1.6. Pollination ecology
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 1.6.1. incentive: Sources of nectar and pollen INCENTIVES FOR VISITING FLOWERS Plants can not move from one location to another, thus many flowers have evolved to attract animals to transfer pollen between individuals in dispersed populations. Flowers that are insect-pollinated are called entomophilous; literally "insect-loving" in Latin. They can be highly modified along with the pollinating insects by co-evolution. Flowers commonly have glands called nectaries on various parts that attract animals looking for nutritious nectar. Birds and bees have color vision, enabling them to seek out "colorful" flowers. Some flowers have patterns, called nectar guides, that show pollinators where to look for nectar; they may be visible only under ultraviolet light, which is visible to bees and some other insects. Flowers also attract pollinators by scent and some of those scents are pleasant to our sense of smell. Not all flower scents are appealing to humans, a number of flowers are pollinated by insects that are attracted to rotten flesh and have flowers that smell like dead animals, often called Carrion flowers including Rafflesia, the titan arum, and the North American pawpaw (Asimina triloba). Flowers pollinated by night visitors, including bats and moths, are likely to concentrate on scent to attract pollinators and most such flowers are white. Still other flowers use mimicry to attract pollinators. Some species of orchids, for example, produce flowers resembling female bees in color, shape, and scent. Male bees move from one such flower to another in search of a mate. http://en.wikipedia.org/wiki/Flower#Attraction_methods Hedera helis, Ivy Nectar presented in open disk
* Allgemeines Hedera helix, Efeu, Araliaceae Nektarsuche Hedera Parnassia palustris, Saxifragaceae attractive pseudo-glands offered, no nectar
Nektarsuche Hedera Parnassia palustris Studentenröschen Saxifragaceae
Parnassia palustris, Pseudonektarien, schlussendlich Selbstbestäubung Ranunculus bulbosus Ranunculaceae
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 various shapes of nectaries with Ranunculaceae Nektarien Ranunculaceae Aquilegia vulgaris, Akelei, Ranunculaceae corolla and nectaries
Aquilegia vulgaris Aquilegia vulgaris nectaries only Ranunculaceae
Aquilegia vulgaris Aconitum napellus Ranunculacaee
Aconitum napellus Aconitum paniculatum, Ranunculaceae, view from the side
Aconitum paniculatum Aconitum paniculatum frontal view Ranunculaceae
Aconitum paniculatum Aconitum paniculatum Ranunculaceae Nectaries prepared
Aconitum paniculatum Nektarien Aconitum septentrionale
Ranunculaceae
Aconitum septentrionale Aconitum vulparia gelber Eisenhut Ranunculaceae Bumblebees digging holes in helmet of flowers in order to steal the pollen from the anthers
Aconitum vulparia, Hummelbesuch Examples of the amazing breadth of pollination ecology: The dove tree (Davidia involucrata ) has “naked” flowers that lack a perianth but are surrounded by two large bracts. These turn from green (a) to white (b) when flowers open which increases pollinator attraction Sun, J.F., Gong, Y.B., Renner, Susanne S., & Huang, S.Q. (2008) Multifunctional Bracts in the Dove Tree Davidia involucrata (Nyssaceae: Cornales): Rain Protection and Pollinator Attraction, Vol. 171, pp. 119-124. The University of Chicago Press for The American Society of Naturalists, http://www.ask-force.org/web/Pollination/Sun-Davidia-Multifunctional-Bracts-2008.pdf Examples of the amazing breadth of pollination ecology: Calypso bulbosa (c), a rewardless orchid, has evolved to deceive naïve bumblebees in the early spring to effect pollination without payment in nectar
Dafni, A. (1983) Pollination of Orchis Caspia--A Nectarless Plant which Deceives the Pollinators of Nectariferous Species from other Plant Families. Journal of Ecology, 71, 2, pp 467-474 http://www.ask- force.org/web/Pollination/Dafni- Pollination-Orchis-Deceives-1983.pdf Ackermann, J.D. (1981) Pollination Biology of Calypso bulbosa var. occidentalis (Orchidaceae), a Food Deception System. Madrono, 28, 3, pp 101-110 http://www.ask-force.org/web/Pollination/Ackermann- Calypso-bulbosa-Food-Deception-1981.pdf The perennial vine, Gelsemium sempervirens (d), contains alkaloids in all plant parts including corollas and nectar which deter floral herbivores and nectar robbers but - in high concentrations - also pollinators
Adler, L.S. & Irwin, R.E. (2005) Ecological costs and benefits of defenses in nectar. Ecology, 86, 11, pp 2968-2978 http://www.ask- force.org/web/Pollination/Adler- http://alabamaplants.com/Yellowopp/Gelsemi Ecological-Costs-Benefits-2005.pdf um_sempervirens_page.html
Adler, L.S. & Irwin, R.E. (2005) Ecological costs and benefits of defenses in nectar. Ecology, 86, 11, pp 2968-2978 http://www.ask-force.org/web/Pollination/Adler-Ecological-Costs-Benefits-2005.pdf Hoverflies (e) visit flowers in a large range of habitats, they even pollinate cultivated plants. Though they are numerous, species-rich and wide-spread, their role as pollinators 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
Rhododendron ponticum (h) is a highly invasive plant in Ireland. Though it provides food resources for generalist bumblebees, effects on native plant pollinator interactions vary with intensity of invasion.
