Palaeoecological perspectives on pattern and process in plant diversity and distribution adjustments: a comment on recent developments*

Two developments in the understanding of the relationship between sedimentary pollen assemblages, vegetation and plant diversity are discussed. The Prentice model of vegetation–pollen relationships has improved our understanding of how and at what scale sedimentary pollen records vegetation. The modelling framework allows improved palaeoecological study designs that may, potentially, give important new insights into processes of plant migrations in response to climate change. Also, reconstructions of plant communities and landscape openness may be improved. Competing hypotheses for the relationship between vegetation and palynological diversity are discussed and it is concluded that more attention should be focused on evenness aspects of palynological diversity and on hypothesis testing.     

Animal-habitat relationships in the Knysna Forest,South Africa: discrimination between forest types by birds and invertebrates

Summary Effects of forest plant species composition and physiognomy on bird and invertebrate communities were investigated in three discrete, relatively undisturbed forest types along a dry-wet soil moisture gradient. Using discriminant function analysis, a 100% floristic and a 78% vegetation structural discrimination were obtained between the three forest types. However, the bird communities of these different forest types were very similar in species composition, and had much lower densities than those normally encountered in other, superficially similar forests. Although an 81% discrimination between forest types was attained through analysis of ground surface invertebrates, measures of litter and aerial invertebrate abundance were also of limited use as discriminators. Historical and biogeographic factors, as well as the low nutritional levels in the soil and vegetation may be the causes of low bird and invertebrate density and diversity. It is concluded that floristics and vegetation structure have, at best, a minor influence on bird community structure, and possibly also on invertebrate community structure in the Knysna Forest.     

Biodiversity and pollen analysis: modern pollen studies and the recent history of a floodplain woodland in S. W. Ireland

One of the problems with biodiversity for palynologists is that their samples come from an unbounded area and source area varies with both pollen/spore type and vegetation type. There have been two broad approaches to the problem of inferring diversity from pollen/spore samples; firstly the use of pollen type richness as a proxy for species richness, and secondly the ‘identification’ of past vegetation communities aided by ecological inference and historical or modern data concerning species affinities and typical diversity. The estimation of biodiversity from pollen analysis depends upon the pollen count, taxonomic precision, source strength for individual types and dispersal/transport of pollen from source areas to the site. A transect of surface pollen samples is used here to test the effect of a vegetation boundary on pollen/spore diversity and compare the pollen/spore diversity with the species richness of the woodland. Palynological richness (at a constant count sum) does, to some extent, reflect changing local vegetation along the transect. Type count curves also reflect the changes in vegetation type along the transect due to the partial influence of species richness on pollen/spore richness. This study suggests that depending upon the woodland composition, the woodland may not entirely drown-out the pollen signal of the surrounding vegetation, and that a small woodland may be more representative (as a sampling site) of the valley floor diversity than a raised mire. The surface transect is also used in the interpretation of Medieval pollen levels from The Gearagh.     

Diversity of Rich Fen Vegetation and Related Plant Specialists in Mountain Refugia of the Iberian Peninsula

In temperate mountains, fens have been reported as relict habitats subject to geographical fragmentation and broad climatic gradients, but few studies have analyzed the influence of these factors on plant diversity. Here we investigate the effect of isolation on the vegetation diversity of rich fens (Caricion davallianae) in the mountains of the Iberian Peninsula, the distribution limit of these habitats in south-western Europe. We used plot-based vegetation data from the Pyrenees and the Cantabrian mountain range to evaluate their regional species-pool, occurrence of specialists, beta-diversity and the effect of geo-climatic variables on their species-richness and species-composition. We found a lower ratio of rare specialists in the Pyrenees than in the Cantabrian range, but similar estimates in the species pools, total species-richness per plot and beta-diversity. The isolation of the two mountain regions resulted in different species assemblages best predicted by summer precipitation and bedrock types, showing region-based differences in the response of vegetation and plant specialists to the environment. The tighter correlation between local climate and diversity estimates in the Cantabrian range suggests relict character of rich fens in that region, where climatic conditions have restricted local distribution of formerly more widely distributed specialists. Although there is no relevant evidence of vegetation impoverishment in that region, historical isolation has probably resulted in the existence of fragmentary plant communities. We conclude that fen vegetation may experience long-time persistence in climatically sub-optimal mountain refugia, but related plant specialists may be sensitive to climatic changes and subject to the extinction of local populations.     

