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Species-specific attraction to pheromonal analogues in orchid bees

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Abstract

Male orchid bees (Euglossini) collect fragrances from flowers and other natural sources, a behaviour that has shaped the euglossine pollination syndrome. Males store such chemicals in hind leg pouches and later expose them during courtship display. In the present study, we show that complex bouquets of two sympatric species of Eulaema, E. meriana and E. bombiformis, are chemically distinct. When exposed during bioassays at display perches individual hind leg extracts rapidly and consistently attracted other males of the correct species, even if derived from males of disparate localities (French Guiana and Panama). Conspecific males as well as females of E. bombiformis arrived at natural perch sites only from downwind, and two copulations were observed. Our findings demonstrate that acquired odours mediate exclusive attraction within species and support the idea that such fragrances are pheromone analogues. Their role in acquiring matings and during male–male interaction is discussed.

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Acknowledgements

We thank Oris Acevedo and the entire BCI staff for continuous support during field seasons. The chemical-analytical part of the study was substantiated by Roman Kaiser who analysed reference samples and identified critical fragrance compounds. His help is gratefully acknowledged, as are the valuable discussions with Klaus Lunau and the members of the Sensory Ecology seminar in Düsseldorf. The study was sponsored by the Deutsche Forschungsgemeinschaft (EL 249/2-1)

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Corresponding author

Correspondence to Thomas Eltz.

Additional information

Communicated by R.F.A. Moritz

Appendix

Appendix

Incidence and relative abundance of chemical compounds detected in hexane extracts of hind legs of Eulaema meriana and E. bombiformis. Abundances are the average of a given compound’s (peak) contribution to the individual GC/MS ion chromatogram (in % of the total ion current). Compounds are indexed as exogenous fragrances (F), labial gland lipids also found in head extracts (H), and other high molecular weight non-fragrance compounds (NF).

RT (min)

Compound

Number of individuals with compound

Average rel. abundance (% of total ion current)

Class

E. bombiformis N=7

E. meriana N=41

Both N=48

E. bombiformis

E. meriana

5.32

Thujene, alpha-

F

0

4

4

0.00

0.01

5.52

Pinene, alpha-

F

0

14

14

0.00

0.04

6.24

Benzaldehyde

F

0

8

8

0.00

0.07

6.56

Sabinene

F

0

34

34

0.00

0.73

6.83

Pinene, beta-

F

0

11

11

0.00

0.05

7.07

Myrcene

F

4

4

8

0.11

0.01

7.47

Menthene, 3-para- ?

F

0

6

6

0.00

0.02

7.75

cyclic monoterpene

F

0

2

2

0.00

0.01

8.02

Terpinene, alpha-

F

0

2

2

0.00

0.00

8.29

Cymene, para- (ortho- ?)

F

0

27

27

0.00

0.32

8.46

Limonene

F

0

10

10

0.00

0.04

8.51

Phellandrene, beta

F

0

3

3

0.00

0.01

8.54

Benzyl alcohol

F

5

7

12

0.50

0.51

8.58

Cineole, 1,8-

F

0

27

27

0.00

0.59

8.66

Ocimene, (Z)-beta-

F

0

2

2

0.00

0.02

9.06

Ocimene, (E)-beta-

F

7

0

7

1.55

0.00

9.50

Terpinene, gamma-

F

0

22

22

0.00

0.23

9.93

Sabinene hydrate, cis- (rel. to OH)

F

0

21

21

0.00

0.68

9.99

?

F

0

17

17

0.00

0.84

10.53

Terpinolene

F

0

7

7

0.00

0.03

11.10

Sabinene hydrate, trans-?

F

0

12

12

0.00

0.05

11.13

?

F

1

0

1

0.04

0.00

11.15

?

F

0

13

13

0.00

0.07

11.35

?

F

0

11

11

0.00

0.02

11.58

Phenylethyl alcohol, 2-

F

5

0

5

2.98

0.00

12.71

Limonene oxide, trans- ?

F

1

15

16

0.03

0.32

12.86

Ipsdienol

F

7

0

7

3.65

0.00

13.71

Benzyl acetate

F

5

8

13

2.74

0.99

14.58

Terpinen-4-ol

F

0

40

40

0.00

4.80

14.95

Methyl salicylate

F

0

22

22

0.00

0.27

15.17

Terpineol, alpha-

F

0

2

2

0.00

0.01

15.36

Dihydro carvone, cis- ?

