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.     

Resilient forest faunal communities in South Africa: a legacy of palaeoclimatic change and extinction filtering?

Aim To examine the influence of climatic extinction filtering during the last glacial maximum (LGM; c. 18,000 yr bp ) and of the subsequent recolonization of forest faunas on contemporary assemblage composition in southern African forests. Location South Africa, Mozambique, Swaziland, Zimbabwe. Methods Data comprised presence/absence by quarter‐degree grid cell for forest‐dependent and forest‐associated birds, non‐volant mammals and frogs. Twenty‐one forest subregions were assigned to one of three previously identified forest types: Afrotemperate, scarp, and Indian Ocean coastal belt. Differences among forest types were examined through patterns and gradients of species richness and endemism, assemblage similarity, species turnover, and coefficients of species dispersal direction. The influence of contemporary environment on assemblage composition was investigated using partial canonical correspondence analysis. Several alternative biogeographical hypotheses for the recolonization of forest faunas were tested. Results Afrotemperate faunas are relatively species‐poor, have low species turnover, and are unsaturated and infiltrated by generalist species. In northern and central regions, communities are supplemented by recolonization from scarp forest refugia, and among frogs by autochthanous speciation in localized refugia. Scarp faunas are relatively species‐rich, contain many forest‐dependent species, have high species turnover, and overlap with coastal and Afrotemperate faunas. Coastal forests are relatively species‐rich with high species turnover. Main conclusions Afrotemperate communities were affected most by climatic extinction filtering events. Scarp forests were Afrotemperate refugia during the LGM and are a contemporary overlap zone between Afrotemperate and coastal forest. Coastal faunas derive from post‐LGM colonization along the eastern seaboard from tropical East African refugia. The greatest diversity is achieved in scarp and coastal forest faunas in northern KwaZulu–Natal province. This historical centre of diversity has influenced the faunal diversity of nearly all other forests in South Africa. The response of vertebrate taxa to large‐scale, historical processes is dependent on their relative mobility: forest birds best illustrate patterns resulting from post‐glacial faunal dispersal, while among mammals and frogs the legacy of climatic extinction filtering remains stronger.     

Palaeodistribution modelling does not support disjunct Pleistocene refugia in several Central American plant taxa

Aim We combine evidence from palaeoniche modelling studies of several tree species to estimate the extent of Central American forest during the Last Glacial Maximum (LGM). In particular, we ask whether the distributions of these species are likely to have changed since the LGM, and whether LGM distributions coincide with previously proposed Pleistocene refugia in this area. Location Central American wet and seasonally dry forests. Methods We developed ecological niche models using two simulations of Pleistocene climate and occurrence data for 15 Neotropical plant species. We focused on palaeodistribution models of three ‘focal’ tree species that occur in wet and seasonally dry Central American forests, where recent phylogeographic data suggest Pleistocene differentiation coincident with previously proposed refugia. We added predictions from six wet‐forest and six seasonally dry‐forest obligate plant species to gauge whether Pleistocene range shifts were specific to habitat type. Correlation analyses were performed between projected LGM and present distributions, LGM distributions and previously proposed refugia. We also asked whether modelled palaeodistributions were smaller than their current extents. Results According to our models, the ranges of the study species were not reduced during the LGM, and did not correlate with refugial models, regardless of habitat type. Relative range sizes between present and LGM distributions did not indicate significant range changes since the LGM. However, relative range sizes differed overall between the two palaeoclimate models. Main conclusions Many of the modelled palaeodistributions of study species were not restricted to refugia during the LGM, regardless of forest type. While constrained from higher elevations, most species found suitable habitat at coastal margins and on newly exposed land due to lowered sea levels during the LGM. These results offer no corroboration for Pleistocene climate change as a driver of genetic differentiation in the ‘focal’ species. We offer alternative explanations for genetic differentiation found in plant species in this area.     

