Palynological and geochemical data of sediment core GeoB3910-2 located offshore Northeast Brazil

High resolution palynological and geochemical data of sediment core GeoB 3910-2 (located offshore Northeast Brazil) spanning the period between 19 600 and 14 500 calibrated year bp (19.6-14.5 ka) show a land-cover change in the catchment area of local rivers in two steps related to changes in precipitation associated with Heinrich Event 1 (H1 stadial). At the end of the last glacial maximum, the landscape in semi-arid Northeast Brazil was dominated by a very dry type of caatinga vegetation, mainly composed of grasslands with some herbs and shrubs. After 18 ka, considerably more humid conditions are suggested by changes in the vegetation and by Corg and C/N data indicative of fluvial erosion. The caatinga became wetter and along lakes and rivers, sedges and gallery forest expanded. The most humid period was recorded between 16.5 and 15 ka, when humid gallery (and floodplain) forest and even small patches of mountainous Atlantic rain forest occurred together with dry forest, the latter being considered as a rather lush type of caatinga vegetation. During this humid phase erosion decreased as less lithogenic material and more organic terrestrial material were deposited on the continental slope of northern Brazil. After 15 ka arid conditions returned. During the humid second phase of the H1 stadial, a rich variety of landscapes existed in Northeast Brazil and during the drier periods small pockets of forest could probably survive in favorable spots, which would have increased the resilience of the forest to climate change.

Negative effects of ocean acidification on calcification vary within the coccolithophore genus Calcidiscus

A large percentage of CO2 emitted into the atmosphere is absorbed by the oceans, causing chemical changes in surface waters known as ocean acidification (OA). Despite the high interest and increased pace of OA research to understand the effects of OA on marine organisms, many ecologically important organisms remain unstudied. Calcidiscus is a heavily calcified coccolithophore genus that is widespread and genetically and morphologically diverse. It contributes substantially to global calcium carbonate production, organic carbon production, oceanic carbon burial, and ocean-atmosphere CO2 exchange. Despite the importance of this genus, relatively little work has examined its responses to OA. We examined changes in growth, morphology, and carbon allocation in multiple strains of Calcidiscus leptoporus in response to ocean acidification. We also, for the first time, examined the OA response of Calcidiscus quadriperforatus, a larger and more heavily calcified Calcidiscus congener. All Calcidiscus coccolithophores responded negatively to OA with impaired coccolith morphology and a decreased ratio of particulate inorganic to organic carbon (PIC:POC). However, strains responded variably; C. quadriperforatus showed the most sensitivity, while the most lightly calcified strain of C. leptoporus showed little response to OA. Our findings suggest that calcium carbonate production relative to organic carbon production by Calcidiscus coccolithophores may decrease in future oceans and that Calcidiscus distributions may shift if more resilient strains and species become dominant in assemblages. This study demonstrates that variable responses to OA may be strain or species specific in a way that is closely linked to physiological traits, such as cellular calcite quota.

Lithology and pollen record of various sediment profiles of postglacial age from Bavaria, Germany

(Einleitung) Im süddeutschen Jungmoränengebiet wurden während der letzten 25 Jahre verschiedene vegetationsgeschichtliche Arbeiten durchgeführt, die der Untersuchung der Späteiszeit galten. Die wichtigsten von ihnen stammen von G. Lang (1952), A. Bertsch (1961), H. Müller (1962) und H. Schmeidl (1971). Ohne Zweifel müssen die dabei gewonnenen Ergebnisse in anderen Landschaften des nördlichen Alpenvorlandes überprüft und verschiedene Probleme weiterhin verfolgt werden, wie z. B. das der Definition und Umgrenzung der Bölling-Zeit und der Älteren Tundrenzeit s. str. und die Abhängigkeit der Vegetationsentwicklung von der Meereshöhe. Die vorliegende Studie ging auch auf die Notwendigkeit zurück, die spätglazialen Ablagerungen bei dem Tonwerk Kolbermoor nahe Rosenheim, einer der klassischen Stätten der Quartärforschung im nördlichen Alpenvorland, einer vegetationsgeschichtlichen Neubearbeitung zu unterziehen. Die Untersuchungen wurden auf benachbarte Seen, den Sims-See und den Hofsrätter See, ausgedehnt, da die Ergebnisse von Kolbermoor faziell beeinflußt schienen (Niedermoore) und an limnischem Material überprüft werden mußten.