A palaeoecological investigation into the role of fire and human activity in the development of montane grasslands in East Africa

Human activity has been widely implicated in the origin and expansion of montane grasslands in East Africa, yet little palaeoecological evidence exists to test whether these grasslands are natural or secondary. Pollen and charcoal data derived from two Holocene records in the Eastern Arc mountains of Tanzania are used as a case study to investigate the supposed secondary nature of montane grasslands in Africa. Fossil pollen data are used to detect vegetation change, and charcoal analysis is used to reconstruct fire history. The pollen data are characterised by stable proportions of local taxa suggesting permanence of grasslands throughout the past ~13,000 years. Recent increases in fire adapted taxa such as Morella point towards the development of a grassland/forest patch mosaic possibly associated with burning. However, robust evidence of human activity is absent from the records, which may be attributed to the late human occupation of the mountains. The records indicate long-term persistence of grasslands which, coupled with a lack of evidence of human activity, suggests that these grasslands are not secondary. These data support the hypothesis that grasslands are an ancient and primary component of montane vegetation in Africa, but that they experienced some expansion during the late Holocene as a result of changing fire regime.     

Effects of time since fire on birds in a plant diversity hotspot

Global changes are influencing fire regimes in many parts of the world. In the Fynbos plant diversity hotspot (Cape Floristic Region, South Africa), fire frequency has increased in protected areas where the mean fire interval went from 12–19 to 6–9 years between 1970 and 2000. Fire is one of the main drivers of plant diversity in the Cape Floristic Region. Too frequent fires threaten the persistence of slow-maturing plant species, and such insights have led to the adoption of fire management principles based on plant responses. The effects of fire on Fynbos fauna are much more poorly understood, and have not generally been considered in depth in Fynbos conservation policies, planning or management. We assessed the response of bird communities to long-term fire-induced vegetation changes using space-for-time substitution. We studied bird communities, vegetation structure and plant functional composition in 84 Fynbos plots burnt between two and 18 years before. Ten of the 14 bird species analysed showed a significant change in their abundance with time since fire. We observed a significant species turnover along the post-fire succession due to changes both in vegetation structure and plant functional composition, with a characteristic shift from non-Fynbos specialists and granivorous species to Fynbos specialists and nectarivorous species.If current trends of increasing fire frequency continue, Fynbos endemic birds such as nectarivores may become vulnerable. Conservation management should thus aim more carefully to maintain mosaics of Fynbos patches of different ages. Future research needs to estimate the proportion of vegetation of different ages and patch sizes needed to support dependent fauna, particularly endemics.     

Genetic consequences of Pleistocene range shifts: contrast between the Arctic, the Alps and the East African mountains

In wide-ranging species, the genetic consequences of range shifts in response to climate change during the Pleistocene can be predicted to differ among different parts of the distribution area. We used amplified fragment length polymorphism data to compare the genetic structure of Arabis alpina, a widespread arctic-alpine and afro-alpine plant, in three distinct parts of its range: the North Atlantic region, which was recolonized after the last ice age, the European Alps, where range shifts were probably primarily altitudinal, and the high mountains of East Africa, where the contemporary mountain top populations result from range contraction. Genetic structure was inferred using clustering analyses and estimates of genetic diversity within and between populations. There was virtually no diversity in the vast North Atlantic region, which was probably recolonized from a single refugial population, possibly located between the Alps and the northern ice sheets. In the European mountains, genetic diversity was high and distinct genetic groups had a patchy and sometimes disjunct distribution. In the African mountains, genetic diversity was high, clearly structured and partially in accordance with a previous chloroplast phylogeography. The fragmented structure in the European and African mountains indicated that A. alpina disperses little among established populations. Occasional long-distance dispersal events were, however, suggested in all regions. The lack of genetic diversity in the north may be explained by leading-edge colonization by this pioneer plant in glacier forelands, closely following the retracting glaciers. Overall, the genetic structure observed corresponded to the expectations based on the environmental history of the different regions.     