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 Heterochelus chiriagricus (Striped monkey beetle), 10mm. [image by M. Picker & C. Griffiths ©, from Field Guide to Insects of South Africa, used with permission]. http://www.biodiversityexplorer.org/beetles/scarabaeidae/r utelinae/hopliini.htm
In the Succulent Karoo of South Mayer, C., Soka, G., & Picker, M. (2006) Africa, monkey beetles (Hopliini) The importance of monkey beetle (Scarabaeidae : Hopliini) pollination for Aizoaceae and Asteraceae in grazed and are important pollinators of many ungrazed areas at Paulshoek, Succulent Karoo, South Africa. plant species Journal of Insect Conservation, 10, 4, pp 323-333 http://www.ask-force.org/web/Pollination/Mayer-Importance- Monkey-Beetle-2006.pdf 1.6.2. Flowers for Pollen Eaters * Pollensammeln, Futterhaare SYNDROME OF FLOWERS VISITED BY POLLEN EATERS considerable amounts of pollen easily accessed as sources, usually offered by flowers without nectar. Typical Flowers: Poppy Papaver, Cistus, Commelina, Rosa, Thalictrum aquilegiifolium sometimes also with intricate deception strategies such as pseudopollinia, see the case of Comelina below. sometimes also fodder hairs and attractive oil sources offered Verbascum nigrum, Lysimachia vulgaris
Papaver rhaeticum, yellow alpine poppy, Papaveraceae
Papaver rhaeticum Cistus albiflorus, visited by pollen eaters
Cistus albiflorus Cistus ladaniferus, nectar and pollen collectors
Cistus ladaniferus Commelina coelestis, yellow and conspicous pseudo-anthers
Commelina coelestis Commelina coelestis, yellow and conspicous pseudo-anthers
Commelina erecta, Staminodien Thalictrum aquilegiifolium coloured anther filaments as attraction for flower visitors
Thalictrum aquilegiifolium Verbascum 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 1.5.3. sexual attraction pseudo-copulation Pseudocopulation describes behaviors similar to copulation that serve a reproductive function for one or both participants but do not involve actual sexual union between the individuals. It is most generally applied to a pollinator attempting to copulate with a flower. Some flowers mimic a potential female mate visually, but the key stimuli are often chemical and tactile.[1] This form of mimicry in plants has been titled Pouyannian mimicry.[2] Orchids commonly achieve reproduction in this manner, secreting chemicals from glands called osmophores located in the sepals, petals, or labellum, that are indistinguishable from the insect's natural pheromones. The pollinator then has a pollinia attached to its body, which it transfers to the stigma of another flower when if it attempts another 'copulation'. Pollinators are often bees and wasps of the order Hymenoptera, and flies. from http://en.wikipedia.org/wiki/Pseudocopulation Ophrys insectifera, pseudo copulation as incentive
* Ophrys insectifera: Pseudocopulation 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 scolopax Orchidaceae
Mimikri Bienen
Ophrys scolopax Ophrys lutea Orchidaceae
Mimikri for attracting male solitary bees for pseudo copulation
Ophrys lutea 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 orchid Leporella fimbriata is pollinated by pseudocopulati on with winged males of the ant Myrmecia urens.
