Systématique et biogéographie du groupe Caesalpinia (famille Leguminosae)

Parmi les lignées des Caesalpinioideae (dans la famille des Leguminosae), l’un des groupes importants au sein duquel les relations phylogénétiques demeurent nébuleuses est le « groupe Caesalpinia », un clade de plus de 205 espèces, réparties présentement entre 14 à 21 genres. La complexité taxonomique du groupe Caesalpinia provient du fait qu’on n’arrive pas à résoudre les questions de délimitations génériques de Caesalpinia sensu lato (s.l.), un regroupement de 150 espèces qui sont provisoirement classées en huit genres. Afin d’arriver à une classification générique stable, des analyses phylogénétiques de cinq loci chloroplastiques et de la région nucléaire ITS ont été effectuées sur une matrice comportant un échantillonnage taxonomique du groupe sans précédent (~84% des espèces du groupe) et couvrant la quasi-totalité de la variation morphologique et géographique du groupe Caesalpinia. Ces analyses ont permis de déterminer que plusieurs genres du groupe Caesalpinia, tels que présentement définis, sont polyphylétiques ou paraphylétiques. Nous considérons que 26 clades bien résolus représentent des genres, et une nouvelle classification générique du groupe Caesalpinia est proposée : elle inclut une clé des genres, une description des 26 genres et des espèces acceptées au sein de ces groupes. Cette nouvelle classification maintient l’inclusion de douze genres (Balsamocarpon, Cordeauxia, Guilandina, Haematoxylum, Hoffmanseggia, Lophocarpinia, Mezoneuron, Pomaria, Pterolobium, Stenodrepanum, Stuhlmannia, Zuccagnia) et en abolit deux (Stahlia et Poincianella). Elle propose aussi de réinstaurer deux genres (Biancaea et Denisophytum), de reconnaître cinq nouveaux genres (Arquita, Gelrebia, Hererolandia, Hultholia et Paubrasilia), et d’amender la description de sept genres (Caesalpinia, Cenostigma, Coulteria, Erythrostemon, Libidibia, Moullava, Tara). Les résultats indiquent qu’il y aurait possiblement aussi une 27e lignée qui correspondrait au genre Ticanto, mais un échantillonage taxonomique plus important serait nécéssaire pour éclaircir ce problème. Les espèces du groupe Caesalpinia ont une répartition pantropicale qui correspond presque parfaitement aux aires du biome succulent, mais se retrouvent aussi dans les déserts, les prairies, les savanes et les forêts tropicales humides. À l’échelle planétaire, le biome succulent consiste en une série d’habitats arides ou semi-arides hautement fragmentés et caractérisés par l’absence de feu, et abrite souvent des espèces végétales grasses, comme les Cactacées dans les néo-tropiques et les Euphorbiacées en Afrique. L’histoire biogéographique du groupe Caesalpinia a été reconstruite afin de mieux comprendre l’évolution de la flore au sein de ce biome succulent. Ce portrait biogéographique a été obtenu grâce à des analyses de datations moléculaires et des changements de taux de diversification, à une reconstruction des aires ancestrales utilisant le modèle de dispersion-extinction-cladogenèse, et à la reconstruction de l’évolution des biomes et du port des plantes sur la phylogénie du groupe Caesalpinia. Ces analyses démontrent que les disjonctions trans-continentales entre espèces sœurs qui appartiennent au même biome sont plus fréquentes que le nombre total de changements de biomes à travers la phylogénie, suggérant qu’il y a une forte conservation de niches, et qu’il est plus facile de bouger que de changer et d’évoluer au sein d’un biome différent. Par ailleurs, contrairement à nos hypothèses initiales, aucun changement de taux de diversification n’est détecté dans la phylogénie, même lorsque les espèces évoluent dans des biomes différents ou qu’il y a changement de port de la plante, et qu’elle se transforme, par exemple, en liane ou herbacée. Nous suggérons que même lorsqu’ils habitent des biomes très différents, tels que les savanes ou les forêts tropicales humides, les membres du groupe Caesalpinia se retrouvent néanmoins dans des conditions écologiques locales qui rappellent celles du biome succulent. Finalement, bien que la diversité des espèces du biome succulent ne se compare pas à celle retrouvée dans les forêts tropicales humides, ce milieu se distingue par un haut taux d’espèces endémiques, réparties dans des aires disjointes. Cette diversité spécifique est probablement sous-estimée et mérite d’être évaluée attentivement, comme en témoigne la découverte de plusieurs nouvelles espèces d’arbres et arbustes de légumineuses dans la dernière décennie. Le dernier objectif de cette thèse consiste à examiner les limites au niveau spécifique du complexe C. trichocarpa, un arbuste des Andes ayant une population disjointe au Pérou qui représente potentiellement une nouvelle espèce. Des analyses morphologiques et moléculaires sur les populations présentes à travers les Andes permettent de conclure que les populations au Pérou représentent une nouvelle espèce, qui est génétiquement distincte et comporte des caractéristiques morphologiques subtiles permettant de la distinguer des populations retrouvées en Argentine et en Bolivie. Nous décrivons cette nouvelle espèce, Arquita grandiflora, dans le cadre d’une révision taxonomique du genre Arquita, un clade de cinq espèces retrouvées exclusivement dans les vallées andines.

