THE DIVERSITY AND ECOLOGICAL IMPACTS OF

BUPRESTID AND CERAMBYCID ON EZEMVELO

NATURE RESERVE, GAUTENG PROVINCE

by

DUNCAN NEIL MACFADYEN

Submitted in partial fulfilment of the requirements for the degree

MAGISTER TECHNOLOGIAE: NATURE CONSERVATION

Department of Nature Conservation

TSHWANE UNIVERSITY OF TECHNOLOGY

Supervisor: Prof. B.K. Reilly

Co Supervisor: Dr C.L. Bellamy

February 2005 DECLARATION

The compilation of this thesis and the work reported on is the result of the author’s original work, unless specifically acknowledged, or stated to the contrary, in the text.

______

2 D.N. MacFadyen September 2004

ABSTRACT

Understanding the extent and cause of diversity on a nature reserve is often seen as a challenge to reserve management. Recent calculations that there may be more than 20 million species of on earth have focused attention on their magnitude and stimulated several new lines of research (although the true figure is now widely thought to be between five and ten million species). This study discusses work based on light trapping, beating and sweeping surveys, plant association, seasonal change and community dynamics of Cerambycidae and , families of the Order Coleoptera. It is argued that progress in estimating insect diversity in understanding insect community dynamics will be enhanced by building local inventories of species diversity, and in descriptive and experimental studies of the structure of communities.

As can be seen from their diaries and notebooks, contemplation of how such wonderful abundance and variety might arise was instrumental in pointing Darwin and especially Wallace to the theory of natural selection (Godfray et al. 1999).

This study was undertaken to investigate the community structure of two families of wood-boring beetles, providing a more systematic and quantitative approach to cataloguing insect diversity in a protected area context.

3 ACKNOWLEDGEMENTS

I would like to thank the following persons and organisations for their input to this study:

The Tshwane University of Technology and E Oppenheimer & Son for financial assistance.

Morgan Hauptfleish (reserve manager) for providing background information regarding the management practices of Ezemvelo Nature Reserve. He is also thanked for logistical support.

Dr C.L. Bellamy and Ruth Müller for taking the time and effort to aid in identification of the numerous specimens. Prof. George Bredenkamp is also thanked for assistance with categorizing the plant features. The following people assisted me in various capacities: Marion Burger, Tersia Perregil, Patrick Wood, Stuart Smith,

Nelius Uys, Willem van der Merwe and Tracey MacFadyen.

My parents, especially my father, Neil MacFadyen, for his support, motivation and faith in me during the course of this study.

Prof. B.K. Reilly for his valued guidance, support and encouragement throughout the course of this study. His tolerance, assistance and patience during the compilation of this thesis was also sincerely appreciated.

Finally, I would like to thank Strilli Oppenheimer, whose great love for insects led to this project being conceived.

4 INDEX

5 PAGE

Declaration ...…………………………………………………………………………… 1

Abstract ….………………………….………………………………………………….. 2

Acknowledgements ...………………………………………………………………...... 3

Index ……………………………………………………………………………………. 4

Chapter One

1. INTRODUCTION……………………………………………………………… 32

1.1 Views of the insect community ……………………………………...... 35

1.2 Rationale for this study ………………………………………………………… 36

1.3 Objectives ………………………………………………………………………. 37

1.4 Hypotheses …………………………………………………..…………………. 37

Chapter Two

2. MATERIALS AND METHODS……………………………………………… 38

2.1 STUDY AREA………………………………………………………………….. 38

2.1.1 Phytosociology………………………………………………………………...... 39

2.1.2 Geology………………………………………………………………………….. 40

2.1.3 Soil………………………………………………………………………………. 40

2.1.4 Climate………………………………………………………………………...... 40

2.1.5 Hydrology……………………………………………………………………….. 40

PAGE

6 2.1.6 Vegetation………………………………………………………………………..6

2.2 METHODS……………………………………………………………………..

2.2.1 Preparatory Work………………………………………………………………...

2.2.2 Reconnaissance…………………………………………………………………..

2.2.3 Stand and quadrant dimensions…………………………………………………..

2.2.4 Positioning of quadrats………………………………………………………......

2.2.5 Field location of quadrats………………………………………………………...

2.2.6 Layout and orientation of quadrats………..……………………………………..

2.3 Field data collection……………………………………………………………..

2.3.1 Abiotic factors………………………………………………………………......

2.3.2 Biotic factors…………………………………………………………………….

2.3.2.1 Plants…………………………………………………………………………….

2.3.2.2 Beetle families…………………………………………………………………..

2.3.3 Collection methods………………………………………………………………

2.3.3.1 Beating method………………………………………………………………….

2.3.3.2 Sweeping method………………………………………………………………..

2.3.3.3 Sheet trap method………………………………………………………………..

2.3.4 Identification of beetles…………………………………………………………..

2.3.5 Data preparation………………………………………………………………….

2.3.6 Statistical analysis………………………………………………………………..

2.3.6.1 Chi Square analysis………..……………………………………………………..

2.3.6.2 Cramér’s V analysis…………………………………………………………...... 2.3.6.2 Cramér’s V analysis…………………………………………………………......

Chapter Three

3. RESULTS………………………………………………………………………

3.1 Cerambycidae data…..………………………………..………………………….

3.1.1 Total Cerambycidae data……………….………………………………......

3.1.2 Quadrat A……………………………………………………..……………...... 7

3.1.3 Quadrat B…………………………………………………………………......

3.1.4 Quadrat C……………………………………………………………………….

3.2 Buprestidae data……………………...………………………………………….

3.2.1 Total Buprestidae data……………………………………………………….....

3.2.2 Quadrat A……..…………………………………………………………….....

3.2.3 Quadrat B…………………………………………………………………….....

3.2.4 Quadrat C…………………………………………………………………….....

3.3 Ecological processes affecting Cerambycidae and Bupresridae on ENR………..

3.4 Cerambycidae/Plant correlation………..……………………………………….

3.4.1 Plant order correlation………………………………………………………….

3.4.2 Plant family correlation…...………………………………………………......

3.4.3 Plant species correlation…………………...…………………………………...

3.4.4 Plant flower size correlation…………………………………………………… 3.4.5 Plant phenology correlation…………………………………………………...

3.4.6 Plant pollination………………………………………………………………...

3.4.7 Plant climate………………………………………………………………….....

3.5 Buprestidae/Plant correlation…………………………………………………….

3.5.1 Plant order correlation………………………………………………………......

3.5.2 Plant family correlation………………………………………………………….

3.5.3 Plant species correlation…………...…………………………………………….

3.5.4 Plant flower size correlation…….………………………..…………………......

3.5.5 Plant phenology correlation……….………………..……………………………

3.5.6 Plant pollination…………………………………………………………………. 8 3.5.7 Plant climate…………………………………………………………………......

3.6 Discussion………………………………………………………………………..

Chapter Four

4. DISCUSSION……………………………………………………………………

4.1 Classification of …………………………………………………...

4.2 Classification of Cerambycidae………………………………………………….

4.3 Factors influencing the abundance and diversity on ENR………………...……..

4.3.1 Cerambycidae abundance and diversity on ENR……………………………......

4.3.2 Buprestidae abundance and diversity of ENR………...…………………………

4.4 Overview………………………………………………………………………… Chapter Five

4 ECOLOGICAL IMPORTANCE AND FUTURE MANAGEMENT OF

CERAMBYCIDAE AND BUPRESTIDAE ON EZEMVELO NATURE

RESERVE…………………………………………………………………………..

Chapter Six

5 CONCLUSION……………………………………………………………………..

Chapter Seven

6 REFERENCES………………………………………………………………………

LIST OF FIGURES

Figure 1: Map of showing Ezemvelo Nature Reserves positioning on the

border of Gauteng and Mpumalanga provinces………………………………. 9

Figure 2: The greater Telperion Nature Reserve with Ezemvelo Nature Reserve on the

western boundary divided by the Wilge River………………………………...

Figure 3: The Wilge river on Ezemvelo Nature Reserve is dominated by rocky

outcrops and woody vegetation……………………………………………… Figure 4.1: Ezemvelo Nature Reserve and quadrats (A, B & C) with

overlaying slope classes…………………………………………………….

Figure 4.2: Ezemvelo Nature Reserve and quadrats (A, B & C)

with overlaying altitude categories………………………………………....

Figure 4.3: Ezemvelo Nature Reserve and quadrats (A, B & C) with

overlaying aspect classes…………………………………………………...

Figure 4.4: Broad vegetation zones of Ezemvelo Nature Reserve

showing placement of quadrats A,B & C…………………………………..

Figure 5.1.1: Transect line A1 within quadrat A in September 2001

on Ezemvelo Nature Reserve………………………………………………

Figure 5.1.2: Transect line A2 within quadrat A in September 2001

on Ezemvelo Nature Reserve………………………………………………

Figure 5.1.3: Transect line A3 within quadrat A in September 2001

on Ezemvelo Nature Reserve……………………………………………… 10

Figure 5.2.1: Transect line B1 within quadrat B in September 2001 Figure 5.2.1: Transect line B1 within quadrat B in September 2001

on Ezemvelo Nature Reserve………………......

Figure 5.2.2: Transect line B2 within quadrat B in September 2001

on Ezemvelo Nature Reserve……………………………………………..

Figure 5.2.3: Transect line B3 within quadrat B in September 2001

on Ezemvelo Nature Reserve……………………………………………..

Figure 5.3.1: Transect line C1 within quadrat C in September 2001

on Ezemvelo Nature Reserve……………………………………………..

Figure 5.3.2: Transect line C2 within quadrat C in September 2001

on Ezemvelo Nature Reserve……………………………………………..

Figure 5.3.3: Transect line C3 within quadrat C in September 2001

on Ezemvelo Nature Reserve……………………………………………...

Figure 6.1.1: The beating method being used on Protea caffra in

quadrat C on Ezemvelo Nature Reserve…………………………………..

Figure 6.1.2: The beating method being used on Acacia caffra in

quadrat A on Ezemvelo Nature Reserve………………………………….. Figure 6.2: The sweeping method being used on Acacia caffra

in quadrat A on Ezemvelo Nature Reserve………………………………..

11

Figure 6.3: The sheet trap method being used in quadrat B

on Ezemvelo Nature Reserve in October 2001…………………………….

Figure 7.1: Anubis clavicornis………………………………………………………………….

Figure 7.2: Anubis mellyi………………………………………………………………………..

Figure 7.3: Anthracocentrus capensis…………………………………………………………

Figure 7.4: Ceroplesis thunbergi……………………………………………………………….

Figure 7.5: Coptoeme krantzi…………………………………………………………………...

Figure 7.6: Crossotus lacunosus………………………………………………………………..

Figure 7.7: Crossotus plumicornis……………………………………………………………..

Figure 7.8: Crossotus stypticus………………………………………………………………… Figure 7.9: Hecyra terrea……………………………………………………………………….

Figure 7.10: Hypoeschrus ferreirae……………………………………………………………

Figure 7.11: Jonthodina sculptilis……………………………………………………………...

Figure 7.12: Lasiopezus longimanus…………………………………………………………..

12

Figure 7.13: Macrotoma natala………………………………………………………………..

Figure 7.14: Macrotoma palmate………………………………………………………………

Figure 7.15: Mycerinicus brevis………………………………………………………………..

Figure 7.16: Nemotragus helvolus……………………………………………………………..

Figure 7.17: Olenecamptus albidus…………………………………………………………….

Figure 7.18: Ossibia fuscata…………………………………………………………………….

Figure 7.19: Pacydissus sp……………………………………………………………………… Figure 7.20: Phantasis giganteus………………………………………………………………

Figure 7.21: Philematium natalense…………………………………………………………...

Figure 7.22: Phryneta spinator…………………………………………………………………

Figure 7.23: Phyllocnema latipes……………………………………………………………...

Figure 7.24: Plocaederus denticornis………………………………………………………….

Figure 7.25: Dalterus degeeri…………………………………………………………………..

Figure 7.26: Dalterus dejeani…………………………………………………………………..

13 Figure 7.27: Prosopocera lactator……………………………………………………………..

Figure 7.28: Alphitopola octomaculata………………………………………………………..

Figure 7.29: Taurotagus klugi…………………………………………………………………..

Figure 7.30: Tithoes maculates…………………………………………………………………

Figure 7.31: Tragiscoschema bertolinii………………………………………………………. Figure 7.32: Xystrocera erosa…………………………………………………………………..

Figure 7.33: Xystrocera dispar…………………………………………………………………

Figure 7.34: Zamium bimaculatum…………………………………………………………….

Figure 7.35: Zamium incultum………………………………………………………………….

Figure 8: Monthly sampling frequencies for the total Cerambycidae collected for 2001

on Ezemvelo Nature Reserve……………………………………………

Figure 9.1: Total Cerambycidae count trend versus average minimum temperature for

the months of the year at Ezemvelo Nature Reserve for January 2001 to

December 2001…………………………………………………………

Figure 9.2: Total Cerambycidae count trend versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve

for January 2001 to December 2001……………………………………… 14

Figure 9.3: Total Cerambycidae count trend versus average monthly

rainfall for the months of the year at Ezemvelo Nature Reserve

for January 2001 to December 2001……………………………………… Figure 10: Monthly sampling frequencies for Cerambycidae collected

in quadrat A for 2001 on Ezemvelo Nature Reserve………………………

Figure 11.1: Cerambycidae count trend in quadrat A versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 to December 2001………………………………...

Figure 11.2: Cerambycidae count trend in quadrat A versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 to December 2001………………………………...

Figure 11.3: Cerambycidae count trend in quadrat A versus average monthly

rainfall for the months of the year at Ezemvelo Nature Reserve between

January 2001 to December 2001…………………………………………..

Figure 12: Monthly sampling frequencies for Cerambycidae collected

in quadrat B for 2001 on Ezemvelo Nature Reserve………………………

Figure 13.1: Cerambycidae count trend in quadrat B versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………... Figure 13.2: Cerambycidae count trend in quadrat B versus average maximum 15 temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………….

Figure 13.3: Cerambycidae count trend in quadrat B versus average monthly rainfall

for the months of the year at Ezemvelo Nature Reserve between January

2001 and December 2001…………………………………………………

Figure 14: Monthly sampling frequencies for Cerambycidae collected in quadrat C for

2001 on Ezemvelo Nature Reserve……………………………………

Figure 15.1: Cerambycidae count trend in quadrat C versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………….

Figure 15.2: Cerambycidae count trend in quadrat C versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………….

Figure 15.3: Cerambycidae count trend in quadrat C versus average monthly rainfall

for the months of the year at Ezemvelo Nature Reserve between January

2001 and December 2001………………………………………………… Figure 16: Monthly sampling frequencies of Cerambycidae between three quadrats on

Ezemvelo Nature Reserve…………………………………………………

Figure 17.1: Acmaeodera aenea………………………………………………………………

Figure 17.2: Acmaeodera albivillosa…………………………...……………………………

… 16

Figure 17.3: Agrilus guerryi………………………..………………………………………….

Figure 17.4: Sphenoptera sinuosa………………………..…………………………………….

Figure 17.5: Acmaeodera punctatissima………………….…………………………………..

Figure 17.6: Acmaeodera inscripta…………..………………………………………………...

Figure 17.7: Acmaeodera ruficaudis……..…………………………………………………….

Figure 17.8: Agrilus sexguttatus………………..……………………..………………………

Figure 17.9: Acmaeodera stellata………………….……….…………………………………..

Figure 17.10: Acmaeodera viridiaenea……………………………………………………… Figure 17.11: Anthaxia sp. 1…………………………………..………………………………...

Figure 17.12: Chrysobothris algoensis………………………………….…………………….

Figure 17.13: Chrysobothris boschismanni………………………………………………….

Figure 17.14: Chrysobothris dorsata…………………………………………………………

Figure 17.15: Evides pubiventris…..…………………...………………………………..

Figure 17.16: Lampetis gregaria……………………..……………………………………….17

Figure 17.17: Phlocteis exasperata…………………………………………………………...

Figure 17.18: Pseudagrilus beryllinus………………………………………………………..

Figure 17.19: Lampetis conturbata…..……………………………………………………….

Figure 17.20: Sphenoptera arrowi……………………………………………………………

Figure 17.21: orissa……………….……………………………….…………….. Figure18: Monthly sampling frequencies for total Buprestidae collected for 2001 on

Ezemvelo Nature Reserve……………………………………………………

Figure19.1: Total Buprestidae count versus average minimum temperature for the

months of the year at Ezemvelo Nature Reserve between January 2001

and December 2001……………………………………………………….

Figure 19.2: Total Buprestidae count versus average maximum temperature for the

months of the year at Ezemvelo Nature Reserve between January 2001

and December 2001……………………………………………………….

Figure 19.3: Total Buprestidae count versus average monthly rainfall for the months of

the year at Ezemvelo Nature Reserve between January 2001 and

December 2001……………………………………………………………

Figure 20: Monthly sampling frequencies for Buprestidae collected in quadrat A for

2001 at Ezemvelo Nature Reserve...... 18

Figure 21.1: Buprestidae count trend in quadrat A versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………….

Figure 21.2: Buprestidae count trend in quadrat A versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001……………………………….

Figure 21.3: Buprestidae count trend in quadrat A versus average monthly rainfall for

the months of the year at Ezemvelo Nature Reserve between January

2001and December 2001………………………………………………….

Figure 22: Monthly sampling frequencies for Buprestidae collected in quadrat B for

2001 at Ezemvelo Nature Reserve………………………………………….

Figure 23.1: Buprestidae count trend in quadrat B versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 to December 2001………………………………….

Figure 23.2: Buprestidae count trend in quadrat B versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 to December 2001………………………………….

Figure 23.3: Buprestidae count trend in quadrat C versus average monthly rainfall for

the months of the year at Ezemvelo Nature Reserve between January 2001

to December 2001………...... 19 Figure 24: Monthly sampling frequencies for Buprestidae collected in quadrat C for

2001 on Ezemvelo Nature Reserve…………………………………………

Figure 25.1: Buprestidae count trend in quadrat C versus average minimum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001………………………………...

Figure 25.2: Buprestidae count trend in quadrat C versus average maximum

temperature for the months of the year at Ezemvelo Nature Reserve

between January 2001 and December 2001……………………………….

Figure 25.3: Buprestidae count trend in quadrat C versus average monthly rainfall for

the months of the year at Ezemvelo Nature Reserve between

January 2001 and December 2001………………………………………...

Figure 26: Monthly sampling frequencies of Buprestidae between the three quadrats

on Ezemvelo Nature Reserve………………………………………………

Figure 27: Monthly sample comparison between Buprestidae and Cerambycidae on Ezemvelo Nature Reserve…………………………………………………..

Figure 28.1: Relationship between Tree Orders and Cerambycidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001…

…………………………………………………………………….. 20

Figure 28.2: Relationship between Tree Family and Cerambycidae species collected

on Ezemvelo Nature Reserve between January 2001and December

2001……………………………………………………………………….

Figure 28.3: Relationship between plant species and Cerambycidae species collected

on Ezemvelo Nature Reserve between January 2001 and December

2001………………………………………………………………………

Figure 28.4: Relationship between flower size and Cerambycidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001…

…………………………………………………………………….

Figure 28.5: Relationship between plant phenology and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001 and

December 2001………………………………………………………….. Figure 28.6: Relationship between plant pollination and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001 and

December 2001……………………………………………………………

Figure 28.7: Relationship between plant climate and Cerambycidae species collected

on Ezemvelo Nature Reserve between January 2001 and December

2001………………………………………………………………………..

Figure 29.1: Relationship between Tree Orders and Buprestidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001… 21 ……………………………………………………......

Figure 29.2: Relationship between plant family and Buprestidae species collected on

Ezemvelo Nature Reserve between January 2001 and December

2001…......

Figure 29.3: Relationship between plant species and Buprestidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001…

……………………………………………………………………..

Figure 29.4: Relationship between flower size and Buprestidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001…

……………...... Figure 29.5: Relationship between plant phenology and Buprestidae species

collected on Ezemvelo Nature Reserve between January 2001 and

December 2001………......

Figure 29.6: Relationship between plant pollination and Buprestidae species collected

on Ezemvelo Nature Reserve between January 2001 and December

2001……………………………………………………………………….

Figure 29.7: Relationship between plant climate and Buprestidae species collected on

Ezemvelo Nature Reserve between January 2001 and December 2001…

……………………………………………………………………..

LIST OF TABLES

22 Table 1. List of species not identified to species level…………………………………...

Table 2: The analysis of the regression indicates a significant linear trend or

insignificant linear trend of the following parameters for Cerambycidae on

Ezemvelo Nature Reserve……………………………………………………..

Table 3: The analysis of the regression indicates the degree of linear trend for the first

six months of the year for Cerambycidae on Ezemvelo Nature

Reserve...... Table 4: The analysis of the regression indicates the degree of linear trend for the last

six months of the year for Cerambycidae on Ezemvelo Nature Reserve………

………………………………………………………………...

Table 5: The analysis of the regression indicates a significant linear trend or

insignificant linear trend of the following parameters for Buprestidae on

Ezemvelo Nature Reserve………………......

Table 6: The analysis of the regression indicates the degree of linear trend for the first

six months of the year for Buprestidae on Ezemvelo Nature Reserve………..

Table 7: The analysis of the regression indicates the degree of linear trend for the last

six months of the year for Buprestidae on Ezemvelo Nature Reserve......

LIST OF APPENDIXES

Appendix A Total Cerambycidae species diversity and abundance for each month of

the year in all quadrats on ENR…………………………………………...23

Appendix B Percentage of Cerambycidae species diversity and abundance for each

month of the year in all quadrats on ENR………………………………… Appendix C Total Cerambycidae species diversity and abundance for each month of

the year in quadrat A on ENR………......

Appendix D Percentage of Cerambycidae species diversity and abundance for each

month of the year in quadrat A on ENR…………………………………..

Appendix E Total Cerambycidae species diversity and abundance for each month of

the year in quadrat B on ENR…......

Appendix F Percentage of Cerambycidae species diversity and abundance for each

month of the year in quadrat B on ENR…………......

Appendix G Total Cerambycidae species diversity and abundance for each month of

the year in quadrat C on ENR…………………......

Appendix H Percentage of Cerambycidae species diversity and abundance for each

month of the year in quadrat C on ENR……………......

Appendix I Total Buprestidae species diversity and abundance for each month of the

year in all quadrats on ENR………………………………………………. Appendix J Percentage of Buprestidae species diversity and abundance for each month

of the year in all quadrats on ENR…………………………………………

24 Appendix K Total Buprestidae species diversity and abundance for each month of the

year in quadrat A on ENR…………………………………………………

Appendix L Percentage of Buprestidae species diversity and abundance for each month

of the year in quadrat A on ENR…………………………………...

Appendix M Total Buprestidae species diversity and abundance for each month of the

year in quadrat B on ENR…………………………………………………

Appendix N Percentage of Buprestidae species diversity and abundance for each month

of the year in quadrat B on ENR…………......

Appendix O Total Buprestidae species diversity and abundance for each month of the

year in quadrat C on ENR…………………………………………………

Appendix P Percentage of Buprestidae species diversity and abundance for each month

of the year in quadrat C on ENR………......

Appendix Q Total Cerambycidae species diversity and abundance for each month and

associated plant Orders in all quadrats on ENR…………………………...

Appendix R Percentage of Cerambycidae species diversity and abundance for each Appendix R Percentage of Cerambycidae species diversity and abundance for each

month and associated plant Orders in all quadrats on ENR……………...

Appendix S Total Cerambycidae species diversity and abundance for associated plant

families in all quadrats on ENR…………………………………………...

Appendix T Percentage of Cerambycidae species diversity and abundance for and

associated plant families in all quadrats on ENR………………………….

25

Appendix U Total Cerambycidae species diversity and abundance for each month and

associated plant species in all quadrats on ENR…………………………..

Appendix V Percentage of Cerambycidae species diversity and abundance for each

month and associated plant species in all quadrats on ENR………………

Appendix W Total Cerambycidae species diversity and abundance for each month and

associated decideous or non-deciduous plants in all quadrats on ENR……

…………………………………………………………………..

