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Pollination Ecology Summary

Prof. em. Klaus Ammann, Neuchâtel [email protected]

June 2013 Ohne den Pollenübertragungs-Service blütenbesuchender Tiere könnten sich viele Blütenpanzen nicht geschlechtlich fortpanzen. Die komplexen und faszinierenden Bestäubungsvorgänge bei Blütenpanzen sind Ausdruck von Jahrmillionen von Selektionsvorgängen, verbunden mit Selbstorganisation der Lebewesen; eine Sicht, die auch Darwin schon unterstützte. Bei vielen zwischenartlichen Beziehungen haben sich zwei oder auch mehrere Arten in ihrer Entwicklung gegenseitig beeinusst. Man spricht hier von sogenannter Coevolution. Deutlich ist die Coevolution auch bei verschiedenen Bestäubungssystemen und -mechanismen, die von symbiontischer bis parasitischer Natur sein können. Die Art-Entstehung, die Vegetationsökologie und die Entstehung von Kulturpanzen sind eng damit verbunden

Veranstalter: Naturforschende Gesellschaft Schaffhausen 1. Pollination Ecology Darwin

http://en.wikipedia.org/wiki/Pollination_syndrome http://www.cas.vanderbilt.edu/bioimages/pages/pollination.htm

Fenster, C.B., Armbruster, W.S., Wilson, P., Dudash, M.R., & Thomson, J.D. (2004) Pollination syndromes and floral specialization. Annual Review of Ecology Evolution and Systematics, 35, pp 375-403 http://www.botanischergarten.ch/Pollination/Fenster-Pollination-Syndromes-2004.pdf invitation to browse in the website of the Friends of Charles Darwin http://darwin.gruts.com/weblog/archive/2008/02/ Working Place of Darwin in Downe Village http://www.focus.de/wissen/wissenschaft/wissenschaft-darwin-genoss-ein-suesses-studentenleben_aid_383172.html Darwin as a human being and as a scientist Darwin, C. (1862), On the various contrivances by which orchids are fertilized by insects and on the good effects of intercrossing The Complete Work of Charles Darwin online, Scanned, OCRed and corrected by John van Wyhe 2003; further corrections 8.2006. RN4 edn. Murray, IS: pp. 366, http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F800&pageseq=1 2. Incentives for flower visitors:

http://en.wikipedia.org/wiki/Pollination_syndrome http://www.cas.vanderbilt.edu/bioimages/pages/pollination.htm

Fenster, C.B., Armbruster, W.S., Wilson, P., Dudash, M.R., & Thomson, J.D. (2004) Pollination syndromes and floral specialization. Annual Review of Ecology Evolution and Systematics, 35, pp 375-403 http://www.botanischergarten.ch/Pollination/Fenster-Pollination-Syndromes-2004.pdf 2.1. Nectar Sources Hedera helis, Ivy Nectar presented in open disk

* Allgemeines palustris, Saxifragaceae attractive pseudo-glands offered, no nectar

Nektarsuche Hedera bulbosus

yellow Corolla actually flower-shaped nectaries green perianth, the actual flower

* Ranunculus bulbosus Ranunculus bulbosus, nektaries flower shaped, Ranunculaceae

* Ranunculus bulbosus, Nektarien Ranunculus repens normal spectrum seen by humans ultraviolett spectrum seen by insects

Ranunculus repens, Nektarmal in UV Aconitum paniculatum Ranunculaceae Nectaries prepared

Aconitum paniculatum Nektarien Aconitum vulparia gelber Eisenhut Ranunculaceae Bumblebees digging holes in helmet of flowers in order to steal the pollen from the anthers

Aconitum vulparia, Hummelbesuch Hoverflies (e) visit flowers in a large range of habitats, they even pollinate cultivated . Though they are numerous, -rich and wide-spread, their role as is often still neglected.

