The adaptive value of web decorations for (Araneae, Araneidae)

Dissertation (kumulativ) Contents

Chapter 1 3

Chapter 2 11

Chapter 3 Argiope bruennichi 22

Chapter 4 Argiope 32 Chapter 5 47

Chapter 6 62

Chapter 7 Argiope bruennichi 74 Chapter 8 85

Chapter 9 90

Literature cited 96

Appendix 106

2

General introduction

3

4

Web attachments

Uloborus Cyclosa Gea Argiope

Silken web decorations Argiope 5

Argiope Argiope Argiope A. lobata A. sector A. ocyaloides

6

Adaptive significance of web decorations in Argiope Argiope Argiope

Argiope

7

Argiope Study structure and aims Argiope

8

Argiope bruennichi A. bruennichi Argiope Argiope Argiope A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. keyserlingi Argiope A. keyserlingi

9

Argiope Argiope A. bruennichi Mantis religiosa Argiope A. bruennichi A. bruennichi

10

Web decorating behaviour in Argiope bruennichi (Araneae, Araneidae): Is short-term variation an indication of a conditional strategy?

Annales Zoologici Fennici

11

Abstract. Argiope Argiope A. bruennichi

12

Introduction . Argiope Argiope Argiope . . .

13

A. bruennichi A. bruennichi A. bruennichi Methods Study site and study A. bruennichi

14

Statistical analyses Results shapes A. bruennichi, n n n

15

² n² n t p

B 32,5% A 39,2% C 1,5%

0 26,9% 0AB C Abruennichi Variation of web decorating behaviour n

16

² χ² p A. bruennichi Sequence Sequence Individuals No. 1 A A A A 2 2 A A B A 1 3 A A B B 4 4 A A B 0 4 5 A A 0 0 2 6 A B A A 1 7 A B A C 1 8 A B B B 1 9 A C 0 B 1 10 B A A A 3 11 B A A B 2 12 B B B B 1 13 B 0 B 0 1 14 B 0 0 B 1 15 B 0 0 0 4 16 0 A B B 2 17 0 B 0 0 1 18 0 0 B 0 1 19 0 0 0 0 1

17

n n n

100 80

60 n = 14 % n = 13 40 n = 4 20 n = 3 0 0 1 2 3 Changes A. bruennichi Web-site relocation and web design n n n n ² χ² p

18

² χ² p 100

80 60 % 40 20 0 web relocations (n=22) without web relocation (n=80) remained web decoration pattern changed web decoration pattern

A. bruennichi Discussion A. bruennichi Argiope .

19

. . A. argentata A. bruennichi Argiope A. trifasciata A. aurantia A. bruennichi Argiope

20

A. bruennichi A. bruennichi Argiope

21

The Argiope bruennichi (Arachnida, Araneidae): is not an obligate life history phase.

Journal of

22

Abstract. Argiope bruennichi A. bruennichi

23

Introduction Argiope Argiope Argiope bruennichi A. bruennichi A. bruennichi

24

A. bruennichi A. bruennichi Argiope A. bruennichi A. trifasciata A. aurantia Argiope Material & methods A. bruennichi n ±

25

n ± ±

26

Drosophila melanogaster n Results ± n ± n p ± ± n n p

27

± n ± n p

A. bruennichi

A. bruennichi

28

A. bruennichi Discussion A. trifasciata A. aurantia Argiope bruennichi

29

A. bruennichi A. bruennichi A. trifasciata A. aurantia A. bruennichi Drosophila melanogaster A. bruennichi Argiope

30

Argiope A. aurantia A. bruennichi Argiope

31

‘Wrap attack’ activates web decorating behavior in Argiope spiders.

Behavioral

32

Abstract. Argiope A. bruennichi A. sector A. keyserlingi Argiope Argiope Argiope

33

Introduction

34

Argiope Argiope Argiope glandulae aciniformes Argiope Argiope

35

Argiope aetheroides Argiope

36

A. bruennichi A. sector A. keyserlingi A. aetheroides Material & methods Study species and experimental design A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. bruennichi A. sector A. keyserlingi A. bruennichi A. sector A. keyserlingi

37

Argiope A. bruennichi A. sector A. keyserlingi Apis mellifera A. bruennichi n Acheta domesticus A. sector n Lucilia A. keyserlingi n Argiope

38

Statistical analyses Results t p A. bruennichi t p A. sector t p A. keyserlingi A. bruennichi n A. sector n A. keyserlingi n

