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(1)

Bestäuberverhalten und Pflanzenevolution

“aktuelle Stunde”

Populations und Evolutionsbiologie, 4.2.04

Florian Schiestl

(2)

Orchid pollination rewarding species

Gymnadenia Platanthera Thelymitra Calochilus

deceptive species

(3)

Important features of pollination by sexual deception:

• Floral odour is most important for attracting the pollinator – sex pheromone mimicry.

• Pollination is highly specific.

(4)

Basic concept of (Batesian) mimicry

Wickler (1968)

Model

Mimic (imitates model-signals)

Operator (dupe)

(5)

Questions:

1) Mechanisms of pollinator attraction

2) Evolutionary dynamics between orchids and

pollinators: impact of orchids on pollinators

3) Odour communication and speciation

(6)

Methods: Gas chromatography with

electroantennograpic detection (GC-EAD); gas

chromatography – mass spectrometry (GC-MS)

(7)

Behavioural tests with pollinator-insects

(8)

1) Mechanisms of pollinator attraction

• Sex pheromone of the pollinator species

• Attractive orchid odour

2) Evolutionary dynamics

3) Speciation

(9)

Where is the female sex pheromone of N.

cryptoides produced?

10 2

10 10

12 17

6 N =

Orchidextr.

Poisongl.

v.d.Vecht's gl.

Dufour's gl.

Cuticle extr.

Headextr.

Control

Mean(+-1 SE) mountingsof males

4

3

2

1

0

-1

*

*

(10)

Wasp,

headextract

Orchid,

labella extracts

Which are the electrophysiologically active compounds in the odour samples?

10 15 20 25 Time (min)

FID EAD

FID EAD

(11)

Compound identification

2-ethyl-5-propyl-1,3-cyclohexandione – „Chiloglottone“

O O

(12)

Chiloglottis - bioassay: attractiveness of Chiloglottone

29 10

N =

Chiloglottone

Orchidextract

Mean (+-1SE) copulation attempts 4,0

3,5 3,0 2,5 2,0 1,5 1,0 ,5 ,0

10

Odourless dummy

*

*

(13)

Summary 1

• In the thynnine wasp N. cryptoides, the female sex pheromone consists of only one (!) compound, which is produced in the head.

• The orchid Chiloglottis trapeziformis produces the

same compound in its floral odour.

(14)

Europe: Ophrys, pollinated by solitary bees

• O. sphegodes: Flowers attract pollinator with 14 compounds (alkanes and alkenes).

• Female bees produce the same compounds, in similar proportions, as sex pheromone.

(Schiestl, Ayasse et al. 1999)

(15)

1) Mechanisms of pollinator attraction 2) Evolutionary dynamics - impact

• How pollinators avoid sexually deceptive orchids.

• What are the impacts for orchid and pollinator fitness?

3) Speciation

(16)

Orchids (presented sequentially in the same spot)

Mountingsof males

8

6

4

2

0

11 10

9 8

7 6

5 4

3 2

1

Wasps do not avoid individual flowers but locations with

flowers

(17)

• Are pollinator-males able to

discriminate between their females and orchid flowers?

• Are males able to find their wingless

females in the presence of orchids?

(18)

Can males discriminate between sex pheromone and

orchid floral odour?

(19)

8 8

N =

Orchid Female

Mean(+-1 SE) mountings

4,0

3,5

3,0

2,5

2,0

1,5

1,0

Can males discriminate between sex pheromone and

orchid floral odour?

(20)

Calling N. cryptoides female

C. trapeziformis flowers

50m 3m

30cm

(21)

9 9

9 9

9 9

N =

+ 10 orchids + 5 orchids

No orchids + 10 orchids

+ 5 orchids No orchids

Mean(+-1 SE) male reactions

12

10

8

6

4

2

0

-2

Approaches Mountings

*

* * *

Do males pick up females that call in an orchid patch?

(22)

selection on increased discrimination in operator refinement of signals in mimic;

Arms race!

(23)

• Europe: pollinators avoid individual flowers;

Australia: pollinators do not avoid individual

orchids, but the areas where orchids flower; they are unable to discriminate between the odour of orchid flowers and their females.

• Orchids may harm their pollinators, since males are not able to find their females within an orchid patch.

Arms race scenario.

Summary 2

(24)

1) Mechanisms of pollinator attraction 2) Evolutionary dynamics - impact

3) Odour communication and speciation

• how is evolutionary change imaginable?

(25)

Important features for speciation:

• Species are interfertile: specificity in

pollination is responsible for reproductive isolation.

• Odour is the key stimulus for specific

pollinator attraction: it is directly linked to

reproductive success and isolation.

(26)

Chiloglottis trapeziformis

Chiloglottis valida

Tow allopatric (with non-overlapping distribution)

orchids with the same pollinator species:

(27)

GC-EAD recordings using floral odour of two allopatric orchid species with one pollinator (Neozeleboria

cryptoides)

C. trapeziformis

C. valida

10 15 20 25 Time (min)

(28)

Two closely related, sympatric (overlapping distribution) orchids with distinct pollinator species

Chiloglottis trilabra Chiloglottis reflexa

(29)

GC-EAD recordings with floral odour of the two sympatric orchid spp. with two pollinators

C. trilabra N. proxima

C. reflexa N. sp. 30

C. reflexa N. proxima

10 15 Time (min)

B,C

A,B

B

(30)

Summary 3

• Allopatric orchids with the same pollinator species may emit the same attractive odour.

• Sympatric orchids with different pollinators may share active compounds but produce distinct compounds as well.

• Small changes in odour emission may trigger

speciation – radiation in orchids may have been a quick process via “stepwise acquiring of new

pollinators”.

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