Biomarkers, stable carbon isotope record and pollen distribution of the soutwest African continental margin

This is part 2 of a study examining southwest African continental margin sediments from nine sites on a north-south transect from the Congo Fan (4°S) to the Cape Basin (30°S) representing two glacial (MIS 2 and 6a) and two interglacial stages (MIS 1 and 5e). Contents, distribution patterns, and molecular stable carbon isotope signatures of long-chain n-alkanes (C27-C33) and n-alkanols (C22-C32) as indicators of land plant vegetation of different biosynthetic types were correlated with concentrations and distributions of pollen taxa in sediments of the same time horizons. Selected single pollen type data reveal details of vegetation changes, but the overall picture is best illustrated by summing pollen known to predominantly derive from C4 plants or C4 plus CAM plants. The C4 plant signals in the biomarkers are recorded in the delta13C data and in the abundances of C31 and C33 n-alkanes, and the C32 n-alkanol. Calculated clusters of wind trajectories for austral summer and winter situations for the Holocene and the Last Glacial Maximum afford information on the source areas for the lipids and pollen and their transport pathways to the ocean. This multidisciplinary approach provides clear evidence of latitudinal differences in leaf wax lipid and pollen composition, with the Holocene sedimentary data paralleling the current major phytogeographic zonations. The northern sites (Congo Fan area and northern Angola Basin) get most of their terrestrial material from the Congo Basin and the Angolan highlands dominated by C3 plants. Airborne particulates derived from the western and central South African hinterland dominated by deserts, semideserts, and savannah regions are rich in organic matter from C4 plants. As can be expected from the present and glacial positions of the phytogeographic zones, the carbon isotopic signatures of n-alkanes and n-alkanols both become isotopically more enriched in 13C from north to south. In the northern part of the transect the relative importance of C4 plant indicators is higher during the glacials than in the interglacials, indicating a northward extension of arid zones favoring grass vegetation. In the south, where grass-rich vegetation merges into semidesert and desert, the difference in C4 plant indicators is small.

Distribution of deep-sea foraminifera in surface sediments east of New Zealand

Paleobathymetric assessments of fossil foraminiferal faunas play a significant role in the analysis of the paleogeographic, sedimentary, and tectonic histories of New Zealand's Neogene marine sedimentary basins. At depths >100 m, these assessments often have large uncertainties. This study, aimed at improving the precision of paleodepth assessments, documents the present-day distribution of deep-sea foraminifera (>63 µm) in 66 samples of seafloor sediment at 90-700 m water depth (outer shelf to mid-abyssal), east of New Zealand. One hundred and thirty-nine of the 465 recorded species of benthic foraminifera are new records for the New Zealand region. Characters of the foraminiferal faunas which appear to provide the most useful information for estimating paleobathymetry are, in decreasing order of reliability: relative abundance of common benthic species; benthic species associations; upper depth limits of key benthic species; and relative abundance of planktic foraminifera. R mode cluster analysis on the quantitative census data of the 58 most abundant species of benthic foraminifera produced six species associations within three higher level clusters: (1) calcareous species most abundant at mid-bathyal to outer shelf depths (<1000 m); (2) calcareous species most abundant at mid-bathyal and greater depths (>600 m); (3) agglutinated species mostly occurring at deep abyssal depths (>3000 m). A detrended correspondence analysis ordination plot exhibits a strong relationship between these species associations and bathymetry. This is manifest in the bathymetric ranges of the relative abundance peaks of many of the common benthic species (e.g., Abditodentrix pseudothalmanni 500-2800 m, Bolivina robusta 200-650 m, Bulimina marginata f. marginata 20-600 m, B. marginata f. aculeata 400-3000 m, Cassidulina norvangi 1000-4500 m, Epistominella exigua 1000-4700 m, and Trifarina angulosa 10-650 m), which should prove useful in paleobathymetric estimates. The upper depth limits of 28 benthic foraminiferal species (e.g., Fursenkoina complanata 200 m, Bulimina truncana 450 m, Melonis affinis 550 m, Eggerella bradyi 750 m, and Cassidulina norvangi 1000 m) have potential to improve the precision of paleobathymetric estimates based initially on the total faunal composition. The planktic percentage of foraminiferal tests increases from outer shelf to upper abyssal depths followed by a rapid decline within the foraminiferal lysocline (below c. 3600 m). A planktic percentage <50% is suggestive of shelf depths, and >50% is suggestive of bathyal or abyssal depths above the CCD. In the abyssal zone there is dramatic taphonomic loss of most agglutinated tests (except some textulariids) at burial depths of 0.1-0.2 m, which negates the potential usefulness of these taxa in paleobathymetric assessments.