Leaf area index for northern and eastern North America at the Last Glacial Maximum: a data–model comparison

Aim To estimate the effects of full‐glacial atmospheric CO₂ concentrations and climate upon leaf area index (LAI), using both global vegetation models and palaeoecological data. Prior simulations indicate lowered LAIs at the Last Glacial Maximum (LGM), but this is the first attempt to corroborate predictions against observations. Location Eastern North America and eastern Beringia. Methods Using a dense surface pollen data set and remotely sensed LAIs from the Moderate Resolution Imaging Spectroradiometer (MODIS) instrument, we evaluate the ability of analogue‐based techniques to reconstruct modern LAIs from pollen data. We then apply analogue techniques to LGM pollen records, calculate the ratio of LGM to modern LAIs (RLAI) and compare reconstructed RLAIs to RLAIs simulated by BIOME4. Sensitivity experiments with BIOME4 distinguish the effects of CO₂ and climate on glacial LAIs. Results Modern LAIs are skilfully predicted (r²= 0.83). Data and BIOME4 indicate that LAIs at the LGM were up to 12% lower than modern values in eastern North America and 60–94% lower in Beringia. In eastern North America, LGM climates partially counteracted CO₂‐driven decreases in LAI, while in Beringia both contributed to lowered LAIs. Main conclusions In both regions climate is the primary driver of LGM LAIs. The decline in eastern North America LAIs is smaller than previously reported, so regional vegetation feedbacks to LGM climate may have been less significant than previously supposed. CO₂ exerts both physiological and community effects upon LAI, by regulating resource availability for leaf production and by influencing the competitive balance among species and hence the composition and structure of plant communities. Pollen‐based reconstructions using analogue methods do not incorporate the physiological effect and so are upper estimates of full‐glacial LAIs. BIOME4 sensitivity experiments indicate that the community and physiological effects together caused 10% to 20% decrease in LAIs at the LGM, so simulated RLAIs that are 80–100% of reconstructed RLAIs are regarded as consistent with data.     

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.     

Simulated effects of low atmospheric CO2 on structure and composition of North American vegetation at the Last Glacial Maximum

1. Physiological experiments have indicated that the lower CO₂ levels of the last glaciation (200 μmol mol^?1) probably reduced plant water-use efficiency (WUE) and that they combined with increased aridity and colder temperatures to alter vegetation structure and composition at the Last Glacial Maximum (LGM). 2. The effects of low CO₂ on vegetation structure were investigated using BIOME3 simulations of leaf area index (LAI), and a two-by-two factorial experimental design (modern/LGM CO₂, modern/LGM climate).3. Using BIOME3, and a combination of lowered CO₂ and simulated LGM climate (from the NCAR-CCM1 model), results in the introduction of additional xeric vegetation types between open woodland and closed-canopy forest along a latitudinal gradient in eastern North America.4. The simulated LAI of LGM vegetation was 25–60% lower in many regions of central and eastern United States relative to modern climate, indicating that glacial vegetation was much more open than today.5. Comparison of factorial simulations show that low atmospheric CO₂ has the potential to alter vegetation structure (LAI) to a greater extent than LGM climate.6. If the magnitude of LAI reductions simulated for glacial North America were global, then low atmospheric CO₂ may have promoted atmospheric warming and increased aridity, through alteration of rates of water and heat exchange with the atmosphere.     

