An extended probabilistic approach of plant vital attributes: an application to European pollen records at 0 and 6 ka

Aim This paper presents a probabilistic method of pollen spectra analysis. The method relies on a pollen taxon characterization using biotic and abiotic plant attribute modes, and their occurrence in a given pollen spectrum at a specific site. This type of analysis can provide an interpretation, which can lead to the reconstruction of the biome and, to an extent, of the abiotic conditions at the site. Methods The analysis has been carried out at the European scale using data provided by the European Pollen Database for about 1000 sites. This dataset contains about 50,000 pollen spectra from the last 21 ka. In these spectra, each pollen taxon has been characterized by a set of 10 chosen attributes. These have been selected with regard to their relevance in biome reconstruction, but also on the basis of available literature. By using the probability of occurrence of each taxon in a given pollen spectrum, it is possible to calculate an affinity index for the spectrum to the attribute considered. To overcome difficulties caused by pollen identification in low diversified pollen spectra, a co‐occurrence concept has been used to give more information. Results The method has been validated on a set of 1327 modern surface samples by comparing the results to the major climatic and environmental variables that control the distribution of the vegetation. A reconstruction exercise on various characteristics of the plants was then carried out on a 6‐ka dataset. This confirmed previous studies by showing a strong dominance of deciduous forest over most of Europe, related to a milder climate than at present in the north and a wetter and colder climate than at present in the south. By analysing the change in pollen/seed dispersion strategies and the light requirement, we show that the history of vegetation dynamics in relation to human influences can be assessed using this method. Main conclusions Our results show that the probabilistic method is an objective tool for pollen assemblage analysis. It allows reconstruction of various characteristics of the vegetation at the continental and global scale for periods and sites with significantly different climate conditions. This method can also be used to compare maps of vegetation attributes for the validation of the new generalized dynamic ecosystems models.     

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–plant–climate relationships in sub-Saharan Africa

Aim  To demonstrate that incorporating the bioclimatic range of possible contributor plants leads to improved accuracy in interpreting the palaeoclimatic record of taxonomically complex pollen types.
Location  North Tropical Africa.
Methods  The geographical ranges of selected African plants were extracted from the literature and geo-referenced. These plant ranges were compared with the pollen percentages obtained from a network of surface sediments. Climate-response surfaces were graphed for each pollen taxon and each corresponding plant species.
Results  Several patterns can be identified, including taxa for which the pollen and plant distributions coincide, and others where the range limits diverge. Some pollen types display a reduced climate range compared with that of the corresponding plant species, due to low pollen production and/or dispersal. For other taxa, corresponding to high pollen producers such as pioneer taxa, pollen types display a larger climatic envelope than that of the corresponding plants. The number of species contained in a pollen taxon is an important factor, as the botanical species included in a taxon may have different geographical and climate distributions.
Main conclusions  The comparison between pollen and plant distributions is an essential step towards more precise vegetation and climate reconstructions in Africa, as it identifies taxa that have a high correspondence between pollen and plant distribution patterns. Our method is a useful tool to reassess biome reconstructions in Africa and to characterize accurately the vegetation and climate conditions at a regional scale, from pollen data.     

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.     

A probabilist model of the relations between pollen and climate and its application to 80 European annual spectra

The relation between pollen and climate is commonly computed by regressing the climatical factor. The disadvantage of this method is that it does not respect the ordinal and intermittent nature of field data. This paper overcomes the artefacts created by this problem by using a probabilist calibration, that quantifies the ecological linkage between a taxon T and a factor with a general parameter, the probability PROX for an abundance A being confined near the rank F of the factor. Confining simulates the effect of the factor upon the concentration of presences and ordering of abundances, and calibrates the climatical behaviour of a taxon with the set of PROX for all possible pairs (A,F). It summarizes a behaviour with the probable position of each abundance A in the range of the factor. Calibration was applied to 130 pollen taxa observed in a network of 80 standardized annual aeropollinic spectra. Spectra were mostly from France, the rest being from a transect stretching from Sweden to Algeria. Spectra were characterized by the values of 10 climatic factors, as well as the presence and abundance of 130 pollen taxa. The influence of geographical climate differences upon pollen content in the atmosphere was quantified by comparing the spectra. Pairs from different localities but the same year were compared. The reliability of indicator taxa was tested by estimating the climate in the 80 spectra using calibration. For all the taxa observed in a spectrum, the envelope of confinings generally followed an unimodal gradient, whose mode was the probable position of the spectrum. Reliability of the estimate was measured by its accuracy, being the agreement between estimates and measures; and by its stability, being the agreement between two estimates from the same climate according to two different flora (the two halves of a spectrum for instance). Average accuracy was 72%, and average stability 87%.     