Appendix X Percentage of Cerambycidae species diversity and abundance for each

month and associated deciduous or non-deciduous plants in all quadrats

on ENR………………………………………………………………….. Appendix Y Total Cerambycidae species diversity and abundance for each month and

associated plant pollination in all quadrats on ENR………………………

Appendix Z Percentage of Cerambycidae species diversity and abundance for each

month and associated plant pollination in all quadrats on ENR……......

Appendix AA Total Cerambycidae species diversity and abundance for each month

and associated plant climate in all quadrats on ENR…......

Appendix AB Percentage of Cerambycidae species diversity and abundance for each

month and associated plant climate in all quadrats on ENR…......

Appendix AC Total Cerambycidae species diversity and abundance for each month

and associated flower size26 in all quadrats on ENR………………………

Appendix AD Percentage of Cerambycidae species diversity and abundance for

associated flower size in all quadrats on ENR…………………………...

Appendix AE Total Buprestidae species diversity and abundance for associated plant

Orders in all quadrats on ENR…………………………………………...

Appendix AF Percentage of Buprestidae species diversity and abundance for

associated plant Orders in all quadrats on ENR……...... Appendix AG Buprestidae species diversity and abundance for associated plant

families in all quadrats on ENR…………......

Appendix AH Percentage of Buprestidae species diversity and abundance for

associated plant families in all quadrats on ENR………………………...

Appendix AI Total Buprestidae species diversity and abundance for associated plant

species in all quadrats on ENR…………………………………………..

Appendix AJ Percentage of Buprestidae species diversity and abundance for associated

plant species in all quadrats on ENR…………………………

Appendix AK Total Buprestidae species diversity and abundance for associated plant

phenology in all quadrats on ENR……………………………………….

Appendix AL Percentage of Buprestidae species diversity and abundance for

associated deciduous or non-deciduous plants in all quadrats on ENR…

………………………………………………………………….27

Appendix AM Total Buprestidae species diversity and abundance for associated plant

pollination in all quadrats on ENR……………………………………...

Appendix AN Percentage of Buprestidae species diversity and abundance for associated plant pollination in all quadrats on ENR……………………

Appendix AO Total Buprestidae species diversity and abundance for associated plant

climate in all quadrats on ENR…………………………………………

Appendix AP Percentage of Buprestidae species diversity and abundance for

associated plant climate in all quadrats on ENR………………………..

Appendix AQ Total Buprestidae species diversity and abundance for associated flower

size in all quadrats on ENR……………………………………......

Appendix AR Percentage of Buprestidae species diversity and abundance for

associated flower size in all quadrats on ENR…………………………...

Appendix AS Cerambycidae light trap results for 2001 on Ezemvelo Nature Reserve…

……………………………………………………………….

Appendix AT Buprestidae light trap results for 2001 on Ezemvelo Nature Reserve……

……………………………………………………………... Chapter One

1. INTRODUCTION

The aim of this study was to quantify the species diversity and abundance of two families of Coleoptera (beetles), the Cerambycidae and Buprestidae on Ezemvelo

Nature Reserve with reference to seasonality, vegetation and climate.

Invertebrates comprise the bulk of global species richness, and the loss of invertebrate species will constitute much of the loss of biodiversity (New 1993; Samways 1994;

New & Yen 1995; Scholtz & Chown 1995). The importance of these two families of wood boring beetles to various ecological processes and the aforementioned biodiversity should not be underestimated. In addition, invertebrates may be used as effective bio-indicators of environmental change (Jansen 1987; Kremen et al. 1993;

New 1993; Kremen 1994; New & Yen 1995; Weaver 1995). Most invertebrate studies have focused mainly on soil-dwelling insects and those considered pests to crops and timber. According to Hull et al. (1998), no studies have examined the

7 distribution of phytophagous insects and the identification of priority areas that would be required to conserve them, although phytophages represent the highest proportion of terrestrial insect species (Lawton & Strong 1981).

Information pertaining to host plants and the association of different species is vital to conservation of the biodiversity of these beetles. It is necessary to calculate the abundance and diversity of beetles on various plants species.

According to Holm & Bellamy (1985), very little is known of the biology of most buprestid taxa, except that they are among the most thermophilic insects known, and the larval stages are often prolonged compared to those of the adult.

In southern Africa, only three studies have sought to investigate areas for conservation of insect taxa (Freitag & Mansell 1997; Muller et al 1997; Hull et al. 1998). Selection of priority conservation areas based on species richness has been shown to be highly insufficient in southern Africa (Kershaw et al. 1994; Freitag & Van Jaarsveld, 1995;

Williams et al. 1996). This study therefore provides an important component to conservation of these two families, especially within the Bankenveld (61) veld type.

According to Noss (1990), Pickett et al. (1992) and Walker (1992), information on phytophagous insects is of importance if priority areas are to be selected on the basis of maintaining ecosystem functioning and not simply species richness.

8 Many populations of insect species have declined markedly over recent years, primarily as a result of agricultural intensification (Aebischer 1991; Feber et al. 1997;

Benton et al. 2002). There are few large and plants left to be discovered in the world (Godfray et al. 1999), yet our ignorance about the number of insects on earth (and, more generally, the total number of all species on earth) is largely due to lack of knowledge regarding certain groups. We know from well studied groups that fewer species are described from smaller insects and not a result of insufficient sampling of the smaller species (Dial & Marzluff 1988).

Classifying species into functional groups that are ecologically relevant (Simberloff &

Dayan 1991; Peeters et al. 2001), allows comparisons and generalizations to be made about insects that are not possible using taxonomic groupings alone. Plant functional groups have been used extensively to determine responses to climate change based on their photosynthetic pathways, plant lifespan, above-ground biomass and geographical location (Bazzaz 1990; Cammell & Knight 1992; Landsberg & Stafford Smith 1992;

Paruelo & Lauenroth 1995; Condit et al. 1996; Diaz & Cabido 1997; Cornelissen et al. 2001; Dormann & Woodin 2002; Epstein et al. 2002; Richardson et al. 2002).

As bemoaned by May (1988, 1990) and others, a century and a half later we have only a rough idea of the actual dimensions of insect species diversity, and an even poorer understanding of the processes through which it is generated and maintained. The sheer weight of the number of species oppresses the whole subject matter (Godfray et al. 1999), and with the exception of a few taxonomic groups, of which butterflies are the most prominent, field identification is impossible or very difficult. But a hundred

9 years ago the situation was not that different for botanists confronted with forests, which might contain several hundred tree species per square kilometre (Godfray et al.

1999). This study suggests that insect ecologists can learn from the success plant ecologists have had in understanding diversity, even if like a puzzle, families of insects can be studied individually and through time a similar situation will prevail as with plants. As suggested by Godfray et al. (1999), entomologists should follow the example of plant ecologists.

The floral inventories are more than just species lists, but include the various characteristics of the plant. They provide a baseline that allows other more poorly known sites to be assessed for species diversity, structure and abundance. This is the massive task insect ecologists have to tackle. This study provides the fundamental answers to diversity and ecological requirements of two families of wood boring beetles on Ezemvelo Nature Reserve.

1.1 Views of the insect community

The populations of plants and phytophagous insects living together in a given environment and interacting with one another to form a distinct living system constitute a biotic community (Storer et al. 1979). Communities are often named for some dominant feature, biotic or physical; e.g., Acacia woodland, Riverine thicket.

The species composition of a community depends on climate and historical

(evolutionary) factors.

10 Some communities have rather sharp boundaries where ranges of some of the more conspicuous species stop, but other communities grade into one another in varying degrees (Storer et al. 1979). Sudden changes often occur when environmental gradients (in temperature, moisture, soil conditions) are steep or change abruptly. The extent of interaction between community members varies. In general, diversity tends to increase in communities with time by the addition of species differing in niche and habitat (Storer et al. 1979). In most communities a few species are dominant over others in numbers or physical characteristics or both. A natural community has been compared by analogy to an organism.

Each species is a separate entity with its own hereditary mechanism responding to natural selection in its own way, although influenced in the community context

(Storer et al. 1979). A community of organisms and their nonliving environment at any one place together constitute an ecosystem (Storer et al. 1979).

1.2 Rationale for this study

A study conducted in 1999 under the auspices of the University of Pretoria provided an overview of the broad vegetation types on Ezemvelo Nature Reserve (ENR), followed by a brief one week of invertebrate study conducted by the Transvaal

Museum in 2004. Unfortunately these studies were very superficial and no connection was made between the two from an ecological point of view. The need for a comprehensive inventory of wood boring beetles and their associated host plants was necessary. Large mammals have had numerous studies undertaken on them, yet the most abundant of all species on the reserve, the invertebrates, were practically

11 untouched from a research perspective. This document will serve as the foundation for future studies in the field of entomology on ENR that has been lacking to date.

The present study investigates species richness, abundance and plant preference of

Cerambycidae and Buprestidae between January 2001 and December 2001. This study provides an estimate of the species richness of these two families in a

Bakenveld (Acocks veldtype 61) reserve. Ezemvelo Nature Reserve has extreme variations in temperature, which is associated with seasonal change. This study also aims to identify plants species that are key to absence or presence of species of beetles.

1.3 Objectives

To quantify species diversity and abundance of beetles of the families

Cerambycidae and Buprestidae on ENR,

To establish correlations between Cerambycidae and Buprestidae and woody plant

species on ENR,

To determine seasonal variation in abundance of Cerambycidae and Buprestidae

on ENR,

To endeavour to predict those factors likely to affect the population densities of

species within these families on ENR,

To determine criteria from which the conservation of these two families can be

incorporated into the reserves management plan.

12 1.4 Hypotheses

The null hypothesis is cerambycid and buprestid beetle species occur

independently of vegetation and time on ENR.

The alternative hypothesis is Cerambycid and Buprestid beetle species occur

dependently of vegetation and time on ENR.

Chapter Two

2. MATERIAL AND METHODS

2.1 STUDY AREA

Ezemvelo Nature Reserve is situated on the border between Gauteng and

Mpumalanga, 25 km North-East of Bronkhorstspruit (Figure 1.).

13 Figure 1: Map of South Africa showing Ezemvelo Nature Reserves positioning

on the border of Gauteng and Mpumalanga provinces.

ENR lies on the farm Elandsfontein 493 JR, between the 25 38` S and 28 53` E. ENR is situated in what is referred to as the Rocky Highveld Grassland or Bankenveld

(Veldtype 61) (Acock’s 1988), in a Grassland Biome. ENR forms the western section of the Telperion Nature Reserve (Figure 2).

N

Figure 2: The greater Telperion Nature Reserve with Ezemvelo Nature

Reserve on the western boundary divided by the Wilge River.

14 2.1.1 Phytosociology

The landscape and topography are dominated by grassy plains, interspersed with rocky outcrops dominated by woody species. The lower lying, more steeply sloped areas tend to be dominated by rocky woodlands (Grobler 1999).

2.1.2 Geology

The area lies on the Wilge River, Ecca and Dwyka formations of the Waterberg and

Karoo groups, which were formed during the Mokolian and Palaeozoic eras respectively (Grobler 1999).

2.1.3 Soil

The lithology is dominated by Arenite-Conglomerate, which produces dystrophic or mesotrophic soils with some red soils, as well as rocky areas with miscellaneous soil.

The Tilite-Arenite produces some rocky areas with miscellaneous soils, as do shale based soils (Grobler 1999).

2.1.4 Climate

The area receives summer rainfall averaging between 650 mm and 700 mm per year.

The temperature reaches a maximum of 39 ˚C and lows of –12 ˚C (Louw & Rebelo

1988). The average minimum and maximum temperatures are 3 ˚C and 28 ˚C. The highest average rainfall is recorded in January. Frost occurs readily in winter from

May to August (Bornman 1995).

15 2.1.5 Hydrology

ENR is bounded by the perennial Wilge river (Figure 3.) and contains three streams that originate from higher lying wetlands and sponge areas (Grobler 1999).

Figure 3: The Wilge River on Ezemvelo Nature Reserve is dominated by rocky

outcrops and woody vegetation.

2.1.6 Vegetation

The grass layer is thought to be maintained by frequent fires, usually not including rocky outcrops. The protection in the rocky areas against frost in winter, also plays an

16 important role in the distribution of the woody plant species (Louw & Rebelo 1988).

Rocky hills and ridges carry bushveld vegetation dominated by Protea caffra, Acacia caffra, A. karoo and Celtis africana.

2.2 METHODS

Cerambycidae and Buprestidae were sampled monthly for one year, from January

2001 (summer) to December 2001 (summer), a total of twelve collections. The basic procedure of the beating method (Holm 1984) was employed on all trees and shrubs within the three pre-selected quadrats. The object of a beating sheet is to capture invertebrates which do not fly readily at low temperatures. The sweeping method

(Holm 1984) is used at high temperatures where insect activity is great. The method of light trapping (Holm 1984) is utilized in the evenings when collecting species attracted to light.

2.2.1 Preparatory Work

2.2.2 Reconnaissance

A reconnaissance is the preliminary inspection or familiarization of the study area prior to sampling which has the object of estimating floristic and environmental variation, familiarization with the flora and to obtain permission from authorities for later work and accommodation possibilities (Westfall 1992; Westfall et al. 1996).

17 This exercise was completed in January 2001 during which time three quadrats were placed randomly in different major vegetation types.

2.2.3 Stand and quadrat dimensions

The concept of stand and sampling plot or quadrat, which are fundamental elements of the Braun-Blauquet approach (Mueller-Dombois & Ellenberg 1974; Westhoff &

Van der Maarel 1980) were placed at each stand of vegetation. A stand is defined as a portion of vegetation that is relatively homogenous in all layers (species composition, growth form and density) and differs from the contiguous types by either quantitative or qualitative characteristics (Daubenmire 1969, In: Panagos 1995). The plot or quadrant refers to a unit that has a measurable area (Panagos 1995) which is placed within the stand from which data is collected (Westhoff & Van der Maarel 1980).

2.2.4 Positioning of quadrats

Grunow (1996), Westfall (1992) and Westfall et al. (1997) have emphasized the need for the reduction in or elimination of observer bias with regard to sampling and classification. For this study, due to the reserve being primarily grassland, the three quadrats were confined to wooded areas along the rocky outcrops and riverine areas.

Three transects were placed in each quadrat to incorporate different plant communities, where host plants representing different species were studied. This included different slope classes (Figure 4.1), altitude (Figure 4.2) and aspect (Figure

4.3) to ensure the entire spectrum was incorporated.

18 Figure 4.1: Ezemvelo Nature Reserve and quadrats (A, B & C) with overlaying

slope classes.

19 Figure 4.2: Ezemvelo Nature Reserve and quadrats (A, B & C) with overlaying

altitude categories.

Figure 4.3: Ezemvelo Nature Reserve and quadrats (A, B & C) with overlaying aspect

classes.

20 KE Y Acacia caffra dominated Acacia karoo/Gymnosporia buxifolia dominated Protea caffra dominated

Figure 4.4: Broad vegetation zones of Ezemvelo Nature Reserve showing placement of

quadrats A, B & C.

2.2.5 Field location of quadrats

Location of predetermined stands was uncomplicated due to the nature of the terrain as areas sampled were islands of wooded vegetation surrounded by grassland (Figure

4.4).

2.2.6 Layout and orientation of quadrats

Three quadrats with nine transect lines were positioned resulting in approximately one kilometre of vegetation being surveyed for beetle activity per quadrat. The three quadrats were randomly placed within each selected stand of vegetation. Each quadrat

21 measured 200 m in length and 100 m in width. In this study, the quadrats were designated A, B and C. Chevron tape was used to mark the positioning of the three transect lines within each quadrat. The three lines were spaced 30 m apart and placed through the quadrat. The transect lines in this study were designated A1, A2, A3, B1,

B2, B3 and C1, C2, C3 respectively (Figure 5.1.1; Figure 5.1.2; Figure 5.1.3; Figure

5.2.1; Figure 5.2.2; Figure 5.2.3; Figure 5.3.1; Figure 5.3.2; Figure 5.3.3). These quadrats were examined both separately and in combination to determine whether the efficiency and identity of area selection for each site changed with changing topography. Because the number of quadrats that can ultimately be selected for surveys is likely to be limited by economic and other considerations, a primary aim in quadrat selection must be to represent all attributes in as small an area as possible

(Kershaw et al. 1994). Monthly calculations of beetles collected within each quadrat were calculated, as was the total number within all quadrats.

Figure 5.1.1: Transect line A1 within quadrat A in September 2001 on Ezemvelo Nature Reserve.

22 Figure 5.1.2: Transect line A2 within quadrat A in September 2001 on Ezemvelo Nature Reserve.

Figure 5.1.3: Transect line A3 within quadrat A in September 2001 on Ezemvelo Nature Reserve.

23 Figure 5.2.1: Transect line B1 within quadrat B in September 2001 on Ezemvelo Nature Reserve.

Figure 5.2.2: Transect line B2 within quadrat B in September 2001 on Ezemvelo Nature Reserve.

24 Figure 5.2.3: Transect line B3 within quadrat B in September 2001 on Ezemvelo Nature Reserve.

Figure 5.3.1: Transect line C1 within quadrat C in September 2001 on Ezemvelo Nature Reserve.

25 Figure 5.3.2: Transect line C2 within quadrat C in September 2001 on Ezemvelo Nature Reserve.

Figure 5.3.3: Transect line C3 within quadrat C in September 2001 on Ezemvelo Nature Reserve.

2.3 Field data collection

2.3.1 Abiotic factors

26 At each quadrat the following data were recorded:

• Topography (crest, mid-slope, foot-slope, riverine);

• Slope in degrees (estimated);

• Aspect (estimated);

• Disturbances (i.e. management road, game path, fire);

• Climate (i.e. rainfall, temperature).

2.3.2. Biotic factors

2.3.2.1 Plants

All trees and shrubs within the quadrats were identified and categorized according to

Order, Family, Species, flower size, phenology, climate and pollination.

Plants that could not be positively identified were forwarded to the National Botanical

Institute in Pretoria for identification. The three quadrats were examined both separately and in combination to determine whether observed differences occurred.

2.3.2.2 Beetle families

2.3.2.2.1 Cerambycidae

The very large family Cerambycidae, generally known as longhorn beetles, contains numerous wood-boring species, but also a fair number that mine the stems and roots of herbaceous and semi woody plants (Skaife 1979). Cerambycids are small to large

(3-100 mm), elongate, cylindrical, sub-cylindrical or flattened beetles with long filiform antennae (Cox 1985). Many species are brightly coloured and hairy. The antennae are usually at least half as long as the body and are often much longer

(especially in males) and are capable of being directed backwards, above and parallel

27 to the body (Cox 1985). Those species occurring on the ground, bark or dead wood are cryptically marked with mottled grey-brown patterns (Cox 1985). Nearly all

Cerambycidae attack trees that are dead or dying, and rarely attack healthy plants.

Savanna ecosystems are often deficient in minerals due to the slow rate of mineralization in the detritus layer due to lack of water (low rainfall) and microbe resistance of leave defence compounds. Wood detritivores, such as beetles, play a vital role in this mineralization process, particularly in the release of phosphorus

(Cowling et al. 1997) This serves in hastening the breakdown of dead wood and the return of nutrients to the soil (Skaife 1979). This family of beetle therefore has a very important role in the functioning of a healthy ecosystem.

2.3.2.2.2 Buprestidae

The Buprestidae, known as jewel beetles, are nearly always metallic or bronzed in colour, some being so beautiful that they are incorporated in jewellery (Skaife 1979).

Buprestids are small to large (1,5-50 mm) torpedo- or wedge-shaped beetles (Holm &

Bellamy 1985). These beetles are very active at the hottest times of the day, but extremely difficult to catch. They often occur on flowers, where some species feed on pollen. Others are found feeding on leaves or bark. These borers gnaw wide galleries between the bark and the sapwood, and often the noise of them chewing is actually audible. Most buprestids attack moribund rather than dead wood, and do not infest seasoned wood (Holm & Bellamy 1985). Usually each species attacks only one or a few genera of plants (Skaife 1979). This family is extremely important from an ecological point of view, as they aid in the process of decomposition in ecosystem functioning.

28 2.3.3 Collection methods

Different sampling methods are required for wood borers due to the variety and habits of the various species. Because all insect capture methods are biased towards catching prey of a certain size, mass, or flight behaviour (Muirhead-Thompson 1991;

Sutherland 1998), a combination of different methods was used.

2.3.3.1 Beating Method (Figure 6.1.1; Figure 6.1.2)

Beetles are collected employing various methods on each quadrat line. The beating method (Holm 1984) is used in the morning and late afternoon, when the insects body temperature is low. This method involves beating the vegetation and capturing insects that are dislodged from foliage (Holm 1984). The beating sheet is made from strong white cloth, square, about 100 cm x 60 cm, with pockets at the corners into which the ends of two diagonal bracing poles were fitted (Fourie 1993). With the poles removed, the beating sheet can be rolled into a neat parcel for easy transport.

29 Figure 6.1.1: The beating method being used on Protea caffra in quadrat C on Ezemvelo Nature Reserve.

Figure 6.1.2: The beating method being used on Acacia caffra in quadrat A on Ezemvelo Nature Reserve.

The vegetation was beaten in a random fashion along the marked transect lines. The individual plants must be identified prior to being beaten and preparation for the capture was taken. A killing bottle (Fourie 1993) was prepared with a label stating information pertaining to the survey. The killing bottle consisted of cotton wool and ethyl acetate to ensure the insects are euthanized quickly and are relaxed (Fourie

1993). Only one or two drops of ethyl acetate were used as excessive amounts could result in condensation on the inner walls of the bottles causing discolouration of specimens (Fourie 1993). The foliage was beaten extensively and inactive insects collected by hand, forceps and an aspirator. The aspirator bottle is a device for collecting small delicate insects individually (Fourie 1993). It can be used to collect insects directly from the beating sheet. An aspirator bottle consists of a bottle (7 cm x

2,5 cm) fitted with a rubber stopper (Fourie 1993). Two holes are drilled through the

30 stopper to take two pieces of hard plastic tubing, each about 7 cm long and about 5 mm in diameter (Fourie 1993). One piece of tubing is pushed through each hole in the stopper, with at least 2 cm showing below and above the stopper. The end of one of the pieces, which is inside the bottle when the stopper is inserted, is covered with a piece of mosquito netting to prevent the insects being sucked into the mouth (Fourie

1993). The air is drawn through the apparatus by sucking on the rubber tube with the mosquito netting and the insects are sucked into the chamber through the other tube, which is pointed towards the insect (Fourie 1993).

2.3.3.2. Sweeping Method (Figure 6.2)

The sweeping method (Holm 1984) should be employed during periods of high temperature. General collecting can be done by sweeping the net back and forth through the foliage (Holm 1984). The insect net is the basic tool of an insect collector

(Fourie 1993). The collecting net used was light and the handle made from a broom handle. The net bag was about 90 cm deep tapered at the bottom as suggested by

Fourie (1993).

31 Figure 6.2: The sweeping method being used on Acacia caffra in quadrat A on Ezemvelo Nature Reserve.

The vegetation is swept in a random fashion along the marked transect lines when insects are active as a result of increased body temperature. The plant to be swept must be identified and a killing bottle prepared with information pertaining to the capture (Fourie 1993). The foliage is then swept from different angles of the plant and the net checked for insect activity. Insects are collected by hand, forceps or an aspirator (Fourie 1993). The best forceps to use are those with prongs that are rounded and with inside surfaces milled (Fourie 1993). The prongs should make contact at the tips only, so that an insect is gripped firmly. The insects in the net are then placed in the labelled killing bottle.

2.3.3.3 Sheet Trap Method (Figure 6.3)

This trap consisted of a light source and a large white sheet placed over the vehicle at night. The sheet was also spread on the ground to catch insects that fall (Fourie 1993).

The light was connected to the vehicle’s battery and a portable generator, where there was no access to electricity.

32 Figure 6.3: The sheet trap method being used in quadrat B on Ezemvelo Nature Reserve in October 2001.

The sheet trap (Holm 1984) is placed at the site just before dark. Insects are attracted to the lamp and settle on the sheet. Insects are collected by hand, forceps or aspirator.

The insects collected from the sheet are placed in a labelled killing bottle (Fourie

1993).