Peakall, R., Beattie, A.J., & James, S.H. (1987) Pseudocopulation of an orchid by male ants: a test of two hypotheses accounting for the rarity of ant pollination. Oecologia, 73, 4, pp 522-524 http://www.ask-force.org/web/Pollination/Peakall-Pseudocopulation-Orchid-Male-Ants-1987.pdf
The sexually attracted male pollinator 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
pr
on the labellum of the orchid Chiloglottisof the orchid on the labellum on the labellum of the orchid Chiloglottisof the orchid trapeziformis on the labellum
Neozeleboria cryptoides Neozeleboria Neozeleboria cryptoides Neozeleboria carrying pollen (pollinia) that was removed during attempted copulation copulation that attempted was during removed (pollinia) pollen carrying
trapeziformis flower a with previous (pseudocopulation) The male pollinator The pollinator male The pollinator male plac take will removal pollen process this During (pseudocopulation). the with flower mating attempted www.anu.edu.au/BoZo/ orchid_pollination/ Australische Beispiele Pseudokopulation 2 Duftstoffe Gaschromatographie 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 Sexually deceptive pollination systems are significantly more efficient than food deceptive systems. But the rewarding flowers most are the most successful in insect pollination.
Scopece, G., S. Cozzolino, S. D. Johnson and F. P. Schiestl (2010), Pollination Efficiency and the Evolution of Specialized Deceptive Pollination Systems, American Naturalist, 175, 1, pp. 98-105,
Figure 1: Comparison of measures of reproductive success among orchids with different pollination strategies (rewarding, food- deceptive, and sexually deceptive). MRS p male reproductive success; FRS p female reproductive success; PE p pollination efficiency. Different letters indicate significant differences (Mann-Whitney U-test, P! .01 after Bonferroni correction). Circles represent outliers. 1.6.4. offering nesting materials Megachile, solitary leaf cutter bees, building nests from cut leaves http://en.wikipedia.org/wiki/Megachile
* Nestbau Mohnbiene Megachile Dalechampia centuncularis Harzquelle Nestbaustoffe Megachile
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 http://en.wikipedia.org/wiki/Coryanthes www.weloennig.de/CorCat.html
1.6.5. summary of pollinator syndromes bee pollination Bee pollination (melittophily) Bee-pollinated flowers tend to fall into two classes: Showy, open, bowl-shaped flowers that are relatively unspecialized (e.g. wild roses, sunflowers) Showy, complicated, non-radially symmetric flowers that are more specialized (e.g. peas, foxgloves) Some bee flowers tend to be yellow or blue, often with ultraviolet nectar guides and scent. Nectar, pollen, or both are offered as rewards in varying amounts. The sugar in the nectar tends to be sucrose-dominated. There are diverse types of bees, however. Honeybees, bumblebees, orchid bees, etc are large groups that are quite distinctive in size, tongue length and behaviour (some solitary, some colonial). Thus generalization about bees is difficult (Fenster at al. 2004.) Some plants can only be pollinated by bees because their anthers release pollen internally, and it must be shaken out by buzz pollination. Bees are the only animals that perform this behaviour http://en.wikipedia.org/wiki/Pollination_syndrome#Bee_pollination_.28melittophily.29 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
1: 3 sepals (outer) and 3 pepals (inner whirl
3: central stamen, Anther, carries 2 pollinia
2: gynostemium or 4: two other stamen reduced to column staminodes http://en.wikipedia.org/wiki/Orchid
5: Rostellum 7: ovary, inferior 9: stigma- covers cave sticky 8: spur, in disks Orchis and 6: Labellum Dactylorhiza without nectar http://www.denniskunk el.com/DK/Plants/893 1B.html
http://www.orchids.co.in/plant-facts/orchid-pollination.shtm In the tribe Orchideae , the pollinia are included in special bags called bursicles ,which slit open upon maturation . As soon as the insect enters the flower, the two viscidia attach themselves to the two sides of the insect’s head. When the insect leaves the flower , the two pollinia are drawn from the bursicles and carried away on the insect’s head. By a special hygrophilous quality of the caudicle ,the pollinia which are at first held upright on the insect’s head slowly bend forward , till it comes to occupy a position at right angles to the insect’s head . In this position ,when the insect visits the next flower , the pollinia immediately come in contact with the stigma . This hygrophilous nature of the caudicle is characteristic of some of the advanced Sarcanthine orchids such as Vanda teres , Luisia sp.etc. Here the caudicle is transparent and ribbon-shaped . Immediately after removal from the anther it appears straight . But while it is being carried on the insect’s head, it absorbs it appears straight . But while it is being carried on the insect’s head , it absorbs water and curves in an S-shaped manner Dactylorrhiza bringing the pollinia forward , so that they are deposited on the stigma maculata of the flower which the insect visits next Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Orchis purpureus, Purpur-Knabenkraut, Orchidaceae, Pollinien-Versuch Reproductive Isolation Orchis mascula / Orichs pauciflora
