Bee-pollination inHibbertia fasciculata (Dilleniaceae)

In direct contrast to mostHibbertia spp., the flowers ofH. fasciculata R. Br. ex D. C. bear only a single whorl of stamens and these stamens are arranged separately (not in typical bundles). The short filaments are appressed to the three carpels so that the inflated, porose and introrsive anthers form a centralized cluster obscuring the three ovaries. The three slender styles emerge at right angles from between the filaments. These styles curve upward and the stigmas form the three points of a triangle; each stigma is approximately one millimeter outside the centralized cluster of anthers. The flowers are nectarless and bear a bright yellow corolla. A pungent and unpleasant fragrance appears to be concentrated within the pollenkitt. When native bees attempt to forage for the pollen, within the cluster of anthers, the ventrally deposited loads of pollen, on the bees' abdomens, contact the outer triangle of stigmas. The major pollinators ofH. fasciculata are female bees in the polylectic genera,Lasioglossum (subgenusChilalictus, Halictidae) andLeioproctus (Colletidae). These bees carry an average of more than two pollen taxa when they are caught foraging onH. fasciculata. 78% of the 47 bees, captured onH. fasciculata carried the pollen from at least one sympatric taxon bearing nectariferous flowers (e.g., genera in theMyrtaceae, Compositae, andEpacridaceae). The pollination biology ofH. fasciculata is assessed in relation to the known radiation of bee-pollinated flowers in the genusHibbertia, and within theDilleniaceae s. l.     

Auxin levels,antiauxin(s) and androgenic plantlet formation in isolated pollen cultures ofNicotiana tabacum

Summary Anthers ofNicotiana tabacum var. Badischer Burley contain endogenous auxins, one of these was identified as indoleacetic acid. At the developmental stage shortly after the first pollen mitosis the anthers contain equivalents of 0.1 mg IAA per kg fresh weight. This endogenous auxin level is maintained during the eight-day preculture of the anthers prior to isolated pollen culture. However, in anthers of short-day plants, which are characterized by a high proportion of embryogenic pollen at the end of preculture (Heberle-Bors andReinert 1979), an increase of the auxin level till the fourth day of culture is detectable.Preculture of anthers in the presence of an inhibitor of auxin synthesis (7-azaindole) and an antiauxin (-(o-chlorophenoxy)-isobutyric acid) results in enhanced plantlet yield by pollen cultures. The significance of these observations for androgenesis is discussed.     

The pollination biology ofHibbertia stricta (Dilleniaceae)

The exines of pollen grains ofHibbertia stricta (DC.)R. Br. exF. Muell. (Sect.Pleurandra) wear an oily, yellow pollen coat that stains positively for lipids. The pollen is collected by asocial bees, exclusively. The most common floral foragers are members of the genusLasioglossum (subgenusChilalictus;Halictidae) and they harvest pollen via thoracic vibration. As these bees cling to the inflated anthers their pollen smeared bodies come in contact with either of the two wet, nonpapillate stigmas. The stigmas respond positively to cytochemical tests for the presence of esterase immediately following expansion of the corolla, indicating the effective pollination period. The foraging patterns of the bees are narrowly to broadly polylectic. AsH. stricta flowers are nectarless, it is not surprising that bees bearing mixed pollen loads always carry the pollen of at least one nectariferous, coblooming plant. The pollination biology ofH. stricta is compared with otherHibbertia spp. and with pollen flowers in general.     

Floral mimesis inThelymitra nuda (Orchidaceae)

Insect pollination occurred inThelymitra nuda R. Br. on sunny days when the ambient temperature exceeded 20 °C. The flower buds on a raceme opened subsynchronously displaying the brightly-colored, actinomorphic perianth and exposed the contrastingly-colored, scented and ornamented column. InT. nuda the staminodes and the filament of the fertile stamen are fused to one another producing an inflated hood over the anther. This staminodal hood is terminated in two non-ornamental, but brightly-colored, central lobes and two terete lateral lobes bearing approximately 400 white trichomes. Each trichome bears a double chain of 30–40 spherical, rugulose cells. Female bees, in the genusLasioglossum (Halictidae), were observed to land directly on the hood and curled their bodies around the four lobed tip of the staminodal complex. The bees attempted to forage on the lobes as if they were collecting pollen from fertile, poricidal stamens. These bees applied thoracic vibration to the yellowish central lobes and actively scraped the trichome clusters (Pseudopollen) with their forelegs. Bees carried the pollinaria ofT. nuda dorsally on their abdomens. Abdominal contact with the rostellum appeared to occur when the female bee disengaged herself from the staminodal hood. Observations made of bees on co-blooming flowers, and analyses of pollen loads collected by bees suggested that the orchid flower mimics the guild of blue-purple flowers that lack floral nectar but offer pollen in poricidal anthers. The models ofT. nuda are co-bloomingLiliaceae in the generaDichopogon andThysanotus. However, nectarless, buzz-pollinated flowers are also extensively distributed over the orchid’s range. More than 30% of the flowers in theT. nuda population had their pollinaria removed, suggesting a high capacity for cross-pollination in an orchid genus usually considered to be self-pollinated via mechanical autogamy. This study confirmed previous predictions that column modifications represented a trend towards pseudanthery.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

