Ecophysiological and bioclimatic foundations for a global plant functional classification

Question: What plant properties might define plant functional types (PFTs) for the analysis of global vegetation responses to climate change, and what aspects of the physical environment might be expected to predict the distributions of PFTs? Methods: We review principles to explain the distribution of key plant traits as a function of bioclimatic variables. We focus on those whole-plant and leaf traits that are commonly used to define biomes and PFTs in global maps and models. Results: Raunkiær's plant life forms (underlying most later classifications) describe different adaptive strategies for surviving low temperature or drought, while satisfying requirements for reproduction and growth. Simple conceptual models and published observations are used to quantify the adaptive significance of leaf size for temperature regulation, leaf consistency for maintaining transpiration under drought, and phenology for the optimization of annual carbon balance. A new compilation of experimental data supports the functional definition of tropical, warm-temperate, temperate and boreal phanerophytes based on mechanisms for withstanding low temperature extremes. Chilling requirements are less well quantified, but are a necessary adjunct to cold tolerance. Functional traits generally confer both advantages and restrictions; the existence of trade-offs contributes to the diversity of plants along bioclimatic gradients. Conclusions: Quantitative analysis of plant trait distributions against bioclimatic variables is becoming possible; this opens up new opportunities for PFT classification. A PFT classification based on bioclimatic responses will need to be enhanced by information on traits related to competition, successional dynamics and disturbance.

Evaluation of coupled ocean-atmosphere simulations of northern hemisphere extratropical climates in the mid-Holocene

We have used the BIOME4 biogeography–biochemistry model and comparison with palaeovegetation data to evaluate the response of six ocean–atmosphere general circulation models to mid-Holocene changes in orbital forcing in the mid- to high-latitudes of the northern hemisphere. All the models produce: (a) a northward shift of the northern limit of boreal forest, in response to simulated summer warming in high-latitudes. The northward shift is markedly asymmetric, with larger shifts in Eurasia than in North America; (b) an expansion of xerophytic vegetation in mid-continental North America and Eurasia, in response to increased temperatures during the growing season; (c) a northward expansion of temperate forests in eastern North America, in response to simulated winter warming. The northward shift of the northern limit of boreal forest and the northward expansion of temperate forests in North America are supported by palaeovegetation data. The expansion of xerophytic vegetation in mid-continental North America is consistent with palaeodata, although the extent may be over-estimated. The simulated expansion of xerophytic vegetation in Eurasia is not supported by the data. Analysis of an asynchronous coupling of one model to an equilibrium-vegetation model suggests vegetation feedback exacerbates this mid-continental drying and produces conditions more unlike the observations. Not all features of the simulations are robust: some models produce winter warming over Europe while others produce winter cooling. As a result, some models show a northward shift of temperate forests (consistent with, though less marked than, the expansion shown by data) and others produce a reduction in temperate forests. Elucidation of the cause of such differences is a focus of the current phase of the Palaeoclimate Modelling Intercomparison Project.

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

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.

Synergistic feedbacks between ocean and vegetation on mid- and high-latitude climates during the mid-Holocene

Simulations with the IPSL atmosphere–ocean model asynchronously coupled with the BIOME1 vegetation model show the impact of ocean and vegetation feedbacks, and their synergy, on mid- and high-latitude (>40°N) climate in response to orbitally-induced changes in mid-Holocene insolation. The atmospheric response to orbital forcing produces a +1.2 °C warming over the continents in summer and a cooling during the rest of the year. Ocean feedback reinforces the cooling in spring but counteracts the autumn and winter cooling. Vegetation feedback produces warming in all seasons, with largest changes (+1 °C) in spring. Synergy between ocean and vegetation feedbacks leads to further warming, which can be as large as the independent impact of these feedbacks. The combination of these effects causes the high northern latitudes to be warmer throughout the year in the ocean–atmosphere-vegetation simulation. Simulated vegetation changes resulting from this year-round warming are consistent with observed mid-Holocene vegetation patterns. Feedbacks also impact on precipitation. The atmospheric response to orbital-forcing reduces precipitation throughout the year; the most marked changes occur in the mid-latitudes in summer. Ocean feedback reduces aridity during autumn, winter and spring, but does not affect summer precipitation. Vegetation feedback increases spring precipitation but amplifies summer drying. Synergy between the feedbacks increases precipitation in autumn, winter and spring, and reduces precipitation in summer. The combined changes amplify the seasonal contrast in precipitation in the ocean–atmosphere-vegetation simulation. Enhanced summer drought produces an unrealistically large expansion of temperate grasslands, particularly in mid-latitude Eurasia.