2.3.4 Identification of collected beetles

Beetles are removed from the killing bottles and placed in labelled envelopes with information pertaining to their capture. The specimens collected were pinned before being identified from reference collections and available literature. Most insects are easy to preserve by air drying. Their external skeletons remain intact while their soft internal tissue desiccates (Fourie 1993). Specimens are pinned on a pinning block, which allows insects to be positioned at standard heights on the pin (Fourie 1993). A pinning block can either be a solid block with holes drilled to different depths or a set

33 of steps with a hole drilled through each level (Fourie 1993). It is always advisable to mount insects within twenty four hours of killing, or use a relaxing jar (Fourie 1993).

A relaxing jar is a dampened airtight jar with a drop of ethyl acetate to prevent fungal infection (Fourie 1993).

Specimens which are medium to large in size are pinned with no. 3 or no. 5 pins

(Fourie 1993). The killing jar is emptied onto a piece of white paper, and individually insects are held between the forefinger and the thumb to be pinned. The label containing the capture details was written on white stiff carding and the data printed in Indian ink, using a drafting pen (Fourie 1993).

2.3.5 Data preparation

The identified specimens were cross referenced to the raw data and tabulated monthly, according to relevant quadrat and plant species. These characteristics were then tabulated with corresponding beetle species collected.

2.3.6 Statistical Analysis

2.3.6.1 Chi-square test

Once the data has been tabulated, statistical techniques are required to analyze the data to determine trends and species associations. The most common method of

34 analyzing frequencies is the chi-square test (Fowler et al. 1998). This involves computing a test statistic which is compared with the chi-square (χ²) distribution of a sample variance, s², unlike that of a sample mean in that it is distributed asymmetrically about the population parameter (Fowler et al. 1998). The left-hand side of the distribution is truncated at the minimum value of zero when all observations in a sample by chance have identical values but the right-hand side may in theory extend to infinity (Fowler et al. 1998). This introduces a positive skew to the distribution. The shape of the distribution of a variance depends on the sample size or, more precisely, the degrees of freedom (Fowler et al. 1998). The larger the size of the replicate samples, the more symmetrical becomes the distribution and for very large samples the distribution converges towards normality (Fowler et al. 1998). If we standardize the horizontal axis by multiplying the variance by the degrees of freedom

(df), thus converting it to a sum in squares and then dividing it by the population variance, we generate densities of probability distributions. There is a separate distribution for each possible number of degrees of freedom (Fowler et al. 1998). The required value for a particular number of degrees of freedom is found in tables. This table showing the distribution of χ² is restricted to critical values at the significance levels we are interested in (Fowler et al. 1998).

Chi-square tests are variously referred to as tests for homogeneity, randomness, association, independence and goodness of fit (Fowler et al. 1998). This application involves the underlying principle of comparing the frequencies we observe and the frequencies we expect on the basis of the Null Hypothesis. If the discrepancy between the observed and expected frequencies is great, then the value of calculated test statistic will exceed the critical value at the appropriate degrees of freedom (Fowler et al. 1998). All versions of chi-square test assume that samples are random and

35 observations are independent. The simplest arithmetical comparison that can be made between an observed frequency and an expected frequency is the difference between them (Fowler et al. 1998). In the test, the difference is squared and divided by the expected frequency. The formula:

χ² = ( 0 – E ) ²

E where 0 is an observed frequency and E is an expected frequency. A series of observed frequencies are compared with corresponding expected frequencies resulting in several components of χ² all of which have to be summed (Fowler et al. 1998).

The limitation with the chi-square test is that the sample size, that is the grand total of observed frequencies (n), should be such that all expected frequencies exceed 5

(Fowler et al. 1998). In marginal cases this can sometimes be achieved by collapsing cells and aggregating the respective observed frequencies and expected frequencies

(Fowler et al. 1998). Most statisticians would not object to some of the expected frequencies being below 5, provided that no more than one-fifth of the total number of expected frequencies is below 5, and none are below 1 (Fowler et al. 1998).

2.3.6.2 Cramér’s V analysis

Failing the assumptions of the χ² test, the Phi (coefficient) and Cramér's V,

Contingency coefficient, Lambda (symmetric and asymmetric lambdas and Goodman and Kruskal's tau), and Uncertainty coefficient are the indicated statistical options

36 (Everitt 1993). Contingency coefficient is a measure of association based on chi- square. The value ranges between zero and 1, with zero indicating no association between the row and column variables and values close to 1 indicating a high degree of association between the variables (Everitt 1993). The maximum value possible depends on the number of rows and columns in a table.

Phi is a chi-square based measure of association that involves dividing the chi-square statistic by the sample size and taking the square root of the result. Cramer's V is a measure of association based on chi-square (Everitt 1993), but is less sensitive to the underlying assumptions. Lambda is a measure of association which reflects the proportional reduction in error when values of the independent variable are used to predict values of the outcome. A value of 1 means that the independent variable perfectly predicts the dependent variable. A value of 0 means that the independent variable is no help in predicting the dependent variable (Everitt 1993).

An uncertainty coefficient is a measure of association that indicates the proportional reduction in error when values of one variable are used to predict values of the other variable (Everitt 1993). For example, a value of 0.83 indicates that knowledge of one variable reduces error in predicting values of the other variable by 83%. All analyses were computed using SPSS ¹.

¹ SPSS Inc., 233 S.Wacker Drive, 11th Floor, Chicago, IL 60606

Chapter Three

3. RESULTS

37 Data for 35 species of Cerambycidae (Figures 7.1-7.35), in three subfamilies in three localities on ENR were used in the analysis.

Cerambycidae

Fig. 7.1: Anubis clavicornis Fig. 7.2: Anubis mellyi Fig. 7.3: Anthracocentrus capensis

Fig. 7.4: Ceroplesis thunbergi Fig. 7.5: Coptoeme krantzi Fig. 7.6: Crossotus lacunosus

38

Fig. 7.7: Crossotus plumicornis Fig. 7.8: Crossotus stypticus Fig. 7.9: Hecyra terrea

Fig. 7.10: Hypoeschrus ferreirae Fig. 7.11: Jonthodina sculptilis Fig. 7.12: Lasiopezus longimanus

Fig. 7.13: Macrotoma natala Fig. 7.14: Macrotoma palmate Fig. 7.15: Mycerinicus brevis

39

Fig. 7.16: Nemotragus helvolus Fig. 7.17: Olenecamptus albidus Fig. 7.18: Ossibia fuscata

Fig. 7.19: Pacydissus sp. Fig. 7.20: Phantasis giganteus Fig. 7.21: Philematium natalense

Fig. 7.22: Phryneta spinator Fig. 7.23: Phyllocnema latipes Fig. 7.24: Plocaederus denticornis

40

Fig. 7.25: Dalterus degeeri Fig. 7.26: Dalterus dejeani Fig. 7.27: Prosopocera lactator

Fig. 7.28: Alphitopola octomaculata Fig. 7.29: Taurotagus klugi Fig. 7.30: Tithoes maculates

Fig. 7.31: Tragiscoschema bertolinii Fig. 7.32: Xystrocera erosa Fig. 7.33: Xystrocera dispar

41

Fig.7.34: Zamium bimaculatum Fig. 7.35: Zamium incultum

There are four species of buprestid and one species of cerambycid that could not be identified to species level and have been sent to the United States of America for identification purposes (Table 1.1).

Table 1: List of species collected at Ezemvelo Nature Reserve not identified to species level. Tota Species l Anthaxia sp. 1 89 Anthaxia sp. 2 102 Anthaxia sp. 3 64 Anthaxia sp. 4 3 Pacydissus sp. 12 Total 270

Both the density of beetle species (the number of species per area) and species richness (the number of species present per number of individuals) (Hurlbert 1971;

Gotelli & Colwell 2001; Magurran 2004) were assessed for samples from the plants.

Estimates of the total number of species for Cerambycidae and Buprestidae were made from sampling.

42 Identification were obtained from specimens in the *Transvaal Museum, South Africa

TMSA².

3.1 Cerambycidae Data

3.1.1 Total Cerambycidae Data

The total number and species diversity of Cerambycidae collected in all three quadrats for the year 2001 totalled 518 specimens and 35 species respectively. Three subfamilies were recorded during this study, namely Cerambycinae (15 species);

Prioninae (4 species) and (6 species). A total of 28 genera were recorded throughout the year, namely Zamium (2 species), Captoeme (1 species), Taurotagus

(1 species), Jonthodina (1 species), Anubis (2 species), Macrotoma (2 species),

Tithoes (1 species), Phantasis (1 species), Dalterus (2 species), Crossotus (3 species),

Olenecamptus (1 species), Anthracocentrus (1 species), Hypoeschrus (1 species),

Plocaederus (1 species), Nemotragus (1 species), Hecyra (1 species), Ceroplesis (1 species), Lasiopezus (1 species), Philematium (1 species), Alphitopola (1 species),

Mycerinicus (1 species), Phryneta (1 species), Tragiscoschema (1 species),

Phyllocnema (1 species), Pacydissus (1 species), Ossibia (1 species), Xystrocera (2 species) and Prosopocera (1 species) (Appendix A).

Cerambycidae were collected at monthly intervals throughout the year. October,

November, December and January were months with the highest activity and species diversity (Fig. 8).

43 * TMSA² Transvaal Museum Coleoptera Department, P.O. Box, Pretoria,

0010

25

20

15 TOTAL 10 CERAMBYCIDAE

5

0 JAN MAR MAY JUL SEP NOV

Figure 8: Monthly sampling frequencies for the total Cerambycidae collected for

2001 on Ezemvelo Nature Reserve.

There is a stronger correlation (r² = 0.60) between total cerambycid numbers and minimum temperature, than the correlation (r² = 0.37) between total cerambycid numbers and maximum temperature (Table 2).

Table 2: Analysis of regression indicating the degree of linear trend parameters for

Cerambycidae on Ezemvelo Nature Reserve.

Minimum Maximum Average Sites Temperature Temperature Rainfall Cerambycidae QA r² = 0.67 r² = 0.47 r² = 0.63 Cerambycidae QB r² = 0.47 r² = 0.20 r² = 0.61 Cerambycidae QC r² = 0.56 r² = 0.37 r² = 0.48 Total Quadrats r² = 0.60 r² = 0.37 r² = 0.37

44 There is a stronger correlation (r² = 0.98) between total cerambycid numbers and summer months January and February, than the correlation (r² = 0.69) between total cerambycid numbers and the winter month of June (Table 3).

Table 3: Analysis of regression indicating the degree of linear trend for the first six months of the year for Cerambycidae on Ezemvelo Nature Reserve.

Months Jan Feb Mar Apri May Jun Cerambycidae QA r² = 0.95 r² = 0.96 r² = 0.86 r² =0.81 r² = 0.49 r² = 1 Cerambycidae QB r² = 0.88 r² = 0.86 r² = 0.85 r² = 1 r² = 0.78 r² = 1 Cerambycidae QC r² = 0.95 r² = 0.90 r² = 0.91 r² = 0.84 r² = 0.76 r² = 0.68 Total Quadrats r² = 0.98 r² = 0.98 r² = 0.96 r² = 0.92 r² = 0.89 r² = 0.69

There is a correlation (r² = 0.99) between total cerambycid numbers and summer months November and December. However the correlation (r² = 1) between total cerambycid numbers and the winter month of July is an exact correlation (Table 4).

Table 4: Analysis of regression indicating the degree of linear trend for the last six months of the year for Cerambycidae on Ezemvelo Nature Reserve.

Months Jul Aug Sep Oct Nov Dec Cerambycidae QA r² = 1 r² = 0.81 r² = 0.93 r² = 0.97 r² = 0.98 r² = 0.98 Cerambycidae QB r² = 1 r² = 0.67 r² = 0.84 r² = 0.90 r² = 0.96 r² = 0.97 Cerambycidae QC r² = 1 r² = 0.47 r² = 0.93 r² = 0.93 r² = 0.98 r² = 0.98 Total Quadrats r² = 1 r² =0.87 r² = 0.98 r² = 0.98 r² = 0.99 r² = 0.99

45 Collection results for months June and July indicate almost no cerambycid activity during this period.

The results indicate that environmental conditions associated with seasonality control the abundance and diversity of cerambycids (Fig. 9.1, Fig. 9.2, Fig. 9.3). The greater percentage of cerambycids were collected in the summer months, December (23%),

November (22%), October (14%), January (13%) and February (10%) (Appendix B).

(Fig. 9.1).

25

20

15 Average M inimim Temperature Beetle Numbers

Numbers 10

5

Average Minimum Temperature /Beetle /Beetle Temperature Minimum Average 0 123456789101112 Months

Figure 9.1: Total Cerambycidae count trend versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve for January 2001 to December 2001.

There is a correlation (r² = 0.60) between the numbers of cerambycids collected and average minimum temperature. This indicates that due to temperatures dropping

46 exceptionally low in winter, these conditions directly affect cerambycid populations and the adult population dies off during this period.

l 30

25

20 Average Maximum 15 Temperature Beetle Numbers Numbers 10

5

Average Maximum Temperature/Beet Average Maximum 0 1 2 3 4 5 6 7 8 9 10 11 12 Months Figure 9.2: Total Cerambycidae count trend versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve for January 2001 to December 2001.

Figure 9.2 indicates that cerambycid numbers and the average maximum temperatures do not correlate (r² = 37), other but not to the same extent as with average minimum temperature.

This would indicate that low temperatures have a greater affect on the cerambycid population dynamics on ENR than high temperatures.

47 160

140

120

100 Average Rainfall 80 Beetle Numbers 60 Average Rainfall Average 40

20

0 123456789101112 Months

Figure 9.3: Total Cerambycidae count trend versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve for January 2001 to December 2001.

Cerambycid numbers indicate a small correlation (r² = 0.37) with average rainfall, numbers decreasing with a decrease in rainfall, while increasing with spring rains

(Fig. 9.3). This relationship does not appear as defined as with temperature.

3.1.2 Quadrat A

The total number and species diversity of Cerambycidae collected in quadrat A for

2001 equalled 203 specimens and 22 species respectively. All three subfamilies were collected in quadrat A, Cerambycinae (8 species), Prioninae (3 species) and Lamiinae

48 (11 species). 19 Genera were recorded in quadrat A, Zamium (1 species), Captoeme

(1 species), Taurotagus (1 species), Jonthodina (1 species), Anubis (1 species),

Macrotoma (1 species), Tithoes (1 species), Phantasis (1 species), Dalterus (2 species), Crossotus (3 species), Olenecamptus (1 species), Anthracocentrus (1 species), Hypoeschrus (1 species), Plocaederus (1 species), Nemotragus (1 species),

Hecyra (1 species), Ceroplesis (1 species), Lasiopezus (1 species) and Philematium (1 species). 13 species were not recorded in quadrat A, Zamium bimaculatum, Anubis mellyi, Alphitopola octomaculata, Mycerinicus brevis, Phryneta spinator,

Tragiscoschema bertolinii, Phyllocnema latipes, Pacydissus sp., Macrotoma natala,

Ossibia fuscata, Xystrocera erosa, Prosopocera lactator and Xystrocera dispar

(Appendix C).

Cerambycids collected in quadrat A follow the same general trend with high beetle numbers being recorded during the summer months with gradual decline towards the winter months (Fig 10). The greatest percentages of cerambycids collected in quadrat

A were collected in the summer months, December & November (20%), October

(17%), January and February (13%) and March and September (6%) (Appendix D).

20

15

10 QUADRAT A

5

0 JAN MAR MAY JUL SEP NOV

49 Figure 10: Monthly sampling frequencies for Cerambycidae collected in

quadrat A for 2001 on Ezemvelo Nature Reserve.

Cerambycid numbers decreased drastically in quadrat A during the winter period.

When average minimum temperatures dropped below 5 degrees, Cerambycid numbers reached zero (Fig. 11.1). This indicates a strong correlation ((r² = 0.67) to minimum temperature.

25

20

15 Average M inimum Temperature Beetle Numbers

Numbers 10

5

Average Minimum Temperature/Beetle Temperature/Beetle Minimum Average 0 123456789101112 Months

Figure 11.1: Cerambycidae count trend in quadrat A versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Cerambycid numbers correlate moderately (r² = 0.47) to average maximum temperature in quadrat A, with numbers increasing with increases in temperature and decreasing with lower average maximum temperatures (Fig. 11.2).

50 30

25

20 Average M aximum Temperature 15 Beetle Numbers Numbers 10

5

Average Maximum Temperature/Beetle Temperature/Beetle Maximum Average 0 123456789101112 Months

Figure 11.2: Cerambycidae count trend in quadrat A versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Minimum temperature fluctuations appear to be a greater limiting factor than maximum temperature fluctuations in quadrat A.

Results indicate that there is a gradual increase in cerambycid numbers with an increase in rainfall in quadrat A (Fig. 11.3). There was, however, very little rainfall in

December, yet cerambycid numbers were not affected negatively. Overall there is, however, a correlation (r² = 0.63) with average rainfall in quadrat A.

51 160

140

120

100 Average Rainfall 80 Beetle Numbers 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months

Figure 11.3: Cerambycidae count trend in quadrat A versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001and December 2001.

3.1.3 Quadrat B

The total number and species diversity of Cerambycidae collected in quadrat B for

2001 equalled 116 specimens and 20 species respectively. Three subfamilies were recorded in quadrat B, i.e. Cerambycinae (9 species), Prioninae (3 species) and

Lamiinae (8 species). 17 genera were recorded within these Subfamilies, Zamium (1 species), Jonthodina (1 species), Anubis (2 species), Macrotoma (2 species), Tithoes

(1 species), Alphitopola (1 species), Crossotus (2 species), Mycerinicus (1 species),

Phryneta (1 species), Dalterus (1 species), Ceroplesis (1 species), Tragiscoschema (1 species), Phyllocnema (1 species), Pacydissus (1 species), Coptoeme (1 species),

Taurotagus (1 species) and Philematium (1 species). 15 species were absent from

52 quadrat B, Zamium incultum, Phantasis giganteus, Olenecamptus albidus,

Anthracocentrus capensis, Hypoeschrus ferreirae, Plocaederus denticornis,

Crossotus stypticus, Nemotragus helvolus, Hecyra terrea, Lasiopezus longimanus,

Dalterus degeeri, Ossibia fuscata, Xystrocera erosa, Prosopocera lactator, and

Xystrocera dispar (Appendix E).

Cerambycid numbers in quadrat B indicate an increase in Cerambycid numbers at the beginning of November into December (Fig. 12). The greatest percentage of cerambycids collected in quadrat B were collected in the summer months, i.e.

November (29%), December (26%), January and October (10%) and February (9%)

(Appendix F).

Cerambycid abundance and diversity in quadrat B appears to decline earlier than in quadrat A, with numbers starting to decline in January.

53 30

25

20

Figure15 12: Monthly sampling frequencies for Cerambycidae collected in quadrat B QUADRAT B 10 for 2001 on Ezemvelo Nature Reserve.

5

0 CerambycidJAN numbers MAR are MAY directly JUL corre SEPlated (r² NOV = 0.47) with average minimum temperature in quadrat B, showing a gradual decrease in numbers and diversity with low minimum temperatures (Fig 13.1). Cerambycid populations appear to follow the trend, and even temporary increases in temperature do affect the cerambycid population. The population starts increasing when temperatures are more stable.

54 35

30

25 Average M inimum 20 Temperature

15 Beetle Numbers Numbers

10

5

Average Minimum Temperature/ Beetle Beetle Temperature/ Minimum Average 0 123456789101112 Months Figure 13.1: Cerambycidae count trend in quadrat B versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Cerambycid numbers show little correlatation (r² = 0.20) with average maximum temperatures in quadrat B, and not as clear a trend as average minimum temperatures

(Fig. 13.2). Cerambycid populations decrease slowly after the decrease in temperature. Figure 13.2 indicates a decrease in temperature in November, resulting in a decrease in beetle numbers in December. Cerambycidae count in quadrat B versus average monthly rainfall for the months of the year at Ezemvelo Nature

Reserve fluctuated considerable and indicated a delayed response to rainfall (Fig.

13.3).

55 35

30

25 Average M aximum 20 Temperature

15 Beetle Numbers Numbers

10

5

Average Maximum Temperature/Beetle 0 123456789101112 Months

Figure 13.2: Cerambycidae count trend in quadrat B versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

160

140

120

100 Average Rainfall 80 Months 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months

Figure 13.3: Cerambycidae count trend in quadrat B versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

56 3.1.4 Quadrat C

The total number and species diversity of Cerambycidae collected in quadrat C for

2001 equalled 199 specimens and 26 species respectively. Three subfamilies were represented in quadrat C, Cerambycinae (13 species), Prioninae (4 species) and

Lamiinae (9 species). 20 genera were recorded within these subfamilies in quadrat C,

Nemotragus (1 species), Crossotus (3 species), Tragiscoschema (1 species),

Anthracocentrus (1 species), Anubis (2 species), Taurotagus (1 species), Coptoeme (1 species), Zamium (2 species), Xystrocera (2 species), Pacydissus (1 species),

Philematium (1 species), Jonthodina (1 species), Macrotoma (1 species), Tithoes (1 species), Phantasis (1 species), Ceroplesis (1 species), Olenecamptus (1 species) and

Prosopocera (1 species). Nine species were not collected in quadrat C, Dalterus dejeani, Hypoeschrus ferreirae, Plocaederus denticornis, Hecyra terrea, Lasiopezus longimanus, Dalterus degeeri, Alphitopola octomaculata, Mycerinicus brevis and

Phryneta spinator (Appendix G).

Cerambycid samples recorded in quadrat C indicate a clear decline from January to

July, with an gradual increase in numbers from August to December (Fig. 14).

November and December clearly are the most environmentally suitable months for adult cerambycid activity. The greatest percentage of cerambycids collected in quadrat

C was in the summer months, December (25%), November (21%), January (14%) and

October (13%) (Appendix H).

57 25

20

15 QUADRAT C 10

5

0 JAN MAR MAY JUL SEP NOV

Figure 14: Monthly sampling frequencies for Cerambycidae collected in

quadrat C for 2001 on Ezemvelo Nature Reserve.

Cerambycid numbers correlate positively with average minimum temperatures in quadrat C (Fig. 15.1). The population appears to decline gradually until temperatures fall below 5 degrees, where there is almost zero activity. The population starts increasing rapidly from September through to December. 30

25

20 Average Minimum Temperature 15 Beetle Numbers

10

5 Average Minimum Temperature Minimum Average

0 123456789101112 Months

Figure 15.1: Cerambycidae count trend in quadrat C versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001and December 2001.

58 Cerambycid numbers correlate moderately to average maximum temperature in quadrat C (Fig. 15.2). The population is controlled to a greater extent by average minimum temperatures.

30

25

20 Average M aximum Temperature 15 Beetle Numbers

10

5 Average Maximum Temperature Maximum Average

0 123456789101112 Months

Figure 15.2: Cerambycidae count trend in quadrat C versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001and December 2001.

Cerambycid numbers are moderately affected by average rainfall, however, this population fluctuation appears to be delayed with an increase or decrease in average rainfall (Fig. 15.3).

59 160

140

120

100 Average Rainfall 80 Beetle Numbers 60 Average Rainfall Average 40

20

0 123456789101112 Months Figure 15.3: Cerambycidae count trend in quadrat C versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Cerambycid samples appear similar between the three quadrats, even though the vegetation in each quadrat differs quite significantly (Fig. 16). Quadrat A had equal numbers collected in November and December. Quadrat B had the highest number of beetles collected in November. Quadrat C had highest sample size recorded in

December. Cerambycid numbers within all quadrats declined to zero between June and July, except for a few records in quadrat C 30

25

20 QUADRAT A 15 QUADRAT B 10 QUADRAT C

5

0 JAN MAR MAY JUL SEP NOV

60 Figure 16: Monthly sampling frequencies of Cerambycidae between the three quadrats on Ezemvelo Nature Reserve.

3.2 Buprestidae data

3.2.1 Total Buprestidae data

Data for 32 species of Buprestidae (Figures 17.1-17.21) in eight subfamilies, 43 genera and three localities on ENR were used in the analysis.