Mayer, C.A., Lynn; Armbruster, W. Scott; Dafni, Amots; Eardley, Connal; Huang, Shuang-Quan; Kevan, Peter G.; Ollerton, Jeff; Packer, Laurence; Ssymank, Axel; Stout, Jane C.; Potts, Simon G. (2011) Pollination ecology in the 21st Century: Key questions for future research. Journal of Pollination Ecology, 3, pp 8-23 http://www.ask- force.org/web/Pollination/Mayer- Pollination-Ecology-21fst-Century- Future-2011.pdf

2.2. Flowers for Pollen Eaters Papaver rhaeticum, yellow alpine poppy, Papaveraceae

Papaver rhaeticum Cistus ladaniferus, nectar and pollen collectors

Cistus ladaniferus Commelina coelestis, yellow and conspicous pseudo-anthers

Commelina erecta, Staminodien nigrum attracts flower visitors with colourful hairs of anther filaments

Verbascum nigrum Futterhaare http://www.elizajewett.com/portfolio/pollination.htm When disruptive vibrational signals were applied to grapevine plants through a supporting wire, mating frequency of the leafhopper pest Scaphoideus titanus dropped to 9 % in semi-field conditions and to 4 % in a mature vineyard. The underlying mechanism of this environmentally friendly pest-control tactic is a masking of the vibrational signals used in mate recognition and location. Because vibrational communication is widespread in insects, mating disruption using substrate vibrations can transform many open field and greenhouse based farming systems.

Eriksson, A., G. Anfora, A. Lucchi, F. Lanzo, M. Virant-Doberlet and V. Mazzoni (2012), Exploitation of insect vibrational signals reveals a new method of pest management PLoS One, 7, 3, pp. e32954, http://www.ask-force.org/web/Regulation/Eriksson-Exploitation-Insect- Vibrational-Signals-2012.pdf 2.3. Sexual attraction pseudo-copulation Ophrys fuciflora Orchidaceae

Mimikri for attracting male solitary bees for pseudo copulation

Ophrys fuciflora Ophrys scolopax Orchidaceae

Mimikri for attracting male solitary bees for pseudo copulation

Ophrys scolopax Ophrys lutea Orchidaceae

Mimikri for attracting male solitary bees for pseudo copulation

Ophrys lutea Andrena nigroaenea male pseudocopulating with a flower of the Early Spider Orchid, Ophrys sphegodes www.unet.univie.ac.at/ ~a8917722/

Andrena nigroaenea Long distance Pollination by sexually Attracted male thynnine wasps

King Spider Photo by Heather Whiting Peakall, R. & Beattie, A.J. (1996) Ecological and genetic consequences of pollination by sexual deception in the orchid Caladenia tentactulata. Evolution, 50, 6, pp 2207-2220 http://www.ask-force.org/web/Pollination/Peakall-Ecological-Genetic-Consequences-Caladenia-1996.pdf The sexually attracted male Neozeleboria cryptoides on the stem of the orchid Chiloglottis trapeziformis

/ www.anu.edu.au/BoZo/ orchid_pollination www.anu.edu.au/BoZo/ Australisches Beispiel Pseudokopulation 1 Duftstoffe Gaschromatographie Ophrys apifera Orchidaceae

auto-pollination despite of perfect mimikri for pseudo copulation

Ophrys apifera autogam Ophrys apifera, auto-pollination despite of perfect mimikri for pseudo copulation

Ophrys apifera autogam Nelson. Autopoiesis proof: H-shaped speculum as ancestral character

Nelson: ursprüngliches Lippenmal Nelson. Autopoiesis proof: compact speculum as evolutionary derived character

Nelson abgeleitetes Lippenmal Megachile Dalechampia centuncularis Harzquelle Nestbaustoffe Megachile

2.4. more tricks http://en.wikipedia.org/wiki/Catasetum

Parfum-Sammler: Prachtbienen der Tropen

Parfum-Sammler http://en.wikipedia.org/wiki/Coryanthes Darwins print, copied from Lindley German translation 6. summary of pollinator syndromes 6.1. bee pollination Darwin, C. (1862) On the various contrivances by which orchids are fertilized by insects and on the good effects of intercrossing, The Complete Work of Charles Darwin online, Scanned, OCRed and corrected by John van Wyhe 2003; further corrections 8.2006. RN4 edn. Murray, London, pp 366 http://darwin- online.org.uk/content/fram eset?viewtype=side&itemI D=F800&pageseq=1 Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Himantoglossum hircinum Bocksriemenzunge Orchidaceae Bienen Cephalanthera rubra, rotes Waldvögelein, Orchidaceae, Bienen, Hummeln, Glockenblumen-Mimikri Asclepias curassavica, Milkweed, Pollinien-Bestäubung Asclepias curassavica Milkweed Pollinienbestäubung

