Bestäubungsökologie: Grundlage der Evolution und Systematik der

Bestäubungsökologie: Grundlage der Evolution und Systematik
der Blütenpflanzen
Prof. em. Klaus Ammann, Neuchâtel
klaus.ammann@ips.unibe.ch
18. März 2013, Schaffhausen
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
Parnassia palustris, Saxifragaceae
attractive pseudo-glands offered, no nectar
Nektarsuche Hedera
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
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 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.askforce.org/web/Pollination/MayerPollination-Ecology-21fst-CenturyFuture-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
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-InsectVibrational-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
www.anu.edu.au/BoZo/ orchid_pollination/
The sexually attracted male
pollinator Neozeleboria
cryptoides on the stem of
the orchid Chiloglottis
trapeziformis
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
centuncularis
Dalechampia
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://darwinonline.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 of pollination 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
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, <Go to
ISI>://WOS:000305278400012 AND
http://www.ask-force.org/web/Pollination/Ward-Modes-Reproduction-Invaisve-2012.pdf
Tibouchina urvilleana, Melastomataceae
Tibouchina urvilleana
Tibouchina urvilleana
Tibouchina urvilleana
Bumblebee
pollination
http://mgonline.com/media/Images/b/butterfly_garden005.jpg
Tibouchina
urvilleana
Lamium orvala, Karawanken, pollination with bumble 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 Verkehrszeichen
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 nearestneighbour rules for optimization of multi-location routes Biology Letters, 8, 1, pp. 13-16, <Go
to ISI>://WOS:000299322200005 AND
http://www.ask-force.org/web/Pollination/Lihoreau-Bees-not-use-nearest-2013.pdf
6.2. butterfly
pollination
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
Lilium bulbiferum
Feuerlilie
Liliaceae
Tagfalter
weiss: Nektar-Röhren
Gentiana 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 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, <Go to ISI>://WOS:000314756900007 AND http://www.askforce.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: Inflorescence and
flower morphology. Note the prominent
style (arrow) among the stamens. 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 selfpollinated 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, <Go to
ISI>://WOS:000314756900007 AND
http://www.askforce.org/web/Pollination/AmorimBeyond-Pollination-Syndrome-NectarEcology-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, <Go to
ISI>://WOS:000302992000009 AND
http://www.askforce.org/web/Pollination/SoundersDiversity-Evolution-PollinationAnnonaceae-2012.pdf
6.5. marsupianpollination
Banksia attenuata
Proteaceae
grobe Bürsten-Blütenstände, robuste
Staubblätter und Narben
mit Beutelmäusen
Banksia integrifolia, Proteaceae, West-Australien
6.8. batpollination
Solanaceae
Cobaea sp.
Underwoods
longtongued
Bat
Hylonycteris
underwoodii
Carpobrothus edulis, Mesembryanthemaceae, Fledermäuse
Solandra nitida, Solanaceae, Fledermäuse
Thunbergia japonica
Acanthaceae
Grosse Hautflügler
Fledermäuse, Vögel
robuste Trompetenform
6.5. birdPollination
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. beetlepollination
Gottsberger, G., I.
SilberbauerGottsberger, 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. 107118, <Go to
ISI>://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
Veronica filiformis, Faden-Ehrenpreis, Scrophulariaceae, Fliegen
Veronica cymbalaria, Fliegenbestäubung
Euphrasia 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 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
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-RafflesiaGigantism-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
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. 40084013,
http://www.pnas.org/content/109/10/4008.abstra
ct AND http://www.askforce.org/web/Evolution/Yashina-Regenerationwhole-plant-Silene-Stenophylla-2012.pdf
30’000 jährige
Silene-Art
aus einer sibirischen
Höhle
wieder zum Leben
erweckta