A baseline appraisal of water-dependant ecosystem services, the roles they play within desakota livelihood systems and their potential sensitivity to climate change

This report forms part of a larger research programme on 'Reinterpreting the Urban-Rural Continuum', which conceptualises and investigates current knowledge and research gaps concerning 'the role that ecosystems services play in the livelihoods of the poor in regions undergoing rapid change'. The report aims to conduct a baseline appraisal of water-dependant ecosystem services, the roles they play within desakota livelihood systems and their potential sensitivity to climate change. The appraisal is conducted at three spatial scales: global, regional (four consortia areas), and meso scale (case studies within the four regions). At all three scales of analysis water resources form the interweaving theme because water provides a vital provisioning service for people, supports all other ecosystem processes and because water resources are forecast to be severely affected under climate change scenarios. This report, combined with an Endnote library of over 1100 scientific papers, provides an annotated bibliography of water-dependant ecosystem services, the roles they play within desakota livelihood systems and their potential sensitivity to climate change. After an introductory, section, Section 2 of the report defines water-related ecosystem services and how these are affected by human activities. Current knowledge and research gaps are then explored in relation to global scale climate and related hydrological changes (e.g. floods, droughts, flow regimes) (section 3). The report then discusses the impacts of climate changes on the ESPA regions, emphasising potential responses of biomes to the combined effects of climate change and human activities (particularly land use and management), and how these effects coupled with water store and flow regime manipulation by humans may affect the functioning of catchments and their ecosystem services (section 4). Finally, at the meso-scale, case studies are presented from within the ESPA regions to illustrate the close coupling of human activities and catchment performance in the context of environmental change (section 5). At the end of each section, research needs are identified and justified. These research needs are then amalgamated in section 6.

Trophic interactions in a changing world: modelling aboveground-belowground interactions

The rate and scale of human-driven changes can exert profound impacts on ecosystems, the species that make them up and the services they provide that sustain humanity. Given the speed at which these changes are occurring, one of society's major challenges is to coexist within ecosystems and to manage ecosystem services in a sustainable way. The effect of possible scenarios of global change on ecosystem services can be explored using ecosystem models. Such models should adequately represent ecosystem processes above and below the soil surface (aboveground and belowground) and the interactions between them. We explore possibilities to include such interactions into ecosystem models at scales that range from global to local. At the regional to global scale we suggest to expand the plant functional type concept (aggregating plants into groups according to their physiological attributes) to include functional types of aboveground-belowground interactions. At the scale of discrete plant communities, process-based and organism-oriented models could be combined into "hybrid approaches" that include organism-oriented mechanistic representation of a limited number of trophic interactions in an otherwise process - oriented approach. Under global change the density and activity of organisms determining the processes may change non-linearly and therefore explicit knowledge of the organisms and their responses should ideally be included. At the individual plant scale a common organism-based conceptual model of aboveground-belowground interactions has emerged. This conceptual model facilitates the formulation of research questions to guide experiments aiming to identify patterns that are common within, but differ between, ecosystem types and biomes. Such experiments inform modelling approaches at larger scales. Future ecosystem models should better include this evolving knowledge of common patterns of aboveground-belowground interactions. Improved ecosystem models are necessary toots to reduce the uncertainty in the information that assists us in the sustainable management of our environment in a changing world. (C) 2004 Elsevier GmbH. All rights reserved.