Distribution of pollen and spores in sediments of ODP Hole 175-1078C

The distribution of pollen in marine sediments is used to record vegetation change on the continent. Generally, a good latitudinal correspondence exists between the distribution patterns of pollen in the marine surface sediments and the occurrence of the source plants on the adjacent continent. To investigate land-sea interactions during deglaciation, we compare proxies for continental (pollen assemblages) and marine conditions (alkenone-derived sea surface temperatures) of two high-resolution, radiocarbon-dated sedimentary records from the tropical southeast Atlantic. The southern site is located West of the Cunene River mouth; the northern site is located West of the Angolan Huambe Mountains. It is inferred that the vegetation in Angola developed from Afroalpine and open savannah during the last Glacial maximum (LGM) via Afromontane Podocarpus forest during Heinrich Event 1 (H1), to an early increase of lowland forest after 14.5 ka. The vegetation record indicates dry and cold conditions during the LGM, cool and wet conditions during H1 and a gradual rise in temperature starting well before the Younger Dryas (YD) period. Terrestrial and oceanic climate developments seem largely running parallel, in contrast to the situation ca. 5° further South, where marine and terrestrial developments diverge during the YD. The cool and wet conditions in tropical West Africa, South of the equator, during H1 suggest that low-latitude insolation variation is more important than the slowdown of the thermohaline circulation for the climate in tropical Africa.

Three species distribution models for the marine diatom Fragilariopsis kerguelensis of the Southern Ocean, links to NetCDF files

Fragilariopsis kerguelensis, a dominant diatom species throughout the Antarctic Circumpolar Current, is coined to be one of the main drivers of the biological silicate pump. Here, we study the distribution of this important species and expected consequences of climate change upon it, using correlative species distribution modeling and publicly available presence-only data. As experience with SDM is scarce for marine phytoplankton, this also serves as a pilot study for this organism group. We used the maximum entropy method to calculate distribution models for the diatom F. kerguelensis based on yearly and monthly environmental data (sea surface temperature, salinity, nitrate and silicate concentrations). Observation data were harvested from GBIF and the Global Diatom Database, and for further analyses also from the Hustedt Diatom Collection (BRM). The models were projected on current yearly and seasonal environmental data to study current distribution and its seasonality. Furthermore, we projected the seasonal model on future environmental data obtained from climate models for the year 2100. Projected on current yearly averaged environmental data, all models showed similar distribution patterns for F. kerguelensis. The monthly model showed seasonality, for example, a shift of the southern distribution boundary toward the north in the winter. Projections on future scenarios resulted in a moderately to negligibly shrinking distribution area and a change in seasonality. We found a substantial bias in the publicly available observation datasets, which could be reduced by additional observation records we obtained from the Hustedt Diatom Collection. Present-day distribution patterns inferred from the models coincided well with background knowledge and previous reports about F. kerguelensis distribution, showing that maximum entropy-based distribution models are suitable to map distribution patterns for oceanic planktonic organisms. Our scenario projections indicate moderate effects of climate change upon the biogeography of F. kerguelensis.

Distribution of marine invertebrate larvae in the southern Kara Sea (Russian Arctic)

Within the generally oligotroph Arctic marine environment river outlets are favoured by many planktonic and benthic organisms due to their high input of organic carbon. The retention of pelagic larvae within nursery grounds and/or the ability to return to their parental grounds prior to settlement is one important factor for the persistence of benthic communities in such river influenced areas. The southern Kara Sea is strongly controlled by high freshwater inputs from the Ob and Yenisei Rivers, which create a pronounced bi-layered pycnocline with a warm fresh/brackish water layer on top and a cold high saline marine layer below. The dispersal of five meroplanktonic species and settled juveniles (the brittle star Ophiocten sericeum, and the polychaetes Micronephtys minuta, Nereimyra aphroditoides, Phyllodoce groenlandica and Prionospio cirrifera) in relation to the adult distribution patterns was investigated. For all apart from P. cirrifera the highest densities of larvae were found in the upper brackish water layer. To assess size-at-settlement, the body sizes of larvae and newly settled juveniles were estimated and compared. Dispersal patterns ranged from virtually no adaption to river run-off as in the common, stenohaline O. sericeum and M. minuta (7 ind./m**3, 459 µm) to local retention as in N. aphroditoides (7 ind./m**3, 541 µm) and P. groenlandica (0.5 ind./m**3, 1121 µm) retained by horizontal eddies created by the outflow. Adults of P. cirrifera, which were exclusively restricted to the estuary of the Yenisei River, showed a well adapted reproductive behaviour to ensure a high retention potential of their progenies. The larvae (1.5 ind./m**3, 1513 µm) were only present in the lower water layers, most probably taking advantage of the prevailing near bottom counter current retaining them within their hatching areas.