Pollen records and lithology of profiles from Lobsigensee, Switzerland

Lobsigensee is a small kettle hole lake 15 km north-west of Bern on the Swiss Plateau, at an altitude of 514 m asl. Its surface is 2ha today, its maximum depth 2.7 m; it has no inlet and the overflow functions mainly during snow melting. The area was covered by Rhone ice during the Last Glaciation (map in Fig.2). Local geology, climate and vegetation are summarized in Figure 3A-C, the history of settlement in Figures 5-7. In order to reconstruct the vegetational and environmental history of the lake and its surroundings pollen analysis and other bio- and isotope stratigraphies were applied to twelve profiles cored across the basin with modified Livingstone corers (Fig.3 D). (1) The standard diagram: The central core LQ-90 is described as the standard pollen diagram (Chapter 3) with 10 local pollen assemblage zones of the Late-Glacial (local PAZ Ll to Ll0, from about 16'000(7) to 10'000 years BP) and 20 PAZ of the Holocene (local PAZ L11 to L30), see Figs. 8-10 and 20-24. Local PAZ L 1 to L3 are in the Late-Glacial clay and record the vegetational development after the ice retreat: L1 shows very low pollen concentration and high Pinus percentages due to long-distance transport and reworking; the latter mechanism is corroborated by the findings of thermophilous and pre-Quaternary taxa. Local PAZ L2 has a high di versi ty of non-arboreal pollen (NAP) and reflects the Late-Glacial steppe rich in heliophilous species. Local PAZ L3 is similar but additionally rich in Betula nana and Sal1x, thus reflecting a "shrub tundra". The PAZ L1 to L3 belong to the Oldest Dryas biozone. Local PAZ L4 to L 10 are found in the gyttja of the profundal or in the lake marl of the littoral and record the Late-Glacial forests. L4 is the shrub phase of reforestation with very high Junlperus and rapidly increasing Betula percentages. L5 is the PAZ with a first, L7 with a second dominance of tree-birches, separated by L6 showing a depression in the Betula curve. L4 to L7 can be assigned to the Balling biozone. Possible correlation of the Betula depression to the Older Dryas biozone is discussed. In local PAZ L8 Plnus immigrates and expands. L9 shows a facies difference in that Plnus dominates over Betula in littoral but not in profundal spectra. L8 and L9 belong to the Allerod biozone. In its youngest part the volcanic ash from Laach/Eifel is regularly found (11,000 BP). The local PAZ Ll0 corresponds to the Younger Dryas blozone. The merely slight increase of the NAP indicates that the pine forests of the lowland were not strongly affected by a cooler climate. In order to evaluate the significance of the littoral accumulation of coniferous pollen the littoral profile LQ-150 is compared to the profundal. Radiocarbon stratigraphies derived from different materials are presented in Figures 13 and 14 and in Tables 2 and 3. The hard-water errors in the gyttja samples and the carbonate samples are similar. The samples of terrestrial plant macrofossils are not affected by hard-water errors. Two plateaux of constant age appear in the age-depth relationship; their consequence for biostratigraphy as well as pollen concentration and influx diagrams are discussed. Radiocarbon ages of the Late-Glacial pollen zones are shown in Table 10. The Holocene vegetational history is recorded in the local PAZ L 11 to L30. After a Preboreal (PAZ L11) dominated by pine and birch the expansions of Corylus, Ulmus and Quercus are very rapid. Among these taxa Corylus dominates dur ing the Boreal (PAZ L 12 and L 1 3), whereas the components of the mixed oak forest dominate in the Older Atlantic (PAZ L14 to L16). In the Younger Atlantic (PAZ L 17 to L 19) Fagus and Alnus play an increasing, the mixed oak forest a decreasing role. During the period of local PAZ L19 Neolithic settlers lived on the shore of Lobsigensee. During the Subboreal (PAZ L20 and L21) and the Older Subatlantic (L22 to L25) strong fluctuations of Fagus and often antagonistic peaks of NAP, Alnus, Betula and Corylus can be interpreted as signs of human impact on vegetation. L23 is characterized not only by high values of NAP (especially apophytes and anthropochorous species) but also by the appearance of Juglans, Castanea and Secale which point to the Roman colonization of the area. For a certain period during the Younger Subatlantic (PAZ L26 to L30) the lake was used for retting hemp (Cannabis). Later the dominance of Quercus pollen indicates the importance of wood pastures. The youngest sediments reflect the wide-spread agricultural grass lands and the plantation of Pinus and Picea. Radiocarbon dates for the Holocene are given in Figure 23 and Table 4, the extrapolated ages of the Holocene pollen zones in Table 15. (2) The cross sections: Figures 25 and 26 give a summary of the litho- and palynostratigraphy of the two cross sections. Based on 