Herbivory in Spiders: The Importance of Pollen for Orb-Weavers

Orb-weaving spiders (Araneidae) are commonly regarded as generalist insect predators but resources provided by plants such as pollen may be an important dietary supplementation. Their webs snare insect prey, but can also trap aerial plankton like pollen and fungal spores. When recycling their orb webs, the spiders may therefore also feed on adhering pollen grains or fungal spores via extraoral digestion. In this study we measured stable isotope ratios in the bodies of two araneid species (Aculepeira ceropegia and Araneus diadematus), their potential prey and pollen to determine the relative contribution of pollen to their diet. We found that about 25% of juvenile orb-weaving spiders’ diet consisted of pollen, the other 75% of flying insects, mainly small dipterans and hymenopterans. The pollen grains in our study were too large to be taken up accidentally by the spiders and had first to be digested extraorally by enzymes in an active act of consumption. Therefore, pollen can be seen as a substantial component of the spiders’ diet. This finding suggests that these spiders need to be classified as omnivores rather than pure carnivores.

Pinus nigra (European black pine) as the dominant species of the last glacial pinewoods in south-western to central Iberia: a morphological study of modern and fossil pollen

Aim Our aim was to discriminate different species of Pinus via pollen analysis in order to assess the responses of particular pine species to orbital and millennial-scale climate changes, particularly during the last glacial period. Location Modern pollen grains were collected from current pine populations along transects from the Pyrenees to southern Iberia and the Balearic Islands. Fossil pine pollen was recovered from the south-western Iberian margin core MD95-2042. Methods We measured a set of morphological traits of modern pollen from the Iberian pine species Pinus nigra, P. sylvestris, P. halepensis, P. pinea and P. pinaster and of fossil pine pollen from selected samples of the last glacial period and the early to mid-Holocene. Classification and regression tree (CART) analysis was used to establish a model from the modern dataset that discriminates pollen from the different pine species and allows identification of fossil pine pollen at the species level. Results The CART model was effective in separating pollen of P. nigra and P. sylvestris from that of the Mediterranean pine group (P. halepensis, P. pinea and P. pinaster). The pollen of Pinus nigra diverged from that of P. sylvestris by having a more flattened corpus. Predictions using this model suggested that fossil pine pollen is mainly from P. nigra in all the samples analysed. Pinus sylvestris was more abundant in samples from Greenland stadials than Heinrich stadials, whereas Mediterranean pines increased in samples from Greenland interstadials and during the early to mid-Holocene. Main conclusions Morphological parameters can be successfully used to increase the taxonomic resolution of fossil pine pollen at the species level for the highland pines (P. nigra and P. sylvestris) and at the group of species level for the Mediterranean pines. Our study indicates that P. nigra was the dominant component of the last glacial south-western/central Iberian pinewoods, although the species composition of these woodlands varied in response to abrupt climate changes.

Survey of pollen and spores in DSDP Holes 96-618 and 619

A preliminary palynological survey of 118 samples from the Mississippi Fan (Sites 615, 616, and 620) and from 2 intraslope basins (Sites 619 and 618) shows pollen in all samples. Reworked pollen is generally abundant, forming over 50% of the sum of pollen and reworked pollen. Concentration of nonreworked pollen is usually low (on the order of tens to hundreds of pollen grains per cubic centimeter wet sediment). Conifers, primarily Pinus, Picea, and Tsuga, dominate Pleistocene marine pollen spectra; significant percentages of Quercus are present in Holocene sediments and in sediments deposited during oxygen-isotope Stage 5.

Spores, pollen and dinoflagellates in Cretaceous sediments of ODP Hole 129-802A

Lower Cretaceous and Jurassic sediments from Ocean Drilling Program Leg 129 (Sites 800, 801, and 802) and Deep Sea Drilling Project Sites 167, 195, 196, and 463 were analyzed for palynomorphs. In contrast to Atlantic occurrences, all Cretaceous pelagic sediments at these sites in the Pacific are barren of preserved palynomorphs. This absence of palynomorphs appears to be independent of facies, sedimentation rate, paleodepth, and paleolatitude. Except for one sample, the dinocyst-bearing sediments also contain spores and pollen grains. The only palynomorphs observed were in redeposited material having sources near former emergent seamounts. Among the dinoflagellate cysts at Site 802, Dingodinium cerviculum, Odontochitina operculata, Canninginopsis colliveri, and Oligosphaeridium complex are the most important species. Based on the presence of these species and their known biostratigraphic ranges, this basal interval of Site 802 is considered to be Aptian/earliest Albian in age. The lack of dinocysts within the Pacific pelagic sediments may be the result of ubiquitous oxygenated bottom waters throughout the Cretaceous or may indicate that open-marine dinoflagellate populations in this ocean did not produce cysts.