http://en.wikipedia.org/wiki/Asclepias description ofpollination see slide 82

http://delta-intkey.com/angio/images/ascle551.gif

http://www.inhs.uiuc.edu/~kenr/prairieplants.A.html Pollination is very unusual in Milkweed, because the pollen is grouped into structures called pollinia rather than being individual grains.. The flowers of the milkweed are very smooth and quite rigid, usually pink or white pink in color, while the are a rounded, oval and waxy to the touch and to view. The insects will slip into notches in the flowers, where the bases of the pollinia which are very sticky, will then attach to their feet, so that they will pull the pollen sacs free when they fly away. Some bees, including some honey bees, only gather nectar from milkweed and since they visit no where else, are not really effective as pollinators for the plants.

Asclepias variegata, White Milkweed, pollinia pollination as Orchids Bakers rule for invasive plants: typically self compatibe pollination

Proportion of fruit-set C: cross-pollination S: self-pollination U: un-manipulated

Ward, M., S. D. Johnson and M. P. Zalucki (2012), Modes of reproduction in three invasive milkweeds are consistent with Baker's Rule Biological Invasions, 14, 6, pp. 1237-1250, ://WOS:000305278400012 AND http://www.ask-force.org/web/Pollination/Ward-Modes-Reproduction-Invaisve-2012.pdf urvilleana,

Tibouchina urvilleana Tibouchina urvilleana

Tibouchina urvilleana pollination Bumblebee urvilleana Tibouchina

http://mgonline.com/media/Images/b/butterfly_garden005.jpg orvala, Karawanken, pollination with bumble bees pratensis, Wiesensalbei left Lamium maculatum, gefleckte Taubnessel right, typical pollination through honey bees Salvia pratensis Wiesensalbei, classic mechanism how bees press anthers on their back Salvia pratensis Wiesensalbei, Lamiaceae

Hebelmechanismus Bienen

Rosmarinus officinalis Rosmarin Lamiaceae

Mechanism female stage Dolchfliege speciosa prächtiger Hohlzahn above: no parking guard rails sidewards yellow and purple nectar traffic sign

Parkverbot oben, Leitplanken seitlich, Saftmal als Verkehrs- zeichen classic honey bee pollination Pedicularis tuberosa knolliges Läusekraut Scrophulariaceae asymmetric flowers only specialists can enter the flowers with a special trick Final remarks about bee pollination Example from the PRRI ASK-FORCE Are GM crops killing honey bees ?

http://pubresreg.org/index.php?option=com_content&task=view&id=63 http://www.spiegel.de/international/world/0,1518,473166,00.html Researchers have found an imported virus that may be associated with the sudden disappearance of honey bees in the United States, known as colony collapse disorder (CCD). This baffling syndrome, which earlier this year made headlines around the world, may have afflicted as many as 23% of beekeepers in the United States and caused losses of up to 90% of hives in some apiaries. The identification of a suspect is an important step, says Nicholas Calderone of Cornell University. “Before, we didn’t even have circumstantial evidence.”

Stokstad, E. (2007) GENOMICS: Puzzling Decline of U.S. Bees Linked to Virus From Australia. Science %R 10.1126/science.317.5843.1304, 317, 5843, pp 1304-1305 http://www.botanischergarten.ch/Bees/Stockstad-IAPV-Culprit-2007.pdf Stokstad, E. (2007) ENTOMOLOGY: The Case of the Empty Hives. Science %R 10.1126/science.316.5827.970, 316, 5827, pp 970-972 http://www.botanischergarten.ch/Bees/Stockstad-Empty-2007.pdf much more information, and stating that the CCD case is still open: http://en.wikipedia.org/wiki/Colony_Collapse_Disorder After extensive training (80 foraging bouts and at least 640 flower visits), bees reduced their flight distances and prioritized shortest possible routes,whilealmost never following nearest-neighbour solutions.

