The Phenologies of Macadamia (Proteaceae) and Thrips (Insecta: Thysanoptera) Communities in Mpumalanga Province, South Africa
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The Phenologies of Macadamia (Proteaceae) and Thrips (Insecta: Thysanoptera) Communities in Mpumalanga Province, South Africa A thesis submitted in fulfilment of the requirements for the Degree of DOCTOR OF PHILOSOPHY of RHODES UNIVERSITY by COLLEEN HEPBURN December 2015 Abstract Macadamia (Proteaceae) is indigenous to Australia; four of the nine species have been used in plant breeding for cultivation and the production of a rapidly growing product. First introduced to Hawaii in the 1880s, macadamias’ potential as a crop was only really considered in 1910. It is due to the early Hawaiian pioneers’ research and the development of Hawaiian cultivars upon which most of the current world’s production is based today. Prior to the 1980s, cultivation in Australia was based mainly on Hawaiian cultivars. Extensive research has been conducted on cultivar development, performance, best practice, production, crop protection and processing. South Africa has become one of the world’s leading producers and exporters of macadamias. The cultivation of macadamia in South Africa began 80 years ago, but the phenology of macadamia has not been rigorously studied under local conditions, and neither has the thrips communities in macadamia. Data collected during this study on the macadamia phenological stages can be used as a foundation on which seasonal data of all insect pests can be plotted, as a basis on which an extensive Integrated Pest Management (IPM) programme can be developed for the macadamia industry in South Africa. Extensive sampling of thrips communities present in macadamias was undertaken over two years, with more than 142,000 individuals collected. The most basic, common trait into which specimens could be sorted were colour groups. Specimens from each colour group were identified through slide-mounting individuals. Fifteen species of thrips were identified to species-level, excluding specimens from the Panchaetothripinae, Haplothrips and Scolothrips which occurred only periodically. The presence of larvae indicated that macadamia is a host plant for some species. Due to their size and morphology, identification i of the total collection could not be processed to species level using a dissection microscope, although Scirtothrips aurantii Faure were distinguishable from other species and their distribution over the seasons observed. Due to logistical constraints and events beyond the author’s control, the results are mainly qualitative. There seems to be phenological variation within and between cultivars. The date of anthesis and duration of immature nut-drop was estimated; some cultivars set out- of-season flowers and nuts; this “secondary crop” could have far-reaching effects on the implementation of an IPM programme. Comparisons of the phenological data of macadamia and thrips showed the highest abundances and diversity of thrips species occurred when flowers were present. An extensive study of abscised nuts showed no significant difference between most cultivars or at the sites where management practices were implemented and those sites where these practices were not. There was however a consistent difference between the two seasons. There was no difference in maturity of abscised nuts, regardless of whether thrips damage to the husk was present or not. Excessive Bathycoelia distincta (Hemiptera: Pentatomidae) damage was found on the kernels of the abscised nuts processed for maturity testing, which is more likely to have been the primary cause of abscission. The Author’s opinion is that thrips damage to the outer husk does not lead to abscission of maturing nuts. Ascertaining the cause of abscission at nut-set and immature nut-drop is more problematic as there are numerous other potential causes. Damage to new flush, especially the late summer flush essential for the build-up of carbohydrates and resources for the following yield, should be carefully monitored and controlled when necessary. ii This thesis is dedicated in loving memory of my late husband, Randall Hepburn, my friend, colleague, teacher, mentor, and champion; the person who shared with me his incomparable knowledge and love of insects... and that’s how it all began. Con te partird. iii Declaration The following thesis has not been submitted to any university other than Rhodes University, Grahamstown, South Africa. The work presented here is that of the author. DATE: 30 March 2017 iv Acknowledgements I wish to acknowledge my late parents, Denis and Shirley Ardren, who laid the foundation and guided me through life to where and who I am today. To my friend and mentor the late Len Hobson; a truly remarkable gentleman, also an alumnus of Rhodes University, Grahamstown. Len fueled my enthusiasm, answered my