Heterostyly in species ofNarcissus (Amaryllidaceae) andHugonia (Linaceae) and other disputed cases

The hypothesis ofHenriques andFernandes that several Iberian species ofNarcissus (Amaryllidaceae) are tristylous is reconsidered. Contrary to the opinion ofBateman and most subsequent authors, we believe that the available evidence indicates that some populations ofN. triandrus andN. fernandesii, at least, are tristylous; other populations ofN. triandrus are distylous.Hugonia cf.penicillanthemum (Linaceae) from new Caledonia is distylous, but it remains possible that other species ofHugonia are tristylous. The disputed occurrence of heterostyly in S. African species ofBauhinia (Leguminosae),Cleome (Capparaceae) andAneilema (Commelinaceae), and inAgelaea (Connaraceae) is discussed.     

Some problems of forest transformation at the transition to the oligocratic/<Emphasis Type="Italic">Homo sapiens</Emphasis> phase of the Holocene interglacial in northern lowlands of central Europe

The paper discusses the main changes in the composition of mixed deciduous forests which occurred mostly between 5,000 and 2,000 b.p., based on selected pollen diagrams from the lowlands of Germany, Denmark and Poland, and including two pollen diagrams from varved sediments, used as reference sites, and on isopollen maps for Poland. The Ulmus retreat is shown on maps, and additional data for its pathogenic origin are presented. Corylus declines at ca. 3,500 b.p. at both reference sites, and its connection with Fagus expansion in the west and Carpinus expansion in the east is discussed. The nature of post-Atlantic transitory shrub–forest communities with dominant Corylus and Quercus is presented. Relationships between the history of Fagus and Carpinus and the development of human settlements are shown. Human impact has been admitted as one of the most important driving forces determining vegetational development since the time of fully developed Neolithic cultures. Other very important abiotic factors were the climate (particularly after 2,500 b.p.), and soil degradation.     

Zur Ultrastruktur und Funktion pollenverbindender Fäden beiEricaceae und anderen Angiospermenfamilien

Viscin threads and other pollen connecting threads of some angiosperm families were investigated, especially those ofEricaceae. According to the definition adopted, viscin threads are ± long exinous processes which consist of exinous material and connect pollen grains or tetrads. Such viscin threads are found within theOnagraceae, Caesalpiniaceae, Ericaceae, andMimosaceae only. While they differ in structure and composition, they always consist of sporopollenin and exhibit a very strong stickiness, even after all viscid substances have been removed by acetolysis. In contrast, the pollen connecting scleroprotein threads ofOrchidaceae and the cellular threads ofStrelitzia reginae Aiton. (Musaceae) are not connected with the exine surface, are destroyed by acetolysis, and thus do not correspond to viscin threads.

     

Die Klebstoffhaare an den Antheren vonCyclanthera pedata (Cucurbitaceae)

The highly specialized, two-celled hairs fringing the anthers ofCyclanthera pedata (L.)Schrad. and its relatives aid in pollination by producing a glue which sticks the coarse pollen grains onto insect visitors. This was proven by direct observations, confirming older assumptions. The adhesive is extruded from the big basal cell which bursts upon the slightest touch to the small top cell.—Turgor and, possibly, swelling pressure of the sticky material, are responsible for plasmoptysis and ejaculation. Cell contraction is due to tension of cuticular clips and, to a minor degree, to elasticity of the pectinic wall. A preformed apical dehiscence line disrupts when the top cell is bent aside.—The anthers hairs of a flower develop synchronously. From light and electron microscopic observations, the adhesive appears not to be a product of internal secretion. It originates from the endopolyploid giant nucleus of the basal cell, which finally degenerates by liquefying and swelling. The numerous peculiar elaioplasts apparently do not contribute to the hair's function.—The anther hairs are homologous to, and possibly have evolved from the so-called explosion hairs occurring on the green parts of manyCucurbitaceae.