Evidence of history in explaining diversity patterns in tropical riverine fish

Aim Documentation of the ongoing effect of rain forest refuges at the last glacial maximum (LGM) on patterns of tropical freshwater fish diversity. Location Tropical South and Central America, and West Africa. Methods LGM rain forest regions and species richness by drainage were compiled from published data. GIS mapping was applied to compile drainage area and contemporary primary productivity. We used multiple regression analyses, applied separately for Tropical South America, Central America and West Africa, to assess differences in species richness between drainages that were connected and disconnected to rain forest refuge zones during the LGM. Spatial autocorrelation of the residuals was tested using Moran's I statistic. We added an intercontinental comparison to our analyses to see if a historical signal would persist even when a regional historical effect (climate at the LGM) had already been accounted for. Results Both area and history (contact with LGM rain forest refuge) explained the greatest proportions of variance in the geographical pattern of riverine species richness. In the three examined regions, we found highest richness in drainages that were connected to the rain forest refuges. No significant residual spatial autocorrelation was detected after considering area, primary productivity and LGM rain forest refuges. These results show that past climatic events still affect West African and Latin American regional and continental freshwater fish richness. At the continental scale, we found South American rivers more species‐rich than expected on the basis of their area, productivity and connectedness to rain forest refuge. Conversely, Central American rivers were less species‐diverse than expected by the grouped model. African rivers were intermediate. Therefore, a historical signal persists even when a regional historical effect (climate at the LGM) had already been accounted for. Main conclusions It has been hypothesized that past climatic events have limited impact on species richness because species have tracked environmental changes through range shifts. However, when considering organisms with physically constrained dispersal (such as freshwater fish), past events leave a perceptible imprint on present species diversity. Furthermore, we considered regions that have comparable contemporary climatic and environmental characteristics, explaining the absence of a productivity effect. From the LGM to the present day (a time scale of 18,000 years), extinction processes should have played a predominant role in shaping the current diversity pattern. By contrast, the continental effects could reflect historical contingencies explained by differences in speciation and extinction rates between continents at higher time scales (millions of years).     

利用孢粉记录定量重建大尺度古植被格局

 古植被定量重建是过去全球变化研究的重点之一, 生物群区化(Biomisation)方法以特征植物功能型来定义生物群区, 通过一种标准化数量方法计算孢粉谱的相似得分, 以此把孢粉谱转变为生物群区类型, 是进行古植被定量重建的一种有效方法。该文在前人综述文章的基础上, 简述了生物群区化方法定量重建古植被格局的发展历史、具体步骤及存在问题, 重点描述了以此方法为基础重建的全新世中期(MH)和末次盛冰期(LGM)的全球古植被分布格局, 以及中国的古植被定量重建工作和古植被格局变化。目前的研究表明, 全新世中期北极森林界线在某些地区有轻微的北移迹象, 北部的温带森林带通常向北远距离迁移, 欧洲的温带落叶林也大范围向地中海地区(向南)和向北扩展, 在北美内陆, 草原侵入到森林生物群区, 但中亚地区却没有此现象, 中国大陆的森林生物群区扩张, 典型撒哈尔植被(如干草原、干旱疏林灌丛和热带干旱森林)进入撒哈拉地区, 而非洲热带雨林却呈减少趋势; 末次盛冰期苔原和草原扩张, 在欧亚大陆北部逐渐混合, 北半球的森林生物群区向南迁移, 北方常绿森林(泰加林)和温带落叶林呈碎片状, 而欧洲和东亚的草原却大范围扩张, 非洲的热带湿润森林(比如热带雨林和热带季雨林)有所减少, 在北美洲的西南地区, 荒漠和草原被开阔针叶疏林所取代。     

Potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum

Aim To identify potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum (LGM). Location Europe. Methods We developed a new approach combining past‐climate simulations with the concept of analogous climate space. Our index gives a continuous measure of the potential of a location to have acted as a source or a sink for species that have shifted their ranges since the LGM. High glacial source potential is indicated by LGM climatic conditions that are widespread now; high post‐glacial sink potential is indicated by current climatic conditions that were widespread at the LGM. The degree of isolation of source and sink areas was calculated as the median distance to areas with analogous climate conditions. Results We identified areas of high glacial source potential in the previously recognized refugial areas in the southern European peninsulas, but also in large areas in central‐western Europe. The most climatically isolated source areas were located in northern Spain, in north‐western Europe and in eastern Turkey. From here species would have had to cover substantial distances to find current climate conditions analogous to LGM conditions of these areas. Areas with high post‐glacial sink potential were mainly located in Fennoscandia and in central and south‐eastern Europe. Some of the most isolated sink areas were located in the Spanish highlands and around the Baltic Sea. Main conclusions Our species‐independent approach successfully identified previously recognized glacial refugial areas with high source potential for species range shifts in southern Europe and in addition highlighted other potential source areas in central Europe. This study offers new insights into how the distribution of past and current climatic conditions may have influenced past species range shifts and current large‐scale biodiversity patterns.     

