Numerical analysis of surface and fossil pollen spectra from northern Fennoscandia

Aim Determination of the main directions of variance in an extensive data base of annual pollen deposition, and the relationship between pollen data from modified Tauber traps and palaeoecological data. Location Northern Finland and Norway. Methods Pollen analysis of annual samples from pollen traps and contiguous high‐resolution samples from a peat sequence. Numerical analysis (principal components analysis) of the resulting data. Results The main direction of variation in the trap data is due to the vegetation region in which each trap is located. A secondary direction of variation is due to the annual variability of pollen production of some of the tree taxa, especially Betula and Pinus. This annual variability is more conspicuous in ‘absolute’ data than it is in percentage data which, at this annual resolution, becomes more random. There are systematic differences, with respect to peat‐forming taxa, between pollen data from traps and pollen data from a peat profile collected over the same period of time. Main conclusions Annual variability in pollen production is rarely visible in fossil pollen samples because these cannot be sampled at precisely a 12‐month resolution. At near‐annual resolution sampling, it results in erratic percentage values which do not reflect changes in vegetation. Profiles sampled at near annual resolution are better analysed in terms of pollen accumulation rates with the realization that even these do not record changes in plant abundance but changes in pollen abundance. However, at the coarser temporal resolution common in most fossil samples it does not mask the origin of the pollen in terms of its vegetation region. Climate change may not be recognizable from pollen assemblages until the change has persisted in the same direction sufficiently long enough to alter the flowering (pollen production) pattern of the dominant trees.     

Differentiation between Neotropical rainforest,dry forest,and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record

Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerradão) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples.Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerradão. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerradão). Anadenanthera was found to be abundant in both the pollen and flora of SDTF (p/v = 3.6), while Poaceae was relatively poorly represented in cerradão (0.2). Moraceae, Cecropia and Schefflera were found to be over-represented palynologically in all ecosystems. Overall, the data demonstrated that no one taxon could be used as a definitive indicator of any of the ecosystems. Instead, associations of taxa were found to be important: METF = Moraceae (> 40%), Cecropia, Hyeronima, Celtis; SDTF = Anadenanthera, Apuleia, Ferdinandusa and non-arboreal Asteraceae, Bromeliaceae, Piper and fern spores; cerradão = Poaceae, Myrtaceae, Borreria, Solanum plus Asteraceae and fern spores. Interpretation of Poaceae pollen was highlighted as problematic, with relatively low abundance in the cerradão (< 20%) in comparison to high abundance in lake environments (c. 30–50%). Re-examination of fossil pollen records from NKMNP revealed that modern vegetation associations were only established in the last few thousand years.     

Quantifying uncertainty from large-scale model predictions of forest carbon dynamics

Linking environmental computer simulation models and geographic information systems (GIS) is now a common practice to scale up simulations of complex ecosystem processes for decision support. Unfortunately, several important issues of upscaling using GIS are rarely considered; in particular scale dependency of models, availability of input data, support of input and validation data, and uncertainty in prediction including error propagation from the GIS. We linked the biogeochemical Forest‐DNDC model to a GIS database to predict growth of Eucalyptus globulus plantations at two different scales (～0.045 ha plot^?1 scale and ～100 ha grid^?1 scale) across Victoria, in south‐eastern Australia. Results showed that Forest‐DNDC was not scale dependent across the range of scales investigated. Reduced availability of input data at the larger scale may introduce severe prediction errors, but did not require adjustment of the model in this study. Differences in the support of input and validation data led to an underestimation of predictive precision but an overestimation of prediction accuracy. Increasing data support, produced a high level of prediction accuracy (^?e%), but a medium level of predictive precision (r²=0.474, ME=0.318) after statistical validation. GIS error contribution could be detected but was not readily or reliably quantified. In a regional case study for 2653 ha of E. globulus plantations, the linked model GIS system estimated a total standing biomass of 95 260 t C for mid‐2003 and a net CO₂ balance of ?45 671 t CO₂‐C yr^?1 for the entire year of 2002. This study showed that regional predictions of forest growth and carbon sequestration can be produced with greater confidence after a comprehensive assessment of upscaling issues.     