Stability of pollination services decreases with isolation from natural areas despite honey bee visits

Sustainable agricultural landscapes by definition provide high magnitude and stability of ecosystem services, biodiversity and crop productivity. However, few studies have considered landscape effects on the stability of ecosystem services. We tested whether isolation from florally diverse natural and semi-natural areas reduces the spatial and temporal stability of flower-visitor richness and pollination services in crop fields. We synthesised data from 29 studies with contrasting biomes, crop species and pollinator communities. Stability of flower-visitor richness, visitation rate (all insects except honey bees) and fruit set all decreased with distance from natural areas. At 1 km from adjacent natural areas, spatial stability decreased by 25, 16 and 9% for richness, visitation and fruit set, respectively, while temporal stability decreased by 39% for richness and 13% for visitation. Mean richness, visitation and fruit set also decreased with isolation, by 34, 27 and 16% at 1 km respectively. In contrast, honey bee visitation did not change with isolation and represented > 25% of crop visits in 21 studies. Therefore, wild pollinators are relevant for crop productivity and stability even when honey bees are abundant. Policies to preserve and restore natural areas in agricultural landscapes should enhance levels and reliability of pollination services.

An assessment of the MODIS collection 5 leaf area index product for a region of mixed coniferous forest

Canopy leaf area index (LAI), defined as the single-sided leaf area per unit ground area, is a quantitative measure of canopy foliar area. LAI is a controlling biophysical property of vegetation function, and quantifying LAI is thus vital for understanding energy, carbon and water fluxes between the land surface and the atmosphere. LAI is routinely available from Earth Observation (EO) instruments such as MODIS. However EO-derived estimates of LAI require validation before they are utilised by the ecosystem modelling community. Previous validation work on the MODIS collection 4 (c4) product suggested considerable error especially in forested biomes, and as a result significant modification of the MODIS LAI algorithm has been made for the most recent collection 5 (c5). As a result of these changes the current MODIS LAI product has not been widely validated. We present a validation of the MODIS c5 LAI product over a 121 km2 area of mixed coniferous forest in Oregon, USA, based on detailed ground measurements which we have upscaled using high resolution EO data. Our analysis suggests that c5 shows a much more realistic temporal LAI dynamic over c4 values for the site we examined. We find improved spatial consistency between the MODIS c5 LAI product and upscaled in situ measurements. However results also suggest that the c5 LAI product underestimates the upper range of upscaled in situ LAI measurements.

Pollen-based biome reconstructions for Latin America at 0, 6000 and 18000 radiocarbon years

The biomisation method is used to reconstruct Latin American vegetation at 6000±500 and 18 000±1000 radiocarbon years before present (14C yr BP) from pollen data. Tests using modern pollen data from 381 samples derived from 287 locations broadly reproduce potential natural vegetation. The strong temperature gradient associated with the Andes is recorded by a transition from high altitude cool grass/shrubland and cool mixed forest to mid-altitude cool temperate rain forest, to tropical dry, seasonal and rain forest at low altitudes. Reconstructed biomes from a number of sites do not match the potential vegetation due to local factors such as human impact, methodological artefacts and mechanisms of pollen representivity of the parent vegetation.