Pollen analysis of a peat profile from the Rieseberger Moor, Lower Saxony, Germany

The Rieseberger Moor is a fen, 145 hectares in size, situated about 20 km east of Brunswick (Braunschweig), Lower Saxony, Germany. Peat was dug in the fen - with changing intensity - since the mid-18th century until around AD 1955. According to Schneekloth & Schneider (1971) the remaining peat (fen and wood peat) is predominantly 1.5 to 2 m thick (maximum 2.7 m). Part of the fen - now a nature reserve (NSG BR 005) - is wooded (Betula, Salix, Alnus). For more information on the Rieseberger Moor see http://de.wikipedia.org/wiki/Rieseberger_Moor. Willi Selle was the first to publish pollen diagrams from this site (Selle 1935, profiles Rieseberger Torfmoor I and II). This report deals with a 2.2 m long profile from the wooded south-eastern part of the fen consisting of strongly decomposed fen peat taken A.D. 1965 and studied by pollen analysis in the same year. The peat below 1.45 m contained silt and clay, samples 1.48 and 1.58 m even fine sand. These samples had to be treated with HF (hydrofluoric acid) in addition to the treatment with hot caustic potash solution. The coring ended in sandy material. The new pollen data reflect the early part of the known postglacial development of the vegetation of this area: the change from a birch dominated forest to a pine forest and the later spreading of Corylus and of the thermophilous deciduous tree genera Quercus, Ulmus, Tilia and Fraxinus followed by the expansion of Alnus. The new data are in agreement with Selle's results, except for Alnus, which in Selle's pollen diagram II shows high values (up to 42% of the arboreal pollen sum) even in samples deposited before Corylus and Quercus started to spread. On contrary the new pollen diagram shows that alder pollen - although present in all samples - is frequent in the three youngest pollen spectra only. A period with dominating Alnus as seen in the uppermost part of Selle's pollen diagrams is missing. The latter is most likely the result of peat cutting at the later coring site, whereas the early, unusually high alder values of Selle's pollen study are probably caused by contamination of the pollen samples with younger peat. Selle took peat samples usually with a "Torfbohrer" (= Hiller sampler). This side-filling type of sampler with an inner chamber and an outer loose jacket offers - if not handled with appropriate care - ample opportunities to contaminate older peat with carried off younger material. Pollen grains of Fagus (2 % of the arboreal pollen sum) were found in two samples only, namely in the uppermost samples of the new profile (0.18 m) and of Selle's profile I (0.25 m). If this pollen is autochthonous, with other words: if this surface-near peat was not disturbed by human activities, the Fagus pollen indicates an Early Subboreal age of this part of the profile. The accumulation of the Rieseberg peat started during the Preboreal. Increased values of Corylus, Quercus and Ulmus indicate that sample 0.78 m of the new profile is the oldest Boreal sample. The high Alnus values prove the Atlantic age of the younger peat. Whether Early Subboreal peat exists at the site is questionable, but evidently none of the three profiles reaches to Late Subboreal time, when Fagus spread in the region. Did peat-growth end during the Subboreal? Did younger peat exist, but got lost by peat cutting or has younger peat simply not yet been found in the Rieseberg fen? These questions cannot be answered with this study. The temporary decline of the curve of Pinus for the benefit of Betula during the Preboreal, unusual for this period, is contemporaneous with the deposition of sand (Rieseberger Moor II, 1.33 - 1,41 m; samples 1.48 and 1.58 m of the new profile) and must be considered a local phenomenon. Literature: Schneekloth, Heinrich & Schneider, Siegfried (1971). Die Moore in Niedersachsen. 2. Teil. Bereich des Blattes Braunschweig der Geologischen Karte der Bundesrepublik Deutschland (1:200000). - Schriften der wirtschaftswissenschaftlichen Gesellschaft zum Studium Niedersachsens e.V. Reihe A I., Band 96, Heft 2, 83 Seiten, Göttingen. Selle, Willi (1935) Das Torfmoor bei Rieseberg. - Jahresbericht des Vereins für Naturwissenschaft zu Braunschweig, 23, 46-58, Braunschweig.