Calcareous dinocysts and grain size distribution along profile Stevns-Klint

The distribution patterns of calcareous dinoflagellate cysts were studied in the classic Cretaceous Tertiary (K-T) boundary section of Stevns Klint, Denmark, focusing mainly on the response of the cyst association to an abrupt environmental catastrophe. A major part of the Fish Clay, which covers the K-T boundary at its base and is exposed in the investigated section, contains fallout produced by an asteroid impact. Calcareous dinoflagenate cysts are the best preserved remains of carbonate-producing phytoplankton in this layer. The potential of this group of microfossils for the analysis of survival strategies and extinction patterns has been underestimated. The cyst species of the investigated section can be grouped into four assemblages that represent victims, survivors, opportunists, and specially adapted forms. The victims (Pithonelloideae) were an extremely successful group throughout the Upper Cretaceous, but were restricted to the narrow outer shelf. This restriction minimized their spatial distribution, which generally should be large to facilitate escape from unfavorable conditions. Spatial restriction optimized the population decrease by mass mortality, disabling a successful recovery. In contrast, the survivors that became the dominating group in the Danian had a wide spatial range from the shelf environment to the oceanic realm. A unique calcareous dinocyst assemblage in the Fish Clay shows that even under the stressed conditions immediately following the impact event, some species flourished due to special adaptation or high ecological tolerance. The ability of these dinoflagellate species to form calcareous resting cysts in combination with their generally wide spatial distribution in a variety of environments appears to be the main reason for a low extinction rate at the K-T boundary as opposed to the high extinction rate of other phytoplankton groups, such as the coccolithophorids.

Pollen distribution in marine sediment samples along the south-western African coast

The distribution of pollen in marine sediments is used to reconstruct pathways of terrigenous input to the oceans and provides a record of vegetation change on adjacent continents. The wind transport routes of aeolian pollen is comprehensively illustrated by clusters of trajectories. Isobaric, 4-day backward trajectories are calculated using the modelled wind-field of ECHAM3, and are clustered on a seasonal basis to estimate the main pathways of aeolian particles to sites of marine cores in the south-eastern Atlantic. Trajectories and clusters based on the modelled wind-field of the Last Glacial Maximum hardly differ from those of the present-day. Trajectory clusters show three regional, and two seasonal patterns, determining the pathways of aeolian pollen transport into the south-eastern Atlantic ocean. Mainly, transport out of the continent occurs during austral fall and winter, when easterly and south-easterly winds prevail. South of 25°S, winds blow mostly from the west and southwest, and aeolian terrestrial input is very low. Generally, a good latitudinal correspondence exists between the distribution patterns of pollen in marine surface sediments and the occurrence of the source plants on the adjacent continent. The northern Angola Basin receives pollen and spores from the Congolian and Zambezian forests mainly through river discharge. The Zambezian vegetation zone is the main source area for wind-blown pollen in sediments of the Angola Basin, while the semi-desert and desert areas are the main sources for pollen in sediments of the Walvis Basin and on the Walvis Ridge. A transect of six marine pollen records along the south-western African coast indicates considerable changes in the vegetation of southern Africa between glacial and interglacial periods. Important changes in the vegetation are the decline of forests in equatorial Africa and the north of southern Africa and a northward shift of winter rain vegetation along the western escarpment.

