Saxifraga odontoloma Piper within the northern half of the Rocky Mountains and foothills in Alberta, which represents the location of modern‐day disjunct populations of these species. Main conclusions Warmer and drier climatic conditions during the Holocene Hypsithermal resulted in the northward displacement of vegetation zones relative to their current distribution patterns. Most of Alberta was probably dominated by grasslands during this period, except the Rocky Mountains and northern highlands. Modern‐day species disjunctions within the Rocky Mountains and Peace River District as well as more northerly areas such as the Yukon Territory occurred when the vegetation receded southward in response to climatic cooling after the Hypsithermal. Wind dispersal was considered an unlikely possibility to explain the occurrence of the disjunct species, as most of the plants lack morphological adaptations for long distance transport and the prevailing winds were from west to east rather than south to north. However, consumption and transport of seeds by northward migrating birds could not be excluded as a possibility.     

Terrestrial biosphere models need better representation of vegetation phenology: results from the North American Carbon Program Site Synthesis

Phenology, by controlling the seasonal activity of vegetation on the land surface, plays a fundamental role in regulating photosynthesis and other ecosystem processes, as well as competitive interactions and feedbacks to the climate system. We conducted an analysis to evaluate the representation of phenology, and the associated seasonality of ecosystem‐scale CO₂ exchange, in 14 models participating in the North American Carbon Program Site Synthesis. Model predictions were evaluated using long‐term measurements (emphasizing the period 2000–2006) from 10 forested sites within the AmeriFlux and Fluxnet‐Canada networks. In deciduous forests, almost all models consistently predicted that the growing season started earlier, and ended later, than was actually observed; biases of 2 weeks or more were typical. For these sites, most models were also unable to explain more than a small fraction of the observed interannual variability in phenological transition dates. Finally, for deciduous forests, misrepresentation of the seasonal cycle resulted in over‐prediction of gross ecosystem photosynthesis by +160 ± 145 g C m^?2 yr^?1 during the spring transition period and +75 ± 130 g C m^?2 yr^?1 during the autumn transition period (13% and 8% annual productivity, respectively) compensating for the tendency of most models to under‐predict the magnitude of peak summertime photosynthetic rates. Models did a better job of predicting the seasonality of CO₂ exchange for evergreen forests. These results highlight the need for improved understanding of the environmental controls on vegetation phenology and incorporation of this knowledge into better phenological models. Existing models are unlikely to predict future responses of phenology to climate change accurately and therefore will misrepresent the seasonality and interannual variability of key biosphere–atmosphere feedbacks and interactions in coupled global climate models.     

Late-glacial and Holocene palaeovegetation zonal reconstruction for central and north-central North America

Aim The purpose of this study is to develop palaeovegetation zonation models for central and north‐central North America, based on late‐Quaternary and Holocene pollen stratigraphic data (n = 246 sites). A secondary purpose was to evaluate an hypothesis ( Strong & Hills, 2003 ) to explain the disjunct distribution of species in western Alberta. Location Hudson Bay‐Lake Michigan to the Rocky Mountains region, north of 36° N to the Arctic Ocean (c. 70° N). Methods Pollen profiles spanning 40 years of palaeoecological research in North America were extracted from published and unpublished archival sources. Individual profiles were subdivided into 1000‐year increments based on the assumption of a constant sedimentation rate between stratigraphic dates (e.g. surface sediments, radiocarbon ¹⁴C dates, tephra layers). The pollen composition among profiles was standardized to 54 commonly recognized taxa, with percentage composition within each stratigraphic sample prorated to 100% prior to analysis. Near‐surface sediments from these profiles were included as analogues of modern vegetation. Cluster analysis was used as a guide to the classification of 2356 temporal stratigraphic samples, which resulted in the recognition of 16 pollen groups. These groups were summarized in terms of their pollen composition, mapped, and used in combination with terrain information and an ecological knowledge of the study area to construct six physiognomically‐based palaeovegetation zonation models at 2000‐year intervals from 14,000 to 4000 yr bp (radiocarbon years before present). Results The 14,000 yr bp model placed Boreal and Cordilleran Forests proximal to the southern glacial front, whereas Arctic tundra dominated the Yukon Territory–Alaska ice‐free zone. Pollen and macrofossil evidence suggests that this Boreal Forest zone contained a mixture of coniferous and deciduous tree species. Grassland was postulated immediately south of the forest zone, with its northern extreme near 49° N latitude in the Alberta–Montana border area. Separation of the Laurentide and Cordilleran glacial fronts about 12,000 yr bp initiated the northward advance of Boreal Forests into western Canada. By the end of the Hypsithermal at about 6000 yr bp , Boreal Forests occurred near the Arctic Ocean, and Grassland and Aspen Parkland zones may have extended to 54° N and 59° N latitude in Alberta, respectively. Between 6000 and 4000 yr bp , a 5° and 1° latitudinal southward shift of the northern Boreal Forest and Grassland/Aspen Parkland boundaries occurred, respectively, near their contemporary positions with corresponding expansions of the Subarctic and Arctic zones. Modern Canadian Cordilleran Forests along the eastern slopes of the Rocky Mountains were interpreted as originating from the north‐central Montana–south‐western Alberta area. Jack pine (Pinus banksiana Lamb.), a common Boreal Forest species, appears to have entered central Canada via the north side of Lake Superior after 11,000 yr bp . Main conclusions Modern vegetation in central Canada evolved from biomes located in the northern USA during the late‐Quaternary. The Boreal Forest biome contained the same arboreal taxa as the modern vegetation, except it lacked jack pine. The proposed regional palaeovegetation models support the hypothesis of Strong & Hills (2003) , but new independent palaeoecological data will be needed for a proper evaluation.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