Scopece, G., A. Croce, C. Lexer and S. Cozzolino (2013), COMPONENTS OF REPRODUCTIVE ISOLATION BETWEEN ORCHIS MASCULA AND ORCHIS PAUCIFLORA, Evolution, 67, 7, pp. 2083-2093,
Orchis Orchis pauciflora mascula http://www.gmpao.org/fichiers/go_crete2.htm http://es.wikipedia.org/wiki/Orchis_mascula Orchis pauciflora x Orchis mascula subsp. mascula (=Orchis colemannii)
http://www.orchis.de/orchis/scripts/fset .php?id=193&bildart=hi Himantoglossum hircinum Bocks-Riemenzunge
Bienen Himantoglossum hircinum Bocksriemenzunge Orchidaceae Bienen Adenostyles leucophylla, filziger Alpendost, Asteraceae, Bienen usw. Cephalanthera rubra, rotes Waldvögelein, Orchidaceae, Bienen, Hummeln, Glockenblumen-Mimikri Orchis morio, kleine Orchis, Orchidaceae, Bienen Orchis coryophora Wanzen-Knabenkraut
Orchidaceae
Bienen Orchis papilionaceae Schmetterlings- Knabenkraut Orchidaceae
Bienen Asclepias curassavica, Milkweed, Pollinien-Bestäubung Asclepias curassavica Asclepias curassavica Milkweed Pollinienbestäubung Asclepias curassavica. One flower with the gynostegium in longitudinal section. Note the gland (corpusculum) and pollinia in the upper left. The corolla tube has been dissected to show the 2 free, superior ovaries, while the top of the gynoecium is connate and then adnate to the androecium to form part of the gynostegium. Asclepias curassavica
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 leaves 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 pollination bee asperula Asclepias www.sbs.utexas.edu/.../asclepias_asperula.htm
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,
Tibouchina urvilleana Tibouchina urvilleana
Tibouchina urvilleana pollination Bumblebee urvilleana Tibouchina
http://mgonline.com/media/Images/b/butterfly_garden005.jpg Lamium orvala, Karawanken, pollination with bumble bees Lamium maculatum gefleckte Taubnessel Lamiaceae Bumble bees and bees Salvia pratensis, Wiesensalbei left Lamium maculatum, gefleckte Taubnessel right, typical pollination through honey bees Salvia pratensis Wiesensalbei, Lamiaceae classic mechanism how bees press anthers on their back Salvia pratensis Wiesensalbei, Lamiaceae
Hebelmechanismus Bienen
Rosmarinus officinalis Rosmarin Lamiaceae
Mechanism female stage Dolchfliege Galeopsis speciosa prächtiger Hohlzahn Scrophulariaceae 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 Walter Haefeker, the German beekeeping official, speculates that "besides a number of other factors," the fact that genetically modified, insect-resistant plants are now used in 40 percent of cornfields in the United States could be playing a role. The figure is much lower in Germany -- only 0.06 percent -- and most of that occurs in the eastern states of Mecklenburg- Western Pomerania and Brandenburg. Haefeker recently sent a researcher at the CCD Working Group some data from a bee study that he has long felt shows a possible connection between genetic engineering and diseases in bees. 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,
http://en.wikipedia.org/wiki/Pollination_syndrome#Bee_pollination_.28melittophily.29 Dianthus 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 Dianthus armeria, rauhe Nelke, Caryophyllaceae, Tagfalter, Cirsium oleraceum Kohldistel Asteraceae viele Bestäuber Lilium bulbiferum Feuerlilie Liliaceae
Tagfalter weiss: Nektar-Röhren Gentiana bavarica, butterfly pollination Parnassius apollo on Centaurea spinosa Kure Mountains, Turkey
www.pbase.com/efratnakash/image/80997440 moth- pollination Moth pollination (phalaenophily) Day-flying sphinx moth nectaring on Brazilian verain Among the more important moth pollinators are the hawk moths (Sphingidae). Their behaviour is similar to hummingbirds: they hover in front of flowers with rapid wingbeats. Most are nocturnal or crepuscular. So moth-pollinated flowers tend to be white, night-opening, large and showy with tubular corollas and a strong, sweet scent produced in the evening, night or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight. Other moths (Noctuids, Geometrids, Pyralids, for example) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads) (Oliveira et al. 2004).