Large inert carbon pool in the terrestrial biosphere at the Last Glacial Maximum

During each of the late Pleistocene glacial–interglacial transitions, atmospheric carbon dioxide concentrations rose by almost 100 ppm. The sources of this carbon are unclear, and efforts to identify them are hampered by uncertainties in the magnitude of carbon reservoirs and fluxes under glacial conditions. Here we use oxygen isotope measurements from air trapped in ice cores and ocean carbon-cycle modelling to estimate terrestrial and oceanic gross primary productivity during the Last Glacial Maximum. We find that the rate of gross terrestrial primary production during the Last Glacial Maximum was about 40±10 Pg C yr−1, half that of the pre-industrial Holocene. Despite the low levels of photosynthesis, we estimate that the late glacial terrestrial biosphere contained only 330 Pg less carbon than pre-industrial levels. We infer that the area covered by carbon-rich but unproductive biomes such as tundra and cold steppes was significantly larger during the Last Glacial Maximum, consistent with palaeoecological data. Our data also indicate the presence of an inert carbon pool of 2,300 Pg C, about 700 Pg larger than the inert carbon locked in permafrost today. We suggest that the disappearance of this carbon pool at the end of the Last Glacial Maximum may have contributed to the deglacial rise in atmospheric carbon dioxide concentrations.

Arid rangeland production systems of Southern Africa

This paper analyses and describes the semi-arid rangelands of southern Africa. These rangelands are found in the grassland, savanna and thicket biomes and comprise all the remaining land which does not support commercial rainfed agriculture, in extent some 778221 km2 (66% of South Africa). Although production is primarily driven by rainfall, rangeland management systems have been developed to cope with the uncertain climate and to ameliorate the impact of inter-annual variation in production. We describe the rangeland types that occur, provide an insight into their management and examine some constraints on livestock production which the socio-economic environment presents. We describe the grazing management systems which apply under the two land tenure systems, namely freehold and leasehold tenure, and discuss how each of these systems effects livestock production, management and resource condition.

The influence of vegetation, fire spread and fire behaviour on biomass burning and trace gas emissions: results from a process-based model

A process-based fire regime model (SPITFIRE) has been developed, coupled with ecosystem dynamics in the LPJ Dynamic Global Vegetation Model, and used to explore fire regimes and the current impact of fire on the terrestrial carbon cycle and associated emissions of trace atmospheric constituents. The model estimates an average release of 2.24 Pg C yr−1 as CO2 from biomass burning during the 1980s and 1990s. Comparison with observed active fire counts shows that the model reproduces where fire occurs and can mimic broad geographic patterns in the peak fire season, although the predicted peak is 1–2 months late in some regions. Modelled fire season length is generally overestimated by about one month, but shows a realistic pattern of differences among biomes. Comparisons with remotely sensed burnt-area products indicate that the model reproduces broad geographic patterns of annual fractional burnt area over most regions, including the boreal forest, although interannual variability in the boreal zone is underestimated.

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.

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.

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.

Climate and biome simulations for the past 21,000 years

This paper reports on a set of paleoclimate simulations for 21, 16, 14, 11 and 6 ka (thousands of years ago) carried out with the Community Climate Model, Version 1 (CCM1) of the National Center for Atmospheric Research (NCAR). This climate model uses four interactive components that were not available in our previous simulations with the NCAR CCM0 (COHMAP, 1988Science, 241, 1043–1052; Wright et al., 1993Global Climate Since the Last Glocial Maximum, University of Minnesota Press, MN): soil moisture, snow hydrology, sea-ice, and mixed-layer ocean temperature. The new simulations also use new estimates of ice sheet height and size from ( Peltier 1994, Science, 265, 195–201), and synchronize the astronomically dated orbital forcing with the ice sheet and atmospheric CO2 levels corrected from radiocarbon years to calendar years. The CCM1 simulations agree with the previous simulations in their most general characteristics. The 21 ka climate is cold and dry, in response to the presence of the ice sheets and lowered CO2 levels. The period 14–6 ka has strengthened northern summer monsoons and warm mid-latitude continental interiors in response to orbital changes. Regional differences between the CCM1 and CCM0 simulations can be traced to the effects of either the new interactive model components or the new boundary conditions. CCM1 simulates climate processes more realistically, but has additional degrees of freedom that can allow the model to ‘drift’ toward less realistic solutions in some instances. The CCM1 simulations are expressed in terms of equilibrium vegetation using BIOME 1, and indicate large shifts in biomes. Northern tundra and forest biomes are displaced southward at glacial maximum and subtropical deserts contract in the mid-Holocene when monsoons strengthen. These vegetation changes could, if simulated interactively, introduce additional climate feedbacks. The total area of vegetated land remains nearly constant through time because the exposure of continental shelves with lowered sea level largely compensates for the land covered by the expanded ice sheets.