Pollen analysis of soil samples from the A.D. 79 level. Boscoreale, Oplontis, and Pompeii

Fifty samples of Roman time soil preserved under the thick ash layer of the A.D.79 eruption of Mt Vesuvius were studied by pollen analysis: 33 samples from a former vineyard surrounding a Villa Rustica at Boscoreale (excavation site 40 x 50 m), 13 samples taken along the 60 m long swimming pool in the sculpture garden of the Villa of Poppaea at Oplontis, and four samples from the formal garden (12.4 x 17.5 m) of the House of the Gold Bracelet in Pompeii. To avoid contamination with modern pollen all samples were taken immediately after uncovering a new portion of the A.D. 79 soil. For comparison also samples of modern Italian soils were studied. Using standard methods for pollen preparation the pollen content of 15 of the archaeological samples proved to be too little to reach a pollen sum of more than 100 grains. The pollen spectra of these samples are not shown in the pollen tables. (Flotation with a sodium tungstate solution, Na2WO4, D = 2.05, following treatment with HCl and NaOH would probably have given a somewhat better result. This method was, however, not available as too expensive at that time.) Although the archaeological samples were taken a few meters apart their pollen values differ very much from one sample to the other. E.g., at Boscoreale (SW quarter). the pollen values of Pinus range from 1.5 to 54.5% resp. from 1 to 244 pine pollen grains per 1 gram of soil, the extremes even found under pine trees. Vitis pollen was present in 7 of the 11 vineyard samples from Boscoreale (NE quarter) only. Although a maximum of 21.7% is reached, the values of Vitis are mostly below 1.5%. Even the values of common weeds differ very much, not only at Boscoreale, but also at the other two sites. The pollen concentration values show similar variations: 3 to 3053 grains and spores were found in 1 g of soil. The mean value (290) is much less than the number of pollen grains, which would fall on 1 cm2 of soil surface during one year. In contrast, the pollen and spore concentrations of the recent soil samples, treated in exactly the same manner, range from 9313 to almost 80000 grains per 1 g of soil. Evidently most of the Roman time pollen has disappeared since its deposition, the reasons not being clear. Not even species which are known to have been cultivated in the garden of Oplontis, like Citrus and Nerium, plant species with easily distinguishable pollen grains, could be traced by pollen analysis. The loss of most of the pollen grains originally contained in the soil prohibits any detailed interpretation of the Pompeian pollen data. The pollen counts merely name plant species which grew in the region, but not necessarily on the excavated plots.

Pollen profile from sediment core Höllerer See

(expanded by Eberhard Grüger, Göttingen) The site "Höllerer See" is a lake in the northern foreland of the Alps, about 30 km north of the city of Salzburg/Austria, situated in the south-western part of Oberösterreich/Austria. A 2 m long piston core from this locality, consisting entirely of calcareous gyttja, was studied by pollen analysis. The three lowermost samples (1.98, 1.95 and 1.92 m) were deposited during the Preboreal when Pinus and Betula were still the dominating forest trees. High pollen values of thermophilous woody species (mainly Corylus and Quercus, but also Ulmus, Tilia, Fraxinus) prove the Boreal age of the next younger sample (1.91 m). The following two pollen spectra attest that Alnus (1.89 m) and - later (1.88 m) - Fagus had become important members of the local (Alnus) and the regional (Fagus) vegetation. From this level up to the top of the profile these two tree taxa contribute - together with Betula - always 50 to 80 % to the arboreal pollen sum. The upper 1.89 m of sediment of the Höllerer See core evidently date from the Subboreal and the Subatlantic. As Preboreal sediment was stated at the base of the profile it must be concluded that most of the Boreal and the Atlantic is - for whatever reason - not represented by sediment in this core. As no radiocarbon dates are available age estimates of the distinguished pollen zones can be achieved only by correlating major changes of the former vegetation with historical events which probably influenced the then contemporary vegetation. The pollen grains of the Triticum and Hordeum type found in samples of zone 2.1 might indicate the growing of cereals in the region during the Late Bronze Age. The first pollen grains of Secale date from the boundary Hallstatt/Latène Age (zone 2.2). The cereal curves become continuous in Bavarian times (Bajuwarenzeit, Middle Ages, zone 3.3). The Plantago laceolata curve, continuous since 1.7 m depth (zone 2.1), points to animal breeding since the Early Subatlantic (Hallstattzeit). This curve reaches its absolute maximum in Roman time (zone 3.1). Roman time forest clearance caused a drastic decrease of tree pollen curves (start of zone 3.1). Values of anthropogenic indicators as high as in zone 3.1 are found again - after a distinct decrease in zone 3.2 - not till the Bavarians settled in the region (6th century). Maximal Fagus values and the simultaneous total lack of anthropogenic indicators mark the Migration Period (zone 3.2). The Younger Subatlantic (zone 4) is characterized by a decrease of deciduous forests due to medieval forest clearance. At the same time the conifers Pinus and Picea gained in importance. The lake was probably used for retting hemp in Medieval times. The distinction of the pollen grains of Cannabis and Humulus might not be certain in all cases. It is known that hemp as well as hop was cultivated in the study area. Markers were added to the samples at the beginning of pollen preparation (13500 Lycopodium spores, sample volume 0.5 cm**3) and counted together with the pollen grains. Therefore pollen concentrations can be calculated: Concentration = C * F / V (with C = number of grains of a particular pollen type, V = volume of the untreated pollen sample, F = marker added/marker counted). F ranges from 39 to 1688. Factors that large are not suited to produce reliably interpretable pollen concentrations. Consequently no use was made of the pollen concentrations in this thesis, although a concentration diagram is added.

