Biological Control of Tecoma Stans L
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Suitability of the leaf-mining fly, Pseudonapomyza sp. (Diptera: Agromyzidae), for biological control of Tecoma stans L. (Bignoniaceae) in South Africa Thesis Submitted in fulfillment of the requirements for the degree of MASTER OF SCIENCE of Fort Hare University By LULAMA GRACIOUS MADIRE Department of Zoology and Entomology University of Fort Hare Private Bag x 1314 Alice 5700 South Africa April 2010 DECLARATIO Suitability of the leaf-mining fly, Pseudonapomyza sp. (Diptera: Agromyzidae), for biological control of Tecoma stans L. (Bignoniaceae) in South Africa I, Lulama Gracious Madire, hereby 1. grant the University of Fort Hare free license to reproduce the above thesis in whole or in part, for the purpose of research; 2. declare that: • The above thesis is my own unaided work, both in concept and execution, and that apart from the normal guidance from my supervisors; I have received no assistance except as stated below. • Neither the substance nor any part of the above thesis has been submitted in the past, or is being, or is to be submitted for a degree in the University or any other university. The thesis has been presented by me for examination for the degree of MSc. 2 ABSTRACT Tecoma stans (L.) Juss. Ex Kunth (Bignoniaceae) also known as yellow bells, has a native distribution from Northern Argentina, central America, Mexico and the Southern USA. In many warm climatic regions of the world, T. stans is commonly planted as an ornamental plant because of its yellow flowers, hence the name yellow bells, and pinnate foliage. As a result, this evergreen shrub has wide distribution in the tropical and subtropical parts of the western hemisphere. As is the case in many other parts of the world, T. stans was introduced into South Africa as an ornamental plant, but escaped cultivation and now invades roadsides, urban open spaces, watercourses, rocky sites in subtropical and tropical areas of five South African provinces; Gauteng, Mpumalanga, Limpopo, KwaZulu-Natal, Eastern Cape and neighboring countries. Tecoma stans has the potential of extending its range because its seeds are easily dispersed by wind. The purpose of this work was to carry out pre-release efficacy studies to determine the host specificity and suitability of Pseudonapomyza sp. (Diptera: Agromyzidae), a leaf-mining fly, as a biological control agent of T. stans . Available information suggests that the fly was brought to South Africa (SA) from Argentina in 2005. In that year a worker collected adult root feeding flea- beetles from T. stans and their eggs by collecting soil around the plants in the Argentinian Province of Jujuy, at San Pedro (24˚12’592”S, 64˚51’328”W). The soil was brought to the SA quarantine laboratory of the Agricultural Research Council, Plant Protection Research Institute (Weeds Division), Pretoria, and placed in a cage containing T. stans plants for flea-beetle larvae to emerge from the eggs. The Pseudonapomyza sp. flies which emerged from that soil were reared to produce a colony of flies used in the study reported here. The feeding behavior of Pseudonapomyza sp. adults is initiated by females which use their ovipositor to puncture holes in the leaf mesophyll and then they feed on the sap oozing from the 3 holes. Since males have no means of puncturing the leaves, they feed from holes made by females. Eggs are laid singly into the tubular leaf punctures. Soon after hatching, the larva feeds on the leaf mesophyll tissue. As the larva feeds within the leaf it creates mines which eventually coalesce to form large blotches. The damaged leaf area reduces the photosynthetic potential of the plant especially when damaged leaves dry and fall off the plants. The potential of Pseudonapomyza sp. as a biocontrol agent is enhanced by the fact that it has a high level of fecundity and a short life cycle. As a result, its populations can build up rapidly to exert a significant impact on T. stans . Host-specificity tests undertaken on 35 plant species in 12 plant families showed that out of the 35 plant species tested, the fly was able to develop on T. stans only . Although Pseudonapomyza sp. adults fed on T. capensis , a South African indigenous ornamental shrub, no larval mines were observed on this plant. This suggests two possibilities; either females of Pseudonapomyza sp. do not oviposit on T. capensis or oviposition takes place but larvae cannot feed and develop on this plant. These studies indicate that this fly is sufficiently host-specific, and can be released against T. stans without posing any threat to either commercial or indigenous plant species grown in South Africa. Experimental designs simulating high populations of Pseudonapomyza sp. showed that the impact of leaf mining fly on T. stans can cause approximately 56% aboveground biomass reduction. Other concurrent studies have also showed that low and high density fly infestations can cause 23% and 48 % belowground biomass reductions, respectively. Based on the available information, it appears that Pseudonapomyza sp. may have the potential to reduce the invasive capacity of T. stans in the affected areas. 