Evaluating the effect of flowering age and forest structure on pollen productivity estimates

Pollen productivity estimates (PPEs) are indispensable prerequisites for quantitative vegetation reconstructions. Estimates from different European regions show a large variability and it is uncertain whether this reflects regional differences in climate and soil or is brought about by different assessments of vegetation abundance. Forests represent a particular problem as they consist of several layers of vegetation and many tree species only start producing pollen after they have attained ages of several decades. Here we used detailed forest inventory data from north-eastern Germany to investigate the effect of flowering age and understory trees on PPEs. Pollen counts were obtained from 49 small to medium sized lakes chosen to represent the different forest types in the region. Surface samples from lakes within a closed forest of Fagus yielded disproportionate amounts of Fagus pollen, increasing its PPE and the variability of all other estimates. These samples were removed from further analysis but indicate a high trunk-space component that is not considered in the Prentice–Sugita pollen dispersal and deposition model. Results of the restricted dataset show important differences in PPEs based on the consideration of flowering age and understory position. The effect is largest for slow growing and/or late flowering trees like Fagus and Carpinus while it is minimal for species that flower early in their development like Betula and Alnus. The large relevant source area of pollen (RSAP) of 7?km obtained in this study is consistent with the landscape structure of the region.     

Relative pollen productivity estimates of major plant taxa and relevant source area of pollen in the warm-temperate forest landscape of northern China

Vegetation History and Archaeobotany - Relative pollen productivity (RPP) is critical for quantitative vegetation reconstruction of past vegetation cover. The Extended R-value (ERV) model is...     

Effects of aggregated classifications of forest composition on estimates of evapotranspiration in a northern Wisconsin forest

Forest management presents challenges to accurate prediction of water and carbon exchange between the land surface and atmosphere, due to its alteration of forest structure and composition. We examined how forest species types in northern Wisconsin affect landscape scale water fluxes predicted from models driven by remotely sensed forest classification. A site‐specific classification was developed for the study site. Using this information and a digital soils database produced for the site we identified four key forest stand types: red pine, northern hardwoods, aspen, and forested wetland. Within these stand types, 64 trees representing 7 species were continuously monitored with sap flux sensors. Scaled stand‐level transpiration from sap flux was combined with a two‐source soil evaporation model and then applied over a 2.5 km × 3.0 km area around the WLEF AmeriFlux tower (Park Falls, Wisconsin) to estimate evapotranspiration. Water flux data at the tower was used as a check against these estimates. Then, experiments were conducted to determine the effects of aggregating vegetation types to International Geosphere– Biosphere Program (IGBP) level on water flux predictions. Taxonomic aggregation resulting in loss of species level information significantly altered landscape water flux predictions. However, daily water fluxes were not significantly affected by spatial aggregation when forested wetland evaporation was included. The results demonstrate the importance of aspen, which has a higher transpiration rate per unit leaf area than other forest species. However, more significant uncertainty results from not including forested wetland with its high rates of evaporation during wet summers.     

Quantitative representation of local forest composition in forest‐floor pollen assemblages

1 We compared modern pollen assemblages from 60 moss polster sites in northern New York with forest composition data within 20–120 m of the sites using extended R ‐value (ERV) models, which correct for non‐linearities arising from use of pollen percentage data. Our sites were concentrated in two regions, one dominated by Tsuga and hardwood ( Acer , Betula , Fagus ) forests, and the other by Tsuga , Pinus , Betula , Acer and Quercus forests.
2 Our results confirm that forest‐floor pollen assemblages are dominated by pollen originating from trees growing more than 20 m from the site of deposition. However, our results suggest that background pollen percentages were overestimated by Jackson & Wong in 1994, owing to unusually high Pinus pollen production in the year of their sampling.
3 Expansion of our vegetation sampling radius from 20 to 120 m resulted in modest but consistent improvement in model fit and a decrease in background pollen percentages.
4 ERV model parameters (slope and background) differed substantially between the two study regions, primarily owing to differences in background pollen productivity and dispersal from regional sources.
5 High background pollen percentages may lead to poor estimation of calibration parameters in regions of complex vegetation patterns. Expansion of the vegetation sampling radius to reduce the background component may lead to better parameter estimates.
6 Calibration of pollen–vegetation relationships requires definition of the vegetation term so that it approximates the vegetation sampled by the pollen assemblages. Critical challenges are to define better the appropriate vegetation sampling area and distance‐weighting functions for application to pollen–vegetation calibration.     

