Modern vegetation–pollen–climate relationships for the Pampa grasslands of Argentina

Aim To analyse the relationships between potential natural vegetation, pollen and climate in order to improve the interpretation of fossil pollen records and provide the background for future quantitative palaeoclimatic reconstructions. Location Pampa grasslands of Argentina, between 33–41° S and 56–67° W. Methods Modern pollen data were obtained from a pollen data base developed by the Grupo de Investigación de Paleoecología y Palinología, Universidad Nacional de Mar del Plata, Argentina (143 surface samples and 17 pollen types). Analysis of pollen and climate data involved multivariate statistics (cluster analysis and principal components analysis), scatter diagrams, Pearson’s correlation and isopoll mapping. Results Vegetation patterns at regional scales (grasslands and xerophytic woodlands) and local scales (edaphic communities) were identified by cluster analysis of pollen surface samples. The main climatic variables that appear to constrain the vegetation distribution and abundance of taxa are mean annual precipitation, annual effective precipitation and summer temperature. Individual pollen types such as Chenopodiaceae, Apiaceae, Cyperaceae, Prosopis, Schinus, Condalia microphylla and other xerophytic taxa are good indicators of moisture regime. Many pollen types are significantly correlated with summer temperature. The modern vegetation–pollen–climate relationships vary in a broadly predictable manner, supporting the contention that fossil pollen assemblages can be related to particular climatic characteristics. Main conclusions An expanded suite of modern analogues facilitated new insights into vegetation–pollen–climate relationships at the regional scale in Pampa grasslands. Relationships between individual pollen types and climate are appraised at a regional scale and new modern analogues are presented. The results provide the basis for improved vegetation and climate reconstruction from fossil records of the study area.     

Modern pollen–vegetation relationships in subarctic southern Greenland and the interpretation of fossil pollen data from the Norse landnám

Aim The objective of this paper is to explore the relationships that exist between vegetation and modern pollen rain in the open, largely treeless landscape of subarctic Greenland. The implications of these results for the interpretation of fossil pollen assemblages from the time of the Norse landnám are then examined. Location The study area is the sheep farming district of Qassiarsuk in the subarctic, subcontinental vegetational and climatic zone of southern Greenland (61° N, 45° W). Between c.ad 1000–1500 this region was contained within the Norse Eastern Settlement. Methods Detrended Correspondence Analysis (DCA) of harmonized plant–pollen data sets is used to compare plant cover in 64 vegetation quadrats with pollen assemblages obtained from moss polsters at matching locations. Presence/absence data are also used to calculate indices of association, over‐ and under‐representation for pollen types. Results Good correspondence between paired vegetation–pollen samples occurs in many cases, particularly in locations where Salix glauca–Betula glandulosa dwarf shrub heath is dominant, and across herbaceous field boundaries and meadows. Pollen samples are found to be poor at reflecting actual ground cover where ericales or Juniperus communis are the locally dominant shrubs. Dominant or ubiquitous taxa within this landscape (Betula, Salix and Poaceae) are found to be over‐represented in pollen assemblages, as are several of the ‘weeds’ generally accepted as introduced by the Norse settlers. Main conclusions Due to their over‐representation in the pollen rain, many of the Norse apophytes and introductions (e.g. Rumex acetosa and R. acetosella) traditionally used to infer human activity in Greenland should be particularly sensitive indicators for landnám, allowing early detection of Norse activity in fossil assemblages. Pteridophyte spores are found to be disassociated with the ground cover of ferns and clubmosses, but are over‐represented in pollen assemblages, indicating extra‐local or regional sources and long residence times in soil/sediment profiles for these microfossils. A pollen record for Hordeum‐type registered in close proximity to a field containing barley suggests that summer temperatures under the current climatic regime are, at least on occasion, sufficient to allow flowering.     

Native woody vegetation in central Argentina: Classification of Chaco and Espinal forests

Question

What are the composition and spatial patterns of native woody plant communities in the southern Great Chaco and Espinal?

Location

Córdoba Province, central Argentina, an area of ca. 161,000 km².

Methods

We collected 351 geo‐referenced relevés representative of the geographic, topographic and ecological variation of the Chaco and Espinal woody vegetation in central Argentina. The relevés were classified into vegetation types using the hierarchical ISOPAM method. Forest and shrubland types were described on the basis of diagnostic species occurrences and their distribution in relation to environmental factors. A map of the actual vegetation derived from remote‐sensed images (Landsat) and field data was used to describe the current distribution and abundance of the different vegetation types.