The incidence and implications of clouds for cloud forest plant water relations

Although clouds are the most recognisable and defining feature of tropical montane cloud forests, little research has focussed on how clouds affect plant functioning. We used satellite and ground‐based observations to study cloud and leaf wetting patterns in contrasting tropical montane and pre‐montane cloud forests. We then studied the consequences of leaf wetting for the direct uptake of water accumulated on leaf surfaces into the leaves themselves. During the dry season, the montane forest experienced higher precipitation, cloud cover and leaf wetting events of longer duration than the pre‐montane forest. Leaf wetting events resulted in foliar water uptake in all species studied. The capacity for foliar water uptake differed significantly between the montane and pre‐montane forest plant communities, as well as among species within a forest. Our results indicate that foliar water uptake is common in these forest plants and improves plant water status during the dry season.     

Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum

Aim We examine several hypotheses emerging from biogeographical and fossil records regarding glacial refugia of a southern thermophilic plant species. Specifically, we investigated the glacial history and post‐glacial colonization of a forest understorey species, Trillium cuneatum. We focused on the following questions: (1) Did T. cuneatum survive the Last Glacial Maximum (LGM) in multiple refugia, and (if so) where were they located, and is the modern genetic structure congruent with the fossil record‐based reconstruction of refugia for mesic deciduous forests? (2) What are the post‐glacial colonization patterns in the present geographical range? Location South‐eastern North America. Methods We sampled 45 populations of T. cuneatum throughout its current range. We conducted phylogeographical analyses based on maternally inherited chloroplast DNA (cpDNA haplotypes) and used TCS software to reconstruct intraspecific phylogeny. Results We detected six cpDNA haplotypes, geographically highly structured into non‐overlapping areas. With one exception, none of the populations had mixed haplotype composition. TCS analysis resulted in two intraspecific cpDNA lineages, with one clade subdivided further by shallower diversification. Main conclusions Our investigation revealed that T. cuneatum survived the LGM in multiple refugia, belonging to two (western, eastern) genealogical lineages geographically structured across south‐eastern North America. The western clade is confined to the south‐western corner of T. cuneatum’s modern range along the Lower Mississippi Valley, where fossil records document a major refugium of mesic deciduous forest. For the eastern clade, modern patterns of cpDNA haplotype distribution suggest cryptic vicariance, in the form of forest contractions and subsequent expansions associated with Pleistocene glacial cycles, rather than simple southern survival and subsequent northward colonization. The north–south partitioning of cpDNA haplotypes was unexpected, suggesting that populations of this rather southern thermophilic species may have survived in more northern locations than initially expected based on LGM climate reconstruction, and that the Appalachian Mountains functioned as a barrier to the dispersal of propagules originating in more southern refugia. Furthermore, our results reveal south‐west to north‐east directionality in historical migration through the Valley and Ridge region of north‐west Georgia.     

Contrasting effects of climate and CO2 on Amazonian ecosystems since the last glacial maximum

The nature of Amazonian ecosystem responses to the large-scale environmental changes characterizing glacial–interglacial cycles is poorly understood. We investigated this issue with a series of transient, continuous 21 000-year simulations using a dynamic process-based ecosystem model. Our results indicate that the Amazon Basin has been dominated by evergreen rain forests since the last glacial maximum (LGM), demonstrating the resilience of this ecosystem to glacial–interglacial environmental change. We find that biome shifts in ecotonal areas since the LGM were driven predominantly by climate change, while coincident, increased ecosystem carbon storage throughout the Amazon Basin was driven largely by CO₂. Our findings imply that recent observed biomass increases in contemporary rain forest plots might be part of a long-term trend driven by the anthropogenic rise in CO₂ over recent centuries.     