Power spectra of extinction in the fossil record

Recent Fourier analyses of fossil extinction data have indicated that the power spectrum of extinction during the Phanerozoic may take the form of 1/f noise, a result which, it has been suggested, could be indicative of the presence of `critical dynamics'' in the processes giving rise to extinction. In this paper we examine extinction power spectra in some detail, using family-level data from two widely available compilations. We find that although the average form of the power spectrum roughly obeys the 1/f law, the spectrum can be represented more accurately by dividing it into two regimes: a low-frequency one which is well fit by an exponential, and a high-frequency one in which it follows a power law with a 1/f² form. We give explanations for the occurrence of each of these behaviours and for the position of the crossover between them.     

Evaluation of uncertainty in alignment tensors obtained from dipolar couplings

Residual dipolar couplings and their corresponding alignment tensors are useful for structural analysis of macromolecules. The error in an alignment tensor, derived from residual dipolar couplings on the basis of a known structure, is determined not only by the accuracy of the measured couplings but also by the uncertainty in the structure (structural noise). This dependence is evaluated quantitatively on the basis of simulated structures using Monte-Carlo type analyses. When large numbers of dipolar couplings are available, structural noise is found to result in a systematic underestimate of the magnitude of the alignment tensor. Particularly in cases where only few dipolar couplings are available, structural noise can cause significant errors in best-fitted alignment tensor values, making determination of the relative orientation of small fragments and evaluation of local backbone mobility from dipolar couplings difficult. An example for the protein ubiquitin demonstrates the inherent limitations in characterizing motions on the basis of local alignment tensor magnitudes.     

Four well-constrained calibration points from the vertebrate fossil record for molecular clock estimates

Recent controversy about the use of the vertebrate fossil record for external calibration of molecular clocks centers on two issues, the number of dates used for calibration and the reliability of the fossil calibration date. Viewing matters from a palaeontological perspective, we propose three qualitative, phylogenetic criteria that can be used within a comparative framework for the selection of well-constrained calibration dates from the vertebrate fossil record. On the basis of these criteria, we identify three highly suitable new fossil calibration dates for molecular clock estimates: the lungfish-tetrapod split (between 419 and 408 Ma), the bird-crocodile split (between 251 and 243 Ma), and the alligator-caiman split (between 71 and 66 Ma). Together with our previously suggested bird-lizard split (between 252 and 257 Ma), these four fossil dates span a range of very different ages. They are, in our opinion, more suitable for molecular clock calibration than the traditionally used mammal-bird split, which is less well constrained. We plea for closer interactions between paleontologists and molecular biologists in studying the timescale of vertebrate evolution.     

Changes in nutritional composition from bee pollen to pollen patty used in bumblebee rearing

Pollen patties are generally supplied to augment the nourishment of honey bee or bumblebee colonies during late winter or early spring. In the present study, we examined nutrient content of bee collected pollen (BP) and pollen patties (PT) prepared from those bee collected pollens in order to understand the chemical compositional change of pollen to pollen patty and figure out the possible benefits for bee’s health. Protein and fat contents of pollen patties were found lower than that reported for the respective bee pollens. Amino and fatty acids followed the same trend for the simple logical reason. In contrast, carbohydrate contents of pollen patties were found much higher than the bee pollen. The addition of sugar solution in the process of pollen patty preparation could presumably explain the change. Another possible determinant factor of bee’s preference of feed protein to fat ratio was found the highest for PT 2 (5.5) followed by PT 1 and BP 2 (4.1), BP Ref. (2.9) and the least value obtained for BP 1 (2.2). Additionally, we discuss the possible health benefits if the bee collected pollen is used as a human food supplement.     