Results

The classification of the 351 plots × 837 species matrix revealed two major clusters comprising seven woody vegetation types corresponding to Chaco lowland and mountain forests and shrublands, Espinal forests and edaphic vegetation. The most important gradients in woody vegetation types are related to elevation, temperature and rainfall variables.

Conclusions

Subtropical seasonally dry woody plant communities from the southern extreme of the Great Chaco and Espinal forests were described for the first time based on complete floristic data. Our results show that lowland Chaco native forests, as well as replacement communities, are still present in its southern distribution range and are well distinguishable from other vegetation types such as the Espinal and mountain forests. Overall, extensive Espinal forests have almost disappeared while Chaco vegetation is highly fragmented and degraded.

     

Modern pollen–vegetation relationships along transects on the Whangapoua Estuary, Great Barrier Island, northern New Zealand

Aims To quantify pollen–vegetation relationships from saline to freshwater in an estuarine gradient from surface samples of the modern pollen rain, to allow more accurate interpretations of the stratigraphic palynological record. Location Whangapoua Estuary, Great Barrier Island, northern New Zealand. Methods Six transects were laid out along a vegetation sequence running from estuarine mud to freshwater swamp. Along these transect lines, 108 plots were sampled for vegetation and surface sediments from wet sand, mud, plant litter or moss (sand and mud sites are inundated by most tides, other sites less frequently). All sediment samples were analysed for pollen. The relationships between plant species frequency and pollen representation were examined at a community scale using twinspan and ordination analyses, and for individual species using fidelity and dispersibility indices, regression and box‐plot analyses. Results The quantitative relationships between source taxon vegetation frequency and its pollen representation varied between species due to differential pollen production and dispersal. twinspan of the surface pollen samples suggests five vegetation types: (A) mangrove (Avicennia marina); (C) Leptocarpus similis salt meadow; (D) Baumea sedges; (E) Leptospermum shrubland; and (F) Typha/Cordyline swamp forest. The (B) Juncus kraussii community is not represented palynologically owing to the destruction of its delicate pollen grains during acetolysis of samples. Detrended correspondence analysis places these communities on an estuarine‐to‐freshwater gradient. However, pollen assemblages at the seaward end of the salinity gradient are less clearly representative of the associated vegetation than those at the landward end, probably because the open vegetation at the former allows the influx of wind‐ and water‐dispersed pollen from surrounding vegetation. Main conclusions The vegetation pattern (zonation) at Whangapoua is reflected in the pollen rain. When the long‐distance and over‐represented pollen types are excluded, five out of six of the broad vegetation communities can be identified by their pollen spectra. Species with high fidelity and low‐to‐moderate dispersibility indices can be used to identify the vegetation types in the sedimentary sequences. The more open vegetation types at the ‘marine end’ of the sequence tend to be ‘overwhelmed’ by regional pollen, but the nature of the sediments and the presence of discriminatory species (e.g. A. marina, Plagianthus divaricatus, Cordyline australis), even in small amounts, will allow correct identification of the local vegetation represented in sedimentary palynological sequences. A box‐plot analysis indicates that the pollen and spore types A. marina (mangroves), Sarcocornia quinqueflora (salt meadow), P. divaricatus (sedges), Gleichenia (shrubland) and C. australis (swamp forest) are highly discriminatory in relation to vegetation type. These discriminatory palynomorphs help with the interpretation of stratigraphic pollen studies. However, salt marsh vegetation communities in the sediments must be interpreted with caution as the marine sediments are easily affected by erosion, bioturbation and tidal inundation effects.     

Coastal dune vegetation and pollen representation in south Buenos Aires Province, Argentina

Aim To establish the relationship between coastal dune vegetation and its pollen representation as an aid to interpret Holocene vegetation dynamics and environmental changes from pollen assemblages. Location The study area is situated on the temperate Atlantic coast of south Buenos Aires Province, Argentina (c. 39° S and 61°20′ W). Methods The vegetation of the active dune area adjacent to the beach was described on the basis of its floristic composition from 25 plots. Classification of the vegetation into distinct zones was carried out by cluster analysis. Surface samples were collected from each vegetation stand and analysed for their pollen composition. Pollen percentage data were analysed using principal components analysis in order to investigate the degree to which the different vegetation units can be distinguished by their pollen spectra. Pollen–vegetation relationships for selected taxa were explored using simple scatter plots and indices of association, under‐ and over‐representation. Indices of floristic diversity and palynological richness were used to assess the representation of the vegetation in the pollen spectra. Results and conclusions Five vegetation zones are defined on the basis of species composition and their quantitative variation: back shore, mobile dunes, slacks, semi‐fixed and fixed dunes. Pollen assemblages from back shore, mobile dunes and slacks are clearly differentiated from semi‐fixed and fixed dunes. Pollen assemblages differ considerably from the associated vegetation composition. Major discrepancies are caused by large differences in pollen and vegetation proportion of Hyalis argentea and Discaria americana. There is a considerable proportion of non‐local pollen in every spectrum. Pollen representation in the coastal dunes at Monte Hermoso is influenced by differences in pollen production, dispersal and preservation of individual taxa as well as by the spatial distribution of the vegetation, the topography of the dune system and the wind pattern. The pollen–vegetation relationship established in this study has important implication for understanding and interpreting fossil pollen records from coastal dune environments.     