Climate and CO2 controls on global vegetation distribution at the last glacial maximum: analysis based on palaeovegetation data, biome modelling and palaeoclimate simulations.

The global vegetation response to climate and atmospheric CO2 changes between the last glacial maximum and recent times is examined using an equilibrium vegetation model (BIOME4), driven by output from 17 climate simulations from the Palaeoclimate Modelling Intercomparison Project. Features common to all of the simulations include expansion of treeless vegetation in high northern latitudes; southward displacement and fragmentation of boreal and temperate forests; and expansion of drought-tolerant biomes in the tropics. These features are broadly consistent with pollen-based reconstructions of vegetation distribution at the last glacial maximum. Glacial vegetation in high latitudes reflects cold and dry conditions due to the low CO2 concentration and the presence of large continental ice sheets. The extent of drought-tolerant vegetation in tropical and subtropical latitudes reflects a generally drier low-latitude climate. Comparisons of the observations with BIOME4 simulations, with and without consideration of the direct physiological effect of CO2 concentration on C3 photosynthesis, suggest an important additional role of low CO2 concentration in restricting the extent of forests, especially in the tropics. Global forest cover was overestimated by all models when climate change alone was used to drive BIOME4, and estimated more accurately when physiological effects of CO2 concentration were included. This result suggests that both CO2 effects and climate effects were important in determining glacial-interglacial changes in vegetation. More realistic simulations of glacial vegetation and climate will need to take into account the feedback effects of these structural and physiological changes on the climate.

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.

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.

Present-day and mid-Holocene biomes reconstructed from pollen and plant macrofossil data from the former Soviet Union and Mongolia

Fossil pollen data supplemented by tree macrofossil records were used to reconstruct the vegetation of the Former Soviet Union and Mongolia at 6000 years. Pollen spectra were assigned to biomes using the plant-functional-type method developed by Prentice et al. (1996). Surface pollen data and a modern vegetation map provided a test of the method. This is the first time such a broad-scale vegetation reconstruction for the greater part of northern Eurasia has been attempted with objective techniques. The new results confirm previous regional palaeoenvironmental studies of the mid-Holocene while providing a comprehensive synopsis and firmer conclusions. West of the Ural Mountains temperate deciduous forest extended both northward and southward from its modern range. The northern limits of cool mixed and cool conifer forests were also further north than present. Taiga was reduced in European Russia, but was extended into Yakutia where now there is cold deciduous forest. The northern limit of taiga was extended (as shown by increased Picea pollen percentages, and by tree macrofossil records north of the present-day forest limit) but tundra was still present in north-eastern Siberia. The boundary between forest and steppe in the continental interior did not shift substantially, and dry conditions similar to present existed in western Mongolia and north of the Aral Sea.

Pollen-based biome reconstructions for China at 0 and 6000 years

Biomization provides an objective and robust method of assigning pollen spectra to biomes so that pollen data can be mapped and compared directly with the output of biomgeographic models. We have tested the applicability of this procedure, originally developed for Europe, to assign modern surface samples from China to biomes. The procedure successfully delineated the major vegetation types of China. When the same procedure was applied to fossil pollen samples for 6000 years ago, the reconstructions showed systematic differences from present, consistent with previous interpretations of vegetation changes since the mid-Holocene. In eastern China, the forest zones were systematically shifted northwards, such that cool mixed forests displaced taiga in northeastern China, while broad-leaved evergreen forest extended c. 300 km and temperate deciduous forestc. 500–600 km beyond their present northern limits. In northwestern China, the area of desert and steppe vegetation was reduced compared to present. On the Tibetan Plateau, forest vegetation extended to higher elevations than today and the area of tundra was reduced. These shifts in biome distributions imply significant changes in climate since 6000 years ago that can be interpreted qualitatively as a response to orbital forcing and its secondary effects on the Asian monsoon.