11 Late-Glacial and 9 Holocene pollen diagrams (in addition to the standard ones), the consistency of the criteria for the definition of the pollen zones is examined in Tables 7 and 8 for the Late-Glacial and in Tables 11 to 14 for the Holocene. Sediment thicknesses across the basin for each pollen zone are presented in these tables as well as in Figures 43 to 45 for the Late-Glacial and in Figures 59 to 65 for the Holocene. Sediment focusing can explain differences between the gyttja cores of the profundal. Focusing is more than compensated for through "stretching" by carbonate precipitation on the littoral terrace. Pollen influx to the cross section are discussed (Chapters 4.1.5. and 4.2.3.). (3) The regional pollen zones: Based on some selected sites between Lake Geneva and Lake Constance regional pollen zones are proposed (Table 16, 17 and 19). (4) Paleoecology: Climatic change in the Late-Glacial can be inferred from Coleoptera, Trichoptera, Chironomidae and d18O of carbonates: a distinct warming is recorded around 12' 600 BP and around 10' 000 BP. The Younger Dryas biozone (10'700-10'000 BP) was the only cooling found in the Late-Glacial. The Betula depression often correlated wi th the Older Dryas biozone was possibl not colder but dryer than the previous period. During the Holocene the lowland site is not very sensitive to the minor climatic changes. Table 22 summarizes climatic and trophic changes before 8'000 BP as deduced from various biostratigraphies studied by a number of authors. Ostracods, Chironomids and fossil pigments indicate that anoxic conditions prevailed during the BoIling (possibly meromixis). Changes in the lake level are illustrated in Figure 74. A first lake-level lowering occurred in the early Holocene (10'000 to 9'000 BP), a second during the Atlantic (about 6'800 to 5'200 BP). The first "shrinking" of the lake volume resulted in a eutrophication recorded by laminations in the profundal and by pigments of Cyanophyceae. The second fall in water level corresponds to an increase of Nymphaeaceae. Human impact can be inferred in three ways: eutrophication of the lake (since the Neolithic), changes of terrestrial vegetation by deforestations (cyclicity of Fagus, see Figures 78 to 80), and enhanced erosion (increasing sedimentation rates by inwashed clay, particularly since the Roman Colonization, see Figures 49 and 81). Summary: This paper was planned as the final report on Lobsigensee. However, a number of issues are not answered but can only be asked more precisely, for example: (1) For the two periods with the highest rates of change, Le. the Bolling and the Preboreal biozones, pollen influx may reflect vegetation dynamics. Detailed investigations of these periods in annually laminated sediments are planned. (2) Biostratigraphies other than palynostratigraphy are needed to estimate the degree of linkage or independence in the development of terrestrial and lacustrine ecosystems. Often our sampling intervals were not identical, thus influencing our temporal resolution. (3) 6180- and 14C-stratigraPhies with high resolution will elucidate the leads and lags of these dynamic periods. Plateaux of constant age in the age-depth relationship have a strong bearing on both biological and geophysical understanding of Late-Glacial and early Holocene developments. (4) Numerical methods applied to the pollen diagrams of the cross section will help to quantify the significance of similari ties and dissimilarities across a single basin (with Prof. Birks). (5) Numerical methods applied to different sites on the Swiss Plateau and on the transect across the Alps will be helpful in evaluating the influence of different environmental factors (with Prof. Birks). (6) A new map 1: 1000 with 50cm-contour lines prov ided by Prof. Zurbuchen will be combined with a grid of cores sampling the transition from lake marl to peat enabling us to calculate paleo-volumes of the lake. This is interesting for the two "shrinking periods" (in Fig. 74A numbers 2-6 and 7-10), both accompanied by eutrophication. The pal eo-volume during the Neoli thic set tlement of the Cortaillod culture linked wi th an est l.mate of trophic change derived from diatoms (Prof. Smol in prep.) could possibly give an indication of the size of the human population of this period. (7) For the period with the antagonism between Fagus peaks and ABC-peaks close collaboration between palynologists, geochemists and archeologists should enable us to determine the influence of prehistoric and historic people on vegetation (collaboration with Prof. Stockli and Prof. Herzig). (8) The core LL-75 taken with a "cold letter box" will be analysed for major and trace elements by Dr. Sturm for 210pb and 137Cs by Prof.von Gunten and for pollen. We will see if our local PAZ L30 really corresponds to the surface sediment and if the small seepage lake reflects modern pollution.