Herrn Professor Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Pollination ofOpuntia lindheimeri and related species

The large, yellow, bowl-shaped flowers ofOpuntia lindheimeri, O. discata, O. phaeacantha major, andO. compressa in Texas are visited by various species of beetles and bees. The beetles and small bees (Perdita, Dialictus) are pollen thieves. The pollinators are the medium-sized and larger bees (Melissodes, Diadasia, Lithurge, Megachile, Agapostemon, etc.). Different species of theOpuntia lindheimeri group have the same pollination system and there is no evidence of any floral isolation between them. The pollination system of these species ofOpuntia in Texas is essentially the same as that ofEchinocereus fasciculatus andFerocactus wislizeni in Arizona.Pollination of North American Cacti, II.—SeeGrant & Grant (1979).     

Pollen diameter and fertility in nine species ofPuccinellia (Poaceae)

Diameter and fertility of pollen grains were surveyed in nine species ofPuccinellia Parl. (Poaceae), mainly from the Iberian Peninsula, allowing information about reproductive behaviour and biological characteristics of pollen of these taxa. Maximum diameter was measured with an image processor and fertility was evaluated by staining the grains with Müntzing's mixture. Significant differences in both characters were found between the taxa. In consequence, maximum diameter and fertility proved to have some taxonomic relevance inPuccinellia.     

Hybridization and evolution inCardamine (Brassicaceae) at Urnerboden, Central Switzerland: Biosystematic and molecular evidence

Hybridization between two diploid (2n = 2x = 16) species ofBrassicaceae, Cardamine rivularis andC. amara, at Urnerboden, Central Switzerland, resulted in the rather unusual triploid hybridC. insueta (2n = 3x = 24), and later on in the amphiploidC. schulzii (2n = 6x = 48). The hybrid and the neopolyploid species colonized successfully some man-made biotopes. Plants ofC. insueta are mostly functional females with non-dehiscent anthers, but true hermaphrodite individuals with partly sterile pollen grains also occur within the population. Analyses of cpDNA and nuclear DNA permitted to establish the parentage of the hybrid: the maternal parent which contributed unreduced egg cells proved to beC. rivularis whereas the normally reduced pollen originated fromC. amara. The pronounced genetic variability inC. insueta revealed by isozyme and RAPD analyses, at variance with the polarized segregation, heterogamy and strong vegetative reproduction of the hybrid, is possibly influenced by recurrent formation ofC. insueta which party results from backcrosses betweenC. insueta andC. rivularis but may also proceed by other pathways. The amphiploidCardamine schulzii has normally developed anthers but its pollen is sometimes highly sterile. The surprisingly uniform genetic make-up of the new amphiploid species might be related to its possible monotopic origin and/or young phylogenetic age but should be further assessed. Site management seems to be very important to a further development of hybridogenous populations and their parent species. In conclusion, the evolution at Urnerboden is discussed in the context of the traditional concept of multiple plant origins.     

Formation of sperm cells inGalium mollugo (Rubiaceae),Trichodiadema setuliferum (Aizoaceae), andAvena sativa (Poaceae)

The formation of sperm cells has been examined ultrastructurally in the tricellular pollen grains ofGalium mollugo L. (Rubiaceae).Trichodiadema setuliferum Schwantes (Aizoaceae), andAvena sativa L. (Poaceae). After detachement from the intine the generative cell of all three species lies free within the vegetative cytoplasm. The two sperm cells are built inTrichodiadema andAvena by a single separating wall, while inGalium mollugo two independent walls are formed. However, both mechanisms separate the two male gametes completely.     

Pollination biology ofOpuntia polyacantha andOpuntia phaeacantha (Cactaceae) in southern Colorado

The phenology, compatibility system, and pollinator fauna ofOpuntia polyacantha Haw. andO. phaeacantha Engelm. in southern Colorado were studied and compared. The total blooming periods overlap, but the peak of blooming differs between the species withO. polyacantha blooming first and for longer. Neither species is apomictic,O. polyacantha is largely self-incompatible andO. phaeacantha is self-compatible but may not automatically fully self-pollinate. Examination of flowering structures showed that the former has larger flowers with more perianth parts and heavier (but fewer) anthers. Further the flowers of the allogamous species are visited by greater diversity (but similar abundance) of pollinators. Medium to large bees of the generaDiadasia, Lithurge, Melissodes, Bombus, Agapostemon andMegachile were found to be effective pollinators of the Opuntias studied. The two species were compared as to their floral attributes, breeding systems, and pollinators, and the possible role of competition for pollinators is discussed with respect to its role in their evolutionary paths.     