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.     

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.     

A new global biome reconstruction and data-model comparison for the Middle Pliocene

Aim To produce a robust, comprehensive global biome reconstruction for the Middle Pliocene (c. 3.6–2.6 Ma), which is based on an internally consistent palaeobotanical data set and a state‐of‐the‐art coupled climate–vegetation model. The reconstruction gives a more rigorous picture of climate and environmental change during the Middle Pliocene and provides a new boundary condition for future general circulation model (GCM) studies. Location Global. Methods Compilation of Middle Pliocene vegetation data from 202 marine and terrestrial sites into the comprehensive GIS data base TEVIS (Tertiary Environmental Information System). Translation into an internally consistent classification scheme using 28 biomes. Comparison and synthesis of vegetation reconstruction from palaeodata with the outputs of the mechanistically based BIOME4 model forced by climatology derived from the HadAM3 GCM. Results The model results compare favourably with available palaeodata and highlight the importance of employing vegetation–climate feedbacks and the anomaly method in biome models. Both the vegetation reconstruction from palaeobotanical data and the BIOME4 prediction indicate a general warmer and moister climate for the Middle Pliocene. Evergreen taiga as well as temperate forest and grassland shifted northward, resulting in much reduced tundra vegetation. Warm‐temperate forests (with subtropical taxa) spread in mid and eastern Europe and tropical savannas and woodland expanded in Africa and Australia at the expense of deserts. Discrepancies which occurred between data reconstruction and model simulation can be related to: (1) poor spatial model resolution and data coverage; (2) uncertainties in delimiting biomes using climate parameters; or (3) uncertainties in model physics and/or geological boundary conditions. Main conclusions The new global biome reconstruction combines vegetation reconstruction from palaeobotanical proxies with model simulations. It is an important contribution to the further understanding of climate and vegetation changes during the Middle Pliocene warm interval and will enhance our knowledge about how vegetation may change in the future.     

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.     

Conservatism of ecological niche characteristics in North American plant species over the Pleistocene-to-Recent transition

Aim  To provide a test of the conservatism of a species' niche over the last 20,000 years by tracking the distribution of eight pollen taxa relative to climate type as they migrated across eastern North America following the Last Glacial Maximum (LGM).
Location  North America.
Methods  We drew taxon occurrence data from the North American pollen records in the Global Pollen Database, representing eight pollen types – all taxa for which ≥5 distinct geographic occurrences were available in both the present day and at the LGM (21,000 years ago ± 3000 years). These data were incorporated into ecological niche models based on present-day and LGM climatological summaries available from the Palaeoclimate Modelling Intercomparison Project to produce predicted potential geographic distributions for each species at present and at the LGM. The output for each time period was projected onto the 'other' time period, and tested using independent known occurrence information from that period.
Results  The result of our analyses was that all species tested showed general conservatism in ecological characteristics over the climate changes associated with the Pleistocene-to-Recent transition.
Main conclusions  This analysis constitutes a further demonstration of general and pervasive conservatism in ecological niche characteristics over moderate periods of time despite profound changes in climate and environmental conditions. As such, our results reinforce the application of ecological niche modelling techniques to the reconstruction of Pleistocene biodiversity distribution patterns, and to project the future potential distribution range of species in the face of global-scale climatic changes.     