The following buprestid species were not photographed Anthaxia bergrothi; Anthaxia sp. 2; Anthaxia sp. 3; Anthaxia sp. 4; Trachys ziziphusii; Brachelytrium transvalense;

Kamosia tenebricosa; Agrilomorpha venosa; Agrilus falcatus; Kamosiella dermestoides; Anthaxia obtectans

Fig. 17.1: Acmaeodera aenea Fig. 17.2: Acmaeodera albivillosa Fig. 17.3: Agrilus guerryi

61

Fig. 17.4: Sphenoptera sinuosa Fig. 17.5: Acmaeodera punctatissima Fig. 17.6: Acmaeodera inscripta

Fig. 17.7: Acmaeodera ruficaudis Fig. 17.8: Agrilus sexguttatus Fig. 17.9: Acmaeodera stellata

Fig. 17.10: Acmaeodera viridiaenea Fig. 17.11: Anthaxia. sp. 1 Fig. 17.12: Chrysobothris algoensis

62

Fig.17.13:Chrysobothris boschismanni Fig 17.14:Chrysobothris dorsata Fig.17.15:Evides pubiventris

Fig. 17.16: Lampetis gregaris Fig. 17.17: Phlocteis exasperata Fig. 17.18: Pseudagrilus beryllinus

Fig. 17.19: Lampetis conturbata Fig. 17.20: Sphenoptera arrowi Fig. 17.21: Sternocera orissa

63 The total number and species of Buprestidae collected in all quadrats for 2001 equalled 805 specimens and 32 species respectively. Five subfamilies were recorded in this study, Polycestinae (12 species), Buprestinae (7 species), Julodinae (1 species),

Agrilinae (9 species) and Chalcophorinae (3 species). 15 genera within these subfamilies were recorded, Acmaeodera (7 species), Sternocera (1 species), Anthaxia

(6 species), Agrilus (3 species), Lampetis (2 species), Chrysobothris (3 species),

Trachys (1 species), Pseudagrilus (1 species), Brachelytrium (1 species), Kamosia (1 species), Sphenoptera (2 species), Agrilomorpha (1 species), Kamosiella (1 species),

Phlocteis (1 species) and Evides (1 species) (Appendix I).

There is a stronger correlation (r² = 0.73) between total buprestid numbers and minimum temperature, than the correlation (r² = 0.52) between total buprestid numbers and maximum temperature (Table 5).

Table 5: Analysis of regression indicating the degree of linear trend of parameters for

Buprestidae on Ezemvelo Nature Reserve.

Minimum Maximum Average Sites Temperature Temperature Rainfall Buprestidae QA r² = 0.77 r² = 0.55 r² = 0.35 Buprestidae QB r² = 0.59 r² = 0.40 r² = 0.24 Buprestidae QC r² = 0.70 r² = 0.51 r² = 0.36 Total Quadrats r² = 0.73 r² = 0.52 r² = 0.34

64 There is a stronger correlation (r² = 0.99) between total buprestid numbers and summer months January, February, March than the correlation (r² = 0.95) between total buprestid numbers and Autumn months April and May (Table 6).

Table 6: Analysis of regression indicating the degree of linear trend for the first six months of the year for Buprestidae on Ezemvelo Nature Reserve.

Months Jan Feb Mar Apri May Jun Buprestidae QA r² = 0.99 r² = 0.99 r² = 0.98 r² = 0.94 r² = 0.87 r² = 0.71 Buprestidae QB r² = 0.98 r² = 0.95 r² = 0.98 r² = 0.79 r² = 0.81 r² = 1 Buprestidae QC r² = 0.98 r² = 0.98 r² = 0.97 r² = 0.86 r² = 0.75 r² = 1 Total Quadrats r² = 0.99 r² = 0.99 r² = 0.99 r² = 0.95 r² = 0.95 r² = 0.69

There is a stronger correlation (r² = 0.99) between total buprestid numbers and summer months October, November, December, than the correlation (r² = 0.47) between total buprestid numbers and winter month of July (Table 7).

Table 7: Analysis of regression indicating the degree of linear trend for the last six months of the year for Buprestidae on Ezemvelo Nature Reserve.

Months Jul Aug Sep Oct Nov Dec Buprestidae QA r² = 0.49 r² = 0.78 r² = 0.96 r² =0.97 r² = 0.98 r² = 0.99 Buprestidae QB r² = 1 r² = 1 r² = 0.65 r² = 0.95 r² = 0.97 r² = 0.98 Buprestidae QC r² = 1 r² = 1 r² = 0.75 r² = 0.97 r² = 0.99 r² = 0.99 Total Quadrats r² = 0.47 r² = 0.75 r² = 0.96 r² = 0.99 r² = 0.99 r² = 0.99

65 Buprestidae were collected at monthly intervals throughout the year. November,

December and January were months with the highest activity and species diversity

(Fig. 18). The greater percentage of buprestids were collected in the summer months,

December (24%), January (19%), November (16%) and February (13%) (Appendix

J). Collection results for months June and July indicates no buprestid activity during this period.

25

20

15 TOTAL 10 BUPRESTIDAE

5

0 JAN MAR MAY JUL SEP NOV

Figure 18: Monthly sampling frequencies for the total Buprestidae

collected for 2001 on Ezemvelo Nature Reserve.

The results indicate that environmental conditions associated with seasonality control the abundance and diversity of buprestids. This is shown in Fig.19.1, Fig.19.2; Fig.

19.3.

66 There appear to be a direct relationship between the number of buprestids collected and average minimum temperature (Fig. 19.1). This indicates that due to temperatures dropping exceptionally low in winter, these conditions directly affect buprestid population dynamics and adult populations die off during this period

30

25

20 Average M inimum Temperature 15 Beetle Numbers Numbers 10

5

Average Minimum Temperature/Beetle 0 123456789101112 Months

Figure 19.1: Total Buprestidae count versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Figure 19.2 indicates that buprestidae numbers and the average maximum temperatures correlate with each other, but not to the same extent as with average minimum temperatures.

This would indicate that low temperatures have a greater affect on buprestid abundance on ENR than high temperatures.

67 30

25

20 Average M aximum Temperature 15 Beetle Numbers Numbers 10

5

Average Maximum Temperature/Beetle Temperature/Beetle Maximum Average 0 123456789101112 Months

Figure 19.2: Total Buprestidae count versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Buprestids numbers indicate a slight correlation with average rainfall, with numbers decreasing with a decrease in rainfall, while increasing with spring rains (Fig. 19.3).

This relationship does not appear as defined as with temperature.

68 160

140

120

100 Average Rainfall 80 Beetle Numbers 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months Figure 19.3: Total Buprestidae count versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

3.2.2 Quadrat A

The total number and species of Buprestidae collected in quadrat A for 2001 equalled

379 specimens and 24 species respectively. Five subfamilies were recorded in quadrat

A, Polycestinae (6 species), Julodinae (1 species), Buprestinae (9 species), Agrilinae

(7 species) and Chalcophorinae (1 species). 12 genera were recorded within these subfamilies in quadrat A, Acmaeodera (6 species); Sternocera (1 species); Anthaxia

(4 species), Agrilus (3 species), Chrysobothris (3 species), Sphenoptera (1 species),

Trachys (1 species), Kamosia (1 species), Agrilomorpha (1 species), Lampetis (1 species), Pseudogrilus (1 species) and Brachelytrium (1 species) (Appendix K). Eight species were absent from quadrat A, Kamosiella dermestoides, Acmaeodera stellata;

Phlocteis exasperata, Psiloptera conturbata, Evides pubiventrus, Sphenoptera sinuosa, Anthaxia obtectans and Anthaxia sp. 4.

Buprestids collected in quadrat A follow a similar trend with high overall beetle numbers being recorded during the summer months, December (21%), January

69 (18%), November (15%) and February (14%) (Appendix L), with gradual decline 25 towards the winter months (Fig 20). 20

15 QUADRAT A 10

5

0 JAN MAR MAY JUL SEP NOV

Figure 20: Monthly sampling frequencies for Buprestidae collected in

quadrat A for 2001 at Ezemvelo Nature Reserve.

Buprestid numbers decreased drastically in quadrat A during the winter period and where average minimum temperatures dropped below 5 degrees, buprestid numbers reached zero (Fig. 21.1).

25

20

15 Average M inimum Temperature Beetle Numbers

Numbers 10

5

Average Minimum Temperature/Beetle 0 123456789101112 Months

70 Figure 21.1: Buprestidae count trend in quadrat A versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Buprestid numbers correlate moderately to average maximum temperature in quadrat

A, numbers increasing with increases in temperature and decreasing with lower average maximum temperatures (Fig. 21.2). Minimum temperature fluctuations appear to be a greater limiting factor than maximum temperature fluctuations in quadrat A.

30

25

20 Average M aximum Temperature 15 Beetle Numbers Numbers 10

5

Average Maximum Temperature/Beetle 0 123456789101112 Months

Figure 21.2: Buprestidae count trend in quadrat A versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001and December 2001.

71 Results indicate that there is a gradual increase in buprestid numbers with an increase in rainfall in quadrat A (Fig. 21.3). There was however very little rainfall in

December, yet buprestid numbers were not affected negatively.

160

140

120

100 Average Rainfall 80 Beetle Numbers 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months

Figure 21.3: Buprestidae count trend in quadrat A versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

3.2.3 Quadrat B

The total number and species of Buprestidae collected in quadrat B for 2001 equalled

153 specimens and 11 species respectively. Four subfamilies were recorded in quadrat

B, Polycestinae (4 species), Buprestinae (5 species), Julodinae (1 species) and

Agrilinae (1 species). Five genera were represented within these subfamilies,

Acmaeodera (4 species), Anthaxia (3 species), Chrysobothris (2 species), Sternocera

(1 species) and Pseudogrilus (1 species) (Appendix M). 21 species were not recorded in quadrat B, Acmaeodera viridiaenea, Anthaxia bergrothi, Agrilus guerryi, Lampetis gregaris, Agrilus sexguttatus, Trachys ziziphusii, Brachelytrium transvalense,

72 Chrysobothris dorsata, Acmaeodera punctatissima, Kamosia tenebricosa,

Agrilomorpha venosa, Agrilus falcatus, Sphenoptera arrowi, Kamosiella dermestoides, Acmaeodera stellata, Phlocteis exasperata, Lampetis conturbata,

Evides pubiventris, Sphenoptera sinuosa, Anthaxia obtectans and Anthaxia sp. 4.

Buprestidae samples from quadrat B indicate a drastic increase in buprestid numbers at the beginning of December (Fig. 22). Buprestid abundance and diversity in quadrat

B appears to decline gradually from March, with no samples collected during June,

July or August. The greater percentage of buprestids collected in quadrat B were collected in summer, December (32%), January (17%), November (14%) and

February (12%) (Appendix N).

35 30 25 20 15 QUADRAT B 10 5 0 FigureJAN MAR22: Monthly MAY sampling JUL SEPfrequencies NOV for Buprestidae collected

in quadrat B for 2001 at Ezemvelo Nature Reserve.

73 Buprestid numbers indicate a correlation (r² = 0.59) with average minimum temperature in quadrat B, showing a gradual decrease in numbers and diversity with low minimum temperatures (Fig 23.1). Buprestid populations appear to follow the trend, and even temporary fluctuations in temperature do not affect the population numbers. The population starts increasing when temperatures increases are more stable.

35

30

25 Average M inimum 20 Temperature

15 Beetle Numbers Numbers

10

5

Average Minimum Temperature/Beetle 0 123456789101112 Months

Figure 23.1: Buprestidae count trend in quadrat B versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001and December 2001.

Buprestidae numbers do not correlate (r² = 0.40) with average maximum temperatures in quadrat B, where as average minimum temperatures (Fig. 23.1) had a greater influence. Buprestid populations decrease gradually after the decrease in temperature.

Figure 23.2 indicates a decrease in temperature in November, resulting in no decrease in beetle numbers in December.

74 35

30

25 Average M aximum 20 Temperature

15 Beetle Numbers Numbers

10

5

Average Maximum Temperature/Beetle 0 1 2 3 4 5 6 7 8 9 10 11 12 Months

Figure 23.2: Buprestidae count trend in quadrat B versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Buprestidae numbers correlate negatively (r² = 0.24) with average rainfall in quadrat

B, however this population fluctuation appears to be delayed with an increase or decrease in average rainfall (Fig. 23.3).

75 160

140

120

100 Average Rainfall 80 Beetle Numbers 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months

Figure 23.3: Buprestidae count trend in quadrat C versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

3.2.4 Quadrat C

The total number and species of Buprestidae collected in quadrat C for 2001 equalled

273 specimens and 19 species respectively. Five subfamilies were recorded in quadrat

C, Polycestinae (3 species), Buprestinae (9 species), Julodinae (1 species), Agrilinae

(3 species) and Chalcophorinae (3 species). Ten genera were represented within these subfamilies, Acmaeodera (3 species), Chrysobothris (2 species), Anthaxia (5 species),

Sphenoptera (2 species), Lampetis (2 species), Sternocera (1 species), Pseudagrilus

(1 species), Phlocteis (1 species), Evides (1 species) and Kamosiella (1 species)

(Appendix 0). 13 species were absent from quadrat C, Acmaeodera albivillosa,

Acmaeodera viridiaenea, Acmaeodera ruficaudis, Agrilus guerryi, Agrilus sexguttatus, Anthaxia sp. 3, Trachys ziziphusii, Brachelytrium transvalense,

Chrysobothris dorsata, Acmaeodera punctatissima, Kamosia tenebricosa,

Agrilomorpha venosa and Agrilus falcatus.

76 Buprestid samples recorded in quadrat C indicate a drastic decline from March to

April, with a rapid increase in numbers from September to December (Fig. 24).

November, December and January clearly are the most environmentally suitable months for buprestid activity. The greater percentage of buprestids collected in quadrat C were collected in the summer months, December (25%), January (19%),

November (18%), March (13%) and February (12%) (Appendix P).

25

20

15 QUADRAT C 10

5

0 JAN MAR MAY JUL SEP NOV

Figure 24: Monthly sampling frequensies for Buprestidae collected in

quadrat C for 2001 on Ezemvelo Nature Reserve.

Buprestid numbers correlate (r² = 0.70) positively with average minimum temperatures in quadrat C (Fig. 25.1). The population appears to decline gradually until temperatures fall below 5, degrees where there is almost zero activity (Fig. 25.1).

The population starts increasing rapidly from September through to December.

77 30

25

20 Average M inimum Temperature 15 Beetle Numbers Numbers 10

5

Average Minimum Temperature/Beetle 0 123456789101112 Months Figure 25.1: Buprestidae count trend in quadrat C versus average minimum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Buprestid numbers correlate (r² = 0.51) moderately to average maximum temperature in quadrat C (Fig. 25.2). It is expected that the population is controlled to a greater extent by average minimum temperatures.

30

25

20 Average M aximum Temperature 15 Beetle Numbers Numbers 10

5

Average MaximumTemperature/Beetle 0 123456789101112 Months

Figure 25.2: Buprestidae count trend in quadrat C versus average maximum temperature for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

78 Buprestid numbers do not correlate (r² = 0.36) with average rainfall, however this population fluctuation appears delayed with an increase or decrease in average rainfall

(Fig. 25.3).

160

140

120

100 Average Rainfall 80 Beetle Numbers 60

40

20 Average Rainfall/Beetle Numbers 0 123456789101112 Months

Figure 25.3: Buprestidae count trend in quadrat C versus average monthly rainfall for the months of the year at Ezemvelo Nature Reserve between January 2001 and December 2001.

Buprestid samples appear similar between the three quadrats, even though the vegetation in each quadrat differs quite significantly (Fig. 26). Quadrat A, Quadrat B and Quadrat C had highest sample size recorded in December. Buprestid numbers within all quadrats declined to zero between June, July and August, except for a few records in quadrat A.

79 35 30 25 20 QUADRAT A 15 QUADRAT B QUADRAT C 10 5 0 JAN MAR MAY JUL SEP NOV

Figure 26: Monthly sampling frequencies of Buprestidae between the three quadrats on Ezemvelo Nature Reserve.

3.3 Ecological processes affecting Cerambycidae and Bupresridae on ENR

Clearly the ecological processes that determine the population structure of both

Cerambycidae and Buprestidae on ENR have a similar affect on both families (Fig.

27). Buprestidae appear to decline later as winter approaches, yet increase slower as summer approaches.

Monthly Differences between Buprestidae and Cerambycidae

25

20

15 Betle Numbers 10 Buprestidae Cerambycidae 5

0 JAN MAR MAY JUL SEP NOV Months

Figure 27: Monthly sample comparison between Buprestidae and Cerambycidae on

Ezemvelo Nature Reserve between January 2001 and December 2001.

80 3.4 Cerambycidae/Plant correlations

The plants surveyed were further categorized according to order, family, species, phenology, pollination type, flower size and climate. Corresponding beetle diversity and abundance was calculated respectively. One method of assessing community structure involves grouping species based on feeding perference (Root 1973).

In addition, some groups are thought to have a very close association with plant taxa on which they oviposit and feed (Gussmann 1994).

3.4.1 Plant order correlation

Statistically there is a relationship between tree order and cerambycid species on ENR

(Cramér’s V = 0.560; p < 0.05; N = 253). The majority of specimens collected were found on trees of the Order Fabales (Appendix Q) (Fig. 28.1).

Celastrales

Ericales Cerambycidae Rosales

Fabales 0 20406080

Figure 28.1: Relationship between tree orders and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

81 The breakdown of the percentage of cerambycids were collected on Fabales (74%),

Rosales (11%) and Sapindales (5%) (Appendix R) (Fig. 28.1).

3.4.2 Plant family correlation

In all quadrats, cerambycid abundance and diversity was greatest on Mimosaceae, which include Acacia caffra and Acacia karoo. Certain plant families had zero species recorded (Appendix S). The greater percentage of cerambycids were collected on Mimosaceae (65%), Rhamnaceae (8%) and Papilionaceae (7%) (Appendix T) (Fig.

28.2).

Olacaceae

Poaceae

Rhamnaceae Cerambycidae Papilionaceae

Mimosaceae 0 20406080

Figure 28.2: Relationship between Tree Family and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

82 3.4.3 Plant species correlation

Statistically there is a relationship between plant species and cerambycid species on

ENR (Cramér’s V = 0.477; p < 0.05; N=243). The majority of specimens collected were found on Acacia species (Appendix U). The greater percentage of cerambycids were collected on Acacia karoo (42%), Acacia caffra (23%), Combretum molle (8%), and Burkea africana (7%) (Appendix V) (Fig. 28.3).

Croton grattismus

Ziziphus Cerambycidae mucronata

Acacia caffra 0 1020304050

Figure 28.3: Relationship between plant species and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

Plants certainly do have some importance as hosts to certain species, with certain plants appearing to have a high attraction to a broad range of species. Certain other species of plants possibly due to high concentrations of browse resistant chemicals, are very unattractive to cerambycids.

83 3.4.4 Plant flower size correlation

Statistically there is a slight relationship between flower size and cerambycid species on ENR (Cramér’s V = 0.267; p < 0.05; N= 506). The majority of specimens collected were found on small flowers (Appendix AC).

The greater percentage of cerambycids were collected on plants with small flowers

(87%), with plants with medium-large flowers representing (13%) (Appendix AD)

(Fig. 28.4).

Cerambycidae Medium Large

Small

0 20406080100

Figure 28.4: Relationship between flower size and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

3.4.5 Plant Phenology

Statistically there is no relationship between plant phenology and cerambycid species on ENR [Cramér’s V = 0.483; 0.002 < 0.05); (p > 0.05)]. The majority of specimens

84 collected were found on deciduous plants (Appendix W). The greater percentage of cerambycids were collected on deciduous plants (83%), with fewer specimens being

collected on non-deciduous plants (17%) (Appendix X). The reason for this is the majority of plants on ENR are deciduous plants (Fig. 28.5).

NO YES

Cerambycidae

0 20 40 60 80 100

Figure 28.5: Relationship between plant phenology and Cerambycidae

species collected on Ezemvelo Nature Reserve between

January 2001 and December 2001.

3.4.6 Plant Pollination

Statistically there is a small relationship between plant pollination and cerambycid species on ENR [(Cramér’s V = 0.436; p < 0.05); (N= 253)]. The majority of specimens collected were found on plants pollinated by insects (Appendix Y). The greater percentage of cerambycids were collected on plants pollinated by insects

85 (79%), with fewer specimens being collected on plants pollinated by insects/wind

(17%) and wind only (3%) (Appendix Z) (Fig. 28.6).

Wind

Insects

Cerambycidae

0 20406080

Figure 28.6: Relationship between plant pollination and

Cerambycidae species collected on Ezemvelo

Nature Reserve between January 2001 and

December 2001.

3.4.7 Plant Climate

Statistically there is no relationship between plant climate and cerambycid species on

ENR [(Cramér’s V = 0.235; p > 0.05); (N=506)]. The majority of specimens collected were found on temperate plants (Appendix AA). The greater percentage of cerambycids were collected on temperate plants (63%), significantly fewer on sub-

86 tropical plants (33%) and medium-temperature plants (4%) (Appendix AB) (Fig.

28.7).

Temperate

Sub- Tropical Cerambycidae

0 20406080 Figure 28.7: Relationship between plant climate and Cerambycidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

3.5 Buprestidae/Plant correlations

3.5.1 Plant order correlation

Statistically there is a relationship between plant order and buprestid species on ENR

[(Cramér’s V = 0.648; p < 0.05); (N=805)]. The majority of specimens collected were found on plants belonging to the order Fabales (Appendix AE). The greater

87 percentage of buprestids were collected on Fabales (79%), Rosales (6%), Sapindales

(4%) and Proteales (4%) (Appendix AF) (Fig. 29.1).

Celastrales

Ericales Buprestidae Rosales

Fabales 020406080

Figure 29.1: Relationship between tree orders and Buprestidae species

collected on Ezemvelo Nature Reserve between January

2001 and December 2001.

3.5.2 Plant family correlation

Statistically there is a relationship between plant family and buprestid species on ENR

[(Cramér’s V = 0.636; p < 0.05); (N=794)]. The majority of specimens collected were found on plants belonging to the Family Mimosaceae (Appendix AG). The greater

88 percentage of buprestids were collected on Mimosaceae (78%), Rhamnaceae (5%),

Celastraceae (4%), Anacardiaceae (3%) and Ebenaceae (3%) (Appendix AH) (Fig.

29.2).

Olacaceae

Poaceae

Rhamnaceae Buprestidae Papilionaceae

Mimosaceae 020406080

Figure 29.2: Relationship between plant family and Buprestidae species

collected on Ezemvelo Nature Reserve between January 2001

and December 2001.

3.5.3 Plant species correlation

Statistically there is a relationship between plant species and buprestid species on

ENR[(Cramér’s V = 0.561; p < 0.05); (N = 805)]. The majority of specimens

89 collected were found on plants belonging to the Acacia (Appendix AI). The greater percentage of buprestids were collected on Acacia karoo (50%), Acacia caffra

(28%), Ziziphus mucronata (5%), Protea caffra (4%), Uclea crispa (3%), Rhus lancea (3%) (Appendix AJ) (Fig. 29.3).

90 Ximenia caffra

Rhus pyroides

Protea caffra Buprestidae

Figure 29.3: Relationship between plant species and Buprestidae species Celtis africanacollected on Ezemvelo Nature Reserve between January 2001 and December 2001. Acacia caffra 0204060 3.5.4 Plant flower size correlation

Statistically there is a relationship between flower size and buprestid species on ENR

[(Cramér’s V = 0.399; p < 0.05); (N = 805)]. The majority of specimens collected were found on plants with small flowers (Appendix AQ). The greater percentage of buprestids were collected on plants with small flowers (94%), with significantly fewer specimens collected on plants with medium-large flowers (Appendix AR) (Fig. 29.4).

91 Buprestidae Medium LargeFigure 29.4: Relationship between flower size and Buprestidae species collected on Ezemvelo Nature Reserve between January Small 2001 and December 2001. 0 20406080100

3.5.5 Plant Phenology

Statistically there is a relationship between plant phenolgy and buprestid species on

ENR [(Cramér’s V = 0.764; p < 0.05); (N =805)]. The majority of specimens collected were found on deciduous plants (Appendix AK). The greater percentage of buprestids were collected on deciduous plants (86%), with significantly fewer specimens collected on non-deciduous plants (14%) (Appendix AL). The reason for this being the majority of the plants on ENR are deciduous plants (Fig. 29.5).