Modern pollen samples from alpine vegetation on the Tibetan Plateau

• 1 A set of 316 modern surface pollen samples, sampling all the alpine vegetation types that occur on the Tibetan Plateau, has been compiled and analysed. Between 82 and 92% of the pollen present in these samples is derived from only 28 major taxa. These 28 taxa include examples of both tree (AP) and herb (NAP) pollen types.
• 2 Most of the modern surface pollen samples accurately reflect the composition of the modern vegetation in the sampling region. However, airborne dust‐trap pollen samples do not provide a reliable assessment of the modern vegetation. Dust‐trap samples contain much higher percentages of tree pollen than non‐dust‐trap samples, and many of the taxa present are exotic. In the extremely windy environments of the Tibetan Plateau, contamination of dust‐trap samples by long‐distance transport of exotic pollen is a serious problem.
• 3 The most characteristic vegetation types present on the Tibetan Plateau are alpine meadows, steppe and desert. Non‐arboreal pollen (NAP) therefore dominates the pollen samples in most regions. Percentages of arboreal pollen (AP) are high in samples from the southern and eastern Tibetan Plateau, where alpine forests are an important component of the vegetation. The relative importance of forest and non‐forest vegetation across the Plateau clearly follows climatic gradients: forests occur on the southern and eastern margins of the Plateau, supported by the penetration of moisture‐bearing airmasses associated with the Indian and Pacific summer monsoons; open, treeless vegetation is dominant in the interior and northern margins of the Plateau, far from these moisture sources.
• 4 The different types of non‐forest vegetation are characterized by different modern pollen assemblages. Thus, alpine deserts are characterized by high percentages of Chenopodiaceae and Artemisia, with Ephedra and Nitraria. Alpine meadows are characterized by high percentages of Cyperaceae and Artemisia, with Ranunculaceae and Polygonaceae. Alpine steppe is characterized by high abundances of Artemisia, with Compositae, Cruciferae and Chenopodiaceae. Although Artemisia is a common component of all non‐forest vegetation types on the Tibetan Plateau, the presence of other taxa makes it possible to discriminate between the different vegetation types.
• 5 The good agreement between modern vegetation and modern surface pollen samples across the Tibetan Plateau provides a measure of the reliability of using pollen data to reconstruct past vegetation patterns in non‐forested areas.

     

黄土高原地区植被与气候的关系

利用地理信息系统技术结合典范对应分析和数量区划的方法。研究了黄土高原地区植被与气候之间的关系。排序结果表明：CCA的第一轴代表黄土高原植被和气候梯度的纬向变化,水分梯度是决定植被分布的最主要气候因子,热量梯度中的全年月平均最低气温、月平均最高气温、年均温也对植被的纬向性分布有较大的影响,黄土高原植被与气候梯度表现出明显的纬向性;CCA的第二轴代表黄土高原植被和气候梯度的经向性变化,热量梯度是决定植被经向性分布的最主要气候因子,水分梯度中的全年最大蒸散量对植被的经向性分布有较大的影响。黄土高原植被与气候梯度表现出明显的经向分布规律性。     

瑞典南部相邻桦树和欧洲云杉林分的地表植被和土壤特征

对瑞曲南部23对相邻的桦树和欧洲云杉林分的地表植被和土壤特征进行了研究。桦树和欧洲云杉林分曲芒发草（Descham0psia flexuosa L.）和苔藓植物的频度几乎没有区别,但桦树林分越桔（Vaccinium myrtillus L.）和黑果越桔（Vac-cinium vitis-idaea L.）的频度远大于欧洲云杉林分的频度。欧洲云杉林分各层土壤（腐殖质层、0－10cm矿物质土壤层和11－30cm矿物质土壤层）的酸度稍大于松树林分各层土壤的酸度。欧洲云杉森分土壤可交换性K、Ca和Mg离子的含量稍大于树林分土壤的含量,但差别不明显,只有可交换性Na离子的含量在欧洲云杉林分土训中明显偏高。由于现在硫沉降明显降低,相对于树冠层而言欧洲云杉冠层而言欧洲云杉冠层较高的阳离子干沉降在一定程度上可能中和土壤酸化对瑞典南部欧洲云杉林的影响。     

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.     