Carbon pool densities and a first estimate of the total carbon pool in the Mongolian forest‐steppe

The boreal forest biome represents one of the most important terrestrial carbon stores, which gave reason to intensive research on carbon stock densities. However, such an analysis does not yet exist for the southernmost Eurosiberian boreal forests in Inner Asia. Most of these forests are located in the Mongolian forest‐steppe, which is largely dominated by Larix sibirica. We quantified the carbon stock density and total carbon pool of Mongolia's boreal forests and adjacent grasslands and draw conclusions on possible future change. Mean aboveground carbon stock density in the interior of L. sibirica forests was 66 Mg C ha^?1, which is in the upper range of values reported from boreal forests and probably due to the comparably long growing season. The density of soil organic carbon (SOC, 108 Mg C ha^?1) and total belowground carbon density (149 Mg C ha^?1) are at the lower end of the range known from boreal forests, which might be the result of higher soil temperatures and a thinner permafrost layer than in the central and northern boreal forest belt. Land use effects are especially relevant at forest edges, where mean carbon stock density was 188 Mg C ha^?1, compared with 215 Mg C ha^?1 in the forest interior. Carbon stock density in grasslands was 144 Mg C ha^?1. Analysis of satellite imagery of the highly fragmented forest area in the forest‐steppe zone showed that Mongolia's total boreal forest area is currently 73 818 km², and 22% of this area refers to forest edges (defined as the first 30 m from the edge). The total forest carbon pool of Mongolia was estimated at ~ 1.5?1.7 Pg C, a value which is likely to decrease in future with increasing deforestation and fire frequency, and global warming.     

Grassland to shrubland state transitions enhance carbon sequestration in the northern Chihuahuan Desert

The replacement of native C₄‐dominated grassland by C₃‐dominated shrubland is considered an ecological state transition where different ecological communities can exist under similar environmental conditions. These state transitions are occurring globally, and may be exacerbated by climate change. One consequence of the global increase in woody vegetation may be enhanced ecosystem carbon sequestration, although the responses of arid and semiarid ecosystems may be highly variable. During a drier than average period from 2007 to 2011 in the northern Chihuahuan Desert, we found established shrubland to sequester 49 g C m^?2 yr^?1 on average, while nearby native C₄ grassland was a net source of 31 g C m^?2 yr^?1 over this same period. Differences in C exchange between these ecosystems were pronounced – grassland had similar productivity compared to shrubland but experienced higher C efflux via ecosystem respiration, while shrubland was a consistent C sink because of a longer growing season and lower ecosystem respiration. At daily timescales, rates of carbon exchange were more sensitive to soil moisture variation in grassland than shrubland, such that grassland had a net uptake of C when wet but lost C when dry. Thus, even under unfavorable, drier than average climate conditions, the state transition from grassland to shrubland resulted in a substantial increase in terrestrial C sequestration. These results illustrate the inherent tradeoffs in quantifying ecosystem services that result from ecological state transitions, such as shrub encroachment. In this case, the deleterious changes to ecosystem services often linked to grassland to shrubland state transitions may at least be partially offset by increased ecosystem carbon sequestration.     

Carbon accumulation in a permafrost polygon peatland: steady long‐term rates in spite of shifts between dry and wet conditions

Ice‐wedge polygon peatlands contain a substantial part of the carbon stored in permafrost soils. However, little is known about their long‐term carbon accumulation rates (CAR) in relation to shifts in vegetation and climate. We collected four peat profiles from one single polygon in NE Yakutia and cut them into contiguous 0.5 cm slices. Pollen density interpolation between AMS ¹⁴C dated levels provided the time span contained in each of the sample slices, which – in combination with the volumetric carbon content – allowed for the reconstruction of CAR over decadal and centennial timescales. Vegetation representing dry palaeo‐ridges and wet depressions was reconstructed with detailed micro‐ and macrofossil analysis. We found repeated shifts between wet and dry conditions during the past millennium. Dry ridges with associated permafrost growth originated during phases of (relatively) warm summer temperature and collapsed during relatively cold phases, illustrating the important role of vegetation and peat as intermediaries between ambient air temperature and the permafrost. The average long‐term CAR across the four profiles was 10.6 ± 5.5 g C m^?2 yr^?1. Time‐weighted mean CAR did not differ significantly between wet depression and dry ridge/hummock phases (10.6 ± 5.2 g C m^?2 yr^?1 and 10.3 ± 5.7 g C m^?2 yr^?1, respectively). Although we observed increased CAR in relation to warm shifts, we also found changes in the opposite direction and the highest CAR actually occurred during the Little Ice Age. In fact, CAR rather seems to be governed by strong internal feedback mechanisms and has roughly remained stable on centennial time scales. The absence of significant differences in CAR between dry ridge and wet depression phases suggests that recent warming and associated expansion of shrubs will not affect long‐term rates of carbon burial in ice‐wedge polygon peatlands.