92 Buprestidae NO

YES

0 20406080100

Figure 29.5: Relationship between plant phenology and Buprestidae species collected on Ezemvelo Nature Reserve between January 2001 and December 2001.

3.5.6 Plant Pollination

Statistically there is a relationship between plant pollination and buprestid species on

ENR [(Cramér’s V = 0.769; p < 0.05); (N = 805)]. The majority of specimens collected were found on plants pollinated by insects (Appendix AM).

The greater percentage of buprestids were collected on plants pollinated by insects

(92%), with significantly fewer specimens being collected on plants pollinated by insects/wind (7%) and only wind (1%) (Appendix AN) (Fig. 29.6).

93 Wind

Buprestidae Insects/wind

Insects

0 20406080100 Figure 29.6: Relationship between plant pollination and Buprestidae

species collected on Ezemvelo Nature Reserve between

January 2001 and December 2001.

3.5.7 Plant Climate

Buprestids were found to occur predominantly on temperate plants (Appendix AO), however significantly more buprestids were found on sub-tropical plants than cerambycids. The greater percentage of buprestids were collected on temperate plants

(69%), with fewer specimens collected on sub-tropical plants (30%) and medium- temperate plants (1%) (Appendix AP) (Fig. 29.7).

94 Temperate

Medium Temperate Buprestidae

Sub-Tropical

020406080

Figure 29.7: Relationship between plant climate and Buprestidae species

collected on Ezemvelo Nature Reserve between January

2001 and December 2001.

3.6 Discussion

The total number of Buprestidae collected for 2001 was 805, of which 631 sampled on either A. caffra or A. karoo, leaving 174 being collected on other plant species.

The total number of Cerambycidae collected for 2001 was 253, of which 165 sampled either on Acacia caffra or Acacia karoo, leaving 88 collected on other plant species.

The genus Acacia is clearly very important to these two families from an ecological perspective. During the winter months both cerambycids and buprestid numbers dropped drastically, in some cases to zero abundance. The greatest total species diversity and abundance were in the summer months, peaking in November and

December. Insects abundance significantly different between quadrats. There were significant correlations between species richness and plant characteristics.

95 Buprestids did not appear to be attracted to light sheet, and only one species,

Acmaeodera albivillosa, was collected on ENR using this method (Appendix AT).

Cerambycids were more readily attracted to the light sheet, with the majority of the specimens collected belonging to the sub-family Lamiinae. The following species were collected at the light trap, Phantasis giganteus, Dalterus dejeani, Crossotus lacunosus, Crossotus plumicornis, Olenecamptus albidus, Alphitopola octomaculata,

Mycerinicus brevis, Phryneta spinator, Tragiscoschema bertolinii, Xystrocera erosa and Prosopocera lactator (Appendix AS).

To assess the relative importance of host plant on beetle species’ distribution, species on more that one host- plant species were classified as cosmopolitan (Nigel et al.

2004), while specialists, e.g. Evides pubiventris, were found only on Lannea discolor.

The number of beetle species collected at each quadrat varied (Figure 26), being highest in quadrat A and C and lower in quadrat B.

Average number of beetles collected per site did not differ as vary in terms of species density. There were no single species occurrences, yet certain species were more common than others. The most scarce cerambycid species were Xystrocera erosa (3 specimens); Prosopocera lactator (3 specimens) and Xystrocera dispar (2 specimens). The most abundant cerambycids species included Jonthodina sculptillis

(41 specimens); Philematium natalense (22 specimens) and Taurotagus klugi (30 specimens). The following buprestids were not collected in large numbers,

Chrysobothris dorsata (2 specimens); Brachelytrium transvalense (3 specimens) and

Anthaxia sp. 4 (2 specimens).

96 Species collected regularly included Pseudagrilus beryllinus (60 specimens) and

Anthaxia sp. 1(89 specimens); Anthaxia sp. 2 (102 specimens) and Anthaxia sp. 3 (64 specimens).

The total number of Cerambycidae and Buprestidae collected was 35 species and 32 species respectively. Common species showed a difference in terms of species richness. Species accumulation and estimated number of species among the different quadrats indicates the common species dataset having lower diversity than quadrat B.

However, there was no significant difference in key species between the different quadrats.

97 Chapter Four

4. DISCUSSION

According to Hull et al. (1998), the South African region may have a greater number of endemic, or range restricted, buprestid fauna compared to . However, this apparent range restriction is impossible to ascertain from the data collected for most of these beetles, and may be a false signal generated by the lack of data for many species. Hull et al. (1998) emphasized the need for additional invetebrate surveys, particularly in undersampled regions of southern Africa (Kremen et al. 1993;

Drinkrow & Cherry 1995). South Africa, high human population densities, greatest extent of land transformation, and often political land claims debate (Khan 1990;

Scholtz & Chown 1993), is in desperate need of information regarding species richness.

In the past, data on insect species richness of particular sites has been collected in two contrasting manners (Coddington et al. 1991; Longino 1994). Systematists collect samples in ways that maximize the number of species collected (Longino 1994), but the unsystematic nature of the sampling means that the ecological generalizations and extrapolations are difficult to make from the resulting inventories. On the other hand, samples collected to answer ecological questions may be more amendable to analysis and extrapolation (Longino 1994), but are often poor representations of the total fauna at a site (Godfray et al. 1999). This project has attempted to combine these two approaches, providing an inventory with sound ecological components included.

98 A number of programmes world-wide with the goal of making structured inventories are now either up and running or in an advanced stage of development (Gamez 1991; di Castri et al. 1992; & Longino 1994). Previous studies aimed at mandatory sites had either, 25 %, 50 %, 75 %, or 100 % of their total area included in conservation areas

(Hull et al. 1998), but excluded private reserves. This project has included the private sector, but excluded government conservation areas.

A number of other issues have been addressed in the design of sampling protocols for species inventories. First, the seasonal component of diversity was measured, since diversity and abundance may or may not be represented at certain times of the year

(DeVries et al. 1997; Richardson et al. 1997). Secondly, care was taken to ensure sampling effort was appropriate in relation to species diversity, since constant sampling effort can generate misleading patterns (Colwell & Coddington 1994;

Colwell & Hurtt 1994). Although it was difficult to estimate species diversity before the start of the inventory, increasingly accurate estimates of total species diversity were obtained as sampling proceeded using a variety of extrapolation techniques

(Codwell & Coddington 1994).

On ENR, this inventory was valuable in determining the magnitude, distribution and taxonomic composition of the biodiversity of these two families, but alone says little about the maintenance and dynamics. To address these questions, we need not only information about numbers and identities of species, but also their interactions.

99 4.1 Classification of Buprestoidea

Only one family, Buprestidae, was classified in this group (Holm & Bellamy 1985).

A new higher classification of the family was proposed in a key by Cobos (1980) in which the tribes of Lacordaire (1857) and the “groups” of Kerremans (1892, 1893a,

1903) are elevated to subfamilies. The monographs of Kerremans (1904-1914) remain the basis for current buprestid . Obenberger (1931b,c) studied various African genera and species. The most recent classification by Bellamy (2003) included two families: Buprestidae and Schizopodidae.

The following 6 subfamilies are represented in South Africa, with 5 subfamilies sampled on ENR:

Julodines are medium-sized to large (10-15 mm), torpedo-shaped beetles (Bellamy

2004). The subfamily is represented by six genera and 41 species (Holm & Bellamy

1985). Ferreira and Ferreira (1958a & b) reviewed the southern African species of

Sternocera occurring on ENR. Sternocera orissa was the only species representing this genus collected on ENR.

Polycestinae are medium-sized (9-25 mm) buprestids (Holm & Bellamy 1985).

The subfamily is represented by 78 genera and hundreds of species

(Bellamy 2004). Holm (1982) revised the African species of Acmaeodera. Seven species representing this subfamily were collected on ENR on a variety of plants.

These species included Acmaeodera albivillosa; Acmaeodera viridiaenea;

Acmaeodera aenea; Acmaeodera ruficaudis; Acmaeodera inscripta; Acmaeodera punctatissima; & Acmaeodera stellata.

100 Members of this subfamily Chalcophorinae are medium to large (15-45 mm) (Holm

& Bellamy 1985). Most are strikingly coloured in metallic shades. The genus

Psiloptera, accounts for the majority of the southern African chalcophorine species

(Holm & Bellamy 1985). The latest revision of southern African species was by

Ferreira and Ferreira (1958b), and Ferreira (1959). The subfamily is well represented in southern Africa with 74 genera and with a large number of species (Bellamy

2003).

Three species were collected on ENR, Lampetis conturbata; Lampetis gregaris &

Evides pubiventris. The later were collected on patches of stunted Lannea discolor in localized locations on a few ridges.

Buprestinae are small to medium-sized (5-16 mm) beetles (Holm & Bellamy 1985).

Most are a dark bronze in colour, but many species of Anthaxia have bright metallic colours (Holm & Bellamy 1985). The subfamily is well represented in southern

Africa with 110 genera and a large number of species (Bellamy 2003). Three species were collected on ENR, however due to their cryptic colouration 4 species could not be identified to species level., these included Anthaxia sp. 1; A. sp. 2; A. sp 3 & A. spp

4. The further 8 species include, A. bergrothi; A.obtectans, Brachelytrium transvalense, Chrysobothris boschismanni; Chrysobothris algoensis & Chrysobothris dorsata. 2 species were collected on ENR, primarily on Protea caffra. These species include Sphenoptera sinuosa & Sphenoptera arrowi. These species are characterized

101 by a waxy coating on the body, forming a colourful pattern which is easily removed when handled.

Agrilinae are small to medium-sized (3-20 mm) cylindrical beetles (Holm & Bellamy

1985). No one has attempted to revise the African Agrilinae, although some major studies such as those by Obenberger (1931a, 1935) and Thery (1929a) have appeared.

There are at least 128 genera and many species, of which 150 species belong to the large genus Agrilus (Bellamy 2004). 9 species were collected on ENR in this study, representing 6 genera. These include Agrilus guerryi; Agrilus falcatus;

Pseudagrilus beryllinus; Kamosia tenebricosa; Agrilomorpha venosa; Kamosiella dermestoides; & Phlocteis exasperata. This subfamily is also represented by

Trachys ziziphusii collected on Ziziphus mucronata on the reserve.

4.2 Classification of Cerambycidae

Cerambycidae are within the superfamily Chrysomeloidea (Cox 1985).

General studies of the southern African cerambycid fauna have been undertaken by

Ferreira and Ferreira (1959 a,b,c) and Tippmann (1959). According to Cox (1985), a comprehensive account of cerambycid biology was given by Duffy (1953) and the larvae and pupae of many of the economically important cerambycids of southern

Africa were described by Duffy (1957).

There are 6 subfamilies of Cerambycidae known to occur in southern Africa, 3 of

102 these subfamilies were represented on ENR.

Parandrinae are regarded as the most primitive group of cerambycids and are poorly represented in southern Africa (only one genus, three species) (Cox 1985). These are medium-sized beetles that are usually found on their hosts or attracted to lights

(Cox 1985). This subfamily was not collected on ENR due to the reserve falling outside the distribution of their host plants.

The Aseminae has been studied by Ferreira (1955), and are not represented on ENR.

Prioninae are medium to large-sized (25-100 mm) beetles and are mostly nocturnal

(Cox 1985). These beetles are usually encountered on the trunks or branches of their hosts or are attracted to artificial light (Cox 1985). Acanthophorus capensis, which was collected at ENR, is one of the largest beetles in southern Africa (Cox 1985).

The Prioninae is represented by a modest number of 13 genera and 22 species (Cox

1985). Southern African prionines have been revised and lists of species given by

Ferreira (1958), Ferreira and Ferreira (1952a,b, 1956, and 1959a,b,c) and Gilmour

(1956). Three genera are represented on ENR, comprising of four species. These included Macrotoma palmata; Macrotoma natala; Anthracocentrus capensis; &

Tithoes maculates. All four species were collected using the beating method.

103 Most Lepturinae are distributed in the northern hemisphere and only a few species are recorded from Africa, and only one species, Otteissa sericea, is known to occur in

South Africa (Cox 1985). This subfamily was not represented on ENR as a result of specific habitat requirements and extremely localized distribution.

The subfamily Cerambycinae is a very large group, which is well distributed in all major faunal areas (Cox 1985). Many species are diurnal and show several interesting mimicry adaptations, species with shortened elyra probably mimic species of Hymenoptera (Cox 1985). The Cerambycinae are well represented in southern Africa, 75 genera and 150 species (Cox 1985). The group was revised by Ferreira (1964). Twelve genera and fifteen species in Cerambycinae were collected on ENR using the beating method (Holm 1985), and were not attracted to light to the same degree as the other subfamilies.

These include Zamium incultum; Zamium. bimaculatum; Anubis clavicornis; Anubis mellyi; Xystrocera erosa; Xystrocera dispar; Captoeme krantzi; Taurotagus klugi;

Jonthodina sculptilis; Hypoeschrus ferreirae; Plocaederus denticornis; Philematium natalense; Phyllocnema latipes; Pacydissus sp. & Ossibia fuscata.

Lamiinae are by far the most successful group of cerambycids and the fauna of southern Africa is no exception (Cox 1985). The diurnal species are often very brightly coloured, with nocturnal species mostly of more sombre colours

(Cox 1985). Certain species cause their host plants to produce galls

(Cox 1985). The subfamily is well represented in southern Africa, 124 genera and 475 species (Cox 1985). Ferreira and Ferreira (1959a & b),

Hunt and Breuning (1956) and Ferreira (1966) compiled lists of the lamiine species.

104 This subfamily is well represented on ENR, with thirteen genera and sixteen species.

These include Crossotus lacunosus; Crossotus plumicornis; Crossotus stypticus;

Dalterus degeeri; Dalterus dejeani; Phantasis giganteus; Olenecamptus albidus;

Nemotragus helvolus; Hecyra terrea; Ceroplesis thunbergi; Lasiopezus longimanus;

Alphitopola octomaculata; Mycerinicus brevis; Phryneta spinator; Tragiscoschema bertolinii & Prosopocera lactator.

4.3 Factors influencing abundance and diversity on ENR

Phytophagous species that are cosmopolitan (in this study, defined as those found in all three quadrats), may be more resilient to local climate changes and changes in the distribution of hosts, and will survive in situ and/or could move with the host plant and potentially expand their range (Nigel et al. 2004). Specialists (defined as species found only on one host species in a single quadrat) would be affected more with changing climate and survival of the host species (Nigel et al. 2004). On ENR, results indicate that many species may be displaced by extreme fluctuations in temperature.

However, the overall community structure of these two phytophagous beetle communities may be resilient. In addition to the clear differences in total number of insects between the quadrats, there was also clearly a difference in species diversity.

When all samples within the various quadrats were pooled, total species richness increased, as expected. According to Nigel et al. (2004) total phytophagous species richness appears to be consistently higher in the tropics than in more temperate zones.

In a comparison of beetle species richness on different host-plant genera, families – see Acacia’s compared to Heteropyxis natalensis and Croton gratissimus.

The majority of the host plants on ENR already extend their range and are pre-adapted to cope with temperature change. However, associated beetles may not be as resilient

105 to the drastic fluctuations in temperature. Environmental gradients are a useful tool for understanding the role of climate in structuring insect communities (Harrison

1993; Hodkinson et al. 1999). May (1990), suggested that the study of food webs might help understand insect richness. Some patterns emerged involving the number of beetle species with different characteristics of different plants.

4.3.1 Cerambycidae abundance and diversity

• Total Cerambycidae collected

Cerambycidae were collected at monthly intervals throughout the year. October,

November, December and January were months with the highest activity and species diversity. Collection results for months June and July, indicate almost no cerambycid activity during this period. The results indicate that environmental conditions associated with seasonality control the abundance and diversity of cerambycids

• Cerambycidae collected in grid A

Cerambycids collected in quadrat A follow the same general trend with high beetle numbers being recorded during the summer months with gradual decline towards the winter months.

• Cerambycidae collected in grid B

Cerambycid numbers in quadrat B indicate an increase in cerambycid numbers at the beginning of November into December. Cerambycid abundance and diversity in quadrat B appears to decline earlier than in quadrat A, with numbers starting to decline in January.

• Cerambycidae collected in grid C

Cerambycid samples recorded in quadrat C indicate a clear decline from January to

July, with a gradual increase in numbers from August to December. November and

106 December clearly are the most environmentally suitable months for cerambycid activity.

• Reserve Climate

There is a direct relationship between the number of cerambycids collected and average minimum temperature. This indicates that due to temperatures dropping exceptionally low in winter, these conditions directly affect cerambycid populations and the adult population dies off during this period, leaving eggs and larvae to continue when spring returns. Cerambycid numbers and the average maximum temperatures correlate with each other but not to the same extent as with average minimum temperature. Cerambycid numbers indicate a slight correlation with average rainfall, numbers decreasing with a decrease in rainfall, while increasing with spring rains. This relationship does not appear as defined as with temperature.

• Plant correlations

Statistically there is a relationship between tree order and cerambycid species on

ENR. The majority of specimens collected were found on trees of the family Fabales.

In all quadrats, cerambycid abundance and diversity was greatest on Mimosaceae, which include Acacia caffra and Acacia karoo. Certain plant families had zero species recorded. Statistically there is a relationship between plant species and cerambycid species on ENR. The majority of specimens collected were found on

Acacia species. Plant species certainly do have some importance as host plants to certain species, however certain plants appear to have a high attraction to a broad range of species. Other species, possibly due to high amounts of chemicals to deter browsers, are very unattractive to cerambycids. Statistically there is a small relationship between flower size and cerambycid species on ENR. The majority of

107 specimens collected were found on small flowers. Statistically there is a significant relationship between plant phenology and cerambycid species on ENR. The majority of specimens collected were found on deciduous plants. Statistically there is no relationship between plant pollination and cerambycid species on ENR. The majority of specimens collected were found on plants pollinated by insects. Statistically there is no relationship between plant climate and cerambycid species on ENR. The majority of specimens collected were found on temperate plants.

4.3.2 Buprestidae abundance and diversity

• Total Buprestidae collected

Buprestidae were collected at monthly intervals throughout the year. November,

December and January were months with the highest activity and species diversity

Collection results for months June and July, indicate almost no buprestid activity during this period.

• Buprestidae collected in grid A

Buprestids collected in quadrat A follow a similar trend with high beetle numbers being recorded during the summer months with gradual decline towards the winter months.

• Buprestidae collected in grid B

Buprestidae samples from quadrat B indicate a drastic increase in buprestid numbers at the beginning of December. Buprestid abundance and diversity in quadrat B appears to decline gradually from March, with no samples collected during June, July or August.

• Buprestidae collected in grid C

108 Buprestidae samples recorded in quadrat C indicate a drastic decline from March to

April, with a rapid increase in numbers from September to December. November,

December and January clearly are the most environmentally suitable months for buprestid activity.

• Reserve Climate

Analysis of regression indicates a significant linear trend (r² =0.73) for minimum temperature parameters and total buprestids collected during 2001 on ENR.

Minimum temperature therefore is expected to have an affect on the total number of buprestids collected in 2001 on ENR. This indicates that due to temperatures dropping exceptionally low in winter, these conditions directly affect buprestid populations and adult populations die off during this period, leaving eggs and larvae to behind. Analysis of regression indicates a significant linear trend (r² = 0.52) for the following parameters, maximum temperature and total buprestids collected during

2001 on ENR. Maximum temperature therefore is expected to have an affect on the number of buprestids collected in 2001 on ENR. Analysis of regression indicates an insignificant linear trend (r² = 0.34) for the following parameters, average rainfall and total buprestids collected during 2001 on ENR Average rainfall is not expected to have an affect on the number of buprestids collected during 2001 on ENR.

• Plant correlations

Statistically there is a relationship between plant order and buprestid species on ENR.

The majority of specimens collected were found on Fabales. Statistically there is also a significant relationship between plant family and buprestid species on ENR. The majority of specimens collected were found on Mimosacease. Statistically there is a

109 relationship between plant species and buprestid species on ENR. The majority of specimens collected were found on plants belonging to the genus Acacia. Statistically there is a relationship between flower size and buprestid species on ENR.

The majority of specimens collected were found on plants with small flowers.

Statistically there is a relationship between plant phenolgy and buprestid species on

ENR. The majority of specimens collected were found on deciduous plants.

Statistically there is a relationship between pollination and buprestid species on ENR.

The majority of specimens collected were found on plants pollinated by insects.

Buprestids were found to occur predominantly on temperate plants, however more buprestids were found on sub-tropical plants than cerambycids.

4.4 Overview

In spite of problems with specifically defining what a rare species is (Gaston 1994), many diversity studies have found that rare species make up a high proportion of the overall species richness (Basset 1993; Fensham 1994; Bürki & Nentwig 1997;

Sárospataki 1999; Novotny & Basset 2000; Magurran & Henderson 2003). On ENR, this does not appear to be the case, and common species seem to make up the highest proportion of overall species richness. According to Coddington et al. (1996) thus indicating that rare species are thought to contribute more to diversity at tropical latitudes than at temperate latitudes. This study indicates that rare species do not appear to have a large role in determining changes in community structure in all quadrats. Changes in species diversity along environmental gradients, such as aspect,

110 slope and altitude, such gradients are in part associated with change in resource availability (Rotenberry 1978; Shmida & Wilson 1985; Stevens & Willig 2002).

The concept of landscapes as complex mosaics of habitats varying in quality with respect to different groups of organisms, has been the subject of a number of recent studies (Wien 1995; Gacon et al. 1999; Ricketts et al. 2001). On ENR, patches of habitat with varying quality are likely to underlie the differences we found in insect abundance and diversity between quadrats.

According to Godfray et al. (1999), sufficient data exists to point to global patterns in insect diversity. More species tend to be found in the canopies of tropical American trees than those in Africa. More data is required, as this does not seem to suggest a correlation between plant and insect diversity. This study proved that host plant specificity can be addressed by studying the diet of what different beetle species. But as a single tree can produce many thousands of individuals of hundreds of species, processing this data would create huge logistical difficulties (Godfray et al. 1999).

Studies of host-specificity and the number of species per tree, are interesting in their own right (e.g. Strong et al. 1984. Futuyma & Moreno 1988). A number of reasons have been put forward to explain both higher and lower host specificity of herbivores

(Janzen 1973; Price 1991). The resource fragmentation hypothesis states that higher diversity lead to lower population densities for individual species and hence less host specificity, as the most specialized species are unable to maintain themselves on the

111 most fragmented resources (Godfray et al. 1999). Fiedler (1998) found no difference in host-plant range (measured by the number of families) between tropical and temperate species. Very little work has been done on the host-plant relationship in phytophagous beetles. No other group of insects matches the butterflies in host-plant data available for a complete fauna (Godfray et al. 1999).

Interestingly, the more widespread host plants, A. caffra and A. karoo, the higher the diversity of insect herbivore species, a pattern also found in temperate tree-feeding herbivores (Southwood 1961). Eggs and immature of the two families were not studied due to time and logistical implication, although predation by birds and other insects would probably have been relevant. Eggs, larval and adults are thought not to have high levels of chemical defense, although there are many exceptions (Brower

1989). It has been suggested that the fractal nature of the world may lead to a greater number of niches at smaller spatial scales (Lawton 1984; Morse et al. 1985, although

Fenchel (1993) and it has been pointed out that if the environment is truly fractal then heterogeneity of resources will be similar at all spatial scales.

It has long been known that insect richness correlates with plant species richness, both at local (e.g. Southwood et al. 1979; Siemann et al. 1998) and regional levels

(Prendergast et al. 1993). Equally interesting at ENR was the variation in numbers, especially within the quadrat B, with low plant diversity. Of course, if the ratio is approximately constant, then the total number of insects follows from the number of plant species (Godfray et al. 1999). Employing this logic gives estimates in the range of three to eight million speices of insects (Gaston 1992).