北京地区植被景观中斑块形状的指数分析

选取４个斑块形状指数（斑块的击长面积比ＳＩ１、斑块周长与等面积的圆周长之比ＳＩ２和身份个斑块分维ＳＩ３和ＳＩ４）并借助ＧＩＳ软件ＡＲＣ／ＩＮＦＯ对北京地区植被景观中的斑块形状进行分析,又地这四个形状指数进行 ｅａｒｓｏｎ相关分析和Ｓｐｅａｒｍａｎ秋相关分析,结果表明,ＳＩ１与ＳＩ４,ＳＩ２与ＳＩ３两两之间均呈显著的正的秩相关,ＳＩ１与ＳＩ２,ＳＩ２与ＳＩ４,ＳＩ３与ＳＩ４两两之间于垢负和秩相关。因     

重庆喀斯特地区现代花粉组合与植被的关系

重庆喀斯特地区4种不同地貌单元的81份表土和苔藓样品的孢粉分析结果表明:(1)研究区植物孢粉组合共由109科属组成。除金佛山外,其他植被退化严重地区样点均以草本和蕨类植物占据优势(剔除石漠化治理大量引种的马尾松、柳杉和侧柏等乔木花粉),其次是乔木植物花粉(主要为马尾松),灌木植物花粉含量很低,显示该区植被退化严重;(2)孢粉组合的PCA分析能够很好地将研究区的现代孢粉组合区分开来,较好地反映了重庆喀斯特地区不同地貌单元的植被特征,特别是石漠化区植被退化严重的孢粉组合特征。生态保护较好、自然植被发育的金佛山地区具有106科属的乔木、灌木与草本孢粉类型,以及高含量的乔木花粉,而喀斯特石漠化地区的鸡公山(46科属)、青木关镇(56科属)和南平镇(50科属)则表现为乔灌木孢粉类型较少;(3)银杉作为金佛山特有的孑遗植物,其花粉含量明显低于松属花粉,说明其花粉不利于远距离传播;(4)随着土地利用强度的增加,现代孢粉组合中乔灌木花粉含量和种类越少,草本和蕨类植物孢粉含量越多,且以耕地杂草为主。因此,土地利用方式的变化是喀斯特石漠化区次生植被及其现代孢粉组合变化的主要影响因素。研究可为重庆喀斯特地区...     

A 3-year airborne pollen and fungal spores record in San Carlos de Bariloche, Patagonia, Argentina

Airborne pollen in San Carlos de Bariloche was sampled from September to March 2001–2004 with a Hirst-type volumetric spore trap placed at a height of 15 m in a city extending from the humid forests, through the mesic forests, to the steppe. The total amount of pollen recorded varied widely from year to year. The pollen index was 4,395 in the sampling period 2001/2002; 9,055 in 2002/2003 and 2,756 in 2003/2004. The main pollen period extended from October to January. In October, pollen concentration was the highest. Sixty-six pollen types were identified. Cupressaceae and Nothofagus were the major contributors. Betula, Prunus, Pinus and Populus, the most abundant ornamental taxa in the city, also contributed to the pollen record. Pollen of Maytenus and Lomatia was representative of the mesic forest, while pollen in the lower layers of the humid forest was present in trace amounts. Cupressaceae, Nothofagus and Betula prevailed during spring (September–December), and Plantago, Rumex and Poaceae during summer (December–March). The association of daily pollen concentration and meteorological variables, temperature (mean, maximum, minimum), dew point, rainfall and wind speed, was significant. Correlations showed to be negative, with the exception of that to wind speed. The total sum of fungi spores increased from 1,771 in 2001/2002; through 8,441 in 2002/2003 to 13,782 in 2003/2004. Relative concentration rose to 29%, 48% and 84% of the total number of pollen and fungal spores recorded during each sampling period.     