The climate and biomes of Europe at 6000 yr BP: comparison of model simulations and pollen-based reconstructions

14C-dated pollen and lake-level data from Europe are used to assess the spatial patterns of climate change between 6000 yr BP and present, as simulated by the NCAR CCM1 (National Center for Atmospheric Research, Community Climate Model, version 1) in response to the change in the Earth’s orbital parameters during this perod. First, reconstructed 6000 yr BP values of bioclimate variables obtained from pollen and lake-level data with the constrained-analogue technique are compared with simulated values. Then a 6000 yr BP biome map obtained from pollen data with an objective biome reconstruction (biomization) technique is compared with BIOME model results derived from the same simulation. Data and simulations agree in some features: warmer-than-present growing seasons in N and C Europe allowed forests to extend further north and to higher elevations than today, and warmer winters in C and E Europe prevented boreal conifers from spreading west. More generally, however, the agreement is poor. Predominantly deciduous forest types in Fennoscandia imply warmer winters than the model allows. The model fails to simulate winters cold enough, or summers wet enough, to allow temperate deciduous forests their former extended distribution in S Europe, and it incorrectly simulates a much expanded area of steppe vegetation in SE Europe. Similar errors have also been noted in numerous 6000 yr BP simulations with prescribed modern sea surface temperatures. These errors are evidently not resolved by the inclusion of interactive sea-surface conditions in the CCM1. Accurate representation of mid-Holocene climates in Europe may require the inclusion of dynamical ocean–atmosphere and/or vegetation–atmosphere interactions that most palaeoclimate model simulations have so far disregarded.

Diversity of condensed tannin structures affects rumen in vitro methane production in sainfoin (Onobrychis viciifolia) accessions

Sainfoin is a non-bloating temperate forage legume with a moderate-to-high condensed tannin (CT) content. This study investigated whether the diversity of sainfoin accessions in terms of CT structures and contents could be related to rumen in vitro gas and methane (CH4) production and fermentation characteristics. The aim was to identify promising accessions for future investigations. Accessions differed (P < 0·0001) in terms of total gas and CH4 productions. Fermentation kinetics (i.e. parameters describing the shape of the gas production curve and half-time gas production) for CH4 production were influenced by accession (P ≤ 0·038), but not by PEG. Accession, PEG and time affected (P < 0·001) CH4 production, but accession and PEG interaction showed only a tendency (P = 0·08). Increase in CH4 due to PEG addition was not related to CT content. Further analysis of the relationships among multiple traits (nutritional composition, CT structure and CH4 production) using principal component analysis (PCA) based on optimally weighted variables revealed differences among accessions. The first two principal component axes, PC1 (57·6%) and PC2 (18·4%), explained 76·0% of the total variation among accessions. Loading of biplots derived from both PCAs made it possible to establish a relationship between the ratio of prodelphinidin:procyanidin (PD:PC) tannins and CH4 production in some accessions. The PD:PC ratio seems to be an important source of variation that is negatively related to CH4 production. These results suggested that sainfoin accessions collected from across the world exhibited substantial variation in terms of their effects on rumen in vitro CH4 production, revealing some promising accessions for future investigations.

Deglacial changes of the southern margin of the southern westerly winds revealed by terrestrial records from SW Patagonia (52 degrees S)