Pollen morphology and functional dioecy inSolanum (Solanaceae)

Dioecy has evolved independently several times in the large, mostly tropical genusSolanum. In all cases of dioecy inSolanum functionally male flowers have normal anthers, normal pollen and reduced stigmas while functionally female flowers have stigmas and anthers that appear normal but contain non-functional, usually inaperturate pollen. The inaperturate pollen has living cytoplasm, but apparently never germinates and it has been hypothesised that the pollen in these functionally female flowers is retained as a pollinator reward. Pollen morphology is compared in twelve of the thirteen known dioecious species ofSolanum, and some stages in the the development of inaperturate pollen in the anthers of functionally female flowers ofSolanum confertiseriatum of Western Ecuador are examined. Observations on the development and morphology of inaperturate pollen in functionally female flowers ofSolanum are related to hypotheses about the evolution of dioecy in the genus.     

Pollenfertility and seed formation in theOrnithogalum umbellatum/angustifolium complex (Liliaceae/Scilloideae)

The pollen fertility and seed formation of six species of theOrnithogalum umbellatum/angustifolium complex and of seven related species were studied. Four types of pollen grains could be recognized. The pollen fertility varied greatly in this complex and is not related to the ploidy level. The seed formation ofO. umbellatum showed an adaptation to a subcontinental-Mediterranean climate, that ofO. angustifolium to an Atlantic climate. In both cases raindrops seem to be important for pollination, in view of the absence of insect pollinators. After open pollination 113 seedlings were obtained in four species. Their chromosome numbers were determined. Nearly all the cultivated seedlings were aneuploid, which points to a positive selection of euploids in nature, because aneuploid individuals are rare in the wild.Biosystematic Studies on theOrnithogalum umbellatum/angustifolium Complex III.—Previous parts of this series are Part I: Taxonomy. Proceeding Kon. Ned. Acad. Wet. series C,85 (4), 563–574 (1982) andvan Raamsdonk (1984).     

Anchusella, a new genus ofBoraginaceae from the Central-Eastern Mediterranean

The two closely related speciesLycopsis variegata andAnchusa cretica, formerly placed inAnchusa subg.Rivinia, were compared with the type species ofLycopsis andAnchusa, on the basis of a set of macro and microcharacters. The presence of only two fertile stamens as well as other peculiar characters in flower structure, androecium, gynoecium, pollen and fruit, supports the institution of the new genusAnchusella, consisting ofA. variegata andA. cretica. Karyological and eco-chorological aspects are consistent with morphological data in pointing to the autonomy of this genus, which appears characterized by autapomorphic, advanced traits.     

The elm decline and the first appearance of plantago maior

Summary The appearance of pollen grains ofPlantago maior andPlantago lanceolata in the pollen diagrams before and contemporary with the elm decline is, unless accompanied by pollen of Cerealia and archaeological material, not necessarily an indication of the presence of Neolithic man in the immediate vicinity, because of the occurrence of the Agropyro-Rumicion crispi alliance, order Plantaginetalia maioris in naturally unstable contact zones.Translated by Dr.J. J. Butler.     

Middle to Late Holocene vegetation history of the Upper Engadine (Swiss Alps): the role of man and fire

To reconstruct the vegetation and fire history of the Upper Engadine, two continuous sediment cores from Lej da Champfèr and Lej da San Murezzan (Upper Engadine Valley, southeastern Switzerland) were analysed for pollen, plant macrofossils, charcoal and kerogen. The chronologies of the cores are based on 38 radiocarbon dates. Pollen and macrofossil data suggest a rapid afforestation with Betula, Pinus sylvestris, Pinus cembra, and Larix decidua after the retreat of the glaciers from the lake catchments 11,000 cal years ago. This vegetation type persisted until ca. 7300 cal b.p. (5350 b.c.) when Picea replaced Pinus cembra. Pollen indicative of human impact suggests that in this high-mountain region of the central Alps strong anthropogenic activities began during the Early Bronze Age (3900 cal b.p., 1950 b.c.). Local human settlements led to vegetational changes, promoting the expansion of Larix decidua and Alnus viridis. In the case of Larix, continuing land use and especially grazing after fire led to the formation of Larix meadows. The expansion of Alnus viridis was directly induced by fire, as evidenced by time-series analysis. Subsequently, the process of forest conversion into open landscapes continued for millennia and reached its maximum at the end of the Middle Ages at around 500 cal b.p. (a.d. 1450).