39

A. bruennichi t p A. keyserlingi t p A. bruennichi ² ² n² ² n A. keyserlingi ² ² n²² n A. sector ² ² n² ² n t p A. bruennichi n n A. sector n n A. keyserlingi n n ² A. bruennichi χ² P A. sector χ² P A. keyserlingi χ² P A. bruennichi ² ²

40

t p A. sector ² ² t p A. keyserlingi ²² t p

41

Argiope A. bruennichi χ² p A. sector χ² p A. keyserlingi χ² p A. keyserlingi t p A. bruennichi t p A. sector t p

42

Discussion Argiope A. keyserlingi A. bruennichi Argiope A. trifasciata A. bruennichi A. sector A. keyserlingi

43

Argiope A. sector A. appensa A. bruennichi A. sector, Argiope Argiope cavaticus

44

Argiope A. keyserlingi Argiope Argiope vice versa

45

Argiope Argiope Argiope Argiope

46

Moulting interferes with web decorating behaviour in Argiope keyserlingi (Araneae: Araneidae).

Journal of Arachnology

47

Abstract. Argiope Argiope keyserlingi A. keyserlingi

48

Introduction Argiope Argiope Octonoba sybotides Argiope

49

per se Argiope Materials & methods Study species and experimental design

Argiope keyserlingi

50

Argiope keyserlingi Lucilia

51

a v d

h

dh c A. keyserlingi

Statistical analyses ² Results Web decorating frequency A. keyserlingi

52

n Web size ² n² n ² ²² ² t p ² n² n r² p r² p n Variation of web decorating behavior A. keyserlingi n n n n n n n n

53

n²χ² p n ² χ² p n t p Number of web decoration bands (decorations, n=433)

24%

2

65% 1

3 6% 4 5% A. keyserlingi

54

Web decorating behavior in context of molting events ² ² n ² n t p ² n ² n ² n F p

55

A. keyserlingi A. keyserlingi

56

A Web decoration size - 12 10 8

6 4

(=100%) interval-output day 2 0

21 referred to the in perccent intensity Decorating -10-9 -8 -7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5 6 7 8 9 10 Day B Web building frequency

12

10

8

% 6

4

2

0 -10-9 -8 -7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5 6 7 8 9 10 Day

A. keyserlingi dotted line

57

Discussion A. keyserlingi A. keyserlingi Argiope Argiope Argiope A. aurantia A. trifasciata A. appensa A. versicolor A. picta A. aetherea Argiope keyserlingi

58

Argiope A. argentata A. savignyi A. flavipalpis A. keyserlingi

A. keyserlingi

59

Nephila clavipes Argiope Argiope N. clavipes

60

Argiope N. clavipes N. clavipes A. aurantia Araneus cavaticus Argiope Argiope

61

Argiope

62

Are web stabilimenta attractive to praying mantids?

Revista Ibérica de Aracnología (Iberian Journal of Arachnology)

63

Abstract. Argiope Archimantis latistylus Argiope keyserlingi Argiope A. keyserlingi A. bruennichi Mantis religiosa M. religiosa Mantis

64

Introduction Argiope Argiope Argiope

65

Archimantis latistylus Argiope keyserlingi A. keyserlingi Argiope

66

Mantis religiosa Argiope bruennichi Material & methods Study and rearing Mantis religiosa Drosophila melanogaster A. bruennichi A. bruennichi Mantis religiosa

67

The Y-maze apparatus Mantis

68

The test scheme M. religiosa

69

Results n n n n ² χ² p p

70

100

80

60 decorated undecorated webs webs 40 Individuals[%] 20

0 "Non- "Approachers" approachers"

M. religiosa n n n A. bruennichi Discussion Argiope bruennichi Mantis religiosa . Argiope keyserlingi Archimantis latistylus Archimantis

71

M. religiosa Archimantis

72

Argiope Mantis A. keyserlingi Mantis Conclusions

73

Argiope Archimantis latistylus

74

Argiope bruennichi shows a drinking like behaviour in web hub decorations (Araneae, Araneidae).

Journal of Ethology

75

Abstract Argiope Argiope bruennichi A. bruennichi A. bruennichi

76

Introduction

Argiope

77

Araneus diadematus Argiope A. trifasciata Argiope A. bruennichi

78

Material & methods Argiope bruennichi A. bruennichi A. bruennichi A. bruennichi A. bruennichi

79

Results Argiope bruennichi . n n n

80

n n p n

81

A. bruennichi

82

37 10 10 Duration 10 34 10 10 10 31 9 9 9 28 9 9 9 25 8 8 8 22 8 8 7 19 7 7 7 16 7 6

Observations 6 13 6 6 5 10 5 5 4 7 4 4 3 4 3 2 2 1 1 0 1 2 3 4 5 6 7 8 9 1011121314 Time [min]