Arctic Siberia: Modern pollen dataset and pollen spectra changes over the last 7,000 years from a lacustrine sediment core (southern Taymyr region)

Palaeoecological investigations in the larch forest-tundra ecotone in northern Siberia have the potential to reveal Holocene environmental variations, which likely have consequences for global climate change because of the strong high-latitude feedback mechanisms. A sediment core, collected from a small lake (radius ~100 m), was used to reconstruct the development of the lake and its catchment as well as vegetation and summer temperatures over the last 7100 calibrated years. A multi-proxy approach was taken including pollen and sedimentological analyses. Our data indicate a gradual replacement of open larch forests by tundra with scattered single trees as found today in the vicinity of the lake. An overall trend of cooling summer temperature from a ~2 °C warmer-than-present mid-Holocene summer temperatures until the establishment of modern conditions around 3000 years ago is reconstructed based on a regional pollen-climate transfer function. The inference of regional vegetation changes was compared to local changes in the lake's catchment. An initial small water depression occurred from 7100 to 6500 cal years BP. Afterwards, a small lake formed and deepened, probably due to thermokarst processes. Although the general trends of local and regional environmental change match, the lake catchment changes show higher variability. Furthermore, changes in the lake catchment slightly precede those in the regional vegetation. Both proxies highlight that marked environmental changes occurred in the Siberian forest-tundra ecotone over the course of the Holocene.

Pollen record from Muggesfelder See and Plussee in Germany

A basaltic tephra layer consisting of brownish-olive glass shards. and about 0.2 mm thick. was found in cores from four lakes in northwest Germany. According to pollen analysis it was deposited during the early Boreal period (corresponding to about 8700 BP). The petrographic properties. the geochemical composition and the age agree with those of the Saksunarvatn tephra. which was first found on the Faroe Islands. The position of the tephra layer in the pollen stratigraphy and in the absolute time-scale is discussed. Procedures for locating the tephra in other cores are suggested.

Palynology of ODP Site 108-658

Ocean Drilling Program Site 658 at 21°N off northwest Africa has a high sedimentation rate and a high concentration of pollen grains and is thus very suitable for detailed pollen analysis. The time scale for the upper 100 m (the last 670 k.y.) of Site 658 is based on biostratigraphic data and isotope stratigraphy. The pollen record has been divided into 34 zones. These are classified into 7 zone types covering a range from very arid to rather humid conditions. The sequence shows a long-term climatic decline: strong glacial stages were found only after 480 k.y. and strong interglacial stages only before 280 k.y. The Site 658 record correlates well with a terrestrial sequence from northern Greece, although both records differ in their response to global climatic change. Spectral analysis shows a 100- and a 42-k.y. period in the curves of pollen brought in by the northwest trade winds and only a 42-k.y. period in the curves of pollen mostly transported by the African Easterly Jet. A 31-k.y. period is found in the curves for Ephedra and Chenopodiaceae-Amaranthaceae. In addition, Ephedra shows a 54-k.y. period.

Biogeochemistry measurements in a mesocosm experiment in the Bay of Hopavagen, Norway, in August 2012