Developmental changes in DNA methylation of the two tobacco pollen nuclei during maturation

Changes in DNA methylation during tobacco pollen development have been studied by confocal fluorescence microscopy using a monoclonal anti-5-methylcytosine (anti-m5C) antibody and a polyclonal anti-histone H1 (anti-histone) antibody as an internal standard. The specificity of the anti-m5C antibody was demonstrated by a titration series against both single-stranded DNA and double-stranded DNA substrates in either the methylated or unmethylated forms. The antibody was found to show similar kinetics against both double- and single-stranded DNA, and the fluorescence was proportional to the amount of DNA used. No signal was observed with unmethylated substrates. The extent of methylation of the two pollen nuclei remained approximately constant after the mitotic division that gave rise to the vegetative and generative nuclei. However, during the subsequent development of the pollen, the staining of the generative nucleus decreased until it reached a normalized value of \documentclass[12pt]{minimal} \usepackage{wasysym} \usepackage{amsmath} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} \begin{equation*}\frac{1}{5}\end{equation*}\end{document} of that of the vegetative nucleus. The use of a confocal microscope makes these data independent of possible focusing artefacts. The anti-histone antibody was used as a control to show that, while the antibody staining directed against 5-methylcytosine changed dramatically during pollen maturation, the histone signal did not. We observed the existence of structural dimorphism amongst tobacco pollen grains, the majority having three pollen apertures and the rest with four. However, the methylation changes observed occurred to the same extent in both subclasses.

Directional Guidance of Nicotiana alata Pollen Tubes in Vitro and on the Stigma1

Pollen tubes navigate the route from stigma to ovule with great accuracy, but the cues that guide them along this route are not known. We reproduced the environment on the stigma of Nicotiana alata by immersing pollen in stigma exudate or oil close to an interface with an aqueous medium. The growth of pollen in this culture system mimicked growth on stigmas: pollen grains hydrated and germinated, and pollen tubes grew toward the aqueous medium. The rate-limiting step in pollen germination was the movement of water through the surrounding exudate or oil. By elimination of other potential guidance cues, we conclude that the directional supply of water probably determined the axis of polarity of pollen tubes and resulted in growth toward the interface. We propose that a gradient of water in exudate is a guidance cue for pollen tubes on the stigma and that the composition of the exudate must be such that it is permeable enough for pollen hydration to occur but not so permeable that the supply of water becomes nondirectional. Pollen tube penetration of the stigma may be the most frequently occurring hydrotropic response of higher plants.

The late Pliocene pollen record of ODP Site 108-658 off Cape Blanc, Atlantic Ocean

A 200 m long marine pollen record from ODP Site 658 (21°N, 19°W) reveals cyclic fluctuations in vegetation and continental climate in northwestern Africa from 3.7 to 1.7 Ma. These cycles parallel oxygen isotope stages. Prior to 3.5 Ma, the distribution of tropical forests and mangrove swamps reached Cape Blanc, 5°N of the present distribution. Between 3.5 and 2.6 Ma, forests occurred at this latitude during irregular intervals and nearly disappeared afterwards. Likewise, a Saharan paleoriver flowed continuously until isotope Stage 134 (3.35 Ma). When river discharge ceased, wind transport of pollen grains prevailed over fluvial transport. Pollen indicators of trade winds gradually increased between 3.3 and 2.5 Ma. A strong aridification of the climate of northwestern Africa occurred during isotope Stage 130 (3.26 Ma). Afterwards, humid conditions reestablised followed by another aridification around 2.7 Ma. Repetitive latitudinal shifts of vegetation zones ranging from wooded savanna to desert flora dominated for the first time between between 2.6 and 2.4 Ma as a response to the glacial stages 104, 100 and 98. Although climatic conditions, recorded in the Pliocene, were not as dry as those of the middle and Late Pleistocene, latitudinal vegetation shifts near the end of the Pliocene resembled those of the interglacial-glacial cycles of the Brunhes chron.

Early Cretaceous spores and pollen of ODP Leg 103 holes

About 80 species of spores and pollen grains were recorded during detailed palynological investigations of selected Lower Cretaceous sections from Holes 638B and 638C and the bottom of Hole 641C. Most of them are long-ranging taxa with worldwide distribution. However, on the Iberian margin and in the southern European basins, Trilobosporites canadensis, Trilobosporites bernissartensis, Parvisaccites amplus, Foveosporites subtriangularis, and Ephedripites multicostatus seem to be index species of the Valanginian to late Aptian interval. Clavatipollenites was not recovered in the Barremian marginal marine sediments.