Dinoflagellate cysts and pollen distribution of marine surface sediments off West Africa

An organic-walled dinoflagellate cyst analysis was carried out on 53 surface sediment samples from West Africa (17-6°N) to obtain insight in the relationship between their spatial distribution and hydrological conditions in the upper water column as well as marine productivity in the study area. Multivariate analysis of the dinoflagellate cyst relative abundances and environmental parameters of the water column shows that sea-surface temperature, salinity, marine productivity and bottom water oxygen are the factors that relate significantly to the distribution patterns of individual species in the region. The composition of cyst assemblages and dinoflagellate cyst concentrations allows the identification of four hydrographic regimes; 1) the northern regime between 17 and 14°N characterized by high productivity associated with seasonal coastal upwelling, 2) the southern regime between 12 and 6°N associated with high-nutrient waters influenced by river discharge 3) the intermediate regime between 14 and 12°N influenced mainly by seasonal coastal upwelling additionally associated with fluvial input of terrestrial nutrients and 4) the offshore regime characterized by low chlorophyll-a concentrations in upper waters and high bottom water oxygen concentrations. Our data show that cysts of Polykrikos kofoidii, Selenopemphix quanta, Dubridinium spp., Echinidinium species, cysts of Protoperidinium monospinum and Spiniferites pachydermus are the best proxies to reconstruct the boundary between the NE trade winds and the monsoon winds in the subtropical eastern Atlantic Ocean. The association of Bitectatodinium spongium, Lejeunecysta oliva, Quinquecuspis concreta, Selenopemphix nephroides, Trinovantedinium applanatum can be used to reconstruct past river outflow variations within this region.

Distribution of organic-walled dinoflagellate cysts in surface sediments of the Benguela upwelling system

To obtain insight in the relationship between the spatial distribution of organic-walled dinoflagellate cysts (dinocysts) and local environmental conditions, fifty-eight surface sediment samples from the coastal shelf off SW Africa were investigated on their dinocyst content with special focus on the two main river systems and the active upwelling that characterise this region. To avoid possible overprint by species-selective preservation, samples have been selected mainly from shelf sites where high sedimentation rates and/or low bottom water oxygen concentrations prevail. Multivariate ordination analyses have been carried out to investigate the relationship between the distribution patterns of individual species to environmental parameters of the upper water column and sediment transport processes. The main oceanographical variables at the surface (temperature, salinity, nutrients chlorophyll-a) in the region show onshore-offshore gradients. This pattern is reflected in the dinocyst associations with high relative abundances of heterotrophic dinocyst species in neritic regions characterised by high chlorophyll-aand low salinity conditions in surface waters. Phototrophic dinocyst species, notably Operculodinium centrocarpum, dominate in the more oceanic area. Differences in the distribution of phototrophic dinocyst species can be related to sea surface salinity and sea surface temperature gradients and to a lesser extent to chlorophyll-a concentrations. Apart from longitudinal gradients the dinocyst distribution clearly reflects regional environmental features. Six groups of species can be distinguished, characteristic for (1) coastal regions (cysts of Polykrikos kofoidii and Selenopemphix quanta), (2) the vicinity of active upwelling (Brigantedinium spp., Echinidinium aculeatum, Echinidinium spp. and Echinidinium transparantum), (3) river mouths (Lejeunecysta oliva, cysts of Protoperidinium americanum, Selenopemphix nephroides and Votadinium calvum), (4) slope and open ocean sediments (Dalella chathamense, Impagidinium patulum and Operculodinium centrocarpum, (5) the southern Benguela region (south of 24°S) (Spiniferites ramosus) and (6) the northern Benguela region (north of 24°S) (Nematosphaeropsis labyrinthus and Pyxidinopsis reticulata). No indication of overprint of the palaeo-ecological signal by lateral transport of allochthonous species could be observed.

Appendix 7: List of species with their distribution ranges restricted to the corresponding phytogeographical region