High-Andean vegetation and environmental gradients in northwestern Patagonia,Argentina

Abstract. We analysed the heterogeneity of high-elevation vegetation on three mountains along a west-east transect at 41 °S lat. in the Andes of northwestern Patagonia, Argentina. In this area, high-Andean vegetation occurs as islands on mountain tops above Nothofagus pumilio forests ? with the timberline at ca. 1700 m a.s.l. We recorded floristic, topographic and substrate data in 166 sites stratified according to longitude, altitude, slope and aspect. Vegetation data were classified with TWINSPAN and ordinated with Detrended Correspondence Analysis. The relationship between environmental and floristic variation was analysed using Canonical Correspondence Analysis. In order of importance, geographical longitude, altitude and aspect were the major determinants of vegetation variation, whereas substrate texture and slope appeared less important. The combination of these factors resulted in two main vegetation gradients. The first gradient is related to a moisture availability gradient, primarily determined by longitude and secondarily to variation in wind exposure (east vs. west aspect). The second vegetation gradient is related to variation in temperature, primarily determined by altitude, and secondarily by variation in insolation related to the contrast between north and south aspects. The four communities obtained with TWINSPAN are therefore associated with four characteristic habitat types: moist-cold, moist-warm, dry-cold and dry-warm. The community of warm and dry environments is the richest and has elements in common with dry steppe communities situated at lower elevations to the east, while the vegetation of the cold-moist habitat type has unique elements that are typical of the southern Andes. Although current climatic factors appear to be the major determinants of high-Andean vegetation gradients, historical events of Pleistocene times probably also affected the vegetation patterns we see today.     

Interdependence of social structure and demography in the southern elephant seal colony of Península Valdés, Argentina

Southern elephant seals breed at Península Valdés (PV, Argentina) along 200 km of coastline. Annual pup counts at peak breeding season for the entire colony increased from 12,113 in 1995 to 14,350 in 2006. Two demographic subunits were identified in the North and South of PV with different trends in births numbers, sex ratios and harem sizes. Birth numbers increased in the South, but decreased sharply in the North. To explain the trends in the colony and subunits, a population model was proposed that integrates social structure (harem size and sex ratio) in a fertility function that quantifies the effects of the social structure on the number of births. We found that a better fit to census data results from our model compared to a linear one  (χ₁²= 4.027, P = 0.045)  . The model was then used to test alternative hypotheses about the role of recruitment and migration on the dynamic of the two subunits. Results indicated the relevance of considering social structure in population models of gregarious and polygynous species, and is an additional tool for comparative studies between populations of elephant seals where long term census are available.     

Phlebotominae of epidemiological importance in cutaneous leishmaniasis in northwestern Argentina: risk maps and ecological niche models

In Argentina, 58.2% out of the 8126 Cutaneous Leishmaniasis (CL) incident cases accumulated from 1954 to 2006 were reported in the provinces of Salta and Jujuy. The aim of this study was to develop an exploratory risk map and a potential distribution map of the vector, in order to offer recommendations for CL prevention. A total of 12 079 Phlebotominae (Diptera: Psychodidae) belonging to the species Lutzomyia neivai (Pinto), Lu. migonei (França), Lu. cortelezzii (Brèthes), Lu. shannoni (Dyar), Lu. quinquefer (Dyar) and Brumptomyia spp. (França & Parrot) were captured. Potential distribution models were created for two species, Lu. neivai (incriminated vector of Leishmania braziliensis) and Lu. migonei, associated with domestic animals in Argentina and that in turn could be involved as a link between zoonotic transmission cycles and anthropozoonotic. The Maximum Entropy Modeling System (MaxEnt) was used. The Jackknife test was performed, and the ‘rainfall of the driest month’ was the variable that best generalized the models. Accuracy was evaluated by the area under the curve (AUC) and validated by the Cohen's kappa index. This approximation provides a new analytical resource of high potential for the prevention of the disease, in order to allocate resources properly and to develop the most suitable strategies for action.     

Soil acidification and vegetation changes in deciduous forest in southern Sweden

Summary Thirtyfour deciduous forest sites in southern Sweden, originally studied in 1949–1970, were resampled in 1984/85. The average pH change in the humus layer was-0.78 and-0.23 in soils originally studied 30–35 and 15–20 years ago, respectively. Cover changes in the field layer were measured and related to pH changes. The number of species had increased in spite of pH decreases, reaching a maximum at pH 4.0–5.0, while the total cover of the field layer was unchanged.Two groups of species showed no correlation with pH decreases in the humus layer. A large number of species had increased in cover on a majority of sites over the entire species specific pH interval, including nitrophilic species (Rubus idaeus, Chamaenerium angustifolium, Aegopodium podagraria, Stellaria nemorum). A few species had decreased in cover on a majority of sites (Polygonatum multiflorum, Pulmonaria officinalis, Dentaria bulbifera). Some species showed a covariation with pH changes, decreasing in cover in sites in the acidic part of the pH interval (Mercurialis perennis, Lamium galeobdolon, Galium odoratum, Oxalis acetosella, Luzula pilosa). Without excluding other factors, this paper suggests that the increased nitrogen deposition and the increased acidity in the humus layer might cause some of the cover changes in the presented species.     

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.