The goal of this work has been to examine the influence of upper ocean food web structure and functioning on both the natural and artificially enhanced sequestration of carbon within the ocean. Data obtained in the mesocosm experiment run in the Bay of Hopavågen in August 2012 are used to assess the extent to which organic matter produced within four different food webs is retained in the upper ocean food web versus remineralized back to carbon dioxide and inorganic nutrients (ammonium, dissolved silicon, phosphate) versus exported from the system in the form of rapidly sinking particles. The experiment was carried out in a set of 12 mesocosms covering, in triplicate, 2 different phytoplankton communities (diatom versus non-diatom) exposed to 2 different zooplankton communities (-copepod and +copepod). These starting conditions were established by first filling the bags, roughly simultaneously, with seawater from the Bay of Hopavågen. Mesozooplankton were then removed to the most complete extent possible immediately removed from half of the mesocosms through repeated vertical hauls of a plankton net (200 µm mesh). Nitrate and phosphate was added to half mesocosms daily to promote the growth of non-siliceous phytoplankton (e.g. dinoflagellates or coccolithophores). To the other half of the mesocosms, nitrate, phosphate, and silicate were added to promote the growth of diatoms. Material was allowed to settle and the two distinct phytoplankton populations were allowed to develop for 4 days, after which copepods collected from the Bay of Hopavågen were added back to the half of the N+P mesocosms and to the half of the N+P+Si mesocosms from which mesozooplankton had not been removed at the beginning. This yielded a set of four initial starting conditions (N+P-copepods, N+P+copepods, N+P+Si-copepods, and N+P+Si+copepods). In the primary mesocosms, samples for a set of core parameters were taken every time the mesocosms were sampled. Samples for particulates (PIC, BSi, POC, PON) were collected on GF/F or 0.4 µm polycarbonate.

Age determination and pollen profile of sediment core Voulkaria 1966, Greece

Lake Voulkaria is situated in northwestern Greece in the Prefecture of Etoloakarnania, 6 km SW of the city of Vonitsa and 10 km east of the northern tip of the island of Levkás (Leukás, Lefkada). The lake is separated from the Ionian Sea on the West by a narrow limestone ridge ca 10 m high and has a size of 940 ha. An almost continuous fringe of Phragmites surrounds the open water. This reed bank is up to 500 m wide along the southern shore of the lake. Water depth is low, predominantly less than 2 m. In the south-eastern part of the lake a maximum depth of 3.1 m was measured in September 1997.

Fossil pollen record of composite sediment core TMD from Tso Moriri, analysis of modern surface pollen samples, mean annual precipitation reconstruction, and digitisation and recalibration of different discussed palaeoclimate proxy records

This paper presents a new fossil pollen record from Tso Moriri (32°54'N, 78°19'E, 4512 m a.s.l.) and seeks to reconstruct changes in mean annual precipitation (MAP) during the last 12,000 years. This high-alpine lake occupies an area of 140 km**2 in a glacial-tectonic valley in the northwestern Himalaya. The region has a cold climate, with a MAP <300 mm, and open vegetation. The hydrology is controlled by the Indian Summer Monsoon (ISM), but winter westerly-associated precipitation also affects the regional water balance. Results indicate that precipitation levels varied significantly during the Holocene. After a rapid increase in MAP, a phase of maximum humidity was reached between ca. 11 to 9.6 cal ka BP, followed by a gradual decline in MAP. This trend parallels the reduction in the Northern Hemisphere summer insolation. Comparison of different palaeoclimate proxy records reveal evidence for a stronger Holocene decrease in precipitation in the northern versus the southern parts of the ISM domain. The long-term trend of ISM weakening is overlaid with several short periods of greater dryness, which are broadly synchronous with the North Atlantic cold spells, suggesting reduced amounts of westerly-associated winter precipitation. Compared to the mid and late Holocene, it appears that westerlies had a greater influence on the western parts of the ISM domain during the early Holocene. During this period, the westerly-associated summer precipitation belt was positioned at Mediterranean latitudes and amplified the ISM-derived precipitation. The Tso Moriri pollen record and moisture reconstructions also suggest that changes in climatic conditions affected the ancient Harappan Civilisation, which flourished in the greater Indus Valley from approximately 5.2 to 3 cal ka BP. The prolonged Holocene trend towards aridity, punctuated by an interval of increased dryness (between ca. 4.5 to 4.3 cal ka BP), may have pushed the Mature Harappan urban settlements (between ca. 4.5 to 3.9 cal ka BP) to develop more efficient agricultural practices to deal with the increasingly acute water shortages. The amplified aridity associated with North Atlantic cooling between ca. 4 to 3.6 and around 3.2 cal ka BP further hindered local agriculture, possibly causing the deurbanisation that occurred from ca. 3.9 cal ka BP and eventual collapse of the Harappan Civilisation between ca. 3.5 to 3 cal ka BP.