Pollen–plant–climate relationships in sub-Saharan Africa

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

Response of pollen diversity to the climate-driven altitudinal shift of vegetation in the Colombian Andes

Change in diversity of fossil pollen through time is used as a surrogate for biodiversity history. However, there have been few studies to explore the sensitivity of the measured pollen diversity to vegetation changes and the relationship between pollen diversity and plant diversity. This paper presents results of a study to assess the relationship between pollen diversity and relative abundance of pollen from different altitudinal vegetation belts (subandean forest, Andean forest, subparamo and grassparamo) in three records from the tropical Andes in Colombia. The results indicated that plant diversity in the vegetation declined with altitude and pollen diversity is positively correlated to the abundance of pollen from lower altitude vegetation belts and negatively correlated to that from the grassparamo. These results, therefore, suggest that pollen diversity coarsely reflects the diversity of the surrounding vegetation. Using this interpretation, we were able to predict changes in plant diversity over the past 430000 years in the Colombian Andes. Results indicated that under warmer climatic conditions, more species-diverse vegetation of low elevation moved upslope to contribute more pollen diversity to the study sites, and under colder conditions, species-poor grassparamo moved downslope and observed pollen diversity was lower. This study concludes that fossil pollen diversity may provide an important proxy to reconstruct the temporal changes in plant diversity.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.     

Site fertility drives temporal turnover of vegetation at high latitudes

Experimental evidence shows that site fertility is a key modulator underlying plant community changes under climate change. Communities on fertile sites, with species having fast dynamics, have been found to react more strongly to climate change than communities on infertile sites with slow dynamics. However, it is still unclear whether this generally applies to high‐latitude plant communities in natural environments at broad spatial scales. We tested a hypothesis that vegetation of fertile sites experiences greater changes over several decades and thus would be more responsive under contemporary climate change compared to infertile sites that are expected to show more resistance. We resurveyed understorey communities (vascular plants, bryophytes, and lichens) of four infertile and four fertile forest sites along a latitudinal bioclimatic gradient. Sites had remained outside direct human disturbance. We analyzed the magnitude of temporal community turnover, changes in the abundances of plant morphological groups and strategy classes, and changes in species diversity. In agreement with our hypothesis, temporal turnover of communities was consistently greater on fertile sites compared to infertile sites. However, our results suggest that the larger turnover of fertile communities is not primarily related to the direct effects of climatic warming. Furthermore, community changes in both fertile and infertile sites showed remarkable variation in terms of shares of plant functional groups and strategy classes and measures of species diversity. This further emphasizes the essential role of baseline environmental conditions and nonclimatic drivers underlying vegetation changes. Our results show that site fertility is a key determinant of the overall rate of high‐latitude vegetation changes but the composition of plant communities in different ecological contexts is variously impacted by nonclimatic drivers over time.     

Ecological consequences of Late Quaternary extinctions of megafauna

Large herbivorous vertebrates have strong interactions with vegetation, affecting the structure, composition and dynamics of plant communities in many ways. Living large herbivores are a small remnant of the assemblages of giants that existed in most terrestrial ecosystems 50 000 years ago. The extinction of so many large herbivores may well have triggered large changes in plant communities. In several parts of the world, palaeoecological studies suggest that extinct megafauna once maintained vegetation openness, and in wooded landscapes created mosaics of different structural types of vegetation with high habitat and species diversity. Following megafaunal extinction, these habitats reverted to more dense and uniform formations. Megafaunal extinction also led to changes in fire regimes and increased fire frequency due to accumulation of uncropped plant material, but there is a great deal of variation in post-extinction changes in fire. Plant communities that once interacted with extinct large herbivores still contain many species with obsolete defences against browsing and non-functional adaptations for seed dispersal. Such plants may be in decline, and, as a result, many plant communities may be in various stages of a process of relaxation from megafauna-conditioned to megafauna-naive states. Understanding the past role of giant herbivores provides fundamental insight into the history, dynamics and conservation of contemporary plant communities.     

