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.     

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

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

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

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

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.     

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

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

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

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

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

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

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

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

Late Quaternary paleoecology from fossil beetle communities in the Awatere Valley, South Island, New Zealand

Aim The research aim is to reconstruct last glacial maximum (LGM) and Holocene vegetation history and ecology from fossil beetle assemblages. Location The LGM and Holocene sites are located in the Awatere Valley, which lies in the tectonically active Marlborough Region in the north east of the South Island of New Zealand. Methods Beetle fossils were extracted from silty organic sediment using the standard kerosene flotation method. Fossils were identified by comparisons made to modern species based on morphology and surface features. The ecology and distribution of modern analogues are extrapolated to reconstruct the fossil environment. Results One hundred and forty‐five beetle species belonging to 33 families were identified. The LGM fossil fauna showed the local vegetation was characterized by a forest patch surrounded by an open tussock/grassland landscape. This Nothofagus (southern beech) forest persisted at the site until mid‐Holocene when it was replaced by a podocarp forest that contained high beetle diversity. Herbivores dominate in the early stage of this zone, indicating a relatively new forest environment. Later in the Holocene, the fauna is dominated by detritivores indicating an older more established forest. The late Holocene is characterized by low diversity and the absence of forest species. This fauna indicates that by 500 years ago, the forest was absent and is associated with an almost compete loss of beetle biodiversity. Main conclusions The fossil beetles provide a unique perspective into the past environment in the Awatere Valley on a local scale. The reconstruction supports regional pollen interpretations of Holocene vegetation by identifying a specific forest patch. Fossil beetles are thus a valuable local proxy for vegetation reconstructions.     

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.     

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.     

Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review

In the fossil deposits of the Late Pleistocene ( c . 115 000–11 500 years ago), five felid species are recorded in Europe: the wildcat Felis silvestris , Eurasian lynx Lynx lynx , Iberian lynx Lynx pardinus , leopard Panthera pardus and cave lion Panthera leo spelaea . In the Holocene, Europe was colonized by F. silvestris , L. lynx and L. pardinus as well as the lion Panthera leo . The status of P. pardus in post-glacial Europe is unknown. So far, only sparse records indicate that P. pardus survived into the early Holocene. During the Late Glacial, both L. lynx and L. pardinus occurred on the Iberian Peninsula. However, from the Holocene, only the Iberian lynx is recorded in this region. There are subfossil records that indicate that L. pardinus also occurred in central and western France until c . 3000 years ago. Surprisingly, with reservations on the determination of the bones (by J. Altuna), both lions and cave lions seem to be recorded in the Iberian Peninsula in the Late Glacial. There are published records of the lion P. leo in the northern Iberian Peninsula from the early Holocene. However, its presence in Europe on the basis of subfossil records was proven initially from the Atlantic period. In Ponto-Mediterranean regions of Europe, the lion is recorded from the Atlantic to the younger sub-Atlantic.     

Sensitivity of African biomes to changes in the precipitation regime

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

Evolution of the eastern Asian-North American biogeographic disjunctions in ferns and lycophytes