F

0

1

1

0.00

0.00

15.60

Dihydro carvone, trans-

F

0

9

9

0.00

0.08

16.25

aromatic compound

F

0

1

1

0.00

0.01

16.93

?

F

1

0

1

0.18

0.00

17.31

Carvone

F

3

27

30

0.27

0.73

17.65

?

F

1

0

1

0.22

0.00

17.71

Phenylethyl acetate, 2-

F

4

0

4

4.27

0.00

17.90

?

F

1

0

1

2.44

0.00

18.00

Carvone epoxide, cis-

F

5

38

4

3.47

12.92

18.49

Geranial

F

1

0

1

0.09

0.00

18.82

Carvone oxide, trans-

F

0

1

1

0.00

0.00

18.97

?

F

0

8

8

0.00

0.12

19.35

Indole

F

3

0

3

0.22

0.00

20.12

?

F

0

5

5

0.00

0.01

20.99

?

F

2

4

6

0.09

0.04

21.16

trans-(trans-Carveol) epoxide?

F

0

12

12

0.00

0.23

21.26

?

F

0

1

1

0.00

0.01

21.47

?

F

0

1

1

0.00

0.00

21.58

?

F

0

1

1

0.00

0.00

21.76

?

F

1

0

1

0.04

0.00

22.22

Eugenol

F

0

1

1

0.00

0.00

23.06

?

F

7

0

7

0.64

0.00

23.21

Copaene, alpha-

F

1

0

1

0.05

0.00

23.27

Geranyl acetate

F

2

0

2

0.11

0.00

23.58

Methyl cinnatamte, (E)-

F

0

8

8

0.00

0.07

23.74

Cubebene, beta-

F

0

1

1

0.00

0.01

23.75

?

F

0

2

2

0.00

0.04

23.89

?

F

0

3

3

0.00

0.04

24.05

?

F

0

2

2

0.00

0.01

24.75

?

F

0

7

7

0.00

0.03

24.95

?

F

0

5

5

0.00

0.04

24.96

?

F

0

1

1

0.00

0.01

24.99

Caryophyllene, (Z)-

F

0

1

1

0.00

0.01

25.02

?

F

0

2

2

0.00

0.02

25.67

Bergamotene, (E-)-alpha-

F

0

2

2

0.00

0.02

25.71

?

F

1

0

1

0.04

0.00

25.72

?

F

1

4

5

0.05

0.02

26.52

?

F

0

1

1

0.00

0.00

26.55

Farnesene, (E)-beta-

F

0

1

1

0.00

0.01

26.61

?

F

1

0

1

0.08

0.00

26.70

?

F

0

1

1

0.00

0.01

27.51

?

F

0

1

1

0.00

0.00

27 53

Dodecanol

H

0

1

1

0 00

0 00

27.72

Germacrene D

F

1

3

4

0.25

0.11

27.83

aff. Farnesene

F

0

1

1

0.00

0.01

28.25

Bicyclogermacrene ?

F

0

2

2

0.00

0.07

28.48

Bulnesene, alpha- ?

F

2

0

2

0.07

0.00

28.55

?

F

0

1

1

0.00

0.01

28.78

Farnesene

F

0

4

4

0.00

0.02

29.00

?

F

0

1

1

0.00

0.01

30.39

?

F

7

20

27

2.86

0.23

31.10

nerolidol, (E)-

F

0

1

1

0.00

0.00

31.65

Spathulenol ?

F

0

1

1

0.00

0.00

32.90

Dodecyl acetate

H

2

12

14

0.19

0.21

33.25

Farnesene epoxide, (E)-alpha-

F

0

6

6

0.00

0.03

33.28

?

F

0

1

1

0.00

0.01

33.31

?

F

0

3

3

0.00

0.04

33.31

Tetradecanal (=Myristylaldehyde)

H

1

5

6

0.35

0.01

34.38

?

F

7

0

7

2.94

0.00

34.56

Methoxynaphtol

F

7

0

7

1.55

0.00

35.28

2-Propenoic acid, 3-[4-Methoxyphenyl]-

F

0

3

3

0.00

0.01

35.65

Tetradecanol

H

1

1

2

0.06

0.00

35.98

?