Multiscale climate change impacts on plant diversity in the Atacama Desert

Comprehending ecological dynamics requires not only knowledge of modern communities but also detailed reconstructions of ecosystem history. Ancient DNA (aDNA) metabarcoding allows biodiversity responses to major climatic change to be explored at different spatial and temporal scales. We extracted aDNA preserved in fossil rodent middens to reconstruct late Quaternary vegetation dynamics in the hyperarid Atacama Desert. By comparing our paleo‐informed millennial record with contemporary observations of interannual variations in diversity, we show local plant communities behave differentially at different timescales. In the interannual (years to decades) time frame, only annual herbaceous expand and contract their distributional ranges (emerging from persistent seed banks) in response to precipitation, whereas perennials distribution appears to be extraordinarily resilient. In contrast, at longer timescales (thousands of years) many perennial species were displaced up to 1,000 m downslope during pluvial events. Given ongoing and future natural and anthropogenically induced climate change, our results not only provide baselines for vegetation in the Atacama Desert, but also help to inform how these and other high mountain plant communities may respond to fluctuations of climate in the future.     

Revealing hidden plant diversity in arid environments

Estimating total plant diversity in extreme or hyperarid environments can be challenging, as adaptations to pronounced climate variability include evading prolonged stress periods through seeds or specialized underground organs. Short‐term surveys of these ecosystems are thus likely poor estimators of actual diversity. Here we develop a multimethod strategy to obtain a more complete understanding of plant diversity from a community in the Atacama Desert. We explicitly test environmental DNA‐based techniques (eDNA) to see if they can reveal the observed and ‘hidden' (dormant or locally rare) species. To estimate total plant diversity, we performed long‐term traditional surveys during eight consecutive years, including El Niño and La Niña events, we then analyzed eDNA from soil samples using high‐throughput sequencing. We further used soil pollen analysis and soil seed bank germination assays to identify ‘hidden' species. Each approach offers different subsets of current biodiversity at different taxonomic, spatial and temporal resolution, with a total of 92 taxa identified along the transect. Traditional field surveys identified 77 plant species over eight consecutive years. Observed community composition greatly varies interannually, with only 22 species seen every year. eDNA analysis revealed 37 taxa, eight of which were ‘hidden' in our field surveys. Soil samples contain a viable seed bank of 21 taxa. Soil pollen (27 taxa) and eDNA analysis show affinities with vegetation at the landscape scale but a weak relationship to local plot diversity. Multimethod approaches (including eDNA) in deserts are valuable tools that add to a comprehensive assessment of biodiversity in such extreme environments, where using a single method or observations over a few years is insufficient. Our results can also explain the resilience of Atacama plant communities as ‘hidden' taxa may have been active in the recent past or could even emerge in the future as accelerated global environmental change continues unabated.     

Natural and land-use history of the Northwest mountain ecoregions (USA) in relation to patterns of plant invasions

Although the Northwest currently has the least proportion of non-native invasive plant species relative to other regions of North America, invasions continue to increase into the mountainous areas of the region. Landscape structure, such as the variation found along the complex gradients of the Northwest mountain ecoregions, affects the expansion of invasive plant species and the invasibility of plant communities. Also, the history of land use and current use patterns affect the expansion of invasive plants, and many of the deteriorated environments in the region's mountains may invite and stabilize plant invasions. We examined the patterns of invasive plant diversity in Northwest mountain ecoregions, as derived from literature sources, to analyze which factors influence plant invasions. Our analysis found altered riparian systems and disturbed forests to be especially vulnerable to plant invasion. Conversely, alpine and wilderness areas are still relatively unaffected by invasive plants. Both riparian and alpine communities, while making up a relatively small area across Northwest mountain ecoregions, have significant ecological importance and deserve special protection from invasive plant introductions. Human settlement at low elevations and intense land use of upland forests will likely continue to enhance invasive plant introductions into Northwest mountain ecosystems. Knowledge of the relationships between biological and environmental factors, disturbance, and human land use will be critical for future management strategies that proactively locate, prevent, or contain plant invasions in the mountains of the Northwest.     