This paper reviews 31 groups in ferns and lycophytes hypothesized to show eastern Asian–North American disjunctions. Fourteen lineages have been supported by recent phylogenetic evidence: Lycopodium nikoense and Lycopodium sitchense; Isoëtes asiatica and the clade of the North American species complex closely allied to I. maritima; Osmundastrum cinnamomeum; Osmunda claytoniana; the Adiantum pedatum complex; the Cryptogramma acrostichoides complex; Diplaziopsidaceae; Cystopteris chinensis and the Cystopteris bulbifera clade; Asplenium rhizophyllum and Asplenium ruprechtii; diploid Phegopteris; Onoclea sensibilis; the Polypodium appalachianum clade; and the Polypodium glycyrrhiza clade. Phylogenetic and/or cytological evidence did not support the biogeographic disjunctions in six cases: (1) Isoëtes asiatica and I. truncata; (2) Botrychium ternatum; (3) Thelypteris beddomei and T. nipponica—Thelypteris noveboracensis and T. nevadensis; (4) Thelypteris glanduligera and Thelypteris. japonica—T. simulata; (5) Woodwardia japonica and W. virginica; and (6) Woodwardia orientalis and Woodwardia fimbriata. Both vicariance and dispersal have been suggested to be the mechanisms for the formation of the disjunct pattern; and the Beringian region has been an active pathway for the migration of ferns and lycophytes between Asia and North America. Disjunctions of ferns and lycophytes reviewed here have been dated in the Tertiary, and are similar to the ages of eastern Asian–North American disjunctions in seed plants, supporting the close biogeographic co‐diversification of ferns and seed plants. Future studies are needed to estimate divergence times and reconstruct biogeographic events in a broad phylogenetic framework, and to test the morphological stasis hypothesis in disjunct ferns and lycophytes.     

Tree mortality in the eastern and central United States: patterns and drivers

Substantial uncertainty surrounds how forest ecosystems will respond to the simultaneous impacts of multiple global change drivers. Long‐term forest dynamics are sensitive to changes in tree mortality rates; however, we lack an understanding of the relative importance of the factors that affect tree mortality across different spatial and temporal scales. We used the US Forest Service Forest Inventory and Analysis database to evaluate the drivers of tree mortality for eastern temperate forest at the individual‐level across spatial scales from tree to landscape to region. We investigated 13 covariates in four categories: climate, air pollutants, topography, and stand characteristics. Overall, we found that tree mortality was most sensitive to stand characteristics and air pollutants. Different functional groups also varied considerably in their sensitivity to environmental drivers. This research highlights the importance of considering the interactions among multiple global change agents in shaping forest ecosystems.     

Collections of nonhuman primate skeletal materials in the United States and Canada

A mail survey was completed of the nonhuman primate skeletal collections (excluding tupaiids) housed in United States and Canadian museums, universities, and other institutions. Survey results are reported by genus for both skulls and postcrania without regard for age or sex. The survey emphasized specimens of wild-caught primates but institutional personnel were not always able to separate animals which died in captivity. Responses were received from 157 of the 212 institutions to which questionnaires were sent. Of those who returned the questionnaires, 75 institutions have wild-caught, nonhuman primate skeletal materials in their collections. Institutions that have either no nonhuman primate skeletal materials (50) or only the remains of captive animals (36) are listed separately. The total number of specimens reported exeeds 21,000 skulls and 6,400 postcrania.     

Environmental influences on spatial and temporal patterns of body-size variation in California ground squirrels (Spermophilus beecheyi)

Aim  In order to understand how ground squirrels ( Spermophilus beecheyi ) may respond to future environmental change, we investigated five biotic and environmental factors potentially responsible for explaining body-size variation in this species across California. We examined the concordance of spatial patterns with temporal body-size change since the last glacial maximum (LGM).
Location  California, western North America.
Methods  We quantified body size of modern populations of ground squirrels ( n  = 81) and used a model-selection approach to determine the best variables (sex, vegetation, number of congeners, temperature and/or precipitation) explaining geographical variation in body size among modern populations. We also quantified body size of one fossil population in northern California ( n  = 39) and compared temporal body-size change in S. beecheyi at this location since the LGM with model predictions.
Results  Body size of modern populations conformed to Bergmann's rule, with larger individuals in northern (wetter and cooler) portions of California. However, the models suggest that precipitation, rather than temperature or other variables, may best explain variation in body size across modern spatial gradients. Our conclusion is supported by the temporal data, demonstrating that the body size of S. beecheyi has increased in northern California since the LGM, concordant with precipitation but not temperature change in the region.
Main conclusions  Precipitation, rather than temperature, vegetation or number of congeneric species, was the main factor explaining both spatial and temporal patterns of body-size variation in S. beecheyi . The integration of space and time provides a powerful mechanism for predicting how local populations may respond to current and future climatic changes.