Upslope migration of Andean trees

Aim Climate change causes shifts in species distributions, or ‘migrations’. Despite the centrality of species distributions to biodiversity conservation, the demonstrated large migration of tropical plant species in response to climate change in the past, and the expected sensitivity of species distributions to modern climate change, no study has tested for modern species migrations in tropical plants. Here we conduct a first test of the hypothesis that increasing temperatures are causing tropical trees to migrate to cooler areas. Location Tropical Andes biodiversity hotspot, south‐eastern Peru, South America. Methods We use data from repeated (2003/04–2007/08) censuses of 14 1‐ha forest inventory plots spanning an elevational gradient from 950 to 3400 m in Manu National Park in south‐eastern Peru, to characterize changes in the elevational distributions of 38 Andean tree genera. We also analyse changes in the genus‐level composition of the inventory plots through time. Results We show that most tropical Andean tree genera shifted their mean distributions upslope over the study period and that the mean rate of migration is approximately 2.5–3.5 vertical metres upslope per year. Consistent with upward migrations we also find increasing abundances of tree genera previously distributed at lower elevations in the majority of study plots. Main conclusions These findings are in accord with the a priori hypothesis of upward shifts in species ranges due to elevated temperatures, and are potentially the first documented evidence of present‐day climate‐driven migrations in a tropical plant community. The observed mean rate of change is less than predicted from the temperature increases for the region, possibly due to the influence of changes in moisture or non‐climatic factors such as substrate, species interactions, lags in tree community response and/or dispersal limitations. Whatever the cause(s), continued slower‐than‐expected migration of tropical Andean trees would indicate a limited ability to respond to increased temperatures, which may lead to increased extinction risks with further climate change.     

Termite diversity in Neotropical dry forests of Colombia and the potential role of rainfall in structuring termite diversity

Termites are ecosystem engineers that play an important role in the biotransformation and re‐distribution of nutrients in soil. The dry forests are endemic repositories, but at same time, they are most threatened by extensive livestock and crop farming, fires, and climate change. In Colombia, the best‐protected dry forests are located in the north. The termite fauna of dry forests are poorly known. The aim was to identify the termite species occurring in tropical dry forests of the Colombian Caribbean coast in relation to diet and precipitation, temperature, elevation, and soil properties. A total of 32 species in 1,103 occurrences were found. Termitidae accounted for 78% of the species richness with the Anoplotermes‐group, Microcerotermes, and Nasutitermes being the dominant genera. Differences in species composition and abundance were found across sites. These differences may be linked to anthropogenic disturbance and polygyny and polydomy. Strikingly, our highest elevation site (334 m) had the highest species richness much higher than the two lower elevation sites. This implies an inversion of the common elevation‐diversity gradient, also found for termites which can be explained by increasing precipitation with elevation in the dry forest. An analysis of termite species richness at the global scale confirms that termite species richness correlates positively with rainfall. Hence, rainfall seems to positively affect termite diversity. In line, the studied Colombian tropical dry forests had low diversity compared to rain forests. A decline of species‐rich soil‐feeding termites with increasing aridity may explain why the highest termite diversity occurs in humid tropical rain forests. Abstract in Spanish is available with online material.     

Identification of winter moth (Operophtera brumata) refugia in North Africa and the Italian Peninsula during the last glacial maximum