Biome reconstruction from pollen and plant macrofossil data for Africa and the Arabian peninsula at 0 and 6000 years

Biome reconstruction from pollen and plant macrofossil data provides an objective method to reconstruct past vegetation. Biomes for Africa and the Arabian peninsula have been mapped for 6000 years bp and provide a new standard for the evaluation of simulated palaeovegetation distributions. A test using modern pollen data shows the robustness of the biomization method, which is able to predict the major vegetation types with a high confidence level. The application of the procedure to the 6000 years data set (pollen and plant macrofossil analyses) shows systematic differences from the present that are consistent with the numerous previous regional and continental interpretations, while providing a more extensive and more objective basis for such interpretations. Madagascar, eastern, southern and central Africa show only minor changes in terms of biomes, compared to present. Major changes in biome distributions occur north of 15°N, with steppe in many low-elevation sites that are now desert, and temperate xerophytic woods/scrub and warm mixed forest in the Saharan mountains. These shifts in biome distributions imply significant changes in climate, especially precipitation, between 6000 years and present, reflecting a change in monsoon extent combined with a southward expansion of Mediterranean influence.     

Surface pollen-vegetation relationships on the Atlantic seaboard: South Uist, Scotland

Studies of modern pollen rain from remote islands have raised a number of interesting issues concerning the spatial precision of present‐day pollen spectra in relation to their parent plant community types. This paper examines the relationships and degree of correlation between a sequence of contemporary vegetation types, environments and their associated surface pollen spectra from a transect across the island of South Uist in the Outer Hebrides of Scotland. Paired data on both contemporary vegetation and associated surface pollen assemblages have been collected and analysed using methods of numerical classification and ordination. In general terms, the modern pollen rain on South Uist reflects the major changes in vegetation pattern and the major community types fairly closely. The major boundary between the alkaline machair sand dune communities and the various acidic upland vegetation types is particularly clear. However, both variability in the vegetation and the effects of the strong prevailing westerly and south‐westerly winds tend to blur the boundaries of the various communities within each of these larger categories. On average 86.6% of the palynomorphs come from in‐community quadrat sources, while only 1.3% are from off‐island sources. The limited present‐day tree distribution on the transect is discussed in the context of the more widespread distribution of arboreal pollen. Overall, there is a strong numerical correspondence between vegetation, pollen and environmental variables. The various problems inherent in examining surface pollen spectra are reviewed.     

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.     

库姆塔格沙漠南缘荒漠植物群落多样性分析

 根据20个样地的调查资料,应用重要值计算多样性指数、均匀度指数、丰富度指数 、优势度指数,对库姆塔格沙漠南缘荒漠植物群落物种多样性进行分析。结果表明: 1）荒漠植物群落分布随其生境地貌不同而不同,山前戈壁上分布有合头草(Sympegma regelii)群落,冲积河道低地分布有荒漠林胡杨(Populus euphratica)、多枝柽柳(Tamarix ramosissima)、胀果甘草（Glycyrrhiza inflata）群落,戈壁沙漠过渡带为梭梭(Haloxylon ammodendron)群落,低海拔的沙山上分布有沙拐枣(Calligonum mongolicum)群落、膜果麻黄(Ephedra rzewalskii)群落和梭梭群落。2）荒漠植物群落物种多样性水平较低,群落结构简单,物种组成单一。群落Shannon_Wiener物种多样性水平表现为合头草群落最高（1.706）,具有草原化荒漠植被类型的成分;梭梭群落、膜果麻黄群落居中（0.875～0.890）,荒漠植被类型特征明显;沙拐枣群落、胡杨群落、多枝 柽柳群落、胀果甘草群落较低（0.079～0.495）,荒漠林、盐地沙生灌丛及盐化草甸植被均有零星分布。3）荒漠植物群落结构层次中,灌木层占居主导地位,群落灌木层物种多样性水平（0.769～1.451）远远大于草本层（0.193～0.254）,且草本层物种多样性受灌木层影响较大。4）荒漠植物群落物种多样性分布格局表现为经向、纬向和海拔梯度的变化,经向、纬向变化为物种多样性水平较高的草原化植物合头草群落（1.706）向物种多样性水平较低的荒漠植物梭梭群落（1.379）和盐化植物多枝柽柳群落（0.376）的过渡,海拔梯度则 呈现低水平的沙拐枣群落（0.819）到高水平的膜果麻黄群落（0.890）向低水平的梭梭群落 （0.645）变化。荒漠植物群落过渡地带一般具有较高的物种多样性和较低的生态优势度。