Lihoreau, M., L. Chittka, S. C. Le Comber and N. E. Raine (2012), Bees do not use nearest- neighbour rules for optimization of multi-location routes Biology Letters, 8, 1, pp. 13-16, ://WOS:000299322200005 AND http://www.ask-force.org/web/Pollination/Lihoreau-Bees-not-use-nearest-2013.pdf 6.2. butterfly pollination glacialis, butterfly pollination, long proboscis Silene (Melandrium) rubrum narrow entrance long proposcis

Butterfly pollination Gymnadenia conopea, Orchidaceae, butterfly pollination: long spur Dianthus armeria Rauhe Nelke

Caryophyllaceae

Tagfalter Lilium bulbiferum Feuerlilie Liliaceae

Tagfalter weiss: Nektar-Röhren bavarica, butterfly pollination 6.4. moth-pollination Platanthera chlorantha print from Darwin

Viscidium in distant position for big moths

Platanthera chlorantha, viscidium distant, for big insect heads (moths) Platanthera chlorantha, experiment with reverse pen, viscidia distant Platanthera chlorantha, experiment with reverse pen, after a few seconds Some moth species, however, are exceptional pollinators. Especially well known are the "hummingbird moths" of the Family Sphingidae.

www.kidsbutterfly.org/faq/general/14 Angraecum sesquipedale Angraecum sesquipedale

Orchidaceae moth pollination predicted by Charles Darwin

Pollinator found later: Xanthopan morgani praedictus http://www.criptozoo.com/absolutenm/templates/skin.asp?articleid=217&zoneid=1 http://darwin.gruts.com/weblog/archive/2008/02/ Charles Darwin famously predicted the existence of a species of Madagascan moth (since aptly named Xanthopan morgani praedicta), based on the shape of the nectaries of a species of orchid (Angraecum sesquipedale). Might it not be possible to take a out of Darwin's book and make similar deductions retrospectively? Could parasitologists not study tetrabothiids and other modern parasites, and make deductions about their extinct ancestral hosts' lifestyles? Beyond the Pollination Syndrome

Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were moreconstant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.

Amorim, F. W., L. Galetto and M. Sazima (2013), Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae) Biology, 15, 2, pp. 317-327, ://WOS:000314756900007 AND http://www.ask- force.org/web/Pollination/Amorim-Beyond-Pollination-Syndrome-Nectar-Ecology-2013.pdf Fig. 2. Inga sessilis flower morphology, visitors and pollen germination in a montane ombrophilous dense forest at Nu´ cleo Santa Virginia, Serra do Mar State Park–SP. A: and flower morphology. Note the prominent style (arrow) among the . B: Glossophaga soricina after visiting a flower. C: Short-tongued Erinnyis ello resting on the stamens while accessing nectar. D: Long-tongued hawkmoth Manduca brasiliensis hovering during a visit. E: A perched female Clytolaema rubricauda while taking nectar. F: A female Clytolaema rubricauda approaching a flower. G: Polyads germinating (asterisks) on the stigma of flowers pollinated under natural conditions. H: Pollen tubes of a self- pollinated flower. I: Pollen tubes of a cross-pollinated flower. Beyond the Pollination Syndrome

Amorim, F. W., L. Galetto and M. Sazima (2013), Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae) Plant Biology, 15, 2, pp. 317-327, ://WOS:000314756900007 AND http://www.ask- force.org/web/Pollination/Amorim- Beyond-Pollination-Syndrome-Nectar- Ecology-2013.pdf The diversity and evolution of pollination systems in Annonaceae

Saunders, R. M. K. (2012), The diversity and evolution of pollination systems in Annonaceae Botanical Journal of the Linnean Society, 169, 1, pp. 222-244, ://WOS:000302992000009 AND http://www.ask- force.org/web/Pollination/Sounders- Diversity-Evolution-Pollination- Annonaceae-2012.pdf 6.5. marsupian- pollination Banksia attenuata Proteaceae grobe Bürsten-Blüten- stände, robuste Staubblätter und Narben mit Beutelmäusen Banksia integrifolia, Proteaceae, West-Australien 6.8. bat- pollination Solanaceae Cobaea sp. Underwoods longtongued Bat Hylonycteris underwoodii