day-flying sphynx moth nectaring on Brazil vervain 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 leaf 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) Plant Biology, 15, 2, pp. 317-327,
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,
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,
Beutler und Vögel Melaleuca quinquenervia Myrtaceae
Säuger und Vögel bat- pollination Saguaro Cereus giganteus Fleder- mäuse Saguaro Cereus giganteus
Fledermäuse Agave sp. Agavaceae Fleder- mäuse Solanaceae Cobaea sp. Underwoods longtongued Bat Hylonycteris underwoodii
Opuntia humifusa, niederliegende Opuntie, Fledermäuse Carpobrothus edulis, Mesembryanthemaceae, Fledermäuse Solandra nitida, Solanaceae, Fledermäuse Dombeya wallichii Sterculiaceae, Madagaskar Fledermäuse Dombeya wallichii, Sterculiaceae, Madagaskar, Fledermäuse Thunbergia japonica Acanthaceae Grosse Hautflügler Fledermäuse, Vögel robuste Trompeten- form 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 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 Anderson, S.H., Kelly, D., Ladley, J.J., Molloy, S., & Terry, J. (2011) Cascading Effects of Bird Functional Extinction Reduce Pollination and Plant Density Science, Science 1199092 Published online 3 February 2011 pp http://www.ask-force.org/web/Pollination/Anderson-Cascading-Effects-Birds-Extinction-2011.pdf AND http://www.sciencemag.org/search?site_area=sci&fulltext=%22dave%20kelly%22&submit=yes Supporting Material http://www.ask-force.org/web/Pollination/Anderson-Cascading-Effects-Birds-Extinction-Supporting-2011.pdf http://www.sciencemag.org/content/early/2011/02/02/science.1199092/suppl/DC1 AND blog material http://www.ouramazingplanet.com/birds-plants-pollination-biodiversity-loss-1058/ A male stitchbird (or hihi) on Tiritiri Matangi Island. This important pollinator vanished off the North Island with the arrival of mammalian predators 140 years ago, but persists on nearby islands. Where stitchbirds and bellbirds survive, the pollination mutualism still works well. Credit: Dave Kelly, University of Canterbury http://www.ouramazingplanet.com/birds-plants-pollination-biodiversity-loss-1058/ for for successful fruit set. [Photo: M Walters] presentation isfinished, so the flowers require bird visitors elongatesbird.stigma The pollena after been visitedbyhas and the ridged fused pollen disc whichismarked theif flower underneath, showing the narrow 10mm long corolla tube, A flower of Rhabdothamnus solandri viewed from
on the mainland, but not on islands (post hoc tests, text). see tests, hoc (post islands on not but mainland, the on worse than hand - significantly did flowers (unmanipulated) natural cases, both In size). from fruit set set fruit % (mean success Pollination (A) high. remain densities bird where offshore islands adjacent on not but extinct, are functionally pollinators bird its wheremainland, Zealand shrub birdpollinated the for limitation pollen strong for Evidence ±
SEM). (B) Seeds per fruit SEM). fruit (B) (mean per Seeds and SEM,estimated Rhabdothamnus solandri on the New the on solandri Rhabdothamnus pollinated flowers
Anderson et al. 2011 significantly fewer seedlings on the mainland ( mainland the on seedlings fewer significantly ( densities adult hadsimilar sites andmainland Island densities). bird (low and the mainland on centered cm) (<30 juvenile on the adult of solandri densities mainland: (≥30 cm and tall) of failure regeneration for Evidence R. solandri adults, on islands (high bird densities) bird (high on islands adults, solandri R. R. solandri plants (mean ± (mean plants solandri R. were there 0.56), but P = Rhabdothamnus
SEM) in plots SEM) in P = 0.020). P =
Anderson et al. 2011 solandri: densitiesseedlingsOctober2008 (mean of in limitationEvidenceseed of for Melicytus ramiflorusGeniostoma or ligustrifolium (see text). significantly highersownplots in for to adjacent unsown plots. Seedling density after 5yearswas augmentationmainlandthe on in plots April2003, compared seed threefrom SEM)20 cm species x sowninto 10 Rhabdothamnus R. solandri, but not forsolandri,not R. but