A survey of modern pollen and vegetation along a south–north transect in Mongolia

Aim This modern pollen‐rain study documents the spatial and quantitative relationships between modern pollen and vegetation in Mongolia, and explores the potential for using this relationship in palaeoclimatic reconstructions. Location East‐central Mongolia. Methods We collected 104 pollen surface samples along a south–north transect across five vegetation zones in Mongolia. Discriminant analysis was used to classify the modern pollen spectra into five pollen assemblages corresponding to the five vegetation zones. Hierarchical cluster analysis was used to divide the main pollen taxa into two major groups and seven subgroups representing the dry and moist vegetation types and the main vegetation communities within them. Results Each vegetation zone along the transect can be characterized by a distinctive modern pollen assemblage as follows: (1) desert zone: Chenopodiaceae–Zygophyllaceae–Nitraria–Poaceae pollen assemblage; (2) desert‐steppe zone: Poaceae–Chenopodiaceae pollen assemblage; (3) steppe zone: Artemisia–Aster‐type–Poaceae–Pinus Haploxylon‐type pollen assemblage; (4) forest‐steppe zone: Pinus Haploxylon‐type–Picea–Artemisia–Betula, montane forb/shrub and pteridophyte pollen assemblage; and (5) mountain taiga zone: Pinus Haploxylon‐type–Picea–Poaceae–Cyperaceae, montane forb/shrub and Pteridophyte pollen assemblage. Main conclusions Based on the ratio between the major pollen taxon groups and subgroups, we propose two pollen–climate indices that represent the precipitation and temperature conditions in the study region. When plotted along our south–north transect, the moisture indices (M) and temperature indices (T) mimic the regional gradients of precipitation and temperature across Mongolia very closely. These pollen–climate indices can be used for palaeoclimatic reconstruction based on fossil pollen data.     

No‐analog climates and shifting realized niches during the late quaternary: implications for 21st‐century predictions by species distribution models

Empirically derived species distributions models (SDMs) are increasingly relied upon to forecast species vulnerabilities to future climate change. However, many of the assumptions of SDMs may be violated when they are used to project species distributions across significant climate change events. In particular, SDM's in theory assume stable fundamental niches, but in practice, they assume stable realized niches. The assumption of a fixed realized niche relative to climate variables remains unlikely for various reasons, particularly if novel future climates open up currently unavailable portions of species’ fundamental niches. To demonstrate this effect, we compare the climate distributions for fossil‐pollen data from 21 to 15 ka bp (relying on paleoclimate simulations) when communities and climates with no modern analog were common across North America to observed modern pollen assemblages. We test how well SDMs are able to project 20th century pollen‐based taxon distributions with models calibrated using data from 21 to 15 ka. We find that taxa which were abundant in areas with no‐analog late glacial climates, such as Fraxinus, Ostrya/Carpinus and Ulmus, substantially shifted their realized niches from the late glacial period to present. SDMs for these taxa had low predictive accuracy when projected to modern climates despite demonstrating high predictive accuracy for late glacial pollen distributions. For other taxa, e.g. Quercus, Picea, Pinus strobus, had relatively stable realized niches and models for these taxa tended to have higher predictive accuracy when projected to present. Our findings reinforce the point that a realized niche at any one time often represents only a subset of the climate conditions in which a taxon can persist. Projections from SDMs into future climate conditions that are based solely on contemporary realized distributions are potentially misleading for assessing the vulnerability of species to future climate change.     

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.     