Hull et al. (1998) recognized the inefficiency of the present southern African conservation network in representing Buprestidae. Some species are represented many

112 times in some reserves while some reserves do not include any known Buprestidae records at all, although this may be due to survey bias (Hull et al. 1998).

It is acknowledged that areas identified as having high or low concentrations

(hotspots and coldspots) may merely reflect biased collection efforts (Gentry 1992).

Examples of such areas are hotspots in close proximity to major towns, cities or research institutions (Gelderblom & Bronner 1995), and/or cold spots in regions of poor sampling (Drinkrow & Cherry 1995).

In this context, the results of this study add to the “bigger picture” of invertebrate conservation on a pristine reserve, reconfirming that the poor management practices of farmers and developers are likely to have a detrimental impact on insects and, particularly, buprestid faunas (Hull et al. 1998).

113 Chapter Five

5. ECOLOGICAL IMPORTANCE AND FUTURE MANAGEMENT OF CERAMBYCIDAE AND BUPRESTIDAE ON EZEMVELO NATURE RESERVE

Cerambycidae and Buprestidae are very important phytophagous insect families from an ecological perspective, due to their role in the breakdown of wood and role in the nutrient cycle.

Fire plays an important role in maintaining and creating suitable conditions for flora and fauna on a reserve (Friend & Williams 1993). Natural and accidental burns occur on the reserve, but it is however difficult to quantify the effect of fire on species diversity and abundance of species on a reserve (Friend & Williams 1993). A study of the amount of activity in the past and present after an area is burnt would give an indication of the effect of the impact. According to Friend & Williams (1993), fire varies in frequency, intensity, extent, season and interactions with other disturbance processes, therefore different fires may have different effects.

Dead or dying wood is high in both Cerambycidae and Buprestidae activity, usually harbouring large numbers of larvae and eggs at varying times of the year. It would therefore be important to note these times, reducing fires to periods when activity may be low, thus minimizing the effect on these insects.

114 Low intensity fires are expected to cause less damage to eggs and larvae as the fires burn faster over an area (Friend & Williams 1993).

According the Holm & Bellamy (1995), life cycles of buprestids can be extremely long, with a case over 35 years on record. According to Friend & Williams (1993), eggs are more susceptible to fire as they are near the surface of the wood, larvae only drill deep into the wood after hatching. Implications are that beetle diversity and abundance would be affected to greater extent with regular burns of high intensity during times when these beetles are laying eggs, namely summer months. ENR is predominantly grassland, with rocky outcrops and fire rarely spreads to these outcrops.

High quantities of wood debris are recognized as an important component of a healthy ecosystem linked to biodiversity and ecosystem processes. According to Friend &

Williams (1993), these areas are high centres of biological interaction and energy exchange symbolizing in many ways the complexity of the ecosystem. Care should be taken when collecting fire wood, such over utilization of dead wood in a non- sustainable manner could lead to decreases in certain species.

Many wood boring beetles only appear particularly damaging to new growth and plants weakened by various causes such as drought, frost damage and defoliation by other leaf feeding insects. According to Holm & Bellamy (1985), most buprestids attack moribund rather than dead wood. Certain cerambycids, however, may oviposit on freshly cut, slightly injured, or decaying wood (Cox 1985).

115 Chapter Six

6. CONCLUSION

Darwin (1889), recently quoted by Longino (1994) wrote: “The number of minute and obscurely coloured beetles is exceedingly great. The cabinets of Europe can, as yet, boast only of the larger species from tropical climates. It is enough to disturb the composure of an entomologist’s mind, to look forward to the dimensions of the complete catalogue”. In the light of this, extrapolation from local inventories to broader geographical areas may provide a way of accurately estimating global species richness (May 1988; Colwell & Codington 1994).

This project was primarily concerned with defining and assessing ecosystem health and providing ecological indicators for ecosystem management. Temperature and seasonal changes in the ecosystem, had a profound impact on the diversity and abundance of both families. Certain species from both families appeared more host- specific to other species from the same genera. Acacia species are the most important host plant, harbouring the highest diversity and abundance of both families. Transects at the three sites significantly differed in the diversity and abundance of cerambycid and buprestid assemblages, with lower diversity and abundance near the Wilge river.

These families showed a gradient in species richness similar to the plants. In transects richer in plant species, there is greater diversity and abundance of buprestids and cerambycids.

116 Inventories are valuable for aiding in conservation related decisions, since land owner’s decisions are often made at local scales and here wood borers and other insects can provide a rich source of data on environmental change (Kremen et al.

1993). According to Godfray et al. (1999), phytophagous insect inventories and their associated host plants reveal the structure and patterning of communities, and generate hypotheses about how component species interact.

The project has acted as an entomological indicator, taking in to consideration two taxonomic groups that reflect the diversity of other insects across a set of environments, thus acting as surrogates for the “wholesale” biodiversity (Gaston

1996a; McGeoch 1998). The conservation value of an area is typically judged using a measure of species richness, or some variant of it (Gaston 1996b; Angermeier &

Winston 1997).

This data suggests that managing reserves to maximize insect abundance, especially these key beetle families, by maintaining diverse and structurally varied habitats, is important in maintaining a healthy ecosystem.

117 Chapter Seven

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134 135 Appendix A

Total Cerambycidae species diversity and abundance for each month of the year in all quadrats on ENR

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Zamium incultum 6 0 401000133321 Coptoeme krantzi 3 5 211000200721 Taurotagus klugi 2 3 002002346830 Jonthodina sculptilis 1 5 1031011118941 Anubis clavicornis 2 2 110000135520 Macrotoma palmata 3 2 000000217318 Tithoes maculates 2 3 310000145827 Phantasis giganteus 2 1 010000151112 Dalterus dejeani 2 1 000000104311 Crossotus lacunosus 0 2 000000223716 Crossotus plumicornis 1 0 100000226921 Olenecamptus albidus 2 2 012101202215 Anthracocentrus capensis 2 2 100000074117 Hypoeschrus ferreirae 0 3 000001041211 Plocaederus denticornis 3 4 110000016218 Crossotus stypticus 2 3 000000033516 Nemotragus helvolus 5 3 200000003114 Hecyra terrea 1 0 00000020115 Ceroplesis thunbergi 6 0 110000354424 Lasiopezus longimanus 0 0 20000010317 Philematium natalense 2 2 310000026622

136 Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Dalterus degeeri 3 1 000000031210 Zamium bimaculatum 3 2 000002215419 Anubis mellyi 2 0 000000028214 Alphitopola octomaculata 2 0 20100200018 Mycerinicus brevis 2 0 00000001104 Phryneta spinator 0 0 10000001215 Tragiscoschema bertolinii 2 3 200000123720 Phyllocnema latipes 1 0 00000012419 Pacydissus sp.0 0 011000112612 Macrotoma natala 3 0 200000002411 Ossibia fuscata 1 0 100000125111 Xystrocera erosa 0 1 10000000103 Prosopocera lactator 0 0 00000010023 Xystrocera dispar 1 0 00000000012 Total 67 50 31 9 11 2 0 9 32 72 115 120 518

137 Appendix B

Percentage of Cerambycidae species diversity and abundance for each month of the year in all quadrats on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Zamium incultum 1 0 10000001114 Coptoeme krantzi 1 1 00000000014 Taurotagus klugi 0 1 00000011126 Jonthodina sculptilis 0 1 00100002228 Anubis clavicornis 0 0 00000001114 Macrotoma palmata 1 0 00000000113 Tithoes maculates 0 1 10000001125 Phantasis giganteus 0 0 00000001002 Dalterus dejeani 0 0 00000000112 Crossotus lacunosus 0 0 00000000113 Crossotus plumicornis 0 0 00000000124 Olenecamptus albidus 0 0 00000000003 Anthracocentrus capensis 0 0 00000001103 Hypoeschrus ferreirae 0 1 00000001002 Plocaederus denticornis 1 1 00000000103 Crossotus stypticus 0 1 00000001113 Nemotragus helvolus 1 1 00000000103 Hecyra terrea 0 0 00000000001 Ceroplesis thunbergi 1 0 00000011115 Lasiopezus longimanus 0 0 00000000101 Philematium natalense 0 0 10000000114

138 Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Dalterus degeeri 1 0 00000001002 Zamium bimaculatum 1 0 00000000114 Anubis mellyi 0 0 00000000203 Alphitopola octomaculata 0 0 00000000002 Mycerinicus brevis 0 0 00000000001 Phryneta spinator 0 0 00000000001 Tragiscoschema bertolinii 0 1 00000000114 Phyllocnema latipes 0 0 00000000102 Pacydissus sp. 0 0 00000000012 Macrotoma natala 1 0 00000000012 Ossibia fuscata 0 0 00000000102 Xystrocera erosa 0 0 00000000001 Prosopocera lactator 0 0 00000000001 Xystrocera dispar 0 0 00000000000 % 13106220026142223100

139 Appendix C

Total Cerambycidae species diversity and abundance for each month of the year in quadrat A on ENR

Species Jany Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Zamium incultum 1 0 401000121313 Coptoeme krantzi 3 1 21000010019 Taurotagus klugi 0 2 000002134416 Jonthodina sculptilis 1 4 000001143216 Anubis clavicornis 0 2 10000010206 Macrotoma palmata 1 0 00000021329 Tithoes maculates 2 0 000000044414 Phantasis giganteus 2 0 01000001004 Dalterus dejeani 2 1 00000010015 Crossotus lacunosus 0 2 00000012027 Crossotus plumicornis 0 0 10000000269 Olenecamptus albidus 2 2 00000100016 Anthracocentrus capensis 0 2 00000002206 Hypoeschrus ferreirae 0 3 000001041211 Plocaederus denticornis 3 4 110000016218 Crossotus stypticus 0 1 00000003127 Nemotragus helvolus 3 0 00000000104

140 Continued

Species Jany Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Hecyra terrea 1 0 00000020115 Ceroplesis thunbergi 2 0 010000122311 Lasiopezus longimanus 0 0 20000010317 Philematium natalense 1 1 100000023210 Dalterus degeeri 3 1 000000031210 Total 2726124100513344041203

141 Appendix D

Percentage of Cerambycidae species diversity and abundance for each month of the year in quadrat A on ENR

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Zamium incultum 0 0 20000001016 Coptoeme krantzi 1 0 10000000004 Taurotagus klugi 0 1 00000101228 Jonthodina sculptilis 0 2 00000002118 Anubis clavicornis 0 1 00000000103 Macrotoma palmata 0 0 00000010114 Tithoes maculates 1 0 00000002227 Phantasis giganteus 1 0 00000000002 Dalterus dejeani 1 0 00000000002 Crossotus lacunosus 0 1 00000001013 Crossotus plumicornis 0 0 00000000134 Olenecamptus albidus 1 1 00000000003 Anthracocentrus capensis 0 1 00000001103 Hypoeschrus ferreirae 0 1 00000002015 Plocaederus denticornis 1 2 00000000319 Crossotus stypticus 0 0 00000001013 Nemotragus helvolus 1 0 00000000002 Hecyra terrea 0 0 00000010002

142 Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Ceroplesis thunbergi 1 0 00000001115 Lasiopezus longimanus 0 0 10000000103 Philematium natalense 0 0 00000001115 Dalterus degeeri 1 0 00000001015 % 13136200026172020100

143 Appendix E

Total Cerambycidae species diversity and abundance for each month of the year in quadrat B on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Zamium bimaculatum 3 2 000001003312 Jonthodina sculptilis 0 1 100000024513 Anubis mellyi 0 0 00000001506 Macrotoma palmata 2 0 00000000103 Tithoes maculates 0 2 00000010025 Alphitopola octomaculata 2 0 20100200018 Crossotus lacunosus 0 0 00000010124 Mycerinicus brevis 2 0 00000001104 Phryneta spinator 0 0 10000001215 Dalterus dejeani 0 0 00000000426 Ceroplesis thunbergi 1 0 00000012004 Tragiscoschema bertolinii 1 1 10000000025 Anubis clavicornis 0 0 00000002316 Phyllocnema latipes 1 0 00000000102

144 Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Pacydissus sp. 0 0 00100010158 Coptoeme krantzi 0 4 00100010028 Taurotagus klugi 0 0 00000001214 Philematium natalense 0 1 10000000114 Macrotoma natala 0 0 00000000224 Crossotus plumicornis 0 0 00000002305 Total 12116030035123430116

145 Appendix F

Percentage of Cerambycidae species diversity and abundance for each month of the year in quadrat B on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Zamium bimaculatum 3 2 000001003310 Jonthodina sculptilis 0 1 100000023411 Anubis mellyi 0 0 00000001405 Macrotoma palmata 2 0 00000000103 Tithoes maculates 0 2 00000010024 Alphitopola octomaculata 2 0 20100200017 Crossotus lacunosus 0 0 00000010123 Mycerinicus brevis 2 0 00000001103 Phryneta spinator 0 0 10000001214 Dalterus dejeani 0 0 00000000325 Ceroplesis thunbergi 1 0 00000012003 Tragiscoschema bertolinii 1 1 10000000024 Anubis clavicornis 0 0 00000002315 Phyllocnema latipes 1 0 00000000102 Pacydissus sp. 0 0 00100010147 Coptoeme krantzi 0 3 00100010027

146 Taurotagus klugi 0 0 00000001213 Philematium natalense 0 1 10000000113

Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Macrotoma natala 0 0 00000000223 Crossotus plumicornis 0 0 00000002304 % 109 5030034102926100

147 Appendix G

Total Cerambycidae species diversity and abundance for each month of the year in quadrat C on ENR.

148 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Nemotragus helvolus 2 3 200000002110 Crossotus stypticus 2 2 00000000239 Tragiscoschema bertolinii 1 2 100000123515 Anthracocentrus capensis 2 0 100000052111 Anubis clavicornis 2 0 01000001048 Anubis mellyi 2 0 00000001328 Phyllocnema latipes 0 0 00000012317 Ossibia fuscata 1 0 100000125111 Taurotagus klugi 2 1 002000200310 Coptoeme krantzi 0 0 00000000044 Zamium incultum 5 0 00000001208 Zamium bimaculatum 0 0 00000121217 Xystrocera erosa 0 1 10000000103 Pacydissus sp.0 0 01000001114 Philematium natalense 1 0 11000000238 Jonthodina sculptilis 0 0 003100051212 Macrotoma palmata 0 2 00000000316 Tithoes maculates 0 1 31000000128 Crossotus lacunosus 0 0 00000000235

Continued

149 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Macrotoma natala 3 0 20000000027 Phantasis giganteus 0 1 00000014118 Crossotus plumicornis 1 0 00000020137 Ceroplesis thunbergi 3 0 10000011219 Olenecamptus albidus 0 0 01210020219 Prosopocera lactator 0 0 00000010023 Xystrocera dispar 1 0 00000000012 Total 2813135720114264149199

Appendix H

Percentage of Cerambycidae species diversity and abundance for each month of the year in quadrat C on ENR

150 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Nemotragus helvolus 1 2 10000000115 Crossotus stypticus 1 1 00000000125 Tragiscoschema bertolinii 1 1 10000011238 Anthracocentrus capensis 1 0 10000003116 Anubis clavicornis 1 0 01000001024 Anubis mellyi 1 0 00000001214 Phyllocnema latipes 0 0 00000011214 Ossibia fuscata 1 0 10000011316 Taurotagus klugi 1 1 00100010025 Coptoeme krantzi 0 0 00000000022 Zamium incultum 3 0 00000001104 Zamium bimaculatum 0 0 00000111114 Xystrocera erosa 0 1 10000000102 Pacydissus sp.0 0 01000001112 Philematium natalense 1 0 11000000124 Jonthodina sculptilis 0 0 00210003116 Macrotoma palmata 0 1 00000000213

Continued

151 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Tithoes maculates 0 1 21000000114 Crossotus lacunosus 0 0 00000000123 Macrotoma natala 2 0 10000000014 Phantasis giganteus 0 1 00000012114 Crossotus plumicornis 1 0 00000010124 Ceroplesis thunbergi 2 0 10000011115 Olenecamptus albidus 0 0 01110010115 Prosopocera lactator 0 0 00000010012 Xystrocera dispar 1 0 00000000011 % 147 7341017132125100

Appendix I

Total Buprestidae species diversity and abundance for each month of the year in all quadrats on ENR.

152 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Acmaeodera albivillosa 7 3 313102135736 Acmaeodera viridiaenea 3 6 200000214220 Acmaeodera aenea 4 2 004000103115 Acmaeodera ruficaudis 2 2 100000020613 Acmaeodera inscripta 3 0 000000113412 Sternocera orisa 125 6410000031041 Anthaxia bergrothi 6 3 511000238938 Agrilus guerryi 6 3 100013122524 Lampetis gregaris 2 0 10000000317 Chrysobothris boschismanni 5 4 100000205825 Chrysobothris algoensis 2 4 020000204923 Agrilus sexguttatus 5 1 000000023112 Anthaxia sp. 1 161098300047131989 Anthaxia sp. 2 11191744000191621102 Anthaxia sp. 3 9 9 6741021451664 Trachys ziziphusii 2 3 230000010415 Pseudagrilus beryllinus 5 5 52100023142360 Brachelytrium transvalense 2 0 00000000103 Chrysobothris dorsata 1 0 00000000102 Acmaeodera punctatissima 0 1 10000002015 Kamosia tenebricosa 2 3 000000003210 Agrilomorpha venosa 3 1 100000113111

Continued

153 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Agrilus falcatus 3 0 10000000116 Sphenoptera arrowi 8 5 5040000461345 Kamosiella dermestoides 3 1 100000002310 Acmaeodera stellata 2 0 402000003213 Phlocteis exasperata 2 2 100000012614 Lampetis conturbata 7 5 500000023527 Evides pubiventrus 6 4 400000003522 Sphenoptera sinuosa 5 2 310000025422 Anthaxia obtectans 4 1 100000220616 Anthaxia sp. 41 0 00000000203 Total 149 104 86 33 27 2 1 7 23 52 126 195 805

Appendix J

154 Percentage of Buprestidae species diversity and abundance for each month of the year in all quadrats on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Acmaeodera albivillosa 1 0 00000000114 Acmaeodera viridiaenea 0 1 00000000002 Acmaeodera aenea 0 0 00000000002 Acmaeodera ruficaudis 0 0 00000000012 Acmaeodera inscripta 0 0 00000000001 Sternocera orisa 1 1 10000000015 Anthaxia bergrothi 1 0 10000000115 Agrilus guerryi 1 0 00000000013 Psiloptera gregaris 0 0 00000000001 Chrysobothris boschismanni 1 0 00000000113 Chrysobothris algoensis 0 0 00000000013 Agrilus sexguttatus 1 0 00000000001 Anthaxia sp. 1 2 1 110000012211 Anthaxia sp. 2 1 2 200000012313 Anthaxia sp. 3 1 1 11000000128 Trachys ziziphusii 0 0 00000000002 Pseudagrilus beryllinus 1 1 10000000237 Brachelytrium transvalense 0 0 00000000000 Chrysobothris dorsata 0 0 00000000000

Continued

155 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Acmaeodera punctatissima 0000000000001 Kamosia tenebricosa 0000000000001 Agrilomorpha venosa 0000000000001 Agrilus falcatus 0000000000001 Sphenoptera arrowi 1110000000126 Kamosiella dermestoides 0000000000001 Acmaeodera stellata 0000000000002 Phlocteis exasperata 0000000000012 Lampetis conturbata 1110000000013 Evides pubiventrus 1000000000013 Sphenoptera sinuosa 1000000000103 Anthaxia obtectans 0000000000012 Anthaxia sp. 4 0000000000000 % 19131143001361624100

Appendix K

156 Total Buprestidae species diversity and abundance for each month of the year in quadrat A on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Acmaeodera albivillosa 52213102125428 Acmaeodera viridiaenea 36200000214220 Acmaeodera aenea 22004000101111 Acmaeodera ruficaudis 1010000002048 Acmaeodera inscripta 30000000113210 Acmaeodera punctatissima 0110000002015 Sternocera orisa 54541000003628 Anthaxia sp. 1 65450000324736 Anthaxia sp. 2 46532000134735 Anthaxia sp. 3 46244102122634 Anthaxia bergrothi 32301000223723 Agrilus guerryi 63100013122524 Agrilus sexguttatus 51000000023112 Agrilus falcatus 3010000000116 Chrysobothris boschismanni 43100000004416 Chrysobothris algoensis 13020000103414 Chrysobothris dorsata 1000000000102 Sphenoptera arrowi 31200000001512 Trachys ziziphusii 23230000010415

Continued

157 Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Kamosia tenebricosa 23000000003210 Agrilomorpha venosa 31100000113111 Lampetis gregaria 1010000000316 Pseudagrilus beryllinus 11010000003410 Brachelytrium transvalense 2000000000103 Total 70 53 34 23 15 2 1 7 15 23 57 79 379

158 Appendix L

Percentage of Buprestidae species diversity and abundance for each month of the year in quadrat A on ENR

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Acmaeodera albivillosa 1110100101117 Acmaeodera viridiaenea 1210000010115 Acmaeodera aenea 1100100000003 Acmaeodera ruficaudis 0000000001012 Acmaeodera inscripta 1000000000113 Acmaeodera punctatissima 0000000001001 Sternocera orissa 1111000000127 Anthaxia sp. 1 2111000011129 Anthaxia sp. 2 1211100001129 Anthaxia sp. 3 1211100101129 Anthaxia bergrothi 1110000011126 Agrilus guerryi 2100000101116 Agrilus sexguttatus 1000000001103 Agrilus falcatus 1000000000002 Chrysobothris boschismanni 1100000000114 Chrysobothris algoensis 0101000000114 Chrysobothris dorsata 0000000000001 Sphenoptera arrowi 1010000000013 Trachys ziziphusii 1111000000014

159 Continued

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Kamosia tenebricosa 1 1 00000000113 Agrilomorpha venosa 1 0 00000000103 Lampetis gregaria 0 0 00000000102 Pseudagrilus beryllinus 0 0 00000000113 Brachelytrium transvalense 1 0 00000000001 % 1814964102461521100

160 Appendix M

Total Buprestidae species diversity and abundance for each month of the year in quadrat B on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Acmaeodera inscripta 0000000000011 Acmaeodera aenea 0000000000101 Acmaeodera albivillosa 2110000001038 Acmaeodera ruficaudis 1200000000025 Anthaxia sp.1 63533000135736 Anthaxia sp 2 47402000037835 Anthaxia sp. 3 534300000231030 Chrysobothris algoensis 0100000010035 Chrysobothris boschismanni 1000000000012 Sternocera orissa 4100000000038 Pseudagrilus beryllinus 312000000051122 Total 26191665000292149153

161 Appendix N

Percentage of Buprestidae species diversity and abundance for each month of the year in quadrat B on ENR

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec % Acmaeodera inscripta 0 0 00000000011 Acmaeodera aenea 0 0 00000000101 Acmaeodera albivillosa 1 1 10000001025 Acmaeodera ruficaudis 1 1 00000000013 Anthaxia sp.1 4 2 322000123524 Anthaxia sp 2 3 5 301000025523 Anthaxia sp. 3 3 2 320000012720 Chrysobothris algoensis 0 1 00000010023 Chrysobothris boschismanni 1 0 00000000011 Sternocera orissa 3 1 00000000025 Pseudagrilus beryllinus 2 1 100000003714 % 17121043000161432100

162 Appendix O

Total Buprestidae species diversity and abundance for each month of the year in quadrat C on ENR.

Species Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec Total Acmaeodera stellata 20402000003213 Acmaeodera inscripta 000000000001 1 Acmaeodera aenea 200000000010 3 Chrysobothris boschismanni 010000002013 7 Chrysobothris algoensis 100000000012 4 Anthaxia bergrothi 31210000015215 Anthaxia sp. 1 420000000245 17 Anthaxia sp. 2 368100000356 32 Anthaxia obtectans 41100000220616 Anthaxia sp. 4 1 0 0 0 0 0 0 0 0 0 2 0 3 Sphenoptera arrowi 54304000045833 Sphenoptera sinuosa 52310000025422 Lampetis conturbata 75500000023527 Lampetis gregaria 100000000000 1 Sternocera orisa 301000000001 5 Pseudagrilus beryllinus 13311000236828 Phlocteis exasperata 22100000012614 Evides pubiventris 64400000003522 Kamosiella dermestoides 31100000002310 Total 53323647 000 6204867273

163 Appendix P

Percentage of Buprestidae species diversity and abundance for each month of the year in quadrat C on ENR.