Carpobrothus edulis, Mesembryanthemaceae, Fledermäuse Solandra nitida, Solanaceae, Fledermäuse Thunbergia japonica Acanthaceae Grosse Hautflügler Fledermäuse, Vögel robuste Trompeten- form 6.5. bird- Pollination Aechmea sp. Bromeliaceae Hochblätter rot Kolibri Cata, Venezuela Aechmea, Bromeliaceae, rote Hochblätter Nektar weisslich eintrocknend, Kolibri, Cata, Venezuela Strelitzia reginae, Musaceae, Südafrika, Vögel stehen auf Antheren Columnea fawcettii, Gesneriaceae, epiphytisch, Jamaica, Vögel botit.botany.wisc.edu/.../ Gesneriaceae/Columnea/ Cattleya aurantiaca, Orchidaceae, Vögel Amherstia nobilis, Caesalpiniaceae, ursprünglich Ostasien, Vögel Cestrum hirtum, Solanaceae, Kolibris 6.9. beetle- pollination Gottsberger, G., I. Silberbauer- Gottsberger, R. S. Seymour and S. Dotterl (2012), Pollination ecology of Magnolia ovata may explain the overall large flower size of the genus Flora, 207, 2, pp. 107- 118, ://WOS:0003019 10900003 Viburnum opulus, gemeiner Schneeball, Käfer usw. Nymphaea alba, weisse Seerose, Nymphaeaceae, Käfer Thalictrum flavum gelbe Wiesenraute Ranunculaceae Canthariden, Weichkäfer 6.10. beetle and fly- pollination Heracleum sphondylium, Bärenklau, Käfer, Fliegen Heracleum sphondylium, Bärenklau, Randblüten mit Schaufunktion Peucedanum ostruthium, Meisterwurz, Blitz in Dämmerung filiformis, Faden-Ehrenpreis, Scrophulariaceae, Fliegen Veronica cymbalaria, Fliegenbestäubung minima, kleiner Augentrost, Fliegen Tozzia alpina, Tozzie, Scrophulariaceae, Fliegen Sarracenia flava karnivor

North Carolina fliegenbestäubt Arum maculatum Spatha, Kolben und Reusenfalle Fliegen Arum maculatum Spatha, Kolben und Reusenfalle Fliegen

Arum maculatum Aronstab Flug- und Gleitbahnen der Fliegen Arum maculatum, Aronstab links: junge Reusen, Anth. geschlossen, Nektartropfen rechts: Reusenhaare verdorrt, Antheren offe The titan arum or Amorphophallus titanum is a with the largest unbranched inflorescence in the world. The largest single flower is borne by the Rafflesia arnoldii; the largest branched inflorescence in the plant kingdom belongs to the Talipot palm (Corypha umbraculifera). The titan arum was originally discovered by an Italian botanist, Odoardo Beccari, in Sumatra in 1878. It thrives at the edges of rainforests near open grasslands. Though found in many botanic gardens around the world it is still indigenous only to the tropical forests of Sumatra. Due to its fragrance, which is reminiscent of the smell of a decomposing mammal,[1] the titan arum is also known as a carrion flower, the "Corpse flower", or "Corpse plant" (in Indonesian, "bunga bangkai" – bunga means flower, while bangkai means corpse or cadaver; for the same reason, the same title is also attributed to Rafflesia which, like the titan arum, also grows in the rainforests of Sumatra).

http://en.wikipedia.org/wiki/Titan_arum Ollerton, J. & Raguso, R.A. (2006) The sweet stench of decay. New Phytologist, 172, 3, pp 382-385 http://www.botanischergarten.ch/Pollination/Ollerton-Sweet-Stench-2006.pdf Stapelia, Aasfliegen Rafflesia pricei, Sabah, Borneo Tambunan Rafflesia Centre in the Crocker Range National Park, Sabah, Malaysia Rafflesia: Big Daddy of the Spurge Family Unassuming cousin, Euphorbia baylissii Photo: Frank Vincentz, via Wiki

Davis, C.C., Latvis, M., Nickrent, D.L., Wurdack, K.J., & Baum, D.A. (2007) Floral Gigantism in Rafflesiaceae. Science, pp 1135260 http://www.botanischergarten.c h/Pollination/Davis-Rafflesia- Gigantism-Evolution-2007.pdf Bulbophyllum careyanum Orchidaceae Aasfliegen Bulbophyllum careyanum Orchidaceae Aasfliegen Bulbophyllum sp. Aasfliegen 6.11. pollinators unknown Altensteinia (Aa), within Sphagnum in Andes of Venezuela tiny 4mm-flowers on, ca. 4000m altitude, pollinator unknown Antelope Orchid Dendrobium minax Orchidaceae Botanic Garden in Bern pollinator ?