F

0

2

2

0.00

0.01

36.48

?

F

0

1

1

0.00

0.00

36.68

Bergamotol, (Z)-alpha-, trans-

F

0

15

15

0.00

0.13

38.87

Benzyl benzoate

F

0

6

6

0.00

0.04

38.89

?

F

0

2

2

0.00

0.01

39.24

Tetradecanoic acid

H

0

1

1

0.00

0.00

40.62

Tetradecyl acetate

H

3

19

2

6.66

1.16

41.08

Hexadecanal

F

1

0

1

0.15

0.00

41.50

?

F

0

1

1

0.00

0.01

41.67

Hexahydrofarnesyl acetone

F

0

10

10

0.00

0.04

42.70

Benzyl salicylate

F

0

1

1

0.00

0.00

43.21

Hexadecanol

H

1

4

5

0.10

0.02

43.41

?

F

0

1

1

0.00

0.00

45.10

Cyclohexadecanolid ?

NF

0

1

1

0.00

0.01

45.70

aromatic compound

F

0

1

1

0.00

0.00

45.97

Hexadecanoic acid?

H

0

2

2

0.00

0.03

47.47

Hexadecyl acetate

H

3

22

25

4.10

1.84

47.53

?

NF

0

2

2

0.00

0.01

47.86

?

NF

2

5

7

0.06

0.02

47.91

?

F

0

6

6

0.00

0.03

48.36

?

F

0

1

1

0.00

0.04

48.37

Falcarinol, (Z)-

F

0

3

3

0.00

0.05

49.38

?

H

0

2

2

0.00

0.02

49.50

?

H

1

36

37

0.02

0.46

49.91

?

H

1

2

3

0.02

0.01

50.44

Heneicosane

H

1

1

2

0.34

0.02

50.62

?

F

0

1

1

0.00

0.03

51.82

?

H

1

6

7

0.50

0.37

53.06

?

F

1

1

2

0.24

0.03

53.76

Octadecanol acetate

H

3

8

11

1.07

0.48

55.58

Tricosene

H

6

40

46

23.83

20.49

55.59

?

H

1

2

3

7.75

1.00

56.47

Tricosane

H

6

37

43

2.71

2.62

58.86

Octadecanoic acid, butyl ester ?

NF

1

1

2

0.20

0.03

61.12

?

NF

0

1

1

0.00

0.04

61.20

Pentacosene

H

1

34

35

0.17

2.28

62.06

Pentacosane

H

7

39

46

3.75

3.16

64.05

?

NF

0

7

7

0.00

0.11

65.96

?

H

1

12

13

0.89

0.42

66.45

?

H

0

1

1

0.00

0.25

66.56

?

H

3

40

43

3.97

6.23

66.91

?

H

1

0

1

0.19

0.00

67.21

Heptacosane

H

2

14

16

0.62

0.36

69.07

?

H

1

9

10

0.18

0.18

70.77

Nonacosadiene ?

H

1

39

40

1.51

6.20

71.03

?

NF

1

10

11

0.08

0.19

71.19

?

H

5

40

45

6.84

10.06

71.95

Nonacosane

NF

2

14

16

0.28

0.30

72.64

?

NF

0

2

2

0.00

0.04

73.19

?

NF

0

3

3

0.00

0.03

74.89

?

H

1

38

39

0.43

4.30

75.38

Untriacontadien ?

NF

0

3

3

0.00

0.07

75.75

?

H

3

19

22

1.09

1.29

76.25

Hentriacontane

H

1

7

8

0.10

0.13

79.49

Tritriacontadiene ?

NF

0

2

2

0.00

0.12

79.51

?

NF

0

14

14

0.00

0.92

80.67

Phytosterole ?

NF

0

6

6

0.00

0.18

81.31

similar beta-Amyrin

NF

0

32

32

0.00

2.50

81.59

similar alpha-Amyrin

NF

0

3

3

0.00

0.17

81.62

Phytosterole

NF

0

13

13

0.00

0.61

82 39

similar alpha-Amyrin

NF

0

38

38

0 00

4 84

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Zimmermann, Y., Roubik, D.W. & Eltz, T. Species-specific attraction to pheromonal analogues in orchid bees. Behav Ecol Sociobiol 60, 833–843 (2006). https://doi.org/10.1007/s00265-006-0227-8

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