±
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,
Käfer, Fliegen usw. Heracleum sphondylium, Bärenklau, Käfer, Fliegen Heracleum sphondylium, Bärenklau, Randblüten mit Schaufunktion Anthriscus silvestris, Wiesenkerbel, Früchte mit Nektar-Scheiben Peucedanum ostruthium, Meisterwurz, Blitz in Dämmerung Veronica filiformis, Faden-Ehrenpreis, Scrophulariaceae, Fliegen Veronica cymbalaria, Fliegenbestäubung Euphrasia minima, kleiner Augentrost, Fliegen Tozzia alpina, Tozzie, Scrophulariaceae, Fliegen Sarracenia flava, karnivor, Insektenfallen, fliegenbestäubt Sarracenia flava karnivor
North Carolina fliegenbestäubt 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 flowering plant 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 Fig. 1 The scent space of sapromyiophilous stapeliads and aroids. Nonmetric multidimensional scaling (NMDS) analysis of the major scent compounds present in the odour profiles of 15 species of stapeliads (Apocynaceae subfamily Asclepiadoideae, tribe Ceropegieae, subtribe Stapeliinae; data from Jürgens et al., 2006), plus 11 species of Arum, 20 species of Amorphophallus and two species of Pseudodracontium (Araceae; data from Kite et al., 1998). NMDS was performed using a binary (presence-absence) matrix of the 48 plant species and the 54 most abundant (≥ 10% volume) compounds from their scent profiles, resulting in a two-dimensional plot using Euclidean distances between species. Unidentified compounds were omitted from the analysis. Several Amorphophallus species have identical oligosulphide odours and thus occupy the same loci in scent space. Chemical structures indicate scent components that contributed most to the separation of clusters in scent space (Jürgens et al., 2006), and are (clockwise from top): hexanoic acid, p-cresol, octanal, dimethyl disulphide and 2-heptanol (centre). Trimethyl amine (upper left) confers a dead-fish odour to Amorphophallus brachyphyllus (Kite & Hetterscheid, 1997), and is predicted to be present in stapeliads such as Huernia thuretii (not yet studied), which would expand the stapeliad domain in scent space.
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 http://www.bulbsociety.org/GALLERY_OF_THE_WOR LDS_BULBS/GRAPHICS/Arum/Arum_palaestinum/Ar um_palaestinum.html
Arum palaestinum
http://picsicio.us/keyword/arum%20palaestinum/ These are vinegar flies trapped in the calyx of the lily Arum palaestinum. Credit: Johannes Stökl, Curr. Biol., Oct. 7, 2010
Deceitful lily fools flies Solomon's lily imitates a yeasty odor to lure vinegar flies into a trap
Stökl, J., Strutz, A., Dafni, A., Svatos, A., Doubsky, J., Knaden, M., Sachse, S., Hansson, B.S., & Stensmyr, M.C. (2010) A Deceptive Pollination System Targeting Drosophilids through Olfactory Mimicry of Yeast. Current Biology, In Press, Corrected Proof, pp http://www.sciencedirect.com/science/article/B6VRT- 51618H5-5/2/4786042e3ee7f68188f5197d1a0d7016 AND http://www.ask-force.org/web/Pollination/Stoekl- Deceptive-Pollination-2010.pdf
-
2010.pdf - 51618H5 - - http://www.ask Pollination - Deceptive - Stökl, J., Strutz, A., Dafni, A., Svatos, A., Doubsky, M., J., Knaden, Doubsky, A., A., Svatos, A., Dafni, J., Strutz, Stökl, (2010) M.C. Stensmyr, & B.S., Hansson, S., Sachse, Olfactory through Drosophilids Targeting System Pollination Deceptive A Proof, In pp Corrected Press, Biology, Current Yeast. of Mimicry http://www.sciencedirect.com/science/article/B6VRT 5/2/4786042e3ee7f68188f5197d1a0d7016 AND force.org/web/Pollination/Stoekl
(C) Headspace odor of A. palaestinum (orange trace) and electro-antennograms (EADs) from females of D. melanogaster and D. simulans. The EAD traces shown are the averages of five runs, respectively. Numbers in the EAD trace refer to peaks that elicited antennal reactions in either of the species; blue numbers denote peaks yielding fully reproducible response: (1) acetoin; (2) 1-hexanol; (3) acetoin acetate; (4) 2,3- butanediolacetate (4i, threo; 4ii, erythro); (5) unknown; (6) 1-propionylethyl acetate; (7) ethyl hexanoate; (8) hexyl acetate; (9) 2,3- butanedioldiacetate; (10) 2-phenethyl alcohol; (11) 2-phenethyl acetate; and (12) geranylacetone. Flowers of Ceropegia arabica (Apocynaceae) are pollinated by small Diptera in common with all other members of the genus studied to date (Ollerton et al. 2009). Photograph by Sage Reynolds