Much of the ongoing discussion regarding synchrony or bipolar asynchrony of paleoclimate events has centered on the timing and structure of the last glacial termination in the southern mid- latitudes, in particular the southwestern Patagonian region (50�e55�S). Its location adjacent to the Drake Passage andnear the southern margin of the southern westerly winds (SWW) allows examining the postulated links between the Southern Oceane SWW coupled system and tmospheric CO2 variations through the last glacial termination. Results from two sites located in the Última Esperanza area (52�S) allow us to infer SWW-driven changes in hydrologic balance during this critical time interval. These findings indicate peatland development under temperate/wet conditions between 14,600 and 14,900 cal yr BP, followed by cooling and a lake transgressive phase that led to a shallow lake during the early part of the Antarctic Cold Reversal (ACR, 13,600-14,600 cal yr BP), followed in turn by a deeper lake and modest warming during Younger Dryas time (YD, 11,800-13,000 cal yr BP), superseded by terrestrialization and forest expansion at the beginning of the Holocene. We propose that the SWW (i) strengthened and shifted northward during ACR time causing a precipitation rise in northwestern and southwestern Patagonia coeval with mid- and high-latitude cooling and a halt in the deglacial atmospheric CO2 rise; (ii) shifted southward during YD time causing a precipitation decline/increase in NW/SW Patagonia, respectively, high-latitude warming, and invigorated CO2 release from the Southern Ocean; (iii) became weaker between 10,000 and 11,500 cal yr BP causing a precipitation decline throughout Patagonia, concurrent with peak mid- and high-latitude temperatures and atmospheric CO2 concentrations.

MODIS-based vegetation index has sufficient sensitivity to indicate stand-level intra-seasonal climatic stress in oak and beech forests

Context: Variation in photosynthetic activity of trees induced by climatic stress can be effectively evaluated using remote sensing data. Although adverse effects of climate on temperate forests have been subjected to increased scrutiny, the suitability of remote sensing imagery for identification of drought stress in such forests has not been explored fully. Aim: To evaluate the sensitivity of MODIS-based vegetation index to heat and drought stress in temperate forests, and explore the differences in stress response of oaks and beech. Methods: We identified 8 oak and 13 beech pure and mature stands, each covering between 4 and 13 MODIS pixels. For each pixel, we extracted a time series of MODIS NDVI from 2000 to 2010. We identified all sequences of continuous unseasonal NDVI decline to be used as the response variable indicative of environmental stress. Neural Networks-based regression modelling was then applied to identify the climatic variables that best explain observed NDVI declines. Results: Tested variables explained 84–97% of the variation in NDVI, whilst air temperature-related climate extremes were found to be the most influential. Beech showed a linear response to the most influential climatic predictors, while oak responded in a unimodal pattern suggesting a better coping mechanism. Conclusions: MODIS NDVI has proved sufficiently sensitive as a stand-level indicator of climatic stress acting upon temperate broadleaf forests, leading to its potential use in predicting drought stress from meteorological observations and improving parameterisation of forest stress indices.

Enhanced biodiversity and pollination in UK agroforestry systems

Monoculture farming systems have had serious environmental impacts such as loss of biodiversity and pollinator decline. The authors explain how temperate agroforestry systems show potential in being able to deliver multiple environmental benefits.

Distance from forest edge affects bee pollinators in oilseed rape fields

Wild pollinators have been shown to enhance the pollination of Brassica napus(oilseed rape) and thus increase its market value. Several studies have previously shown that pollination services are greater in crops adjoining forest patches or other seminatural habitats than in crops completely surrounded by other crops. In this study, we investigated the specific importance of forest edges in providing potential pollinators in B. napus fields in two areas in France. Bees were caught with yellow pan traps at increasing distances from both warm and cold forest edges into B. napus fields during the blooming period. A total of 4594 individual bees, representing six families and 83 taxa, were collected. We found that both bee abundance and taxa richness were negatively affected by the distance from forest edge. However, responses varied between bee groups and edge orientations. The ITD (Inter-Tegular distance) of the species, a good proxy for bee foraging range, seems to limit how far the bees can travel from the forest edge. We found a greater abundance of cuckoo bees (Nomada spp.) of Andrena spp. and Andrena spp. males at forest edges, which we assume indicate suitable nesting sites, or at least mating sites, for some abundant Andrena species and their parasites (Fig. 1). Synthesis and Applications. This study provides one of the first examples in temperate ecosystems of how forest edges may actually act as a reservoir of potential pollinators and directly benefit agricultural crops by providing nesting or mating sites for important early spring pollinators. Policymakers and land managers should take forest edges into account and encourage their protection in the agricultural matrix to promote wild bees and their pollination services.