A. bruennichi Discussion Argiope bruennichi A. bruennichi Argiope

83

A. bruennichi Argiope A. bruennichi A. diadematus Cyrtophora moluccensis Frontinella pyramitela A. bruennichi Argiope

84

Argiope trifasciata Argiope Argiope A. bruennichi Argiope

85

Synthesis

86

Argiope Argiope Argiope

87

Argiope A. bruennichi A. keyserlingi A. bruennichi A. bruennichi

88

A. keyserlingi Archimantis latistylus Mantis religiosa A. bruennichi A. bruennichi Conclusion Argiope Argiope

89

Argiope Argiope Argiope

90

Zusammenfassung

91

Argiope Argiope Argiope Argiope A. keyserlingi

92

Argiope A. bruennichi A. keyserlingi A. bruennichi

93

A. bruennichi A. keyserlingi Archimantis latistylus A. bruennichi Mantis religiosa M. religiosa A. bruennichi Argiope

94

A. bruennichi A. bruennichi Argiope Argiope Argiope

95

Argiope

96

Literature cited

Argyope bruennichi Entomologische Nachrichten und Berichte 30 Argiope bruennichi J Ethol 24 Argiope bruennichi Zool Anz 210 Cyrtophora moluccensis J Arachnol15 Micranthena gracilis Ecology 62 Naturwissenschaften 91 Proc Roy Soc Lond B 265 Argiope aurantia Argiope trifasciata J Zool (Lond) 246 Behav Ecol. 10 Behaviour 138 In Ecophysiology of Spiders Proc Roy Soc Lond B 270 The Comity of Spiders J Zool 269 J Evol Biol 14 Biol Letters 1 Argiope J Nat Hist 39 Naturwissenschaften 88 Argiope keyserlingi Anim Behav 60 Am Nat 110 Behav Ecol 18 97

The spider book In Extinction Rates Tenodera aridifolia sinensis J Exp Biol 148 J Arachnol 11 Funct Ecol 5 Anim Behav 47 Spiderwebs and Ecology 71 Argiope argentata Evolution 55 Argyope bruennichi Mitteilungen aus dem Zoologischen Museum in Berlin 37 J Exp Biol 29 J Arachnol 13 In Ecophysiology of Spiders Pirata sedentarius J Arachnol 31 Argiope bruennichi Arachnol Mitt 20 Uloborus diversus J Zool (Lond) 171 J Arachnol 15 J Arachnol 16 An Rev Ecol Sys 21 Allocyclosa bifurca Behaviour 140 Proc Biol Sci 248

98

Argiope flavipalpis Argiope trifasciata In Proc. 9 th Int. Congr. Arachnol., Panama 1983 Water balance in land Science 219 Argiope aetherea Austr J Ecol 21 Argiope keyserlingi Proc Roy Soc Lond B 267 Argiope aurantia Am Nat 90 Ecology 55 Argiope Am Nat 94 Argiope flavipes J Nat Hist 6 Biology of spiders Argiope bruennichi Argiope bruennichi In Proceedings of the 15 th European Colloquium of Arachnology, České Budějovice 1994 Argyope bruennichi Mitteilungen der Naturhistorischen Gesellschaft Nürnberg 2 Arch Naturgesch 90 Argiope bruennichi Verhandlungen des naturwissenschaftlichen Vereins Hamburg (NF) 23 Cyclosa fililineata Cyclosa morretes Ethology 111 J Exp Biol 202 Zool J Linn Soc 123

99

Argiope bruennichi Bonner zoologische Beiträge 30 Ethol Ecol Evol10 Anim Cogn 2 Argiope bruennichi De Levende Natuur 84 Argiope keyserlingi Austr J Zool 48 Biol Rev 75 Argiope keyserlingi Evol Ecol Res 2 J Arachnol 28 Argiope keyserlingi Austral Ecology 28 Ethology 111 Nephila clavipes J Behav 3 Funct Ecol 10 Nephila clavipes Physiol Entomol 24 Nephila clavipes Funct Ecol 15 Proc Zool Soc Lond 28 Am Nat 149 Psyche 87 Neogea Bull Brit Arachnol Soc 9 Mem Qd Mus 33 In The praying mantids