Assessing the quality of molecular divergence time estimates by fossil calibrations and fossil-based model selection

Estimates of species divergence times using DNA sequence data are playing an increasingly important role in studies of evolution, ecology and biogeography. Most work has centred on obtaining appropriate kinds of data and developing optimal estimation procedures, whereas somewhat less attention has focused on the calibration of divergences using fossils. Case studies with multiple fossil calibration points provide important opportunities to examine the divergence time estimation problem in new ways. We discuss two cross-validation procedures that address different aspects of inference in divergence time estimation. 'Fossil cross-validation' is a procedure used to identify the impact of different individual calibrations on overall estimation. This can identify fossils that have an exceptionally large error effect and may warrant further scrutiny. 'Fossil-based model cross-validation' is an entirely different procedure that uses fossils to identify the optimal model of molecular evolution in the context of rate smoothing or other inference methods. Both procedures were applied to two recent studies: an analysis of monocot angiosperms with eight fossil calibrations and an analysis of placental mammals with nine fossil calibrations. In each case, fossil calibrations could be ranked from most to least influential, and in one of the two studies, the fossils provided decisive evidence about the optimal molecular evolutionary model.     

Structure of a corystosperm fossil forest from the Late Triassic of Argentina

A palaeoecological study of a standing Late Triassic forest containing 150 silicified stumps from the Río Blanco Formation of Mendoza province, Argentina is described. A mapped portion of the forest floor provides quantitative data — tree density, mean separation of trees and basal area per stump — which, along with taxonomic and sedimentologic information, allowed the reconstruction of a plant community that grew along river banks and within proximal floodplain environments. Analysis of architectural and phenological data from monotypic forest indicates an evergreen community composed of a corystosperm genus with a canopy height of c. 13–21 m. The corystosperm taxon: Elchaxylon, like Rhexoxylon, has polyxylic axes with centripetal secondary xylem but does not generate perimedullar bundles and the centrifugal secondary xylem produces an undivided solid pycnoxylic cylinder. Vegetation analysis shows that the forest has a clustered distribution pattern with high density. Forest density ranges between 727 and 1504 trees/ha but there are first order clusters with elevated density (mean nearest neighbour distance of 1.85 m). The histogram of diameter classes based on 131 stumps suggests an earlier colonization by few older pioneers (the largest ones) followed by establishment of a large younger cohort of coeval trees. Based on 9 series and 139 rings, the mean ring width and mean sensitivity (MS) were 3.47 mm and 0.31 respectively. MS values and the presence of false rings indicate the forest community responded to stressed ecosystems. The growth rings are very erratic and range from 0.27 to 8.94 mm. For fossil growth analysis it was assumed that wider rings would suggest a warmer climate and the considerable range in growth rates would indicate variability in the limiting factors among subsequent cycles.     

Robustness and uncertainty in estimates of butterfly abundance from transect counts

Many butterfly populations are monitored by counting the number of butterflies observed while walking transects during the butterfly’s flight season. Methods for estimating population abundance from these transect counts are appealing because they allow rare populations to be monitored without capture–recapture studies that could harm fragile individuals. An increasingly popular method for estimating abundance from transect counts relies on strong assumptions about the counting process and the processes that govern butterfly population dynamics. Here, we study the statistical performance of this method when underlying model assumptions are violated. We find that estimates of population size are robust to departures from underlying model assumptions, but that the uncertainty in these estimates (i.e., confidence intervals) is substantially underestimated. Alternative bootstrap and Bayesian methods provide better measures of the uncertainty in estimated population size, but are conditional upon knowledge of butterfly detectability. Because of these requirements, a mixed approach that combines data from small capture–recapture studies with transect counts strikes the best balance between accurate monitoring and minimal injury to individuals. Our study is motivated by monitoring studies for St. Francis satyr (Neonympha mitchelli francisci), a rare and relatively immobile butterfly occurring only in the sandhills region of south-central North Carolina, USA.     