Definition of grassland biomes from phytoliths in West Africa

Aim In order to enhance the effectiveness of comparisons between modelled and empirical data for present and past vegetation, it is important to improve the characterization of tropical grass‐dominated biomes reconstructed from fossil tracers. This study presents a method for assigning phytolith assemblages to tropical grass‐dominated biomes, with the objective of offering a new tool for combining pollen and phytolith data in the reconstruction of tropical biomes. Location The West African latitudinal transect studied here extends from 12° N (southern Senegal) to 23° N (southern Mauritania), passing through the Guinean, Sudanian, Sahelian and Saharan bioclimatic zones. Methods Modern phytolith assemblages were extracted from 59 soil surface samples taken throughout the study area and allocated, a priori, to three current biomes: (1) desert C₄ grassland, (2) short grass savanna, and (3) tall grass savanna. Five out of nine phytolith types identified were used as predictors in a discriminant analysis (with calibration and validation steps) for assigning phytolith assemblages to biomes. In addition, 74 modern pollen spectra from the West African transect, acquired from the African Pollen Database ( http://medias.obs‐mip.fr/apd ), were processed by the biomization method. This mathematical procedure involves assigning palynological taxa to one or more plant functional types, which represent broad classes of plants. The plant functional types, in turn, are combined to define biomes following a specific set of algorithms and rules. The resulting maps of the phytolith biomes thus derived were compared with maps of pollen biomes and of contemporary ecosystem classes. Results In the calibration and validation steps, 91.5% and up to 83%, respectively, of the phytolith samples were assigned to the correct biome. The short grass savanna and tall grass savanna biomes were assigned with similar accuracy by both the phytolith and pollen biomization methods, but the phytolith method gave substantially superior results for the desert C₄ grassland biome, providing seven out of seven correct assignments, compared with just one out of four by pollen biomization. Comparisons between an existing ecosystem map and the maps created from phytolith estimation showed close correspondence for desert C₄ grassland, short grass savanna and tall grass savanna, the latter providing correct assignments in 88, 62 and 91% of cases, respectively. Main conclusions The phytolith discriminant analysis method presented here accurately estimates three C₄ grass‐dominated biomes that are widespread in West Africa. Complementarities between the phytolith method and pollen biomization are highlighted. Combining complementary phytolith and pollen data would provide more accurate assignments of C₄ grass‐dominated biomes than pollen biomization alone.     

Late Quaternary biomes of Canada and the eastern United States

Pollen data have been used to construct biome maps for today, 6000 ¹⁴C yr bp and 18,000 ¹⁴C yr bp for Canada and the eastern United States. The inferred modern biome distributions agree well with independent reconstructions of North American vegetation prior to European settlement. Some discrepancies between the pollen data and the modern potential vegetation are caused by post‐settlement clearing of the landscape and the consequent increase of herbaceous types in the recent pollen record. Biome distributions at 6000 ¹⁴C yr bp reflected the warmer and drier conditions then prevalent in the continental interior, but the overall position of biomes was similar to that of today. The boreal treeline in North America was not significantly north of its present position, in contrast to the 100–200 km shift reported for Siberia. At the last glacial maximum (18,000 ¹⁴C yr bp ), steppe and tundra were prevalent in the Midwest and north‐western Canada, and coniferous forests and woodlands grew in eastern North America. The open vegetation at 18,000 ¹⁴C yr bp was probably due to drier conditions and/or lower concentrations of atmospheric CO₂. The composition and physical structure of biomes is not constant over time. Mid‐Holocene biomes were similar in structure to those of today, but shifts in the relative importance of individual plant functional types are large enough that the physical properties of biomes, such as albedo, canopy conductance and surface roughness, are likely to have varied even during the Holocene. Last glacial maximum biomes were structurally different from their modern counterparts. The biome maps therefore may obscure significant vegetational changes in space and time during the late Quaternary. The difference between the highest and next highest affinity scores for each sample measures how strongly affinity scores discriminate among biomes. For many biomes, the difference is not large, and affinity score ties are not uncommon, highlighting the importance of tie‐break procedures when using the biomization method.     

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.     

The mid‐Holocene vegetation of the Mediterranean region and southern Europe,and comparison with the present day