4 In order to exert more herbivore pressure on T. stans , it is suggested that agents belonging to other feeding guilds, such as root-, stem- and seed-feeding insects, be considered for release to complement the leaf-feeding of Pseudonapomyza sp. An application to release this fly in SA has been submitted to one of the two regulatory authorities. 5 ACKOWLEDGEMETS I am so grateful to Professor S. Waladde my supervisor and Dr David Simelane, my co-supervisor, for their guidance and support. I’m also thankful to Weeds Division (Pretoria) staff for their support and technical assistance. I would like to thank the staff from South African National Collection of Insects (PPRI) for their help towards identification of the leafminer. I would also like to thank Ms V. April and Mr T. Chauke for their technical assistance during the study. I also want to thank my kids and husband for their prayers and encouragement. I would like to thank ARC-PPRI and WfW for funding the study. Lastly, I would like thank my God who gave me strength and courage during my studies, without Him this would not be possible. 6 TABLE OF COTETS TITLE PAGE 1 DECLARATION 2 ABSTRACT 3 ACKNOWLEDGEMENTS 6 TABLE OF CONTENTS 7 LIST OF TABLES 10 LIST OF FIGURES 11 1. INTRODUCTION 12 1.1 Statement of the problem 12 1.1.1 Justification for the study 12 1.1.2 Study objectives 13 2. LITERATURE REVIEW 14 2.1 Alien plants 14 2.2 Distribution and impact of Tecoma stans 15 2.3 Biology and ecology of Tecoma stans 20 2.4 Control methods 21 2.4.1 Mechanical 21 2.4.2 Chemical 21 2.4.3 Biological control 21 2.5 Biology of the leaf-mining fly, Pseudonapomyza sp. 23 2.6 Host-specificity tests on Pseudonapomyza sp. 25 2.7 Potential impact of Pseudonapomyza sp. on Tecoma stans 27 3. MATERIALS AND METHODS 31 7 3.1 Laboratory conditions and test plants 31 3.1.1 Pseudonapomyza sp. colony 32 3.1.2 Pre-oviposition period 32 3.1.3 Developmental and generation periods 33 3.1.4 Reproductive output 33 3.2 Field surveys in South Africa 34 3.3 Host-specificity studies 34 3.3.1 No-choice tests 34 3.3.2 Multi-choice tests 35 3.4 Potential impact of Pseudonapomyza sp. on Tecoma stans 39 3.4.1 Larval densities and adult emergence at different inoculation densities 40 3.4.2 Effect of leaf mining on growth of T. stans 40 3.4.3 Effect of leaf mining on T. stans biomass 41 3.5 Data analysis 41 4. RESULTS 43 4.1 Biology of the leafminer, Pseudonapomyza sp. 43 4.1.1 Appearance, feeding and oviposition behaviour of Pseudonapomyza sp. 43 4.1.2 Pre-oviposition period 44 4.1.3 Reproductive output 44 4.1.4 Developmental period (from egg to adult) 44 4.1.5 Generation period and number of generations per year 44 4.2 Host-specificity of the leafminer, Pseudonapomyza sp. 48 4.2.1 Field surveys in South Africa 48 4.2.2 No-choice tests 49 4.2.3 Multi-choice tests 49 8 4.3 Potential impact of the leafminer, Pseudonapomyza sp. on T. stans 52 4.3.1 Larval densities and adult emergence at different fly densities 52 4.3.2 Effect of leaf mining on growth of T. stans 53 4.3.3 Effect of leaf mining on T. stans biomass 58 5 DISCUSSION 61 5.1 Origin of the Pseudonapomyza sp. 61 5.2 Potential and limitations of the leafminer as a biocontrol agent 61 5.3 Host-specificity 66 5.4 Impact of Pseudonapomyza sp. 67 5.5 Release of multiple biological control agents on T. stans 69 6. CONCLUSION 71 REFERENCES 72 9 LIST OF TABLES 1. List of test plants used during no-choice tests on Pseudonapomyza sp. 37 2. Randomly arranged test plants in five rows in a walk-in cage 38 3. Field survey for Pseudonapomyza sp. in South Africa 50 4. Feeding damage of Pseudonapomyza sp. on various test plant species in no-choice tests 51 5. Response of Pseudonapomyza sp. on closely related Bignoniaceae plant species in multi-choice tests 52 10 LIST OF FIGURES 1 Distribution map of Tecoma stans in South USA 16 2. Tecoma stans plant with flowers and open pods . 17 3. Tecoma stans infestation at Wonderboom, Pretoria 18 4. Distribution map of Tecoma stans in South Africa 19 5. Small cages used for no-choice bioassays done inside the quarantine glasshouse 38 6.