Ollerton, J., Masinde, S., Meve, U., Picker, M., & Whittington, A. (2009) Fly pollination in Ceropegia (Apocynaceae: Asclepiadoideae): biogeographic and phylogenetic perspectives. Annals of Botany, 103, 9, pp 1501-1514 http://www.ask- force.org/web/Pollination/Ollerton- Fly-Pollination-Ceropegia-2009.pdf
Stapelia, Aasfliegen Stapelia, Aasfliegen Stapelia, Aasfliegen Rafflesia pricei, Sabah, Borneo Tambunan Rafflesia Centre in the Crocker Range National Park, Sabah, Malaysia http://lemberk.vcm.cz/cestopisy/rafflesia2.htm provincie Benkulu ve střední Sumatře a Západní Kalimantan (Indonésie), západní Sarawak (Malajsie) Rafflesia arnoldii Rafflesia arnoldii is a member of the genus Rafflesia. It is noted for producing the largest individual flower on earth. There are some plants with larger flowering organs, the Titan Arum and Talipot palm, but these are technically clusters of many flowers. It occurs only in the rainforests of Sumatra and Borneo in the Indonesia Archipelago.Molecular studies in 2007 have resulted in the final taxonomy of Rafflesia arnoldii which has been assigned to the family of Euphorbiaceae (Davis et al. 2007).DescriptionSeveral species of Rafflesia grow in the jungles of southeast Asia, many of them threatened or endangered. Rafflesia arnoldii is the largest; its flower attains a diameter of nearly a meter (3 ft) and can weigh up to 11 kilograms (24 lb).It lives as a parasite on the Tetrastigma vine, as its host, which grows only in primary (undisturbed) rainforests. While many parasites appear like normal plants, Rafflesia lacks any observable leaves, roots, or even stems. Likened to fungi, Rafflesia individuals grow as thread-like strands of tissue completely embedded within and in intimate contact with surrounding host cells from which nutrients and water are obtained. Perhaps the only part of Rafflesia that is identifiable as distinctly plant-like are the flowers; although, even these are bizarre because they attain massive proportions and are usually reddish-brown and stink of rotting flesh. The flower is pollinated by flies attracted by its scent.Specimen of Rafflesia arnoldii at the Kyoto Botanical Garden.EnlargeSpecimen of Rafflesia arnoldii at the Kyoto Botanical Garden.Rafflesia arnoldii is rare and fairly hard to locate. It is especially difficult to see in flower; the buds take many months to develop and the flower lasts for just a few days. How many of these strange plants still survive is unknown, but as the remaining primary forests of Borneo and Sumatra disappear, it can only be assumed that their numbers are dwindling. Many are known to be nearing extinction. Some environmentalists are thinking of a way to recreate the species’ environment, in an effort to stimulate a recovery in the population of this endangered species. This has proved unsuccessful so far, but the efforts have continued. However, steps are being taken to conserve the forests of Sumatra and Borneo.Pollination is a rare event due to several factors. The flowers are unisexual and single sites usually produce either male or female flowers. Therefore, in order to have effective pollination, male flowers must be near to, and open at the same time as, the female flowers so that flies can transfer the pollen.While male and female individuals could be closely spaced, flower bud mortality is 80-90% per site thereby reducing the chance of co-flowering two individuals. Furthermore, the flower lifespan is only 5-7 days. Also, the current population distributions are fragmented due to habitat destruction. So, successful reproduction relies on the unlikely event that a male and female would bloom during the same 5-7 days and that a fly could carry pollen between the often widely separated populations.