Aim To contribute to the intense debate surrounding the relative influence of climate and humans on Mediterranean‐region land cover over the past 6000 years, we assess the Holocene biogeography and vegetation history of southern Europe by means of an extensive pollen record dataset. Location The Mediterranean biogeographical zone and neighbouring parts of Iberia, the Alps and Anatolia, between 30° N, 48° N, 10° W and 45° E. Methods We compiled a southern European pollen record dataset using available pollen databases (124 sites) and other sources (74 sites), with improved spatial coverage and dating control compared with earlier studies. We used only those sites that had pollen data for both 0 ka and 6 ka. We reconstructed mid‐Holocene and present‐day biomes, arboreal pollen percentages and distribution and relative abundance of 11 key woody taxa, with anomaly maps. Results Northern temperate forest biomes extended further south at the mid‐Holocene than at present, but not as far as earlier studies suggested. Sclerophyllous vegetation occurred along the Mediterranean coast throughout the region at 6 ka. Arboreal pollen percentages were up to 50% higher than at present. At 6 ka, Olea, Fagus and Juniperus had smaller distributions and/or abundances; Abies, Cedrus and both deciduous and evergreen Quercus had larger distributions and/or abundances; Phillyrea, Pistacia and Cistus showed minimal difference; and Pinus showed a cosmopolitan distribution with variable abundance. Main conclusions Temporal difference analysis is more meaningful when only sites containing samples for all time slices are analysed. During the mid‐Holocene, southern Europe was more heavily forested with temperate vegetation than it is at present, but drought‐tolerant xeric vegetation was still widespread along the southern margins of the region. Although human land use may have caused the degradation of land between the mid‐Holocene and the present, the mere presence of xeric vegetation in the Mediterranean region does not require human impact. This challenges the commonly held belief that modern Mediterranean vegetation represents a ‘degraded’ state.     

Biodiversity surrogacy: indicator taxa as predictors of total species richness in Brazilian Atlantic forest and Caatinga

The use of indicator taxa as biodiversity surrogates has received widespread attention in conservation planning, but remains a highly contentious issue. Here we assess biodiversity surrogacy in the two most important biomes of tropical northeastern Brazil, Atlantic forest and Caatinga. We examine the extent to which species richness is correlated among taxonomic groups, and assess relationships between the richness of individual groups, and combinations of groups and total species richness. We introduce a new approach to dealing with autocorrelation between focal taxon richness and total species richness, using standardized data such that each taxon is given equal weight. Our Atlantic forest data covered seven taxa (bryophytes, pteridophytes, trees, ants, euglosine bees, birds, and mammals; total of 768 species) sampled from 12 sites; in Caatinga it was four taxa (trees, spiders, beetles and ants; total of 184 species) from 25 sites. Our results showed that: (1) in nearly all cases the species richnesses of individual taxa were significantly correlated with each other; (2) the species richnesses of most individual taxa were significantly correlated with total species richness in both biomes; (3) only two taxa were required for excellent (R ² > 80%) surrogacy of total species richness in both biomes; and (4) the same two taxa (trees and ants) can provide reasonable (R ² > 60%) surrogacy for total richness in these contrasting biomes. Our findings therefore suggest that the ‘shopping basket of taxa’ required for effective biodiversity surrogacy may not only need to be very small (two taxa), but may also be very limited in composition.     

Within‐taxon niche structure: niche conservatism,divergence and predicted effects of climate change

Several studies have observed that taxa below the level of species can vary in the degree to which they differ from one another in the environmental space they occupy. These patterns of within‐species niche variation raise the question of whether these differences should be considered when developing models for predicting the potential effects of climate change on species distributions. We address this question with two divergent datasets, one on sister species and subspecies from the European herpetofauna, the other on subspecies of breeding birds in North America. Atlas and observation data come from the Atlas of Amphibians and Reptiles in Europe and the North American Breeding Bird Survey, respectively. We develop boosted regression tree models of climate–distribution relationships and project the predicted geographic range of each taxon using interpolated weather station data and modeled climate for the year 2080. We find differences between models that distinguish the contributions of subtaxa and those that do not, in terms of prediction of both current and future distributions. In comparison to models that ignore sub‐taxon structure, models that incorporate this structure generally predict larger areas of suitable conditions, consistently perform better, if only marginally, as measured by cross‐validated AUC, and can reveal divergent potential effects of climate change on subtaxa. Differences in niche occupancy and predicted distribution appear between closely related taxa regardless of their phylogenetic distinctness. For these reasons, information on subtaxon membership and phylogeographic structure should be included in modeling exercises when available, in order to identify both the contribution of these units to the niche occupancy of species and the potentially distinct responses of subtaxa to climate change.