6.12. closing Measurements of mating success. Left: the self-fertilization allele is shown as closed circles and outcrossing allele as open circles. Transmission pathways from parent to offspring are shown as arrows; solid arrows represent gene transmission to progeny by the parent capable of self-fertilization, while dashed arrows represent transmission pathways for the outcrossing parent. Fitness is measured as the sum of gene copies transmitted by pollen and ovules. Assuming that the number of pollen grains produced is large relative to the number of ovules and that all the ovules are fertilized, a selfing genotype enjoys a 50% advantage in gene transmission relative to a outcrossing genotype. This results in the cost of outcrossing (Fisher, 1941). Right: the large circles represent individual parents and their progeny. Arrows represent the production of discrete offspring by parents, with solid arrows representing pollinator-independent offspring production, and dashed arrows representing pollinator-dependent offspring production. Mating system is often equated with the number of seeds produced. Selfing ensures offspring production when lack of cross-pollinators limits seed-set (Darwin, 1876). Cheptou, P. O. (2012), Clarifying Baker's Law Annals of Botany, 109, 3, pp. 633-641, ://WOS:000300488700013 AND http://www.ask-force.org/web/Pollination/Cheptou-Clarifying-Bakers-Law-2012.pdf Cheptou, P. O. (2012), Clarifying Baker's Law Annals of Botany, 109, 3, pp. 633-641, ://WOS:000300488700013 AND http://www.ask-force.org/web/Pollination/Cheptou-Clarifying-Bakers-Law-2012.pdf Joint evolution of dispersal and self-fertilization in a temporally heterogeneous pollination environment (from Cheptou and Massol, 2009; Massol and Cheptou, 2011). The model assumes an infinite island model where each patch is prone to temporally heterogeneous pollination because of the stochastic presence of pollinator (see Kalisz et al., 2004, for empirical support). We consider a basic mechanism of self-fertilization: plants self-fertilize a proportion, s, of their ovules while the proportion 1 – s can be fertilized by outcrossing only if pollinators are present, which occur with a probability (1 – e). Otherwise, the fraction 1 – s is left unfertilized ( probability e). Seeds produced disperse at random among patch at rate d but only a fraction q survive to dispersal. Progeny produced by self-fertilization suffer from inbreeding depression (d).

Model outcome: the mathematical analysis (invasion analysis) reveals that only two syndromes of traits are possible: the ‘dispersal/outcrossing’ syndrome (d* ¼ (e/[1 – q(1 – e)], s* ¼ 0) and the ‘no-dispersal/selfing’ syndrome (d* ¼ 0, s* ¼ (2e/[2d + e – 1])) (see Cheptou and Massol, 2009, for the influence of parameters). Pollination fluctuations suffered by out-crossers create among-patch variance and thus favour dispersal (see Comins et al., 1982). In contrast, selfers do not suffer from pollination heterogeneity, thus cancelling selection pressure for dispersal.

Cheptou, P. O. (2012), Clarifying Baker's Law Annals of Botany, 109, 3, pp. 633-641, ://WOS:000300488700013 AND http://www.ask-force.org/web/Pollination/Cheptou-Clarifying-Bakers-Law-2012.pdf Mass-flowering crops enhance wild bee abundance

Fig. 1 The mean number (±SE) of Red Mason Bee brood cells in trap nests in a oilseed rape fields adjacent to grassland (oilseed rape ? grassland; n = 17) versus isolated oilseed rape fields (oilseed rape - grassland; n = 17; model with presence–absence data), b grasslands adjacent to oilseed rape grassland ? oilseed rape; n = 17) versus isolated grasslands (grassland - oilseed rape; n = 16; model with number of brood cell data)

Holzschuh, A., C. Dormann, T. Tscharntke and I. Steffan-Dewenter (2013), Mass-flowering crops enhance wild bee abundance Oecologia, 172, 2, pp. 477-484, http://dx.doi.org/10.1007/s00442-012-2515-5 AND http://www.ask-force.org/web/Pollination/Holzschuh-Mass-Flowering-Bee-Abundance-2013.pdf Flowering Phenology as functional Trait in Prairie Communities

Fig. 1 (a) Patterns of first flowering dates (FFDs) for Konza herbaceous flora expressed as a fraction of all recorded species, flowering per d or number of species (S) flowering per d (S d–1).