100

Ecology 76 Phil Trans Roy Soc B 352 Paratenodera angustipennis Oecologia 56 J Arachnol 2 Argiope appensa J Zool 229 Behav Ecol Sociobiol 10 Argiope bruennichi Entomologisk Tidskrift 120 Argiope appensa Pacific Sci 47 Argiope bruennichi Zoologisches Jahrbuch für Systematik 114 Mantis religiosa Empusa fasciata J Insect Behav 12 Gea Argiope Bull Mus Comp Zool Harv 136 Argiope Gea Neogea Bull Mus Comp Zool Harv 150 J Arachnol 30 Proc Biol Sci 272 Behaviour 140 Anim Behav 67 Anim Behav 68 Natur und Landschaft 54 Argiope argentata J Arachnol 2 In Spiders: webs, behavior, and evolution

101

Ecology 74 Z vgl Physiol 56 Argiope bruennichi Zool Jb Syst 117 Bull Brit Arachnol Soc 1 Tenodera angustipennis In The praying mantids American spiders and their spinning work Ecol Entomol 4 Oecologia 98 Argiope bruennichi Mem Qd Mus 33 Cyclosa Ethology 110 Oecologia 56 Argiope Verh Dtsch Zool Ges 79 In Ecophysiology of spiders Argiope argentata Zool Anz 221 The Biology of Spiders With especial Reference to the Danish Fauna Mitt Schweiz Entomol Ges 60 Anz Schädlingskde Pflanzenschutz Umweltschutz 54 J Appl Entomol 107 Ecology 61

102

Oecologia 49 Oecologia 49 Ecology 63 Funct Ecol 12 Biol Behav 9 Comp Biochem Physiol 19 Araneus diadematus J Exp Zool 176 Z Morphol Ökol Tiere 42 Zool Jb Physiol 97 Nature 411 Oecologia 76 In The praying mantids Tenodera aridifolia sinensis Sphodromantis lineola Brain Behav Evol 39 In The praying mantids Zool Stud 45 Argiope bruennichi Biologia 60 In Ecophysiology of spiders Argiope Bulletin des Reserves Naturelle et Ornithologiques de Belgique 4 Argiope trifasciata Heredity 83 Argiope trifasciata J Arachnol 31 Argiope keyserlingi Aust J Zool 55

103

Oecologia Pardosa Oecologia 5 Argiope Zool J Linn Soc 54 Argiope argentata Symp Zool Soc Lond 42 In Function and Evolution in Behavior Argiope argentata Can Ent 102 Nephila clavipes Psyche 80 Zool J Linn Soc 64 J Anim Ecol 46 Argiope bruennichi. Veröff Naturhist Mus Schleusingen 6 Argiope bruennichi Hercynia (NF) 26 Argiope bruennichi Entomol Nachr Ber 34 Zool J Linn Soc 120 Argiope bruennichi Entomologiske Meddelelser 65 Argiope argentata Behav Ecol Sociobiol 31 Proc Roy Soc Lond B 268 Argiope versicolor J Zool (Lond)258 Stegodyphus Oecologia 82 Histoire Naturelle des Araignées Argiope bruennichi Newsl Brit Arachnol Soc 87

104

Ann Zool Fennici 39 Frontinella pyramitela J Arachnol 13 J Arachnol 27 Agyrodes fissifrons Agelena limbata J Arachnol 12 Phonognatha graeffei Bull Brit Arachnol Soc 10 Nature 411 Araneus cavaticus J Insect Physiol 32 Argiope aurantia Argiope trifasciata Psyche 82 Argiope trifasciata Argiope aurantia Psyche 84 Argiope trifasciata Oikos Araneus cavaticus J Arachnol 12 J Exp Biol 209 Argiope trifasciata Anim Behav 52 Behaviour 135 Argiope trifasciata Am. Nat 141 Argiope aetheroides Behaviour 141 Nat. Mus 59 In Ecophysiology of Spiders In Ecophysiology of Spiders Nature 410 Argiope bruennichi J Arachnol 33

105

Argiope Behav Ecol Argiope bruennichi J Ethol Octonoba sybotides Proc Roy Soc Lond B 267 Ethol Ecol Evol 5 Z Morph Ökol Tiere 8 Z Morph Ökol Tiere 11 In Die Tierwelt Deutschlands und der angrenzenden Meeresteile Biological Invasions In Cambridge studies in ecology J Arachnol 33

106

Appendix

107

Danksagung Argiope keyserlingi Argiope sector

108

Curriculum vitae

Contact information Private University Personal information Education

109

Diploma thesis: Argiope bruennichi Argiope bruennichi PhD thesis: Argiope Publications Argiope bruennichi J Arachnol 33 Argiope Behav Ecol Argiope bruennichi J Ethol Revista Ibérica de Aracnologia Argiope keyserlingi J Arachnol Argiope bruennichi Ann Zool Fenn

110

Erklärung

111