Species Jan Feb Mar Apr May Jun July Aug Sept Oct Nov Dec % Acmaeodera stellata 1010100000115 Acmaeodera inscripta 0000000000000 Acmaeodera aenea 1000000000001 Chrysobothris boschismanni 0000000010013 Chrysobothris algoensis 0000000000011 Anthaxia bergrothi 1010000000215 Anthaxia sp. 1 1100000001126 Anthaxia sp. 2 12300000012212 Anthaxia obtectans 1000000011026 Anthaxia sp. 4 0000000000101 Sphenoptera arrowi 21101000012312 Sphenoptera sinuosa 2110000001218 Lampetis conturbata 32200000011210 Lampetis gregaria 0000000000000 Sternocera orisa 1000000000002 Pseudagrilus beryllinus 01100000112310 Phlocteis exasperata 1100000000125 Evides pubiventris 2110000000128 Kamosiella dermestoides 1000000000114 % 19121313000271825100

164 Apppendix Q

Total Cerambycidae species diversity and abundance for each month and associated plant orders in all quadrats on ENR

165 Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal Total Zamium incultum 8 0 020000000010 Coptoeme krantzi 9 0 200000000011 Taurotagus klugi 9 0 050000000014 Jonthodina sculptilis 190 110000000021 Anubis clavicornis 110 001000000012 Macrotoma palmata 7 0 00000000007 Tithoes maculates 120 130000000016 Phantasis giganteus 6 0 00000000006 Dalterus dejeani 3 0 01000000004 Crossotus lacunosus 3 0 00030000006 Crossotus plumicornis 8 0 00000000008 Olenecamptus albidus 100 020000000012 Anthracocentrus capensis 6 0 231000000012 Hypoeschrus ferreirae 1 0 10000000002 Plocaederus denticornis 3 3 00100100008 Crossotus stypticus 5 0 00000020007 Nemotragus helvolus 6 0 02000000008 Hecyra terrea 0 1 00000000001 Ceroplesis thunbergi 101 100000000012 Lasiopezus longimanus 1 0 00000000001 Philematium natalense 100 020001000013 Dalterus degeeri 1 0 01000001003 Zamium bimaculatum 6 1 030002100013 Anubis mellyi 7 0 012000000010 Continued

Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal Total

166 Alphitopola octomaculata 0 0 20000000002 Mycerinicus brevis 0 0 00000010001 Phryneta spinator 0 1 00000000001 Tragiscoschema bertolinii 100 200000000012 Phyllocnema latipes 4 0 00000000004 Pacydissus sp.4 0 00000000004 Macrotoma natala 3 0 02000000005 Ossibia fuscata 6 0 00000000006 Xystrocera erosa 0 0 00000100001 Prosopocera lactator 0 0 00000000000 Xystrocera dispar 0 0 00000000000 Total 1887 122853054100253

Fabales 188 Myrtales 7 Sapindales 12 Rosales 28 Gentialanes 5 Santalales 3 Ericales 0 Malvales 5 Proteales 4 Celastrales 1 Poales 0 Malpighiales 0 Appendix R

167 Percentage of Cerambycidae species diversity and abundance for each month and associated plant orders in all quadrats on ENR

Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal % Zamium incultum 3 0 01000000004 Coptoeme krantzi 4 0 10000000004 Taurotagus klugi 4 0 02000000006 Jonthodina sculptilis 8 0 00000000008 Anubis clavicornis 4 0 00000000005 Macrotoma palmata 3 0 00000000003 Tithoes maculates 5 0 01000000006 Phantasis giganteus 2 0 00000000002 Dalterus dejeani 1 0 00000000002 Crossotus lacunosus 1 0 00010000002 Crossotus plumicornis 3 0 00000000003 Olenecamptus albidus 4 0 01000000005 Anthracocentrus capensis 2 0 11000000005 Hypoeschrus ferreirae 0 0 00000000001 Plocaederus denticornis 1 1 00000000003 Crossotus stypticus 2 0 00000010003 Nemotragus helvolus 2 0 01000000003 Hecyra terrea 0 0 00000000000 Ceroplesis thunbergi 4 0 00000000005 Lasiopezus longimanus 0 0 00000000000

Continued

168 Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal % Philematium natalense 4 0 01000000005 Dalterus degeeri 0 0 00000000001 Zamium bimaculatum 2 0 01000100005 Anubis mellyi 3 0 00100000004 Alphitopola octomaculata 0 0 10000000001 Mycerinicus brevis 0 0 00000000000 Phryneta spinator 0 0 00000000000 Tragiscoschema bertolinii 4 0 10000000005 Phyllocnema latipes 2 0 00000000002 Pacydissus sp.2 0 00000000002 Macrotoma natala 1 0 01000000002 Ossibia fuscata 2 0 00000000002 Xystrocera erosa 0 0 00000000000 Prosopocera lactator 0 0 00000000000 Xystrocera dispar 0 0 00000000000 % 743 51121022000100

Appendix S

169 Total Cerambycidae species diversity and abundance for associated plant family in all quadrats on ENR

Mi Co An Ca Pa Lo Ul Sa Rh St Pr Ce Po Eu He Species m m a e p g m p a e o l a pEbetOlaTotal Zamium incultum 8000000020000000010 Coptoeme krantzi 8001000200000000011 Taurotagus klugi 9000000050000000014 Jonthodina sculptilis 150 0400 01100000000 21 Anubis clavicornis 100 0101 00000000000 12 Macrotoma palmata 70000000000000000 7 Tithoes maculates 120 1000 30000000000 16 Phantasis giganteus 40020000000000000 6 Dalterus dejeani 30000000100000000 4 Crossotus lacunosus 10020000000000003 6 Crossotus plumicornis 60002000000000000 8 Olenecamptus albidus 100 0000 20000000000 12 Anthracocentrus capensis 5001010230000000012 Hypoeschrus ferreirae 10100000000000000 2 Plocaederus denticornis 33000100010000000 8 Crossotus stypticus 50000000002000000 7 Nemotragus helvolus 60000010100000000 8 Hecyra terrea 01000000000000000 1 Ceroplesis thunbergi 101 0000 01000000000 12

170 Continued

Species Mim Co Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola Total m Lasiopezus longimanus 10000000000000 0 0 01 Philematium natalense 90010020010000 0 0 013 Dalterus degeeri 10000000100100 0 0 03 Zamium bimaculatum 51001000321000 0 0 013 Anubis mellyi 70000200100000 0 0 010 Alphitopola octomaculata 00200000000000 0 0 02 Mycerinicus brevis 00000000001000 0 0 01 Phryneta spinator 01000000000000 0 0 01 Tragiscoschema bertolinii 70230000000000 0 0 012 Phyllocnema latipes 40000000000000 0 0 04 Pacydissus sp. 00022000000000 0 0 04 Macrotoma natala 30000000200000 0 0 05 Ossibia fuscata 50010000000000 0 0 06 Xystrocera erosa 00000000010000 0 0 01 Prosopocera lactator 00000000000000 0 0 00 Xystrocera dispar 00000000000000 0 0 00 Total 165 7 6 18 5 5 8 6 20 5 4 1 0 0 0 0 3 253

171 Appendix T

Percentage of Cerambycidae species diversity and abundance for and associated plant family in all quadrats on ENR

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola % Zamium incultum 3 0000000100000 0 0 0 4 Coptoeme krantzi 3 0000001000000 0 0 0 4 Taurotagus klugi 4 0000000200000 0 0 0 6 Jonthodina sculptilis 6 0020000000000 0 0 0 8 Anubis clavicornis 4 0000000000000 0 0 0 5 Macrotoma palmata 3 0000000000000 0 0 0 3 Tithoes maculates 5 0000010000000 0 0 0 6 Phantasis giganteus 2 0010000000000 0 0 0 2 Dalterus dejeani 1 0000000000000 0 0 0 2 Crossotus lacunosus 0 0010000000000 0 0 1 2 Crossotus plumicornis 2 0001000000000 0 0 0 3 Olenecamptus albidus 4 0000010000000 0 0 0 5 Anthracocentrus capensis 2 0000001100000 0 0 0 5 Hypoeschrus ferreirae 0 0000000000000 0 0 0 1 Plocaederus denticornis 1 1000000000000 0 0 0 3 Crossotus stypticus 2 0000000001000 0 0 0 3 Nemotragus helvolus 2 0000000000000 0 0 0 3 Hecyra terrea 0 0000000000000 0 0 0 0 Ceroplesis thunbergi 4 0000000000000 0 0 0 5

172 Continued

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola % Lasiopezus 00000000000000000 0 longimanus Philematium 40000010000000000 5 natalense Dalterus degeeri 00000000000000000 1 Zamium 20000000110000000 5 bimaculatum Anubis mellyi 30000100000000000 4 Alphitopola 00100000000000000 1 octomaculata Mycerinicus brevis 00000000000000000 0 Phryneta spinator 00000000000000000 0 Tragiscoschema 30110000000000000 5 bertolinii Phyllocnema latipes 20000000000000000 2 Pacydissus sp. 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 2 Macrotoma natala 10000000100000000 2 Ossibia fuscata 20000000000000000 2 Xystrocera erosa 00000000000000000 0 Prosopocera 00000000000000000 0 lactator Xystrocera dispar 00000000000000000 0 % 65 3 2 7 2 2 3 2 8 2 2 0 0 0 0 0 1 100

173 Appendix U

Total Cerambycidae species diversity and abundance for each month and associated plant species in all quadrats on ENR

Species A caf A kar B afr M ser S pun C afr P cap Z muc C mol D rot P caf C ery G P aus C R pyr R lan U L dis H nat X caf Total bux gra cri Zam inc 3 50 0 000 2 00 000 0000000010 Cop kra 1 71 0 002 0 00 000 0000000011 Tau klu 4 50 0 000 5 00 000 0000000014 Jon scu 0154 0 00 1 1 00 000 00000000 21 Anu cla 2 81 0 100 0 00 000 0000000012 Mac pal 0 70 0 000 0 00 000 00000000 7 Tit mac 6 60 0 030 0 00 000 0001000016 Pha gig 1 32 0 000 0 00 000 00000000 6 Dal dej 0 30 0 000 1 00 000 00000000 4 Cro lac 1 02 0 000 0 00 000 00000003 6 Cro plu 2 40 2 000 0 00 000 00000000 8 Ole alb 5 50 0 020 0 00 000 0000000012 Ant cap 2 31 0 102 3 00 000 0000000012 Hyp fer 1 00 0 000 0 00 000 00100000 2 Plo den 3 00 0 100 0 21 010 00000000 8 Cro sty 2 30 0 000 0 00 200 00000000 7 Nem hel 3 30 0 010 1 00 000 00000000 8 Hec ter 0 00 0 000 0 10 000 00000000 1 Cer thu 3 70 0 001 0 10 000 0000000012 Las lon 0 10 0 000 0 00 000 00000000 1

174 Continued

Species A caf A kar B afr M ser S pun C afr P cap Z muc C mol D rot P caf C ery G P aus C R pyr R lan U L dis H nat X caf Total bux gra cri Phi nat 3 61 0 020 0 01 000 0000000013 Dal deg 0 10 0 000 1 00 001 00000000 3 Zam bim 3 20 1 000 3 12 100 0000000013 Anu mel 5 20 0 200 1 00 000 0000000010 Alp oct 0 00 0 000 0 00 000 00020000 2 Myc bre 0 00 0 000 0 00 100 00000000 1 Phr spi 0 00 0 000 0 10 000 00000000 1 Tra ber 2 53 0 000 0 00 000 0002000012 Phy lat 1 30 0 000 0 00 000 00000000 4 Pac spp. 0 02 2 000 0 00 000 00000000 4 Mac nat 2 10 0 000 2 00 000 00000000 5 Oss fus 3 21 0 000 0 00 000 00000000 6 Xys ero 0 00 0 000 0 01 000 00000000 1 Pro lac 0 00 0 000 0 00 000 00000000 0 Xys dis 0 00 0 000 0 00 000 00000000 0 Total 58 107 18 5 5 8 6 20 6 5 4 1 1 0 0 1 5 0 0 0 3 253

Appendix V

175 Percentage of Cerambycidae species diversity and abundance for each month and associated plant species in all quadrats on ENR

Species A caf A kar B afr M ser S pun C afr P cap Z muc C mol D rot P caf C ery G P aus C R pyr R lan U L dis H nat X caf % bux gra cri Zam inc 1 20 0 000 1 00 000 00000000 4 Cop kra 0 30 0 001 0 00 000 00000000 4 Tau klu 2 20 0 000 2 00 000 00000000 6 Jon scu 0 62 0 000 0 00 000 00000000 8 Anu cla 1 30 0 000 0 00 000 00000000 5 Mac pal 0 30 0 000 0 00 000 00000000 3 Tit mac 2 20 0 010 0 00 000 00000000 6 Pha gig 0 11 0 000 0 00 000 00000000 2 Dal dej 0 10 0 000 0 00 000 00000000 2 Cro lac 0 01 0 000 0 00 000 00000001 2 Cro plu 1 20 1 000 0 00 000 00000000 3 Ole alb 2 20 0 010 0 00 000 00000000 5 Ant cap 1 10 0 001 1 00 000 00000000 5 Hyp fer 0 00 0 000 0 00 000 00000000 1 Plo den 1 00 0 000 0 10 000 00000000 3 Cro sty 1 10 0 000 0 00 100 00000000 3 Nem hel 1 10 0 000 0 00 000 00000000 3 Hec ter 0 00 0 000 0 00 000 00000000 0 Cer thu 1 30 0 000 0 00 000 00000000 5 Las lon 0 00 0 000 0 00 000 00000000 0

Continued

176 Species A caf A kar B afr M ser S pun C afr P cap Z muc C mol D rot P caf C ery G P aus C R pyr R lan U L dis H nat X caf % bux gra cri Phi nat 1 20 0 010 0 00 000 00000000 5 Dal deg 0 00 0 000 0 00 000 00000000 1 Zam bim 1 10 0 000 1 01 000 00000000 5 Anu mel 2 10 0 100 0 00 000 00000000 4 Alp oct 0 00 0 000 0 00 000 00010000 1 Myc bre 0 00 0 000 0 00 000 00000000 0 Phr spi 0 00 0 000 0 00 000 00000000 0 Tra ber 1 21 0 000 0 00 000 00010000 5 Phy lat 0 10 0 000 0 00 000 00000000 2 Pac spp. 0 01 1 000 0 00 000 00000000 2 Mac nat 1 00 0 000 1 00 000 00000000 2 Oss fus 1 10 0 000 0 00 000 00000000 2 Xys ero 0 00 0 000 0 00 000 00000000 0 Pro lac 0 00 0 000 0 00 000 00000000 0 Xys dis 0 00 0 000 0 00 000 00000000 0 % 23427 2 23 2 8 22 200 00020001100

Appendix W

177 Total Cerambycidae species diversity and abundance for each month and associated plant phenology in all quadrats on ENR

Species Yes No Total Zamium incultum 8210 Coptoeme krantzi 9211 Taurotagus klugi 9514 Jonthodina sculptilis 19 2 21 Anubis clavicornis 11 1 12 Macrotoma palmata 707 Tithoes maculates 15 1 16 Phantasis giganteus 606 Dalterus dejeani 314 Crossotus lacunosus 336 Crossotus plumicornis 808 Olenecamptus albidus 12 0 12 Anthracocentrus capensis 6612 Hypoeschrus ferreirae 202 Plocaederus denticornis 718 Crossotus stypticus 527 Nemotragus helvolus 718 Hecyra terrea 101 Ceroplesis thunbergi 11 1 12 Lasiopezus longimanus 101

Continued

178 Species Yes No Total Philematium natalense 13 0 13 Dalterus degeeri 123 Zamium bimaculatum 9413 Anubis mellyi 7310 Alphitopola octomaculata 022 Mycerinicus brevis 011 Phryneta spinator 101 Tragiscoschema bertolinii 10 2 12 Phyllocnema latipes 404 Pacydissus sp. 4 0 4 Macrotoma natala 325 Ossibia fuscata 606 Xystrocera erosa 101 Prosopocera lactator 000 Xystrocera dispar 000 Total 209 44 253

Appendix X

179 Percentage of Cerambycidae species diversity and abundance for each month and associated deciduous or non-deciduous plants in all quadrats on ENR

Species Yes No % Zamium incultum 314 Coptoeme krantzi 414 Taurotagus klugi 426 Jonthodina sculptilis 818 Anubis clavicornis 405 Macrotoma palmata 303 Tithoes maculates 606 Phantasis giganteus 202 Dalterus dejeani 102 Crossotus lacunosus 112 Crossotus plumicornis 303 Olenecamptus albidus 505 Anthracocentrus capensis 225 Hypoeschrus ferreirae 101 Plocaederus denticornis 303 Crossotus stypticus 213 Nemotragus helvolus 303 Hecyra terrea 000

Continued

180 Species Yes No % Ceroplesis thunbergi 405 Lasiopezus longimanus 000 Philematium natalense 505 Dalterus degeeri 011 Zamium bimaculatum 425 Anubis mellyi 314 Alphitopola octomaculata 011 Mycerinicus brevis 000 Phryneta spinator 000 Tragiscoschema bertolinii 415 Phyllocnema latipes 202 Pacydissus sp. 2 0 2 Macrotoma natala 112 Ossibia fuscata 202 Xystrocera erosa 000 Prosopocera lactator 000 Xystrocera dispar 000 % 83 17 100

Appendix Y

Total Cerambycidae species diversity and abundance for each month and associated plant pollination in all quadrats on ENR

181 Species Insects Insects/wind Wind Total Zamium incultum 82010 Coptoeme krantzi 92011 Taurotagus klugi 95014 Jonthodina sculptilis 19 2 0 21 Anubis clavicornis 11 1 0 12 Macrotoma palmata 7007 Tithoes maculates 12 1 3 16 Phantasis giganteus 6006 Dalterus dejeani 3104 Crossotus lacunosus 6006 Crossotus plumicornis 8008 Olenecamptus albidus 10 0 2 12 Anthracocentrus capensis 66012 Hypoeschrus ferreirae 1102 Plocaederus denticornis 4408 Crossotus stypticus 7007 Nemotragus helvolus 6118 Hecyra terrea 0101

Continued

Species Insects Insects/wind Wind Total

182 Ceroplesis thunbergi 10 2 0 12 Lasiopezus longimanus 1001 Philematium natalense 11 0 2 13 Dalterus degeeri 2103 Zamium bimaculatum 94013 Anubis mellyi 73010 Alphitopola octomaculata 0202 Mycerinicus brevis 1001 Phryneta spinator 0101 Tragiscoschema bertolinii 10 2 0 12 Phyllocnema latipes 4004 Pacydissus sp. 4 0 0 4 Macrotoma natala 3205 Ossibia fuscata 6006 Xystrocera erosa 1001 Prosopocera lactator 0000 Xystrocera dispar 0000 Total 201 44 8 253

Appendix Z

Percentage of Cerambycidae species diversity and abundance for each month and associated plant pollination in all quadrats on ENR

183 Species Insects Insects/wind Wind % Zamium incultum 3104 Coptoeme krantzi 4104 Taurotagus klugi 4206 Jonthodina sculptilis 8108 Anubis clavicornis 4005 Macrotoma palmata 3003 Tithoes maculates 5016 Phantasis giganteus 2002 Dalterus dejeani 1002 Crossotus lacunosus 2002 Crossotus plumicornis 3003 Olenecamptus albidus 4015 Anthracocentrus capensis 2205 Hypoeschrus ferreirae 0001 Plocaederus denticornis 2203 Crossotus stypticus 3003 Nemotragus helvolus 2003 Hecyra terrea 0000 Ceroplesis thunbergi 4105

Continued

Species Insects Insects/wind Wind % Lasiopezus longimanus 0000 Philematium natalense 4015

184 Dalterus degeeri 1001 Zamium bimaculatum 4205 Anubis mellyi 3104 Alphitopola octomaculata 0101 Mycerinicus brevis 0000 Phryneta spinator 0000 Tragiscoschema bertolinii 4105 Phyllocnema latipes 2002 Pacydissus sp. 2002 Macrotoma natala 1102 Ossibia fuscata 2002 Xystrocera erosa 0000 Prosopocera lactator 0000 Xystrocera dispar 0000 % 79 17 3 100

Appendix AA

Total Cerambycidae species diversity and abundance for each month and associated plant climate in all quadrats on ENR

185 Species Sub- Medium Temperat Total Tropical Temperate e Zamium incultum 30710 Coptoeme krantzi 22711 Taurotagus klugi 401014 Jonthodina sculptilis 411621 Anubis clavicornis 40812 Macrotoma palmata 0077 Tithoes maculates 601016 Phantasis giganteus 3036 Dalterus dejeani 0044 Crossotus lacunosus 6006 Crossotus plumicornis 2068 Olenecamptus albidus 50712 Anthracocentrus capensis 42612 Hypoeschrus ferreirae 1012 Plocaederus denticornis 4138 Crossotus stypticus 2057 Nemotragus helvolus 3058 Hecyra terrea 0011

Continued

Species Sub- Medium Temperat Total Tropical Temperate e Ceroplesis thunbergi 31812 Lasiopezus longimanus 0011

186 Philematium natalense 41813 Dalterus degeeri 0033 Zamium bimaculatum 32813 Anubis mellyi 70310 Alphitopola octomaculata 0022 Mycerinicus brevis 0011 Phryneta spinator 0011 Tragiscoschema bertolinii 50712 Phyllocnema latipes 1034 Pacydissus sp. 2 0 2 4 Macrotoma natala 2035 Ossibia fuscata 4026 Xystrocera erosa 0101 Prosopocera lactator 0000 Xystrocera dispar 0000 Total 84 11 158 253

Appendix AB

Percentage of Cerambycidae species diversity and abundance for each month and associated plant climate in all quadrats on ENR

Species Sub- Medium Temperat % Tropical Temperate e

187 Zamium incultum 1034 Coptoeme krantzi 1134 Taurotagus klugi 2046 Jonthodina sculptilis 2068 Anubis clavicornis 2035 Macrotoma palmata 0033 Tithoes maculates 2046 Phantasis giganteus 1012 Dalterus dejeani 0022 Crossotus lacunosus 2002 Crossotus plumicornis 1023 Olenecamptus albidus 2035 Anthracocentrus capensis 2125 Hypoeschrus ferreirae 0001 Plocaederus denticornis 2013 Crossotus stypticus 1023 Nemotragus helvolus 1023 Hecyra terrea 0000

Continued

Sub- Medium Temperat Species Tropical Temperate e % Ceroplesis thunbergi 10 35 Lasiopezus longimanus 00 00 Philematium natalense 20 35

188 Dalterus degeeri 00 11 Zamium bimaculatum 11 35 Anubis mellyi 30 14 Alphitopola octomaculata 00 11 Mycerinicus brevis 00 00 Phryneta spinator 00 00 Tragiscoschema bertolinii 20 35 Phyllocnema latipes 00 12 Pacydissus sp. 1 0 1 2 Macrotoma natala 10 12 Ossibia fuscata 20 12 Xystrocera erosa 00 00 Prosopocera lactator 00 00 Xystrocera dispar 00 00 % 33 4 63 100

Appendix AC

Total Cerambycidae species diversity and abundance for each month and associated flower size in all quadrats on ENR

189 Species Small Medium Total Large Zamium incultum 10 0 10 Coptoeme krantzi 10 1 11 Taurotagus klugi 14 0 14 Jonthodina sculptilis 17 4 21 Anubis clavicornis 11 1 12 Macrotoma palmata 707 Tithoes maculates 16 0 16 Phantasis giganteus 426 Dalterus dejeani 404 Crossotus lacunosus 426 Crossotus plumicornis 628 Olenecamptus albidus 12 0 12 Anthracocentrus capensis 11 1 12 Hypoeschrus ferreirae 202 Plocaederus denticornis 718 Crossotus stypticus 527 Nemotragus helvolus 808 Hecyra terrea 101 Ceroplesis thunbergi 12 0 12