On the interpretation of fossil Poaceae pollen in the lowland humid neotropics

Poaceae pollen is abundant in fossil records and is often used as a paleoclimatic indicator. A common interpretation is to link increases in Poaceae pollen abundance to increased regional aridity. However, the representation of Poaceae pollen is influenced by a number of factors, such as the proportion of other plants in the flora that are anemophilous, the size of local marshes, and the influence of humans on the landscape. Abundant anemophilous trees are likely to mask the contribution of Poaceae pollen, whereas floras that are primarily entomophilous are likely to produce a pollen spectrum containing an over-representation of Poaceae. As most fossil pollen data are drawn from flooded settings, it is critically important that palynologists recognize the Poaceae pollen contribution derived from floating grasses and marshes that surround their coring site. Interpretations that ignore changes in effective lake size and assume that Poaceae percentage is a simple indicator of regional vegetation change are likely to overstate ‘dry’ episodes and transitions from wet forest to scrub environments. Human occupation of a site is sometimes manifested in increased Poaceae pollen abundance in lake sediments. The duration, intensity and land-use associated with occupation are all variables that influence Poaceae pollen representation. Very high (50–90%) abundances of Poaceae pollen provide a strong indicator of savanna habitats, but trying to determine transitional vegetation types between savanna and wet forest is best determined using other taxa. Indeed, reliance on Poaceae abundance as an indicator of paleoprecipitation is potentially very misleading when it is in the fossil record at moderate abundances.     

Bootstrap estimates of the statistical accuracy of thresholds obtained from psychometric functions

Estimates of the accuracy of a threshold obtained from a psychometric function are often based on asymptotic theory. When the number of trails is small, however, these estimates may be untrustworthy. A computer program is described that uses a move reliable bootstrap approach to obtaining estimates of the standard deviation and confidence limits of a threshold and of the slope and spread of the psychometric function for any criterion level of performance.     

Ephedroid fossil pollen from the Lower Cretaceous (Upper Albian) of Hokkaido,Japan

This paper presents eight ephedroid palynomorphs from the Lower Cretaceous (Upper Albian) of Hokkaido, Japan. The ephedroid palynomorphs are ellipsoid and polyplicate pollen grains that show a wide range of variation in pollen size, shape, and plication. These ephedroid palynomorphs suggest a wide range of diversity in Gnetales at mid- or high-paleolatitude in the eastern side of Lauresia during the Upper Albian.     

Numerical analysis of modern and fossil pollen spectra as a tool for elucidating the nature of fine-scale human activities in boreal areas

Modern pollen assemblages from the major vegetation units (both natural and anthropogenic) on the island of Hailuoto, Finland are studied from 29 surface moss samples. A total of 59 pollen and spore taxa are recorded. The pollen data-set is related by redundancy analysis (RDA) to six external synthetic variables that characterize in general terms the different major ecological situations relevant to the land-use on the island (sand, humus, forest, field, deforested, treeless). Weighted average (WA) optima are calculated to identify which pollen taxa are most indicative of four of these external variables. Two fossil pollen diagrams from Hailuoto are re-interpreted by positioning their samples on the RDA plot of the modern data-set and by classifying the modern and fossil spectra together in a minimum-variance cluster analysis. The RDA plot shows that the strongest features in the modern pollen assemblages are the contrast between spectra from dry sandy soils and those from damper soils with more organic content, and between assemblages from forested and unforested areas. For anthropogenic situations the numerical analyses detect a distinction in the pollen assemblages characterizing fields from those representing farms and trackways but is less successful in separating the latter two. The WA optima indicate that most pollen taxa considered as indicator types primarily represent a mixed field/farm category of land-use. The RDA and minimum-variance cluster analysis confirm that the two fossil sites have had different vegetational histories. In neither is there any phase equivalent to the modern farm situation on Hailuoto. The initial phase in each diagram represents shore meadows and fields but because Poaceae pollen is identifiable only to family level it is not possible to separate these two habitat types.