Biogenic flux and primary production of sediment traps in the Atlantic and the Southern Ocean

Fluxes of organic carbon normalised to a depth of 1000 m from 18 sites in the Atlantic and the Southern Ocean are presented, comprising nine biogeochemical provinces as defined by Longhurst et al. (1995. Journal of Plankton Research 17, 1245-1271). For comparison with primary production, we used a recent compilation of primary production values derived from CZCS data (Antoine et al., 1996. Global Biogeochemical Cycles 10, 57-69). In most cases, the seasonal patterns stood reasonably well in accordance with the carbon fluxes. Particularly, organic carbon flux records from two coastal sites off northwest and southwest Africa displayed a more distinct correlation to the primary production in sectors (1 x 1°) which are situated closer to the coastal environments. This was primarily caused by large upwelling filaments streaming far offshore, resulting in a cross-shelf carbon transport. With respect to primary production, organic carbon export to a water depth of 1000 m, and the fraction of primary production exported to a depth of 1000 m (export fraction=EF1000), we were able to distinguish between: (1) the coastal environments with highest values (EF1000=1.75-2.0%), (2) the eastern equatorial upwelling area with moderately high values (EF1000=0.8-1.1%), (3) and the subtropical oligotrophic gyres that yielded lowest values (EF1000=0.6%). Carbon export in the Southern Ocean was low to moderate, and the EF1000 value seems to be quite low in general. Annual organic carbon fluxes were proportional to primary production, and the export fraction EF1000 increased with primary production up to 350 gCm**-2 yr**-1. Latitudinal variations in primary production were reflected in the carbon flux pattern. A high temporal variability of primary production rates and a pronounced seasonality of carbon export were observed in the polar environments, in particular in coastal domains, although primary production (according to Antoine et al., 1996. Global Biogeochemical Cycles 10, 57-69), carbon fluxes, and the export fraction remained at low.

Downward particle fluxes at the oligotrophic and mesotrophic site of the EUMELI program

A two year record of downward particle flux was obtained with moored sediment traps at several depths of the water column in two regions characterized by different primary production levels (mesotrophic and oligotrophic) of the eastern subtropical North Atlantic Ocean in the framework of the EUMELI program. Settling particles were collected with multisample conical sediment-traps moored at 1000 and 2500 depths in the water column. Time-series samples were obtained between February 1991 and November 1992. During this time, sampling intervals varied from 8 to 10 d and were synchronized at all depths and also between the oligotrophic and mesotrophic moorings. Sediment-trap sampling procedures were consistent with JGOF and described elsewhere. The data shown here are mass, particulate organic carbon (POC), particulate inorganic carbon (PIC), coccolithophore, opal, and lithogenic downward fluxes obtained during the entire sediment-trap deployments at both sites.

Palynological record of Lake Nero in Russia

This collection prepared to IX Congress of INQUA containes 25 articles concerning general and regional problems of Pleistocene. The chronological scale of the Late Pliocene and Pleistocene, climatical cycles and methods of the absolute dating are considered. Some data obtained by means ef paleomagnetic, thermoluminescence and radiocarbon methods at several point sections (Likhvin, Rostov-Jarosiavsky, Priasovje, Ob-garm, Chagan, Pryobskoje Plateau, Lower Volga) are given.

Pollen and spore deposition in different areas of the Lena Delta

Studies of the annual pollen and spore deposition in different areas of the Lena Delta were undertaken for the first time in the Asian sector of the Arctic during the Russian-German ''LENA 98'' and ''LENA 99'' expeditions in the framework of the International ''Laptev Sea System-2000'' Project. To achieve this objective, three spore-pollen traps were set up along the meridional delta profile in accordance with the European Pollen Monitoring Programme for the period July 1998 to August 1999. A comparison between the results of spore-pollen analysis of the contents of traps and the surrounding vegetation was performed. The results confirmed the current spore-pollen spectra are comprised both of pollen and spores of the local plants and of long-distance pollen and spores. The dependence of the long-distance pollen deposition on the character of the wind regime of the region was established. The prevailing southerly and southeasterly wind direction determines the main pollen influx of tree species from the areas of their growth south of the delta. The features of the morphological structure and fossilization of pollen and the features of the productive capability and plant growing conditions are of large significance in the pollen transfer and deposition.