Continued

Species Small Medium Total Large Lasiopezus longimanus 101

190 Philematium natalense 11 2 13 Dalterus degeeri 303 Zamium bimaculatum 9413 Anubis mellyi 10 0 10 Alphitopola octomaculata 202 Mycerinicus brevis 011 Phryneta spinator 101 Tragiscoschema bertolinii 9312 Phyllocnema latipes 404 Pacydissus sp. 044 Macrotoma natala 505 Ossibia fuscata 516 Xystrocera erosa 011 Prosopocera lactator 000 Xystrocera dispar 000 Total 221 32 253

Appendix AD

Percentage of Cerambycidae species diversity and abundance for associated flower size in all quadrats on ENR

191 Species Small Medium % Large Zamium incultum 404 Coptoeme krantzi 404 Taurotagus klugi 606 Jonthodina sculptilis 728 Anubis clavicornis 405 Macrotoma palmata 303 Tithoes maculates 606 Phantasis giganteus 212 Dalterus dejeani 202 Crossotus lacunosus 212 Crossotus plumicornis 213 Olenecamptus albidus 505 Anthracocentrus capensis 405 Hypoeschrus ferreirae 101 Plocaederus denticornis 303 Crossotus stypticus 213 Nemotragus helvolus 303 Hecyra terrea 000 Ceroplesis thunbergi 505

Continued

Species Small Medium % Large Lasiopezus longimanus 000 Philematium natalense 415 Dalterus degeeri 101

192 Zamium bimaculatum 425 Anubis mellyi 404 Alphitopola octomaculata 101 Mycerinicus brevis 000 Phryneta spinator 000 Tragiscoschema bertolinii 415 Phyllocnema latipes 202 Pacydissus sp. 0 2 2 Macrotoma natala 202 Ossibia fuscata 202 Xystrocera erosa 000 Prosopocera lactator 000 Xystrocera dispar 000 Total 87 13 100

Appendix AE

Total Buprestidae species diversity and abundance for associated plant orders in all quadrats on ENR

Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal Total Acmaeodera albivillosa 350000100000036 Acmaeodera viridiaenea 200000000000020 Acmaeodera aenea 150000000000015

193 Acmaeodera ruficaudis 8 0000000050013 Acmaeodera inscripta 120000000000012 Sternocera orissa 410000000000041 Anthaxia bergrothi 2300150000000038 Agrilus guerryi 0 00000230000124 Lampetis gregaria 7 000000000007 Chrysobothris 250000000000025 boschismanni Chrysobothris algoensis 230000000000023 Agrilus sexguttatus 2 00100000000012 Anthaxia sp. 1 890000000000089 Anthaxia sp. 210200000000000102 Anthaxia sp. 3 640000000000064 Trachys ziziphusii 0 00150000000015

Continued

194 Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal Total Pseudagrilus beryllinus 600000000000060 Brachelytrium transvalense 3 000000000003 Chrysobothris dorsata 2 000000000002 Acmaeodera punctatissima 5 000000000005 Kamosia tenebricosa 6 0012100000010 Agrilomorpha venosa 9 0001100000011 Agrilus falcatus 3 002000001006 Sphenoptera arrowi 8 150000102000145 Kamosiella dermestoides 100000000000010 Acmaeodera stellata 130000000000013 Phlocteis exasperata 140000000000014 Psiloptera conturbata 200070000000027 Evides pubiventris 0 02200000000022 Sphenoptera sinuosa 1 30000011600122 Anthaxia obtectans 140110000000016 Anthaxia sp. 4 2 010000000003 Total 636429513323113660 3805

Appendix AF

Percentage of Buprestidae species diversity and abundance for associated plant orders in all quadrats on ENR

195 Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal % Acmaeodera albivillosa 40 00000000004 Acmaeodera viridiaenea 20 00000000002 Acmaeodera aenea 20 00000000002 Acmaeodera ruficaudis 10 00000001002 Acmaeodera inscripta 10 00000000001 Sternocera orissa 50 00000000005 Anthaxia bergrothi 30 02000000005 Agrilus guerryi 00 00003000003 Lampetis gregaria 10 00000000001 Chrysobothris boschismanni 30 00000000003 Chrysobothris algoensis 30 00000000003 Agrilus sexguttatus 00 01000000001 Anthaxia sp. 1110000000000011 Anthaxia sp. 2130000000000013 Anthaxia sp. 3 80 00000000008 Trachys ziziphusii 00 02000000002 Pseudagrilus beryllinus 70 00000000007 Brachelytrium transvalense 00 00000000000

Continued

196 Species Fab Myr Sap Ros Gen Sant Eri Mal Pro Cel Poa Mal % Chrysobothris dorsata 0000000000000 Acmaeodera punctatissima 1000000000001 Kamosia tenebricosa 1000000000001 Agrilomorpha venosa 1000000000001 Agrilus falcatus 0000000000001 Sphenoptera arrowi 1010000120006 Kamosiella dermestoides 1000000000001 Acmaeodera stellata 2000000000002 Phlocteis exasperata 2000000000002 Lampetis conturbata 2001000000003 Evides pubiventris 0030000000003 Sphenoptera sinuosa 0000000020003 Anthaxia obtectans 2000000000002 Anthaxia sp. 4 0000000000000 % 791460031410099

Appendix AG

Buprestidae species diversity and abundance for associated plant family in all quadrats on ENR

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola Tot Acm alb 35000000000000000136

197 Acm vir 20000000000000000020 Acm ae 15000000000000000015 Acm ruf 8000000000050000013 Acm ins 12000000000000000012 Ste ori 41000000000000000041 Ant ber 23 0 0 0 0 0 0 0 15 0 0 0 0 0 0 0 0 38 Agr gue 00000000000001230024 Lam greg 700000000000000007 Chr bos 25000000000000000025 Chr algo 23000000000000000023 Agr sex 20000010000000000012 Ant sp. 1 89 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 89 Ant sp. 2 102 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 102 Ant sp. 3 64 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 64

Continued

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola Tot Tra ziz 00000000150000000015 Pse ber 60000000000000000060 Bra tra 300000000000000003

198 Chr dor 100100000000000002 Acm pun 500000000000000005 Kam ten 6000020010000000110 Agr ven 9000010000000000111 Agr fal 100020200001000006 Sph arr 815000000102000100045 Kam der 10000000000000000010 Acm ste 13000000000000000013 Phl exa 14000000000000000014 Lam con 20000000070000000027 Evi pub 00220000000000000022 Sph sin 13000000011600100022 Ant obt 14000000110000000016 Ant sp. 4 0 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 3 Totals 631 4 28 3 2 3 12 1 39 11 36 6 0 3 23 0 3 805

Appendix AH

Percentage of Buprestidae species diversity and abundance for associated plant family in all quadrats on ENR

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola Tot Acm alb 400000000000000004 Acm vir 200000000000000002

199 Acm ae 200000000000000002 Acm ruf 100000000001000002 Acm ins 100000000000000001 Ste ori 500000000000000005 Ant ber 300000002000000005 Agr gue 000000000000003003 Lam greg 100000000000000001 Chr bos 300000000000000003 Chr algo 300000000000000003 Agr sex 000000100000000001 Ant sp. 111000000000000000011 Ant sp. 213000000000000000013 Ant sp. 3800000000000000008 Tra ziz 000000002000000002 Pse ber 700000000000000007

Continued

Species Mim Com Ana Cae Pap Log Ulm Sap Rha Ste Pro Cel Poa Eup Ebe Het Ola Tot Bra tra 000000000000000000 Chr dor 000000000000000000 Acm pun 100000000000000001 Kam ten 100000000000000001 Agr ven 100000000000000001 Agr fal 000000000000000001

200 Sph arr 101000000120000006 Kam der 100000000000000001 Acm ste 200000000000000002 Phl exa 200000000000000002 Lam con 200000001000000003 Evi pub 003000000000000003 Sph sin 000000000020000003 Ant obt 200000000000000002 Ant sp. 4000000000000000000 Totals78030001051410030096

Appendix AI

Total Buprestidae species diversity and abundance for associated plant species in all quadrats on ENR

201 Species A caf A kar B afr M ser S pun Ce afr P cap Z C mol D rot P C G P aus C R pyr R U cri L dis H X caf Tot muc caf ery bux gra lan nat Acm alb 2015000000000000000000136 Acm vir 155000000000000000000020 Acm ae 510000000000000000000015 Acm ruf 5 3000000000050000000013 Acm ins 7 5000000000000000000012 Ste ori 392000000000000000000041 Ant ber 1580000015000000000000038 Agr gue 0 00000000000001002300024 Lam greg 0 700000000000000000007 Chr bos 223000000000000000000025 Chr algo 1013000000000000000000023 Agr sex 2 0 0 0 01000000000000000012 Ant sp. 13653000000000000000000089 Ant sp. 220820000000000000000000102 Ant sp. 32044000000000000000000064 Tra ziz 0 00000015000000000000015

Continued

202 Species A caf A kar B afr M ser S pun Ce afr P cap Z muc C D P caf C G P C R R U cri L dis H nat X caf Tot mol rot ery bux aus gra pyr lan Pse ber 258000000000000000000060 Bra tra 0300000000000000000003 Chr dor 0110000000000000000002 Acm pun 1400000000000000000005 Kam ten 15002001000000000000110 Agr ven 36001000000000000000111 Agr fal 0102020000001000000006 Sph arr 2600000001020100150000045 Kam der 46000000000000000000010 Acm ste 211000000000000000000013 Phl exa 311000000000000000000014 Lam con 812000007000000000000027 Evi pub 000000000000000000220022 Sph sin 010000002116100100000022 Ant obt 410000011000000000000016 Ant sp. 4 0020000000000000100003 Totals2264053231213921136260351232203805

Appendix AJ

Percentage of Buprestidae species diversity and abundance for associated plant species in all quadrats on ENR

203 Species A caf A kar B afr M ser S pun Ce afr P cap Z muc C D P caf C G P C R R U cri L dis H nat X caf % mol rot ery bux aus gra pyr lan Acm alb 2200000000000000000004 Acm vir 2100000000000000000002 Acm ae 1100000000000000000002 Acm ruf 1000000000001000000002 Acm ins 1100000000000000000001 Ste ori 5000000000000000000005 Ant ber 2100000200000000000005 Agr gue 0000000000000000030003 Lam greg 0100000000000000000001 Chr bos 0300000000000000000003 Chr algo 1200000000000000000003 Agr sex 0000010000000000000001 Ant sp. 1 4 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 11 Ant sp. 2 2 10 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 13 Ant sp. 32 5 0 0 0 0 0 0000000000 00 008 Tra ziz 0000000200000000000002 Pse ber 0700000000000000000007

Continued

Species A caf A kar B afr M ser S pun Ce afr P cap Z muc C D P caf C G P C R R U cri L dis H nat X caf % mol rot ery bux aus gra pyr lan Bra tra 0 000000000000000000000

204 Chr dor 0 000000000000000000000 Acm pun 0 000000000000000000001 Kam ten 0 100000000000000000001 Agr ven 0 100000000000000000001 Agr fal 0 000000000000000000001 Sph arr 0 100000001200001000006 Kam der 0 100000000000000000001 Acm ste 0 100000000000000000002 Phl exa 0 100000000000000000002 Lam con 1 100000100000000000003 Evi pub 0 000000000000000003003 Sph sin 0 000000000200000000003 Ant obt 0 100000000000000000002 Ant sp. 40000000000000000000000 % 2850000105014010010330097

Appendix AK

Total Buprestidae species diversity and abundance for associated plant phenology in all quadrats on ENR

Species Yes No Total

205 Acmaeodera albivillosa 35 1 36 Acmaeodera viridiaenea 20 0 20 Acmaeodera aenea 15 0 15 Acmaeodera ruficaudis 85 13 Acmaeodera inscripta 12 0 12 Sternocera orissa 41 0 41 Anthaxia bergrothi 23 15 38 Agrilus guerryi 024 24 Lampetis gregaria 70 7 Chrysobothris boschismanni 25 0 25 Chrysobothris algoensis 23 0 23 Agrilus sexguttatus 12 0 12 Anthaxia sp. 1 89 0 89 Anthaxia sp. 2 102 0 102 Anthaxia sp. 3 64 0 64 Trachys ziziphusii 015 15 Pseudagrilus beryllinus 60 0 60 Brachelytrium transvalense 30 3

Continued

Species Yes No Total Chrysobothris dorsata 20 2 Acmaeodera punctatissima 50 5 Kamosia tenebricosa 64 10 Agrilomorpha venosa 92 11

206 Agrilus falcatus 51 6 Sphenoptera arrowi 24 21 45 Kamosiella dermestoides 10 0 10 Acmaeodera stellata 13 0 13 Phlocteis exasperata 14 0 14 Lampetis conturbata 20 7 27 Evides pubiventris 22 0 22 Sphenoptera sinuosa 517 22 Anthaxia obtectans 14 2 16 Anthaxia sp. 4 2 1 3 Total 690 115 805

Appendix AL

Percentage of Buprestidae species diversity and abundance for associated deciduous or non-deciduous plants in all quadrats on ENR

Species Yes No % Acmaeodera albivillosa 404 Acmaeodera viridiaenea 202

207 Acmaeodera aenea 202 Acmaeodera ruficaudis 112 Acmaeodera inscripta 101 Sternocera orissa 505 Anthaxia bergrothi 325 Agrilus guerryi 033 Lampetis gregaria 101 Chrysobothris boschismanni 303 Chrysobothris algoensis 303 Agrilus sexguttatus 101 Anthaxia sp. 1 11 0 11 Anthaxia sp. 2 13 0 13 Anthaxia sp. 3 8 0 8 Trachys ziziphusii 022 Pseudagrilus beryllinus 707 Brachelytrium transvalense 000

Continued

Species Yes No % Chrysobothris dorsata 000 Acmaeodera punctatissima 101 Kamosia tenebricosa 101 Agrilomorpha venosa 101 Agrilus falcatus 101 Sphenoptera arrowi 336

208 Kamosiella dermestoides 101 Acmaeodera stellata 202 Phlocteis exasperata 202 Lampetis conturbata 213 Evides pubiventris 303 Sphenoptera sinuosa 123 Anthaxia obtectans 202 Anthaxia sp. 4 0 0 0 % 86 14 100

Appendix AM

Total Buprestidae species diversity and abundance for associated plant pollination in all quadrats on ENR

Species Insects Insects/wind Wind Total Acmaeodera albivillosa 36 0 0 36 Acmaeodera viridiaenea 20 0 0 20

209 Acmaeodera aenea 15 0 0 15 Acmaeodera ruficaudis 13 0 0 13 Acmaeodera inscripta 12 0 0 12 Sternocera orissa 41 0 0 41 Anthaxia bergrothi 23 15 0 38 Agrilus guerryi 24 0 0 24 Lampetis gregaria 7007 Chrysobothris 25 0 0 25 boschismanni Chrysobothris algoensis 23 0 0 23 Agrilus sexguttatus 201012 Anthaxia sp. 1 89 0 0 89 Anthaxia sp. 2 102 0 0 102 Anthaxia sp. 3 64 0 0 64 Trachys ziziphusii 015015 Pseudagrilus beryllinus 60 0 0 60 Brachelytrium transvalense 3003

Continued

Species Insects Insects/wind Wind Total Chrysobothris dorsata 2002 Acmaeodera punctatissima 5005 Kamosia tenebricosa 73010 Agrilomorpha venosa 10 1 0 11 Agrilus falcatus 4026 Sphenoptera arrowi 39 6 0 45

210 Kamosiella dermestoides 10 0 0 10 Acmaeodera stellata 13 0 0 13 Phlocteis exasperata 14 0 0 14 Lampetis conturbata 20 7 0 27 Evides pubiventris 22 0 0 22 Sphenoptera sinuosa 19 3 0 22 Anthaxia obtectans 14 2 0 16 Anthaxia sp. 4 2 1 0 3 Total 740 53 12 805

Appendix AN

Percentage of Buprestidae species diversity and abundance for associated plant pollination in all quadrats on ENR

Species Insects Insects/wind Wind % Acmaeodera albivillosa 4004 Acmaeodera viridiaenea 2002 Acmaeodera aenea 2002 Acmaeodera ruficaudis 2002 Acmaeodera inscripta 1001

211 Sternocera orissa 5005 Anthaxia bergrothi 3205 Agrilus guerryi 3003 Lampetis gregaria 1001 Chrysobothris 3003 boschismanni Chrysobothris algoensis 3003 Agrilus sexguttatus 0011 Anthaxia sp. 1 11 0 0 11 Anthaxia sp. 2 13 0 0 13 Anthaxia sp. 3 8 0 0 8 Trachys ziziphusii 0202 Pseudagrilus beryllinus 7007 Brachelytrium transvalense 0000

Continued

Species Insects Insects/wind Wind % Chrysobothris dorsata 0000 Acmaeodera punctatissima 1001 Kamosia tenebricosa 1001 Agrilomorpha venosa 1001 Agrilus falcatus 0001 Sphenoptera arrowi 5106 Kamosiella dermestoides 1001 Acmaeodera stellata 2002

212 Phlocteis exasperata 2002 Lampetis conturbata 2103 Evides pubiventris 3003 Sphenoptera sinuosa 2003 Anthaxia obtectans 2002 Anthaxia sp. 4 0 0 0 0 % 92 7 1 100

Appendix AO

Total Buprestidae species diversity and abundance for associated plant climate in all quadrats on ENR

Species Sub- Medium Temperat Total Tropical Temperate e Acmaeodera albivillosa 21 0 15 36 Acmaeodera viridiaenea 15 0 5 20 Acmaeodera aenea 501015 Acmaeodera ruficaudis 50813 Acmaeodera inscripta 70512

213 Sternocera orissa 39 0 2 41 Anthaxia bergrothi 15 0 23 38 Agrilus guerryi 102324 Lampetis gregaria 0077 Chrysobothris 202325 boschismanni Chrysobothris algoensis 10 0 13 23 Agrilus sexguttatus 201012 Anthaxia sp. 1 36 0 53 89 Anthaxia sp. 2 20 0 82 102 Anthaxia sp. 3 20 0 44 64 Trachys ziziphusii 001515 Pseudagrilus beryllinus 205860 Brachelytrium transvalense 0033

Continued

Species Sub- Medium Temperat Total Tropical Temperate e Chrysobothris dorsata 1012 Acmaeodera punctatissima 1045 Kamosia tenebricosa 40610 Agrilomorpha venosa 50611 Agrilus falcatus 0066 Sphenoptera arrowi 3103245 Kamosiella dermestoides 40610 Acmaeodera stellata 201113

214 Phlocteis exasperata 301114 Lampetis conturbata 801927 Evides pubiventris 002222 Sphenoptera sinuosa 112022 Anthaxia obtectans 411116 Anthaxia sp. 4 2 0 1 3 Total 238 12 555 805

Appendix AP

Percentage of Buprestidae species diversity and abundance for associated plant climate in all quadrats on ENR

Species Sub- Medium Temperate % Tropical Temperate Acmaeodera albivillosa 30 24 Acmaeodera viridiaenea 20 12 Acmaeodera aenea 10 12 Acmaeodera ruficaudis 10 12 Acmaeodera inscripta 10 11 Sternocera orissa 50 05

215 Anthaxia bergrothi 20 35 Agrilus guerryi 00 33 Lampetis gregaria 00 11 Chrysobothris 00 33 boschismanni Chrysobothris algoensis 10 23 Agrilus sexguttatus 00 11 Anthaxia sp. 1 4 0 7 11 Anthaxia sp. 2 2 0 10 13 Anthaxia sp. 3 2 0 5 8 Trachys ziziphusii 00 22 Pseudagrilus beryllinus 00 77 Brachelytrium transvalense 00 00

Continued

Species Sub- Medium Temperat % Tropical Temperate e Chrysobothris dorsata 0000 Acmaeodera punctatissima 0001 Kamosia tenebricosa 0011 Agrilomorpha venosa 1011 Agrilus falcatus 0011 Sphenoptera arrowi 0146 Kamosiella dermestoides 0011 Acmaeodera stellata 0012 Phlocteis exasperata 0012

216 Lampetis conturbata 1023 Evides pubiventris 0033 Sphenoptera sinuosa 0023 Anthaxia obtectans 0012 Anthaxia sp. 4 0 0 0 0 % 30 1 69 100

Appendix AQ

Total Buprestidae species diversity and abundance for associated flower size in all quadrats on ENR

Species Small Medium Total Large Acmaeodera albivillosa 36 0 36 Acmaeodera viridiaenea 20 0 20 Acmaeodera aenea 15 0 15 Acmaeodera ruficaudis 13 0 13 Acmaeodera inscripta 12 0 12 Sternocera orissa 41 0 41

217 Anthaxia bergrothi 38 0 38 Agrilus guerryi 24 0 24 Lampetis gregaria 707 Chrysobothris 25 0 25 boschismanni Chrysobothris algoensis 23 0 23 Agrilus sexguttatus 12 0 12 Anthaxia sp. 1 89 0 89 Anthaxia sp. 2 102 0 102 Anthaxia sp. 3 64 0 64 Trachys ziziphusii 15 0 15 Pseudagrilus beryllinus 60 0 60 Brachelytrium transvalense 303

Continued

Species Small Medium Total Large Chrysobothris dorsata 112 Acmaeodera punctatissima 505 Kamosia tenebricosa 10 0 10 Agrilomorpha venosa 11 0 11 Agrilus falcatus 426 Sphenoptera arrowi 15 30 45 Kamosiella dermestoides 10 0 10 Acmaeodera stellata 13 0 13 Phlocteis exasperata 14 0 14 Lampetis conturbata 27 0 27

218 Evides pubiventris 22 0 22 Sphenoptera sinuosa 51722 Anthaxia obtectans 16 0 16 Anthaxia sp. 4 1 2 3 Total 753 52 805

Appendix AR

Percentage of Buprestidae species diversity and abundance for associated flower size in all quadrats on ENR

Species Small Medium % Large Acmaeodera albivillosa 404 Acmaeodera viridiaenea 202 Acmaeodera aenea 202 Acmaeodera ruficaudis 202 Acmaeodera inscripta 101 Sternocera orissa 505

219 Anthaxia bergrothi 505 Agrilus guerryi 303 Lampetis gregaria 101 Chrysobothris 303 boschismanni Chrysobothris algoensis 303 Agrilus sexguttatus 101 Anthaxia sp. 1 11 0 11 Anthaxia sp. 2 13 0 13 Anthaxia sp. 3 8 0 8 Trachys ziziphusii 202 Pseudagrilus beryllinus 707 Brachelytrium transvalense 000

Continued

Species Small Medium % Large Chrysobothris dorsata 000 Acmaeodera punctatissima 101 Kamosia tenebricosa 101 Agrilomorpha venosa 101 Agrilus falcatus 001 Sphenoptera arrowi 246 Kamosiella dermestoides 101 Acmaeodera stellata 202 Phlocteis exasperata 202 Lampetis conturbata 303 Evides pubiventris 303

220 Sphenoptera sinuosa 123 Anthaxia obtectans 202 Anthaxia sp. 4000 %946100

Appendix AS

Cerambycidae light trap results for 2001 on Ezemvelo Nature Reserve

Ja Ju Ju Se Species nFebMarAprMayn lAugpOctNovDecTotal Phantasis giganteus 20 30 10001 1 13 12 Dalterus dejeani 00 01 10001 1 12 7 Crossotus lacunosus 30 20 00000 0 20 7 Crossotus plumicornis 10 10 00001 2 11 7 Olenecamptus albidus 01 12 00000 0 11 6 Alphitopola octomaculata 00 00 31123 0 32 15

221 Mycerinicus brevis 10 00 00000 0 01 2 Phryneta spinator 22 10 00000 1 41 11 Tragiscoschema bertolinii 03 00 00010 3 12 10 Xystrocera erosa 01 11 00000 1 60 10 Prosopocera lactator 03 00 00000 3 33 12 Total 9 10 9 4 5 1 1 3 6 12 23 16 99

Appendix AT

Buprestidae light trap results for 2001on Ezemvelo Nature Reserve

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Acmaeodera albivillosa 0100000000000 Total 010 0 0 00 0 0 0 00 1

222 .

223 224 225