many questions and handed over to me his macadamia literature and correspondence for safekeeping. I am extremely fortunate to have had Len’s knowledge, guidance and friendship. My grateful thanks also go to: Martin Villet, Department of Zoology & Entomology, Rhodes University, Grahamstown, South Africa, for once again being my supervisor, for his assistance, guidance and support over the years. Michael Stiller, of the Agricultural Research Council, Plant Protection Research, Biosystematics Division, Pretoria, South Africa, for sharing his knowledge, helping me learn about thrips identification and identifying the thrips communities in macadamias. His enthusiasm, encouragement and patience is much appreciated. Prof Sarah Radloff, Department of Statistics, Rhodes University, Grahamstown, South Africa, long-time friend and colleague, for her encouragement, support and help with the statistical analyses. Keenan van Zuydam, my eldest son, whose help with the extensive field work for this project, and the countless other entomology projects he’s been involved in over the many years, has proved invaluable; a big thank you my boy. Keenan has been exceptionally supportive and encouraging, questioning and discussing ideas, and even when obstacles were placed in the way, he’s pushed me to keep going. The late Johannes Ngomane, who was involved from the beginning with the field work for this project; he was of enormous help. He and I worked shoulder to shoulder throughout the seasons; his presence is sorely missed. Ngiyabonga Johannes, hamba kahle. Neil van Zuydam, my younger son, thank you for your encouragement and support, lifting my spirits and making me smile. To my extended family and friends, Kirsten Hepburn, David and Liz Hepburn, Ashton and Sara Hepburn, Rowan Hepburn, Cynthia and Charles Boudreaux, Catherine Stewart, Stuart and Sanet Ardren, Christian and Catherine Pirk, Jessica Cockburn, Tarryn v Goble, Adrian and Cheryl Craig, a big thank you for all your love, support and encouragement. Candice Marshall, Manager: Administration and Technical Support Sections, Department of Zoology & Entomology, Rhodes University, Grahamstown, South Africa, thank you for the administrative assistance over the past years, helping me run this project from a distance. John Gillam, Manager: Postgraduate Funding, Research Office, Rhodes University, Grahamstown, South Africa, for his amazing commitment and ability to sort out funding glitches, thank you John. Eileen Sheppard, Senior Librarian Science Faculty, Rhodes University Library, Grahamstown, South Africa, and her staff for accessing and obtaining copies of numerous publications I needed. Dr Schalk Schoeman, Agricultural Research Council, Institute of Tropical and Subtropical Crops, Nelspruit, South Africa, for his encouragement and help with study sites. The owner and his manager of the Commercial Farm where some of the study sites were located, thank you for access to your orchards. Citrus Research International (CRI), Nelspruit, South Africa, for kind permission to use their equipment to enable us to photograph and identify the various species of thrips. South African Macadamia Growers’ Association (SAMAC) for partially funding this research. Scholarship funding provided by The Ernst & Ethel Eriksen Trust, Bryanston, South Africa, for 2011, 2012 and 2013 is gratefully acknowledged. Partial funding was provided by the National Research Foundation (NRF) of South Africa. Any opinion, findings and conclusions or recommendations expressed in this material are those of the author and do not necessarily reflect the views of the National Research Foundation. vi Table of Contents THE PHENOLOGIES OF MACADAMIA (PROTEACEAE) AND THRIPS (INSECTA: THYSANOPTERA) COMMUNITIES IN MPUMALANGA PROVINCE, SOUTH AFRICA I ABSTRACT..............................................................................................................................................I DECLARATION...................................................................................................................................IV ACKNOWLEDGEMENTS...................................................................................................................V TABLE OF CONTENTS....................................................................................................................VII LIST OF TABLES...............................................................................................................................XII LIST OF FIGURES............................................................................................................................XIV LIST OF ABBREVIATIONS..........................................................................................................XXII DISCLAIMER.................................................................................................................................XXIV