See also http://www.parasiticplants.siu.edu/Rafflesiaceae/Raff.arn.page.html
Video from BBC Archive: Flies pollinating Rafflesia http://www.arkive.org/rafflesia/rafflesia-spp/video-09.html 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 dayanum Mimikry: decaying animals attracting flyes feeding on cadavres Bulbophyllum sp. Aasfliegen Bulbophyllum sp. Aasfliegen 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 ? Ant- pollination Ant pollination systems are remarkably rare. We show that pollen exposed to ants for brief periods exhibits reduced viability, reduced percent germination, and shorter pollen tubes relative to control pollen. Pollination with ant-borne pollen also results in lower seed-set than pollination with untreated pollen. This disruption of pollination processes must have exerted a powerful selection pressure against the evolution of ant- pollination systems. It is suggested that the nest-building and brood-rearingh abits of ants require that they secrete large amounts of antibiotics to combat pathogenic microorganisms. It is these secretions that disrupt pollen function. Bees and wasps exhibit very different nesting behavior, consequently there are no chemical barriers to their coevolving with flowers as pollinators. Beattie, A.J., Turnbull, C., Knox, R.B., & Williams, E.G. (1984) ANT INHIBITION OF POLLEN FUNCTION - A POSSIBLE REASON WHY ANT POLLINATION IS RARE. American Journal of Botany, 71, 3, pp 421-426
Euphorbia (Chamaesyce) repens
http://blog.wildaboutants.com/2012/04/01/sandmat-and-ants-pollination/ Glaux maritima
Strand- Milchkraut
Ant – Pollination?
Possible Ant Pollination ? See also: Dubinsky, Z., Seckbach, J., Rostas, M., & Tautz, J. (2011) Ants as Pollinators of Plants and the Role of Floral Scents. In All Flesh Is Grass (eds Seckbach J. & Dubinsky .), Vol. 16, pp. 149-161. Springer Netherlands Cellular Origin, Life in Extreme Habitats and Astrobiology http://dx.doi.org/10.1007/978-90-481-9316-5_6 The ant Formica sericea and F. Schaufussii move from flower to flower seeking nectar from the bases of the antipetalous stamens.
Proven Ant Pollination
Wyatt, R. (1981) Ant-Pollination of the Granite Outcrop Endemic Diamorpha smallii (Crassulaceae). American Journal of Botany, 68, 9, pp 1212-1217 http://www.jstor.org/stable/2443043 AND http://www.ask-force.org/web/Pollination/Wyatt-Ant-Pollination-Diamorpha-Smallii-1981.pdf FIG. 1. Flowers and pollinators of Cytinus hypocistis: (A) Cytinus hypocistis, (B) Aphaenogaster senilis, (C) Crematogaster auberti, (D) Pheidole pallidula, (E) Tetramorium semilaeve and (F) Oplisa aterrima. Scale bars ¼ 5 mm. de Vega, C., Arista, M., Ortiz, P.L., Herrera, C.M., & Talavera, S. (2009) The ant-pollination system of Cytinus hypocistis (Cytinaceae), a Mediterranean root holoparasite. Annals of Botany, 103, 7, pp 1065-1075 http://aob.oxfordjournals.org/content/103/7/1065.abstract AND http://www.ask-force.org/web/Pollination/deVega-Ant-Pollinatioin-System- Cytinus-2009.pdf de Vega, C., Arista, M., Ortiz, P.L., Herrera, C.M., & Talavera, S. (2009) The ant-pollination system of Cytinus hypocistis (Cytinaceae), a Mediterranean root holoparasite. Annals of Botany, 103, 7, pp 1065-1075 http://aob.oxfordjournals.org/content/103/7/1065.abstract AND http://www.ask-force.org/web/Pollination/deVega-Ant-Pollinatioin-System- Cytinus-2009.pdf
Fig. 3.Time spent by different pollinators on female and male flowers of Cytinus hypocistis during the day. Note that for each species the y- axis differs, and the x-axis is also different for the nocturnal Camponotus pilicornis. Aphaenogaster senilis spent more time on male flowers (χ2 = 11·83; P = 0·0006), whereas P. schmitzii (χ2 = 22·01; P < 0·0001) and T. nigerrima (χ2 = 22·01; P < 0·0001) spent more time on female flowers. In the other species no differences were found. All species are ants except those marked with an asterisk.
de Vega, C., Arista, M., Ortiz, P.L., Herrera, C.M., & Talavera, S. (2009) The ant-pollination system of Cytinus hypocistis (Cytinaceae), a Mediterranean root holoparasite. Annals of Botany, 103, 7, pp 1065-1075 http://aob.oxfordjournals.org/content/103/7/1065.abstract AND http://www.ask-force.org/web/Pollination/deVega-Ant-Pollinatioin-System- Cytinus-2009.pdf
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