Also shown are FFDs compared between: (b, c) functional groups (C3 Cyperaceae, C3 , C4 euDicots, C3 Poaceae, and C4 Poaceae);

(d, e) native and nonnative species; and (f, g) life history groups (annual, biennial, and perennial species).

Craine, J. M., E. M. Wolkovich, E. G. Towne and S. W. Kembel (2012), Flowering phenology as a functional trait in a tallgrass prairie New Phytologist, 193, 3, pp. 673-682, ://WOS:000298984900013 AND http://www.ask- force.org/web/Pollination/Craine- Flowering-Phenology-Tallgrass- 2011.pdf Modularity of the network of interactions considering the pollination syndrome in high- altitude grasslands at Serra da Bocaina, southeastern Brazil.

Interactions were not equally predicted by different pollination syndromes, and the accuracy of the prediction was strongest for ornithophily and melittophily.

White circles: bees and plants related to melittophily, black: flies and plants related to myophily, ash: butterflies and plants related to psychophily, and striped: hummingbirds and plants related to ornithophily.

(A) Modules considering all the interactions:

(A) Modules considering all the interactions: 1a - 14 species (6 melittophilous plants, 8 bees); 2a - 45 species (18 melittophilous plants, 2 psychophilous plants, 1 miophilous plant; 17 bees, 4 butterfl ies, 2 wasps, 1 fl y); 3a - 35 species (17 melittophilous plants, 2 miophilous plants, 9 bees, 4 fl ies, 3 wasps); 4a - 30 species (7 melittophilous plants, 2 psychophilous plants, 1 miophilous plant, 3 bees, 5 butterfl ies, 8 fl ies, 4 wasps); 5a - 26 species (8 melittophilous plants, 3 psychophilous plants, 8 bees, 1 wasp, 3 fl ies, 3 butterfl ies); 6a - 25 species (5 Danieli-Silva, A., J. M. T. de Souza, A. J. Donatti, R. P. Campos, J. miophilous plants, 3 melittophilous plants, 8 wasps, Vicente-Silva, L. Freitas and I. G. Varassin (2012), Do pollination 8 fl ies,1 butterfl y); 7a - 29 species (7 miophilous plants, 4 melittophilous plants, 11 fl ies, 4 syndromes cause modularity and predict interactions in a pollination wasps, 3 bees); 8a - 7 species (2 melittophilous network in tropical high-altitude grasslands? Oikos, 121, 1, pp. 35-43, plants, 4 bees, 1 fl y); 9a - 40 species (10 miophilous plants, 1 melittophilous plant, 22 ://WOS:000298484200004 AND http://www.ask- wasps, 4 fl ies, 2 bees, 1 butterfl y); 10a - 2 species (1 melittophilous plant, 1 bee); 11a - 10 species (5 ornithophilous plants, 5 hummingbirds); force.org/web/Pollination/Danieli-Silva-Pollination-Syndromes- 12a - 3 species (1 miophilous plant, 1 fl y, 1 Grassland-2012.pdf wasp); 13a - 2 species (1 miophilous plant, 1 wasp). Dangles, O. & Casas, J. (2012) The bee and the turtle: a fable from Yasuni National Park. Frontiers in Ecology and the Environment, 10, 8, pp 446-447 http://www.esajournals.org/doi/abs/10.1890/1540-9295-10.8.446 AND http://www.ask-force.org/web/Pollination/Dangles-Bee-Turtle-Yasuni-NP-2012.pdf

Träne im Saugrüssel Eine Einsiedlerbiene saugt Tränenflüssigkeit aus dem Auge einer Schienen schildkröte im ecuadorianischen Yasuní- Nationalpark. Die salzigen Tränen dienen dem Insekt vermutlich als Natriumquelle. Yashina, S., S. Gubin, S. Maksimovich, A. Yashina, E. Gakhova and D. Gilichinsky (2012), Regeneration of whole fertile plants from 30,000-y-old fruit tissue buried in Siberian permafrost Proceedings of the National Academy of Sciences, 109, 10, pp. 4008- 4013, http://www.pnas.org/content/109/10/4008.abstra ct AND http://www.ask- force.org/web/Evolution/Yashina-Regeneration- whole-plant-Silene-Stenophylla-2012.pdf

30’000 jährige Silene-Art aus einer sibirischen Höhle wieder zum Leben erweckta