Understanding species distribution patterns and the corresponding environmental determinants is a crucial step in the development of effective strategies for the conservation and management of plant communities and ecosystems. Therefore, a central prerequisite is the biogeographical and macroecological analysis of factors and processes that determine contemporary, potential, as well as future geographic distribution of species. This thesis has been conducted in the framework of the BIOMAPS-BIOTA project at the Nees Institute of Biodiversity of Plants, which was funded by the German Federal Ministry of Education and Research (BMBF). The study investigated patterns of plants species richness and phytogeographic regions under contemporary environmental conditions and forecasted future climate change in the area of West Africa covering five countries: Benin, Burkina Faso, Côte d'Ivoire, Ghana and Togo. Firstly, geographic patterns of vascular plant species richness have been depicted at a relatively fine spatial resolution based on the potential distribution of 3,393 species. Species richness is closely related to the steep climatic gradient existing in the region with a high concentration of species in the most humid areas in the south and decreases towards the northern drier areas. The investigation of the effectiveness of the existing network of protected areas shows an overall good coverage of species in the study area. However, the proportion of covered species is considerably lower at national extent for some countries, thus calling for more protected areas in order to cover adequately a maximum number of plants species in these countries. Secondly, based on the potential distribution range of vascular plant species, seven phytogeographic regions have been delineated that broadly reflect the vegetation zones as defined by White (1983). However notable differences to the delineation of White (1983) occur at the margins of some regions. Corresponding to a general southward shifted of all regions. And expansion of the Sahel vegetation zone is observed in the north, while the rainforest zone is decreased in the very south.This is alarming since the rainforest shelters a high number of species and a high proportion of range-restricted or endemic species, despite their relatively small extent compared to the other regions. Finally, the evaluation of the potential impact of climate change on plant species richness in the study area, results in a severe loss of future suitable habitat for up to 50% of species per grid cell, particularly in the rainforest region. Moreover, the analysis of the possible shift of phytogeographic regions shows in general a strong deterioration of the West African rainforest. In contrast the drier areas are expanding continuously, although a slight gain in species number can be observed in some particular regions. The overall lesson to retain from the results of this study is that the West African rainforest should be fixed as a high priority area for the conservation of biodiversity of plants, since it is subject to severe contemporary and projected future threats.

Vertical distribution of pelagic polychaetes in Kuril-Kamchatka Trench plankton in August 1966

Vertical distribution patterns of four mass species of pelagic polychaetes (with separate consideration of the adult, juvenile and larval stages) in plankton of the Kuril-Kamchatka region from the surface to depth of 7000 m are analyzed. This material, the author's personal data on the Norwegian and Barents Seas, and literature data on other seas are used to distinguish some general patterns in vertical distribution and reproductive ecology of pelagic polychaetes in the World Ocean as a whole. Mass pelagic polychaetes are also compared to other bathypelagic animals with respect to these ecological features and an estimate is given of their abundance in ocean plankton.

Skill distribution and comparative advantage: a comparison of China and India

This paper empirically examines the different comparative advantages of two emerging economic giants, China and India, in relation to the different skill distribution patterns in each country. By utilizing industry export data on China and India from 1983 to 2000, we find that a country with a greater dispersion of skills (i.e., India, especially in the earlier years) has higher exports in industries with shorter production chains, whereas a country with a more equal dispersion of skills (i.e., China, especially in the later years) is found to have higher exports in industries with longer production chains. The causal relationship is fairly robust across different specifications. This empirical evidence supports our assumption that the likely mechanism for these results is the negative impact of low-skilled workers on input quality, which accumulates and becomes larger as the length of production chains and the proportion of low-skilled workers in the economy increase.

Saproxylic Beetle Assemblage Selection as Determining Factor of Species Distributional Patterns: Implications for Conservation

The knowledge of the distributional patterns of saproxylic beetles is essential for conservation biology due to the relevance of this fauna in the maintenance of ecological processes and the endangerment of species. The complex community of saproxylic beetles is shaped by different assemblages that are composed of species linked by the microhabitats they use. We evaluate how different the species distribution patterns that are obtained can be, depending on the analyzed assemblage and to what extent these can affect conservation decisions. Beetles were sampled using hollow emergence and window traps in three protected areas of the Iberian Peninsula. Species richness, composition, and diversity turnover were analyzed for each sampling method and showed high variation depending on the analyzed assemblage. Beta diversity was clearly higher among forests for the assemblage captured using window traps. This method collects flying insects from different tree microhabitats and its captures are influenced by the forest structuring. Within forests, the assemblages captured by hollow emergence traps, which collect the fauna linked to tree hollows, showed the largest turnover of species, as they are influenced by the characteristics of each cavity. Moreover, the selection of the forest showing the highest species richness strongly depended on the studied assemblage. This study demonstrates that differences in the studied assemblages (group of species co-occurring in the same habitat) can also lead to significant differences in the identified patterns of species distribution and diversity turnover. This fact will be necessary to take into consideration when making decisions about conservation and management.