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The Ecology of Raoiella Indica (Hirst: Tenuipalpidae) In The ecology of Raoiella indica (Hirst) (Acari:Tenuipalpidae) in India and Trinidad The ecology of Raoiella indica (Hirst: Tenuipalpidae) in India and Trinidad: Host plant relations and predator: prey relationships Arabella Bryony K. Taylor (CID: 00459677) PhD Thesis June 2017 Imperial College London Department of Life Sciences CABI Egham, UK 1 The ecology of Raoiella indica (Hirst) (Acari:Tenuipalpidae) in India and Trinidad Copyright declaration The copyright of this thesis rests with the author and is made available under a Creative Commons Attribution Non-Commercial No Derivatives licence. Researchers are free to copy, distribute or transmit the thesis on the condition that they attribute it, that they do not use it for commercial purposes and that they do not alter, transform or build upon it. For any reuse or redistribution, researchers must make clear to others the licence terms of this work. I certify that the contents of this thesis are my own work and the works by other authors are appropriately referenced. Some of the work described in chapter 4 of this thesis has been previously published in Taylor et al. (2011). 2 The ecology of Raoiella indica (Hirst) (Acari:Tenuipalpidae) in India and Trinidad Abstract Red Palm Mite, Raoiella indica (Acari:Tenuipalpidae) (RPM), an Old World species first recorded in India (1924), was reported historically on a small number of host species of Arecaceae (palms) throughout Asia and the Middle East. In 2004, the mite invaded the New World resulting in high population densities and apparent new host associations- including Musa spp. (bananas and plantains). Subsequently, RPM has become widely established in the tropical Americas. The aim of the work was to understand some of the factors underlying the differences in RPM population status between the two regions by comparing morphological and molecular characteristics, investigating Old World host relations and comparing predator–prey interactions in India and Trinidad. Field population densities on main host Cocos nucifera were significantly higher in Trinidad compared to India. Substantial RPM field densities and breeding colonies were confirmed on Musa spp. in Trinidad, but not India; although RPM was shown to survive and lay eggs on Musa spp. in vitro in India. Arthropod predator diversity in India was higher on C. nucifera and lower on Musa spp. compared to Trinidad. Surveys on both host species only recorded predatory mites in Trinidad, whereas predatory insects and mites were recorded in India. Although molecular studies were unsuccessful, morphological studies showed that, independent of host plant, RPM had significantly longer lateral setae in the Caribbean compared to India. Adventive range populations may have resulted from a genetic bottleneck upon introduction. Assays in Trinidad showed that RPM with droplets on dorsal setae were contacted and consumed by Amblyseius largoensis on fewer occasions than those without. Setae may also play a role in defence against A. largoensis. Studies confirmed differences in RPM severity and status on host plants between regions and suggest natural enemies play a role in maintaining a narrow host range in India. 3 The ecology of Raoiella indica (Hirst) (Acari:Tenuipalpidae) in India and Trinidad Contents Copyright declaration................................................................................................................. 2 Abstract ...................................................................................................................................... 3 Contents ..................................................................................................................................... 4 1.0 Introduction .......................................................................................................................... 8 2.0 Raoiella indica: A literature review ................................................................................... 12 2.1 Mites, the Tenuipalpidae and the genus Raoiella .......................................................... 12 2.2 Raoiella indica: life cycle .............................................................................................. 13 2.3 Host plants of Raoiella indica........................................................................................ 17 2.4 Biogeography ................................................................................................................. 18 2.5 Predator relations ........................................................................................................... 22 2.6 Raoiella indica- Population dynamics ........................................................................... 27 2.7 Invasion ecology theories .............................................................................................. 28 2.8 Host association, host races and biotypes. ..................................................................... 31 2.9 Questions relating to observations of invasive populations of RPM ............................. 33 2.9.1 Has RPM expanded its host range in the New World and if so, what are the factors behind this expansion? ..................................................................................................... 33 2.9.2 How accurate are the reports relating to the increase in RPM population density between Old and New World regions? If accurate, what factors may be driving the differences? ...................................................................................................................... 34 3.0 The morphological and molecular characterisation and comparison of RPM from India and the Caribbean .................................................................................................................... 36 3.1 Introduction .................................................................................................................... 36 3.2 General approaches ........................................................................................................ 38 3.2.1 Morphological comparison ..................................................................................... 38 3.2.2 Molecular techniques .............................................................................................. 40 3.3 Materials and Methods ................................................................................................... 42 3.3.1 Selection of populations .......................................................................................... 42 3.3.2 Collection of mites .................................................................................................. 42 3.3.3 Morphological measurements ................................................................................. 43 3.3.4 Statistical analysis of morphological measurements .............................................. 43 3.3.5 Molecular analysis .................................................................................................. 46 4 The ecology of Raoiella indica (Hirst) (Acari:Tenuipalpidae) in India and Trinidad 3.3.6 DNA extraction ....................................................................................................... 46 3.3.7 PCR methodology ................................................................................................... 47 3.3.8 Gel electrophoresis.................................................................................................. 47 3.3.9 Sequencing .............................................................................................................. 47 3.3.10 Improved PCR methodology ................................................................................ 48 3.4 Results ............................................................................................................................ 49 3.5 Discussion ...................................................................................................................... 72 4.0 Host plant relations in the Old World ................................................................................ 77 4.1 Introduction .................................................................................................................... 77 4.2 Methods.......................................................................................................................... 81 4.2.1 Hypothesis 1............................................................................................................ 81 4.2.2 Hypothesis 2............................................................................................................ 89 4.2.3 Analyses .................................................................................................................. 90 4.3 Results ............................................................................................................................ 92 4.3.1 Hypothesis 1............................................................................................................ 92 4.3.2 Hypothesis 2.......................................................................................................... 108 Survival and reproduction of RPM on natal host compared to alternative host ............ 108 4.4 Discussion .................................................................................................................... 115 5.0 A comparison of RPM and related predator complex between the adventive range (Trinidad) and
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