Numerous studies have shown that the genetic diversity of species inhabiting temperate regions has been shaped by changes in their distributions during the Quaternary climatic oscillations. For some species, the genetic distinctness of isolated populations is maintained during secondary contact, while for others, admixture is frequently observed. For the winter moth (Operophtera brumata), an important defoliator of oak forests across Europe and northern Africa, we previously determined that contemporary populations correspond to genetic diversity obtained during the last glacial maximum (LGM) through the use of refugia in the Iberian and Aegean peninsulas, and to a lesser extent the Caucasus region. Missing from this sampling were populations from the Italian peninsula and from North Africa, both regions known to have played important roles as glacial refugia for other species. Therefore, we genotyped field‐collected winter moth individuals from southern Italy and northwestern Tunisia—the latter a region where severe oak forest defoliation by winter moth has recently been reported—using polymorphic microsatellite. We reconstructed the genetic relationships of these populations in comparison to moths previously sampled from the Iberian and Aegean peninsulas, the Caucasus region, and western Europe using genetic distance, Bayesian clustering, and approximate Bayesian computation (ABC) methods. Our results indicate that both the southern Italian and the Tunisian populations are genetically distinct from other sampled populations, and likely originated in their respective refugium during the LGM after diverging from a population that eventually settled in the Iberian refugium. These suggest that winter moth populations persisted in at least five Mediterranean LGM refugia. Finally, we comment that outbreaks by winter moth in northwestern Tunisia are not the result of a recent introduction of a nonnative species, but rather are most likely due to land use or environmental changes.     

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.     

Quaternary Ice-Age dynamics in the Colombian Andes: developing an understanding of our legacy

Pollen records from lacustrine sediments of deep basins in the Colombian Andes provide records of vegetation history, the development of the floristic composition of biomes, and climate variation with increasing temporal resolution. Local differences in the altitudinal distribution of present-day vegetation belts in four Colombian Cordilleras are presented. Operating mechanisms during Quaternary Ice-Age cycles that stimulated speciation are discussed by considering endemism in the asteraceous genera Espeletia, Espeletiopsis and Coespeletia. The floristically diverse lower montane forest belt (1000-2300 m) was compressed by ca. 55% during the last glacial maximum (LGM) (20 ka), and occupied the slopes between 800 m and 1400 m during that period. Under low LGM atmospheric pCO2 values, C4-dominated vegetation, now occurring below 2200 m, expanded up to ca. 3500 m. Present-day C3-dominated paramo vegetation is therefore not an analogue for past C4-dominated vegetation (with abundant Sporobolus lasiophyllus). Quercus immigrated into Colombia 478 ka and formed an extensive zonal forest from 330 ka when former Podocarpus-dominated forest was replaced by zonal forest with Quercus and Weinmannia. During the last glacial cycle the ecological tolerance of Quercus may have increased. In the ecotone forests Quercus was rapidly and massively replaced by Polylepis between 45 and 30 ka illustrating complex forest dynamics in the tropical Andes.     

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.     

Effects of Pleistocene climate changes on species ranges and evolutionary processes in the Neotropical Atlantic Forest

The effects of global glaciations on the distribution of organisms is an essential element of many diversification models. However, the empirical evidence supporting this idea is mixed, in particular with respect to explaining tropical forest evolution. In the present study, we evaluated the impacts of range shifts associated with Pleistocene global glacial cycles on the evolution of tropical forests. In particular, we tested the predictions: (1) that population genetic structure increases with fragmentation variation between the present and the Last Glacial Maximum (LGM) and also (2) with geographical range instability; and (3) that genetic diversity increases with range stability and (4) decreases with fragmentation variation between periods. To address our predictions, we studied population genetic structures and modelled present and past distributions of 15 Atlantic Forest (AF) endemic birds. Afterwards, we evaluated the relationship of population genetic parameters with metrics of species range shifts between the present and the LGM. We found that geographical ranges of AF birds changed in concert with Pleistocene glacial cycles but, unexpectedly, our findings suggest the novel idea that ranges during glacial maxima were slightly larger on average, as well as equally fragmented and displaced from the interglacial ranges. Our findings suggest that range shifts over the late Pleistocene impacted on the diversification of forest organisms, although they did not show that those range shifts had a strong effect. We found that a combination of fragmentation variation across time, small current range size, and range stability increased population genetic structure. However, neither fragmentation, nor range stability affected genetic diversity. Our study showed that evolutionary responses to range shifts across AF birds have a high variance, which could explain the mixed support given by single‐species studies to the action of Pleistocene range shifts on population evolution.     

Improving predictions of tropical forest response to climate change through integration of field studies and ecosystem modeling

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species‐specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model‐data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate.