Citrus Mites: Identification, Bionomy and Control

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CITRUS MITES

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CITRUS MITES Identiﬁcation, Bionomy and Control

Vincenzo Vacante Professor of General and Applied Entomology Department OASI, Mediterranean University Reggio Calabria, Italy

CABI is a trading name of CAB International CABI Head Ofﬁce Nosworthy Way Wallingford Oxfordshire OX10 8DE UK

CABI North American Ofﬁce 875 Massachusetts Avenue 7th Floor Cambridge, MA 02139 USA

Tel: +44 (0)1491 832111 Fax: +44 (0)1491 833508 E-mail: [email protected] Website: www.cabi.org

Tel: +1 617 395 4056 Fax: +1 617 354 6875 E-mail: [email protected]

© V. Vacante 2010. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Vacante, Vincenzo. Citrus mites : identiﬁcation, bionomy and control / Vincenzo Vacante. p. cm. Includes bibliographical references and index. ISBN 978-1-84593-498-9 (alk. paper) 1. Citrus—Diseases and pests. 2. Mites. I. Title SB608.C5V33 2010 634′.30496542–dc22 2009022588 ISBN-13: 978 1 84593 498 9 Typeset by AMA DataSet, Preston, UK. Printed and bound in the UK by the MPG Books Group. The paper used for the text pages in this book is FSC certiﬁed. The FSC (Forest Stewardship Council) is an international network to promote responsible management of the world’s forests.

In memory of Carlo Vidano, Master of Entomology and Ethics

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Contents

Preface Acknowledgements

x xiv

Part I Introduction 1

Citriculture and Injurious Mites 1.1 Introduction 1.2 Citriculture 1.2.1 Species and varieties cultivated 1.2.2 World production 1.2.3 Fruit exports 1.2.4 Juice production 1.3 Citrus Mites and Their Economic Importance

2

Introduction to Acari 2.1 Introduction 2.2 Morphology and Structure 2.2.1 Body division and external morphology 2.2.2 Gnathosoma 2.2.3 Idiosoma 2.2.4 Legs 2.3 Classiﬁcation 2.3.1 Higher classiﬁcation 2.3.2 Suborder Prostigmata

3 3 3 4 7 7 8 9 11 11 11 11 14 16 16 17 17 19

vii

viii

Contents

3

Methods and Techniques 3.1 Collecting 3.1.1 Collecting from plants 3.1.2 Collecting in substrate 3.2 Preserving 3.3 Preparing 3.3.1 Clearing/maceration 3.3.2 Temporary mounts 3.3.3 Permanent mounts 3.4 Rearing

20 20 20 21 21 22 22 22 23 24

4

Plant Damage 4.1 Feeding Mechanisms 4.2 Feeding Symptoms 4.3 Plant Damage 4.3.1 Local damage 4.3.2 General alterations

25 25 27 28 28 30

5

Control 5.1 Chemical Control 5.2 Side Effects of Chemicals 5.3 Biological Control 5.4 Integrated Pest Management

31 33 33 35 36

Part II Citrus Mites 6

Key to the Identiﬁcation of Families, Subfamilies, Tribes, Genera and Species

41

7

Phytoptidae Murray 7.1 Introduction 7.2 Morphological Characteristics and Systematic Outline 7.3 Phytoptinae Murray

55 55 55 56

8

Eriophyidae Nalepa 8.1 Introduction 8.2 Morphological Characteristics and Systematic Outline 8.3 Eriophyinae Nalepa 8.3.1 Aceriini Amrine et Stasny 8.4 Cecidophyinae Keifer 8.4.1 Colomerini Newkirk et Keifer 8.5 Nothopodinae Keifer 8.5.1 Nothopodini Keifer 8.6 Phyllocoptinae Nalepa 8.6.1 Anthocoptini Amrine et Stasny 8.6.2 Calacarini Amrine et Stasny 8.6.3 Phyllocoptini Nalepa

58 58 58 61 61 67 67 68 68 70 70 84 87

Contents

ix

9

Diptilomiopidae Keifer 9.1 Introduction 9.2 Morphological Characteristics and Systematic Outline 9.3 Diptilomiopinae Keifer

101 101 101 102

10

Tarsonemidae Canestrini et Fanzago 10.1 Introduction 10.2 Morphological Characteristics and Systematic Outline 10.3 Pseudotarsonemoidinae Lindquist 10.3.1 Pseudotarsonemoidini Lindquist

104 104 104 107 107

11

Tenuipalpidae Berlese 11.1 Introduction 11.2 Morphological Characteristics and Systematic Outline 11.3 Brevipalpinae Mitrofanov 11.4 Tenuipalpinae Mitrofanov

113 113 113 117 155

12

Tuckerellidae Baker et Pritchard 12.1 Introduction 12.2 Morphological Characteristics and Systematic Outline 12.3 Tuckerella Womersley

163 163 163 166

13

Tetranychidae Donnadieu 13.1 Introduction 13.2 Morphological Characteristics and Systematic Outline 13.3 Bryobiinae Berlese 13.3.1 Bryobiini Reck 13.3.2 Hystrichonychini Pritchard et Baker 13.3.3 Petrobiini Reck 13.4 Tetranychinae Berlese 13.4.1 Tenuipalpoidini Pritchard et Baker 13.4.2 Eurytetranychini Reck 13.4.3 Tetranychini Reck

172 172 172 177 177 183 188 194 194 195 219

14

Conclusions 14.1 Systematics 14.2 Bio-ecology 14.3 Pest Status 14.4 Natural Enemies 14.5 Means of Control 14.6 Horticultural Practices 14.7 Prevention 14.8 Integrated Pest Management

298 298 299 299 300 301 302 302 303

References

305

Index

371

Preface

An exhaustive treatment of injurious mites associated with economic plants gives rise to the necessity, according to the large number of species involved and their particular bionomics, of an organic presentation divided into systematic categories and individual crops. This would facilitate the presentation of the subject and avoids the consolidated and reprehensible practice, which tends to simplify, either for convenience or opportunism, the problem of mites injurious to citrus in regard to the small number of traditionally known species, legitimizing inappropriate phytoiatric choices on the basis of decisions based on macroscopic examination and/or the colour of an organism rather than on more scientiﬁc parameters (morphological, biological, etc.), to the economic, toxicological and ecological detriment of the entire ﬁeld and societies that directly or indirectly enter into a physical relationship with farming and/or its products. The present work is the result of this reﬂection and aims to assess the problem of injurious mites associated with citrus in the world. The work basically consists of a bibliographical research of the phytophagous mites recorded on citrus throughout the world, integrated with the author’s knowledge. Various researchers (Quayle, 1938; Bodenheimer, 1951; Ebeling, 1959; Chapot and Delucchi, 1964; Talhouk, 1975; Jeppson, 1978, 1989; Smith and Peña, 2002) have brieﬂy dealt with the subject on both a regional and global scale. The list of species associated with citrus in southern California published by McGregor (1956) is well known, together with those of Muma (1975) for Florida, Vacante et al. (1989) for the Mediterranean area and Dhooria et al. (2005) for India. A more organic work was published by Jeppson et al. (1975), who outlined a global picture of the problem of mites injurious to different economic plants and illustrated with a wealth of details the case of citrus. Recently, Gerson (2003) presented a list of species known for citrus throughout the world. However, an updated work in the ﬁeld is required that is able suitably to guide personnel (researchers, technicians and managers x

Preface

xi

of phytopathological departments) interested in the problem of citrus mites. In total 104 species ascribed to the Phytoptidae Murray, Eriophyidae Nalepa, Diptilomiopidae Keifer, Tarsonemidae Kramer, Tenuipalpidae Berlese, Tuckerellidae Baker et Pritchard and Tetranychidae Donnadieu families were treated. All species quoted are formally phytophagous, although a number of them pose no problem to citrus groves (Bryobiinae, some Tuckerellidae and Tetranychidae and Eriophyidae) and were included in the list of injurious mites for completeness and out of respect for the bibliographical information. However, treatment of these latter species allows for the discrimination of mites that occasionally infest citrus from those that are potentially or realistically harmful, assisting the work of phytosanitary services. The book presents the fundamental elements of the external morphology of the mites with the aim of providing a tool that helps in their identiﬁcation, of the high systematics of the species dealt with and the means and methods of collecting mites from plants and substrate, together with their preservation and preparation for study. The structure of the mouthparts of mites is brieﬂy dealt with to introduce the fundamental aspects of symptomatology and damage. A key is presented for the identiﬁcation of the basic systematic categories (families, subfamilies, tribes, genera) and species, brief information on the morphology and systematics of each category, the main elements of morphology and bionomics (geographical distribution, bio-ecology, natural enemies, symptomatology and damage) of the different species. The morphological description of each species is mostly correlated by original black and white drawings produced at the time by the various authors and, where possible, information is given about web sites (Table 4.1) where the natural colours and features of leaving injurious mites and their damages on citrus can be observed. Under no circumstances did the author discuss the merits of decisions of a taxonomical nature. Morphological descriptions of the various species were dealt with by the contributions of specialists; generally they are reported exactly and in some cases were simpliﬁed and adapted to the needs of the text. The information on natural enemies mainly regards the species collected on citrus, or introduced into this crop from other regions and/or investigated for the control of injurious mites. Among these, Phytoseiidae mites represent the greatest number of species, according to the catalogue of de Moraes et al. (2004). This choice does not involve any position of the author on their taxonomic status but only the attempt towards a given minimum order to the discussion, resulting in the ideas of different specialists on the systematics of Phytoseiidae mites being an open problem. Symptomatology and damage are reported only in the case of sufﬁcient certainty, supported by appropriate bibliographic references. In any case, for each family, several tables are presented summarizing the pest status and geographical distribution of each species. For each species a paragraph on control examines the chemical and biological means and the control strategies (chemical, biological and integrated pest management) available in the different regions of the world.

xii

Preface

The information given is taken from the ofﬁcial bibliography and where possible has been integrated with the author’s experience. In this respect, the information on acaricides refers to their chemical groups in order to avoid giving information exceeded by the rapid evolution of the market and/or licence to use, commonly distinct by the withdrawal of various substances from the market and the entry of other substances. The suggested data do not have a general relevance, since the use of various acaricides is subject to the laws of individual countries. It is therefore the responsibility of the reader to verify their applicability in different regions. The complexity of the subject and the considerable physical dimensions of the citrus crop, spread throughout the subtropical and tropical regions of the world, do not facilitate its treatment and make reference to ecological, horticultural and socio-economic aspects, which often differ from one another. In this context, environmental factors are very important and may directly inﬂuence the bio-ecology of the different species and require strategic choices, which may vary from one region to the next for the same species, as in the citrus rust mite, Phyllocoptruta oleivora (Ashmead), which is the most important phytophagous mite pest in the warm and humid areas of Florida, together with the Texas citrus mite, Eutetranychus banksi (McGregor), while the citrus bud mite, Aceria sheldoni (Ewing), is not an economic problem; on the other hand, in the warm and arid areas of California, the citrus red mite, Panonychus citri (McGregor), is the most feared followed by A. sheldoni and Ph. oleivora and is an economic problem only in coastal areas (Childers et al., 1996). Similar situations have occurred in Mediterranean areas and in other humid regions of the world for the pink citrus rust mite, Aculops pelekassi (Keifer). Of equal importance are the productive choices, which vary between regions and continents. Indeed, the control of pests of products destined for fresh consumption generally requires more interventions than that designated for processing. With the former, cosmetic appearance is a priority, while with the latter, the development and abscission of the fruit are unaffected until 50–75% of the surface of the fruit is damaged (Allen and Stamper, 1979). As a result, production in Florida destined for the fresh market is treated with plant protection products three to four times a year, compared with up to two treatments for that designated for processing, with one petroleum oil and one acaricide treatment (McCoy, 1985; Browning, 1992). From this point of view, Mediterranean citrus crops, fundamentally directed at satisfying the needs of a market orientated towards fresh consumption, is considerably different from that of the USA and is therefore different in terms of control strategies. The FAO (FAOSTAT, 2008) estimated for citrus a disparity in yield values (hectogram/ha) between different regions of the world, linked to various environmental and socio-economic causes, including variable availability of technical means and appropriate knowledge and/or specializations in the various ﬁelds of agricultural production. From this point of view, the knowledge of biotic adversities of crops and fruits helps to optimize quality and improve yield. In India, more than 30% of citrus production is lost every year

Preface

xiii

as a result of damage by insect and mite pests (Pruthi and Mani, 1945; Butani, 1979a, b). The above-mentioned facts may inﬂuence the agrarian economy of a region and do not allow for generalizations. Indeed, control of Ph. oleivora in Florida requires an annual expenditure of between 75 and 100 million dollars (McCoy, 1996b), while it poses no problem in other important citrus areas such as Italy where it is not present. Equally important is the case of the reddish black ﬂat mite, Brevipalpus phoenicis (Geijskes), responsible for the transmission in the American continent of ‘Lepra esplosiva’ or ‘Leprosis’ (Childers et al., 2001, 2003b), a feared viral disease (CiLV), prevention of which costs approximately 100 million dollars per year in Brazil alone (Rodrigues et al., 2003) but which is non-existent and does not spread from the mite in the Mediterranean region. In lemon groves in Italy and Spain, up to three acaricide treatments are carried out annually with a cost per hectare of approximately 450 per treatment. The control may have toxicological and ecological consequences, as in the case of the use of dithiocarbamates, aldicarb or other substances, whose social cost is difﬁcult to quantify. In conclusion, it is likely that in some cases the abnormality of the adaptations of one or more species may have escaped our attention, in which case we offer our apologies and willingness to compare with other authors involved in the subject who would like to contribute with their suggestions to improve this work. Vincenzo Vacante Reggio Calabria, May 2009

Acknowledgements

I am particularly grateful to Professor Uri Gerson of the Hebrew University of Jerusalem, Rehovot (Israel), for his critical review of the text and kind suggestions. In addition, I would like to thank Professor Enrico De Lillo of the University of Bari (Italy), Professor Stefano Colazza of the University of Palermo (Italy), Professor Tetsuo Gotoh of Ibaraki University Ami, Ibaraki (Japan), Professor Michel Bertrand, Professor Serge Kreiter and Professor Alain Migeon of the University of Montpellier and of INRA (France), Professor Carlos H.W. Flechtmann, of the University of São Paulo (Brazil), Professor Carlo Duso of the University of Padova (Italy), Professor Eduard A. Uckermann of ARC-Plant Protection Research Institute (South Africa), Professor Kazuhiro Tanaka, Editor-in-Chief of Acta Arachnologica (Japan), Doctor Kim Hogeland, Permissions Administrator, University of California Press (USA) and Evert E. Lindquist of Agriculture and Agri-Food, Canada, who have helped me by sending bibliographic material or for giving permission to use several drawings. I would also like to thank Professor Franco Zagari and Piero Donin of the OASI Department of the Mediterranean University of Reggio Calabria (Italy), and colleagues in the same department, who have been helpful ﬁnancially toward the editorial initiative. Lastly, I especially thank Miss Sarah Hulbert of CABI Publishing for her availability and for the invaluable suggestions that led to the publication of this book.

xiv

I

Introduction

●

● ●

● ●

Overview of the world’s citriculture, species and varieties cultivated, and the importance of injurious mites. Introduction to morphology and classiﬁcation of mites. Methods and techniques for collecting, preserving, preparing and rearing mites. Feeding mechanisms, symptoms and plant damage. Biological, chemical and integrated control.

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1

Citriculture and Injurious Mites

1.1 INTRODUCTION A brief overview of the fundamental aspects (harvested areas, cultivated species and varieties, yield, production quantity, economic importance, etc.) of global citriculture helps to illustrate its economic importance and yield data facilitating the presentation of limiting factors, including biotic adversity such as pathogens, insects and injurious mites.

1.2 CITRICULTURE According to Webber (1967), ‘the various species of the genus Citrus are all believed to be native to the subtropical and tropical regions of Asia and the Malay Archipelago, and to have spread from there to other sections of the world. They have been cultivated from remote ages, and prototype forms of the most important species are not deﬁnitely known’. The harvested area of citrus extends from 40° parallel north to 40° south (Chapot, 1975). It has been calculated that about 140 countries grow citrus and the FAO estimated a total harvested area of 8,322,605 ha for 2007 (FAOSTAT, 2008). In the American continent, the main producing countries are Brazil (915,056 ha), Mexico (524,000 ha), USA (376,050) and Argentina (148,500 ha), where orange, lemon, lime, grapefruit and several mandarins are cultivated. In Asia, China possesses the largest harvested area in the world (2,008,700 ha) with different varieties of citrus and particularly mandarins and pomelos, followed by India (690,100 ha), Iran (243,500 ha), Pakistan (192,700 ha), Thailand (97,600 ha), Iraq (68,750 ha) and Japan (64,730 ha), where orange, grapefruit, mandarin, lemon, lime and other citrus species are produced. © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

3

4

Chapter 1

In Africa, Nigeria has the largest citrus-growing area (732,000 ha), followed by Egypt (137,370 ha), Morocco (79,300 ha) and South Africa (69,980 ha), where orange, lemon, mandarin, grapefruit and lime are cultivated. In the Mediterranean region, Spain is the leading producer (313,850 ha), followed by Italy (173,009 ha), Turkey (94,600 ha), Greece (57,250 ha) and Israel (18,965 ha). In this area, the most common citruses are orange, lemon, grapefruit and mandarin; other citrus fruits, such as lime, lemon, bergamot and bitter orange, have a localized distribution.

1.2.1 Species and Varieties Cultivated The orange (Citrus sinensis (Linnaeus) Osbeck) is the most widespread species in the world, cultivated from the equator to the colder environments of the distribution area of citrus. The FAO estimated that in 2007, worldwide orange cultivation covered a total of 3,905,780 ha variously distributed in different continents (FAOSTAT, 2008). In the same year, it was calculated that the total worldwide harvested area of lemon (Citrus limon (Linnaeus) Burman) and lime (Citrus aurantifolia (Christmann) Swingle) was 911,726 ha; the harvested area of lemon involves mainly the Mediterranean area, California and Argentina. Limes are common in tropical areas, where they effectively replace lemon and are concentrated in a number of countries such as Mexico (140,000 ha in total). The harvested area of grapefruit (Citrus paradisi Macfadyen) and pomelo (Citrus grandis (Linnaeus) Osbeck) amounted to 289,248 ha and is widespread in China, the USA, Cuba, Mexico, South Africa and other countries. The pomelo is present throughout tropical and subtropical Asia. Other citrus fruits have a localized distribution, like the Italian bergamot (Citrus bergamia Risso) and citron (Citrus medica Linnaeus) in Italy and Puerto Rico, or bitter orange (Citrus aurantium Linnaeus) in Andalusia (Spain). The harvested area of tangerines, mandarins (Citrus nobilis Lour.) and clementines (Citrus reticulata Blanco) covers approximately 2,146,597 ha. 1.2.1.1 Orange The FAO has estimated that in the American continent, Brazil possesses the largest harvested area of orange (799,356 ha) (FAOSTAT, 2008). In this country, the most common varieties are Pera, Valencia and Natal, full of juice and principally used for the production of juice. Some cultivars of Navel (Bahia, Bahianinha) are destined for the fresh market. In addition, they are cultivated fruit varieties with low acidity (Laranja lima, Piralima, Lima tardia). Mexico occupies fourth place with 325,000 ha (mostly cultivar Valencia) and production is largely for fresh consumption and juices. The USA occupies ﬁfth place with a cultivated area of about 270,000 ha, most of which is used by the processing industry for juice production. Florida accounts for 82% of production, with a prevalence of the Valencia cultivar. In California, different cultivars of Navel and Valencia are used for fresh consumption. Argentina, where there is a tendency to replace the old varieties (Hamlin, Calderon, etc.)

Citriculture and Injurious Mites

5

with new cultivars of the Navel and Valencia group, occupies second position with a total harvested area of 59,000 ha. As regards the Asian continent, India (cultivar Mosambi, etc.) has an orange-producing area of 440,000 ha and represents the second largest producing country in the world, followed by China with 385,500 ha in total, where the best oranges are grown in the central areas and in modern crops oranges of Navel type are grown with imported cultivars (Robertson, etc.). Iran is the sixth largest producer with 150,000 ha, Pakistan is eighth with 135,000 ha and Indonesia 11th with 72,400 ha. Among the African countries of the Mediterranean Basin, Egypt has an orange-growing area of 85,000 ha and constitutes the tenth largest producer in the world, followed by Morocco with 50,000 ha and South Africa with 40,000 ha, which has abandoned the production of old varieties of orange (Tomango, Hamlin, Premier) and where production is now oriented towards cultivars from international markets (Navel, etc.). In the Mediterranean region, Spain has a growing area of 140,000 ha and is the sixth largest producer of oranges, largely Navel (cultivars Navelina, New Hall, Washington Navel, Navelate, Lanelate), whereas Italy has 105,334 ha, with production fundamentally characterized by a predominance of red pulp varieties such as the Tarocco (Tarocco nucellare, Scirè, Gallo, Tapi, Messina, Meli, Ippolito), the Moro and Sanguinelli. Other Navel orange types occupy niche positions. A secondary interest has other cultivar blondes without Navel, such as the Oval Calabrese and Belladonna. The Valencia orange is more important. Greece and Turkey both have growing areas of 40,000 ha, Tunisia has 12,500 ha, Lebanon 9700 ha, Portugal 7400 ha and Israel (cultivar Shamuti, etc.) 5540 ha (Damigella and Tribulato, 1980; Reforgiato Recupero and Russo, 2009).

1.2.1.2 Lemon and lime According to FAO data, in 2007, the global harvested area of lemon and lime was approximately 911,726 ha (FAOSTAT, 2008). In the USA, the cultivation of lemon (cultivars Lisbon, Eureka, Rosemberg, Ross) covers 25,000 ha, mostly located in California. In Argentina 45,000 ha of lemons (cultivars Genoa, Villafranca) are cultivated, marketed as fresh products or processed, and placed in the global market, including Europe. In the Mediterranean region, the main producers of lemons are Spain (cultivars Fino, Verna, Real) with 46,500 ha, Italy (cultivars Femminello, Monachello, Interdonato, Lunario, Lemox) with 30,046 ha and Turkey (cultivars Kütdiken, Interdonato, Italyan Memeli, Lamas, Molla Mehmet, Kibris) with 20,000 ha. Greece (cultivars Magalene, Karystos, Adamopoulos) has 10,000 ha, Israel 1735 ha and Portugal (cultivar Gallego) approximately 1000 ha (Calabrese and Crescimanno, 1980; Calabrese and Barone, 2009b). As regards the limes, although FAO data are combined with lemon, it is possible to draw conclusions on the main producing countries. In the American continent, Mexico is the major lime-producing country (cultivar West Indian

6

Chapter 1

or Mexican or Key lime), with a cultivated area of 140,000 ha, followed by Brazil with 50,440 ha, partly cultivated with lemon (cultivar Femminello and other), Peru with 21,000 ha and the USA in South Florida (cultivars Persiana, Key, Tahiti, Bearss). In Asia, India (cultivars West Indian or Mexican or Key lime, Mitha Nimbu, Indiana or Palestine) is the largest producing country in the world with 230,000 ha, followed by China with 63,450 ha, partly cultivated with lemon, Iran with 41,000 ha and Thailand with 26,700 ha. In the Mediterranean region, in Egypt (cultivars Key, Indiana or Palestine) about 15,000 ha of limes are grown. In Morocco 1100 ha of a lime acid (cultivar Limûn Boussera) are cultivated (Continella, 1980; Calabrese and Barone, 2009a).

1.2.1.3 Grapefruit and pomelo FAO estimates indicate that in 2007 the worldwide harvested area of grapefruit and pomelo was equal to 289,248 ha (FAOSTAT, 2008). In the American continent, the major producers of grapefruit are the USA (62,000 ha), Mexico (16,000 ha), Argentina (12,500 ha) and Cuba (17,000 ha). Florida produces about 78% of US grapefruit and a large share of production is destined for the fresh domestic market and exports, especially pink-pigmented fruits (Red Ruby, Burgundy, Star Ruby, Ray Ruby). In Mexico, the favourable climatic conditions give the pigmented varieties a ﬁne organolectic quality and a satisfactory coloration that promote their export to Europe and Argentina. In South Africa, 14,000 ha of grapefruit are cultivated, widely bred and destined for export. In Asia, the main producing countries of grapefruit and pomelo are China (62,500 ha), the largest producer in the world of clear or pink ﬂesh pomelo (cultivars Mato, Banpeyu, Red Shaddock, Webber, Dirado Buntan), Thailand (12,000 ha) (cultivars Kao Panne, Kao Phuang, Thong Dee), Syria (10,000 ha), India (8,100 ha) and Bangladesh (6000 ha). In the Mediterranean region, the most important country for grapefruit production is Israel (5370 ha), where a gradual replacement has started of the Marsh variety, characterized by its clear ﬂesh, with the Star Ruby, marked by its more or less intense pink pigmentation (Continella, 1980; Calabrese, 2009).

1.2.1.4 Mandarin and mandarin-like In 2007, the FAO estimated a total mandarin and mandarin-like cultivated area of 2,146,597 ha (FAOSTAT, 2008), characterized by different cultivars throughout the world, such as clementines in the Mediterranean, satsumas in Japan, hybrids in California, Ponkan in China and other countries of South-east Asia. In the American continent, the main producing countries are Brazil (61,000 ha), Argentina (32,000 ha), Mexico (30,000 ha) and the USA (18,000 ha), where 72% of tangerines (mandarins and mandarin-like) are produced in Florida. In California, excellent tangerines and tangelos are produced in the southern areas. In Brazil, the second most important species of citrus is the mandarin (C. reticulata), whose most common cultivar is the Cravo (Laranja Cravo).

Citriculture and Injurious Mites

7

In Asia, China (1,411,000 ha) is the largest producer in the world, and modern satsumas as well as Mediterranean mandarins and tangerines of the Dancy variety originate from Whenzhou. The Ponkan is common throughout South-east Asia. In Japan, 51,000 ha are cultivated, largely satsumas and mandarins, in Pakistan 50,000 ha, in Iran 45,000 ha and 37,200 ha in Thailand. In Africa, the most important countries are Egypt (37,000 ha), Morocco (27,000 ha) and South Africa (5100 ha). In the Mediterranean region, clementines represent about 70% of the production, whereas the cultivation of satsumas is declining and that of lateripening hybrids (variety Fortune) has increased in recent years, especially in Spain. The main producing countries are Spain (125,000 ha), the second largest producer in the world, following by Italy (35,829 ha), Turkey (30,800 ha), Israel (5320 ha) and Portugal (4200 ha). In Spain, there is a prevalence of clementines, which constitute 25% of all citrus production, and major cultivars (Marisol, Fina, Nules, Oroval, Hernandina) are selected. In Portugal, the Encore cultivar, used only in local markets, is well known. In Italy, there has been a strong decrease in the Havana mandarin as well as a crisis with the Tardivo of Ciaculli cultivar (Calabrese and Crescimanno, 1980; Calabrese and Pensabene Bellavia, 2009).

1.2.2 World Production The FAO estimated that in 2007 worldwide production of citrus amounted to 115,650,545 tonnes (t) in total (FAOSTAT, 2008). The ten largest producers are Brazil (20,682,309 t), China (19,617,100 t), the USA (10,017,000 t), India (6,286,000 t), Spain (5,703,600 t), Iran (3,739,000 t), Italy (3,579,782 t), Nigeria (3,325,000 t), Turkey (3,102,414 t) and Indonesia (2,600,000 t). The estimated yield (hectogram/ha) placed in descending order Indonesia (359,116), Turkey (327,950), the USA (266,374), Brazil (226,022), Italy (206,913), Spain (181,730), Iran (153,552), China (97,660), India (91,088) and Nigeria (45,423). The disparity in the values of yield among different regions of the world is linked to various environmental and socio-economic causes, including the variable availability of technical means, and appropriate knowledge and/or specializations in the various ﬁelds of agricultural production. This directly affects the quantitative and qualitative standard and the export opportunities and/or consumption of fresh products. From this point of view, knowledge of biotic adversities of crops and fruits helps to optimize quality and improve yield. In different countries, production is destined in varying degrees for fresh consumption on the local and national markets, and processed into juice also for export.

1.2.3 Fruit Exports Citrus fruits are present in all world markets and are widely present in countries with developed economies. A strong global demand feeds major exports

8

Chapter 1

from producing countries. In 2002, citrus fruit per capita consumption was calculated to be approximately 22 kg per capita per year (UNCTAD from FAO data). The FAO estimate for 2005 showed that the export of citrus fruits in the world was equal to 12,088,535 t (10.45% of total) for a value of US$6,935,692,000. The greatest exporter of fresh citrus in the world is Spain with 3,021,194 t, followed by South Africa with 2,041,225 t, the USA with 936,048 t, Turkey with 894,493 t, Argentina with 644,384 t, Morocco with 559,170 t and China with 450,553 t (FAOSTAT, 2008). The global export of lemon and lime was calculated to be 2,143,935 t, with a value of US$1,220,184,000. Mexico is the largest exporter of limes with 387,196 t for a value of US$26,300,000, whereas Argentina is the largest exporter of lemons with 369,483 t for a value of US$151,830,000, followed by Spain, the third largest exporter of lemons with 362,577 t for a value of US$281,220,000, Turkey with 355,656 t for a value of US$169,395,000, South Africa with 135,014 t for a value of US$57,012,000, the USA with 111,408 t for a value of US$86,878,000 and Italy with 40,584 t for a value of US$31,553,000 (FAOSTAT, 2008). As regards oranges, in 2005 worldwide exports amounted to 5,270,262 t for a value of US$2,589,246,000. The main exporting countries were South Africa with 1,235,027 t for a value of US$272,7764,000, Spain with 1,116,274 t for a value of US$863,876,000, Morocco with 256,160 t with a value of US$115,387,000, Egypt with 214,165 t for a value of US$74,914,000, Greece with 209,821 t for a value of US$100,326,000, Turkey with 193,538 t for a value of US$75,918,000, Argentina with 169,359 t for a value of US$46,503,000 and Italy with 100,742 t for a value of US$65,372,000. The USA and China occupy the ninth and tenth places with 583,471 and 55,867 t, respectively, and with a value equal to US$384,016,000 and US$18,730,000, respectively (FAOSTAT, 2008). In 2005, the export of grapefruits and pomelos worldwide amounted to 1,354,128 t for a value of US$619,590,000. The main exporting countries were South Africa with 585,675 t for a value of US$98,649,000, the USA with 219,385 t for a value of US$154,995,000, Turkey with 98,962 t with a value of US$50,217,000, Israel with 78,068 t with a value of US$47,939,000 and Argentina with 34,103 t with a value of US$12,881,000 (FAOSTAT, 2008). As regards tangerines, mandarins and clementines, total global exports for 2005 were estimated to be 3,320,210 t for a total value of US$2,506,672. The main exporting countries were Spain with 1,512,619 t for a value of US$1,475,896,000, China with 372,131 t for a value of US$10,522,000, Morocco with 302,452 t for a value of US$221,943,000, Turkey with 246,337 t for a value of US$109,312,000 and South Africa with 85,509 t with a value of US$54,621,000 (FAOSTAT, 2008).

1.2.4 Juice Production Citrus fruit processing accounts for approximately one third of total citrus fruit production. More than 80% of it is orange processing, mostly for orange juice production. According to the FAO, in 2005 exports of citrus juice equalled

Citriculture and Injurious Mites

9

5,200,753 t for an export value of US$3,930,898,000. The most important producers are Brazil with 1,778,506 t for a value of US$1,111,305,000, primarily interested in the production of orange juice and exported totally; the USA with 432,771 t for a value of US$372,975,000, largely consisting of orange juice and grapefruit, with 90% consumed in the internal market; and Spain with 240,825 t for a value of US$168,925,000 and consisting largely of orange juice (FAOSTAT, 2008). In the USA, Florida contributes signiﬁcantly to the production of orange juice and is the major producer of grapefruit. Brazil controls the international market for orange juice (1,777,599 t) and Argentina that of lemon (50,703 t), leading both the international price of the product reference in the remaining countries of the world. Chinese production is now equal to about 6108 t, whereas Italian production (74,902 t) is generally characterized by the pigmented red variety, with a demand of juice in several foreign markets. Italian lemons are most sought after for their essences rather than for the juice. Equally important are the essential oils of mandarin, whereas there is less demand for the juices and essences of clementines.

1.3 CITRUS MITES AND THEIR ECONOMIC IMPORTANCE In addition to pathological aspects related to viruses (Tristeza, Leprosis, etc.), fungi (Phoma tracheiphila (Petri) Kantachveli et Gikachvili, Phtophtora spp., etc.) and other injurious organisms, a variety of vertebrate and invertebrate animals commonly attack citruses in the different regions of the world, causing serious damage to the crops and/or the harvest. In this context, arthropods are very important and among them the mites and insects (Quayle, 1938, 1941; Ebeling, 1950, 1959; Bodenheimer, 1951; Reuther et al., 1989; Bedford et al., 1998). Mites and insects possess both appendages articulate, but the ﬁrst are eight-legged and the last six-legged. Insects show a greater degree of ecological ﬁtness, deriving from the richness of the faunistic structure and from numerous evolutionary adaptations. Little is known about mites, at least in some countries, but they are no less important in the ecological and phytopathological context. The problem has been studied in the different citrus regions of the world (McGregor, 1956; Gerson, 1971; Rasmy et al., 1972; Muma, 1975; Jeppson, 1978; Mijuskovic, 1973a, b; Mijuskovic and Tomasevic, 1975; Vacante and Nucifora, 1985; Garcia Marí et al., 1986; Vacante et al., 1989). Nevertheless, with the exception of the contribution of Jeppson et al. (1975) on mites injurious to economic plants, including the citrus, and a recent publication by Gerson (2003), there has not been a contribution that presents organic and up-to-date information on mites injurious to citrus in the world. In marginal productive areas or in less developed areas, this want may signiﬁcantly interfere with pest control and the prevention of the accidental introduction of pests from other countries of the world, with economic, ecological and toxicological disadvantages. The complexity of the subject and the wide physical expanse of citrus crops, spread throughout the subtropical and tropical regions of the world, do not facilitate its treatment and make reference to ecological, horticultural

10

Chapter 1

and socio-economic aspects which often differ from one another. Environmental factors are very important and may directly inﬂuence the bio-ecology of the different species and require strategic choices, which may vary from one region to another for the same species, as in the case of the different behaviour of the citrus rust mite, Ph. oleivora, Texas citrus mite, E. banksi, and citrus bud mite, A. sheldoni in the warm and humid areas of Florida and in the warm and arid areas of California (Childers et al., 1996). In India, several factors contribute towards the decline in the yield of citrus trees, and the threat from different insects and mites has been considered one of the most important factors (Bindra, 1970); the losses related to injurious mites are quite substantial, especially during years when climatic conditions are more favourable for their development (Dhooria et al., 2005). The productive choices, varying among regions and continents, inﬂuence mite control. Production destined for fresh markets generally requires more interventions than that destined for processing juices. In the former, cosmetic appearance represents a priority, whereas the latter permits greater levels of tolerance (Allen and Stamper, 1979). In Florida, production destined for the fresh market is treated with chemicals three to four times per year, whereas that intended for processing is treated only up to twice per year, with one petroleum oil and one acaricide treatment (McCoy, 1985; Browning, 1992). Mediterranean citrus crops, primarily destined for the fresh market, are considerably different from that of the USA. These facts have serious economic importance and do not allow for generalizations, as in the case mentioned in the Preface, of the control of Ph. oleivora in Florida, which requires an annual expenditure of US$75–100 million (McCoy, 1996a, b), whereas it poses no problem in other important citrus areas where it is not present. Similarly, the reddish black ﬂat mite, B. phoenicis, responsible for the transmission in the American continent of ‘Lepra esplosiva’ or ‘Leprosis’ (Childers et al., 2001b), a viral disease (CiLV), necessitates a prevention that costs approximately US$100 million per year in Brazil alone (Rodrigues et al., 2003), but is non-existent and does not spread from the mite in the Mediterranean region. In lemon groves in Italy and Spain, up to three acaricide treatments are carried out per year with a cost per hectare of approximately ?450 per treatment.

2

Introduction to Acari

2.1 INTRODUCTION Acari are small animals, the adults of which range from 300 to 500 μm in body length, except for some eriophyoids that are approximately 100 μm long or certain ticks whose female measures about 30,000 μm. From a systematic point of view, they represent a subclass of Arachnida, subphylum Chelicerata, and are distinguished from insects, with which they are often wrongly associated, for the lack of antennae, mandibles and maxillae and the presence in the adults of four pairs of legs, with the exception of eriophyoids, which possess two pairs of legs. Their biological cycle develops through the stages of egg, larva and nymph. The larva is six-legged and the nymph eight-legged. At present, approximately 48,000 species have been described (Halliday et al., 1999), but it is believed that the group will grow to number over 1,000,000 species (Walter and Proctor, 1999). As regards ecological adaptation, Acari have colonized most of the available aquatic and terrestrial habitats, and possess a capillary ability to exploit the trophic resources available thanks to different dietary regimes (phytophagous, parasitic, predaceous, mycophagous, saprophagous, coprophagous, necrophagous). Some species are phoretic (Krantz, 1978; Lindquist, 1984; Walter and Proctor, 1999; Krantz, 2009b).

2.2 MORPHOLOGY AND STRUCTURE 2.2.1 Body Division and External Morphology Acari have a globular or subglobular body, sometime fusiform or worm-like, pale in colour, rarely lively, with or without inconspicuous abdominal segmentation and are divided into two principal morphological regions, © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

11

12

Chapter 2

ta ti ge

fe Gnathosoma v2

tr

sc1

Prodorsum sc2

IDIOSOMA

c3

c2

Opisthosomal dorsum d2

e2

c1

d1

e1 f2

f1

h1

Fig. 2.1. Tetranychus urticae Koch. Dorsal aspect of the female showing the different regions of the body (from Vacante and Nucifora, 1985, partially modiﬁed); fe, femur; ge, genu; ta, tarsus; ti, tibia; tr, trochanter. The setal notation is explained in the text on p. 176.

respectively called gnathosoma and idiosoma (Fig. 2.1). The limits between these regions of the body are not always well deﬁned and are sometimes identiﬁed by the existence of scarcely discernible sutures (circumcapitular, disjugal, sejugal, abjugal) (Krantz, 1978; Alberti and Coons, 1999). Every group possesses a typical chaetotaxy, with a number of setae, characterized by shape, length, distance between their bases and function (Fig. 2.2).

Introduction to Acari

13

A ag C g1

B g2 ps2 h3

ps1

F E

D

G H

I

Fig. 2.2. (A) Ano-genital regions of female of Tetranychus urticae Koch (explained in the text) (from Vacante and Nucifora, 1985, partially modiﬁed); (B) dorsal tactile seta of Bryobia praetiosa Koch; (C) duplex setae of tarsus IV of B. praetiosa (solenidion at left and tactile seta at right) (from Vacante, 1985, partially modiﬁed); (D) solenidion and famulus of Thyreophagus cooremani Fain; (E) supracoxal seta of Tyrophagus palmarum Oudemans (from Vacante, 1989); (F) trichobothrium of Humerobates rostrolamellatus Grandjean (from Vacante and Nucifora, 1985); (G) distal anasthomosis of peritreme of B. praetiosa; (H) distal anasthomosis of peritreme of Bryobia rubrioculus (Scheuten); (I) peritreme of Panonychus citri (McGregor) (from Vacante, 1985). The setal notation is explained in the text on p.176.

14

Chapter 2

Respiration may be cuticolar or tracheal, through paired stigmata or with peritremes of various shape and structure (Fig. 2.2). The tracheae may be placed dorsolaterally, ventrolaterally, anteriorly to the bases of chelicerae or near the base of the legs. Some characters (number, length, shape, location) of the stigmata are used in systematics. The body presents a number of sensory organs, most of which are setae, primarily tactile or chemotactile (Fig. 2.2). The presence of ocelli and photosensitive organs is well known. The presence of several cuticolar pores is attributable to that of an excretion organ. In addition, coxal glands are involved in the water balance and regulation of ion concentration (Krantz, 1978).

2.2.2 Gnathosoma The gnathosoma or capitulum is the ﬁrst region of the body and derives from embrional cheliceral segment, and the derived biramous appendages of the second somite. From the fusion of the palpcoxae arises the subcapitulum, more or less conical and with a dorsal and longitudinal wrinkle hosting the chelicerae; it also possesses the preoral wrinkle, mouth and pharynx. The palps are located dorsolaterally. The dorsal anterior part of the propodosoma forms the prodorsum (Fig. 2.1).

Proterosoma

Hysterosoma Opisthosoma

Prosoma

Precheliceral I, II

Gnathosoma

III–IV

V–VI

Prop

Met

VII–XVI

Podosoma

Fig. 2.3. Diagram of primitive segmentation in the Acari (according to Coineau, 1974); prop, propodosoma; met, metapodosoma.

Introduction to Acari

15

B C A

md

p

st

D

lab

E

F

fd ig

pa md

md

is as

ss

Fig. 2.4. Gnathosomal structures. (A) Dorsal view of gnathosoma of Tetranychus urticae Koch; (B) dorsal view of gnathosoma and prodorsum (partially designed) of Brevipalpus phoenicis (Geijskes); (C) palp of Bryobia sp. (from Vacante and Nucifora, 1985); (D) gnathosoma with extended chelicerae of Tarsonemella africanus (Hirst); (E) apex of gnathosoma and palp of Asiocortarsonemus malayi Fain (from Lindquist, 1986); (F) diagram of transverse section of cheliceral and associated structures at level near apices of stylets (according to Lindquist, 1996); as, auxiliary stylet; fd, ﬁxed digit; ig, infracapitular guide; is, infracapitular stylet; lab, labrum; md, movable digit; p, peritreme; pa, palp; ss, stylet sheath; st, stylophore.

The chelicerae are considered as derivative endopodal appendages of the ﬁrst somite and are placed dorsally compared with the opening mouth and commonly possess three segments called cheliceral base, digitus ﬁxus and digitus mobilis, respectively. The ﬁrst is basal and bears dorsally the digitus ﬁxus articulated with the distal digitus mobilis. The cheliceral bases may be partially or completely fused to each other and form, as in the case of tetranychids and other similar groups, the stylophore. The chelicerae may have a different morphology, as in the tetranychids that present the digitus mobilis modiﬁed in a stylet ﬁt for piercing the cell wall (Fig. 2.4).

16

Chapter 2

The palps are postoral appendices, consisting of a variable number of articles (trochanter, femur, genu, tibia, tarsus and apotele), including from one or two to six (Fig. 2.4). The two coxae merge with each other and form the subcapitulum, which opens the mouth, bounded dorsally by the labrum and laterally by various structures (lateral labra, rutella, malae). The function of the palps is primarily related to the search for and handling of food, and usually the distal segment’s leading sensorial setae (mechanoreceptors, etc.), which acts as a support for the trophic activity and may provide the outlet of silk glands (Krantz, 1978, 2009a).

2.2.3 Idiosoma The idiosoma is formed by the anterior propodosoma and posterior hysterosoma. The propodosoma is anterior to the dorsal disjugal furrow, anterior to the ventral sejugal furrow, and includes the pairs of legs I and II. The hysterosoma is posterior to the dorsal disjugal furrow to ventral sejugal furrow and includes legs III and IV (Fig. 2.3). The area bearing the legs derives from four embryonal somites and is also called podosoma; the part deriving from the ﬁrst two somites shelters the ﬁrst two pairs of legs and is the propodosoma, while the last part deriving from two other somites presents the other two pairs of legs and is called metapodosoma (Fig. 2.3) (Coineau, 1974; Krantz, 1978; Alberti and Coons, 1999; Krantz, 2009a). The part of the hysterosoma excluding legs III and IV constitutes the opisthosoma and derives, according to the systematic groups, from six to 13 embryonal somites and shelters the genital and anal openings (Fig. 2.3) (Coineau, 1974). Its shape is from ovoid to fusiform and presents a cuticle variously striate, with furrows or covered by plates or shields, of shape and size variable in the different systematic groups. Male and female genital openings open both anteriorly or posteriorly. The sperm transfer may take place directly through the intrusion of the male copulatory organ or aedeagus (Fig. 2.5) in the female genital opening or in special extragenital structures located on the rear end of the idiosoma (Fig. 2.2). It is also known how indirect transfer of the sperm through the chelicerae of the male or the deposition of spermatophores on the substrate are subsequently recovered by the female through structures of the genital apparatus or epigynium. In various species, the female produces sexual pheromones (Krantz, 1978, 2009a).

2.2.4 Legs The legs possess seven segments; coxa, trochanter, femur, genu, tibia, tarsus and pretarsus (Fig. 2.1). The coxae may be free or fused with podosoma venter and the femur may be divided into basifemur and telofemur. The various segments present a precise number of setae, tactile or specialized (solenidia, eupathidia, famuli, microsetae, trichobothria), organized in whirls and set in

Introduction to Acari

17

A

B

C

E D

F

Fig. 2.5. Different shapes of aedeagus. (A) Petrobia tunisiae Manson; (B) Tetranychus urticae Koch; (C) Panonychus citri (McGregor) (from Vacante, 1985). Different shapes of receptaculum seminis; (D) Aplonobia histricina (Berlese); (E) Petrobia latens Müller (from Vacante, 1985); (F) Petrobia harti (Ewing) (from Meyer Smith, 1987).

dorsal, ventral and lateral positions. The distal end of each tarsus may bear the ambulatory appendages, consisting of a pair of lateral claws and a central empodium, with variations in different groups (Fig. 2.6). The pretarsus is also called ambulacrum when it possesses meolian elements (Krantz, 1978, 2009a).

2.3 CLASSIFICATION 2.3.1 Higher Classiﬁcation The latest studies consider the Acari a subclass of the class Arachnida (Krantz, 1978; Johnston, 1982; Lindquist, 1984; Evans, 1992; Walter and Proctor, 1999; Zhang, 2003; Lindquist et al., 2009). Recently, Zhang (2003) has partially modiﬁed the view of Johnston (1982) and Evans (1992) and has divided the subclass into three superorders (Opilioacariformes, Parasitiformes and

18

Chapter 2 C B

A

E D F

Fig. 2.6. Different pretarsi. (A) Pretarsus I of female of Tyrophagus tropicus Robertson (from Vacante, 1989); (B) pretarsus I of female of Bryobia kissophila van Eyndhoven; (C) pretarsus I of female of Panonychus citri (McGregor); (D) pretarsus I of female of Eotetranychus rubiphilus (Reck); (E) pretarsus I of female of Tetranychus urticae Koch (from Vacante, 1985, partially modiﬁed); (F) empodium of Aculops pelekassi (Keifer) (from Keifer, 1959a, partially modiﬁed).

Acariformes) and seven orders. The superorder Opilioacariformes number the order Opilioacaridida, the superorder Parasitiformes, the orders Holothyrida, Mesostigmata and Ixodida and the superorder Acariformes number the orders Prostigmata, Astigmata and Oribatida. Recently, Lindquist et al. (2009) recognized in the subclass only the superorders Parasitiformes and Acariformes. The ﬁrst includes the order Opilioacaridida and the latter the orders Trombidiformes and Sacroptiformes. The species treated in this book belong to the suborder Prostigmata of the order Trombidiformes. From a systematic point of view, the subclass Acari numbers currently 540 families, 124 superfamilies, 5500 genera and 1200 subgenera (Lindquist et al., 2009).

Introduction to Acari

19

2.3.2 Suborder Prostigmata The Prostigmata include about 17,170 described species, belonging to 1348 genera and 131 families (Walter and Proctor, 1999) characterized by different feeding habits (phytophagous, predaceous, mycophagous, etc.). The typical morphological character consists of the stigmata, set on the anterior margin of the prosoma or between the bases of chelicerae. Furthermore, the palp segments may be blended or reduced. In different groups, the palptarsus may displace to the base of the tibia and form the ‘thumb claw complex’ (Fig. 2.4). The digitus ﬁxus of the chelicerae may be missing or regressed and the digitus mobilis may be transformed into a stylet (Fig. 2.4). The cheliceral bases may form a stylophore or be fused to the subcapitulum (Fig. 2.4). The prosoma and opisthosoma often bear plates or dorsal shields (Zhang, 2003; Lindquist et al., 2009). The species reported for citrus belong to the families Phytoptidae, Eriophyidae, Diptilomiopidae, Tarsonemidae, Tenuipalpidae, Tuckerellidae and Tetranychidae.

3

Methods and Techniques

3.1 COLLECTING In citrus crops, the Acari normally live on the plants, in the soil and/or in supports of a different nature and sometimes in all three habitats. The collecting method therefore varies depending on the behaviour of different species and the needs of the operator. Indeed, the needs of the control may be very different from those of a laboratory rearing or a systematic study. Regardless, identiﬁcation is desirable in order to collect the largest number of specimens and preferably in all biological stages, noting their colour, damage symptoms and host plant, besides the date, location of collection and name of the collector. In this contribution, the techniques used for mites injurious to cultivated plants and citrus in particular are treated. However, several reasons may justify the examination of beneﬁcial mites and for this, information from Gerson et al. (2003) is applicable to the pest species.

3.1.1 Collecting from Plants Various techniques allow the collection of immature and adult stages of Acari on plants (Jeppson et al., 1975; Upton, 1991; Amrine and Manson, 1996; Perring et al., 1996; Walter and Krantz, 2009). Many phytophagous or predaceous species living on the canopy can be collected by shaking the foliage with a stick or with hands on to a white or black support (tray, paper, etc.), depending on individual needs, and then using a needle or brush to collect them. Generally, the most common method involves keeping the manually collected plant material in sealed plastic or paper bags, storing it in the refrigerator at 10–15°C and examining it within a short time in the laboratory through a dissecting microscope. This avoids deterioration or death of specimens collected as a result of curing or predation. 20

© V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

Methods and Techniques

21

Working in this way, Acari are kept alive and can be used in the laboratory for biological studies and various kinds of analyses (morphological, biochemical, etc.). The ﬁeld collection of Acari on the various organs of plants (leaves, branches, fruits, portions of bark) can also be carried out using pins mounted on sticks of various sizes, pipettes or ﬁne hair brushes and using a hand lens, preserving them alive for a short time on empty vials or vials containing a preservative liquid such as 70% alcohol or Oudeman’s ﬂuid (see below). The method is not suitable for all systematic groups and is normally used for not very mobile species and those of middle or large size and still sufﬁciently well known. For highly mobile and medium- or large-sized Acari, an aspirator can be used, consisting of a glass vial, or one made from another transparent material, with a rubber bung and two tubes, one of which is used as a sucking tube, served behind a ﬁlter to avoid the Acari being sucked up by the operator, and the other with a thin tip used for collecting the specimens from substrates. The eriophyoids vagrant on leaves and fruits can be collected by pins and brushes or by pouring a syrup of sorbitol mixed with a 25% solution of isopropyl alcohol with a few iodine crystals on to the leaves and fruits, taking care to collect everything in a small container and observe the material through a dissecting microscope. It is possible to wash the leaves and other parts of plants in hot water and separate the mites in the water by pouring the water through a sieve.

3.1.2 Collecting in Substrate Mites on the soil surface can be collected with a hand-operated vacuum apparatus. The specimens collected on the ﬁlter can be observed directly with a hand lens or with a dissecting microscope, or you can pour the contents on a coloured mat, after washing with water or not. A Berlese– Tullgren funnel can be used to collect mites from the ground vegetation and to extract living mites from soil and litter. A mass collection of mites may be conducted by placing attacked vegetation into a Berlese funnel. It can also be done by the canopy fogging method, involving the fumigation of vegetation or of part of the canopy with insecticides applied with fog (Krantz, 1978; McSorely and Walter, 1991; Upton, 1991; Walter and Krantz, 2009).

3.2 PRESERVING The mites are usually kept in small vials or glass containers containing 70–80% alcohol with the addition of 5% glycerol to prevent alcohol evaporation (Evans et al., 1961). Oudeman’s ﬂuid (a mixture of 87 parts of 70% alcohol, ﬁve parts of glycerol and eight parts of glacial acetic acid) can be used as an alternative, which has the advantage in cases of long preservation

22

Chapter 3

of clarifying (in varying degrees according to the species) preserved specimens. Before conserving, it is necessary to immerse the mites in hot water, forcing them to extend their legs and other appendages, which facilitates their subsequent morphological examination; it is possible to ﬁx the mites in a solution of methanol and acetic acid in the quantity of two parts each and one part of distilled water, transferring them within a week to the preservative medium (Evans, 1992; Saito and Osakabe, 1992).

3.3 PREPARING 3.3.1 Clearing/Maceration The microscopic examination of the mites by optical phase contrast and interference system demands the specimens be cleared and mounted on slides. It is possible to use different media (Krantz, 1978; Walter and Krantz, 2009), including the very common lactophenol, produced by adding in sequence 50 parts lactid acid, 25 parts phenol crystal and 25 parts distilled water. To mitigate the aggressiveness of lactophenol on the most sensitive mites, a water solution of 50–95% lactic acid is used. In one way or another, depending on the size and nature of the species, it is possible to clear a specimen in a week at room temperature. Heating the mites in the clearing medium on a hot plate produces a rapid maceration within a period ranging from a few minutes to approximately 1 h. Cleared specimens must be washed with distilled water before mounting on slides.

3.3.2 Temporary Mounts The mounting slide may be temporary or permanent and it is possible to consult various specialist contributions for this (Singer, 1967; Krantz, 1978; Gutierrez, 1985a; Evans, 1992; Amrine and Manson, 1996; Walter and Krantz, 2009). Lactic acid can be used for temporary mounts, applying a drop at the centre of a glass slide and placing the mite at the centre with the help of an appropriately sized needle or spatula. It is possible to intervene with the same tools better to guide the specimen, paying attention to the position of the legs and avoiding them settling under the body. Subsequently, it is possible to dispose above the mite a coverslip, of appropriate shape and size. Again we must act carefully, ensuring the mite is at a suitable position; the outcome depends by the experience of the operator, the size of specimens and the slip covering the specimen. Some specialists use a concave glass slide, covering partially with the coverslip in order to guide the specimen in the study. This method is commonly applied in the study of particular groups (e.g. Oribatida).

Methods and Techniques

23

3.3.3 Permanent Mounts Permanent mounts require the use of a slide, some drop of medium and a coverslip of different size and shape, depending on the various species and groups of mites. Common media for permanent mounts are lactophenol media, resinbased media and Hoyer’s medium (Krantz, 1978; Saito et al., 1993; Upton, 1993; Amrine and Manson, 1996; Walter and Krantz, 2009). Among the lactophenol media, Heinze’s PVA is commonly employed, used as Hoyer’s medium and produced by mixing the following substances: ● ● ● ● ● ●

Polyvinyl alcohol Distilled water Lactic acid (85–92%) Phenol 1% aqueous solution Glycerol Chloral hydrate

10 g 40–60 g 35 ml 25 ml 10 ml 100 g

Resin-based media include Canada Balsam and Euparal (and the opinion of acarologists on their use is divided), and these possess modest optical properties, enabling them completely to macerate specimens and dehydrate those already treated. Hoyer’s medium is widely used and basically derives from Berlese ﬂuid, based on the use of arabic gum and chloral hydrate, and can be used for clear, weakly sclerotized specimens; it also has good optical properties, even if it is not always considered a permanent medium. Since this is a toxic mixture, its use raises the need to take precautions, e.g. avoiding contact with skin and eyes. Moreover, it quickly degrades those specimens clariﬁed with lactic acid and not sufﬁciently washed with distilled water. It is produced by mixing in sequence the following substances: ● ● ● ●

Distilled water Arabic gum Chloral hydrate Glycerine

25 ml 15 g 100 g 10 ml

We must use arabic gum from the crystalline source, and not from powdered form, ﬁltering the mixture several times. The methodology of mounting the specimen in special positions was investigated in a number of cases, such as in tetranychid males, whose study requires that they be mounted sideways to see the male aedeagus on a lateral view. It is advisable to spread a small drop of Hoyer’s medium on the slide, placing the mite in the centre and laterally, and drying in a drying oven at 40°C for 3 h, or for a longer period at room temperature. Then, place a new drop of Hoyer’s medium on the specimen and on the layer of Hoyer’s medium that was semi-dry and the two layers should combine (Henderson, 2001).

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The permanent mounts require a drying process and hooping of the slides to identify the location of the mite in the slide. In particular, the use of water-soluble media demands that the specimen be fully dried in a hot oven at 40–50°C for 1–2 weeks. If the conservation environment of the dried slides does not possess a low humidity, the slides must be sealed around the coverslip with a sealant. For this purpose, insulating paint is used (Glyceel, Euparal, Glyptal), ringing the coverslip with a small paintbrush and repeating the coatings several times (Travis, 1968; Tribe, 1972; Fain, 1980).

3.4 REARING Summarizing the information on the rearing methods of mites is not simple, since varying techniques according to the systematic groups and aims of study are employed. A comprehensive review appears in Krantz (1978) and, according to systematic groups, different contents can be used. For eriophyoids refer to Oldﬁeld and Perring (1996), for tarsonemids to Liang (1980) and Xu et al. (1994) and for tetranichids to Helle and Overmeer (1985a).

4

Plant Damage

Many morphological, physiological and behavioural adaptations for feeding on plants favour the Trombidiform mites. Thus, over 4000 species of Trombidiformes belonging to about 300 genera are recorded for their obligate phytophagy (Amrine and Stasny, 1994; Bolland et al., 1998), including some important pests of economic plants. Mite pests can cause different degrees of serious damage to citrus trees and fruits according to various environmental factors. Thus, whereas LeClerg (1965) recorded a 2.5% loss in citrus fruit production related to damage by pest mites alone in the USA, more than 30% of production is damaged every year in India by insect and mite pests (Pruthi and Mani, 1945; Butani, 1979a, b). Knowledge of feeding mechanisms and feeding symptoms helps in understanding the nature and type of damage, to recognize the pest responsible and place the control in a context of rationality. Feeding symptoms are correlated to morphological and functional adaptations and the bio-ecology of the injurious species, aside from the attacked organ, the host plant and its phenology. The feeding activity of mites causes local damage and general alterations. In order to aid understanding of the symptoms and identiﬁcation of various species, a number of websites have been listed, which contain photographs of the most important species and their damage on citrus (Table 4.1).

4.1 FEEDING MECHANISMS In the Tetranychoidea (Tetranychidae, Tenuipalpidae, Tuckerellidae) the stylophore, the chelicerae and the palps are involved in the feeding mechanisms (Fig. 2.4). The stylophore is mobile (retractable and extrudable) and lies on the dorso-medial surface of the infracapitulum; it consists of the bases of chelicerae consolidated between them, curves ventrally and is divided by a median and vertical septum and penetrated in two basal chelicerae, each of which hosts a stylet, attached at the base of a cheliceral lever that activates it © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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Table 4.1. Some web sites on which it is possible to ﬁnd information on injurious citrus mites, to observe and their natural colours and see citrus damage. http://cekern.ucdavis.edu http://www.extento.hawaii.edu http://gardenresourceguide.com http://www.inra.fr http://kcc-weslaco.tamu.edu http://www.ipm.ucdavis.edu http://www.forestryimages.org http://nassau.ifas.uﬂ.edu http://ipm.ncsu.edu http://www.sel.barc.usda.gov

on a vertical plane. The chelicerae possess a uniformly long and thick digitus mobilis, except at the distal end where it is sharpened and slightly notched and, if protracted and juxtaposed with that of the other side, forms an empty tube. The stylophore pushes the chelicerae into plant tissues and in retraction probably leaves an opening through which the cellular ﬂuids may be pushed out by turgidity pressure exerted by the same cell. Immediately behind the stylets, the palps press the plant surface and the pharyngeal pump located at the distal side sucks the cellular contents. The stylets penetrate to a depth of between 70 and 120 μm and, once inserted into the plant tissue, push and withdraw continuously causing mechanical damage. As soon as they have consumed that part of the tissue, they move and look for a site that has not been attacked. The depth of damage depends on the length of stylets, feeding time and population density (Jeppson et al., 1975; Sances et al., 1979; Mothes and Seitz, 1982; André and Remacle, 1984; Lindquist, 1985). In eriophyoid mites, the cheliceral stylets (Fig. 2.4) are connected to the underlying motivator. They are rigid, separated from each other for its entire length and not forming a food channel; their function consists only of an alternate longitudinal movement. On the sides of cheliceral stylets, two auxiliary stylets of uncertain function are implanted. In addition, a single long stylet called a labrum, and below which is a gutter, is connected to the pharynx and propelled by strong muscles. After selecting a suitable site for nutrition, the mite stops, applies the rostrum to the host surface and contracts the telescopic segments of the palp, which allows the protrusion of cheliceral stylets over a short distance. In a few seconds, the stylets mechanically penetrate the epidermal cells. It seems that the limited time available does not allow the saliva issued to dissolve the cell wall enzymatically, but only to notch it. Also, it is difﬁcult for the mite to maintain its cheliceral stylets within the wound, given that it does not establish a food tube. It is possible that the mite goes backwards and that the action of the terminal segments of palp allows the assimilation of the food. The movement of the food in the gnathosoma has not been deﬁnitively clariﬁed, but it is possible that the adaptation of the labrum in the gap on the cell wall is followed by a funnelling of the food along its ventral gutter to the pharynx. The regular

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27

contractions of the pharyngeal pump allow active ingestion (Lindquist, 1996; Nuzzaci and Alberti, 1996; Westphal and Manson, 1996). In Tarsonemids, the mouth appendices apparently seem unable to penetrate the plant tissues. The palp is simple and consists of two portions adapted to tactile function or, in some cases, driven to eating solid food particles, making certain species capable of swallowing whole fungal spores. The chelicerae are simple, stylet-like and a little stretched (Fig. 2.4). They seem more suited to stinging the cell walls of fungi and of soft tissues rather than ligneous tissues such as leaves and mature shoots. It is believed, however, that the toxins injected with saliva during feeding can alter the normal ontogenesis of tissues. Some species, like the citrus silver mite, Polyphagotarsonemus latus (Banks), are also capable of feeding on hardened leaves, probably through their ability to produce a degree of toxaemia (Jeppson et al., 1975; Lindquist, 1986).

4.2 FEEDING SYMPTOMS In tetranychoid mites, the feeding symptoms derive from the removal of the contents of cells of palisade tissue involving the disappearance of chloroplasts and the clotting of cellular residues, these taking on the appearance of small, amber-coloured masses. In the palisade layer, only the attacked cells are damaged and the adjacent cells are spared, as well as the elements of vascular leaf ribs. The small punctures present a light colour and, with prolonged exposure, assume the appearance of irregular white or greyish spots. The density of the punctures is a function of the size and duration of the attack and of the injurious species. The density of punctures/cm2 of leaf surface can be high as in the case of Eutetranychus banksi, where 873 feeding punctures/cm2 have been counted on the leaves of Valencia orange (Hall and Simms, 2003). Sometimes, the alterations can become consistent and be yellowish (Tetranychus urticae Koch), grey (Eutetranychus orientalis (Klein), Meyernychus emeticae (Meyer), Schizotetranychus baltazari Rimando) or yellow-bronze (Oligonychus coffeae (Nietner)); the attack may involve foliar curlings (Polyphagotarsonemus latus, Aculush pelekassi), necrosis of young leaves (Calacarus citrifolii Keifer, A. pelekassi, M. emeticae, Eotetranychus sexmaculatus (Riley)), stalks and shoots (Citrus citrifolii, A. pelekassi, E. orientalis, E. yumensis (McGregor)) or even leaf burning and defoliation (C. citrifolii, Phyllocoptruta oleivora, A. pelekassi, E. africanus (Tucker), E. orientalis, E. sexmaculatus, E. yumensis, O. coffeae) and/or fruit drop (Ph. oleivora, E. yumensis). The silk production of many tetranychid mites and webbing may be retained as an aspect of the symptomatology of the attacks, if it is well known that it has different functions (Gerson, 1985a). In Eriophyoid and Tarsonemid mites, the feeding symptoms differ from those of tetranychids, consisting of morphological and chromatic alterations of various organs, and varying with the level and duration of the attack. Several Eriophyoids (Cosella ﬂeschneri (Keifer), Circaces citri Boczek, Phyllocoptruta spp., A. pelekassi, Tegolophus spp., Diptilomiopus assamica Keifer) produce russeting on the leaves and fruit, probaby due to the tarsonemid P. latus; different tetranychids (P. citri, E. lewisi McGregor, E. yumensis) and tenuipalpids

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(Brevipalpus californicus (Banks), B. obovatus Donnadieu, B. phoenicis) are responsible for citrus russets. The feeding symptoms are related to the bio-ecology of each species. Thus, while the sex-spotted spider mite, E. sexmaculatus, prefers the coastal wetlands, generally lives on the lower leaf surface and does not attack the fruit, the Yuma spider mite, E. yumensis, adapts to less conﬁned areas and infests leaves, fruit and the young spurs of young lemon plants, or as in the case of the Lewis spider mite, E. lewisi, is unable to live in conditions of aridity and is usually only harmful to lemon fruit in the ripening period. Colonies of the two-spotted spider mite, T. urticae, usually develop on a well-deﬁned portion of the surface of the lower leaves and fruit, while the oriental red mite, E. orientalis, the citrus red mite, P. citri, or the yellow citrus mite, Eotetranychus cendanai Rimando, while attacking both foliar surfaces, prefer the upper leaf; the citrus silver mite, P. latus develops on young lemons until they reach the size of a walnut, while the pink citrus rust mite, A. pelekassi, appears at an even later stage (Jeppson et al., 1975). In general, the population density and feeding interval needed to cause appreciable damage are inﬂuenced by plant vigour, nutrition and water provided by the roots and by transpiration, which is in turn inﬂuenced by general water conditions (Jeppson et al., 1975). A typical symptom of the attack, not connected with any trophic action, is related to citrus grey mite, C. citrifolii, which secretes white wax on the upper side of the leaves.

4.3 PLANT DAMAGE Jeppson et al. (1975) use the term ‘plant disease’ instead of ‘plant damage’ and write that ‘the term plant disease in its broadest meaning includes all injuries or abnormalities generated from sources outside the plant regardless of the cause’. There is therefore a link between mites and diseases sometimes transmitted, and the damage may be local, wider or general – involving the entire plant, due to the injection of systemic or persistent toxins or viruses. The reaction of a plant to the mite attack therefore consists of a range of external symptoms resulting from mechanical damage (colour changes, shape alterations, reductions in growth and losses of ﬂowers and production) and internal alterations of a biochemical nature. Understanding these reactions requires the analysis of mutual interaction between the host plant and pest (Tomczyk and Kropczyn´ska, 1986).

4.3.1 Local Damage Local damage consists of the removal of cellular contents and involves an economic loss if a sufﬁcient amount of plant material is subtracted for a sufﬁciently long time and with a high population of mites (Jeppson et al., 1975). It may affect individual cells attacked and also those adjacent, and occurs at different depths, depending on the length of stylets, feeding time and the

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29

density of the pest population, and involves aspects of a morphological, histological and cytological nature, which vary depending on the harmful species and the host plant. The alterations to load cells not directly affected may depend on the rupture of the cell as a result of the vacuum created during the trophic process, by the deposited saliva of the mite on adjacent cells and by a process of degeneration of unaffected cells activated by damaged tissues (Tanigoshi and Davis, 1978; Sances et al., 1979; Mothes and Seitz, 1982). At a microscopic level, a signiﬁcant amount of cellular contents pass through the intestinal tract of mites, estimated by McEnroe (1963) to be a volume equal to 1.2 × 10−2 ml/mite/h and corresponding to approximately 50% of the mass of an adult female of T. urticae. Leisering (1960) has calculated that the mite stings and empties 100 foliar cells photosynthetically active in 1 min. Mothes and Seitz (1981) reported that after nutrition, only granules of thylakoid can be found in the intestinal tract, derived from grain thylakoid, of fundamental importance in plant cells and made up of 45–50% of proteins, 50–55% of lipids and small amounts of RNA and DNA (Noggle, 1983). Water and other small quantities of substances are excreted directly (McEnroe, 1963). Sometimes it is difﬁcult to separate the inﬂuence of mechanical damage from that of local toxins (Jeppson et al., 1975). The small size of the mouth parts of eriophyoids (7–30 μm) forces them to feed on the epidermic tissues, while the vagrant species attack a single cell and shortly after leave to attack another cell, leading at high density to the destruction of epidermal tissue. In areas where the epidermal cells are destroyed, a layer of callous tissue rich in lignin is formed above the parenchyma, and it is likely that the typical russeting or silvering of eriophyids arises from its oxidation (Jeppson et al., 1975; McCoy and Albrigo, 1975; Royalty and Perring, 1996). In orange fruits infested by Ph. oleivora, the damage also affects cell layers not directly affected by the cheliceral and auxiliary stylets of the eriophyoid, and the attacked cells exhibit lignin, callose or tannin and emit ethylene; in July and August, the fruit exhibits a peridermic wound below the layer of epidermic dead cells (McCoy and Albrigo, 1975) and the infested fruits of Valencia late orange have less juice, more soluble solids and acids and a higher concentration of acetaldehydes and ethanol (McCoy et al., 1976a). On the fruit of Citrus sinensis, the feeding adults of the pink citrus rust mite, A. pelekassi, produce about 20 punctures, each approximately 1 μm in diameter, in the epidermal cell (10 μm × 7μm). The depth of penetration is about 20 μm and reaches as far as the second and third layer of the fruit epidermis (Tagaki, 1981). The damage may consist of changes to tissue development, as in the case of the citrus bud mite, A. sheldoni, which feeds within the buds and may cause the destruction of embryonic tissue producing alterations of sprouts, leaves and fruit, and symptoms similar in appearance to that of attack by virus, systemic toxin or other mites. A number of tetranychid species such as E. sexmaculatus or T. urticae develop on the lower surface of citrus leaves, and their trophic action involves a delay in developmental tissue determining the appearance on the upper side, of chlorotic blisters of a yellowish colour.

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4.3.2 General Alterations General alterations include viral diseases such as ‘Lepra explosive’ or ‘Leprosis’, transmitted by tenuipalpids of the genus Brevipalpus (Childers et al., 2001, 2003b); biochemical abnormalities caused by different agents inoculated with saliva during the trophic process, including plant hormone substances (Storms, 1971); saliva responsible for symptoms of toxicogens (concentric rings blotch), as in the case of the eriophyoid C. citrifolii (Doidge, 1925; Dippenaar, 1958a, b, c); or the citrus bud mite that causes an increase in the host’s level of phenols in the tissues of the buds, with a simultaneous decline in activity of auxine, and alterations in these and of the RNase (Ishaaya and Sternlicht, 1969, 1971). Cytological and chemical changes may be accompanied in the infested plants by changes in physiological processes such as photosynthesis and transpiration. A detailed overview of aspects related to eriphyoid damage was published by McCoy (1996a). In Ph. oleivora the result of the attack on the lower surfaces of the leaves (abaxial) is different to that on the upper side (adaxial); in fact, attacks on the upper side are conﬁned to the epidermal cells and present as symptoms similar to the condition of russeting in immature fruit (Albrigo and McCoy, 1974). In the case of serious attacks, the cuticle may lose its shine and become bronze, with yellowish spots in the areas degreened by the release of ethylene in the production process of the wound (McCoy and Albrigo, 1975). In the lower surface this can lead to appreciable ‘mesophyll collapse’ as de-greened yellowish spots that later become necrotic (Albrigo and McCoy, 1974). A severe attack on the lower surfaces can increase transpiration and stimulate a consequent defoliation (McCoy, 1976). As regards the fruit, infestations of A. pelekassi affect the diameter, volume and weight of fruit, with those of damaged fruit being less than undamaged fruit. In particular, the sugar content of juice is higher in damaged fruit, showing that concentration of soluble solids through water loss is caused by mite attack (Tono et al., 1978). Severe infestations of P. citri lead to substantial changes in the transpiration and photosynthesis of the plants. In particular, transpiration increases during these attacks and decreases with a low mite density, and the quantity of chlorophyll may be reduced to 60% (Wedding et al., 1958). Heavy infestations of E. orientalis accompanied by poor irrigation, drought conditions and strong ventilation may induce defoliation (Jeppson et al., 1975). In general, attacks may disrupt the normal physiological processes and inﬂuence growth intensity, ﬂowering and production. The developmental delay of all attacked organs is very common. The sprouts may be shorter and accumulate less dry matter, with repercussions also in subsequent years. This scenario occurs frequently with many injurious mites, including the citrus red mite, P. citri. Productivity may suffer accordingly, with changes in performance appreciable even 2 or 3 years later. With regard to control, we need to consider seriously the importance of systems for damage assessment (McCoy et al., 1976a; Walker et al., 1992) and models for forecasting losses (Allen, 1981; Royalty and Perring, 1996).

5

Control

In general, the development of mite populations on crops may adversely affect the economy of a farm. This raises the need to understand the bioecology of the pest involved and of factors regulating the development of their populations, in order to place any decision in a context of wide rationality. It is not possible to generalize on knowledge acquired elsewhere, standardizing control measures and strategies in every region of the world. Each species must be examined in relation to its environment and conditions. This suggestion does not deny the value of information in your possession, but calls for each case to be placed in its ecological and productive context. Indeed, environmental conditions can variously inﬂuence the behaviour of mite populations. The populations of the same species may take different attitudes within the same geographical region or in different climatic zones and require different technical choices. This is the case for Panonychus citri and Eotetranychus kankitus Ehara in China, which were found to be thermophilic, causing damage on citrus shoots during warm, dry weather as the shoots grow (Li, 1990), or of Phyllocoptruta oleivora, Aceria sheldoni and Eutetranychus banksi in Florida and California. In hot humid areas of Florida, Ph. oleivora is the most harmful species, A. sheldoni is of secondary importance and E. banksi the most injurious pest, while in hot but arid areas of California, the most harmful species is P. citri, A. sheldoni is important and Ph. oleivora is an economic problem only in the coastal areas (Childers et al., 1996). The behaviours pointed out may interfere with important technical choices and have serious economic implications (see Chapter 1, p.10). The prevention of the accidental introduction of a new injurious species plays a fundamental role and there are calls to put into place all the mechanisms that help to intercept it on time and eradicate the ﬁrst outbreaks of development. In certain areas of the world, the interception of harmful or potentially © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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Chapter 5

harmful species, either directly or as vectors of serious diseases, is very important (Childers and Rodrigues, 2005) and for obvious reasons their control cautions against the use of chemicals. From a practical point of view, any decision is empowered to establish the real need for control. Nevertheless, the monitoring of the populations is difﬁcult because variables such as climate, location and mite chemical control can affect the economic threshold. McMurtry (1985) reported that the successful monitoring of mite populations is based more on experience than on the use of economic thresholds, i.e. more citrus red mites can be tolerated in California during spring and summer than during the autumn, when the dry winds may cause stress to the trees. In any case, each decision requires a reliable estimate of the attack, relating when and where possible the dynamics of population of the pest, with biotic and abiotic factors affecting it and with the economic implications that its evolution may involve in terms of damage (damage assessment) (Allen, 1981; Rabbinge, 1985). Independently of this reﬂection, the above-mentioned aspects indicate that the knowledge and application of a correct system of sampling and monitoring of populations, of economic injury level (EIL) and economic threshold (ET) of involving species, play a important part in the success of a control programme. There are various protocols relating to the most harmful species, such as Ph. oleivora in Florida, where Childers et al. (2007) suggested a sampling method based on rust mite density in processed fruit (rust mites/cm2), in late spring or summer, of six rust mites/cm2 as a planned threshold where pesticide treatment may be required within 10–14 days, and ten rust mites/cm2 would be an action threshold where treatment would be required as soon as possible; while in fresh fruit it is necessary to monitor mite populations every 10–14 days, and an average of two mites/cm2 is an action threshold. In the case of the citrus red mite, P. citri, Song et al. (2003) produced sequential sampling plans for adults of the citrus red mite, developing a decision-making citrus red mite population level based on the different action threshold levels (2.0 ± 0.25, 2.5 ± 0.25 and 3.0 ± 0.25 for mites/leaf). The maximum number of trees and required number of trees sampled on a ﬁxed-sample size plan on 2.0, 2.5 and 3.0 thresholds with a 0.25 precision level were 19, 16 and 15, and their critical values were 554, 609 and 659, respectively. Childers et al. (2007) found a relationship between the average number of Texas citrus mite, E. banksi, or citrus red mite, P. citri, and the percentage of leaves attacked across 10-acre (4 ha) blocks of young orange trees. An average of ﬁve spider mites/leaf correspond to 70–80% infestation levels, and this value constitutes a treatment threshold for processing fruit. Like pests injurious to other crops, the control of mites harmful to citrus is achieved through the use of biological, chemical and other means (horticultural, physical, etc.) according to three strategies of control: chemical, biological and integrated control, or IPM (integrated pest management).

Control

33

5.1 CHEMICAL CONTROL In citriculture, biological control has actually solved many entomological problems (Bennett et al., 1976) but, even today, the use of chemicals may be a necessity (McCoy, 1996a), taking into account environmental conditions, as in the case of the citrus rust mite, where chemical control frequently arises, especially in wetlands and crop varieties for fresh consumption (McCoy et al., 1989). Moreover, in this case, the rapid development of populations, the increase in the damage, the small size of the mite and the lack of natural enemies make monitoring difﬁcult (McCoy, 1996a). Chemical control is based on the use of various kinds of substances belonging to different chemical groups, commonly known as ‘acaricides’ and sometimes active also against insects. Among the oldest recorded is sulfur, used as a suspension in water or powder by the end of 19th century (Hubbard, 1885) and as lime sulfur by the beginning of last century (Yothers, 1915). Petroleum oils were employed for the next 20 years, followed by synthetic organic compounds, including dinitrophenol, widely used between 1930 and 1950, either alone or in combination with oils. From 1940, a series of acaricides belonging to different chemical categories began to appear, such as dithiocarbamates (zineb, maneb, mancozeb), diphenyl carbinols (chlorobenzilate, bromopropylate, dicofol), organochlorines (endosulfan), the sulfur-bridged (tetradifon, propargite), oxythioquinox, amitraz, organotins (cyhexatin, fenbutatin-oxide), organophosphates (ethion, pirimiphos-methyl, monocrotophos, phosalone, methamidophos), carbamates (carbaryl, carbosulfan, oxamyl, aldicarb), pyrethroids (fenpropathrin, ﬂuvalinate, ﬂucythrinate), clofentezine, hexythiazox, ﬂubenzimine, benzoylphenylureas (diﬂubenzuron, teﬂubenzuron, ﬂufenoxuron, ﬂucycloxuron), abamectin and the more recent pyridaben, fenazaquin, tebufenpyrad, brofenprox, spirodiclofen and other substances (Childers et al., 1996; Walsh, 2002; Bell et al., 2005; Childers et al., 2007). Some of these substances also act as a fungicide and insecticide. In each country the use of these chemicals is subject to speciﬁc legislative constraints and is included in ‘pesticide technology’, involving several aspects recently summarized by Jepson (2000), such as the efﬁciency of active ingredients, dose, distribution, farm monitoring and the detection of ecological impact. With regard to acaricides, the effectiveness of their action (Reed et al., 1967; McCoy et al., 1982; Helle and Overmeer, 1985b; Childers and Peregrine, 1986; Childers, 1996), toxicological and ecotoxicological aspects, such as the selectivity (Overmeer, 1985; Childers et al., 1996), dose and distribution and relating the treatments to the risk of selection of resistant strains (Jeppson et al., 1975; Georghiou and Saito, 1983; Cranham and Helle, 1985; Messing and Croft, 1996; Childers et al., 2007) have also been examined.

5.2 SIDE EFFECTS OF CHEMICALS The additional side-effects of various substances commonly used in crop protection (Duncombe, 1972; Coates, 1974; McCoy, 1977a, b; Beattie et al.,

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1991; Smith and Papacek, 1991) or the selection of resistant strains to various chemicals (Swirski et al., 1967; Jeppson et al., 1975; Herne et al., 1979; Omoto et al., 1994) stimulating the development of mite populations are well known. In this context, the negative roles of organophosphates (Attiah and Wahba, 1973), synthetic pyrethroids (Gerson and Cohen, 1989) or the contradictory role of carbamates, which primarily control major pests such as the citrus rust mite (Johnston et al., 1957) and secondarily help the development of the two-spotted spider mite (Boykin and Campbell, 1982; Childers, 1990) but inhibit the beneﬁcials (Buskovskaya, 1976; Kashio and Tanaka, 1979), are widely veriﬁed. Some carbamates such as aldicarb are effective against eriophyoid and tetranychid mites injurious to citrus (French and Timmer, 1981; Childers et al., 2007) but possess high mammalian toxicity, persistence in plants and are a risk to groundwater contamination, and thus their use has been restricted (Childers et al., 1996). Some new principles allowed today provide appropriate answers to different technical needs, allowing the realization of concrete IPM programmes (Childers et al., 1996, 1997; McCoy, 1996a). Understanding the side-effects of chemicals used in agriculture against injurious mites is crucial from an ecological and practical point of view, allowing both pest control and the stability of agro-ecosystems through the preservation of beneﬁcial mite populations. In this context, the research of the working group ‘Pesticides and Beneﬁcial Organisms’ of the International Organization for Biological Control (IOBC), which periodically produces reports on the inﬂuence of new chemicals (insecticides, acaricides, fungicides, herbicides), is highly valuable. With regard to predatory mites, an interesting report was published on the inﬂuence of 20 substances on three important species of phytoseiids (Sterk et al., 1999). Detailed information in this regard has been published by Croft (1990) and Kostiainen and Hoy (1996). Gerson et al. (2003) recently presented an update on the knowledge gained on the inﬂuence of various substances on the principal groups of beneﬁcial mites. Examining the factors that affect the impact of chemicals on beneﬁcial mites, they display the role of formulation, solvents and application modes, the persistence of various active ingredients, the mode of action, the effects of chemicals on plants and the biology of beneﬁcial chemicals. With reference to the side-effects, they examined the importance of increased fecundity (Grout et al., 1997), the removal of competitors and alternative prey, repellence and dispersal (Hislop et al., 1981). Equally important in this context is the selection of strains of pesticide-resistant mites, known for some time (Huffaker and Kennett, 1953) and investigated in recent decades (Gerson et al., 2003). The issues brieﬂy illustrated clearly indicate that the use of chemicals against mites (and other pests) injurious to citrus crops or other crops requires adequate knowledge about the side-effects that their use entails, without which no programme of biological control and/or IPM can be carried out.

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5.3 BIOLOGICAL CONTROL The ecology of mite populations is inﬂuenced by abiotic factors, such as climatic conditions, and biotic, like the potential biotic of the species, the interaction with the host plant, the natural enemies and the agricultural practices in general (Jeppson et al., 1975; Zamorra and Nasca, 1985). Of these factors, natural enemies, whose action in citrus crops has been known for a long time, play an important role (DeBach and Bartlett, 1951; DeBach, 1969, 1974). The natural enemies of mites include pathogens such as viruses, bacteria and fungi (Shaw et al., 1968; Reed et al., 1972; Jeppson et al., 1975; Muma et al., 1975; Simanton, 1976; van der Geest, 1985; McCoy, 1996b; Paz et al., 2009a); predatory mites, including Phytoseiidae (Muma et al., 1975; Chant, 1985; Helle and Sabelis, 1985; Sabelis, 1996; Grout, 1998), Stigmaeidae (Santos and Laing, 1985; Thistlewood et al., 1996), Anystidae, Cheyletidae, Erythraeidae, Bdellidae, Tarsonemidae, Tydeidae and Cunaxidae (Jeppson et al., 1975; Muma et al., 1975; Gerson, 1985b; Vacante, 1986; Gerson and Vacante, 1993; Perring and McMurtry, 1996; Grout, 1998); spiders (Carroll, 1980; Gerson, 1985b; Mansour and Whitcomb, 1986; Benfatto et al., 1992; Yan et al., 1992; Rodriguez Almaraz and Contreras-Fernandez, 1993; Dippenaar Schoeman, 1998); and various insects including the Coleoptera Coccinellidae and Staphylinidae, Neuroptera, Thysanoptera, the Diptera Cecidomyiidae and other species, Dermaptera, Hemiptera Anthocoridae, Miridae, Nabidae and Lygaeidae (Jeppson et al., 1975; Muma et al., 1975; Chazeau, 1985). Of the above-cited organisms, the importance of the Acari as biocontrol agents has been proposed for some time (Hoy et al. 1983; Gerson and Smiley, 1990) and Gerson et al. (2003) have recently dealt with the fundamental aspects that inﬂuence the technical choice, examining about 30 families of beneﬁcial mites variously involved in the biological control of injurious mites and/or other pests, and calling attention to scientiﬁc and practical implications that help the implementation of the method. In particular, Gerson et al. (2003) gave a realistic and modern deﬁnition of the biological control of pests by mites based on acarine actions, reducing pest density and/or the extent of their damage to below economic injury level. The choice includes natural and applied reductions, and permits the use of three basic strategies, such as importation, preservation and augmentation, that introduce IPM. With regard to injurious mites infesting citrus, the action of natural enemies seems more promising in the biological control of tetranychids (McMurty, 1985) and less in that of eriophyoids (McCoy, 1996a) with some exceptions, such as those reported for Queensland (Australia) by Smith and Papacek (1991) for Euseius victoriensis (Womersley). Regardless of this constraint, the action of natural enemies, if properly managed, could allow the implementation of concrete IPM programmes (McCoy, 1985, 1996a).

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5.4 INTEGRATED PEST MANAGEMENT Integrated pest management (IPM) was proposed between the late 19th and early 20th centuries. However, in the last 50 years the method has been deﬁned in a more scientiﬁcally appropriate way (Stern et al., 1959). Of the authors who have investigated the strategy, Flint and van den Bosch (1981) adopted probably the most comprehensive deﬁnition and report that integrated pest control is based on the factors of plant resistance and natural mortality, such as that caused by natural enemies, and take control techniques that disturb as little as possible to these factors. The method does not exclude the use of chemicals but uses them as a last resort after a systematic monitoring of pest populations and if the natural control factors will pose the need. An integrated program considers all possible control measures, including the option not to act, and assesses the potential interaction between different techniques, cultivation practices, the development climate, the various pests and the crop.

A deﬁnition of IPM more adherent to our needs we should probably be that of Rosen (1986), who writes that: Integrated pest management (IPM) provides a reasonable compromise, taking into account both the desirability of biological control and the need for some form of chemical control. Like diplomacy, IPM is the art of the possible. It represents a holistic approach, recognizing the unit of the ecosystem and harmonizing all available measures in an attempt to optimize pest control and crop production. Effective IPM may be achieved through the development of a vigorous program of applied biological control, in combination with a relatively judicious use of selective pesticides, only when absolutely necessary and in the least disruptive modes of application. Other selective tactics should be incorporated into the program whenever applicable.

The strategy aims to control pest populations using all possible means (chemical, biological, biotechnical, horticultural, etc.) that do not impede the action of indigenous and artiﬁcially introduced natural enemies, and in agreement with the ecological, toxicological and economic requirements of agricultural production. The method involves both their economic and ecological mechanisms, deﬁning a strong phytoiatric vision, resulting in multiple technologies and adaptations to speciﬁc areas of cultivation. IPM is a fundamental prerequisite for the realization of ‘integrated production’, meaning a holistic approach to the production system. The farm represents a base unit and the agro-ecosystem a central point of intervention, in which the preservation and improvement of soil fertility and diversity of the environment are essential. Biological, technical and chemical methods must be balanced and respectful of environmental protection, income and social needs (El-Titi et al., 1995). The strategy aims to link the total cost of citrus crop defence with the effectiveness of the control programme, aiming to escape the constraint of dealing with every adversity separately and the consequent high cost of control. Its application becomes economically viable in large areas, loading the economic cost of operations on to a larger number of farmers and covering

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all adversities (Graebner et al., 1984). Furthermore, the implementation of a programme requires adequate technical support for the mass production of beneﬁcials, used mainly in augmentative biological control programmes (Luck and Forster, 2003). It is possible to ﬁnd further information in the most signiﬁcant experiences gained in this regard on citrus in different works (McCoy, 1985, 1996a; McMurtry, 1985; Browning, 1992; Childers et al., 1996; Bedford, 1998a; Grout, 1998).

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II

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Citrus Mites

Key to the identiﬁcation of subfamilies, tribes, genera and species. Morphological characters, systematic outline, bio-ecology, natural enemies, symptomatology, damage and control of known species of Phytoptidae, Eriophyidae, Diptilomiopidae, Tarsonemidae, Tenuipalpidae, Tuckerellidae and Tetranychidae.

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6

Key for the Identiﬁcation of Subfamilies, Tribes, Genera and Species1

1. Body vermiform or fusiform with two pairs of legs … 2 - Body rounded with more than two pairs of legs … 4 2. Prodorsal shield without anterior setae (vi or ve); scapular setae (sc) present or absent; never with tibial solenidion (f); opisthosoma never with subdorsal seta (c1); female spermathecal tubes always short, either projecting laterally or diagonally caudal … 3 - Prodorsal shield with one to ﬁve setae, always with anterior setae present (paired or unpaired vi and/or paired ve); opisthosoma with all usual setae, and some species with subdorsal seta pair; female spermathecal tubes usually long (three to ﬁve or more times longer than spermathecal tubes in Eriophyidae and Diptilomiopidae), often extending diagonally forward then recurving caudally … family Phytoptidae Murray … 7 3. Gnathosoma commonly small compared with body; when large, with chelicerae straight or slightly curved; terminal segments of palps short and truncated and enclosing the short oral stylet; female genital coverﬂap commonly with ridges … family Eriophyidae Nalepa … 8 - Gnathosoma large compared with body; chelicerae abruptly curved and bent down near base; palps attenuated, enclosing the long oral stylet; female coverﬂap commonly smooth, less often with ridges … family Diptilomiopidae Keifer … 24 1The

Eriophyoidea key is based on the works of Lindquist and Amrine (1996) and Amrine et al. (2003), the Tenuipalpidae key on the works of Meyer (1979), the Tuckerellidae key on the works of Meyer and Ueckermann (1997) and the Tetranychidae key on the works of Meyer (1987) and Bolland et al. (1998). The key is fundamentally based on adult morphological characteristics, with the exception of Brevipalpus jordani Dosse and Brevipalpus phoenicoides Gonzalez, of which the juvenile stages are also taken into account. ‘A catalogue of Tenuipalpidae (Acari) of the World with a key to genera’ (Mesa et al., 2009) was very recently published and therefore missed being included in this book. The reader may be interested to consult this work.

© V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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4. Palps without thumb–claw complex … 5 - Palps with thumb–claw complex … 6 5. Body not ﬂattened; female with one pair of prodorsal bothridia and with pseudostigmatic organs; ﬁve dorsal idiosomal tergites; chelicerae not markedly long; tarsal claws without tenent hairs; leg IV of female 3-segmented … family Tarsonemidae Canestrini et Fanzago … 25 - Body ﬂattened; any prodorsal bothridia with pseudostigmatic organs, and dorsal idiosomal tergites; chelicerae long, recurved and stilet-like; tarsal claws with tenent hairs; leg IV of the female with more than three segments … family Tenuipalpidae Berlese … 26 6. Tarsi I and II without duplex setae and with distal sensorial setae; female genitalia without folds; histerosoma with 36 fan-shaped or palmate setae and a row of ﬂagellate caudal setae … family Tuckerellidae Baker et Pritchard … 48 - Tarsi I and II with duplex setae; female genitalia with folds; histerosoma maximum with 12 pairs of setae, and devoid of ﬂagellate setae … family Tetranychidae Donnadieu … 53 7. Prodorsal shield with two anterior setae (ve) present; interior verticals (vi) absent; setae sc rarely minute or absent; spermathecal tubes short; body vermiform, with opisthosomal annuli narrow and subegual dorsoventrally; setae sc pointing up if short, forward if long; opisthosomal seta pair c1 present (subfamily Phytoptinae Murray); femora and genua separate, not fused; prodorsal shield lacking gland; setae sc short or long, but not minute; microtubercles distributed normally on dorsal annuli (genus Phytoptus Dujardin); sides of the prodorsal shield and coxae I granulate … Phytoptus ﬁcivorus (Channabasavanna) 8. Tibiae reduced or completely fused with tarsi; tibiae without seta … 9 - Tibiae always distinct from tarsi, tibial seta usually present … 10 9. Spatulate projections absent from tarsi, legs of average thickness; coxae of legs I often fused across centre line with sternal line faint or absent; empodia relatively small (subfamily Nothopodinae Keifer); coxal setae 1b absent; coxae and tibiae of leg I variable (tribe Nothopodini Keifer); prodorsal shield tubercles ahead of rear margin, scapular setae (sc) directed posterolaterally; coxae I more or less fused with subcapitulum (genus Cosella Newkirk et Keifer); opisthosoma with about 45 dorsal annuli and 50 ventral annuli; microtubercles elongate, obscure or absent dorsally; setae h1 absent; genital coverﬂap granular; empodia four-rayed … Cosella ﬂeschneri (Keifer) (Fig. 8.3) 10. Female genitalia appressed to coxae separating coxae more than normal, and, in lateral view, usually noticeably projecting from venter; coxae I usually narrowly connate at centre line, sternal line shortened; coxae often with curved lines outlining tubercles of setae, especially setae 1a; female genital apodeme bent up and shortened, commonly appearing as a heavy transverse line in ventral view; female genital coverﬂap typically in two uneven ranks (subfamily Cecidophyinae Keifer); scapular tubercles and setae present (tribe Colomerini Newkirk et Keifer); opisthosoma with broad dorsal annuli and narrow ventral annuli; dorsal annuli with elongate microtubercles, except faint on about ﬁve or six annuli before setae f (genus Circaces Keifer); scapular setae (sc) 4 μm long and direct to rear and diverging; opisthosoma

Key for Identiﬁcation

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with 28 dorsal annuli, smooth and evenly arched and 44 microtuberculate ventral annuli; microtubercles elongated; setae h1 absent; genital coverﬂap granulate … Circaces citri Boczek Female genitalia not appressed to or separating coxae, and, in lateral view, lying more on a level with venter; female genital apodeme commonly extending moderate distance forward, does not appear as a heavy transverse bar in ventral view; female genital coverﬂap variably sculptured, ridges rarely occurring in two ranks … 11 11. Body vermiform, annuli subequal dorsoventrally, at least on anterior half to two-thirds of opisthosoma; prodorsal shield typically lacking a frontal lobe, or with a slight projection over gnathosoma base; if frontal lobe is present, then it is narrow, basally ﬂexible and combined with narrow annuli … subfamily Eriophyinae Nalepa … 12 Body commonly more fusiform; prodorsal shield commonly with a broad-based and rigid frontal lobe over gnathosoma; opisthosoma typically divided into broad and stout dorsal annuli, and narrow, microtuberculate ventral annuli; if frontal lobe is absent or only a slight one present, then annuli differ dorsoventrally, at least in larger dorsal microtubercles; if annuli are subequal, and broad frontal lobe is absent, the annuli are as broad as the genital coverﬂap is long … subfamily Phyllocoptinae Nalepa … 13 12. Prodorsal shield tubercles on, or very near, rear shield margin with transverse basal axes, setae directed to rear, usually divergently (tribe Aceriini Amrine et Stasny); opisthosoma lacking a broad longitudinal dorsal furrow; posterior opisthosoma with annuli continuous and subequal dorsoventrally; gnathosoma of normal length; tibial setae present; coxal seta 1b present (genus Aceria Keifer); opisthosomal microtubercles elliptical and touching the edge of the annuli; female genital coverﬂap from ten to 12 longitudinal ribs; empodia ﬁve-rayed … Aceria sheldoni (Ewing) (Fig 8.2) 13. Scapular setae (sc) absent, scapular tubercles present or absent; empodia entire (tribe Calacarini Amrine et Stasny); dorsum with three ridges; middorsal ridge narrow, extends as far as lateral ridges; opisthosoma tapers evenly toward rear; tibial setae present (genus Calacarus Keifer); scapular setae (sc) missing; opisthosoma with about 60 dorsal annuli and 65–70 ventral annuli; with ﬁve white longitudinal wax bands, gradually diminishing in distinctness caudally; genital coverﬂap with many ﬁne longitudinal lines placed in two ranks; empodia ﬁve-rayed … Calacarus citrifolii Keifer (Fig. 8.8) Scapular setae (sc) and tubercles present … 14 14. Scapular setae (sc) usually with well-formed, often plicate, tubercles placed ahead of rear margin, directing setae forward, up or central; if tubercles and setae are near rear shield margin, then tubercles are subcylindrical and bent forward or the alignment of their bases is longitudinal or diagonal to the body (tribe Phyllocoptini Nalepa); prodorsal shield with a distinct frontal lobe over the gnathosoma; opisthosoma with a wide mid-dorsal longitudinal furrow, with edges longitudinal and parallel; coxal setae 1b present … genus Phyllocoptruta Keifer) … 15 Scapular setae (sc) with tubercles on or very near the rear shield margin, directing setae to rear, commonly divergently; scapular tubercles either

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subcylindrical, or the alignment of their bases is transverse to the body … tribe Anthocoptini Amrine et Stasny … 17 15. Empodia ﬁve-rayed … 16 Empodia four-rayed; prodorsal shield with scapular setae (sc) backward and upward; opisthosoma without subdorsal ridges; female genital coverﬂap with ten longitudinal ribs … Phyllocoptruta paracitri Hong et Kuang (Fig. 8.11) 16. Female genital coverﬂap with basal area granular and a median longitudinal line, and apical area with 14–16 longitudinal ribs … Phyllocoptruta oleivora (Ashmead) (Fig. 8.10) Female genital coverﬂap with 12 longitudinal scorelines … Phyllocoptruta citri Soliman et Abou Awad … (Fig. 8.9) 17. Dorsal opisthosoma evenly rounded (a slight mid-dorsal ridge may be present, anteriorly) … 18 Dorsal opisthosoma with distinct mid-dorsal furrow or ridge … 20 18. Prodorsal shield frontal lobe acuminate, frequently ending in a sharp point; never with spinules under front edge … genus Aculops Keifer … 19 Prodorsal shield frontal lobe broad and rounded, not acuminate; some species with two to four small spines or spinules projecting forward from under front edge (genus Aculus Keifer); opisthosoma with 47–50 dorsal annuli, and 57–58 ventral annuli, dorsal annuli smooth and ventral annuli microtuberculate; genital coverﬂap weakly marked with ﬁve or six furrows; empodia four-rayed … Aculus advens (Keifer) (Fig. 8.5) 19. Admedian lines complete, extending from the sides of the frontal lobe to the rear margin, somewhat sinuate, and furthest apart the rear; admedian also receive the cross-lines at half and two-thirds, which connect them with the median; microtubercles on the dorsal annuli faint, beadlike on the ventral annuli, and rest on the ring margins; setae d 40 μm, on ventral annulus 16 … Aculops pelekassi (Keifer) (Figs 8.1 and 8.4) A short transverse line running clearly across the shield between anterior part of admedian line near the frontal lobe of process, setae d especially long and reaching 82.8 μm, and the microtubercles of the last six ventral annuli on the opisthosoma long and thin and regularly arranged … Aculops suzhouensis Xin et Dong 20. Opisthosoma with mid-dorsal longitudinal furrow … 21 Opisthosoma with mid-dorsal longitudinal ridges … 22 21. Prodorsal shield without posterior plate; coxal setae 1b present; genu II setae present; scapular tubercles near midline, sc strongly recurved laterally; subdorsal ridges distinct (genus Paratetra Channabasavanna); shield design with a pattern of ridges with a prominent broad V-shaped ridge between the dorsal tubercles; opisthosoma with 30–32 smooth, broad dorsal annuli, and about 70 microtuberculate ventral annuli; ﬁrst three to four dorsal annuli narrowed and not depressed and the rest form a broad, longitudinal trough ﬂanked by a narrow ridge on either side; the last three to four dorsal annuli form complete rings; the microtubercles on the dorsal annuli are tiny and beadlike, set along rear margins and those on the last 8–10 ventral annuli microstriate … Paratetra murrayae Channabasavanna

Key for Identiﬁcation

45

22. Opisthosoma with three ridges; mid-dorsal opisthosomal ridge stronger than lateral ridges but fades caudally, not ending in a furrow; coxal setae 1b present … genus Tegolophus Keifer … 23 23. Dorsocentral ridge beginning at third dorsal annulus and fading on 16th dorsal annulus; ﬁrst dorsal annulus broad and rough; female genital coverﬂap with 18 longitudinal ribs … Tegolophus australis Keifer (Fig. 8.6) Dorsocentral ridge extending throughout the entire opisthosoma; ﬁrst dorsal annulus similar to other annuli; female genital coverﬂap with 12–14 (maximum 16) longitudinal ribs … Tegolophus brunneus Flechtmann (Fig. 8.7) 24. Empodium divided; usually deeply (subfamily Diptilomiopinae Keifer); scapular setae (sc) absent; genu absent from both legs; coxal setae 1b absent; tibial setae absent (genus Diptilomiopus Nalepa); dorsal tubercles, minus scapular setae (sc), present within rear shield margin; opisthosoma with about 55–60 dorsal annuli, and 70–75 ventral annuli; setae c2 missing; setae h1 absent; genital coverﬂap smooth; empodia ﬁve-rayed … Diptilomiopus assamica Keifer (Fig. 9.1) 25. Metapodosomal venter of adult with three or four pairs of setae; coxisternal plates III commonly with two or three pairs of setae, and plates IV with one pair (subfamily Pseudotarsonemoidinae Lindquist); cheliceral stylets short to moderate length, when retracted not occupying most of the length of gnatosomal capsule and not recurved; coxisternal plates III with three pairs of setae, and plates IV with one pair of seate (tribe Pseudotarsonemoidini Lindquist); femur I and tarsus II of the female lacking seta l’’ and spinelike seta pl’’, respectively; female with gently curved, sessile claw and reduced, contiguous unguinal setae on leg I; prodorsal shield of the female unenlarged not covering the stigmata; dorsal idiosomal setae slender, with setae sc2 longer than other setae; female with tarsal setae pv’ usually enlarged, spine-like on leg I, and only two setae on tibiotarsus of leg IV (genus Polyphagotarsonemus Beer et Nucifora); both sexes with short body setae; female with a claw on tibiotarsus of leg I, and with four pairs of coxal setae and spherical pseudostigmatic organs; male with three pairs of propodosomal setae and four pairs of metapodosomal ventral setae; tibia and tarsus IV fused in a tibiotarsus ending in a button-like claw; coxae III and IV closely associated … Polyphagotarsonemus latus (Banks) (Fig. 10.1) 26. Without ventral plates … subfamily Tenuipalpinae Mitrofanov … 27 With ventral plates … subfamily Brevipalpinae Mitrofanov … 32 27. Without plates in the genito-anal area, with three pairs of setae near to genital opening and three pairs near to anal opening; palp four-segmented (genus Ultratenuipalpus Mitrofanov); ﬁrst pair of dorsolateral hysterosomal setae lanceolate and minute … Ultratenuipalpus gonianensis Sadana et Sidhu (Fig. 11.19) With plates in the genito-anal area; penultimate pair of hysterosomal dorsolateral setae commonly ﬂagellate; if these last present are of normal length, the podosoma is very wide and the opisthosoma is very narrow … genus Tenuipalpus Donnadieu … 28 28. Opisthosomal dorsal setae c1 and d1 strongly spatulate and longer than distances between their bases … Tenuipalpus caudatus (Dugès)

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Opisthosomal dorsal setae c1 and d1 minute and shorter than the distances between their bases … 29 29. Prodorsal setae sc2 broad, lanceolate and strongly longer than other prodorsal setae … Tenuipalpus orilloi Rimando (Fig. 11.18) Prodorsal setae sc2 longer than other prodorsal setae but not strongly … 30 30. Prodorsum with transverse broken striae and with broken thick lines laterally; dorsal opisthosoma with irregular, broken, transverse striae, and with a few longitudinal striae to the caudal end … Tenuipalpus mustus Chaudhri Prodorsum without transverse broken striae … 31 31. Prodorsum with mediodorsal area outlined by longitudinal striae, and with irregular striae inside this area; lateral prodorsum irregularly striate; dorsal opisthosoma with transverse to irregular striae … Tenuipalpus emeticae Meyer (Fig. 11.17) Prodorsum with a pair of L-shaped ridges back to back along midline, both Ls sometimes broken at foot, the rest of the dorsum broken up by innumerable small ridges; dorsal opisthosoma with an oval area of small, more or less longitudinal ridges bounded anteriorly and posteriorly by stronger ridges and laterally by the pores, the rest of the dorsum broken up by small ridges … Tenuipalpus sanblasensis De Leon (Fig. 11.17) 32. Hysterosoma without dorsosublateral setae or not more than one pair; penultimate pair of dorsolateral setae commonly of normal length; genital and anal plates well deﬁned and well separated; ventral plate rectangular … genus Brevipalpus Donnadieu … 33 Hysterosoma with more than 11 pairs of dorsal setae and with fewer than four pairs of dorsosublateral setae; palp with more than two segments; rostral shield when present always with broad lobes; female metapodosoma not separated from opisthosoma by transverse furrow (genus Pentamerismus McGregor); hysterosoma with six pairs of dorsolateral setae, all setiform; dorsum evenly striate … Pentamerismus tauricus Livshitz et Mitrofanov 33. Tarsus II with one solenidion … 34 Tarsus II with two solenidia … 42 34. With nine opisthosomal dorsal setae … 35 With ten opisthosomal dorsal setae … 39 35. Dorsal body more or less uniformly areolate … 36 Dorsal body not uniformly areolate … 37 36. Rostral shield not extending beyond the base of femur I; central area of prodorsum changing from smooth to rugose, and medial area uniformly reticulate … Brevipalpus obovatus Donnadieu (Fig. 11.14) Rostral shield extending beyond the base of femur I; central area of prodorsum and opisthosoma uniformly reticulate … Brevipalpus chilensis Baker (Fig. 11.5) 37. Rostral shield extending distinctly beyond the base of femur I … Brevipalpus mcgregori Baker (Fig. 11.13) Rostral shield not extending distinctly beyond the base of femur I … 38 38. Irregular broken striae cover the major part of the body, with a number of mediolateral reticulations; striae fade away laterally, and the area in the middle is devoid of any ornamentation … Brevipalpus amicus Chaudhri (Fig. 11.3)

Key for Identiﬁcation

47

Prodorsal reticulate pattern with a median ridge and broad lateral margin irregularly rugose … Brevipalpus deleoni Pritchard et Baker (Fig. 11.8) 39. Rostral shield extending beyond the base of femur I … 40 Rostral shield not extending beyond the base of femur I … 41 40. Anterior half of dorsal opisthosoma reticulate; posteriorly the mediolateral portion is reticulate; longitudinal, broken striae anterior and posterior to setae e1; striae directed marginally on lateral sides … Brevipalpus dosis Chaudhri, Akbar et Rasool (Fig. 11.9) Dorsal opisthosoma reticulate in the middle, with broader cells in the sides … Brevipalpus cucurbitae Mohanasundaram (Fig. 11.6) 41. The dorsal reticulate pattern may be irregular; the lateral areolae on the prodorsum longer than they are wide and on the anterior part, where the pattern becomes dorsal, the elements are wider than they are long; on the posterior part of the prodorsum the pattern does not seem to tend dorsally; the opisthosomal pattern is more indeﬁnite and on the dorsal surface the areolae are broken and wider than they are long; on lateral sides just outside the dorsal setae the areolae are longer than they are wide … Brevipalpus cuneatus (Canestrini et Fanzago) (Fig. 11.7) The prodorsal and opisthosomal reticulate pattern does not meet dorsally and with areolae longer than they are wide … Brevipalpus lewisi (McGregor) (Fig. 11.12) 42. With ten opisthosomal dorsal setae … Brevipalpus californicus (Banks) (Fig. 11.4) With nine opisthosomal dorsal setae … 43 43. Prodorsal shield twice as narrow as its length … Brevipalpus jambhiri Sadana et Balpreet Prodorsal shield more than twice as wide as its length … 44 44. Prodorsal shield with broken striae … 45 Prodorsal striae without broken striae … 46 45. Prodorsum with irregular and broken striae in the middle and laterally, and reticulate mediolaterally; opisthosoma with longitudinal wavy lines meeting medially and caudally, and lateral striae directed marginally … Brevipalpus rugulosus Chaudhri, Akbar et Rasool (Fig. 11.16) Prodorsum with dim reticulations in middle, and reticulate mediolaterally; opisthosoma with irregular, longitudinal striae, thick, meeting caudally in the middle, wavy, forming cells caudally, and the lateral striae directed marginally … Brevipalpus karachiensis Chaudhri, Akbar et Rasool (Fig. 11.11) 46. Prodorsal shield with ﬁrst and second lateral projections little and near … Brevipalpus tinsukiaensis Sadana et Gupta First and second lateral projections of prodorsal shield different … 47 47. Protonymph with opisthosomal setae d3 and e3 smaller than other opisthosomal dorsal setae … Brevipalpus phoenicis (Geijskes) (Figs 11.1, 11.2) Protonymph with opisthosomal setae d3 and e3 of the same size as other opisthosomal dorsal setae … Brevipalpus jordani Dosse (Fig. 11.10) and Brevipalpus phoenicoides Gonzalez (Fig. 11.15) 48. Idiosoma with 44 dorsal setae fan-shaped or palmate; several setae of legs I and II fan-shaped … genus Tuckerella Womersley … 49

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Chapter 6

49. Opisthosoma with ﬁve pairs of ﬂagellate caudal setae … 50 Opisthosoma with six pairs of ﬂagellate caudal setae … 51 50. Opisthosomal ﬂagellate caudal setae equal in length; opisthosomal setae f1–2 about equal in length; f1 situated slightly anteriad of f2 … Tuckerella ornata (Tucker) (Figs 12.1, 12.4) 51. Opisthosomal setae f1–2 with rounded tips … 52 Opisthosomal setae f1–2 with pointed tips, subequal in length … Tuckerella nilotica Zaher et Rasmy (Figs 12.2, 12.4) 52. Opisthosoma setae f1–2 subequal in length; opisthosomal setae c4 about half as long as distances between their setal bases and those of prodorsal setae sce; reticulations on posterior part of opisthosoma tend to be irregular … Tuckerella knorri Baker et Tuttle (Fig. 12.3) Opisthosoma setae f1–2 unequal, f2 larger in size than f1; prodorsal setae vi speculated, about as long as broad … Tuckerella pavoniformis (Ewing) (Figs 12.2 and 12.3) 53. Empodium with tenent hairs; female with two or three pairs of pseudanal setae (ps1–3) and male with ﬁve pairs of genital (g1–2) and pseudanal setae (ps1–3) … subfamily Bryobiinae Berlese … 54 Empodium without tenent hairs or absent; female with one or two pairs of pseudanal setae (ps1–2) and male with four pairs of genital and pseudanal setae (ps1–2, g1–2) … subfamily Tetranychinae Berlese … 56 54. True claw uncinate; empodium pad-like (tribe Bryobiini Reck); opisthosoma with 12 pairs of dorsal setae; prodorsum with four anterior projections and four pairs of setae; empodia II–IV with more than one pair of tenent hairs … genus Bryobia Koch … 67 True claw pad-like; empodium pad-like or uncinate … 55 55. True claw and empodium pad-like (tribe Hystrichonychini Pritchard et Baker); prodorsum with three pairs of setae and without anterior projections; opisthosoma with ten pairs of setae; setae f1 in normal position; tarsus I with two series of duplex setae; some dorsal setae or all on strong tubercles … genus Aplonobia Womersley … 70 True claw pad-like and empodium uncinate (tribe Petrobiini Reck); prodorsum without projections over gnathosoma; body with a maximum of 15 pairs of dorsal setae; three pairs of medioventrals; setae h2–3 in ventral position; empodium curved distally and with two rows of tenent hairs; true claw without hair-like processes … genus Petrobia Murray … 72 56. Empodium when present is claw-like; tarsus I with loosely associated setae or with one pair of duplex setae; when tarsus I possesses two pairs of duplex setae, tarsus II is devoid … tribe Eurytetranychini Reck … 57 Empodium claw-like or split distally; tarsus I with two pairs of duplex setae and tarsus II with one pair … 59 57. Two pairs pseudanal setae (ps1–2) … 58 One pair of pseudanal setae; ten pairs of dorsal opisthosomal setae … genus Aponychus Rimando … 75 58. Ten pairs of dorsal opisthosomal setae … genus Eutetranychus Banks … 76

Key for Identiﬁcation

49

Nine pairs of dorsal opisthosomal setae (c2 absent) (genus Meyernychus Mitrofanov); dorsal opisthosomal setae vary from subspatulate to spatulate and some are set on small tubercles; dorsal opisthosomal setae e1 and f1 form a rectangle and the striae between setae d1 and e1 are longitudinal; peritremes bilobed distally; receptaculum seminis depicted … Meyernychus emeticae (Meyer) (Fig. 13.20) 59. Opisthosoma with setae f1 in marginal position or absent (tribe Tenuipalpoidini Pritchard et Baker); setae f1 in marginal position; empodium consisting of a simple claw; tarsus II with distal setae of duplex setae consisting of a long and tapered solenidion (genus Tenuipalponychus Channabasavanna et Lakkundi); dorsal body setae set on strong tubercles, serrate spatulate or sometimes tapering and not longer than the distances between their bases; scapular setae sc2 longer than other prodorsal setae; central area of the prodorsum with reticulate pattern consisting of polygonal elements; opisthosoma feebly striate, with striae formed by discontinuous elements and transversal except between the dorsal opisthosomal setae e1, where they are longitudinal; opisthosomal setae f2 as long as half f1 … Tenuipalponychus citri Channabasavanna et Lakkundi Opisthosoma with f1 in normal dorsal position … tribe Tetranychini Reck … 60 60. Two pairs of setae h (h2–3) … 61 One pair of setae h … 66 61. Empodium claw-like, entire or split in two claw-like structures … 63 Empodium split distally or ending in a tuft of hairs … 62 62. Empodium split distally; dorsal setae long and strong, set on tubercles; one pair of pseudanal setae (genus Acanthonychus Wang); idiosoma striate transversally, also on tubercles; dorsal propodosomal setae, except the second pair, set on small circular tubercles; dorsal opisthosomal setae, except the c3, f1, f2 and h1, set on strong, prominent conical tubercles; all dorsal setae strong, and deeply serrated … Acanthonychus jiangfengensis Wang (Fig. 13.21) Empodium split near the middle in three pairs of hairs; opisthosomal integument striate transversally; dorsal body setae as long or longer than the intervals between their bases; two pairs of pseudanal setae (ps1–2) … genus Eotetranychus Oudemans … 83 63. Empodium a single claw-like structure … 64 Empodium split into two claw-like structures and commonly with hairs … 65 64. Empodium lacking proximoventral hairs; empodial claw strong; dorsal setae stout; reticulate pattern on the integument; ten pairs of dorsal opisthosomal setae … genus Mixonychus Meyer et Ryke … 91 Empodium with proximoventral hairs, as long or longer than proximoventral hairs arising at right angles from the claw … genus Panonychus Yokoyama … 92 65. Opisthosoma with ten pairs of dorsal setae … genus Schizotetranychus Trägårdh … 94

50

Chapter 6

66. Empodium split distally, commonly into three pairs of hairs; duplex setae of tarsus I well separated; empodial spur commonly visible; peritreme recurved distally … genus Tetranychus Dufour … 98 Empodium claw-like, as long or longer than the proximoventral hairs; duplex setae of tarsus I distal and adjacent; opisthosoma with ten pairs of dorsal setae; two pairs of pseudanal setae (ps1–2) … genus Oligonychus Berlese … 112 67. Prodorsal projections without little ridges pointed … 68 Prodorsal projections may bring little ridges pointed … 69 68. Prodorsal central projections partially jointed between them and as large as high, and the medial larger than high and rounded or triangular … Bryobia graminum (Schrank) (Fig. 13.1) Prodorsal central projections higher than medial and the base wider than high, and the medial projections are higher than large … Bryobia praetiosa Koch (Fig. 13.2) 69. Body approximately 650 μm long and distal anasthomosis of peritreme approximately 20 μm long … Bryobia rubrioculus (Scheuten) (Fig. 13.3) 70. Dorsal body setae expanded distally and receptaculum seminis large and oval … Aplonobia citri Meyer (Fig. 13.4) Dorsal body setae tapering distally … 71 71. Setae c3 lateral to setae c2 and receptaculum seminis broadly rounded … Aplonobia honiballi Meyer (Fig. 13.5) Setae c3 more or less in a line with dorsolateral setae and receptaculum seminis large and oval … Aplonobia histricina (Berlese) (Fig. 13.6) 72. Peritreme ending simply … 73 Peritreme ending with distal anasthomosis … 74 73. Dorsal setae set on strong tubercles and the receptaculum seminis is from oval to more rounded and has an irregular outline … Petrobia harti (Ewing) (Fig. 13.7) Dorsal setae not set on tubercles; receptaculum seminis tubular, smooth on one side and crested on the other … Petrobia tunisiae Manson (Fig. 13.9) 74. Peritreme ends in a terminal anasthomosis about 50 μm long and the receptaculum seminis is tubular and smooth … Petrobia latens (Müller) (Fig. 13.8) 75. Prodorsal setae sc1 short and less than half in length as setae v2 … Aponychus chiavegatoi Feres et Flechtmann (Fig. 13.10) Prodorsal setae sc1 double, about as long as setae v2 … Aponychus spinosus (Banks) (Fig. 13.11) 76. Fourth pair of dorsocentral setae (f1) set normally; opisthosoma without wart-like elevations between dorsocentral setae … 77 Fourth pair of dorsocentral setae (f1) set marginally; opisthosoma with wart-like elevations between dorsocentral setae … Eutetranychus cratis Baker et Pritchard … (Fig. 13.15) 77. Tibia II with ﬁve tactile setae … 78 Tibia II with six or seven tactile setae … 80 78. Most opisthosomal setae shorter than the distances between consecutive setae … 79

Key for Identiﬁcation

51

Opisthosomal setae longer than distances between consecutive setae … Eutetranychus pantopus (Berlese) (Fig. 13.18) 79. Dorsal body setae relatively slender and slightly enlarged distally; setae c1, d1, e1, f1 and h1 relatively long, reaching to bases of setae next behind; prodorsum with medio dorsal striae tortuous, forming crescentic ﬁgures … Eutetranychus pyri Attiah (Fig. 13.19) Dorsal body setae spatulate; dorsolateral setae c2, d2, e2 and f2, relatively long, considerably longer than dorsocentral c1, d1, e1, f1 and h1; stylophore rounded anteriorly … Eutetranychus citri Attiah (Fig. 13.14) 80. Genu III with two setae; tibia III with ﬁve setae … 81 Genu III with two setae; tibia III with six setae … 82 81. Tarsus I with solenidion of loosely associated setae less than half as long as proximal tactile seta; tarsus II with solenidion about two-thirds as long as proximal tactile setae … Eutetranychus eliei Gutierrez et Helle (Fig. 13.16) Tarsus I with solenidion of loosely associated setae about three-quarters as long as proximal tactile seta; tarsus II with solenidion about as long as proximal tactile seta or very little or shorter … Eutetranychus banksi (McGregor) (Fig. 13.13) 82. Coxa II with one seta … Eutetranychus orientalis (Klein) (Fig. 13.17) Coxa II with two setae … Eutetranychus africanus (Tucker) (Fig. 13.12) 83. Genital ﬂap striate transversally … 84 Genital ﬂap striate longitudinally … 90 84. Area anterior to genital ﬂap striate longitudinally … 85 Area anterior to genital ﬂap striate transversally … 87 85. Tarsus I of female with ﬁve tactile setae proximal to the duplex setae … 86 Tarsus I of female with four tactile setae proximal to the duplex setae … Eotetranychus yumensis (McGregor) (Fig. 13.29) 86. Tibia I and II of female with nine and eight tactile setae, respectively … Eotetranychus kankitus Ehara (Fig. 13.23) Tibia I and II of female with eight and eight tactile setae, respectively … Eotetranychus limoni Blommers et Gutierrez (Fig. 13.25) 87. Tibia I of female with nine tactile setae … 88 Tibia I of female with eight tactile setae … 89 88. Tibia II of female with eight tactile setae … Eotetranychus lewisi (McGregor) (Fig. 13.24) Tibia II of female with six tactile setae … Eotetranychus mandensis Manson (Fig. 13.26) 89. Tibia II of female with ﬁve tactile setae … Eotetranychus cendanai Rimando (Fig. 13.22) Tibia II of female with six tactile setae … Eotetranychus limonae Karuppuchamy et Mohanasundaram 90. Aedeagus short, direct posteriorly and ending bluntly … Eotetranychus pamelae Manson (Fig. 13.27) Aedeagus curved dorsally near the middle of the shaft, and distally direct caudoventrally with the tip deﬂexed … Eotetranychus sexmaculatus (Riley) (Fig. 13.28)

52

Chapter 6

91. Prodorsal vertical setae v2 slightly shorter than the scapular setae sc1; dorsal body setae strong, serrate and rod-like, most slightly shorter than distances between them … Mixonychus ziolanensis (Lo et Ho) Prodorsal vertical setae v2 shorter than the scapular setae sc1; all strongly setiform, pubescent, set on prominent tubercles, each slightly enlarged near the distal end, most longer than distances between them … Mixonychus ganjuis Qian, Yan et Ma 92. Dorsocentral setae h1 and dorsolateral setae f2 of equal length … 93 Dorsocentral setae h1 about a third of the length of dorsocentral setae f1; dorsolateral setae f2 about two-thirds of the length of dorsocentral setae f1 … Panonychus ulmi (Koch) (Fig. 13.36) 93. Aedeagus with distal sigmoid region about 1.5 times as long as the dorsal margin of the shaft … Panonychus citri (McGregor) (Fig. 13.34) Aedeagus with distal sigmoid region about twice as long as the dorsal margin of shaft … Panonychus elongatus Manson (Fig. 13.35) 94. Distal part of the aedeagus turns dorsally to form a sigmoid curve, slightly hooked at the tip, and the axis of the knob or barb is almost parallel to the shaft … Schizotetranychus hindustanicus (Hirst) (Fig. 13.38) Aedeagus not as above or unknown … 95 95. Tibia I of female with six tactile setae … Schizotetranychus baltazarae Rimando (Fig. 13.37) Tibia I of female with more tactile setae … 96 96. Tibia I of female with nine tactile setae … Schizotetranychus lechrius Rimando Tibia I of female with seven tactile setae … 97 97. Dorsal opisthosomal setae awl-shaped and almost nude, with c1, d1 and e1 slightly less than half as long as the distances between them longitudinally; opisthosomal setae c3, f2 and h1 longer than the oblate setae, slender, and serrate … Schizotetranychus spiculus Baker et Pritchard (Fig. 13.39) Dorsal body setae slightly serrate, with c1, d1 and e1 fusiform, broad at middle length and narrow and tapering distally; dorsal opisthosomal setae h1 not tapering and h2–3 slender and tapering … Schizotetranychus youngi Tseng 98. Aedeagus with distal knob … 100 Aedeagus without distal knob … 99 99. Aedeagus long, slender, tapering, and curved dorsally, and about eight times as long as its largest width at base … Tetranychus ﬁjiensis Hirst (Fig. 13.41) Aedeagus broadly bent dorsally to form a distal narrow part of subequal width, with caudal end truncate, and about three times as long as their largest width at base … Tetranychus taiwanicus Ehara 100. Knob of aedeagus with anterior and/or posterior rounded projections … 101 Knob of aedeagus without rounded projections and with projections of different size … 106 101. Knob of aedeagus indented dorsally … 102 Knob of aedeagus not indented dorsally … 103

Key for Identiﬁcation

53

102. Anterior projection of knob rounded and posterior forming an acute angle … Tetranychus gloveri Banks (Fig. 13.42) Anterior and posterior projections of knob rounded … Tetranychus neocaledonicus André (Fig. 13.46) 103. Anterior projection of knob rounded and posterior forming an acute angle … 104 Anterior projection of knob forming an acute angle and posterior rounded … Tetranychus ludeni Zacher (Fig. 13.44) 104. Axis of shaft parallel to that of knob … 105 Axis of shaft slightly inclined toward dorsal and forming a small acute angle with axis of knob … Tetranychus turkestani (Ugarov et Nikolski) (Fig. 13.42) 105. Axis of knob about as half as the largest width of shaft, and with anterior projections strong, rounded and protruse … Tetranychus kanzawai Kishida (Fig. 13.42) Axis of knob shorter as half as the largest width of shaft, and with anterior projections rounded but not strongly protruse … Tetranychus tumidus Banks (Fig. 13.48) 106. Anterior and posterior projections of knob small and subequal; axis of knob parallel to that of the shaft and about three times as long as the greatest width of shaft … Tetranychus urticae Koch (Figs 2.1, 2.2, 2.4, 2.5, 2.6, 13.42) Anterior and posterior projections of knob of different size, with the posterior larger … 107 107. Dorsal margin of knob convex … 108 Dorsal margin of knob slightly sigmoid and with hollow in the anterior part and with tip of posterior projection direct ventrally … Tetranychus paraguayensis Aranda (Fig. 13.42) 108. Posterior projection of knob curves downwards strongly … Tetranychus desertorum Banks (Fig. 13.40) Posterior projection of knob does not curve downwards strongly … 109 109. Axis of knob forming an angle with that of shaft … 110 Axis of knob parallel to that of shaft … 111 110. Axis of knob about two times longer than the width of shaft … Tetranychus paciﬁcus McGregor (Fig. 13.47) Axis of knob about as long as the width of shaft … Tetranychus salasi Baker et Pritchard 111. Empodia without mediodorsal spur … Tetranychus lambi Pritchard et Baker (Fig. 13.43) Empodia with a large mediodorsal spur … Tetranychus mexicanus (McGregor) (Fig. 13.45) 112. Dorsal setae shorter than distances between their bases … Oligonychus peruvianus (McGregor) (Fig. 13.33) Dorsal setae as long as or longer than the distances between their bases … 113 113. Tarsus I with three tactile setae proximal to duplex setae … Oligonychus coffeae (Nietner) (Fig. 13.31)

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Chapter 6

Tarsus I with four tactile setae proximal to the duplex setae … 114 114. Aedeagus curves dorsally and forms at the tip a large knob longer than dorsal margin; the anterior projection is small and acute and the posterior projection is long and curves downwards; the axis of the knob is parallel to the axis of the shaft and the dorsal margin of the knob is convex … Oligonychus biharensis (Hirst) (Fig. 13.30) Aedeagus narrows distally and curves dorsally approximately at a right angle; distally, presents a large knob with a small anterior projection and a long undulate posterior projection, and the tip is direct ventrally … Oligonychus gossypii (Zacher) (Fig. 13.32)

7

Phytoptidae Murray

7.1 INTRODUCTION The Phytoptidae Murray family has been recorded on citrus in India, with the species Phytoptus ﬁcivorus (Channabasavanna) (Prasad, 1974; Dhooria and Gupta, 1998; Dhooria et al., 2005) producing uncertain damage.

7.2 MORPHOLOGICAL CHARACTERISTICS AND SYSTEMATIC OUTLINE The general morphological characters are similar to those of Eriophyidae mites discussed in the next chapter. The Phytoptidae bear from one to ﬁve setae with anterior setae on the prodorsum, present (paired or unpaired vi and/or paired ve). The gnathosoma may be of various sizes, often large, and the chelicerae are straight or slightly and evenly curved; the palp is commonly short and truncate and enclosing the oral stylet. The legs bear the usual setae and often the solenidion I on tibia I. All empodia are undivided. The opisthosoma presents the usual setae, and the setae h1 are sometimes long. The female genital coverﬂap is devoid of ridges; the anterior female apodeme always extending forward a moderate distance, and the spermatecal tubes are commonly long, sometimes extending diagonally forward then recurving caudally. The Phytoptidae number ﬁve subfamilies (Prothricinae Amrine, Novophytoptinae Roivainen, Nalepellinae Roivainen, Phytoptinae Murray, Sierraphytoptinae Keifer) and various tribes (Lindquist and Amrine, 1996; Amrine et al., 2003). Phytoptus ﬁcivorus belongs to subfamily Phytoptinae. © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

55

56

Chapter 7

7.3 PHYTOPTINAE MURRAY The body is typically vermiform, and has opisthosomal annuli that are narrow and more or less equal dorsoventrally. Moreover, the seta sc points up if short and forward if long and the opisthosomal setae c1 are present (Lindquist and Amrine, 1996; Amrine et al., 2003).

7.3.1 Phytoptus Dujardin The femur and the genu are well distinct; the prodorsal shield is devoid of glands and the setae sc not are minute. The microtubercles are commonly distributed on the dorsal annuli (Amrine et al., 2003).

7.3.1.1 Phytoptus ﬁcivorus (Channabasavanna) Common name Unknown. Diagnostic characteristics FEMALE. The body is wormlike, 120–150 μm long, 25–40 μm thick and whitish coloured. The gnathosoma is 20–22 μm long, projecting forward and slightly bent down around the middle. The prodorsal shield is 28 μm wide and 21–25 μm long. The median line is bold, on basal third; the admedian lines are bold, complete, gradually diverging up to a third and converging at the rear shield margin; the ﬁrst submedian lines diverging gradually up to front of dorsal tubercles, passing just mesad of dorsal tubercles and converging to rear margin; a short diagonal, diverging fork to ﬁrst submedian just ahead of dorsal tubercles; the second submedian is short, not touching the apical shield margin, ending at two-thirds; the third submedian is almost complete, diverging to the rear; the sides of shield are granular, with a few indistinct, longitudinal lines, granular area extending up to second submedian lines. The dorsal tubercles are slightly ahead of rear shield margin, and 12 μm apart. The scapular setae are 12 μm long, and directed cephalad. The coxae I are ornamented with granules and short strokes. The empodium is ﬁverayed. The opisthosoma presents 58–62 annuli, uniformly microtuberculate, with microtubercles close-set, oval in outline, occupying almost the entire annulus width from the rear margin; the annuli beyond the setae f with obscure microtubercles dorsally and linear ventrally. The setae c2 are 17–19 μm long, on annulus 9; the setae d are 45–55 μm long, on annulus 20; the setae e are 7–11 μm long, on annulus 34; the setae f are 16–20 μm long, on annulus six from rear; the setae h2 are 50–70 μm long. The internal genitalia are 4–16 μm wide, and 8–9 μm long, the genital coverﬂap has eight bold longitudinal stripes; the seta is 8 μm long (Channabasavanna, 1966).

Phytoptidae Murray

57

MALE.

It is quite common, 120 μm long; with internal genitalia 13 μm wide, and seta 6 μm long (Channabasavanna, 1966). Geographical distribution India (Channabasavanna, 1966; Prasad, 1974; Dhooria and Gupta, 1998; Dhooria et al., 2005). Bio-ecology Phytoptus ﬁcivorus is a typical bud mite collected on Ficus infectoria Roxb. Several mites in different stages of development are to be found underneath the outer scales of buds. The species is generally found along with Aceria infectoriae Channabasavanna in the buds. On this host plant, the infested branches generally show paler, reduced leaves, which are also malformed (Channabasavanna, 1966). This eriophyoid is also recorded for citrus (Dhooria and Gupta, 1998; Dhooria et al., 2005) damaged with uncertainty and not requiring any control measures.

8

Eriophyidae Nalepa

8.1 INTRODUCTION Thirteen species of Eriophyidae have been recorded on citrus worldwide (Amrine and Stasny, 1994; Childers and Achor, 1999; Gerson, 2003; Dhooria et al., 2005), of which the pest status of almost six species is unknown (Gerson, 2003) (Table 8.1). In regions with a humid climate, outbreaks of the citrus rust mite, Phyllocoptruta oleivora, and the pink citrus rust mite, Aculops pelekassi, are particularly feared for the damage caused to fruit destined for fresh consumption (Ebeling, 1959; Jeppson et al., 1975; McCoy and Albrigo, 1975; McCoy, 1996a; Vacante and Bonsignore, 2009). The brown citrus rust mite, Tegolophus australis Keifer, and Cosella ﬂeschneri are considered to be of secondary importance in their areas of origin (Australia, India) and their infestation does not normally require any control measures (McCoy, 1996a). A. pelekassi, T. australis and C. ﬂeschneri may live together on the leaves and fruit of citrus with Ph. oleivora (Jeppson et al., 1975; Seki, 1981; Smith and Papacek, 1991; McCoy, 1996a). The citrus bud mite, Aceria sheldoni, prefers lemon (also orange in some regions of the world), and the importance of its attacks and its ecological role have been analysed (Hare et al., 1999; Vacante et al., 2007). In addition to A. sheldoni, Ph. oleivora and A. pelekassi, a number of rare vagrant species belonging to the Eriophyidae (Acaricalus sp., Tegolophus sp., Abacarus sp.) and Diptilomiopidae (Rhynacus sp.) families have been found on Citrus in Florida (Childers and Achor, 1999).

8.2 MORPHOLOGICAL CHARACTERISTICS AND SYSTEMATIC OUTLINE The Eriophyidae are very small (approximately 200 μm), worm-like or fusiform mites. Their body is made up of the gnathosoma, the prodorsum and an 58

© V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

Eriophyidae Nalepa

59

Table 8.1. Mites of the families Phytoptidae Murray, Eriophyidae Nalepa, Diptilomiopidae Keifer and Tarsonemidae (only the injurious species) Canestrini et Fanzago collected on citrus worldwide. Families and species Phytoptidae Phytoptus ﬁcivorus (Channabasavanna) Eriophyidae Aceria sheldoni (Ewing) Circaces citri Boczek Cosella ﬂeschneri (Keifer) Aculops pelekassi (Keifer)

Pest status

Distribution

U

India

Ma U U Ma, Me

Aculops suzhouensis Xin et Ding Aculus advens (Keifer) Paratetra murrayae Channabasavanna Tegolophus australis Keifer Tegolophus brunneus Flechtmann Calacarus citrifolii Keifer

U U U Mi Mi MI

Phyllocoptruta citri Soliman et Abou-Awad Phyllocoptruta oleivora (Ashmead) Phyllocoptruta paracitri Hong et Kuang Diptilomiopidae Diptilomiopus assamica Keifer Tarsonemidae Polyphagotarsonemus latus (Banks)

U Ma U

Worldwide Thailand India, Taiwan Croatia, Greece, Italy, Japan, Paraguay, Taiwan, Thailand, USA (Florida) China USA (California) India (Bangalore) Australia Brazil Angola, Kenya, Mozambique, Nigeria, South Africa, Zambia, Zimbabwe Egypt Worldwide China

U

Australia, India

Ma, Me

Worldwide

According to Gerson (2003), the pest status is indicated as either Ma (major), Me (medium), Mi (minor) or U (unknown). The references are indicated in the text.

extended and annular opisthosoma. The prodorsal shield is devoid of anterior setae (vi or ve), and with scapular setae sc present or absent (Figs 8.1, 8.2). Generally, the gnathosoma is prognathous and direct anteriorly, and possesses two stumpy palps, the infracapitulum and the chelicerae. The palps are parallel to the infracapitulum, each consisting of a base and three segments. The mouth is surrounded by a series of stylets derived from digits of the chelicerae, the oral stylet or labrum and the infracapitulum or rostrum; the cross-section of the latter is U-shaped and its walls form a membranous sheath in front and externally, with edges that overlap dorsally concealing from seven to nine stylets (Fig. 2.4). The latter consist of one pair of cheliceral shafts divided towards the apex in a dorsal digitus ﬁxus and ventral digitus mobilis from the labrum, and by one pair of auxiliary stylets, also known as inner infracapitular stylets. The basal segments of the chelicerae are close to each other and to the motivator, which provides a basal support for the chelicerae (Nuzzaci and Alberti,

60

Chapter 8 A

PRODORSUM

OPISTHOSOMA an

sc h2

dt

GNATHOSOMA

h1 e 3a

B

d

c2

f

1a 1b

2a C

aa

st s 3a

Fig. 8.1. Morphological features of Eriophyoid mites. (A) Lateral view of adult of Aculops pelekassi (Keifer) (from Vacante and Nucifora, 1985); (B) coxal and genital regions; (C) internal genitalia of female (from Keifer, 1959); aa, anterior apodeme; an, annuli; dt, dorsal tubercle; s, spermatheca ; sc, scapular setae; st, spermathecal tube. The notation of the different setae is explained in the text.

1996). The opisthosomal surface presents a series of transverse rings, or annuli, and never with subdorsal seta c1; the other setae are present; occasionally one or two pairs of setae (c2, d, e) and the setae h1 are sometimes absent. The coxal region bears setae and plate I may be devoid of setae 1b (Figs 8.1, 8.2). They have two pairs of legs; the chaetotaxy leg is complete and femoral setae I and II, genual seta II, tibial seta I and both tarsal setae ft’ and u’ of legs I and II are sometimes absent; tibia I without solenidion (Fig. 8.2). The tarsal empodium (Fig. 8.2) is sometimes thickened or of varied shape, rarely deeply divided. The

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female genital coverﬂap or epigynium is usually adorned, often with one or two rows of linear striae, sometimes with granules (Figs 8.1, 8.2); the female spermathecal tubes are always short, projecting either laterally or diagonally caudally (Figs 8.1, 8.2) (Lindquist, 1996; Lindquist and Amrine, 1996; Amrine et al., 2003). The Eriophyidae numbers six subfamilies (Eriophyinae Nalepa, Phyllocoptinae Nalepa, Nothopodinae Keifer, Aberoptinae Keifer, Cecidophyinae Keifer, Ashieldophyinae Mohanasundaram) (Lindquist and Amrine, 1996; Amrine et al., 2003) and 12 tribes (Amrine et al., 2003). Known species for citrus are variously distributed in the above-mentioned systematic groups. C. ﬂeschneri belongs to the Nothopodinae subfamily and Nothopodini Keifer tribe, Circaces citri to the Cecidophyinae subfamily and Colomerini Newkirk et Keifer tribe; A. sheldoni is attributed to the Eriophyinae subfamily and Aceriini Amrine et Stasny tribe; the Phyllocoptinae subfamily includes Calacarus citrifolii in the Calacarini Amrine et Stasny tribe, Ph. oleivora, Ph. citri Soliman et Abou-Awad and Ph. paracitri Hong et Kuang in the Phyllocoptini Nalepa tribe and A. pelekassi, A. suzhouensis Xin et Ding, Aculus advens (Keifer), T. australis, T. brunneus Flechtmann and Paratetra murrayae Channabasavanna in the Anthocoptini Amrine et Stasny tribe (Amrine and Stasny, 1994; Childers and Achor, 1999; Amrine et al., 2003; Gerson, 2003). Table 4.1 contains information on a number of websites, that contain images of living mites with their identifying colours and the damage they cause to citrus.

8.3 ERIOPHYINAE NALEPA The body is vermiform, with annuli subequal dorsoventrally, at least on the anterior half or on two-thirds of opisthosoma; the prodorsal shield is devoid of frontal lobe, or presents a slight projection over the base of the gnathosoma; if the frontal lobe is present it is narrow, basally ﬂexible and related to narrow annuli (Amrine et al., 2003).

8.3.1 Aceriini Amrine et Stasny The tubercles of the prodorsal shield are set on, or very near, the rear shield margin with transverse basal axes; the setae are directed to rear, and commonly divergently between them (Amrine et al., 2003).

8.3.1.1 Aceria Keifer The gnathosoma is curved downward and devoid of spine-like process on the anterior shield; the opisthosma is without a dorsal smooth area and is arched in cross-section and the posterior has continuous annuli and is subequal dorsoventrally; the tibia bears setae, and the coxal seta 1b is present (Amrine et al., 2003).

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Chapter 8 A

B

1a

sl

1b ml 2a al fgc dt

sc

3a taso

D

tase

tae C

aa

ta

E pts

ti ge

ags F

bfs

s

st

fe

tr

G H h2 h1 f

Fig. 8.2. Aceria sheldoni (Ewing). (A) Prodorsal shield; (B) coxal and genital regions; (C) internal genitalia; (D) leg I; (E) lateral opisthosoma; (F) empodium; (G) lateral view of prodorsum; (H) distal opisthosoma (from Keifer et al., 1982); aa, anterior apodeme; ags, antaxial genual setae; al, admedian line; bfs, basiventral femoral setae; dt, dorsal tubercle; fe, femur; fgc, female genital coverﬂap; ge, genu; ml, median line; pts, paraxial tibial setae; s, spermatheca; sc, scapular setae; sl, submedian lines; st, spermathecal tube; ta, tarsus; tae, tarsal empodium; tase, tarsal setae; taso, tarsal solenidion; ti, tibia; tr, trochanter. The setal notation is explained in the text.

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8.3.1.1.1 Aceria sheldoni (Ewing) (Fig. 8.2) Common name Citrus bud mite. Diagnostic characteristics body is vermiform, 170–180 μm long and from hyaline white to yellowish or pink–orange in colour. The prodorsal shield has a variety of markings, with indistinct or quite marked lines; in its most deﬁned condition the median line is broken and ends as a dart-shaped mark close to the rear shield margin. The admedian lines are complete; the submedian lines extend back towards the dorsal tubercles, in front of which they encounter transversally curved lines. The dorsal tubercles are 17 μm apart, and the scapular setae are 16 μm long and project backward. The opisthosoma has about 65–70 annuli. Each annulus is 2 μm wide, is strongly microturculate (they touch the rear rim of the annulus) and the last annuli are microstriate. The setae c2 are 19 μm long and about 8–10 annuli. The setae e are 30 μm long and about 23–25 annuli. The setae f are 16.5 μm and 5 or 6 annuli from the rear. The setae h2 are 40–45 μm long and the h1 setae are 3.5 μm long. The coxae I have a short sternal line and are marked with granules. The female genital coverﬂap bears 10–12 longitudinal ribs. The empodia are ﬁve-rayed (Keifer, 1938). FEMALE. The

Body worm-like, 180 μm long, with external features similar to female, except for the genital structures (Keifer, 1938).

MALE.

Geographical distribution Worldwide distribution (Jeppson et al., 1975). Bio-ecology The citrus bud mite lives on different species of citrus (lemon, orange, lime, grapefruit, tangerine, bergamot, citron, etc.), and of these prefers lemon for its large buds, which offer greater protection, as well as a number of varieties of orange (Valencia and Navel). It is common in coastal citrus-growing areas and is probably present in every lemon-growing area of the world with adequate relative humidity (RH) for its development. Their populations develop inside both wood and ﬂower buds or under the fruit rosette and shelter in ﬂowering buds as soon as fruit growth begins, and individuals sometimes shelter in the points of contact between two young lemon fruits, avoiding those sites most exposed to light (Jeppson et al., 1975). During the year, varying peaks in population are recorded from one season to another depending on various ecological factors (the growth of new vegetation, climatic trends, etc.) and the highest densities are observed in the buds in the coldest months with the lowest in spring–summer (Schwartz, 1975b; Vacante, 1986; Vacante et al., 2007). Population dynamics of the mite appears stable if compared with that of other mites (Searle, 1978). As soon as A. sheldoni populations become

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exposed during new bud growth, they move in search of new buds, spreading over the plant. Dispersion occurs throughout the year, reaching a peak during spring and at the beginning of summer, coinciding with the growth of new vegetation and increased air currents, and it is also helped forward by nursery material, insects and birds (Jeppson et al., 1975). The eriophyoid searches for new buds during the day, even with a low population density, and is more marked with high densities; reduced migration is observed in the morning or evening. The reproduction is parthenogenetic arrhenotokous. During its life the male deposits 25–100 spermatophores (2–15 per day), randomly distributing them on the plant; the latter, exposed to temperature extremes of 5°C or 38°C for an hour, do not fertilize the eggs (Sternlicht, 1970). A small percentage of males are found on lemon in the winter period and a high percentage in spring and autumn (Sternlicht and Goldenberg, 1971). The eggs hatch in 3–14 days and the length of a generation (egg to egg) is 12–33 days; hatching is optimal at 25°C and 98% RH but reduces with lower values (35–40%). The threshold of embryonic development is 9°C and 12.5°C for completion of the life cycle. Mean fertility is six eggs (four to eight) and increases up to 12 (5–19) if the female feeds from the bud during larval development. Survival of the population is inﬂuenced by extreme temperature values; at +30°C and −15°C, 50% of a population dies within 30 minutes (Sternlicht, 1970). In Australia, development from egg to adult takes 10 days in the ﬁeld in summer and approximately 15 days in autumn. Mortality begins at 43.3°C and is complete at 47.7°C (Boyce and Korsmeier, 1941). In Israel, it has been calculated that the citrus bud mite can produce 15 generations per year on fruit (Sternlicht, 1970). With regard to natural enemies, the Hirsutella thompsonii Fisher var. synematosa fungus (Clavicipitaceae) has been found to develop on the citrus bud mite (Searle, 1973; McCoy, 1981; Gomez and Nasca, 1983; McCoy, 1996a, b). Among the predatory mites, a number of stigmaeids have been recorded in different regions of the world, such as Agistemus africanus (Meyer et Ryke) (Searle, 1973; Searle and Meyer Smith, 1998), Agistemus collyerae Gonzalez (Vacante, 1986), Agistemus exsertus Gonzalez-Rodriguez (Sternlicht, 1969, 1970), Agistemus tranatalensis Meyer (Searle, 1973; Searle and Meyer Smith, 1998), Eupalopsellus brevipilis (Meyer et Rike) (Searle, 1973; Searle and Meyer Smith, 1998), Zetzellia graeciana Gonzalez (Vacante, 1986) and Zetzellia mali (Ewing) (Vacante, 1986). The Cheyletid mites include Cheletogenes ornatus (Canestrini et Fanzago) (Searle, 1973; Vacante, 1986; Searle and Meyer Smith, 1998) and Cheletomimus berlesei (Oudemans) (Sternlicht, 1969, 1970). Among the Phytoseiids mites are included Typhlodromips swirskii (Athias Henriot) (Sternlicht, 1969, 1970), Typhlodromus pyri Scheuten (Mijuskovic, 1973a) and other species. Symptomatology and damage Aceria sheldoni infests buds of all ages, including those dormant in old wood. Adults, immature stages and eggs are found under the axillary buds, at the base of the petioles near the buds, under the bracts, inside ﬂowering or fruiting buds and under the sepals. Trophic activity of the mite within the

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65

axillary leaf buds of lemon is the principal cause of damage and structural modiﬁcations. Attacked buds, recognizable by the presence of brownish or black bracts, often die and encourage multiple gemmation on the infested bud; the percentage of infested axillary buds varies with the age of the shoot. Mite populations progressively infest the occasional buds that emerge in reaction to the initial attack, resulting in a widespread emergence of buds, which impedes shoot growth and ﬂower development and negatively conditions vegetative growth. Even small amounts of damage reduce the vegetative integrity of the plant, and branches emerging from damaged buds may turn out to be shorter, larger and ﬂattened; in lemon trees rosetted growths occasionally derive from bud proliferation (Ebeling, 1959; Jeppson et al., 1975). In the northern hemisphere, south-facing trees are infested to a greater extent than north-facing ones, with levels of infestation between buds of different shoots usually greater than that recorded among buds of the same shoot (Walker et al., 1992). The opposite occurs in the southern hemisphere (Searle and Meyer Smith, 1998). Leaves originating from damaged buds have unusual shapes, with curled and twisted foils, divided and divergent at the tip; ﬂowers are misshapen and stunted, with deformed or aborted reproductive organs. Altered blossoms fall and may produce fruit that falls early or has a peculiar shape (Jeppson et al., 1975). Distortion of the fruit results from the attack on the embryonic tissue of the fruit in the bud and, once it emerges, the fruit is not affected by further attack from the mite (Walker et al., 1992). Depending on the entity of the attack and blossom damage, lemons may acquire a rounded rather than ellipsoid shape. The symptoms on oranges (cultivars Valencia and Navel) are the same as those observed on lemon except for a less evident deformity of the branches, leaves and fruit. Oranges may ﬂatten vertically, have crests and scars on the skin or small openings in the stylar cicatrix (Jeppson et al., 1975). Trophic activity of the citrus bud mite causes an increase in phenol levels in the bud tissue, with a simultaneous fall in auxin activity with changes in the activity of the same and the RNase (Ishaaya and Sternlicht, 1969, 1971). The extent of blossom and fruit abscission signiﬁcantly increases with the distortion caused by mite attack. Distortion of the fruit is signiﬁcantly linked to that of the blossoms from which they develop. It is probable that the increase in blossom and fruit abscission may depend on a decrease in auxin activity and other biochemical changes in the axillary buds infested by the mite (Phillips and Walker, 1997). A density of between one and three mites per bud results in the delayed development of the foliage (Ishaaya and Sternlicht, 1969). The level and economic importance of the attack depends on the age of the crop, the cultivated species, the variety and the extent of infestation. A high-density attack on young groves may seriously disrupt the development of lemon and orange plants (Di Martino, 1985; Searle and Meyer Smith, 1998). Swarms of mite may signiﬁcantly delay the vegetative development of Navel orange. The degree and quality of vegetative shoots determine the ﬂowering time and fructiﬁcation of the following season. The fall in lemon production may

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prove to be greater than that of Navel orange (Schwartz, 1972). Mite swarms also affect Navel orange in New Zealand (Harty et al., 2004). However, Vacante and Nucifora (1984) have established that in lemon groves in eastern Sicily, a level of 70% of buds infested by mites still guarantees a satisfactory gross marketable production. Findings in California have not provided signiﬁcant differences in terms of volume, quality or value in production between lemon groves treated once or twice a year with oils and untreated lemon groves, to the extent of recording greater production levels in untreated lots (Hare et al., 1999). Recently, Vacante et al. (2007) have recorded similar results in two lemon groves in southern Italy. Control Control may involve the use of several acaricides, insecticides and/or fungicides, resulting in the biological control being inadequate. BIOLOGICAL CONTROL. No classical biological or microbial control programmes have been initiated (McCoy, 1996a).

During the last 50 years, different principles have been used and a detailed account is given by Childers et al. (1996). Some of the most effective active ingredients have been found to be the banned and no longer available organochlorines (Jeppson et al., 1955, 1958; Attiah and Wahba, 1973; Mathew, 1973; Schwartz, 1975c; Brown and Jesser, 1981; Di Martino, 1985; Costilla et al., 1987; Lacasa et al., 1990); oil emulsion (Schwartz and Riekert, 1967; Mijuskovic, 1973a, b; Di Martino, 1985); sulfur, such as polysulphide sulfur or lime sulfur (Sternlicht, 1966; Mathew, 1973; Jeppson et al., 1975; Fourie, 1988); and petroleum oils (Boyce and Korsmeier, 1941; Boyce et al., 1942; Jeppson et al., 1975; Vacante, 1986; Atkins et al., 1987; Sale, 1988; Vacante, 1995; Salas and Macian, 2003; Vacante, 2009a), distributed in summer or in winter, according to their efﬁciency, the environmental conditions and the use permission of each single country. Normally, in nurseries and young crops, one treatment per season may be necessary. In adult crops with high infestation, two applications of acaricides are required in late winter or early spring and in summer. Instead, only one application is necessary in adult crops with a medium infestation (Searle and Meyer Smith, 1998). The treatments must be carried out when there is the highest density of the citrus bud mite population in the buds, before vegetative growth. Winter treatments with petroleum oils are also effective against the armoured scale insects and, in summer, it is necessary to use summer petroleum oils in those regions, like Sicily (Italy), where it is possible to apply a method for the summer production of lemons based on a limited stoppage of irrigation and after the ﬁrst irrigation and before vegetative growth (Di Martino, 1985; Vacante, 1986, 1995, 2009a). McCoy (1996a) reports that two applications per year are carried out in May and June and/or from September to November. In Greece, Papaioannou Souliotis et al. (1999) regard the periods from mid-May to mid-July and from mid-August to midSeptember as the most effective time for intervention.

CHEMICAL CONTROL.

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67

INTEGRATED PEST MANAGEMENT. Petroleum oils are selective for the beneﬁcials (Phytoseiids, Stigmaeids, etc.) and allow the realization of IPM programmes (Rosen, 1986; Vacante, 1986; McCoy, 1996a).

8.4 CECIDOPHYINAE KEIFER The female genital apodemes are shortened and bent up, and from a ventral view commonly appear as heavy transverse lines; the female coverﬂap present the ridges in two uneven ranks, and the female genitalia are appressed to coxae, separating coxae more than normal, and from a lateral view, commonly noticeably project from the venter; the coxae I narrowly connate at the centre line, and the sternal line is shortened; the coxae often present curved lines outlining tubercles of setae, particularly the setae 1a (Amrine et al., 2003).

8.4.1 Colomerini Newkirk et Keifer The Colomerini possess scapular tubercles and setae (Amrine et al., 2003).

8.4.1.1 Circaces Keifer The body is vermiform, and the frontal lobe may be absent. The scapular setae are projected to the rear. The opisthosoma has broad dorsal annuli with elongate microtubercles, except faint on about ﬁve or six annuli before the setae f, and narrow ventral annuli. The tibial seta is present (Amrine et al., 2003).

8.4.1.1.1 Circaces citri Boczek Common name Unknown. Diagnostic characteristics FEMALE. The body is fusiform, 118 μm long, 60 μm wide and translucent white in colour. Prodorsal shield 29 μm long, 51 μm wide and suboval, forming a pattern of seven ﬁelds; without frontal lobe over gnathosoma; dorsal tubercles 32 μm apart and situated on rear shield margin; scapular setae (sc) 4 μm long and directed to the rear and diverging. Opisthosoma with 28 dorsal annuli, smooth and evenly arched and 44 microtuberculate ventral annuli. The microtubercles are elongated. Setae c2 15 μm long, on ventral annulus six; setae d 40 μm long, on ventral annulus 16; setae e 5 μm long, on ventral annulus 28; setae f 11 μm long, on six annuli from rear; setae h1 absent. The genital coverﬂap granulate; proximal setae on coxisternum III 3a 6 μm long and 15 μm apart. The empodia are ﬁve-rayed (Boczek and Chandrapatya, 1996).

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Chapter 8 MALE. Unknown

(Boczek and Chandrapatya, 1996).

Geographical distribution Thailand (Boczek and Chandrapatya, 1996). Bio-ecology The mite has been collected on leaves of Citrus hystrix Dc. (Boczek and Chandrapatya, 1996), and its bio-ecology is unknown. Circaces citri causes rusting of the lower surface of leaves (Boczek and Chandrapatya, 1996), but its true pest status is unknown (Gerson, 2003).

8.5 NOTHOPODINAE KEIFER The tibiae are reduced or fused with tarsi, and devoid of seta. The tarsi lack spatulate projections. The coxae I are often fused across the centre line, and the sternal line is faint or absent. The empodia are small (Amrine et al., 2003).

8.5.1 Nothopodini Keifer The species of the tribe Nothopodini lack coxal seta 1b, and coxae and tibiae I are variable (Amrine et al., 2003).

8.5.1.1 Cosella Newkirk et Keifer The prodorsal shield bears a short frontal lobe over the gnathosoma, and the scapular setae are directed posterolaterally. The coxae I are fused with no sternal line, and more or less fused with the subcapitulum (Amrine et al., 2003).

8.5.1.1.1 Cosella ﬂeschneri (Keifer) (Fig. 8.3) Common name Unknown. Diagnostic characteristics body is spindleform, 130–140 μm long and probably yellowishwhite coloured. Prodorsal shield 32 μm long, 40 μm wide, anterior and lateral margins forming a semicircle with frontal lobe almost non-existent. Median line strong and present on rear three-quarters; admedian lines complete, slightly diverging, connected to median by two cross-lines; submedian

FEMALE. The

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69

A

G

C B

D

E F

Fig. 8.3. Cosella ﬂeschneri (Keifer). (A) Lateral view of adult; (B) prodorsal shield; (C) lateral view of prodorsum; (D) coxal and genital regions; (E) lateral opisthosoma; (F) internal genitalia of the female; (G) empodium (from Keifer, 1959b).

and lateral lines curving and forming a loose network. Dorsal tubercles 25 μm apart, well ahead of rear margin; scapular setae (sc) 7 μm long, projecting upwards and converging. Coxae I fused together and suboral plate, all coxae and this plate granular, tubercles I absent. Opisthosoma with about 45 dorsal annuli and 50 ventral annuli; microtubercles elongate, obscure or absent

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dorsally. Setae c2 22 μm long, on about ventral annulus ﬁve; setae d 30 μm long, on about ventral annulus 15; setae e 13 μm, on about ventral annulus 26; setae f 18 μm long, on ventral annulus seven from rear. Setae h1 absent. The genital coverﬂap is granular. The empodia are four-rayed (Keifer, 1959b). MALE.

Unknown (Keifer, 1959b).

Geographical distribution India (Keifer, 1959b; Jeppson et al., 1975; Seki, 1981; Smith and Papacek, 1991; McCoy, 1996a; Dhooria et al., 2005); Taiwan (Huang and Wang, 2003). Bio-ecology This species is vagrant, and has been reported on Citrus reticulata, lemon and orange (Navel and Valencia) (Keifer, 1959b), often associated with Ph. oleivora (Seki, 1981; Smith and Papacek, 1991; McCoy, 1996a) and A. pelekassi (Huang and Wang, 2003); its bio-ecology is unknown. Keifer (1959b) reports that the mite rusts the undersides of the leaves and occasionally attacks the fruit. The pest status of this species is unknown (Gerson, 2003) and its presence on citrus does not require any control measures (McCoy, 1996a).

8.6 PHYLLOCOPTINAE NALEPA The body is commonly fusiform. The prodorsal shield has a broad base and a rigid frontal lobe over the gnathosoma. The opisthosoma has broad and stout dorsal annuli, and narrow microtuberculate ventral annuli. When the frontal lobe is absent or only a slight one is present, the annuli differ dorsoventrally, at least in dorsal microtubercles. If the annuli are subequal and the frontal lobe is lacking, the annuli are as broad as the genital coverﬂap is long (Amrine et al., 2003).

8.6.1 Anthocoptini Amrine et Stasny The scapular setae are set on tubercles on or very near the rear shield margin, and directed towards the rear, commonly divergently. The scapular tubercles are subcylindrical, or the alignment of their bases is transverse to the body. The empodium is entire (Amrine et al., 2003).

8.6.1.1 Aculops Keifer The body is fusiform, often ﬂattened, with dorsal annuli wider than ventral annuli. The opisthosoma bears the setae e. The frontal lobe is acuminate, often ends in a sharp point and never has spinules under the front edge. The tibial setae and coxal setae 1b are also present. The empodia frequently have fewer than seven rays (Amrine et al., 2003).

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A

B

C

D

E

F

G

Fig. 8.4. Aculops pelekassi (Keifer). (A) Lateral view of adult; (B) prodorsal shield; (C) lateral view of prodorsum and gnathosoma; (D) coxal and genital regions; (E) lateral opisthosoma; (F) internal genitalia of the female; (G) empodium (from Keifer, 1959a).

8.6.1.1.1 Aculops pelekassi (Keifer) (Figs 8.1, 8.4) Common name Pink citrus rust mite, Japanese citrus mite. Diagnostic characteristics body is spindleform, 140–150 μm long and pinkish in colour. The gnathosoma is 24 μm long and projects diagonally downwards. The prodorsal shield is 36 μm long, 40 μm wide, subtriangular in anterior outline, with bulging sides, and with a moderately acute but terminally rounded frontal

FEMALE. The

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lobe over the gnathosoma. The shield pattern is a more or less obscure network of lines. The median lines are present on the rear half or two-thirds and receives transverse lines at the one-third and two-thirds positions. The line at one-third extends laterally around to rear shield margin. A subparallel line, below the transverse line at one-third, runs from the side of the frontal lobe to the shield margin above the coxae. The admedian lines are complete, extending from the sides of the frontal lobe to the rear margin, somewhat sinuate, and furthest apart at the rear; admedian also receive the cross-lines at half and two-thirds, which connect them with the median. The dorsal tubercles are 26 μm apart, on the rear margin; the scapular setae (sc) are 9 μm long, and diverging to the rear. The opisthosoma has about 36 dorsal annuli and 50 ventral annuli. The microtubercles on the dorsal annuli are faint, bead-like on the ventral annuli and rest on the ring margins. The setae c2 are 26 μm long, set on ventral annulus six; the setae d are 40 μm, set on ventral annulus 16; the setae e are 7 μm long, set on ventral annulus 30; the setae f are 21 μm long, set on ventral annulus six from the rear; the setae h1 are 2 μm long. The genital coverﬂap has two basal transverse lines with granules; the longitudinal ribs may have two forms, about either ten longitudinal ribs or transversely curved ribs. The empodia is four-rayed (Keifer, 1959a). MALE. Similar

to female, except for the genital structures (Keifer, 1959a).

Geographical distribution Croatia (Mijuskovic and Kosac, 1972); Greece (Keifer, 1959a); Italy (Costantino, 1962); Japan (Ehara, 1964a, b); Paraguay (Flechtmann and Aranda, 1970); Taiwan (Huang and Wang, 1997); Thailand (Keifer and Knorr, 1978) and the USA (Florida) (Denmark, 1962; Burditt et al., 1963). Bio-ecology Aculops pelekassi may be easily confused with Ph. oleivora. However, the former is pink and the latter yellow, and the egg of the former species is translucent white and laid scattered, while that of the latter species is opaque and laid in the hollows along the ribs. The populations of Ph. oleivora are also found on mature leaves and placed on the leaf surface while those of A. pelekassi prefer young leaves and are more numerous on the margins of the leaves and, for this reason, can easily disperse with the wind. Phyllocoptruta oleivora is not stimulated by light, while A. pelekassi reacts to the stimulation of ﬂuorescent light (Burditt et al., 1963). The pink citrus rust mite is vagrant on leaves, twigs and fruits of Citrus. The sexes are separated and the sex ratio changes seasonally, with 84% of females in October and 100% in December (Ashihara et al., 2004). The maximum oviposition (21.8 eggs/female) occurs at 25°C and stops when temperatures drop to 15°C. The time of development from egg to adult varies with temperature and ranges from 14.9 days at 20°C to 6.3 at 30°C (Seki, 1979); in the summer (with variables temperatures) it requires 6–7 days and in spring 8–13 days, while with a constant temperature of 35°C it takes 5 days, and

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18–19 days at 18°C. In Croatia, the eriophyoid produces 18–22 generations per year in the ﬁeld between May and October. The female lives for 5–32 days and fertility is equal to 55 (average 31.5) (Mijuskovic and Tomasevic, 1975). At 27°C, the female life span is 6.4 ± 11.22 days and the intrinsic rate of increase (rm) is 0.192 individuals/female/day. Mortality from egg to adult is equal to 45.59% at 27°C and to 32.25% at 17°C. The optimum conditions for the development of the species are between 22° and 27°C, with 75–77% RH. The thermal threshold for development is equal to 11°C, and at 10°C oviposition stops (Ebrahim, 2000). The developmental zero and effective heat units from egg to adult are 10.6°C and 119 degree-days, respectively (Seki, 1979). Although the species can reproduce in all seasons (Nucifora, 1969; Jeppson et al., 1975), it does not normally reproduce during the winter and shelters between the bracts of buds (Ciampolini and Rota, 1963; Seki, 1981), under the rosettes of the fruits, in the gorges of the plants and branches (Martelli, 1964), under the empty follicles of scale insects (Nucifora, 1967) and in the stylar scars of the fruits as adult female or different biological stages (Nucifora, 1967; Di Martino, 1985; Vacante, 1995). At the end of spring the mite begins to reproduce and completes the ﬁrst generation in the buds, while the subsequent generation develops on young sprouts, new leaves and fruits that it infests from the month of June (Ashihara et al., 2004). The density of the mite populations on the leaves increases rapidly in early summer, and reaches a peak in late July. Summer generations of the pink citrus rust mite move from the leaves to the immature fruits, on which they feed, reproduce and cause russet fruit. Adult males began to crawl from the leaves and fruits to the buds in early October (Seki, 1981). In Florida, A. pelekassi begins to feed on small fruit during April or May; in this region it has been observed that at the end of June or in early July, the populations of pink citrus rust mite rapidly decrease and the citrus rust mite, Ph. oleivora, begins to increase (Childers and Achor, 1999). In Japan, high densities of populations are related to higher spring temperatures and low precipitation in early summer; similar conditions occur in Florida (Childers and Achor, 1999). The prohibition of dithiocarbamate use, active against a number of eriophyoids (Johnston et al., 1957; Martelli and Di Martino, 1963; Swirski et al., 1967) has recently created a problem of this pest in some areas of eastern Sicily (Italy), where the lemon crop is important and these chemicals were widely used for the control of serious diseases such as the ‘Mal secco’, caused by the fungus Phoma tracheiphila (Vacante, 2009a; Vacante and Bonsignore, 2009). The dispersion of the pink citrus rust mite relies on weather conditions, the use of propagating infested material, the transport of infested fruit and farmers who involuntarily transport it from one environment to another. High population densities of the pest have been observed in the sunspot areas of fruit at midday, or afternoon as they dispers. This behaviour differs from that observed with Ph. oleivora, where avoidance of direct sunlight occurs. The pink citrus rust mite forms a golden yellow to pinkish dust patch on the fruit during the dispersal phase as a result of their large aggregations (Childers and Achor, 1999).

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Natural enemies include pathogens and predatory mites. Among these the fungus Hirsutella thompsonii (Clavicipitaceae) (McCoy, 1996b) has been recorded. In different countries of the world, different phytoseiid mites are collected, such as Amblyseius andersoni (Chant) (Kim and Paik, 1999), Amblyseius eharai Amitai et Swirskii (Ashihara et al., 2004), Galendromus occidentalis (Nesbitt) (Kim and Paik, 1999), Neoseiulus fallacis (Garman) (Kim and Paik, 1999), Neoseiulus longispinosus (Evans) (Ashihara et al., 2004), Neoseiulus womersleyi (Schicha) (Kim and Paik, 1999), Iphiseiodes quadripilis (Banks) (Villanueva and Childers, 2007) and T. pyri (Kim and Paik, 1999), some ascid mites such as Proctolaelaps pygmaeus (Müller) (Ehara, 1964a) and the stigmaeid mite Agistemus terminalis (Quayle) (Ashihara et al., 2004). Symptomatology and damage The species is potentially most harmful of Ph. oleivora and attacks all citrus, of which it seems to prefer orange, mandarin and clementine. The symptomatology can be confused with that of viral diseases. The damage to the host plant is similar to that of Ph. oleivora and affects leaves, green stems and fruit. In early spring, the mites move from overwinter sites and develop on mature leaves, where they feed on the lower surface. The feeding activity produces along the margins and the median ribbings of the developing leaves irregular brown blemishes, different degrees of distortion, curling under the margins, crinkling of tissues, burn and leaf dieback. Densities of 200 or more mites per leaf are destructive. Later the mites move into the fruit and back to the leaves (Childers et al., 2007). The green stems have corky peridermic alterations, which, depending on the attack, become rust and cracks. The damage is most common in young nurseries. In the ﬁeld, it is most rare to ﬁnd these symptoms, which could affect the fruit only (Burditt and Reed, 1963; Nucifora, 1967; Jeppson et al., 1975; Pennisi et al., 1975; Di Martino, 1985; Vacante, 1995, 2009a; Vacante and Bonsignore 2009). On the fruit of C. sinensis, each feeding adult of A. pelekassi produces about 20 punctures, each about 1 μm in diameter, per epidermal cell (10 μm long and 7 μm wide). The depth of penetration is about 20 μm and reaches the second and third layer of the fruit epidermis (Tagaki, 1981). Callus formation on the peel surface of Citrus unshiu Marcowich fruit occurs at early infestation in August when cells are dividing. Late attacks from September to October cause bronzing injury. Generally, injury during the middle stage of fruit growth results in both types of symptoms near the oil glands (Kato, 1977). Diameter, volume and weight of damaged fruit are less than those of undamaged fruit. The sugar content of juice is higher in damaged fruit, because the concentration of soluble solids increases through water loss caused by mite attack (Tono et al., 1978). At the beginning of the attack, the fruits lose their natural shine and are scattered with a thin and delicate brown powder consisting of thousands of individual mites. The lemon fruits have a green–bronze colour that later turns

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to silver and umbrine, more or less intense. The attack may be confused with that of the citrus silver mite, P. latus. Nevertheless, the eriophyoid infests the fruit at all stages of development, and the tarsonemid until reaching the size of a walnut; P. latus searches for fruit sites in the shade within the foliage, while A. pelekassi prefers those exposed to direct sunlight. Moreover, while the skin surface of ripe fruit infested by P. latus is cracked and desquamated, that of fruit infested by A. pelekassi doe not desquamate (Nucifora, 1967). Orange and tangerine fruit initially show an easing of brightness and the skin gradually becomes green–bronze, with tones more or less intense depending on the severity of the attack in the glandular concavities. The mandarins infested early take on at ripening a colour that becomes dark, and then may become olivaceous or fuliginous. The symptoms can be confused with that of the red and black ﬂat mite, B. phoenicis. However, in the case of tenuipalpids, the pericarp dimples have crater-shaped cracks while that of A. pelekassi shows generally longitudinal cracks that sometimes intersect each other (Nucifora, 1967). If the attack affects only a part of the fruit, spots are observed that are more or less numerous and dark orange in colour, delicate and with undeﬁned contours. They gradually converge with each other as they head towards the part of the fruit most ﬁrmly attached (Nucifora, 1967). Control Strict monitoring is fundamental for the control of the mite populations. Some factors should trigger the need to monitor the mite on the leaves and fruit, namely mild winter temperatures and/or low rainfall during the previous year, or a recent infestation of the mite in ﬁeld. Monitoring should then be initiated into early spring (Childers and Achor, 1999). Guidelines for control of the pink citrus rust mite in the citrus groves of the Mediterranean region suggested general inspection of the fruit on 10% of the plants sampled. The threshold should be kept within the limits of 2–3% of infested fruit (Cavalloro and Prota, 1983). Childers et al. (2007) proposed a sampling method and thresholds for processed fruit and fresh fruit (see Ph. oleivora). Hall et al. (2007) investigated binomial sampling to estimate rust mite densities on orange fruit, and concluded that binomial sampling for a general estimate of mite densities seemed to be a viable alternative to absolute counts of mites per sample for a grower using a low management threshold, such as two or three mites per sample. BIOLOGICAL CONTROL.

They are no known practical experiences of biological

control. CHEMICAL CONTROL. The use of sulfur such as lime sulfur, sulfur dust or wettable sulfur is well known (Mijuskovic, 1973a; Seki, 1981; Childers et al., 2007), and is also used in a mixture with dithiocarbamates (Ciampolini and Rota, 1963; Martelli and Di Martino, 1963; Nucifora, 1967; Di Martino and Benfatto, 1973; Tsuchiya, 2002); the use of sulfur, however, negatively inﬂuences the populations of natural enemies (McCoy, 1977a ; Smith and Papacek, 1991).

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Petroleum oils can also be used alone or with acaricides (Childers et al., 2007). Several acaricides, such as organochlorine, thiazolidine, diphenyl oxazoline and quinoline, are ineffective (Martelli and Di Martino, 1963; Costantino, 1968; Nucifora, 1969; Di Martino and Benfatto, 1973; Ashihara et al., 2004). The activity of some amidine and pyridazinone derivates (Tsuchiya, 2002) has recently been conﬁrmed. Tetronic acid derivate in combination with petroleum oils provides excellent initial and residual control of the mite (Childers et al., 2007), no less than spraying with horticultural oil for avermectins and pyridazinone derivates (Bell et al., 2005). Control is currently entrusted to the use of acaricides which, when correctly employed, permit the carrying out of IPM programmes. INTEGRATED PEST MANAGEMENT.

8.6.1.1.2 Aculops suzhouensis Xin et Ding Common name Unknown. Diagnostic characteristics FEMALE. Similar

to A. pelekassi, from which it may be distinguished by its more robust body, a short transverse line running clearly across the shield between the anterior part of the admedian line near the frontal lobe of process, the setae d especially long and reaching 82.8 μm, and the microtubercles of the last six ventral annuli on the opisthosoma, which are long and thin and regularly arranged (Xin and Ding, 1982).

MALE.

Unknown (Xin and Ding, 1982).

Geographical distribution China (Xin and Ding, 1982). Bio-ecology The species was reported as a pest on the leaf and fruit of orange in China (Xin and Ding, 1982), but the symptomatology and damage are unknown.

8.6.1.2 Aculus Keifer The prodorsal shield possesses a broad and rounded frontal lobe that is not acuminate. Some species bear from two to four small spines or spinules projecting forward from under the front edge. The tibial setae and the coxal setae 1b are present (Amrine et al., 2003).

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A E

B

F

C

D

G

Fig. 8.5. Aculus advens (Keifer). (A) Lateral view of adult; (B) ventral view of adult; (C) prodorsal shield; (D) lateral view of prodorsum and gnathososma; (E) genital region; (F) lateral opisthosoma; (G) empodium (from Keifer, 1938).

8.6.1.2.1 Aculus advens (Keifer) (Fig. 8.5) Common name Unknown. Diagnostic characteristics FEMALE. The body is about 130 μm long, 58–60 μm wide, rather short, robust, curved and cone-shaped. The gnathosoma is 19 μm long, projecting

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downwards. The prodorsal shield is broad, subtriangular, 42 μm long and 51 μm wide, with a slightly over hanging gnathosoma base, and a disc with some inconspicuous lines, sides smooth; the prodorsal tubercles are of moderate size, on the rear margin, 25 μm apart; the scapular setae (sc) are 19 μm long and project backwards. The sternum is indistinctly forked. The opisthosoma possesses 47–50 dorsal annuli and 57–58 ventral annuli, with dorsal annuli smooth and ventral annuli microtuberculate. The dorsal annuli are 3.5 μm wide. The setae c2 are 21 μm long, set on ventral annulus eight; the setae d are 29 μm long, set on ventral annulus 22; the setae e are about 20 μm long, set on ventral annulus 39; the setae f are 34 μm long, on ventral annulus ﬁve from rear; the setae h1 are 3 μm long, and setae h2 45 μm long. The genital coverﬂap is weakly marked with ﬁve or six furrows; glands rather small, short-stalked; setae 3a about 25 μm long. The empodia are four-rayed (Keifer, 1938). MALE. Unknown

(Keifer, 1938).

Geographical distribution USA (California) (Keifer, 1938). Bio-ecology The species was collected under the fruit button of Citrus limonia (Pomona lemons) in California; its bio-ecology is unknown. The mites were evidently overwintering and no damage to the lemon was observed (Keifer, 1938).

8.6.1.3 Paratetra Channabasavanna The scapular tubercles are set near the midline, with setae sc strongly recurved laterally. The opisthosoma has a mid-dorsal longitudinal furrow. The subdorsal ridges are distinct. The coxal setae 1b and the setae of genu II are present (Amrine et al., 2003).

8.6.1.3.1 Paratetra murrayae Channabasavanna Common name Unknown. Diagnostic characteristics FEMALE. The body is fusiform, yellowish-white, 135–145 μm long, 55 μm wide and 50 μm thick. The gnathosoma is 20 μm long, projecting downwards. The prodorsal shield is subtriangular, 46 μm wide and 25 μm long. The frontal lobe is thick, prominent obliquely curved downwards over the base of the

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gnathosoma, with a short, thin spine projection from the lower margin from the side view. The rear margin of the shield is obscure. The shield design has a pattern of ridges with a prominent, broad V-shaped ridge between the dorsal tubercles. The median lines are absent; the admedian lines are uniformly arched, diverging both anteriorly and posteriorly, arising at base of the frontal lobe and meeting the two ends of the V-ridge; the submedian lines rise almost at the apex of frontal lobe, diverging sharply to base of the lobe and continuing posteriorly to the base of the dorsal tubercles in the form of ridges. The sides of the shield have a few faint longitudinal lines of granules. The dorsal tubercles are large, 8 μm long, a little in front of the rear shield margin, inclined slightly backwards, forming an angle with the shield surface, 17 μm apart; the dorsal setae are 24 μm long, and directed upwards and laterally or posterolaterally. The coxae are almost smooth, short and stout; the coxae I moderately connatat. The empodium is ﬁverayed. The opisthosoma possesses 30–32 smooth, broad dorsal annuli, and about 70 microtuberculate ventral annuli. The ﬁrst three to four dorsal annuli are narrowed and not depressed, and the rest form a broad, longitudinal trough ﬂanked by a narrow ridge on either side; the last three to four dorsal annuli form complete rings. The microtubercles on the dorsal annuli are tiny and beadlike, set along the rear margins and those on the last 8–10 ventral annuli microstriate. The setae c2 are 18 μm long, on about ventral annulus 12; the setae d are 55 μm long, on about ventral annulus 26; the setae e are 10 μm long, on about ventral annulus 46; the setae f are 16 μm long, on ventral annulus ﬁve from rear; the setae h2 are 55 μm long; the setae h1 are hardly visible. The internal genitalia are 21 μm wide, 11 μm long and the coverﬂap has about 11–16 longitudinal stripes; the seta is 9 μm long (Channabasavanna, 1966). MALE. The body is 115 μm long, with internal genitalia 16 μm wide and seta 7 μm long (Channabasavanna, 1966).

Geographical distribution India (Bangalore) (Channabasavanna, 1966; Dhooria et al., 2005). Bio-ecology This Eriophyoid has been collected in India on Murraya koenigii (Linnaeus) Sprengel. The male is rare. It is a vagrant species on tender shoots, including leaves, and may cause some browning (Channabasavanna, 1966).

8.6.1.4 Tegolophus Keifer The opisthosoma has three ridges and the mid-dorsal ridges are stronger than the lateral ones, but fades caudally and does not end in a furrow. The coxal setae 1b are present (Amrine et al., 2003).

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A

C

D B E

F

G

Fig. 8.6. Tegolophus australis Keifer. (A) Dorsal view of adult; (B) lateral view of prodorsum, gnathosoma and opisthosoma; (C) coxal and genital regions; (D) lateral opisthosoma; (E) internal genitalia of female; (F) leg I; (G) empodium (from Keifer, 1964).

8.6.1.4.1 Tegolophus australis Keifer (Fig. 8.6) Common name Brown citrus rust mite. Diagnostic characteristics FEMALE. The body is fusiform, 180–190 μm long, 75 μm wide, rough and brownish in colour. The gnathosoma is 27 μm long. The prodorsal shield is 58 μm long, 65–70 μm wide, coarsely pitted, with the frontal lobe moderately acuminate over the gnathosoma. The shield surface is rough and largely obliterates the basic pattern, although admedian lines are faintly discernible, with an amphora-shaped median boss with handles. The rear edge of the shield between the dorsal tubercles is partially furrowed. The dorsal tubercles are small, 42 μm apart, the scapular setae (sc) are 5 μm long and do not

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reach across the ﬁrst dorsal annulus. The opisthosoma has about 22 dorsal annuli, broken laterally, and 50–55 ventral annuli, with more small and elongate ventral microtubercles. The ﬁrst dorsal annulus is rather broad and rough with longitudinal lines; the second dorsal annulus is narrow dorsally; the central ridge begins at the third dorsal annulus and fades on the 16th dorsal annulus. The setae c2 are 8 μm long, on about ventral annulus ﬁve; the setae d are 55 μm long, on about ventral annulus 16; the setae e are 4 μm long, on about ventral annulus 29; the setae f are 14 μm long, on about ventral annulus ﬁve from rear; the setae h1 are absent. The genital coverﬂap bears in the basal part a pair of subcircular areas containing short dashes, and about 161 longitudinal ribs. The empodia are four-rayed (Keifer, 1964). MALE. Unknown

(Keifer, 1964).

Geographical distribution Australia (Keifer, 1964). Bio-ecology The brown citrus rust mite has been found in Australia on orange (C. sinensis) (Keifer, 1964). In New South Wales, the mite infestations do not constitute a serious problem for orange and grapefruit, and the dynamics of the population is marked by peaks clashing with an increase and decrease in seasonal temperature and RH recorded in summer/autumn and spring/early summer, and with the stasis of reproduction of natural enemies. The decreases observed in winter are generally associated with lower values of RH and temperature, particularly in July when the average temperature is equal to 10°C. The density of mobile forms does not appear to be affected by climatic adversities (Beattie et al., 1991). Natural enemies include mites and insects. The former number the phytoseiids Amblyseius deleoni Muma et Denmark, Amblyseius lentiginosus Denmark et Schicha (Beattie and Gellatley, 1986), Euseius elinae (Schicha) and Euseius victoriensis (Smith and Papacek, 1991). The insects are the coccinellids Halmus chalybeus (Boisduval) and Serangium bicolor Blackburn (Beattie and Gellatley, 1986). Symptomatology and damage Tegolophus australis is a minor pest of orange and grapefruit. The symptomatology of the attack may be confused with that of Ph. oleivora; this latter species in New South Wales prefers to shelter inside the fruit and inner fruit surface, producing a slightly rough and greyish-brown injury, with deeper brown margins (Hely, 1968). The feeding injury of the brown citrus rust mite consists of smooth and brown blemishes on the fruit located on the outer part of the canopy. In smooth blemishes, the rind looks as though it has been polished with dark tan boot polish (Jeppson et al., 1975). 1According

to Flechtmann (1999), there are 18.

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Control It is possible to employ biological and chemicals means, but commonly the need for control is rare. With regard to biological control, the phytoseiid mite, E. victoriensis produces a good control of the pest and in spring a density of 10–20 individuals/100 leaves increases to just 100 individuals/100 leaves in mid-summer, bringing infested fruit below the economic threshold of 10%. As a supplementary food source the predatory mite uses the pollen of Rhodes grass, Chloris gayane Kunth, and the presence of this host plant for the production of pollen increases the phytoseiid population (Smith and Papacek, 1991).

BIOLOGICAL CONTROL.

CHEMICAL CONTROL. In Queensland, the use of avermectins mixed with petroleum oils controlled both Ph. oleivora and T. australis for 20 weeks in a trial conducted on Valencia late orange during January–June 1997; some organotine is also effective (Smith et al., 1998).

8.6.1.4.2 Tegolophus brunneus Flechtmann (Fig. 8.7) Common name New brown citrus rust mite. Diagnostic characteristics body is fusiform, 178 μm long, 63 μm wide and brownish-purple in colour. The prodorsal shield is 66 μm long, 54 μm wide, with the frontal lobe 10 μm long, declivitous in side view, with a ventral peripheral rim. The shield surface is rough, with a median longitudinal boss, anteriorly narrow and large oval body, as large as the distance between scapular setae. The outlines of this boss resemble admedian lines, anteriorly parallel and posteriorly arched outward. The dorsal tubercles are 41 μm apart, directing scapular setae (sc) divergently backwards, 6 μm long, and reaching the second tergite. The opisthosoma possesses a dorsomedian ridge, extending throughout the entire length, with 28 smooth dorsal annuli and 54 microtuberculate ventral annuli; the ventral annuli are anterior to setae d with small, rounded microtubercles and become elongate posteriorly; at level of setae e the microtubercles are thin, elongate, half as long as the annulus and at level of setae f as long as the annulus. The setae c2 are 18 μm long, set on ventral annulus three; the setae d are 53 μm long, set on ventral annulus 15; the setae e are 15 μm long, set on ventral annulus 32; the setae f are 18 μm long, set on ventral annulus 48 or ventral annulus ﬁve from rear; the setae h1 missing, and setae h2 55 μm long. The coxae are smooth. Genital coverﬂap with 14 longitudinal ribs on distal half, with irregular dashes at bases; setae 3a 11 μm long. The empodia are four-rayed (Flechtmann, 1999). FEMALE. The

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83

B

A

C

E D

F

G

Fig. 8.7. Tegolophus brunneus Flechtmann. (A) Dorsal view of adult; (B) lateral view of prodorsum and gnathosoma; (C) coxal and genital regions; (D) genital region; (E) lateral detail of opisthosomal annuli; (F) leg I; (G) empodium (from Flechtmann, 1999).

MALE. Smaller than the female, 135–145 μm long and 48–53 μm wide (Flechtmann, 1999).

Geographical distribution Brazil (Flechtmann, 1999). Bio-ecology Vagrant species collected on leaves and fruit of orange and mandarin in different areas of São Paulo State in Brazil. It produces rust on leaves but mainly on fruits (Flechtmann, 1999).

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8.6.2 Calacarini Amrine et Stasny The scapular setae are absent, and the scapular tubercles are present or absent. The empodium is entire (Amrine et al., 2003).

8.6.2.1 Calacarus Keifer The dorsum presents three ridges, and the mid-dorsal ridge is narrow and extends as far as the lateral ridges. The opisthosoma tapers evenly towards the rear. The tibial setae are present (Amrine et al., 2003).

8.6.2.1.1 Calacarus citrifolii Keifer (Fig. 8.8) Common name Citrus grey mite or citrus blotch mite. Diagnostic characteristics FEMALE. The body is fusiform, solid, 185–200 μm long, 70 μm thick and grey to purplish in colour. The gnathosoma is 40 μm long, and curved downwards. The prodorsal shield is 60 μm long, and 65 μm wide, with a broad and rounded frontal lobe. The shield pattern is a network, with admedian lines without small vertical dashes along them. The median line is present on the frontal lobe, and slightly indicated on the rear half of the shield. The admedian lines are strongly sinuate and undulating, arising from the sides of the frontal lobe, arching back to a lateral cross line at about one-third, converging centrally to a slight cross-line at about half, then curving out three-quarters, where they meet another lateral line and then fork. The inner fork lines of the admedians meet centrally a little ahead of the rear shield margin; the outer part of the bifurcation arches back and recurves outwardly at the rear margin. The submedian line, with branches, forms three large cells on the outside of the admedian. The scapular setae sc are missing. The opisthosoma possesses about 60 dorsal annuli and 65–70 ventral annuli; with ﬁve white longitudinal wax bands, gradually diminishing in distinctness caudally. The setae c2 are 40 μm long, on about ventral annulus 11; the setae d are 45 μm long, on about the ventral annulus 27; setae e 40 μm long, on about ventral annulus 46; the setae f are 25 μm long, on ventral annulus eight from rear; setae h1 absent. The coxae I narrowly meet centrally, and are ornamented with many curved lines. The genital coverﬂap presents many ﬁne longitudinal lines placed in two ranks. The empodia are ﬁve-rayed (Keifer, 1955).

MALE.

Unknown (Keifer, 1955).

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85

A

B

C

E

D

F G

Fig. 8.8. Calacarus citrifolii Keifer. (A) Lateral view of adult; (B) prodorsal shield; (C) coxal and genital regions; (D) lateral opisthosoma; (E) leg I; (F) empodium; (G) internal genitalia of the female (from Keifer, 1955).

Geographical distribution Angola (De Carvalho and Cardoso, 1972); South Africa (Keifer, 1955; Meyer Smith, 1996; Bedford, 1998b); Zimbabwe; Kenya; Mozambique; Zambia; and Nigeria (Meyer Smith, 1996; Bedford, 1998b). Bio-ecology The citrus grey mite is vagrant on young leaves, twigs and fruits of different species of citrus. On leaves, it normally prefers the upper surface, where immature and adult stages secrete a white wax that partly protects them and appears as a greyish-looking powder when the leaves are oblique to incident light. The mite is active throughout the year and has been reported on 16

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different dicotyledonous plants of 11 families in nine botanical orders, including Citrus (Van der Merwe and Coates, 1965; Meyer Smith, 1996). Of their host plants, the pest prefers Rhus pyroides Burch., the poinsettia and papaya, and can attack the fruit of the banana (Schwartz, 1971, in Bedford, 1998b); in Mozambique it has been found on the poinsettia and in Angola on Citrus, papaya and granadilla (Bedford, 1998b). Its reproduction is parthenogenetic. At 20° and 27°C, the citrus grey mite develops a generation (from egg to egg) in 13 and 7 days, respectively. The female begins to lay after 24 h and at 27°C lays on average three eggs per day for a total of 33 eggs in 10 days; at 30°C fertility is equal to 43 eggs (van der Merwe and Coates, 1965). The oviposition occurs in the hours of darkness, with little light or on cloudy days, mainly in depressions over the oil glands. At 27°C the eggs hatch in 5–6 days and at 20°C in 13 days. The mite produces several generations per year and there are no known wintering stages. Nymphal stages I and II live 1–2 and 3–5 days, respectively. The climatic conditions and the vegetative state of the crop can inﬂuence the population dynamics of the mite, whose seasonal ﬂuctuations are generally unpredictable (Dippenaar, 1958a). The density can reach 200 mites/leaf and the highest peaks are observed in the warmest months (van der Merwe and Coates, 1965). Symptomatology and damage The infestation of citrus grey mite induces on most host plants the appearance of leaf browning or necrotic yellow blotches, while on Citrus it transmits a serious disease, commonly known as ‘concentric ring blotch’, reported for the ﬁrst time in South Africa (Doidge, 1925) and extensively studied by Dippenaar (1958a, c). The disease is caused by the transmission of a toxic saliva of the mite, but it has been observed that high levels of attack are not always associated with disease (Rossouw and Smith, 1963), which affects fundamentally the young tissues and in strong growth involves the young leaves, shoots, branches and fruits, while the mature leaves and shoots can be exempt. It becomes particularly serious in nurseries and young plants, and the symptomatology is appreciated more frequently on the parts of the canopy exposed to the sun. The disease has a discontinuous presence over the years. Leaves and fruit show plots marked by ring blotch, hence the term ‘concentric ring blotch’, which signiﬁcantly varies in number, form, size and colour; other characteristics include a general chlorosis of the leaf surface, partial or general defoliation, dieback of shoots and shedding of fruit reaching maturity. On the leaves, there are two types of plots, one consisting of an atypical chlorotic blotch and concentrically marked and the other of a necrotic plot with or without concentric markings, surrounded by halo chlorotic tissue. The blotches on fruit are map-like outlines or circular, whitish-green in colour, with the damaged area of variable size and expanding with the development of the fruit until reaching a diameter of 50 mm and above. On shoots, concentric point-like, individual or multiple blotches are observed. The young lesions are chlorotic or green, clearer than the surrounding surface. From these areas on shoots and branches, necrosis, longitudinal cracks and abundant emissions

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of rubber may arise. A severe level of attack can lead to death of the entire shoot or the branch (Dippenaar, 1958a, b; Jeppson et al., 1975; Bedford, 1998b). In concentric alterations of the infested leaves, a high concentration of microorganisms such as Spiroplasma has been found (Kotze et al., 1987). On orange, the mite attack may initially go unnoticed (Grout and Schutte, 2004). Control Biological control is inadequate and mite infestations are treated with chemicals. In the nursery, it is possible to treat every 3 or 4 months, while in the ﬁeld a ﬁrst treatment can be carried out at petal fall and a second at a later stage (Dippenaar, 1958c). On Navel orange and lemon, young plants of Valencia orange and in the cultivar with an average age of over 10 years, only one treatment in autumn generally allows the implementation of IPM programmes of both the citrus bud mite and the citrus blotch mite (Bedford, 1968, 1998b). Among the chemicals used, in addition to older acaricides such as various organochlorines and some organohalogens, it has been reported that various formulations of sulfur (lime sulfur, sulfur dust and wettable sulfur) have been effective against both mobile forms and eggs of the mite (Dippenaar, 1958a, c; Krause et al., 1996; Meyer Smith, 1996; Bedford, 1998b). On banana, a good response to some organophosphate has been reported (Jones, 1979).

CHEMICAL CONTROL.

8.6.3 Phyllocoptini Nalepa The scapular setae are commonly set on well formed, often plicate, tubercles ahead of the rear margin of the shield, and the setae are directed forward, upward or centrally. When the tubercles and the setae are set near the rear of the shield margin, the tubercles are subcylindrical and bent forward or the alignment of their bases is longitudinal or diagonal to the body (Amrine et al., 2003).

8.6.3.1 Phyllocoptruta Keifer The prodorsal shield presents a distinct frontal lobe over the gnathosoma, and the opisthosoma possesses a wide mid-dorsal longitudinal furrow. The coxal setae 1b are present (Amrine et al., 2003).

8.6.3.1.1 Phyllocoptruta citri Soliman et Abou-Awad (Fig. 8.9) Common name Unknown.

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D

C

E F

Fig. 8.9. Phyllocoptruta citri Soliman et Abou Awad. (A) Dorsal view of adult; (B) prodorsal shield; (C) lateral view of prodorsum and gnathosoma; (D) coxal and genital regions; (E) lateral opisthosoma; (F) empodium (from Soliman and Abou-Awad, 1978).

Diagnostic characteristics FEMALE. The body is cone-shaped, ﬂattened dorsally, somewhat curved from a

lateral view, 117.5–167.5 μm long and yellow in colour. Prodorsal shield subtriangular, 42 μm long and 54 μm wide, with front reduced and overlying the gnathosoma. The shield pattern stands out sharply, giving the shield a very rough appearance from the side; the median line is incomplete, with two admedian lines longitudinally oblique, forked posteriorly, with lateral limb curved and meeting with that of the other line; anterolateral side cellshaped. The dorsal tubercles are 26 μm apart, well ahead of the rear margin, the scapular setae (sc) are 8 μm long and project anteriorly. All coxae are blank. The opisthosoma with two slight subdorsal longitudinal ridges fades out posteriorly, with 32 dorsal annuli and 58 ventral annuli. The dorsal annuli are much wider than ventral annuli, without microtubercles, and the ventral annuli are heavily tuberculated with micro-rounded tubercles. The setae c2 are about 20 μm long, on about ventral annulus six; the setae d 40 μm, set on ventral annulus 20; the setae e are 7 μm long, set on ventral annulus 37; the setae f are 13 μm long, on about ventral annulus 54; the setae h1 and h2 arise

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from a lobe behind the last annulus. The genital coverﬂap has 12 longitudinal scorelines, bowl-shaped, with proximal setae on coxisternum III (3a) 37 μm long, set on large tubercle, and surpassing setae d. The empodia are ﬁve-rayed (Soliman and Abou-Awad, 1978). MALE. Similar to female, 162.5 μm long and 57.5 μm wide (Soliman and AbouAwad, 1978).

Geographical distribution Egypt (Soliman and Abou-Awad, 1978). Bio-ecology The mite has been collected on orange and mandarin. It is a vagrant species on leaves and fruit of orange (Navel and Valencia) and mandarin, attacking in descending order and causing rusting symptoms (Soliman and AbouAwad, 1978). Its real pest status is unknown.

8.6.3.1.2 Phyllocoptruta oleivora (Ashmead) (Fig. 8.10) Common name Citrus rust mite. Diagnostic characteristics FEMALE. The body is fusiform, ﬂattened, 150–165 μm long, 53 μm wide and yellow to straw-coloured. Prodorsal shield with moderate frontal lobe, with a transverse furrow across the front edge. The admedian lines rise from each side of the frontal lobe, and curve out to the area just past the lobe base; from there they curve inward and meet a slight cross-line at one-third, then curve slightly outwards past the inner side of the dorsal tubercles at two-thirds, and end in transverse arcs ahead of the rear shield margin. Dorsal tubercles moderately large, well ahead of the rear shield margin, 23 μm apart, scapular setae (sc) 9 μm long and projecting dorsoanteriorly. The opisthosoma has a broad longitudinal trough delimited by two longitudinal ridges, with about 31 dorsal annuli and 58 ventral annuli. Microtubercles are present only on ventral annuli, and each dorsal annulus covers two or three ventral annuli. Setae c2 25 μm long, on about ventral annulus ﬁve; setae d 35 μm long, on ventral annulus 17; setae e 8 μm long, on about ventral annulus 33; setae f about 15 μm long, on ventral annulus ﬁve from rear; setae h1 very short and setae h2 nearly 40 μm long. The genital coverﬂap is granular at the base, with a median longitudinal line, and apically with 14–16 ribs. The empodia are ﬁve-rayed (Keifer, 1938). MALE. Body fusiform, 135 μm long, 54 μm wide, with same external features as the female except the genital structures (Keifer, 1938).

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A

C D

B

E

F

G

Fig. 8.10. Phyllocoptruta oleivora (Ashmead). (A) Dorsal view of adult; (B) lateral view of adult; (C) empodium; (D) internal genitalia of the female; (E) lateral opisthosoma; (F) coxal and genital regions; (G) legs I and II (from Keifer, 1982).

Geographical distribution Worldwide distribution (Jeppson et al., 1975). Bio-ecology The citrus rust mite is a serious pest of citrus in many areas of the world, injuring by its feeding the quality of fruit (Jeppson et al., 1975). It is a vagrant species, living on leaves and fruit of all citrus species and varieties, and reproducing by arrhenotokous parthenogenesis (Swirski and Amitai, 1958). The ovoviviparity of the species has been well observed (Hall, 1967). The female lays eggs in the depressions of the pericarp of the fruits or leaves. At

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32°C embryonic development is completed in 3 days and the Ist nymphal stage in 1.8 days; at 22°C the egg develops in 5.5 days, Ist nymphal stage in 4.3 and 2nd in 6.4 days. In summer a generation develops in 7–10 days and in winter in about 14 days. The female lives less than 20 days and lays a total of 20–30 eggs (Yothers and Mason, 1930; Ebeling, 1959). With constant temperatures between 14° and 31°C the development threshold is 29°C (Allen et al., 1995). At 32°C and with 88% RH the lifespan of the female is equal to 3.2 ± 0.673 days, with an intrinsic rate of increase (rm) of 0.350 individuals/ female/day. The thermal threshold of development is equal to 11°C and oviposition ends at 10°C. Mortality from egg to adult is equal to 53.06% at 17°C and decreases to 25.81% and 26% at 30° and 32°C, respectively. The optimum temperature for development is between 30° and 32°C (Ebrahim, 2000). The development from egg to adult requires 330.07 degree-days, with a temperature of between 23° and 32°C and a photoperiod of 12/12 h of light/dark (Li et al., 1989). The species is active in all seasons and can develop a large number of generations during the year, as in Suriname where approximately 40 have been observed (Van Brussel, 1975) and in Israel up to 28 or more (Swirski and Amitai, 1958). Although the mite develops well with high values of RH and particularly after rain (Pratt, 1957; Dean, 1959), in South Africa it has been observed that with high values of RH the mortality of nymphs increases and oviposition decreases (Van Brussel, 1975). In Florida, the mite is active all year and its population increases in the spring and during the summer (Grifﬁths and Thompson, 1957). The density of the populations increases in May–July and then declines in late August, but can increase again in late October or early November (Childers et al., 2007). The behaviour of the mite with exposure to direct sunlight and with the different parts of the canopy is unclear. Indeed, Van Brussel (1975) reports that this eriophyoid does not tolerate direct sunlight and usually prefers the lower surface of leaves and areas in the shade of the fruit; on young plants it can be found on the lower surface of the leaves and on the upper surface of trees older than 1 year. According to McCoy (1979), populations migrate to the new shoots and settle on the lower surface of the leaves before moving to reproduce towards the upper surface and then on fruit. Allen and McCoy (1979) analysed the distribution of populations on individual fruit and over the canopy, and report that the citrus rust mite refuses direct exposure to sunlight. Populations are generally more abundant on fruit and leaves on the outer margins of the tree canopy. The northern foot of the tree canopy is preferred, and the least favourable conditions for population increase occur in the southern top of the tree canopy (Childers et al., 2007). In Israel, it has been observed that the mite prefers the lower surface of the leaves and the outside of the canopy rather than internally, and prefers the fruit to leaves; it also develops better in citrus groves with a high density of transplantation, attacks the fruit of high branches and not of the low branches; and also attacks lemons in spite of the presence of other species and varieties of citrus (Swirski, 1962). At night and on cloudy days, pest populations pour out on both leaf surfaces, tend to disperse across the foliage and are easily observed during

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Table 8.2.

Phyllocoptruta oleivora (Ashmead) natural enemies (fungi, molluscs and insects).

Families and species Clavicipitaceae Beauveria bassiana (Balsamo) Vuillemin Hirsutella thompsonii Fisher Hirsutella kirchneri (Rostrup) Minter et al. Hirsutella nodulosa Petch Exobasidiomycetidae Acaromyces ingoldii Boekhout et al. Meira geulakonigii Boekhout et al. Meira argovae Boekhout et al. Mycosphaerellaceae Mycosphaerella citri Whiteside Orthalicidae Drymaeus dormani (Binney) Cecidomyiidae Cecidomyiids Feltiella sp. Lestodiplosis sp. Coccinellidae Pentilia castanea Mulsant Coniopterygidae Semidalis vicina Hagen Coniopteryx westwoodi Fitch Psychidae Cryptothelea gloverii (Packard) Thripidae Leptothrips mali (Fitch)

Reference(s)

Alves et al., 2005a Spears and Yothers, 1924; McCoy and Kanavel, 1969; McCoy, 1996a Cabrera and Dominguez, 1987 Cabrera and Dominguez, 1987 Paz et al., 2009a Sztejnberg et al., 2004; Paz et al., 2009a Paz et al., 2009a Hernandez and Escobio, 1988 Bledsoe and Minnick, 1982 Muma et al., 1975 Villanueva et al., 2006 Villanueva et al., 2006 Van Brussel, 1975 Muma, 1967 Muma, 1967 Villanueva et al., 2005 Grifﬁths and Thompson, 1957

the sunniest days, with the greatest levels of attack on the outside of the canopy (Jeppson et al., 1975). The dispersion of the mite from one environment to another is facilitated by the wind, with shifts of up to 135 m away and with higher peaks between mid-September and mid-October (Bergh and McCoy, 1997). Several natural enemies develop on the citrus rust mite, including fungi, molluscs, insects and acari. Insects include a number of coniopterygids, cecidomyiids, thrips and coccinellids (Table 8.2), and mites include a number of stigmaeids, cheyletids and tydeids, and several phytoseiids (Table 8.3). Symptomatology and damage The citrus rust mite is a serious problem in all citrus areas of the world, with the exception of a number of regions such as New South Wales (Australia), where it is not always harmful to orange (Beattie et al., 1991), or Italy, where it was not found until recently.

Eriophyidae Nalepa Table 8.3.

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Phyllocoptruta oleivora (Ashmead) natural enemies (mites).

Families and species Cheyletidae Cheletogenes ornatus (Canestrini et Fanzago) Hemicheyletia wellsi (Baker) Phytoseiidae Amblyseius herbicolus (Chant) Euseius citri (van der Merwe et Ryke) Euseius citrifolius Denmark et Muma Euseius elinae (Schicha) Euseius mesembrinus (Dean) Euseius stipulatus (Athias Henriot) Euseius victoriensis (Womersley) Euseius sp. Galendromus helveolus (Chant) Neoseiulus cucumeris (Oudemans) Iphiseiodes zuluagai Denmark et Muma Iphiseoides quadripilis (Banks) Iphiseius degenerans (Berlese) Typhlodromalus peregrinus (Muma) Typhlodromips swirskii (Athias Henriot) Typhlodromus athiasae Porath et Swirski Typhlodromus rickeri Chant Stigmaeidae Agistemus cyprius Gonzalez Agistemus ﬂoridanus Gonzalez Agistemus exsertus Gonzalez Rodriguez Tydeidae Tydeus sp.

Reference(s)

Rezk and Gadelhak, 1996 Feres et al., 2005 Argov, 1993; Matioli et al., 1998; Argov et al., 2002 Schwartz and Meyer Smith, 1998 Gravena et al., 1993 Argov, 1993; Argov et al., 2002 Abou-Setta and Childers, 1989; Flores et al., 1996b McMurtry et al., 1992; Argov et al., 2002 Smith and Papacek, 1991; Argov, 1993; Argov et al., 2002 Matioli et al., 1998 Caceres and Childers, 1991 Zhang et al., 2003 Gravena et al., 1993; Matioli et al., 1998 Villanueva and Childers, 2007 Harpaz and Rosen, 1971; Palevsky et al., 2003 Peña, 1992 Harpaz and Rosen, 1971; Palevsky et al., 2003 Harpaz and Rosen, 1971 Argov et al., 2002 Palevsky et al., 2003 Muma and Selhime, 1971 Rezk and Gadelhak, 1996 Chen et al., 1988

The eriophyoid feeds on leaves and fruit of all citrus species and prefers lemon. On the upper leaf, surface damage involves the epidermal cells and appears as slightly brownish patches to the back. In severe cases, the cuticle loses its lustre, takes on a bronze-like colour and may shows patches of cells yellowish in russet areas de-greened by ethylene release during the attack (Yothers and Mason, 1930; McCoy and Albrigo, 1975). On Valencia orange and Marsh grapefruit, because of the wound, the leaf surfaces presents a periderm, consisting on the leaves of Sunburst tangerine of three layers (phellem, phelloderm and phellogen) and the leaves of other cultivars not from the phellem; the phelloderm consists of up to seven layers of cells, but

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only two to three in other cultivars. The lower leaf surface may present a mesophyll collapse that appears ﬁrst as yellow-degreened patches and later as necrotic spots (Albrigo and McCoy, 1974). The probability of defoliation increases with the amount of damage, and the relationship between the size of the area involved and that of the leaves not shows an apparent effect on leaf abscission (McCoy, 1976). The mite attacks early fruits of the size of a pea, and symptoms appear until they reach the size of a walnut. At harvest, fruits infested in the early, mid- or late stage of growth and development show different levels of injury and symptoms indicated as ‘sharkskin’, ‘russet’ or ‘bronzing’. The stylar feeding of the mite on fruits causes the collapse of the epidermic cells, probably as a result of saliva inoculation. Since this is a vagrant species characterized by continuous probing of the leaf surface and fruit, the destruction of a large quantity of cells that have the appearance of brownish-black patches is produced (Albrigo and McCoy, 1974). Affected tissues emit ethylene, which can stimulate an early degreening of the leaves and fruits (McCoy and Albrigo, 1975). The discoloration of the fruit surface appears to be associated with the formation of lignin and likely oxidation of cytoplasmatic substances. According to variety and maturity of fruits, the injury may vary; if it concerns the growth phase before fruit maturity, it leads to a breaking up of the dead epidermis and the formation of a peridermal wound beneath the epidermis. Cracks in the epidermis appear but the layer epidermis is slotted in patches. The cracks form a rough texture that can be polished, while the oxidized cell contents give the fruit a brownish-black appearance. This earlier injury phase is deﬁned as a ‘russet condition’. On grapefruit, lemon, lime and orange, occasionally the ‘russet condition’ assumes a sharkskin appearance (Albrigo and McCoy, 1974). This form of ‘early russet’ derives from the sloughing of the dead epidermis that exposes a wound periderm. The injury on the ripe fruits differs from ‘early russet’. The epidermal cells die and have a very brownish-black colour; there are small cracks on the epidermal layer and periderm wounds. These fruits are polished for the presence of the natural cutin and the wax layer, and present a condition named as ‘bronzing’ (Albrigo and McCoy, 1974). Severe attacks produce a reduced size of the fruits (Yothers and Mason, 1930; Schwartz, 1975a), fruit water loss (McCoy et al., 1976a; Allen, 1979), fruit drop (Allen, 1978) and alterations in juice quality (McCoy, 1976). Although the aesthetic damage to fruits involves commercial damage, the content in total soluble solids does not appear to be vitiated, and production can be addressed without drawbacks for industrial use (Chandramani et al., 2004). Examination of the frequency distribution of the damaged fruits showed that those sites in the northern quadrant present a higher average area of damage, followed by the eastern, southern and western quadrants. The distribution of frequency of the damaged surface of fruit changes with the level of average damage. Indeed, with a low average damage, most of the fruits present no damage, whereas with increased average damage to the fruits, the proportion of fruit without damage decreases (Yang et al., 1995). On orange, a linear positive correlation between the cumulative population and the

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percentage of damaged surface of the fruit was found, showing that a single mite damages 0.048% of the surface of a fruit every day (Sanchez Salas and Padron Chavez, 1985). The increase in damage to fruits can lead to a small increase in fruit drop, with a slight negative relationship between the size of fruits and the amount of damage (Yang et al., 1994). Control Control of the citrus rust mite is basically entrusted to the chemicals applied in a context of IPM programmes (McCoy, 1996a). The choice of strategy depends on the productive trend. Indeed, the control of production destined for fresh consumption normally requires more treatments than that destined for juice production, making the cosmetic appearance a priority in fresh consumption, while in the fruit processed for juice the development and abscission of the fruits are not affected until 50–75% of the fruit surface is damaged (Allen and Stamper, 1979). In Florida, production destined for the fresh market is treated with pesticides three to four times a year – in April, June, August and October, while production designated for processing is treated up to twice, one with petroleum oil and one with acaricides (McCoy, 1985; Browning, 1992). With regard to the economic threshold, Allen (1981) writes that there is no single threshold value, with a range of damage thresholds equivalent to the cost of the treatment, which depends on value of production, time of the pest attack, harvest and temperature. Nascimento et al. (1982) concluded that populations of the citrus rust mite can be accurately sampled by taking ﬁve leaves per tree from 1.5% of the trees in the grove and counting the number of mites in an area of 2 cm2 on the upper surface of each leaf, 2 cm from the leaf base. Oliveira et al. (1982) established that population levels were approximately the same over the whole tree, and therefore the samples could be taken from any cardinal point. At any level of infestation, the results obtained by counting the mites present on a predetermined limited area of a leaf (about 4.9 cm2 on the lower surface of the leaf at about 2 cm from the petiole) were as accurate as counting the numbers on the whole of both surfaces of the entire leaf. A leaf-brushing machine provided fairly accurate results when the infestation level was low, but not when it was high. If a precise determination of the number of mites is required, the ‘limited area’ method is to be preferred, but any of the established methods can be used for comparative studies. Manzur (1989) reports that Ph. oleivora showed a negative binomial distribution on grapefruit tree leaves, and the economic thresholds of pest density were set as: 0.5 mites/leaf demanding no insecticide treatment, and greater than 2 mites/leaf requiring treatment. Rogers et al. (1994) applies the modiﬁed Horsfall–Barratt system for estimating citrus rust mite populations in an integrated crop management programme. In South Africa, the ﬁrst discovery of the mites on leaves allow for treatments (Schwartz and Meyer Smith, 1998). Hall et al. (2005) have investigated the consequence of reducing sample size on the accuracy and precision of estimates of citrus rust mite. Hall et al. (2007) investigated binomial

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sampling to estimate rust mite densities on orange fruit; and concluded that a binomial sampling for a general estimate of mites densities seemed to be a viable alternative to absolute counts of mites per sample for a grower using a low management threshold, such as two or three mites per sample. In Florida, Childers et al. (2007) suggested one sampling method based on rust mite density (rust mites/cm2), in late spring or summer, in processed fruit trees selected at random and within uniformly distributed areas throughout a 10–40-acre block representing a single variety with uniform horticultural practices. Avoid sampling adjacent trees. Fruit should be sampled at random, representing the four quadrants of the tree and taken midway in the canopy. One fruit surface area should be examined midway between the sun and shade areas. The number of rust mites per cm2 should be recorded and averaged for 10 acres (4 ha), represented by 20 trees with four fruit per tree, or 80 readings for 10 acres. Six rust mites/cm2 would be a planning threshold where pesticide treatment may be required within 10–14 days; ten rust mites/ cm2 would be an action threshold where treatment would be required as soon as possible. In fresh fruit, it is necessary to monitor mite populations every 10–14 days, and an average of 2 mites/cm2 as an action threshold. The use of microorganisms has been investigated for a long time. H. thompsonii Fisher (Clavicipitaceae) was reported for the ﬁrst time in association with Ph. oleivora (Spears and Yothers, 1924) and described as a new species by Fisher (1950); the pathogen gives a clear pathogenicity to the citrus rust mite (McCoy and Kanavel, 1969) and is effective with values of RH that can guarantee the development of propagules on leaves and fruit (McCoy, 1996a). Hirsutella kirchneri (Rostrup) Minter, Brady et Hall and Hirsutella nodulosa Petch (Cabrera and Dominguez, 1987), Beauveria bassiana (Balsamo) Vuillemin (Alves et al., 2005a) and the Mycosphaerellaceae Mycosphaerella citri Whiteside (Hernandez and Escobio, 1988) have also been examined. The Exobasidiomycetidae Meira geulakonigii Boekhout et al.2 produces a control of 100% of Ph. oleivora, and also develops at the expense of a number of tetranychid mites such as P. citri and the pathogenic fungus of cucumber, Sphaerotheca fusca (Fr.) Blumer (Sztejnberg et al., 2004). Unfortunately, H. thompsonii is active, with a high density of attack and with the presence of damage to the plant (McCoy, 1981). The action of a protein (irsutellina A) extracted from the substrate of a culture of H. thompsonii has recently been observed capable of inhibiting mite fecundity (Omoto and McCoy, 1998). The effectiveness of the fungus has been veriﬁed in several citrus areas throughout the world, including Florida (McCoy et al., 1971; McCoy and Selhime, 1977; McCoy and Couch, 1982), China (Yen, 1974; Chen and Chen, 1986; Chen et al., 1987), Suriname (Van Brussel, 1975), Cuba (Cabrera et al., 1981), Argentina (Gomez and Nasca, 1983), Brazil (Almeida et al., 1981; Santos da Silva et al., 1981), Thailand (Saowanit et al., 1999) and Iran (Aghajanzadeh et al., 2006). The pathogen

BIOLOGICAL CONTROL.

2These

last three species have been described by Boekhout et al. in 2003, and complete indications are reported in the bibliography (Boekhout et al., 2003).

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gives results comparable with those of chemical control but, for a number of reasons, including the impossibility of its continued and rapid development in the ﬁeld, its mass production has been stopped (McCoy, 1996b). Recently, Paz et al. (2009a) have evaluated in the laboratory the role of three Exobasidiomycetidae (Meira argovae Boekhout et al., Meira geulakonigii Boekhout et al. and Acaromyces ingoldii Boekhout et al.) in the control of Ph. oleivora, observing that after 14 days the three pathogens guarantee mortality rates of 87.1, 91.1 and 89.1%, respectively. These good results have encouraged veriﬁcation in the ﬁeld of the action of M. geulakonigii on grapefruit infested from Ph. olievora. It has been observed that one application of the pathogen and monthly applications produced 87.4 and 87.2% of non-damaged fruits, respectively, while in the control the percentage of damaged fruits was 77.2%. The fungus is a beneﬁcial endophyte of grapefruits that colonizes the fruit’s peel, protecting it from the attack of Ph. oleivora (Paz et al., 2009b). Unfortunately, at present no practical indications are known for the use of these pathogens. With regard to predators, Schicha (1987) and Smith and Papacek (1991) report that in Australia the phytoseiid E. victoriensis has controlled an attack of the Eriophyoid to fruit less than 5%. In Israel the phytoseiids Amblyseius herbicolus (Chant), E. victoriensis, E. elinae, Euseius stipulatus (Athias Henriot) and Typhlodromus rickeri Chant have been introduced from different geographical areas (Argov, 1993; Argov et al., 2002). In China, Neoseiulus cucumeris (Oudemans) have been released at the rate of 900,000 larvae/ha for the control of P. citri and Ph. oleivora, and its efﬁcacy against mites on citrus trees was 85–95% 30 days after release and pests were controlled for 6 months (Zhang et al., 2003). As regards insects, in Florida the Psychidae Cryptothelea gloverii (Packard) preyed on the citrus rust mite and consumed the eggs and adults of both Ph. oleivora and P. citri (Villanueva et al., 2005), and two species of Cecidomyiidae of the genera Feltiella and Lestodiplosis are both efﬁcient predators of Ph. oleivora eggs, larvae and nymphs (Villanueva et al., 2006). Nevertheless, different obstacles limit the use of these means, and today the control of mite infestations is fundamentally entrusted to the use of chemicals, sometimes in a context of IPM too. Chemical control is needed, especially in wetlands and crop varieties for the fresh market (McCoy et al., 1989). In this respect, the speedy development of mite populations, the rapid increase in the damage, the small size of the mite, which makes difﬁcult the monitoring, and the lack of natural enemies (McCoy, 1996a) are important. The inﬂuence of biotic and abiotic factors and horticultural practices (Jeppson et al., 1975; McCoy, 1977a; Smith and Papacek, 1981; Zamorra and Nasca, 1985) can inﬂuence population density; also, the side-effects of many chemicals can stimulate the development of populations (McCoy, 1977a, b; Smith and Papacek, 1981; Beattie et al., 1991) or the selection of resistant strains (Swirski et al., 1967; Jeppson et al., 1975; Herne et al., 1979; Omoto et al., 1994). The strategy employs various substances, and a detailed account of the topic is given by Childers et al. (1996). Included among the most effective

CHEMICAL CONTROL.

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substances are avermectins (French, 1982, 1984; McCoy et al., 1982; Childers, 1985; Zamorra et al., 1989; French and Villarreal, 1990; Moreira, 1992; Iskander, 1993; Chiaradia and Cruz, 1996; Smith et al., 1998; Bergh et al., 1999; Lu, 2000; Grout and Stephen, 2002); organochlorines (Dean, 1979; French and Gage, 1983; Oliveira et al., 1985; Childers, 1990); organotins (Childers and Konsler, 1980; French, 1982; Brown and Jones, 1983; McCoy et al., 1990; Oliveira et al., 1993; Childers, 1994a; Drishpoun et al., 1995; Moraes et al., 1995a, b; Wu, 1995); dithiocarbamates (Schwartz, 1977a; Childers, 1990; Ibrahim, 1992; Iskander, 1993; Drishpoun et al., 1995; Rezk and Gadelhak, 1996; Kalaisekar et al., 2000, 2003; Ying et al., 2000; Ran et al., 2001; Grout and Stephen, 2002); narrowrange petroleum oils (Selhime, 1984; Knapp, 1991; Iskander, 1993; Ibrahim, 1994; Rae et al., 2000; Knapp et al., 2001; Cai et al., 2001; Lei et al., 2003); and sulfur (Grifﬁths and Thompson, 1957; Brown and Jones, 1983; Ibrahim, 1992; Moraes et al., 1995a, b; Chiaradia and da Cruz, 1996; Scarpellini and Santos, 1997; Scarpellini and Clari, 1999; Kalaisekar et al., 2000; Huang, 2005). Avermectins are widely used in various regions of the world, either alone or in combination with petroleum oils, and are considered partially selective towards the beneﬁcials (McCoy et al., 1982). The use of sulfur was proposed between the end of the 19th century (Hubbard, 1885) and the beginning of the 20th century (Yothers, 1915), but it is not very selective for natural enemies (McCoy, 1977a; Smith and Papacek, 1991). Medium-range petroleum oils have been used worldwide since 1940 (Jeppson et al., 1975) and are effective, but have a modest persistence and are largely selective (McCoy et al., 1976b; Vacante, 1986). An alternative approach is to use 435–66 FC, FC 455–88 or 470 petroleum oils as a fungicide for sooty mould control and to suppress pest mites (Childers et al., 2007). Acaricides can be used in combination with other principles (insecticides, fungicides), and it is a good rule to assess their mutual interaction and selectivity (McCoy, 1996a). Recently, Bell et al. (2005) have reported that a new acaricide – tetronic acid derivate in combination with horticultural oil spray – provided excellent initial and residual control of the citrus rust mite, equal or superior to avermectins and pyridazinone derivate. Nevertheless, the selectivity of this last acaricide is unclear. INTEGRATED PEST MANAGEMENT. The above-mentioned aspects require that control is placed in a context of IPM (McCoy, 1996a), with particular attention to pertinent technical aspects (choice of chemicals, threshold values, sampling techniques, etc.). Equally important is the canopy density, which has an effect on the ability of populations to increase within a short period of time (Childers et al., 2007). In China, in the context of prevention, irradiation is suggested as a quarantine treatment (Hu et al., 2004).

8.6.3.1.3 Phyllocoptruta paracitri Hong et Kuang (Fig. 8.11) Common name Unknown.

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A

C

D

Fig. 8.11. Phyllocoptruta paracitri Hong et Kuang. (A) Lateral view of adult; (B) prodorsal shield; (C) coxal and genital regions; (D) empodium (from Hong and Kuang, 1989).

Diagnostic characteristics FEMALE. The body is fusiform, 163 μm long and 68 μm wide. The gnathosoma is 18 μm long, projecting downwards. The prodorsal shield is 35 μm long and 60 μm wide, with a frontal lobe. The median line is incomplete, the admedian lines and submedian lines are complete and form a complex design, including three cells on each side along the anterior margin of the shield. The dorsal tubercles are 33 μm apart, ahead of the rear margin; the scapular setae (sc) are 8 μm long, pointing and inclined dorsally. The coxae are smooth, and the sternal line is present. The opisthosoma possesses 27 smooth dorsal annuli, and 46–50 ventral annuli bearing round microtubercles; the dorsocentral furrow is shallow and broad. The setae c2 are 12 μm long, set on ventral annulus ten; the setae d are 50 μm long, set on ventral annulus 21; the setae e are 8 μm long, set on ventral annulus 34; the setae f are 20 μm long, on the ring ﬁve from the rear; the setae h1 are present. The empodia are four-rayed. The genital coverﬂap has ten longitudinal ribs and the proximal setae on coxisternum III (3a) are 22 μm long (Hong and Kuang, 1989). MALE. Similar to female, 138 μm long, 53 μm wide, genitalia 15 μm wide, with granules in the anterior part, and seta 18 μm long (Hong and Kuang, 1989).

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Geographical distribution China (Hong and Kuang, 1989). Bio-ecology Vagrant species collected on lower leaf surface of Poncirus sp. and Citrus sp. (Hong and Kuang, 1989).

9

Diptilomiopidae Keifer

9.1 INTRODUCTION The Diptilomiopidae family is present on Citrus with the species Diptilomiopus assamica Keifer, recorded only for India (Keifer, 1959a, b; Chakrabarti and Mondal, 1983; Dhooria et al., 2005) and Australia (Knihinicki and Boczek, 2002).

9.2 MORPHOLOGICAL CHARACTERS AND SYSTEMATIC OUTLINE The general morphological characters are similar to those of Eriophyidae. The prodorsal shield possess two or no setae, the scapular setae (sc) are present or absent, and the unpaired setae vi and ve are largely absent. The gnathosoma is sharply bent towards the base, with cheliceral stylets folded in the same way and with long oral stylets. The opisthosoma is frequently devoid of setae c1, the remaining setae are present or sometimes any one of the setae c2 or d or setae h1 are absent. The chetotaxy of the coxal plate is complete, plate I is sometimes without setae 1b and rarely with setae 1a. The chetotaxy of the legs is complete and may miss the basiventral femoral setae I and II, the antaxial genual seta of genu II, the paraxial tibial seta of tibia I and both the paraxial and fastigial tarsal setae (ft’) or the paraxial and unguinal tarsal setae (u’) of the legs I and II; tibia I is devoid of solenidium; the tarsal empodium may be thick and is commonly divided. Sometimes the genital coverﬂap may have one or two rows of ridges and have spots or semilunar granules (Lindquist 1996; Lindquist and Amrine, 1996; Amrine et al., 2003). The family includes the subfamilies Diptilomiopinae Keifer and Rhynchaphytoptinae Roivainen. The species D. assamica is ascribed to the former subfamily (Lindquist and Amrine, 1996; Amrine et al., 2003). © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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9.3 DIPTILOMIOPINAE KEIFER Diptilomiopinae are marked from the main axis of the empodium divided, usually deeply, and leg II is without basiventral femoral seta (Amrine et al., 2003).

9.3.1 Diptilomiopus Nalepa The mite lacks scapular setae and the genu is absent from both legs. The coxal setae 1b are absent (Amrine et al., 2003).

9.3.1.1 Diptilomiopus assamica Keifer (Fig. 9.1) Common name Unknown. Diagnostic characteristics FEMALE. The body is fusiform, elongate, 215–230 μm long, 60 μm thick, light yellowish-white in colour. The gnathosoma is 43 μm long, and moderately attenuate. The prodorsal shield is 31 μm long, 53 μm wide, with a design network of raised lines. Almost no anterior shield frontal lobe, median shield line present at rear only, admedian lines complete, strongly angled at crossline connections and joined by three transverse lines, the latter two joining the median line; lower front and sides of shield a pattern of vertically elongate cells. The dorsal tubercles, minus scapular setae (sc), are present within the rear shield margin, 20 μm apart. The opisthosoma possesses about 55–60 dorsal annuli and 70–75 ventral annuli; a shallow longitudinal furrow extending on each side of the mid-dorsum, fading caudally; small ventral microtubercles on ventral annuli touching ring margins, absent dorsally. The setae c2 missing; setae d 18 μm long, on about ventral annulus 26; setae e 15 μm long, on about ventral annulus 45; setae f 23 μm long, on about ventral annulus nine from rear; setae h1 absent. The genital coverﬂap is smooth. The empodia are ﬁve-rayed (Keifer, 1959b). MALE.

Unknown (Keifer, 1959b).

Geographical distribution Australia (Knihinicki and Boczek, 2002); and India (Keifer, 1959b; Chakrabarti and Mondal, 1983; Dhooria et al., 2005). Bio-ecology Diptilomiopus assamica is a leaf vagrant species, commonly associated together other eriophyoids (Aculus pelekassi, Tegolophus australis, Cosella ﬂeschneri) with

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A

B

C

D E

G F

Fig. 9.1. Diptilomiopus assamica Keifer. (A) Lateral view of adult; (B) prodorsal shield; (C) lateral view of prodorsum and gnathosoma; (D) coxal and genital regions; (E) lateral opisthosoma; (F) internal genitalia of female; (G) empodium (from Keifer, 1959b).

Phyllocoptura oleivora (Seki, 1981; Smith and Papacek, 1991). The species has been collected on Citrus limon and several varieties of orange in India (Keifer, 1959b) and Australia (Knihinicki and Boczek, 2002). Keifer (1959b) has reported that D. assamica is a rust mite but does not produce serious damage. McCoy (1996a) indicates that the leaves show diffused russeting but that control is unnecessary, while Gerson (2003) reports that the pest status of the mite on citrus is unknown.

10

Tarsonemidae Canestrini et Fanzago

10.1 INTRODUCTION The Tarsonemidae recorded on citrus are mostly mycophagous and only the citrus silver mite, Polyphagotarsonemus latus, produces severe damage to citrus and other cultivated plants (Jeppson et al., 1975; Gerson, 1992; Nucifora and Vacante, 2004); it is probable that the Tarsonemidae represent a complex species (Lindquist, 1986).

10.2 MORPHOLOGICAL CHARACTERS AND SYSTEMATIC OUTLINE The body of Tarsonemidae is formed by a gnathosoma, prodorsum and opisthosoma (Fig. 10.1). The gnathosoma possesses a stylophore, palps, infracapitulum and chelicerae (Fig. 2.4). The stylophore derives from the merged bases of chelicerae and, integrating with the infracapitulum, forms the gnathosomal capsule. The palp has two articles and the chelicerae are retractable and of different length in the various species. The prodorsum of the female has a sclerotized dorsal shield variously ornate, one pair of stigmata, two pairs of setae (v1, sc2), one pair of bothridia with its pseudostigmatic organs and one pair of pits (v2) (Fig. 10.1). The prodorsal shield of the male has vertical (v1, v2) and scapular setae (sc1, sc2) variously expressed. The opisthosoma presents dorsally ﬁve tergites (C, D, EF, H and Ps), related to each other in a telescopic way. The tergite C of the female is usually entire, well separated from the next tergite D and bearing setae c1 and c2. Tergites C and D of the male are fused together as tergite CD. Tergite D of the female is usually entire, with one pair of setae (d) and cupules (ia). In the female, tergite EF derives from tergites E and F fused together, and it bears 104

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G pa s ap 1

b

P 1b

v1 v2

1a 2a

sc2

ap 2 2b

ap 3 C 3c

c1

ap 4

c2

3b

ia D d

im f

EF e

h Ps

H

Fig. 10.1. Polyphagotarsonemus latus (Banks). Dorsal view (right) and ventral view (left) of female (from Nucifora and Vacante, 2004); ap 1–4, apodemes 1–4; b, bothridium; C, D, EF, H, Ps, opisthosomal tergites; G, gnathosoma; P, prodorsum; pa, prosternal apodeme. The setal notation is explained in the text.

two pairs of setae (e, f) and one pair of cupules (im). In the male, the tergite EF normally has one pair of setae and one pair of pits, except in some species such as P. latus, which has two setae (Fig. 10.2). Tergite H bears the setae h and the cupules ih. The pseudanal tergite Ps is a small structure, caudal or

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A ps CD

B

EF

Fig. 10.2. Polyphagotarsonemus latus (Banks). (A) dorsal (right) and ventral (left) views of male; (B) leg I of male (from Nucifora and Vacante, 2004); CD, tergite CD ; EF, tergite EF; ps, prodorsal shield.

ventro-caudal, few scleriﬁed pseudanal tergite Ps, with one pair of caudal setae (ps) in the adult female (Fig. 10.1) and none in the male (Fig. 10.2). On the ventral side, the female has an expanded and consistent (absent in the male) aggenital shield, which sometimes has one pair of aggenital setae. The ventral podosoma consists of four different coxisternal plates, generally not ornate. The ﬁrst two plates are fused to each other in the middle position and internally form a prosternal apodema and the apodemes one and two; they stretch around the edge of the rear trochanters I and II, in the dorsolateral position above the legs, have one pair of setae (1a, 2a) and a pit each (1b, 2b). The coxisternal metapodosomal plates III

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and IV have three apodemes (3, 4 and poststernal); those of the fourth pair are normally associated with the poststernal, which is unequal and median. Two pairs of setae (3c, 3b) are set near apodemes three and four (Fig. 10.1). The legs have six articles. The tibia and tarsus I of the female are fused into a single article (in the male they are always separated). In most species, leg I of the male and female bears a single claw; the latter in the female may suffer profound changes and is sessile, large and uncinate. Pretarsus I, unlike pretarsi II and III, lacks a true empodium (Fig. 10.2). Trochanter III of the female is long, subelliptic and ﬁxed; femur III is fused with the genu. Leg IV of the male generally has four articles, with the trochanter short and femur and patella fused. Leg IV of the female consists of three articles, with the trochanter short, the second resulting from the fusion of the femur and genu and the last by the fusion of the tibia with the tarsus, ending with a long seta and devoid of distal claw (Lindquist, 1986). The family is divided into three subfamilies (Pseudotarsonemoidinae Lindquist, Acarapinae Schaarschmidt and Tarsoneminae Canestrini et Fanzago) and different tribes. Polyphagotarsonemus latus belongs to the subfamily Pseudotarsonemoidinae and to tribe Pseudotarsonemoidini Lindquist (Lindquist, 1986). Table 4.1 contains information on a number of web sites that have pictures of living mites showing their colours and natural features and the damage they can cause to citrus.

10.3 PSEUDOTARSONEMOIDINAE LINDQUIST The prodorsal stigmata of the adult female are posterolateral of vertical setae. The prodorsal botridia and their sensilla correspond to setae sc1. The prodorsum of the male has three or four pairs of setae. Plate EF of the male has one or two pairs of setae. The metapodosomal venter of the adult has three or four pairs of setae (Lindquist, 1986).1

10.3.1 Pseudotarsonemoidini Lindquist The cheliceral stylets are of short to moderate length, when retracted they do not occupy most of the length of the gnathosomal capsule and are not recurved. The coxisternal plates III possess three pairs of setae, and plates IV each have one pair of setae (Lindquist, 1986).

1Formally,

it is treated with reference to citrus as a single subfamily and the morphological references seem unnecessary. However, they are given to avoid any confusion with other subfamilies present on citrus but are not relevant to this discussion.

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10.3.1.1 Polyphagotarsonemus Beer et Nucifora The prodorsal shield of the female is unenlarged and not covering the stigmata; the dorsal idiosomal setae are slender, and the setae sc2 are longer than other setae. Femur I and tarsus II of the female lack seta l’’ and spindle-like seta pl’’, respectively; the female has a gently curved, sessile claw and reduced, contiguous unguinal setae on leg I; female with tarsal setae pv’ usually enlarged, spine-like on leg I, and only two setae on tibiotarsus of leg IV (Lindquist, 1986).

10.3.1.1.1 Polyphagotarsonemus latus (Banks) (Figs 10.1 and 10.2) Common name On citrus, the species is commonly known as the citrus silver mite, but the most widely used name is broad mite, followed by other common names such as yellow tea mite, white mite or tropical mite. Diagnostic characteristics FEMALE. The body is oval, about 200 μm long and white amber to yellow or pale greenish coloured, sometimes with a white median spot, intense and with faint contours. The body setae are short, and the pseudostigmatic organs are spherical in shape with four pairs of ventral metapodosomal setae. The tibiotarsus of leg I bears a claw. MALE.

The body is 170 μm long, and similar in colour to the female. It has three pairs of prodosomal setae and four pairs of ventral metapodosomal setae; tibia and tarsus IV are fused into a tibiotarsus ending with a knob-like claw; the coxae III and IV are intimately associated. Geographical distribution Cosmopolitan and typical of tropical and subtropical areas and greenhouses of temperate and subtropical countries (Jeppson et al., 1975). Bio-ecology The citrus silver mite is injurious to many plants belonging to about 57 botanical families (Gerson, 1992), both herbaceous and arboreous trees, including citrus (Nucifora, 1961; Jeppson et al. 1975; Schwartz, 1977b; Meyer Smith, 1981; Di Martino, 1985; Meyer Smith and Schwartz, 1998a). Reproduction is parthenogenetic arrhenotokous and the optimum development of populations typically requires tropical climatic conditions (Jones and Brown, 1983). The species is active throughout the year and its reproductive activity slows during the coldest months (Jeppson et al., 1975; Schwartz, 1977b). It generally attacks the tender tissues of leaves just issued and young citrus fruits and, between these, prefers lemon; sometimes it is found in

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nurseries. The female oviposits in hollows of the lower leaf surface and younger fruit. The males carry the quiescent nymphs, also known as pharate females, from older leaves on the new sprouts (Lavoipierre, 1940; Moutia, 1958). Optimum development occurs at 25°C and with RH between 90% and 100% (Jones and Brown, 1983); with 80 of RH the lethal thermal threshold is 43.6°C, and at 25°C the intrinsic rate of increase of population is 0.3283 (Li, 1990). At 27.5 ± 0.5°C, with 67.6 ± 1.3% RH and a constant photoperiod, development of the young stages of female and male on lemon occurs in about 3.7 ± 0.1 and 3.6 ± 0.1 days, respectively, with 100% survival. Preoviposition lasts 1.0 ± 0.2 days and the female lays 5.6 ± 0.5 eggs per day for 10.5 ± 0.9 days. Longevity is equal to 13.4 ± 1.0 days for the female and 12.0 ± 2.4 days for the male. The intrinsic rate of increase is equal to 0.359, the ﬁnished rate of increase is equal to 1.42 individuals/female/day, the average time of a generation is 10.34 days and the net reproductive rate amounts to 41.0 (Vieira and Chiavegato, 1999). On Persian limes (Citrus latifolia (Tanaka ex Yu. Tanaka) Tanaka) at 22.43°C and 63.0% RH, the development from egg to adult stage takes 3.95 days and mortality is 17% (Ramos and Alvarez, 1987). The species disperses through the male, wind, farmers and insects. Notable among the latter is the action of the tobacco whiteﬂy, Bemisia tabaci (Gennadius) (Flechtmann et al., 1990) and the greenhouse whiteﬂy, Trialeurodes vaporariorum (Westwood) (Parker and Gerson, 1994). In this context, Palevsky et al. (2001) determined the speciﬁcity of the association between the mite and its phoretic hosts, observing that also attached to a citrus pest were the whiteﬂies Dialeurodes citri (Asmehad) and other Aleurodidae. It is likely that infestations on lemon also derive from weeds or vegetables near or associated with citrus groves. The natural enemies of the citrus silver mite include fungi and mites. Among the former are known Beauveria bassiana, Hirsutella thompsonii (Peña et al., 1996), Paecilomyces fumosoroseus (Wise) Brown and Smith (Clavicipitaceae) (Peña et al., 1996; Smitha and Giraddi, 2006) and Verticillium lecanii (Zimmermann) Viégas (Ascomycota) (Smitha and Giraddi, 2006). Predators number several phytoseiid mites such as Amblyseius largoensis Muma (Rodriguez and Ramos, 2000, 2004), Euseius stipulatus, Euseius hibisci (Chant) (Brown and Jones, 1983), Euseius nicholsi (Ehara et Lee) (Wu, 1984), Euseius ovalis (Evans) (Moutia, 1958; Manjunatha et al., 2001), Euseius victoriensis (Smith and Papacek, 1991), Neoseiulus agrestis (Karg) (Kolodochka and Prutenskaya, 1987), Neoseiulus barkeri Hughes (Fan and Petitt, 1994), Neoseiulus californicus (McGregor) (Peña and Osborne, 1996), Neoseiulus cucumeris (Prabaningrum et al., 1999; Cantliffe et al., 2004), Neoseiulus longispinosus (Hariyapa and Kulkarni, 1988), Typhlodromalus peregrinus (Muma) (Peña, 1992) and Typhlodromus athiasae Porath et Swirski (Karut et al., 1998). Symptomatology and damage Although the mite can infest all species of citrus it is normally seriously harmful only for the lemon, and in summer and autumn attacks young

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unripe fruits and leaves of the last growth, usually those younger and closer to the vegetative apex; these show distortions and curling, sometimes so pronounced as to prevent the development of the sprout. The lower surface of the young leaves can present a shining, silver appearance. The tarsonemids infest the fruit as soon as it has transformed from ﬂower to fruit and until they reach the size of a walnut, and their populations develop more frequently on young fruit sites in the shade and inside the apical share of the canopy; the attack causes a silvering of the pericarp with irregular contours and sometimes on the entire fruit surface; with the progress of the infestation and development of the fruit, the alteration of the pericarp evolves in russeting, with irregular contour and variable colour from white–silver to brown (Nucifora, 1961; Jeppson et al. 1975; Schwartz, 1977b; Meyer Smith, 1981; Di Martino, 1985; Meyer Smith and Schwartz, 1998a; Vacante, 2009a). The damage to lemon fruits is similar to that of Phyllocoptruta oleivora and may result in a reduction of commercial value of up to 100%. In South Africa, lemon losses of between 30% and 70% have been reported (Meyer Smith and Schwartz, 1998a). Control There is no information on biological control programmes, and the information produced fundamentally regards experimental experiences. Also for this mite, correct chemical control forms a part of IPM programmes and the chemicals should be applied on the ﬁrst symptoms of the attack. Leonel et al. (1999) reported that when lemon fruits in Argentina had ten mites/cm2 and chemical treatment which is carried, it was successful and provided goodquality fruit, using just one acaricide spray. In India, the treatment is carried when 5% of the fruit is infested (Dhooria et al., 2005). Curvilinear and simple linear regressions represent the relationship between broad mite-days and percentage fruit surface damage, and between mite-days and percentage damaged fruit per tree. Economic injury levels of 9–15 broad mite-days causing yield losses were established (Peña, 1990). In Florida, economic injury level per lime tree lies between 42 and 45 mite-days in the spring and summer, respectively (Peña et al., 2002). BIOLOGICAL CONTROL. Tested biological means include the Clavicipitaceae B. bassiana, H. thompsonii and P. fumosoroseus, the ﬁrst of these appearing to offer the best results (Peña et al., 1996; Ihsan and Yusof, 2004). The action of the Ascomycota V. lecanii and P. fumosoroseus has been recently evaluated in India (Smitha and Giraddi, 2006). The phytoseiid mite E. stipulatus has been introduced on citrus in California and offers a more convincing control, especially in spring, than that of E. hibisci (Brown and Jones, 1983). The former species, also reported as N. californicus, has lowered – along with other indigenous predatory mites – the density of attack of P. latus on lime fruit to below the levels of economic damage (Peña and Osborne, 1996). Wu (1984) has found that in southern China E. nicholsi predates 10.5 P. latus per day and guarantees good control on citrus.

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In Florida, the action of T. peregrinus has been evaluated, with promising results (Peña, 1992). Equally interesting was the response of N. barkeri (Fan and Petitt, 1994) and N. cucumeris (Prabaningrum et al., 1999; Cantliffe et al., 2004; Mizobe and Tamura, 2004). The response of E. ovalis has been known for a long time (Moutia, 1958) and, if the predator is employed at a ratio of 1 mite/20 prey, it provides good control of the tarsonemid (Manjunatha et al., 2001); recently, this action has been reviewed in India, together that of N. longispinosus (Smitha and Giraddi, 2006), the response to which seemed of practical value (Hariyapa Kulkarni, 1988), but less satisfactory was that of N. agrestis (Kolodochka and Prutenskaya, 1987). In Queensland (Australia), E. victoriensis has provided effective pest control (Smith and Papacek, 1991). In Cuba, Rodriguez and Ramos (2000) have assessed the contribution of A. largoensis and in the Czech republic the response to T. athiasae was positively evaluated (Karut et al., 1998). In the Canary Islands, the role of Typhlodromus swirskii (Athias Henriot) has been examined (Castañe and Sanchez, 2006). CHEMICAL CONTROL. With regard to chemical control since the late 1970s, several active ingredients (acaricides, insecticides and fungicides) have been experimented with and proposed, individually and variously mixed with each other, on citrus and/or herbaceous crops. Efﬁcacious among acaricides are avermectins (Gavioli et al., 1988; Herron et al., 1996; Leonel et al., 1999; Scarpellini, 1999; Misra, 2003; Sarkar et al., 2006; Venzon et al., 2006); chloronicotinyls (Reddy et al., 2005); organochlorines (Ingram, 1960; Nucifora, 1961; Lafﬁ, 1982, Brown and Jones, 1983; Di Martino, 1985; Buergo et al., 1986; Hugon and Chaupin, 1986; Gough and Qayyum, 1987; Raj and Sexena, 1988; Costilla et al., 1994; Herron et al., 1996 ; Karmakar et al., 1996; Mohapatra, 1996; Misra, 2003; Srinivasan et al., 2003; Panda and Sarangi, 2004; Reddy et al., 2005; Sarkar et al., 2006); and quinazolines (Sarkar et al., 2006). Insecticides include organochlorines (Lafﬁ, 1982; Hugon and Chaupin, 1986; Gough and Qayyum, 1987; Raj and Sexena, 1988; Costilla et al., 1994; Herron et al., 1996; Karmakar et al., 1996; Mohapatra, 1996; Reddy et al., 2005); organophosphates (Mohapatra, 1996; Mallapur et al., 2001; Misra, 2003; Panda and Sarangi, 2004; Smitha and Giraddi, 2006); neonicotinoids (Panda and Sarangi, 2004; Reddy et al., 2005); and phenylpyrazoles (Reddy et al., 2005). Sulfur is among the fungicides recorded (Cranham et al., 1962; Wolfenbarger, 1974; Jeppson et al., 1975; Brown and Jones, 1983; Buergo et al., 1986; Hugon and Chaupin, 1986; Fourie, 1989; Peña, 1989; Karmakar et al., 1996; Mohapatra, 1996; Venzon et al., 2006). INTEGRATED PEST MANAGEMENT.

In this context, selective acaricides and other substances such as sulfur are employed, which is particularly efﬁcacious against the adult and immature stages of the broad mite and used as sulfur dust, or wettable sulfur or lime sulfur, distributed in cases where no other chemicals are used, at least twice a year (Krause at al., 1996; Meyer Smith and Schwartz, 1998a). Dhooria et al. (2005) suggested that infested plants can be

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dusted with sulfur two to three times at 5-day intervals. The use of sulfur, however, negatively inﬂuences the populations of natural enemies (McCoy, 1977a, b; Smith and Papacek, 1991), but its persistence is feeble. Furthermore, sulfur cannot be employed in high ambient temperatures or in a mixture with mineral oils.

11

Tenuipalpidae Berlese

11.1 INTRODUCTION Tenuipalpidae are often confused macroscopically with Tetranychidae mites because of the orange to red colour of their body, but unlike the latter they are slow moving and do not produce silk webbing and for this reason they are also called ‘false spider mites’. Twenty-four species have been collected on citrus in different regions of the world (Table 11.1). Only a few species are injurious to citrus in some regions of the world such as Central and Southern America, where they transmit serious disease (Knorr et al., 1960, 1968; Kitajima et al., 1972; Carter, 1973; Childers et al., 2001, 2003a, b, c), while in other regions, such as the Mediterranean area (Vacante, 2009a) or South Africa, the same species are of less importance. In certain species, this behaviour is explained by the phenomenon of clones (Weeks et al., 2000; Rodrigues et al. 2003). Irrespective of this, tenuipalpids are usually regarded as secondary plant pests, but the recent emergence of a number of injurious species as vectors of plant viruses (Childers et al. 2003a) has necessitated an updated review of this relatively little-known family (Gerson, 2008).

11.2 MORPHOLOGICAL CHARACTERS AND SYSTEMATIC OUTLINE The body of the Tenuipalpidae is dorsoventrally ﬂattened, 200–400 μm long, commonly orange to red in colour and with ridges and reticulations, and sometimes yellowish-red, green, reddish-green or reddish-black. It is formed of a gnathosoma and a idiosoma (Fig. 11.1). Like the Tetranychidae the gnathosoma has a stylophore, chelicerae and palps (Fig. 11.2). The stylophore is extrudable and derived partially by fusion of © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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Table 11.1. Mites of the families Tenuipalpidae Berlese and Tuckerellidae Baker et Pritchard collected on citrus worldwide.

Families and species

Pest status

Tenuipalpidae Brevipalpus amicus Chaudhri Brevipalpus californicus (Banks) Brevipalpus chilensis Baker Brevipalpus cucurbitae Mohanasundaram Brevipalpus cuneatus (Canestrini et Fanzago) Brevipalpus deleoni Pritchard et Baker Brevipalpus dosis Chaudhri et al. Brevipalpus jambhiri Sadana et Balpreet Brevipalpus jordani Dosse Brevipalpus karachiensis Chaudhri et al. Brevipalpus lewisi McGregor Brevipalpus mcgregori Baker Brevipalpus obovatus Donnadieu Brevipalpus phoenicis (Geijskes) Brevipalpus phoenicoides Gonzalez Brevipalpus rugulosus Chaudhri et al. Brevipalpus tinsukiaensis Sadana et Gupta Pentamerismus tauricus Livshitz et Mitrofanov Tenuipalpus caudatus (Dugès)

U Me, Mi Mi U U U U U U U Mi U Ma, Mi Ma, Mi U U U U U

Tenuipalpus emeticae Meyer Tenuipalpus mustus Chaudhri Tenuipalpus orilloi Rimando Tenuipalpus sanblasensis De Leon Ultratenuipalpus gonianensis Sadana et Sidhu Tuckerellidae Tuckerella knorri Baker et Tuttle

U U U U U

Tuckerella nilotica Zaher et Rasmy Tuckerella ornata (Tucker) Tuckerella pavoniformis (Ewing)

U U Mi

Ma, Mi

Distribution

India, Pakistan Worldwide Chile, India India Italy India , USA (Florida) India, Pakistan India (northern) Egypt, Lebanon, Tanzania India, Pakistan Worldwide USA (California) Worldwide Worldwide Thailand India (northern), Pakistan India Crimea France, Greece, Italy, Portugal South Africa India, Pakistan Indonesia, Philippines Mexico India

China, Costa Rica, Iran, Philippines, Thailand Egypt Worldwide Worldwide

According to Gerson (2003), the pest status is indicated as either Ma (major), Me (medium), Mi (minor) or U (unknown). The references are indicated in the text.

the bases of the chelicerae; furthermore, the latter has a long, stylet-like digitus mobilis, which is strongly recurved proximally. The palp is simple, with one to ﬁve segments and the penultimate has no claw (Fig. 11.2). Respiration occurs through two tracheae direct anteriorly and ending in a bulb that may be related with the longitudinal folds of the invagination of the stylophore (Fig. 11.2).

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ps

PRODORSUM

GNATHOSOMA

v2

prp

sc1 sc2

PROPODOSOMA

dd

IDIOSOMA op c3

HYSTEROSOMA

c1

d1

e1

d3

e3

f3

OPISTHOSOMA

h2 h1

Fig. 11.1. Brevipalpus phoenicis (Geijskes). Dorsal view of female (from Vacante and Nucifora, 1985); dd, dorsal disjugal suture; op, opisthosomal pore; prp, prodorsal pore; ps, prodorsal shield. The setal notation is explained in the text.

The dorsal disjugal and the ventral sejugal sutures divide the idiosoma into an anterior propodosoma and a posterior hysterosoma. The dorsal surface of the body is divided into the prodorsum, corresponding to the anterodorsal part of the propodosoma, and the opisthosoma, coinciding with the hysterosoma excluding legs III and IV (Alberti and Coons, 1999; Welbourn et al., 2003) (Fig. 11.1). The prodorsum bears an anterior projection or rostral shield of different shape and size and with different projections (central, medial), three pairs of setae – including one pair of vertical (v2) and two pairs of

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1a vss B

ch

3a C

ps

tr 4a

vp

ag

gp

g2 st

ap

g1 ps1

ps2

E

e

D s

Fig. 11.2. Brevipalpus phoenicis (Geijskes). (A) Ventral view of female (partially designed); (B) prodorsum and gnathosoma (partly designed); (C) palp (from Vacante and Nucifora, 1985); (D) pretarsus of Brevipalpus recki Livshits et Mitrofanov (partially modiﬁed from Livshits and Mitrofanov, 1967); (E) distal end of tarsus II of B. phoenicis (partially modiﬁed, from Livshits and Mitrofanov, 1967); ap, anal plates; ch, chelicerae; e, eupathidium; gp, genital plate; ps, prodorsal shield; s, solenidion; st, stylophore; tr, trachea; vp, ventral plate; vss, ventral sejugal suture. The setal notation is explained in the text.

scapular (sc1, sc2) – a pair of eyes, a variable ornamentation from smooth to reticulate and a pair of pores (Welbourn et al., 2003). The opisthosoma hosts the genital and anal openings and a number of plates (ventral, genital, anal) and a maximum of 15 pairs of setae, excluding the genital and aggenital setae, of which 13 pairs are dorsal (Lindquist, 1985; Welbourn et al., 2003).

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The maximal opisthosomal chaetotaxy consists of three pairs each in the C, D and E rows and two pairs each in the F and H rows, and the PS are ventral to ventrocaudal. In the genus Brevipalpus, a maximum of ten pairs of dorsal opisthosomal setae are present and two pairs each in the C (c1, c3), D (d1, d3), E (e1, e3), F (f2, f3) and H (h1, h2) rows and a pair of opisthosomal pores (Fig. 11.1). The ventral opisthosoma has two pairs of pseudanal setae (ps1, ps2), two pairs of genital setae (g1, g2) and one pair of aggenital setae (ag) (Fig. 11.2). The dorsal and ventral surfaces have variable reticulations. The legs are articulate with the podosoma, corresponding for the ﬁrst two pairs to the posterior part of propodosoma and for pairs III and IV to the anterior part of the hysterosoma. The legs are short, wrinkled and with ﬁve free segments (Fig. 11.1). The coxae are fused to the venter and form a coxal ﬁeld. The coxal ﬁeld of legs I has three setae (1a, 1b, 1c), and the coxal ﬁeld of legs II–IV each has two setae (2a, 2b; 3a, 3b; 4a, 4b) (Fig. 11.2). As regards the chaetotaxy of other segments in the genus Brevipalpus, the trochanters I–IV bear one, one, two and one setae; the femurs I–IV four, four, two, and one setae; the genua I–IV three, three, one and one setae; the tibias I–IV ﬁve, ﬁve, three and three setae and tarsi I–IV eight, eight, ﬁve and ﬁve setae. Depending on the species, tarsi I and II have a different number and type of setae (normal, unguinals, euphatidials, tectals, fastigials, solenidia) (Fig. 11.2). The pretarsi have two claws and the empodium has tenent hairs (Fig. 11.2) (Welbourn et al., 2003). The Tenuipalpidae are divided into three subfamilies (Brevipalpinae Mitrofanov, Tenuipalpinae Mitrofanov and Tegopalpinae Smiley et Gerson) (Ghai and Shenhmar, 1984; Smiley and Gerson, 1996; Smiley et al., 1996), and the species collected on citrus belong to the ﬁrst and second subfamilies, More than 900 species are known, ascribed to 30 genera (Kane, 2003). Most of these species are included in the subfamily Brevipalpinae and genus Brevipalpus Donnadieu and in the subfamily Tenuipalpinae and genus Tenuipalpus Donnadieu (Jeppson et al., 1975; Meyer Smith, 1979, 1993; Ghai and Shenhmar, 1984; Sepasgosarian, 1990). The genus Brevipalpus shelters around 300 species (Welbourn et al., 2003), and 928 hosts of 513 genera and 139 botanical families have been recorded for only B. californicus, B. obovatus and B. phoenicis (Childers et al., 2003a). Table 4.1 has information on a number of web sites that show pictures of living mites with their colours and natural features and their damages on citrus.

11.3 BREVIPALPINAE MITROFANOV The Brevipalpinae have a ventral plate (Mitrofanov, 1973a, b).

11.3.1 Brevipalpus Donnadieu The prodorsum possesses a prodorsal shield and three pairs of setae (v2, sc1, sc2). The opisthosomal dorsal setae have a maximum of ten pairs, two pairs

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each in the C (c1, c3), D (d1, d3), E (e1, e3), F (f2, f3), H (h1, h2) rows and a pair of opisthosomal pores. The ventral opisthosoma bears two pairs of pseudanal setae (ps1, ps2), two pairs of genital setae (g1, g2) and one pair of aggenital setae (ag). The dorsal and ventral surfaces are variously reticulate (Welbourn et al., 2003).

11.3.1.1 Brevipalpus amicus Chaudhri (Fig. 11.3) Common name Unknown.

A B

D

C

Fig. 11.3. Brevipalpus amicus Chaudhri. Female. (A) Dorsal view; (B) ventral view; (C) rostral shield; (D) palp (from Chaudhri, 1972a).

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Diagnostic characteristics FEMALE. The body is 245 μm long and 153 μm wide. The rostral shield has one pair of central, three pairs of adjacent and two pairs of lateral projections. The prodorsum has three pairs of marginal setae including one pair of vertical (v2) and two pairs of scapular (sc1, sc2), short, lanceolate and serrate, with two eyes per side set between the scapular setae. Irregular broken striations cover the majority of the body, with a few mediolateral reticulations. The striations fade away laterally and the area in the middle is devoid of any ornamentation. The opisthosoma possesses nine pairs of setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2), short, lanceolate serrate, with dorsocentral simple and shorter. The striations directing marginally lateral to the groove, a few reticulation along the inner margin of groove and caudally. A few dim reticulations medially, posterior to the suture. Irregular, somewhat transverse, broken, striations in the middle; the area near centrals II is bare. The ventral plate and the genital plate each have reticulations that are wider than they are long and have one pair and two pairs of setae, respectively. Reticulations marginally, lateral to ventral plate. The tarsus II has one solenidion (Chaudhri, 1972a). MALE.

Unknown (Chaudhri, 1972a).

Geographical distribution Pakistan (Chaudhri, 1972a); India (Ghai and Shenhmar, 1984). Bio-ecology The species has been collected on Andrachne sp. in Pakistan (Chaudhri, 1972a), and on other herbaceous and arboreous host plants, including Citrus sp. in India (Ghai and Shenhmar, 1984; Gerson, 2003). Its bio-ecology is unknown.

11.3.1.2 Brevipalpus californicus (Banks) (Fig. 11.4) Common name Citrus ﬂat mite. Diagnostic characteristics FEMALE. The body is ovoid, 240 μm long and 156 μm wide, red coloured with a dark pattern in the central area. The rostral shield extends beyond the base of femur I, the central rostral projection is long and pointed and the division between the central and medial rostral projections is deep. The prodorsum bears three pairs of marginal setae, including one pair of verticals (v2) and two pairs of scapular (sc1, sc2), short, lanceolate and serrate, one pair of eyes per side, one pair of prodorsal pores. The central area of the prodorsum has irregular to uniform reticulations, the medial region has uniform reticulations and the lateral is wrinkled. The opisthosoma bears ten pairs of dorsal

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A

B

Fig. 11.4. Brevipalpus californicus (Banks). Female. (A) Dorsal view (from Jeppson et al., 1975); (B) ventral view (from Baker, 1949).

setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), marginal short, lanceolate and serrate. The furrow between setae c1 and c3, extending to setae h2 is indistinct. The central area between setae c1 and d1 has uniform reticulations. The central area between setae d1 and e1 has an indistinct groove starting at setae d1 and ending at setae e1; the area between grooves is rugose. The central area between setae e1 and h2 has irregular reticulations. The median area between setae c1, d1, e1 and the lateral furrow to f3 is reticulate. The lateral area between the lateral furrow and setae c3, d3 and e3 is rugose. The genital plate is areolate to colliculate. Tarsus II has two solenidia (Baker, 1949; Welbourn et al., 2003). MALE. Similar to female, but smaller and less common. The metapodosoma pres-

ents reticulations mediocaudally on the venter (Pritchard and Baker, 1958). Geographical distribution Worldwide distribution (Jeppson et al., 1975). Bio-ecology The citrus ﬂat mite has been collected on 316 plants belonging to 67 genera and 33 botanical families (Childers et al., 2003a). It feeds on leaves, fruits,

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twigs, petioles and developing buds of all citrus in varying degrees (Sadana and Joshi, 1979) and with a different symptomatology according to cultivar. On Valencia orange and grapefruit the fruit are preferred to twigs and leaves, and the green fruit to ripe fruit (Schwartz, 1970). Like B. obovatus and B. phoenicis, it is considered an important citrus pest in different areas of the world (Lewis, 1949; Dean and Maxwell, 1967; Jeppson et al., 1975; Schwartz, 1977c; Childers, 1994b; French and Rakha, 1994; Ochoa et al., 1994; Childers et al., 2003c), and for orchids also (Ochoa et al., 1994). The female has two chromosomes and reproduction may be parthenogenetic thelytokous (Helle and Bolland, 1972). The bacterial symbiont Cardinium sp. manipulates the reproduction of the mite, feminizing it (Chigira and Miura, 2005; Groot and Breeuwer, 2006). With temperatures of between 18° and 24°C and 55% RH, egg development takes about 9 days, and with temperatures of between 21° and 30°C, the development of larval, protonymphal and deutonymphal stages requires 8.6, 6.2 and 7.0 days, respectively, and the quiescent stages 3.6 days each. The life cycle takes about 3 weeks, oviposition begins about 3 days after last moult and the female lays one egg per day over 25 or more days (Manglitz and Cory, 1953). A decrease in temperature from 30.5° to 15.3°C increases developmental durations (Nehru and Bhagat, 2006). On Azalea sp. at 23 ± 1°C and 60 ± 5% RH, the life cycle from egg to adult develops in 26.5 days and at 27°C in 21 days (Trindade and Chiavegato, 1994). On citrus, the density of populations is commonly low and the greatest densities are observed in the cracks and crevices of fruit, driven also by the wind or hail, where juvenile and adult stages feed around the edges, emphasizing the outlines. In the early infestation, the mite populations are most abundantly at the stem end of the fruit near or under the fruit button (Dean and Maxwell, 1967; Meyer Smith and Schwartz, 1998b). Preferential distribution of the mite on the leaves of Citrus sinensis (cultivar Jaffa) revealed that the tender leaf growth stage has a signiﬁcantly low population of the mite (1.4) compared with the mature leaf (5.0). The preference of the mite in decreasing order is mature > young > tender leaf growth stage. The midrib region of tender, young and mature leaves had maximum mite population compared with apex, base, margin, veins of lamina and petiole. The abaxial leaf surface harboured the maximum mite population (Nehru et al., 2005). In Egypt, the populations of B. californicus reach their greatest densities on orange and mandarin in August; they prefer the eastern part of canopy and the lower branches and especially the fruits. The mite is present in larger populations on oranges compared with mandarins (El-Halawany, 1991). The use of phosphorus compounds may cause an increase in population (Attiah and Wahba, 1973), and the pruning and sequence of dry weather followed by light rainfall may favour their development (Baptist and Ranawerra, 1955). Natural enemies include a number of phytoseiid mites such as Euseius hibisci (Gupta et al., 1971), Euseius mesembrinus (Dean) (Badii et al., 1993), Euseius scutalis (Athias Henriot) (El-Halawany et al., 1993), Galendromus helveolus (Chant) (Chen et al., 2006), Typhlodromus doreenae Schicha (James and Whitney,

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1993), and Typhlodromus rhenanus (Oudemans) (Gupta et al., 1971). A decline of almost 90% in the numbers of B. californicus was achieved after releasing 50 E. scutalis per citrus tree in Egypt (El-Halawany et al., 1993). Symptomatology and damage The citrus ﬂat mite damages, directly or indirectly, leaves, twigs, branches and fruit of various citrus species and varieties. In the seedlings of sour orange, mite feeding produces severely stunted rounded leaves, with marginal necrosis and tip burn. The infestation also results in severe stunting of new shoot growth with the formation of corky, swollen buds. Leaves fail to develop from these gall-like buds. Damage has also been observed on citrus seedlings in Venezuela and Florida, identiﬁed as Brevipalpus gall (Knorr and Denmark, 1970), and the affected plants do not produce new leaves (Childers et al., 2003c). In Australia, it produces a silvering of the fruit, particularly lemon, known as ‘silver mite’ (Jeppson et al., 1975). In Texas, injury to citrus is prevalent on the inside of fruit in the lower tree canopy, below 2 m. The presence of B. californicus and B. phoenicis has been associated with the rind spotting of grapefruit with high density of populations. The fruit lesions ﬁrst appear as very slight yellowish and circular discoloured areas in depressions on the fruit surfaces of grapefruit or oranges. A central brown necrotic area or spot develops in these lesions and slowly becoming darker and corky. Irregularly shaped spots appear and vary in size from 1 to 12 mm or larger, and are most common ﬁrst on the side and stylar end of the fruit but, as feeding continues, may cover the fruit. Initially the spots are level with the surface of the peel, but on storage tend to become raised and darker in colour (Dean and Maxwell, 1967; French and Rakha, 1994). The irregularly damaged areas on the fruit gradually become necrotic and result in the fruit being rejected for the fresh market. In South Africa, the feeding activity of B. californicus produces brown to bronze-coloured and corky, scab-like spots on the rind of sweet orange and, on the Valencia orange, alterations are found in the depressions of the rind, essentially on the exposed side of the fruit; on grapefruit these scab-like spots have an irregular shape and size and are not restricted to any particular side of the fruit rind, mainly in the middle of the trees (Schwartz, 1977c; Meyer Smith and Schwartz, 1998b). In Italy B. californicus is regarded as a secondary pest, sometimes responsible for the silvering of fruit of various citrus (Di Martino, 1985). The mite may transmit a toxin or virus to the leaves and fruit of citrus; this can strongly limit the development of citrus groves in certain areas of the world (Knorr et al., 1968). On grapefruit and orange varieties in Texas and Florida, smaller necrotic lesions form on the infested surface of leaves and fruits, known as ‘leprosis-like spotting’ or ‘nail head rust’ (Dean and Maxwell, 1967; Jeppson, 1989; French and Rakha, 1994). In Florida it has been recorded on twigs and branches and is called ‘Florida scaly bark’, and in Argentina both symptoms are known as ‘lepra esplosiva’ or ‘leprosis’. At present, leprosis is the major viral disease of citrus

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in Brazil (Chagas et al., 2001), but the only species vector of the disease is B. phoenicis (Rodrigues et al., 2003). On the leaves and fruit of sweet orange, feeding mites produce small platelets of dry gum material; similar alterations are formed on twigs. When the injury extends and the twig’s growth is affected, the alterations increase and resemble scrofulous shelling of bark (Knorr et al., 1960). Control The need for control varies between different countries of the world. In some South American countries it is a priority, in others not. The same condition has also been registered in the Mediterranean region: thus, whereas in Italy the need for control is rare and a good horticultural management and/or a rational use of chemicals against pests is usually sufﬁcient, in Egypt this need appears stronger and El-Halawany (1991) has recommended that acaricides should be applied before August to the lower level and eastern side of trees. In South Africa, treatment is recommended during midsummer at the ﬁrst signs of a build-up on twigs and/or fruit (Meyer Smith and Schwartz, 1998b). There is little information regarding biological control and natural enemies alone are generally unable to contain the development of pest populations. For this reason, control is entrusted to the use of chemicals. Among the latter, avermectins (Iskander, 1993; Iskander et al., 1993 ; French and Rakha, 1994); organochlorines (Jeppson et al., 1975; El-Kady et al., 1977; Hanna et al., 1977; Iskander et al., 1993); petroleum oils (El-Kady et al., 1977; Hanna et al., 1977; Iskander, 1993); and sulfur (Jeppson et al., 1975; Ibrahim, 1992) have been cited in the literature.

11.3.1.3 Brevipalpus chilensis Baker (Fig. 11.5) Common name Grape ﬂat mite. Diagnostic characteristics FEMALE. The body is ovoid and red in colour, 267 μm long and 167 μm wide. The rostral shield extends beyond the base of femur I, with the central rostral projections broader than the two lateral projections and the posterior central area reticulate. The prodorsum bears three pairs of setae (v2, sc1 and sc2), short, lanceolate and apparently not serrate, and one pair of eyes per side. The opisthosoma possesses nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2), marginal short, lanceolate and apparently not serrate and one pair of pores. The central area of the prodorsum and opisthosoma is uniformly reticulate, with areolae more long than wide. The reticulate pattern of the genital plate and anterior ventral plate bears areolae wider than long, and the more lateral areolae are as long as wide. The posterior medioventral setae 4a extend beyond the ventral sejugal suture between the propodosoma and

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A B

C

Fig. 11.5. Brevipalpus chilensis Baker. Female. (A) Dorsal view; (B) ventral view (from Baker, 1949). Male. (C) Dorsal view (from Pritchard and Baker, 1958).

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hysterosoma. Tarsus II has one solenidion. The dorsal setae of femora I and II are lanceolate serrate and about half as long as the width of the segment (Baker, 1949). MALE.

Similar to female but smaller (Pritchard and Baker, 1958).

Geographical distribution Chile (Baker, 1949); India (Ghai and Shenhmar, 1984). Bio-ecology The grape ﬂat mite infests different plants of fruit (citrus, ﬁg, almond), ornamentals (Chrysanthemum sp., Geranium sp.), forest trees, annual weeds and is destructive for grapevines in Chile (Jeppson et al., 1975). It lives on the lower surface of the leaves and overwinters as a fecund female on various shelters of the bark. The greatest development occurs on the higher part of the canopy and oviposition in October; depending on the climatic conditions, the mite completes a generation in 25.3 days (with a range of 18–59 days) and from three to six generations per year (Gonzalez, 1968). Natural enemies include the Coccinellid Stethorus histrio Chazeau (Aguilera, 1987) and the phytoseiid mites Neoseiulus chilenesis (Dosse) (Gonzalez, 1983), Neoseiulus californicus and Typhlodsomus pyri; the latter predators appear able to regulate the development of mite populations (Vargas et al., 2005). Symptomatology and damage Feeding injury of B. chilensis on citrus is not severe and well known. On lemon, it results in a roughened silvering of the rind (Ripa and Rodriguez, 1999). Control Natural enemies are unable to contain the development of pest populations (Hernera Villamil, 1958). Control is entrusted to the use of chemicals, and applications should be started as early as possible in spring (Jeppson et al., 1975). Chemicals cited in the literature include avermectins (Gonzalez, 1999, 2001; Gonzalez and Barria, 1999); pyrethroids (Gonzalez, 2001); organochlorines (Jeppson et al., 1975); petroleum oils (Curkovic et al., 1994; Montano, 1995); sulfur (Jeppson et al., 1975); or other acaricides (quinoxalines, organosulphurs, pyridazinones) (Curkovic et al., 1994) used as spring or summer treatments. Jadue et al. (1996) tried to control the mite by low temperatures.

11.3.1.4 Brevipalpus cucurbitae Mohanasundaram (Fig. 11.6) Common name Unknown.

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B

A

C

Fig. 11.6. Brevipalpus cucurbitae Mohanasundaram. Female. (A) Dorsal view; (B) ventral view; (C) palp (from Mohanasundaram, 1982).

Diagnostic characteristics FEMALE. The body is 265 μm long, 158 μm wide and red in colour. The rostral shield reaches more than half the femur I, bifurcate and with the shield tip blunt. The palp has four segments, and the last bears three long setae, with

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two spines and one sensory seta at its tip. The prodorsum has a polygonal reticulation, three pairs of prodorsal setae (v2, sc1 and sc2), with sc2 longer and sc1 shorter. The opisthosoma is reticulate in the middle, with broader cells in the sides, and bears ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), lanceolate and serrate. On the venter, the ventral setae (3a) are short and the posterior (4a) long. The ventral plate bears a pair of aggenital setae (ag). The genital plate bears two pairs of genital (g1, g2) and the anal plate two pairs of pseudanal setae (ps1, ps2). All ventral setae are simple, except the genital lanceolate and serrate. Tarsi I and II bear one solenidion each (Mohanasundaram, 1982). MALE.

Unknown (Mohanasundaram, 1982).

Geographical distribution India (Mohanasundaram, 1982; Sadana and Balpreet, 1995). Bio-ecology The species has been collected on squash (Cucurbita maxima Duchesne) (Mohanasundaram, 1982) and on Key lime (Citrus aurantifolia (Sadana and Balpreet, 1995)). There is no information on its bio-ecology.

11.3.1.5 Brevipalpus cuneatus (Canestrini et Fanzago) (Fig. 11.7) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid, 267 μm long and 200 μm wide. The rostral shield has central rostral projections of normal length, the medial projections are small and the posterior area is reticulate. The prodorsum bears three pairs of setae (v2, sc1 and sc2) short, lanceolate and serrate, and one pair of eyes per side. The opisthosoma has ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), short, lanceolate and serrate with setae f and h clearly paired. Setae c1, d1 and e1 are short, lanceolate and apparently not serrate. The dorsal reticulate pattern may be irregular; the lateral areolae on the prodorsum longer than it is wide and on the anterior part where the pattern becomes dorsal the elements are wider than they are long; on the posterior part of the prodorsum the pattern does not appear to tend dorsally. The opisthosomal pattern is more indeﬁnite and on the dorsal surface the areolae are broken and are wider than they are long. On the lateral sides just outside the dorsal setae the areolae are longer than they are wide. The ventral surface is covered by a reticulate pattern, with the areolae of the genital plate wider than they are long, and those of the ventral plate are slightly wider than they are long. Other areolae, except those on the posterior centre of the propodosoma, tend

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A

B

Fig. 11.7. Brevipalpus cuneatus (Canestrini et Fanzago). Female. (A) Dorsal view; (B) ventral view (from Baker, 1949).

to be longer than they are wide. The posterior ventral setae 4a are short, not longer than ventral setae 3a. Tarsus II has a long solenidion. The dorsal setae of femora I and II are short, lanceolate serrate, about half as long as the width of the segment (Baker, 1949). MALE.

Unknown (Pritchard and Baker, 1958).

Geographical distribution Italy (Canestrini and Fanzago, 1876; Baker, 1949). Bio-ecology The species was ﬁrst described by Canestrini and Fanzago (1876) on specimens collected on a hedge of unknown host plant in northern Italy. Successively, the mite has been recorded by Cavarra and Mollica (1903) and Tardo (1960) in Sicily and Calabria (Italy), on lemon and other citrus, respectively. Di Martino (1985) treated the tenuipalpid as a citrus pest, and subsequent

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research carried out by Vacante and Nucifora (1985) on the citrus mites in Italy has not conﬁrmed the presence of the species. Di Martino (1985) reports that the mite is present during the year as both adult and egg, with high densities in the depressions of the rind, along the midrib of leaves and in the cracks and crevices of the fruit. On lemon fruits the feeding activity of the mite causes silvering of the rind; the damage is marked on fruits of mandarin and produces a break of glands of the essential oils, with hardening of the superﬁcal layers of the rind and crevices. Further evaluation of this mite species is required.

11.3.1.6 Brevipalpus deleoni Pritchard et Baker (Fig. 11.8) Common name Unknown.

Fig. 11.8. Brevipalpus deleoni Pritchard et Baker. Dorsal view of female (from Pritchard and Baker, 1958).

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Diagnostic characteristics FEMALE. The body is 263 μm long and 167 μm wide. The rostral shield has striae, the central projections do not extend beyond the base of femur I and the ﬁrst medial projection is very small. The prodorsum bears three pairs of setae (v2, sc1 and sc2), very short, narrowly lanceolate and sparsely serrate, and two eyes per side; the prodorsal reticulate pattern has a median ridge and broad lateral margin irregularly rugose; the mediolateral depression has irregular reticulations. The opisthosoma possesses nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2), and the setae c1, d1 and e1 are smooth. The ventral and genital plates are areolate. The ventral setae 3a are shorter than the posterior ventral setae 4a. Tarsi I and II each have a single solenidion. The dorsal setae of femur I are broadly lanceolate serrate (Pritchard and Baker, 1958). MALE.

Unknown (Pritchard and Baker, 1958).

Geographical distribution USA (Florida) (Pritchard and Baker, 1958); India (Sadana and Balpreet, 1995). Bio-ecology The species has been collected on Petrea sp. in Florida (Pritchard and Baker, 1958) and on Citrus jambhiri Lsh., C. aurantifolia and Ficus carica Linnaeus in northern India (Sadana and Balpreet, 1995). Its bio-ecology is unknown.

11.3.1.7 Brevipalpus dosis Chaudhri, Akbar et Rasool (Fig. 11.9) Common name Unknown. Diagnostic characteristics FEMALE. The body is 245 μm long and 133 μm wide. The palp has four segments, distally bearing two setae and one sensory seta. The rostral shield has a few reticulations at the base, a deep narrow median notch, one median and three lateral projections on each side. The mediolateral and anterior part of the propodosoma is reticulate, and the middle surface is devoid. The prodorsum bears three pairs of setae (v2, sc1 and sc2) serrate, with sc2 slightly longer, and two eyes per side. The anterior half of the opisthosoma is reticulate, and posteriorly the mediolateral portion is reticulate. Longitudinal, broken striae anterior and posterior to setae e1; striae are directed marginally on the lateral sides. The opisthosomal setae c1, d1 and e1 are minute, the c3 are serrate and the lateral opisthosomal setae (d3, e3, f2, f3, h1 and h2) are lanceolate and serrate; the seta h1 cross the distance h1–h2. The ventral seta 3a is 10 μm long and

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A B

E

C

D

Fig. 11.9. Brevipalpus dosis Chaudri, Akbar et Rasool. Female. (A) Dorsal view; (B) ventral view; (C) prodorsal shield; (D) dorsal seta; (E) palp (from Chaudri et al., 1974).

the posterior ventral seta 4a 34 μm long. The ventral plate is reticulate at the junction of the anterior and lateral sides, with one pair of simple setae (ag). The genital plate has transverse, wavy striations and two pairs of serrate setae (g1, g2). The anal plate bears two pairs of pseudanal setae (ps1, ps2), of which one pair are serrate. Tarsus II possesses one solenidion (Chaudhri et al., 1974).

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Unknown (Chaudhri et al., 1974).

Geographical distribution Pakistan (Chaudhri et al., 1974); India. Bio-ecology The species has been collected on Olea europea Linnaeus in Pakistan (Chaudhri et al., 1974) and on lime from India by the writer.

11.3.1.8 Brevipalpus jambhiri Sadana et Balpreet Common name Unknown. Diagnostic characteristics FEMALE. The body is 203 μm long and 133 μm wide. The rostral shield has one central and four medial projections on each side, and the ﬁrst medial close to central is very small. The prodorsum has three pairs of short lanceolate setae (v2, sc1 and sc2) and two eyes per side. The opisthosoma bears nine pairs of short lanceolate dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2). The reticulations mediolaterally of the prodorsal fade away laterally. The reticulate pattern is constituted by longitudinal, wavy lines mediolaterally, turning oblique posteriorly and meeting caudally, with faint, broken striations laterally. The posterior ventral setae 4a are about four times as long as the ventral setae 3a. The ventral plate is reticulated, with areolae wider than they are long, and genital plate with transverse striations. Tarsus II has two solenidia. The dorsal setae of femora I and II are not longer than the width of the segment (Sadana and Balpreet, 1995).

MALE.

Unknown (Sadana and Balpreet, 1995).

Geographical distribution Northern India (Sadana and Balpreet, 1995). Bio-ecology The species has been collected on Citrus jambhiri in Punjab in northern India (Sadana and Balpreet, 1995). Its bio-ecology is unknown.

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11.3.1.9 Brevipalpus jordani Dosse (Fig. 11.10)1 Common name Unknown. Diagnostic characteristics FEMALE. The body is similar to that of B. phoenicis, and is not possible to distinguish dorsally between the two species. The prodorsum bears three pairs of setae (v2, sc1 and sc2), two eyes per side and two pores, small and barely outlined. The palp has a long seta, pointed and small accessory setae. The hysterosoma possesses ﬁve pairs of lateral setae and two well-developed pores. Tarsus II has two solenidia (Dosse, 1972). MALE.

Similar to female, smaller and rare (Dosse, 1972).

LARVA AND NYMPHS. The larva has the opisthosomal dorsal setae c1, d1 and e1 minute, and e1 approximate between them, while the setae c3, d3, e3, f3 and h2 are well developed and the same length and slightly serrate, and h2 are very long and simple. The nymphs have the opisthosomal dorsal setae c1, d1 and e1 are minute and the ﬁrst and second pairs simple and e1 approximate and slightly serrate; the other dorsal opisthosomal setae are well developed, slightly serrate and the same length. In B. phoenicis the opisthosomal dorsal setae d3 and e3 are very short and f3, h2 and h1 are developed and the same length (Dosse, 1972). This latter aspect has been conﬁrmed by Welbourn et al. (2003).

Geographical distribution Lebanon; Tanzania; Egypt (Dosse, 1972). Bio-ecology The species has been described by Dosse (1972) on specimens collected on Eriobotrya japonica japonica (Thunb.) Lindley, lemon and orange in Jordan. The same author reports that he collected the mite in Tanzania on Passiﬂora sp. in September 1966, and that the specimens described by Attiah (1956) as B. phoenicis in Egypt are the same species. Reproduction is parthenogenetic (Dosse, 1972).

1The

diagnostic characteristics of nymphs stated by Dosse (1972) for B. jordani and by Gonzalez (1975) for B. phoenicoides shown a probable synonymy. Nevertheless, any conclusion depends on the study of the specimens of the two species. For this reason, in this work the two species are treated separately.

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B A

Fig. 11.10. Brevipalpus jordani Dosse. (A) Dorsal view of left side of larva; (B) dorsal view of right side of nymph (from Dosse, 1972, partially modiﬁed).

11.3.1.10 Brevipalpus karachiensis Chaudhri, Akbar et Rasool (Fig. 11.11) Common name Unknown. Diagnostic characteristics FEMALE. The body is 204 μm long, and 102 μm wide. The rostral shield has a narrow, deep notch, one median and three lateral projections on each side.

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A

B

C

Fig. 11.11. Brevipalpus karachiensis Chaudri, Akbar et Rasool. Female. (A) Dorsal view; (B) ventral view; (C) prodorsal shield (from Chaudri et al., 1974).

The prodorsum is reticulate mediolaterally, with faint reticulations in the middle, and bears three pairs of setae (v2, sc1 and sc2) lanceolate, serrate, about the same length and one pair of eyes per side. The opisthosoma has thick, irregular, longitudinal striations, meeting caudally in the middle, wavy, forming cells caudally, and the lateral striations are directed marginally. The opisthosoma bears nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2). The setae c3 are lanceolate and serrate, the setae c1 are longer than d1 and e1, and other opisthosmal setae are lanceolate, with h1 shorter. The ventral setae 3a are 8 μm long, and the posterior ventral setae 34 μm long. The ventral plate is reticulate, fading away in the middle, with one pair of simple setae (ag). The genital plate has transverse, wavy striations, and two pairs of simple setae (g1, g2). The anal plate bears two pairs of pseudanal setae (ps1, ps2), of which one pair are serrate. Tarsus II bears two solenidia (Chaudhri et al., 1974).

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Unknown (Chaudhri et al., 1974).

Geographical distribution Pakistan (Chaudhri et al., 1974); India (Ghai and Shenhmar, 1984; Sadana and Sidhu, 1990). Bio-ecology Collected on different host plants (Chaudhri et al., 1974; Ghai and Shenhmar, 1984) and also in India on C. sinensis (Sadana and Sidhu, 1990). Its bio-ecology is unknown.

11.3.1.11 Brevipalpus lewisi McGregor (Fig. 11.12) Common name Citrus ﬂat mite, ﬂat mite, scab mite or bunch mite.

Fig. 11.12. Brevipalpus lewisi (McGregor). Female. Dorsal view (from Jeppson et al., 1975).

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Diagnostic characteristics FEMALE. The body is ovoid and light reddish brown to bright red in colour, 246 μm long and 153 μm wide. The rostral shield has two long central rostral projections, and the medial projections are large but smaller and notched in some specimens. The prodorsum bears three pairs of setae (v2, sc1 and sc2), short, slightly lanceolate and serrate and one pair of eyes per side. The opisthosoma has ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), short, slightly lanceolate and serrate and one pair of opisthosomal pores. The prodorsal and opisthosomal reticulate pattern does not meet dorsally and has areolae longer than they are wide. The areolae of the genital plate are wider than long; anterior at plates, the reticulate pattern is faint towards central area; the other part of pattern with areolae longer than wide. The distal palpal segment possesses three setae. Tarsus II has one solenidion. The dorsal setae of femora I and II are lanceolate serrate and about half as long as the width of the segment (Baker, 1949). MALE. Similar to female, with idiosomal dorsal setae shorter than the longitudinal intervals between them. Propodosoma without mediodorsal reticulations. Hysterosomal pores are present and strongly tubular (Pritchard and Baker, 1958).

Geographical distribution Worldwide distribution (Jeppson et al., 1975). Bio-ecology The citrus ﬂat mite is a pest on citrus (Lewis, 1949; Elmer and Jeppson, 1957), grapes (Buchanan et al., 1980; Arias Giralda and Nieto Calderon, 1985), pistachio (Rice and Weinberger, 1981), pomegranates (Ebeling and Pence, 1949), walnuts (Michelbacher, 1956) and many ornamental plants (Jeppson et al., 1975). On citrus it infests twigs and leaves, and prefers green to ripe fruit, and fruit at any stage to the leaves. On the leaves of Vitis champini Planchon (cv Dog Ridge) the development of immature stages takes from 16.8 days at 34°C and 35% RH to 27.9 days at 22°C and 70% RH. The mean fecundity ranges from 5.7 eggs per female per day at 34°C and 35% RH to 18.8 eggs per female per day at 22°C and 70% RH (Buchanan et al., 1980). The maximum value of rm is 0.04/day, at 28°C and 35% RH. The optimal net reproductive rate (Ro) of 4.82 is observed at 22°C and 70% RH. A ﬁeld study of seasonal abundance and intra-vine distribution of the mite showed that populations increased about 60 times during a season (Buchanan et al., 1980). In central California, the mite overwinters as an adult, but in Imperial Valley and Coachella Valley on citrus they are constantly active throughout the year, and on grapes are inactive in the winter. The peaks of populations are reached in the warmest months because the high temperatures and low RH do not negatively affect the populations. On

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grapes in Spain, the citrus ﬂat mite abandons overwintering sites when temperatures reach 20°C. The winter females migrate to the growth shoots over a period of 1–2 months, ﬁnishing in mid-May. Four generations develop between May and September. The most dangerous period for fruit production is the mites’ colonization of the nodes where the bunches are situated, and this critical period starts at the beginning of June (Rodriguez et al., 1987). Natural enemies include the phytoseiid mites (Euseius victoriensis (James and Whitney, 1993), Phytoseius plumifer (Canestrini et Fanzago) (Yousef, 1970), Metaseiulus mcgregori (Chant) (Rice and Weinberger, 1981), Neoseiulus reticulatus (Oudemans) (Buchanan et al., 1980), Typhlodromus pyri (Yousef, 1970) and Typhlodromus doreenae (Smith and Papacek, 1991; James and Whitney, 1993)); the tydeid (Homeopronematus anconai (Baker)) (Hessein and Perring, 1988); and the stigmaeid mite (Agistemus exsertus) (Yousef, 1970). Symptomatology and damage On citrus, B. lewisi lives on green fruit, which are preferred to ripe fruit and leaves, and the populations are abundant at the stem end of the fruit or under the button. Attacks frequently occur on areas of fruits injured by leafhoppers (Cicadellidae) or citrus thrips. On ripe fruit these areas normally become inconspicuous, but if infested develop in conspicuous, scab-like, isolated depressions where the mites live, and the rind-blemished areas evolve as scars. High population densities may cause large alterations on the surface of the fruit. The damage may produce a reduction in the quality of fruit. The mite does not cause damage to leaves or wood, and the scab-like scars observed on most varieties of citrus rarely occur on grapefruit (Elmer and Jeppson, 1957). It has been observed that the attack of B. lewisi on tangerines causes grade-reducing scarring on 17–28% of the fruit, on 16–21% of Navel oranges and on 18–35% of grapefruits (Elmer, 1968). On Lisbon lemon fruits, scarring may affect over 25% of the fruit (Lewis, 1949). Control In south-eastern Australia, B. lewisi is controlled in grapevines by the phytoseiids T. doreenae and E. victoriensis, thus facilitating a reduction in acaricide applications (James and Whitney, 1993). Moreover, natural enemies do not always guarantee a sufﬁcient control of infestations. Chemical control is spearheaded by several acaricides. The latter include organohalogens (Goodwin, 1982); carbinols (Iacob, 1978); sulphonates (Iacob, 1978); organochlorines (Telliev and Geshev, 1980; Goodwin, 1982; Kerns, 2006); petroleum oils (Goodwin, 1982); organosulphurs (Telliev and Geshev, 1980); tetronic acid derivates (Kerns, 2006); sulfur (Chakrov, 1973; Goodwin, 1982); and organosulphurs alone or in various mixtures (Goodwin, 1982). On citrus, B. lewisi is very susceptible to sulfur and the best results are obtained with applications in late winter and early spring; during the summer plant damage may be prevented by organochlorine sprays (Jeppson et al., 1975). In Spain, insecticide treatments are recommended on grapes before June (Rodriguez et al., 1987).

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In Arizona, organochlorines and tetronic acid derivates ensure satisfactory control of mites on citrus for about 1 month (Kerns, 2006).

11.3.1.12 Brevipalpus mcgregori Baker (Fig. 11.13) Common name Unknown. Diagnostic characteristics FEMALE. The body is 246 long and 166 μm wide. The rostral shield extends beyond the base of femur I, the central rostral projection is long and pointed and the medial rostral projections are small and the central part has longitudinal striae. The prodorsum has three pairs of marginal setae, including one pair of verticals (v2) and two pairs of scapular (sc1, sc2) short, broadly lanceolate and serrate, and one pair of eyes per side. The opisthosoma possess nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2) short, broadly lanceolate

A

B

Fig. 11.13. Brevipalpus mcgregori Baker. Female. (A) Dorsal view; (B) ventral view (from Baker, 1949).

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and serrate, with the exception of c1, d1 and e1, which are very small and simple. The prodorsum and opisthosoma have a reticulate pattern made up of longitudinal striae that in the opisthosoma form elongated areolae. The genital plate has areolae wider than long, those of the ventral plate less so; other areolae tend to be longer than they are wide. Tarsus II has one solenidion. The dorsal setae of femora I and II are broadly lanceolate, serrate, about half as long as the width of the segment (Baker, 1949). MALE.

Unknown (Baker, 1949).

Geographical distribution USA (California) (Baker, 1949). Bio-ecology The species has been collected on buds of lemon in California (Baker, 1949). Its bio-ecology is unknown.

11.3.1.13 Brevipalpus obovatus Donnadieu (Fig. 11.14) Common name Ornamental ﬂat mite, privet mite. Diagnostic characteristics FEMALE. The body is 250–300 μm long, and its colour ranges from light orange to dark red with various patterns of dark pigmentation. The rostral shield does not extend beyond the base of femur I, it is blunt and with a shallow division between central and medial projections. The prodorsum has three pairs of marginal setae, including one pair of verticals (v2) and two pairs of scapular (sc1, sc2) lanceolate serrate, one pair of eyes per side and one pair of prodorsal pores. The central area of the prodorsum changes from smooth to rugose, the medial area is uniformly reticulate and the lateral area from smooth to rugose. The opisthosoma bears nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2), lanceolate serrate and one pair of opisthosomal pores. The furrow between setae c1 and c3, extending to setae h2, is narrow. The central area between setae c1 and d1 has large reticulations. The central area between the setae d1 and e1 has well-developed transverse reticulations. The central area between setae e1 and h2 has irregular reticulations, more ‘U’-shaped posteriorly. The median area between setae c1, d1, e1 and the lateral furrow to setae f3 are reticulate, and posterior to setae f3 are more wrinkled. The lateral area between lateral furrow and setae c3, d3 and e3 is rugose, with the posterior region between setae f3 and h1 more wrinkled. The ventral, genital and anal plates are uniformly colliculate. Tarsus II has one solenidion (Jeppson et al., 1975; Welbourn et al., 2003).

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Fig. 11.14. Brevipalpus obovatus Donnadieu. Female. Dorsal view (from Jeppson et al., 1975). MALE.

Similar in colour to female but smaller. The dorsomedian area of the propodosoma is irregularly striate or reticulate. The dorsocentral hysterosomal setae are very short, and the opisthosoma possesses four pairs of dorsolateral setae (Pritchard and Baker, 1958). Geographical distribution Worldwide distribution (Jeppson et al., 1975). Bio-ecology The ornamental ﬂat mite has been collected on 451 plants belonging to 119 genera and ﬁve botanical families (Childers et al., 2003a) and is common on

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privet and citrus (Jeppson et al., 1975). Like B. californicus and B. phoenicis, it is considered an important citrus pest in different areas of the world (Lewis, 1949; Dean and Maxwell, 1967; Jeppson et al., 1975; Schwartz, 1977c; Childers, 1994b; French and Rakha, 1994; Ochoa et al., 1994; Childers et al., 2003c). The female is aploid and possesses two chromosomes, the males are rare and reproduction is fundamentally parthenogenetic thelytokous (Helle and Bolland, 1972; Helle et al., 1980); the male mates but does not fertilize the female and some male specimens are clearly intersex (Pijnacker et al., 1981; Weeks et al., 2003). The bacterium Cardinium sp. induces thelytoky by feminizing unfertilized haploid eggs. Moreover, some clones of B. obovatus contain neither Cardinium nor any other bacteria that could induce thelytoky. Females treated with antibiotics produce no males, suggesting that in these lines thelytoky is probably a genetic property of the species itself (Groot and Breeuwer, 2006). The optimum temperatures for development are between 20° and 27°C (Jeppson et al., 1975); according to Goyal et al. (1985), 25°C is optimum, and the greatest development occurs at 30°C, the least at 20°C and extreme temperature conditions of 15 and 35°C are unﬁt; the highest mortality occurs at 20°C, with the lowest at 25°C. On Azalea sp. at 23 ± 1°C and 60 ± 5% RH, the life cycle from egg to adult takes 27.8 days and, at 27°C, 21.5 days (Trindade and Chiavegato, 1994). On citrus leaves at 25 ± 2°C and 65 ± 5% RH, the incubation period occurs in 3.07 days, with a hatching percentage of 87.49%. The larval, protonymphal and deutonymphal stages take 2.76, 2.63 and 3.21 days, respectively (Rezk, 2001). According to Jeppson et al. (1975), at 20° and 27°C, a total of three and 54.3 eggs per female are produced over an adult life span of 67 and 38.1 days, respectively; according to Rezk (2001), at 25 ± 2°C and 65 ± 5% RH, the lifespan is 34.26 days. In citrus crops, B. obovatus overwinters mostly as adult stages, in the lower parts of plants or sheltered on the lower surface of the leaves, sometimes with eggs and immature stages (Jeppson et al., 1975). In Egypt, the principal population peaks on citrus have been observed in early spring and autumn, with an average of 0.52 ± 0.04 and 0.61 ± 0.02 mites/leaf, respectively (Rezk, 2001). The factors limiting the development of populations include pathogens and predators. Among the fungi is reported H. thompsonii (Rosas Acevedo and Sampedro Rosas, 2000). Predators include the phytoseiids E. scutalis (Donia et al., 1995) and Neoseiulus idaeus Denmark et Muma (Tamai et al., 1997); the tydeids Tydeus californicus (Banks); the stigmaeids A. exsertus; the and cheyletid mite Cheletogenes ornatus (Rizk et al., 1983; Rezk and Gadelhak, 1996). Symptomatology and damage The mite infests the leaves, fruit and twigs of citrus. Its feeding activity removes cell substances, kills cells and injects toxic saliva into the tissues, and on sweet orange leaves causes large chlorotic spots (Morishita, 1954). The symptomatology of feeding on leaves and fruit is similar to that illustrated in Texas for B. californicus and B. phoenicis (Dean and Maxwell, 1967;

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Childers, 1994b; French and Rakha, 1994; Childers et al., 2003c). In the USA (Jeppson et al., 1975), South Africa (Meyer Smith and Schwartz, 1998c) and Mediterranean regions, the ornamental ﬂat mite does not cause serious damage to citrus. The pest is considered responsible for the transmission of citrus leprosis, but its ability to transmit the virus is not proved (Childers et al., 2001); the disease is actually absent from the USA (Childers et al., 2003b), where there have been no problems for a long time (Morishita, 1954). In Argentina, mite feeding produces on the leaves of sweet orange large chlorotic spots, with a concentric ring of resinous, chestnut-coloured substance, covering up to a third of the leaf surface. Similar but smaller changes are produced on fruits. Ring spots also affect the twigs. In later stages, these alterations erupt to produce scaling of the bark, as in scaly bark psorosis. This injury is known as ‘lepra esplosiva’ or ‘leprosis’ (Vergani, 1945). In Venezuela, the ornamental ﬂat mite is associated with ‘halo scab’ on the leaves of sour orange seedlings grown in humid structures. When B. obovatus and B. phoenicis are present on the same plant, the injury to the leaves and stems is more severe and primarily due to the latter species (Knorr et al., 1960). Control Natural enemies alone are unable to control the pest and its control is entrusted to the use of acaricides, including organohalogens (Rezk, 2001); organochlorines (Jeppson et al., 1975; Childers, 1994b; Rezk and Gadelhak, 1996); organotins; pyrazoles (Rezk, 2001); petroleum oils; pyridazinones or high rates of sulfur (Jeppson et al., 1975; Childers, 1994b).

11.3.1.14 Brevipalpus phoenicis (Geijskes) (Figs 11.1, 11.2 and 11.15) Common name Red and black ﬂat mite, reddish black ﬂat mite, leprosis mite, scarlet mite, red crevice mite. Diagnostic characteristics FEMALE. The body is oval, about 300 μm long and red coloured with a central dark pattern in older specimens. The rostral shield does not extend beyond the base of femur I, with a long and pointed central projection and a shallow division between the central and medial projections. The prodorsum has three pairs of marginal setae, including one pair of vertical (v2) and two pairs of scapular (sc1, sc2) lanceolate and from serrate to ﬁnely serrate, one pair of eyes per side, and one pair of occasionally obscure prodorsal pores. The central area of the prodorsum is smooth and with isolate areolae, the medial area has elongated reticulations and is smooth anteriorly and the lateral area is rugose. The opisthosoma possesses nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f3, h1 and h2), lanceolate and from serrate to ﬁnely serrate and one pair of

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B C

D

E

Fig. 11.15. Brevipalpus phoenicoides Gonzalez. Female. (A) Dorsal view; (B) solenidia of tarsus II; (C) dorsal seta; (D) deutonymph; (E) deutonymph of Brevipalpus phoenicis (Geijskes) (from Gonzalez, 1975).

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opisthosomal pores. The furrow between setae c1 and c3, extending to setae h2 is well developed. The central area between setae c1 and d1 is rugose. The central area between setae d1 and e1 with distinct groove starting at setae d1 and extending to setae e1. The central area between setae e1 and h2 has six to eight ‘V’- to ‘U’-shaped ridges and reduced posteriorly. The median area between setae c1, d1, e1 and lateral furrow to setae f3 is reticulate. The lateral area between the lateral furrow and setae c3, d3 and e3 is rugose, with the posterior region between setae f3 and h1 more wrinkled. The ventral and anal plates are uniformly verrucose and the genital is areolate to colliculate. Tarsus II bears two solenidia (Meyer Smith, 1981; Welbourn et al., 2003). MALE.

Similar to female, smaller and rare. The propodosoma has mediolateral reticulations dorsally and the opisthosoma bears four pairs of dorsolateral setae (Pritchard and Baker, 1958). Geographical distribution Worldwide distribution (Jeppson et al., 1975; Meyer Smith, 1979). Bio-ecology

The red and black ﬂat mite is highly polyphagous, and the list of host plants includes 486 species belonging to 118 genera and 64 botanical families (Childers et al., 2003a). It is commonly associated with B. californicus and B. obovatus on leaves, fruit and twigs of different citrus, and with the latter is considered an important citrus pest in different areas of the world (Lewis, 1949; Dean and Maxwell, 1967; Jeppson et al., 1975; Schwartz, 1977c; Childers, 1994b; French and Rakha, 1994; Ochoa et al., 1994; Childers et al., 2003c). It is also a pest on coffee in Brazil (Chagas et al., 2001), tea in Asia (Oomen, 1982), passion fruit (Kitajima et al., 1997) and various ornamental plants worldwide (Jeppson et al., 1975). Their populations usually live on the fruit and lower leaf surface and aggregate along the mid-vein or major lateral veins, and normally move to another plant when resources become limited (Haramoto, 1969). The female is aploid, possesses two chromosomes and reproduction is parthenogenetic telithockous (Helle et al., 1980; Pijnacker et al. 1981; Weeks et al., 2003); the aploid condition is guided by a bacterium of the Wolbachia genus (Otto and Jarne, 2001) and, like other Brevipalpus spp., the bacterium Cardinium sp. induces thelytoky by feminizing unfertilized haploid eggs (Groot and Breeuwer, 2006). At 25°C, 60% RH and LD 14:10, on the leaves and fruit of Valencia and Pera Rio orange the preoviposition period is 2.34 days long; the females lay an average of 22.5 eggs each; the mean incubation period lasts 7.7 days; the developmental period of larvae and nymphs is 1.6 days and 9.9 days, respectively; and the life span of adults is 22.2 days (Chiavegato, 1986). On fruits of Citrus aurantifolia, the thresholds for egg and post-embryonic development and the pre-oviposition period are 9.1°, 15.5° and 11°C, respectively, and the sums of effective temperatures for the completion of the egg stage, postembryonic development, pre-oviposition period and complete life cycle are

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135, 145, 54 and 330.4 day-degrees, respectively. Under constant temperatures, the average length of a generation ranges from 27.5 days at 24°C to 18.3 days at 30°C, while in naturally ﬂuctuating temperatures it varies from 30.8 days at 21–23°C to 19.4 days at 30–32°C (Prieto Trueba, 1975). At 27.5°C, the life cycle on citrus takes 21 days and at 13.8°C is 96 days; about ten generations a year occur in Egypt (Zaher et al., 1970). The pest has a low growth rate (rm = 0.127) and high rates of survival, and in constant conditions the net reproduction rate is 33.2 eggs/mite, the generation time is 27.6 days and the population doubles in 5.5 days. The stable age proﬁle of the population is distributed as 74.9%, 19.24% and 6.07%, respectively, of eggs, immature stages and adults (Kennedy et al., 1996). On citrus fruits B. phoenicis has a higher reproductive rate compared with B. californicus and B. obovatus (Trindade and Chavegato, 1990). With regard to dispersion, the mite moves less than 1 cm per day. A wind speed of 23 km/h is not enough to trigger mite dispersal, and wind speeds of 30 and 40 km/h are capable of triggering the dispersal of less than 1% of mites on fruits. Mite dispersal studies with the use of sticky traps under ﬁeld conditions also showed that B. phoenicis dispersal is limited when compared with other mite species from citrus groves (Alves et al., 2005b). The natural enemies include pathogenic fungi and several predators (Table 11.2). Symptomatology and damage Like other false spider mites, B. phoenicis injects toxic saliva into the fruit, leaf, stem and bud tissues of citrus, and prefers the green fruit to ripe fruit, and the latter to leaves and twigs. On the leaves, the mite normally lives on the lower surface along the mid-vein or principal lateral veins, causing a yellow blistering of the leaf surface opposite feeding mite aggregations. It also eats the stems or green twigs (Haramoto, 1969). The damaged areas slowly become necrotic, with consequent leaf drop, coinciding with high infestations (Childers, 1994b). However, the phenomenon is not common (Childers et al., 2003c). On grapefruit or oranges, the injury is similar to that previously described for B. californicus and B. obovatus in Texas (Dean and Maxwell, 1967; French and Rakha, 1994; Childers et al., 2003c) and in South Africa (Schwartz, 1970, 1977c; Meyer Smith and Schwartz, 1998d). Oranges infested by B. phoenicis are usually lighter, and their weight is inversely proportional to the degree of infestation; furthermore, affected trees may lose 50% of their yield (Rodrigues et al., 2003). In Italy, Di Martino (1985) has observed greyish scabby patches and medial cracks on the apical epidermis of mandarin fruit. Many lesions are located on the oleiferous glands, which result in these being emptied and dried. The orange fruit shows rounded and reddish patches. The greatest risk from mite attack is from the transmission of severe diseases such as ‘leprosis’ or ‘lepra esplosiva’ (Kitajima et al., 1972; Carter, 1973; Boaretto and Chiavegato, 1994; Rodrigues et al., 1997; Childers et al., 2003c),

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Table 11.2. Brevipalpus phoenicis Geijskes: natural enemies (fungi, insects and mites). Families and species Ascomycotaa Verticillium lecanii (Zimmermann) Viégas Clavicipitaceae Aschersonia sp. Hirsutella thompsonii Fisher Metarhizium anisopliae (Metschnikoff) Sorokin Paecilomyces sp. Coccinellidae Sticholotis punctata Crotch Cheyletidae Mexecheles hawaiiensis (Baker) Phytoseiidae Amblyseius acalyphus Denmark et Muma Amblyseius largoensis (Muma) Euseius alatus De Leon Euseius alstoniae (Gupta) Euseius citrifolius Denmark et Muma Euseius delhiensis (Narayan et Kaur) Iphiseiodes zuluagai Denmark et Muma Metaseiulus camelliae (Chant et Yoshida-Shaul) Phytoseiulus macropilis (Banks) Typhlodromips cananeiensis Gondim Jr. et Moraes Typhlodromips swirskii (Athias Henriot) Typhlodromus homalii (Gupta) Stigmaeidae Agistemus sp. Zetzellia malvinae Matioli et al. Zetzellia javanica Ehara et Oomen-Kalsbeek Tydeidae Pronematus elongates Baker Pronematus ubiquitus (McGregor) Pronematus sp. Tydeus californicus (Banks) aThis

Reference(s) Albuquerque et al., 2000

Albuquerque et al., 2000 Cabrera, 1978; Rosas Acevedo and Sampedro Rosas, 2000; Rossi Zalaf and Alves, 2006 Albuquerque et al., 2000; Magalhães et al., 2005 Albuquerque et al., 2000 Haramoto, 1969 Haramoto, 1969 De Vis et al., 2006 Haramoto, 1969 Gravena et al., 1994; Yamamoto and Gravena, 2001; Reis et al., 2003 Kumari and Sadana, 1991 Gravena et al., 1994; Yamamoto and Gravena, 2001; Reis et al., 2003; De Vis et al., 2006 Neena and Sadana, 2000 Gravena et al., 1994; Yamamoto and Gravena, 2001; Reis et al., 2003; De Vis et al., 2006 De Vis et al., 2006 Haramoto, 1969; Jaime de Herrero, 1985 De Vis et al. , 2006 Zaher et al., 1971 Sandhu et al., 1979 Oomen, 1982 De Vis et al., 2006 Oomen, 1982

Gupta, 2001 Zaher et al., 1971 Borthakur, 1981; De Vis et al., 2006 Zaher et al., 1971

species has not been included in a particular family, resulting in the systematics of the group being unclear.

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recorded in Argentina, Brazil, Paraguay, Venezuela and also recently in Panama. In Florida the disease has been known since the late 1800s, but we have no updates since 1960 (Morishita, 1954; Childers et al., 2001), and the subsequent research excludes the presence of the virus in Florida and Texas (Childers et al., 2003b). With regard to the presence and distribution of the virus there is some confusion, and clear differences between feeding damage and leprosis virus infection are few (Childers et al., 2003c). The responsible agent is Citrus Leprosis Rhabdovirus (CiLV), which occurs in two forms: the more common CiLV-C, infecting the cytoplasm of infected plant cells, and CiLV-N, occurring in the cell nucleus; the prevention costs in Brazil alone are about US$100 million per year (Rodrigues et al., 2003). The damage from the disease negatively affects yield; and more than US$60 million per season are spent on chemical sprays to control the mite vector (Rodrigues, 2006). Bassanezi and Laranjeira (2007) recently calculated that acaricide applications for the annual control of infestations of the mite cost the Brazilian citrus industry $ 62 million, constituting 35% of total agrochemical costs and 14% of total production costs in mature orchards. Like Brevipalpus spp. present on citrus, only B. phoenicis transmits the virus throughout the different biological stages, but not transovarially, and each mite acquires them separately in order to become infective. The complete nucleotide sequence and genomic organization of CiLV-C has recently been established (Pascon et al., 2006). The virus is mechanically transmitted from plant to plant and from some herbaceous plants belonging to the genera Atriplex, Beta and Chenopodium (Chenopodiaceae), Gomphrena (Amaranthaceae) and Tetragonia (Tetragoniaceae), and the trials of puriﬁcation and characterization have been unsuccessful (Childers et al., 2001). The virus attacks mainly sweet orange, but also citrange, citron, Cleopatra mandarin, grapefruit, lemon, mandarin, sour orange and tangor (Lovisolo, 2001). The principal symptomatology consists of lesions on fruit, leaves, twigs and small branches, with early fruit drop, leaf drop, death of twigs and branches leading to decline of tree vitality (Rodrigues et al., 2003). Localized lesions commonly, occur; those deriving from CiLV-N are usually smaller than those related to CiLV-C (Bastianel et al., 2006). The lesions result in a long-term decline in the quality of sweet orange trees, bringing about fruit drop and even death. Weeks et al. (2000) report that clones, corresponding to isofemale lines and deriving from single females of the mite, are collected from citrus number 17 genotype lines. Other various clones of B. phoenicis – from Sao Paolo and Florida, and varying along geographical lines by a mitochondrial DNA analysis – differ in their capacity to transmit CiLV (Rodrigues et al., 2003), and the ﬁtness of the mite became limited when it was displaced to new host plants. According to Groot et al. (2005), the phenomenon can be explained by the fact that the species consists of diverse, host-specialized clones instead of one generalist form, each adapted to different environments and host plants. The taxonomic inferences based on the mitochondrial COI sequences in B. phoenicis do not help the classical taxonomy based on morphology, and beg a taxonomic revision of the group (Groot and Breeuwer, 2006).

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In Florida another disorder is described as ‘a diffuse laminar chlorosis and a marginal chlorosis of the leaves’, known as ‘phoenicis blotch’, but does not represent a true leprosis or stem cankering. This disorder is similar to the early stages of leprosis but it is not accompanied by a separation of the parts affected – resinous or corky. The damage is linked to large populations of the mite and limited to plants previously defoliated by stress factors (Jeppson et al., 1975). In Venezuela on sour orange, the mites are associated with the scabs caused by the fungus Elsinoe fawcetti Bitanic et Jenkins (Elsinoaceae), and they produce a disorder known as ‘halo scab’, with symptoms ranging from spotting of the leaves to death of the plants; the leaves attacked by the scab fungus alone do not drop, but leaves attacked by both drop – with defoliation and death of plants (Knorr and Malaguti, 1960; Chiavegato and Kharfan, 1993). In Honduras, B. phoenicis is associated with E. fawcettii on sour orange (C. aurantifolia) (Evans et al., 1993). A similar interaction between two mite species and the fungus Sphaceloma fawcetti Jenkins has been reported in Costa Rica (Ochoa et al., 1994). In Brazil, the mite prefers citrus fruits already infected by sweet orange scab, a fungal disease caused by the fungus Elsinoe australis Bitancourt et A.E. Jenkins (Chiavegato and Kharfan, 1993) and, according to Gravena et al. (1994), the action of the phytoseiid Euseius citrifolius (Denmark et Muma) on the pest is limited on oranges with scab, protecting the scab from populations of B. phoenicis. Another disorder, before B. californicus was considered, also attributed in Venezuela to B. phoenicis and named ‘Brevipalpus galls’, may cause the death of 60% of orange seedlings. Gall-like protuberances on the main stem of the seedlings are observed, which are woody and appear like axils that have proliferated to the point where bud-studded cushions develop. The axils occupied by these cushions do not produce leaves and, if all the buds are replaced by cushions, the trees become devoid of leaves and soon die (Knorr et al., 1960, 1968; Knorr and Denmark, 1970). Control As regards the monitoring and sampling of mite populations, Netto (1987), examining the causes of the failure of control of the red and black ﬂat mite on citrus in Brazil, recommended acaricide spraying when only 10% of the fruit are infested with at least 30 mites/cm2. Goes et al. (1985) report that random samples should be taken from two of every 100 trees (ﬁve fruits showing signs of attack being taken from each of the two), and treatment should begin when mites are found on 2% or more of the total number of fruits examined. In South Africa, the use of acaricides is recommended in summer when the ﬁrst adults are observed on fruit (Krause et al., 1996). Sequential sampling for leprosis on orange trees is usually based on binomial distribution. However, Bassanezi and Laranjeira (2007) concluded from a study of the spatial patterns of leprosies and of B. phoenicis that the patterns of either the diseased or the infested plants deviate from a binomial distribution. Hence, estimates of disease or mite incidence may not be precise.

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The control was entrusted to different methods: quarantine, horticultural and chemical, biological and integrated pest control. QUARANTINE.

The quarantine method provides enforcement of a strict quarantine regime for citrus plants that may harbour the three vectors of plant viruses, and is probably the best way to control these diseases. However, such regimes are seldom fully enforced, and the diseases would therefore be expected to spread from their present foci, and quarantine would not be useful if a Tenuipalpid is suspected of being windborne (Gerson, 2008).

HORTICULTURAL PRACTICES. These are intended to limit the spread of leprosis and the Tenuipalpid using resistant varieties, disease- and/or mite-free plants; removing or pruning infected trees or parts; avoiding hedge and windbreak plants that are heavily infested or that are alternate host plants for the mites; and removing susceptible weeds and other plants from the area (Maia and Oliveira, 2004). The problem of resistance has been investigated over a long period (Sadana and Joshi, 1979; Grewal, 1993; Bastianel et al., 2005, 2006; Rodrigues, 2006), and recent studies have suggested that inheritance of resistance to leprosis may be controlled by only a few genes (Bastianel et al., 2006). BIOLOGICAL CONTROL.

Unfortunately, biological control alone does not solve the problem. Albuquerque et al. (2000) have reported that Metarhizium anisopliae, Aschersonia sp. and Paecilomyces sp. (Clavicipitaceae) and V. lecanii (Ascomycota) not give solid control, while Magalhães et al. (2005) – using M. anisopliae var. acridum – observed 90% adult mortality after 8 days. H. thompsonii gives a good response (Rosas Acevedo and Sampedro Rosas, 2000), and some isolates caused high mortality with indices higher than 90% (Rossi Zalaf and Alves, 2006), but we have no evidence of any practical application. The same disadvantage has also been observed for predatory mites recorded in different countries of the world. Gravena et al. (1994) investigated predation of the phytoseiid Euseius citrifolius on the pest on oranges. In Israel, Galendromus occidentalis has been introduced from California (Swirski and Dorzia, 1969).

CHEMICAL CONTROL. If necessary, control is entrusted, as in Central and South America, to chemicals that include organochlorines (Jeppson et al., 1975; Schwartz, 1975a, b; Suplicy Filho et al., 1977; Takematsu et al., 1979; Srivastava et al., 1980; Myazaki et al., 1982; Oliveira et al., 1983; Goes et al., 1985; Mariconi et al., 1989; Raga et al., 1990; Alves et al., 2000a, b; Omoto et al., 2000); organotins (Sato et al., 1991, 1994; Raga et al., 1996; Oliveira et al., 1998; Scarpellini and Santos, 1999; Alves et al., 2000b); petroleum oils (Sato et al., 1991; French and Rakha, 1994; de Moraes et al., 1995b; Oliveira et al., 2003); and sulfur (Elmer and Jeppson, 1957; Ebeling, 1959; Knorr, 1959; Busbillo Pardey and Saldarriaga Velez, 1970; Jeppson et al., 1975; Suplicy Filho et al., 1977; Takematsu et al., 1979; Myazaki et al., 1982; de Moraes et al., 1995b; Chiaradia and da Cruz, 1996), either alone or in various mixtures. Also, in this case, it is necessary to razionalize the use of various chemicals to contain the common problem of the selection of resistant strains (Omoto, 1998; Alves et al., 2000a;

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Campos and Omoto, 2002, 2006). It has been observed that applications of fenbutatin oxide reduce the numbers of B. phoenicis on citrus by 75–100% for as long as 4 months post-treatment (Oliveira et al., 1998). However, the large number of sprays required for this purpose is leading to the development of resistance to both organochlorines and phenyl pyrazoles. INTEGRATED PEST MANAGEMENT. To limit the use of chemicals, it has been suggested

in Brazil to direct control towards IPM (Gravena, 1998), respecting the technical characteristics of the main acaricides registered for citrus and the detailed procedures of a laboratory bioassay conducted to evaluate the efﬁcacy of acaricides against citrus leprosis mite (Gravena et al., 2005).

11.3.1.15 Brevipalpus phoenicoides Gonzalez (Fig. 11.15) Common name Unknown. Diagnostic characteristics FEMALE. The body is 300 μm long. The median and propodosomal area lack reticulations, instead having distinctly elongated alveoli. The mediolateral area has three to four rows of irregular, large, polygonal areolae, about eight cells per row. The prodorsal pores are present. The hysterosomal reticulation is without deﬁned polygonal areolae. The median area has contorted, elongated elements converging in the median posterior section; the mediolateral area is without reticulation, replaced by oblique, interrupted lines. The opisthosomal pores are set behind strong arcs. The hysterosomal marginal setae gradually diminish in length. Tarsus II bears lateral and mesal solenidia, 6.0 and 4.5 μm long respectively (Gonzalez, 1975). It is difﬁcult to separate B. phoenicoides from B. jordani Dosse, and the ﬁrst species is treated for completeness of information; the solution of the problem will be given by future research. MALE.

The propodosomal pattern is similar to that of the female. The opisthosomal pores are large and the opisthosomal reticulation is made up of longitudinal elements (Gonzalez, 1975). Geographical distribution Thailand (Gonzalez, 1975). Bio-ecology Brevipalpus phoenicoides has been collected in Thailand on C. reticulata, C. aurantifolia, Pandanus sp. and Synzygium jambos (Linnaeus) Alst. (Gonzalez, 1975). Its bio-ecology is unknown.

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11.3.1.16 Brevipalpus rugulosus Chaudhri, Akbar et Rasool (Fig. 11.16) Common name Unknown. Diagnostic characteristics FEMALE. The body is 214 μm long and 112 μm wide. The distal segment of the palp has two setae and one sensory seta. The rostral shield has a deep, narrow notch with longitudinal, broken striations, and one median and four projections on each side. The prodorsum possesses irregular and broken striations

A

B

E

D C

Fig. 11.16. Brevipalpus rugulosus Chaudhri, Akbar et Rasool. Female. (A) Dorsal view; (B) ventral view; (C) prodorsal shield; (D) dorsal seta; (E) palp (from Chaudri et al., 1974).

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in the middle and laterally with reticulations mediolaterally and bears three pairs of setae (v2, sc1, sc2) slightly lanceolate, serrate and with sc1 slightly longer and has one pair of eyes per side. The opisthosoma presents longitudinal wavy lines meeting medially and caudally, and lateral striations directed marginally. The dorsal opisthosomal setae are nine pairs (c1, c3, d1, d3, e1, e3, f2, f3 and h1), with setae c1, d1 and e1 simple, c3 serrate and the others lanceolate serrate. The ventral setae 3a are 8 μm long, and the posterior 4a 26 μm. The ventral plate is reticulate, with one pair of simple setae (ag), and with areolae lateral to shield. The genital plate is reticulate with two pairs of serrate setae (g1, g2). The anal plate bears two pairs of serrate pseudanal setae (ps1, ps2). Tarsus II has two solenidia (Chaudhri et al., 1974). MALE.

Unknown (Chaudhri et al., 1974).

Geographical characteristics Pakistan (Chaudhri et al., 1974); northern India (Sadana and Balpreet, 1995). Bio-ecology Brevipalphus rugulosus has been collected on different host plants in Pakistan (Chaudhri et al., 1974), and occurs in India on fruit trees, vegetables, medicinal and ornamental plants, oilseeds and weeds (Sadana and Rajinder Sharma, 1989). In India it has also been collected on C. sinensis (cultivar Muscambi), C. medica (cultivar Acida) and Citrus lemon (Ghai and Shenhmar, 1984; Sadana and Balpreet, 1995). Development occurs from 20° to 30°C and stops at 15° and 35°C. At 30°C the life cycle of the female is shorter (18.24 ± 1.74 days), while fecundity (12.15 ± 1.23), daily rate of eggs (2.33) and longevity (18.72 ± 3.14) are greater. At this temperature, the viability of eggs is 82.43%, and the lowest mortality rate is observed (Sadana and Rajinder Sharma, 1989).

11.3.1.17 Brevipalpus tinsukiaensis Sadana et Gupta Common name Unknown. Diagnostic characteristics FEMALE. The body is 300 μm long and 150 μm wide. The rostral shield is deeply notched, with one central and four lateral projections on each side. The prodorsum has three pairs of marginal setae, including one pair of vertical (v2) and two pairs of scapular (sc1, sc2), short, lanceolate and serrate, one pair of eyes per side and with reticulations mediolaterally, bare centrally and laterally. The opisthosoma bears nine pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3 and h1) lanceolate and serrate, with reticulations meeting caudally, incomplete reticulations centrally, oblique striations laterally. The ventral and

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genital plates are reticulated. Tarsus II possesses two solenidia (Sadana and Gupta, 1983). MALE.

The podosoma is broad anteriorly, and the opisthosoma is narrowed posteriorly. The dorsum is divided into propodosoma, metapodosoma and opisthosoma by two transverse sutures (Sadana and Gupta, 1983). Geographical distribution India (Sadana and Gupta, 1983; Sadana and Balpreet, 1995). Bio-ecology The species has been collected on C. limon and other host plants in Assam (Sadana and Gupta, 1983), and on C. medica (var. Acidula) and C. jambhiri in different areas of northern India (Sadana and Balpreet, 1995). Its bio-ecology is unknown.

11.3.2 Pentamerismus McGregor The palp is ﬁve-segmented. The prodorsum bears three pairs of setae (v2, sc1, sc2) and two pairs of eyes on each side. The dorsal opisthosoma bears 12 pairs of setae (c1, c2, c3, d1, d2, d3, e1, e3, f1, f3, h1, h2). There is a genital and a ventral plate. Sometimes the ventral plate is not invariably distinguishable (Meyer Smith, 1979).

11.3.2.1 Pentamerismus tauricus Livshits et Mitrofanov Common name Unknown. Diagnostic characteristics FEMALE.

The body is 279 μm long, 156 μm wide and broadly ovate. The dorsum is evenly striate. The prodorsum has three pairs of setae (v2, sc1, sc2) and two eyes per side. The striae between the setae v2 and sc2 are longitudinal. The opisthosoma bears 12 pairs of dorsal setiform setae (c1, c2, c3, d1, d2, d3, e1, e3, f2, f3, h1 and h2), and transverse reticulations between setae c1, d1 and e1. The dorsal opisthosomal setae c2, d2 and f2 are longer, and e1 shorter. The ventral body is evenly striate except into genital and anal plates that are smooth; the setae 1a and 4a are longer than setae 3a. The ventral plate is lacking, and one pair of aggenital setae is present (ag); the genital plate possesses two pair of genital setae (g1, g2), and the anal plate has three pairs of setae (Livshits and Mitrofanov, 1970). MALE. Similar to female, smaller, 210 μm long and 115 μm wide (Livshits and Mitrofanov, 1970).

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Geographical distribution Crimea (Livshits and Mitrofanov, 1970). Bio-ecology Pentasmirenus tauricus has been collected on Cistus tauricus J. Presl et C. Presl in the Crimea (Livshits and Mitrofanov, 1970). Ghai and Shenhmar (1984) reported it for citrus in the former USSR. Its bio-ecology is unknown.

11.4 TENUIPALPINAE MITROFANOV Tenuipalpinae are devoid of ventral plates (Mitrofanov, 1973a, 1973b)

11.4.1 Tenuipalpus Donnadieu The opisthosomal dorsal setae h2 are ﬂagellate. If these ﬂagellate setae are absent, the podosoma is very broad and the opisthosoma very narrow. The ventral and genital plates may be fused together to form a genitoventral plate, or they may be separated. If the ventral plate is distinct, it is not quadrangular in shape (Meyer Smith, 1979).

11.4.1.1 Tenuipalpus caudatus (Dugès) Common name Unknown. Diagnostic characteristics FEMALE. The body is approximately 320 μm long and red coloured. The rostral shield is deeply cleft medially, and presents lateral angulations. The prodorsum is striate longitudinally on the median area except posteriorly, where it is irregularly striate. The posterolateral corners are angulate. The prodorsal setae include one pair of verticals (v2) and two pairs of scapular (sc1, sc2), with v2 and sc1 setiform and minute and sc2 greatest and spatulate. The opisthosoma bears ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), with setae c1, d1 and e1 spatulate and c1 reaching to the bases of setae next behind; the setae e3, f2, f3 and h1 are non-ﬂagellate and h2 ﬂagellate. On the opisthosoma there is a distinct expansion anterior to coxa III, and longitudinal striae on the median portion are anterior to the second dorsocentral setae. A pair of opisthosomal pores is present. The ﬁrst and second pairs of dorsocentral setae are spatulate and the third is minute and serrate. Medioventral setae are present on the ventral surface (Meyer Smith, 1979; Ehara and Masaki, 2001).

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The body is about 250 μm long, similar to the female, with posterior opisthosoma smaller and narrower. The opisthosomal dorsum presents oblique striae on the anterior half. The dorsocentral setae are minute and slightly serrate (Ehara and Masaki, 2001). Geographical distribution France (Dugès, 1834); Italy (Canestrini and Fanzago, 1876); Greece (Hatzinikolis, 1986a); and Portugal (Ehara and Masaki, 2001). Gerson (2003) reports a worldwide distribution. Bio-ecology

The species has been described from specimens collected in France on laurestinus (Viburnum tinus Linnaeus) (Dugès, 1834). It is well documented on Acacia, Calligonum, Citrus, Laurus, Malus and Olea (Ehara and Masaki, 2001). Its bio-ecology is unknown. 11.4.1.2 Tenuipalpus emeticae2 Meyer (Fig. 11.17) Common name Unknown. Geographical distribution South Africa (Meyer Smith, 1979). Diagnostic characteristics FEMALE. The body is 260–280 μm long and 160–170 μm wide. The prodorsum has three pairs of marginal setae, including one pair of verticals (v2) and two pairs of scapular (sc1, sc2), setiform to lanceolate and ﬁnely serrate, with setae sc2 longest, and one pair of eyes per side. The mediodorsal area of the prodorsum is outlined by longitudinal striae and has transverse striae in this area; the lateral propodosoma possesses longitudinal to irregular striae. The dorsal opisthosoma presents transverse to irregular striae, and bears ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), setiform to lanceolate and ﬁnely serrate, with setae e3 shorter, setae h2 ﬂagelliform, and the other opisthosomal setae shorter than the distance between their bases, and one pair of opisthosomal pores. The ﬁrst medioventral setae are short and the posterior medioventral setae long. The ventral and genital plates are inconspicuously separated from each other. Tarsi I and II have one solenidi on each. The femur I with inner dorsal seta lanceolate, serrate and about a third of the length of

2Meyer

Smith (1993) reports that Tenuipalpus citri Meyer, 1979 is synonymous with T. emeticae, 1979.

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A

B

E

C

D

Fig. 11.17. Tenuipalpus emeticae Meyer. Female. (A) Dorsal view; (B) genitoventral regions; (C) palp; (D) femur I (from Meyer Smith, 1979). Tenuipalpus sanblasensis De Leon. Female. (E) Dorsal view (right half) (from De Leon, 1957).

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the segment; the inner distal seta is slightly shorter than the dorsal one (Meyer Smith, 1979). MALE.

Unknown (Meyer Smith, 1979).

Bio-ecology Meyer Smith (1979) has studied the morphology of six specimens (females) collected by Schwartz on Citrus sp. in South Africa, four specimens (females) collected on Trichilia emetica Vahl. and two specimens on an unidentiﬁed wild tree. The bio-ecology of the mite is unknown. 11.4.1.3 Tenuipalpus mustus Chaudhri Common name Unknown. Diagnostic characteristics FEMALE.

The body is 241 μm long, and 130 μm wide. The palp is threesegmented, with the distal segment bearing a rod, and the second a long, barbed seta. The rostral shield is deeply bifurcate with projections on each side. The prodorsum has transverse broken striations, with broken, thick lines laterally, and three pairs of setae including one pair of verticals (v2) and two pairs of scapular (sc1, sc2), with the ﬁrst and second pairs minute and the third 18 μm long and serrate, and one pair of eyes per side. The dorsal opisthosoma has irregular, broken, transverse striations, a few longitudinal striations in the caudal end. There are ten pairs of dorsal opisthosomal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2), all serrate and falling short of the distances between their bases, from 8 to 12 μm long, except h2, which is longer and ﬂagellate. The anterior medioventral setae are 13 μm long, and the posterior 47 μm, crossing the base of ventral plate setae. The ventral and genital plates are transverse, broken striate. The ventral plate bears one pair of simple setae (ag) and does not cross the base of genital plate setae. The genital plate bears two pairs of simple setae (g1, g2), and the anal plate also possesses two pairs of simple pseudanal setae (ps1, ps2) (Chaudhri, 1972b). MALE.

Unknown (Chaudhri, 1972b).

Geographical distribution Pakistan (Chaudhri, 1972b); India (Ghai and Shenhmar, 1984). Bio-ecology Tenuipalpus mustus has been collected on several host plants, including Citrus sp., in Pakistan (Chaudhri, 1972b) and India (Ghai and Shenhmar, 1984). Its bio-ecology is unknown.

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11.4.1.4 Tenuipalpus orilloi Rimando (Fig. 11.18) Common name Unknown. Diagnostic characteristics FEMALE. The length of the body is 296 μm, and the greatest width 187 μm. The prodorsum bears three pairs of setae (v2, sc1, sc2), of which the ﬁrst and second pairs are short and ovate, and the third pair long, broadly lanceolate. The median portion of the prodorsum is clearly deﬁned to form a shieldshaped structure, and the lateral areas have rather indistinct diagonal striae. The median area of the opisthosoma as far as the third dorsocentrals is strongly deﬁned, transverse striae occurring within this area. The opisthosoma bears ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2, f3, h1 and h2) and one pair of opisthosomal pores. The setae c1, c3, d1, d3, e1 and e3 are minute lanceolate, and e1 is longer than c1 and d1; the setae f2, f3 and h1 are ovate, and h2 whip-like and long (Manson, 1963a). MALE. The body is 238 μm long and the greatest width is 138 μm. Similar to the female, except that the shield on the hysterosoma is not quite as extensive (Manson, 1963a).

Geographical distribution Indonesia (Manson, 1963a) and the Philippines (Rimando, 1962a, b). Bio-ecology Tenuipalpus orilloi has been collected on different host plants, including Citrus sp., in the Philippines (Rimando, 1962a, b) and Spathiphyllum sp. in Indonesia (Manson, 1963a). We do not have enough information about its bio-ecology.

11.4.1.5 Tenuipalpus sanblasensis De Leon (Fig. 11.17) Common name Unknown. Diagnostic characteristics FEMALE.

The body is oval and red in colour. The palp is made up of three segments. The prodorsum bears three pairs of marginal setae, including one pair of verticals (v2) and two pairs of scapular (sc1, sc2), short setiform lanceolate and ﬁnely serrate, with setae sc2 lightly longer, and one pair of eyes per side; with a pair of L-shaped ridges back to back along the midline, both Ls sometimes broken at foot, the rest of the dorsum broken up by innumerable small ridges. The opisthosoma bears ten pairs of dorsal setae (c1, c3, d1, d3, e1, e3, f2,

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A

B

C

Fig. 11.18. Tenuipalpus orilloi Rimando. Female. (A) Dorsal view; (B) ventral view. Male. (C) Dorsal view (from Manson, 1963a).

f3, h1 and h2), short setiform to lanceolate and ﬁnely serrate, with setae h2 ﬂagelliform and with an oval area of small, more or less longitudinal ridges bounded anteriorly and posteriorly by stronger ridges and laterally by the pores; the rest of the dorsum is broken up by small ridges. One pair of

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anterior and one pair of posterior medioventral setae. The leg setae are setiform and small. Coxa III bears a seta on the anterior side. Genua I and II each have a seta on the anterior and posterior side; genua III and IV are bare; tarsi I and II possess a posterodistal solenidion and an overlying normal seta (De Leon, 1957). MALE.

Unknown (De Leon, 1957).

Geographical distribution Mexico (De Leon, 1957). Bio-ecology Tenuipalpus sanblasensis has been collected on ‘naranjilla’ (Solanum quitoense Lam.) in Mexico (De Leon, 1957), and Gerson (2003) has also reported it on citrus. Its bio-ecology is unknown.

11.4.2 Ultratenuipalpus Mitrofanov The palp is four-segmented. The prodorsum bears a rostral shield, three pairs of setae (v2, sc1, sc2) and two pairs of eyes per side. The dorsal opisthosoma possesses ten pairs of setae (c1, c2, c3, d1, d3, e1, e3, f3, h1, h2). The ano-genital region is not differentiated into plates. The genital plate is not demarcated and has three pairs of setae (g1, ag1, ag2), and the anal plate has one pair of anal setae and two pairs of pseudanal setae (ps1, ps2) (Meyer Smith, 1979). 11.4.2.1 Ultratenuipalpus gonianensis Sadana et Sidhu (Fig. 11.19) Common name Unknown. Diagnostic characteristics FEMALE. The body is 216 μm long and 132 μm wide. The palp is four-segmented,

the distal with a sensory rod and two setae. The rostral shield reaches up to the middle of femur I. The propodosoma is bare, with a few striations at the posterolateral corners and has three pairs of setae (v2, sc1, sc2), 14, 14 and 15 μm long, respectively. The hysterosoma bears ten pairs of setae; it is bare and has a few faint transverse striations close to setae d1 and e1; setae c3 are serrate and 14 μm long; setae c1, d1 and e1 are 12, 3 and 13 μm long, respectively, whereas c1 and e1 are serrate and d1 is simple. Setae c2 are 2 μm long, d3 15 μm, e3 14 μm, f3 15 μm, h2 14 μm and h1 15 μm long. All setae except d1 are serrate and spatulate. The venter of the propodosoma and hysterosoma have a ﬁngerprint-like pattern throughout. Ventral propodosomal setae 1a are long and ﬁliform, 26 μm long; the anterior medioventral metapodosomal

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C

B

Fig. 11.19. Ultratenuipalpus gonianensis Sadana et Sidhu. Female. (A) Dorsal (left half) and ventral (right half) view; (B) prodorsal shield; (C) palp (from Sadana and Sidhu, 1990).

setae 3a are minute, 6 μm long; ventral plate along with its setae absent; the genital plate is not demarcated and has three pairs of setae (g1, ag1, ag2); the anal plate has one pair of minute anal setae, 3 μm long, and two pairs of pseudanal setae (ps1, ps2) 3 μm long. Tarsi I–IV have one solenidion each (Sadana and Sidhu, 1990). MALE.

Unknown (Sadana and Sidhu, 1990).

Geographical distribution India (Sadana and Sidhu, 1990). Bio-ecology The species has been collected on C. limon (Sadana and Sidhu, 1990). Its bio-ecology is unknown.

12

Tuckerellidae Baker et Pritchard

12.1 INTRODUCTION The Tuckerellidae number few species and are commonly plant-feeding mites. Some species have been collected from moss on boulders bordering a saltwater harbour soil (Collier, 1969), others from the roots of plants (Baker and Tuttle, 1975; McDaniel et al., 1975) from on aerial habitats (Jorgensen, 1967; Zaher and Rasmy, 1969; Ehara, 1975; Jeppson et al., 1975; McDaniel et al., 1975; Meyer Smith, 1981; Meyer Smith and Ueckermann, 1997), feeding on some 25 plant families. Four species have been recorded on citrus worldwide (Table 11.1), but their damage potential is doubtful and unclear. According to Gerson (2003), they do not cause a consistent economic level of injury nor require control measures. Until recently these species were considered only potential pests, but Ochoa et al. (1994) report that Tuckerella pavoniformis (Ewing) and Tuckerella knorri (Baker et Tuttle) are serious pests in Central America for fruit and citrus plants.

12.2 MORPHOLOGICAL CHARACTERS AND SYSTEMATIC OUTLINE The body of the Tuckerellidae is made up of the propodosoma and hysterosoma. The dorsal surface presents the prodorsum, corresponding to the anterodorsal part of the propodosoma, with the opisthosoma formed from the hysterosoma, excluding legs III and IV. The prodorsum bears four pairs of setae described as vertical internal (vi), vertical external (ve), scapular internal (sci) and scapular external (sce) (Fig. 12.1). The opisthosoma possesses 36 pairs of dorsal palmate setae, with ﬁve or six pairs of ﬂagelliform caudal setae set on four distinct dorsal plates (C, D, EF, H). The ﬁrst plate (C) © V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

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vi e

cI

ve sci sce

c4

c II

c1

c2

c3

ic1 P

C

c5

c6

ic3

c7

c III d1

d2

D

d3 d5

d4

e2

e3 e4

f2

c IV ic4

e1 f1

EF

h6 h4 h2 h8 h7 h5 h3 h1 H

Fig. 12.1. Tuckerella ornata (Tucker). Diagram of dorsal (left) and ventral (right) chaetotaxy of the female idiosoma (according to Quirós Gonzalez and Baker, 1984); e, eyes; C I-IV, coxae; C, D, EF, H, dorsal plates; P, prodorsum. The setal notation is explained in the text. The pores are not indicated.

possesses two pores and a setal row (c1–c7); the second plate (D) two pores and a setal row (d1–d5); the third plate (EF) four pores and two setal rows (e1–e4) and (f1–f2) and the last plate (H) two pores and a caudal row of setae (h1–h5 + h7 or h1–h8). Setal rows c, d and e form an ‘L’ pattern, whereas rows f and h form a transverse pattern. The palp is elongate and bears a thumb– claw complex. The distal end of the peritreme is similar to that of tetranychids (Fig. 12.2). The female genital area possesses two pairs of aggenital setae (ag1, ag2), four pairs of genital setae (g1–g4) and three pairs of pseudanal setae (ps1–ps3) (Fig. 12.2). The male has only one pair of genital setae (g1) (Fig. 12.2). Both sexes have three pairs of intercoxal setae (ic1, ic3, ic4). The coxa possesses 2–2–1–1 setae, the trochanter 1–1–2–1 setae, the femur 7–7–2–1

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A

B

C

D

ag1

g4

E

ag1

g3 ag2 g2 g1 ag2

ps3

g1

ps3

ps2 ps2 ps1 ps1

Fig. 12.2. Tuckerella pavoniformis (Ewing). (A) Palp; (B) distal end of tarsus I; (C) distal end of peritreme (from McGregor, 1950). Tuckerella nilotica Zaher et Rasmy; (D) genito-anal region of female (from Rasmy and Abou-Awad, 1984, partially modiﬁed); (E) diagram of genito-anal region of male of Tuckerella ornata (Tucker) (according to Quirós Gonzalez and Baker, 1984). The setal notation is explained in the text.

setae, the genu 5–4–1–0 and the tibia 8–5–5–4 setae; tibia I has one dorsal solenidion. Tarsi I–IV bear 14, 11, eight and eight tactile and sensorial setae, respectively. The pretarsi has two true claws and and a pad-like empodium with tenent hairs (Fig. 12.2) (Quirós-Gonzalez and Baker, 1984). The Tuckerellidae family belongs to the Tetranychoidea superfamily. It is monogeneric and hosts the single genus Tuckerella Womersley, with more than 20 species worldwide (Meyer Smith and Ueckermann, 1997).

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Table 4.1 contains information of a number of websites displaying images of living mites with their colours and natural features, in addition to details of the damage they cause on citrus.

12.3 TUCKERELLA WOMERSLEY According to Meyer Smith and Ueckermann (1997), the diagnostic characters of the family can be applied for the genus Tuckerella.

12.3.1 Tuckerella knorri Baker et Tuttle (Fig. 12.3) Common name Ornamented mite.

D

A

B

C

Fig. 12.3. Tuckerella knorri Baker et Tuttle. Female. (A) Dorsal view; (B) ventral view of marginal opisthosomal seta; (C) dorsal view of marginal opisthosomal seta (from Baker and Tuttle, 1975); Tuckerella pavoniformis (Ewing). Adult. (D) Dorsal view (from Jeppson et al., 1975).

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Diagnostic characteristics FEMALE. The body is oval, about 400 μm long and 200 μm wide, colour in life red with white setae and well reticulated. The prodorsal setae ve are about two-thirds the length of setae vi. The opisthosoma has setae f1–2 subequal in length, setae c4 about half as long as the distances between their bases and those of the setae sce and six pairs of ﬂagellate caudal setae; the posterior opisthosoma presents irregular reticulations. The distal solenidion on tarsus I is longer than the proximal solenidion (Baker and Tuttle, 1975). MALE.

Unknown (Baker and Tuttle, 1975).

Geographical distribution China (Lin, 1982); Costa Rica (Ochoa, 1989a, 1994); Iran (Kamali, 1990); the Philippines (Corpuz Raros, 1989); Thailand (Baker and Tuttle, 1975). Bio-ecology Baker and Tuttle (1975) described the ornamented mite on specimens collected on Pandanus odoratissimus Linnaeus and Achras zapota (Linnaeus) in Thailand. In Costa Rica, T. knorri has been collected on Citrus limon (var. Mesina), Citrus sinensis and Mangifera indica Linnaeus (Ochoa, 1989a, 1994). Its life cycle develops through three nymphal stages and it has been veriﬁed that the species is paedogenetic, with the eggs emerging from the anal region in mounted tritonymphs (Ochoa, 1989b). On lemon, the mite generally lives on branches, and it feeds and moves in small depressions or cracks where it is also protected. It uses the crevices for feeding, protection and oviposition (Ochoa et al., 1994). Ochoa (1989a) has reported that when predatory mites approach T. knorri, it moves its ﬂagellate setae forward. Its bio-ecology is unknown. Symptomatology and damage In Costa Rica, T. knorri is a serious pest of citrus and occurs in association with the fungus Sphaceloma fawcettii (Ochoa, 1989a). The mite is considered a causative agent of the cracking of citrus fruits (Aguilar and Gonzalez, 1990); it produces irregular cracking of the epidermis of fruit where the fungus S. fawcettii is present and changes the light brown colour of the fungus to brownish-black. When the fungus is widespread, the ornamented mite is common around the lesions, occasionally changing the colour of the fungus to reddish-brown. The combination of T. knorri and the fungus causes damage that is more severe than that of the fungus and Brevipalpus phoenicis (Ochoa, 1989a). On lemon, a heavy infestation of T. knorri caused a signiﬁcant reduction in yield in Costa Rica. The mite also attacks C. sinensis (var. Valencia), and occasionally the symptoms are mistaken for fungal disease (Ochoa et al., 1994).

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Control Insufﬁcient information is available, except in Costa Rica where there is need for control.

12.3.2 Tuckerella nilotica Zaher et Rasmy (Figs 12.2, 12.4) Common name Unknown.

A

B

Fig. 12.4. Tuckerella ornata (Tucker). Female. (A) Dorsal view (from Baker and Pritichard, 1953a). Tuckerella nilotica Zaher et Rasmy. Female. (B) Dorsal view (from Rasmy and Abou-Awad, 1984).

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169

Diagnostic characteristics FEMALE. The body is oval and red in colour, about 400 μm long and 200 μm wide. The dorsum is reticulate with typical fan-shaped or palmate setae. The prodorsum has four pairs of palmate setae and the opisthosoma has 18 pairs of palmate setae, seven laterals and 11 in four transverse rows (four, three, two and two); the two posterior opisthosomal are the smallest, with the outer pair subequal to the inner, the last four lateral large and elongate. The caudal region bears six pairs of long, ﬂagellate setae as long as the body, arising from dorsal tubercles arranged in a straight line and extending caudally. There are two small foliaceous setae in a mediocaudal position between the long, ﬂagellate setae and a similar single seta between the second and third ﬂagellate setae on either side. Tarsus I has the anterior solenidion slightly longer than posterodistal; tarsus II with anterior solenidion very much longer than posterior (Rasmy and Abou-Awad, 1984). MALE.

Unknown (Rasmy and Abou-Awad, 1984).

Geographical distribution Egypt (Zaher and Rasmy, 1969; Rasmy and Abou-Awad, 1984). Bio-ecology Tuckerella nilotica has been collected on the fruit and buds of orange trees in Egypt (Zaher and Rasmy, 1969; Rasmy and Abou-Awad, 1984). Its life cycle evolves by three nymphal stages, and the bio-ecology is unknown.

12.3.3 Tuckerella ornata (Tucker) (Figs 12.1, 12.4) Common name Unknown. Diagnostic characteristics FEMALE. The body is oval and dark red in colour, about 400 μm long and 200 μm wide. The dorsum is reticulate, the prodorsum has four pairs and the opisthosoma 18 pairs of leaf-shaped and silvery-white setae; the caudal opisthosoma bears ﬁve pairs of ﬂagellate caudal setae; opisthosomal setae f2 are situated slightly anterior of f1, and f1–2 are about equal in length (Meyer Smith, 1981; Meyer Smith and Ueckermann, 1997). The last four leaf-shaped prodorsal setae with the outer pairs smaller than the inner pair and placed caudally of the inner pair. The anterodistal solenidion on tarsus I is as long as but more slender than the posterodistal solenidion. Tarsus II bears a slender solenidion anterodistally. Leg IV has slender setae dorsally (Baker and Pritchard, 1953a).

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Chapter 12 MALE.

Unknown (Baker and Pritchard, 1953a).

Geographical distribution The mite has a worldwide distribution and is recorded for Brazil (Flechtmann, 1979); California (USA) (Baker and Pritchard, 1953a); China (Lin, 1982); Costa Rica (Ochoa, 1989a); Florida (Baker and Pritchard, 1953a; De Leon, 1955); Guatemala (Baker and Pritchard, 1953a); Hawai (Garrett and Haramoto, 1967); Kenya (Meyer Smith and Ueckermann, 1997); Namibia (Meyer Smith and Ueckermann, 1997); the Philippines (Baker and Pritchard, 1953a; Corpus Raros, 1989); Reunion Island (Meyer Smith and Ueckermann, 1997); South Africa (Tucker, 1926; Baker and Pritchard, 1960; Meyer Smith and Ueckermann, 1997); Thailand; Venezuela (Quirós Gonzalez and Baker, 1984); and Zimbabwe (Meyer Smith and Ueckermann, 1997). Bio-ecology The species was originally collected by Tucker (1926) on orange fruits in the Western Cape Province and Transvaal. Successively, it has been recorded by Baker and Pritichard (1960) on citrus from Natal and by Meyer Smith (1981) and Meyer Smith and Ueckermann (1997) in South Africa on various plants, including C. sinensis and C. limon. De Leon (1955) reported the mite in Florida on Citrus mitis and Citrus maxima (Burman) Merril, and Ochoa (1989a) in Costa Rica on the fruit of C. limon. With regard to natural enemies, the phytoseiid mite Amblyseius largoensis has been collected in association with T. ornata in the Philippines (Corpus Raros, 1989). The population density is generally low and the injury caused is unclear (Meyer Smith, 1981). Tucker (1926) has reported that the mite is probably associated with scarring of citrus fruit. Control is not usually required.

12.3.4 Tuckerella pavoniformis (Ewing) (Figs 12.2, 12.3) Common name Peacock spider mite. Diagnostic characteristics FEMALE. The body is oval, red with whitish-orange setae, about 400 μm long and 200 μm wide. The dorsum is reticulate, the prodorsum has four pairs of setae and the opisthosoma 18 pairs of foliaceous or palmate setae. The caudal region of the opisthosoma possesses six pairs of long ﬂagellate and pectinate setae. The prodorsum has the last pair of lateral palmate setae (sce) larger than inner pair (sci) and anterior pair (vi) wide as long. The anterodistal solenidion of tarsus I is very short in comparison to the posterodistal solenidion. Tarsus II bears one short solenidion (Baker and Pritchard, 1953a). MALE.

Unknown (Baker and Pritchard, 1953a).

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171

Geographical distribution The mite has a worldwide distribution and has been recorded in Australia (Womersley, 1940); Brazil (Mineiro et al., 2005); California (McGregor, 1950; Baker and Pritchard, 1953a); Costa Rica (Ochoa, 1989a); Florida (Jeppson et al. 1975); Georgia (Reck, 1959); Hawai (Ewing, 1922); Mauritius (Jeppson et al., 1975); Formosa; and Japan (Ehara, 1966). Bio-ecology The peacock spider mite has been collected on different cultivated plants and on citrus (McGregor, 1950; Jeppson et al., 1975). Like T. knorri, it lays its eggs in the cracks of host plants, where adults and other mobile forms hide (Ochoa et al., 1994). Its bio-ecology is unknown. The mite has never been associated with serious injury to citrus. It has been recorded in Central America as a pest on fruits and citrus plants (Ochoa et al., 1994). The need for control is not widespread and at present seems to arise only in Central America (Ochoa et al., 1994).

13

Tetranychidae Donnadieu

13.1 INTRODUCTION Tetranychidae mites are commonly known as ‘spider mites’ because of the tendency of many species to make webs. They are typically phytophagous and include species more harmful to cultivated plants or those of economic importance. Many species are polyphagous. The family has a worldwide distribution and includes about 1250 species belonging to 73 genera (Pritchard and Baker, 1955; Gutierrez, 1985b; Meyer Smith, 1987; Bolland et al., 1998; Migeon and Flechtmann, 2004; Migeon and Dorkeld, 2006). Sixty species have been recorded on citrus in different regions of the world (Tables 13.1–13.3), some of which have been accidentally collected and do not cause any injuries to crops (Bryobia praetiosa Koch, Aplonobia citri Meyer, Aplonobia histricina (Berlese), Aplonobia honiballi Meyer, Petrobia tunisiae Manson, P. harti (Ewing), etc.), whereas other species are distinctly injurious. Sometimes, the diffusion of some species depends on their ability to live under precise ecological conditions or through worldwide distribution, strong polyphagy or remarkable survival capacities that determine a relevant phytopathological problem (Panonychus citri, Tetranychus urticae, Eutetranychus orientalis, etc.).

13.2 MORPHOLOGICAL CHARACTERS AND SYSTEMATIC OUTLINE The body of the Tetranychidae is soft and more or less ovoid or round. It is from 350 to 1000 μm long, variously coloured (red, orange, green or yellow) and formed of a gnathosoma and idiosoma. The dorsal disjugal and the 172

© V. Vacante 2010. Citrus Mites: Identiﬁcation, Bionomy and Control (V. Vacante)

Tetranychidae Donnadieu Table 13.1.

173

Mites of the family Tetranychidae Donnadieu collected on citrus worldwide. Pest status

Distribution

Bryobia graminum (Schrank) Bryobia praetiosa Koch Bryobia rubrioculus (Scheuten) Aplonobia citri Meyer Aplonobia honiballi Meyer Aplonobia histricina (Berlese) Petrobia harti (Ewing) Petrobia latens (Müller) Petrobia tunisiae Manson Tenuipalponychus citri Channabasavanna et Lakkundi Aponychus chiavegatoi Feres et Flechtmann Aponychus spinosus (Banks)

U U U U U U U U U U

Worldwide Worldwide Worldwide Australia, South Africa South Africa Australia, Israel, Italy, South Africa Worldwide Worldwide Iran, Israel, Italy, Spain, Tunisia India

U

Brazil

U

Eutetranychus africanus (Tucker)

Mi

Eutetranychus banksi (McGregor)

Ma

Eutetranychus citri Attiah Eutetranychus cratis Baker et Pritchard Eutetranychus eliei Gutierrez et Helle Eutetranychus orientalis (Klein) Eutetranychus pantopus (Berlese) Eutetranychus pyri Attiah Meyernychus emeticae (Meyer) Acanthonychus jiangfengensis Wang

Mi U U

Brazil, Canada, Paraguay, Philippines, USA Australia, Comoros, Egypt, India, Japan, Madagascar, Mauritius, Mozambique, Myanmar Burma, Papua New Guinea, Philippines, Réunion Island, South Africa, Thailand Egypt, Hawaii, India, North, Central and South America, Portugal, Spain Egypt, India Republic of the Congo, Congo (RDC, ex-Zaïre), Nigeria Madagascar

Ma U Mi Mi U

Worldwide Australia, Egypt, Sudan Egypt Angola, South Africa China

Species

According to Gerson (2003), the pest status is indicated as Ma (major), Me (medium), Mi (minor) or U (unknown). The references are indicated in the text.

ventral sejugal sutures divide the idiosoma into an anterior propodosoma and a posterior hysterosoma. The dorsal surface of the body presents the prodorsum, corresponding to the anterodorsal part of the propodosoma, and the opisthosoma, identiﬁable with the hysterosoma, excluding legs III and IV (Fig. 2.1). The gnathosoma has a stylophore and two chelicerae (Fig. 2.4). The latter possess a long and evenly thick digitus mobilis, except at the distal end where it is pointed and, if protracted and juxtaposed with that of the other side,

174 Table 13.2.

Chapter 13 Mites of the family Tetranychidae Donnadieu collected on citrus worldwide.

Species

Pest status Distribution

Eotetranychus cendanai Rimando

Mi

Eotetranychus kankitus Ehara Eotetranychus lewisi (McGregor)

Mi Mi

Eotetranychus limonae Karuppuchamy et Mohanasundaram Eotetranychus limoni Blommers et Gutierrez Eotetranychus mandensis Manson Eotetranychus pamelae Manson Eotetranychus sexmaculatus (Riley)

U

Cambodia, China, Philippines, Taiwan, Thailand China, India, Japan Bolivia, Chile, Colombia, Costa Rica, El Salvador, Guatemala, Hawaii, Honduras, Libya, Madeira Island, Mexico, Nicaragua, Panama, Peru, South Africa, Taiwan, USA India

Mi

Madagascar

U U Ma

Eotetranychus yumensis (McGregor) Mixonychus ganjuis Qian et al. Mixonychus ziolanensis (Lo et Ho) Oligonychus biharensis (Hirst) Oligonychus coffeae (Nietner) Oligonychus gossypii (Zacher)

Mi U U U U U

Oligonychus peruvianus (McGregor)

Mi

Panonychus citri (McGregor) Panonychus elongatus Manson

Ma Mi

Panonychus ulmi (Koch) Schizotetranychus baltazari Rimando

U Mi

Schizotetranychus industanicus (Hirst) Schizotetranychus lechrius Rimando Schizotetranychus spiculus Baker et Pritchard Schizotetranychus youngi Tseng

Mi U U

India India Australia, China, Formosa, Hainan Island, Hawai, Korea (Rep. South), India, Iraq, Japan, Korea, New Zealand, Okinawa Island, Taiwan, USA Mexico, USA China Taiwan Worldwide Worldwide Angola, Benin, Brazil, Cameroun, Central Africa Rep., Colombia, Congo, Congo (Rep. of the) (RDC, ex-Zaire), Costa Rica, Ecuador, Ethiopia, Guinea-Bissau, Honduras, Kenya, Madagascar, Nigeria, Sao Tome and Principe, Senegal, Sierra Leone, Tanzania, Togo, Uganda, Venezuela Colombia, Costa Rica, Ecuador, Guatemala, Mexico, Peru, Trinidad and Tobago, USA, Venezuela Worldwide Australia, China, Korea, Myanmar (Burma), Papua New Guinea, Taiwan, Thailand Worldwide China, Hong Kong, India, Indonesia, Myanmar (Burma), Philippines, Taiwan, Thailand India Indonesia, Philippines, Taiwan India, Kenya, Congo (RDC, ex-Zaire)

U

Taiwan

According to Gerson (2003), the pest status is indicated as Ma (major), Me (medium), Mi (minor) or U (unknown). The references are indicated in the text.

Tetranychidae Donnadieu Table 13.3.

175

Mites of the family Tetranychidae Donnadieu collected on citrus worldwide. Pest status

Distribution

Tetranychus desertorum Banks Tetranychus ﬁjiensis Hirst

U

Worldwide

Mi

Tetranychus gloveri Banks

Mi

Tetranychus kanzawai Kishida Tetranychus lambi Pritchard et Baker

Mi

Australia, Carolina Islands, China, Fiji, Hainan Island, India, Kiribati, Malaysia, Marianas Northern, Marshall Islands, Micronesia Federated States, New Caledonia, Papua New Guinea, Philippines, Seychelles, Sri Lanka, Taiwan, Thailand American Samoa, Australia, Bermuda, Brazil, Colombia, Costa Rica, Cuba, French Polynesia, French West Indies, Greece, Guadeloupe, Guam Island, Hawaii, Honduras, Les Saintes, Marianas Northern, Mexico, Panama, Paraguay, Peru, Puerto Rico, Samoa (American), Suriname, Trinidad and Tobago, USA, Venezuela Worldwide

Tetranychus ludeni Zacher Tetranychus mexicanus (McGregor)

Mi Mi

Tetranychus neocaledonicus André Tetranychus paciﬁcus McGregor Tetranychus paraguayensis Aranda Tetranychus salasi Baker et Pritchard Tetranychus taiwanicus Ehara Tetranychus tumidus Banks

Mi

Australia, Cook Islands, Fiji, French Polynesia, Iran, New Caledonia, New Zealand, Papua New Guinea, Samoa (American), Samoa (Western), Taiwan, Tasmania, Tonga, Vanuatu, Wallis and Futuna Worldwide Argentina, Brazil, China, Colombia, Costa Rica, Cuba, El Salvador, Guadeloupe, Honduras, Les Saintes, Mexico, Nicaragua, Paraguay, Peru, USA, Uruguay, Venezuela Worldwide

Mi

Canada, Mexico, USA

U

Paraguay

U

Costa Rica, Nicaragua

Mi

China, Hainan Island, Taiwan, Thailand

Mi

Tetranychus turkestani (Ugarov et Nikolski) Tetranychus urticae Koch

U

Colombia, Cuba, Greece, Guam, Panama, Puerto Rico, Thailand, USA Worldwide

Ma, Mi

Worldwide

Species

U

According to Gerson (2003), the pest status is indicated as Ma (major), Me (medium), Mi (minor) or U (unknown). The references are indicated in the text.

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forms an empty tube. The cheliceral bases are consolidated together, curving ventrally form to the stylophore, which is capable of protraction and retraction and receiving at the base two stigmata associated with two peritremes (Fig. 2.4) leading anteriorly to a membranous prodorsum. The lower surface of the stylophore matches, slipping to the length, the posterior medial surface of the infracapitulum, conformed as a longitudinal hollow. The ventral surface of the infracapitulum shrinks to the distal end and constitutes the buccal cone, which contains the mouth. The palp has four to ﬁve segments, including the ‘thumb–claw’, and possesses simple setae, eupathidia and solenidia (Fig. 2.4) (Lindquist, 1985). The prodorsum bears three or four pair of setae named v1 (internal vertical), v2 (external vertical), sc1 (internal scapulars) and sc2 (external scapulars), and between the scapular setae are two eyes per side. The integument can be striate, dotted or reticulate, or furnished by one to four projections ornate with setae (subfamily Bryobiinae). The setae may be arranged on tubercles. The opisthosoma bears up to 14 pairs of dorsal setae, associated with segments C (c1–3), D (d1–3), E (e1–3), F (f1–2) and H (h1–3). Except for the setae h2–3 the opisthosomal setae are generally similar to the prodorsals; they can be ﬁnely barbulate, pointed, setiform, clavate, spatulate or lanceolate, all or partially set on tubercles. The integument is commonly similar to that of the prodorsum. The opisthosomal ventral region of the female has one pair of aggenital setae (ag), the genital opening, two pairs of lateral genital setae (g1–2) and the anal opening, with one to three pairs of pseudanal setae (ps) (Fig. 2.2). The oriﬁce of the bursa copulatrix is located between the anal and genital openings, joined within through a thin duct to the receptaculum seminis (Fig. 2.5). Between the genito-anal valvae is found the male aedeagus (Fig. 2.5); the shape and size of this are used in systematics for the identiﬁcation of species (Lindquist, 1985). The ventral podosomal region commonly bears three pairs of simple or intercoxal ventral setae (1a, 3a and 4a). The legs are made up of ﬁve segments (trochanter, femur, genu, tibia and tarsus), ﬁrst pair bearing, dorsally to the coxae, the supracoxal setae e. Commonly, the maximum number of coxisternal setae for the four legs is 2–2–1–1. The trochanter bears 1–1–1–1 setae. On the femur and genu are found additional setae, and it is not possible to determine the correct notation of the two segments. The standard condition of chetotaxy of the tibia of the Tetranychinae subfamily is equal to 9 (+ one solenidion)–7–6–7, whereas in the Bryobiinae subfamily it ranges from 8 (+ one solenidion)–5–5–5 to 15 (+ 15 solenidia)–11–11–11, with some variation in males. The chetotaxy of the tarsus of Tetranychinae is variable and ranges from between 15 (+ three solenida)–14 (+ two solenidia)–9 or 10 (+ one solenidion)–10 (+ one solenidion) to 13 (+ three solenidia)–11 (+ two solenidia)–9 (+ one solenidion)–9 (+ one solenidion) and in the Bryobiinae there is greater variation; tarsi I and II bear the duplex setae (Fig. 2.2) formed by a solenidion and a tactile seta. The ambulacrum consists of a short, ﬂexible pretarsus (Fig. 2.6), consisting of a pair of lateral structures deriving from the true claws and a median empodium. In different systematic categories these structures may change (Lindquist, 1985).

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The family is divided into the Bryobiinae Berlese and Tetranychinae Berlese subfamilies; the former includes the Bryobiini Reck, Hystrichonychini Pritchard et Baker and Petrobiini Reck tribes, and the latter Eurytetranychini Reck, Tenuipalpoidini Pritchard et Baker and Tetranychini Reck tribes (Gutierrez, 1985b; Meyer Smith, 1987; Bolland et al., 1998). Of the tetranychid mites recorded for citrus worldwide, nine species are ascribed to the Bryobiinae subfamily and Bryobiini, Hystrichonychini and Petrobiini tribes, and 51 species to the subfamily Tetranychinae and Tenuipalpoidini, Eurytetranychini and Tetranychini tribes. Moreover, it is possible that the species Eotetranychus aurantii (Targioni Tozzetti) – according to the etymology of its name – could be included among those collected over time on citrus, even if formally it has been collected on Quercus ilex Linnaeus (Targioni Tozzetti, 1878; Berlese, 1886). Irrespective of this, its identity could be better investigated (Migeon and Flechtmann, 2004). Table 4.1 contains the addresses of a number of web sites, which contain images of living mites with their colours and natural features and their damages on citrus.

13.3 BRYOBIINAE BERLESE The empodium has tenent hairs; the female has two or three pairs of pseudanal setae (ps1–3) and the male ﬁve pairs of genito-pseudanal setae (ps1–3, g1–2) (Bolland et al., 1998).

13.3.1 Bryobiini Reck The true claws are uncinate, and the empodium pad-like (Bolland et al., 1998).

13.3.1.1 Bryobia Koch The prodorsum bears four pairs of setae, and prominent lobes over the gnathosoma. The opisthosoma has 12 pairs of setae. Setae h2–3 are present. Empodia II–IV have more than one pair of tenent hairs (Bolland et al., 1998). 13.3.1.1.1 Bryobia graminum (Schrank) (Fig. 13.1)1 Common name Unknown.

1The

species has been redescribed by van Eyndhoven (1957) as Bryobia cristata Dugès and then by Livshitz and Mitrofanov (1971) as B. graminum.

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C B

D E

F

Fig. 13.1. Bryobia graminum Schrank. Female. (A) Dorsal view; (B) prodorsal projections; (C) dorsal seta. Male. (D) Dorsal view; (E) aedeagus; (F) pretarsus I (partially modiﬁed from van Eyndhoven, 1957, as Bryobia cristata Dugès).

Diagnostic characteristics FEMALE. The body is about 800 μm long and dark red coloured. The external prodorsal projections present a large base, are conic-like and with the inner side outside. The inner and outer projections are well separated. The inner

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projections are obtuse (see Fig. 13.1). The apex of setae of prodorsal projections reach the same line. The peritremes end in a well-deﬁned anasthomosis. All dorsal setae are set on tubercles and are spatulate serrate and longer than they are wide. Pretarsus I has two true claws bearing one pair of tenent hairs each and the median empodium possesses two tenent hairs. Tarsi III and IV bear two duplex setae, with the solenidon longer than the tactile seta (van Eyndhoven, 1957). MALE.

The male is about 425 μm long, with the inner projections fused with each other and the outer projections of the same shape and size. The dorsal chaetotaxy is equal to that of the female. The peritreme is small, with few anasthomoses and is bulbous. The dorsal setae are long and serrate. The aedeagus is long and lanceolate (van Eyndhoven, 1957). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology

Bryobia graminum has been recorded on 27 different host plants (Migeon and Dorkeld, 2006), herbaceous and fruit plants, including Prunus dulcis Miller, Pyrus communis Linnaeus and Citrus aurantium (Bolland et al., 1998; Migeon and Dorkeld, 2006). In Germany, it is occasionally injurious to apples and pears in the late summer. The species overwinters as eggs or in all stages in the lower parts of trunks of apple and pear trees, or among lichens or fences near the plants. The hatching of eggs starts in late March, and the ﬁrst generations feed on grasses and the summer generations on leaves of fruit trees. This latter generation develops in 27–57 days, longevity ranges from 18 to 103 days and fecundity from 30 to 35 eggs. The mite develops seven generations per year. Another biological strain lives only on grasses (Gäbele, 1959). Its presence on citrus is occasional.

13.3.1.1.2 Bryobia praetiosa Koch (Fig. 13.2) Common name Clover mite. Diagnostic characteristics FEMALE. The body is 750–800 μm long and is dark red in colour, sometimes greenish, with lighter edges and legs. The prodorsal projections are subtriangular, well separated from each other, with the central higher than medial and the base wider than it is high; the medial projections are higher than they are wide. All dorsal setae are arranged on tubercles, spatulate serrate and longer than they are wide. The peritremes end in an anasthomosis about 50 μm long. The pretarsus I bears two true claws, with a pair of tenent hairs each and a

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C B A

D

E

F

G

Fig. 13.2. Bryobia praetiosa Koch. Female. (A) Prodorsal projections; (B) hysterosomal dorsal seta; (C) pretarsus I; (D) palp; (E) distal anasthomosis of peritreme; (F) duplex setae of tarsus III; (G) duplex setae of tarsus IV (partially modiﬁed from Vacante, 1985).

median empodium with two tenent hairs. Tarsi III and IV possess two duplex setae, with the solenidion longer than the tactile seta (Vacante, 1985). MALE.

Unknown (Vacante, 1985).

Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The clover mite has been recorded on 265 host plants (Migeon and Dorkeld, 2006), herbaceous trees and also on trees such as Citrus reticulata, Citrus sinensis and Citrus sp. (McGregor, 1950, 1956; Jeppson et al., 1975; Meyer Smith, 1981; Vacante and Nucifora, 1986; Bolland et al., 1998; Migeon and Dorkeld, 2006). The species reproduces through parthenogenesis thelytokous and possesses a haploid condition of four chromosomes (Helle et al., 1970). The most favourable conditions for development occur with temperatures ranging

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between 21° and 24°C (Mori, 1961), and oviposition and hatching of eggs from 8° to 18°C. Development decreases below 7°C and increases above 18°C and a little hatching has been observed above 30°C (Meyer Smith, 1981). In warmer regions, the clover mite overwinters as egg, immature and adult stages. In spring the winter eggs hatch, becoming the spring generation, at the same time as the generation deriving from the eggs laid from overwintering females. Overwintered larvae and nymphs mature in spring and their progeny represents a delayed portion of the spring generation. The species oversummers in the egg stage, laid in cavities and cracks of the terrain and in the walls of dwellings or on the bark of trees; the hatching of eggs occurs when the temperature decreases (late September) and continues until early winter, when all stages overwinter (English and Snetsinger, 1957). With temperatures between −2° and 8°C the eggs may hatch after a week, and eggs overwintering with temperatures ranging between 18° and 24°C may hatch in 12–18 h. A small percentage of eggs laid in spring do not oversummer and hatch in early summer, giving continuous generations; the oversummered eggs laid in May and June may hatch in late summer or at the ﬁrst decrease in temperature; the peak of hatching may follow the ﬁrst decrease in temperature (Anderson and Morgan, 1958; Gäbele, 1959). On citrus the clover mite occasionally occurs as an egg laid at the base of the trunk or as immature and adult stages on leaves of lower foliage, but does not cause any damage. Sometimes it is confused with other tetranychids and incorrectly treated with acaricides. In Sicily (Italy), the mite has been collected on herbaceous plants (Vacante, 1985) and grasses under the canopy of citrus or on leaves of the lower foliage of lemon (Vacante and Nucifora, 1986) and orange trees. As regards the natural factors limiting the populations of the clover mite in Illinois (USA), the bdellid Bdella depressa Ewing alone represents over 75% of the agents of biological control of the mite (Snetsinger, 1956). In Tasmania, the coccinellid Stethorus vagans Blackburn is reported (Chazeau, 1979).

13.3.1.1.3 Bryobia rubrioculus (Scheuten) (Fig. 13.3) Common name Brown mite. Diagnostic characteristics FEMALE. The length of the body ranges from 500 to 650 μm and is dark in colour, sometimes greenish, with deeper patterns in the central area, and projections and legs light. The medial prodorsal projections are subtriangular, small and separated from the central projections, which are thin and higher. Both projections may have small pointed ridges. The dorsal setae are set on small tubercles, and are spatulate. The peritremes end with a small anasthomosis 20 μm long. Pretarsus I possesses two true claws with a pair of tenent hairs each and a median empodium with two tenent hairs. The

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D

E

F

Fig. 13.3. Bryobia rubrioculus (Scheuten). Female. (A) Prodorsal projections; (B) distal anasthomosis of peritreme; (C) pretarsus IV; (D) hysterosomal dorsal seta; (E) duplex setae of tarsus III; (F) duplex setae of tarsus IV (partially modiﬁed from Vacante, 1985).

solenidion and the tactile seta of tarsus III are about the same length; in tarsus IV the solenidion and the tactile seta are well separated, with the solenidion proximal and shorter than the tactile seta (Vacante, 1985). MALE.

Unknown (Vacante, 1985).

Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The brown mite has been recorded on 62 host plants (Migeon and Dorkeld, 2006), primarily arboreous, frequently on fruit trees (apple, pear, peach, deciduous fruits) and on C. reticulata (Bolland et al., 1998; Migeon and Dorkeld, 2006).

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The mite overwinters as winter eggs laid on the bark of spurs, along the undersides of twigs and forks of branches and on the side of the trunk and branches, commonly close together in folds and crevices. In spring the eggs hatch and the larvae migrate to buds where they attack the leaves, and hatching is completed before the fruit bloom, with best hatch at a temperature range of 19° to 26°C, and the percentage decreases as RH reaches 80%. The ﬁrst eggs hatch in 6–18 days (Anderson and Morgan, 1958; Georgala, 1958; Dosse, 1963). In Nova Scotia B. rubrioculus develops two generations per year and one partial third (Herbert, 1965). In Lebanon Dosse (1963) has observed the progress of four generations per year, and in South Africa the mean generation time varies from 22 to 33 days, with six generations being produced per year (Georgala, 1955). On citrus, the species is occasionally present but does not cause any damage.

13.3.2 Hystrichonychini Pritchard et Baker The claws and the empodium are pad-like (Bolland et al., 1998).

13.3.2.1 Aplonobia Womersley The prodorsum bears three pairs of setae, and is without lobes over the gnathosoma. The dorsal opisthosoma possesses ten pairs of setae, some or all set on strong tubercles. The opisthosomal seta f1 is set in normal dorsal position. Tarsus I has two sets of duplex setae, and the coxal formula does not exceed 4–3–2–2 (Bolland et al., 1998).

13.3.2.1.1 Aplonobia citri Meyer (Fig. 13.4) Common name Unknown. Diagnostic characteristics FEMALE.

The body is about 1000 μm long and dark red in colour. All dorsal setae are set on strong tubercles, serrate and expanded distally, about as long as or longer than the distances between their bases. The prodorsum is punctate in the central area and has longitudinal striae laterally. The opisthosoma is transversely striate. The peritremes end in well-developed anasthomosis. The receptaculum seminis is oval. Tarsus I bears 11 tactile setae and six solenidia proximal to the duplex setae; tarsus II possesses seven or eight tactile setae and one or two solenidia proximal to the duplex setae (Meyer Smith, 1974, 1987).

MALE.

Unknown (Meyer Smith, 1974, 1987).

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B

C D

Fig. 13.4. Aplonobia citri Meyer. Female. (A) Dorsal view; (B) distal anasthomosis of peritreme; (C) pretarsus I; (D) receptaculum seminis (from Meyer Smith, 1974, 1987).

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Geographical distribution Australia (Gutierrez and Schicha, 1983) and South Africa (Meyer Smith, 1974, 1987). Bio-ecology In South Africa, the mite has been collected, together with A. honiballi, on the trunk of C. sinensis covered by the eggs of two mite species (Meyer Smith, 1974). In Australia, it has been found on C. limon and Prunus persica (Linnaeus) Batsch (Gutierrez and Schicha, 1983). Its bio-ecology is unknown.

13.2.2.1.2 Aplonobia honiballi Meyer (Fig. 13.5) Common name Unknown. Diagnostic characteristics FEMALE. The body is about 1000 μm long and dark red in colour. All dorsal setae

are set on strong tubercles, tapering and reach about as far as the distances to the bases of consecutive setae or slightly shorter. The central area of the prodorsum is punctate and presents laterally longitudinal striae. The opisthosoma has mostly transverse striae. The peritremes end with an ovate anastomosis. The receptaculum seminis is broadly rounded. Numerous setae are borne proximal to duplex setae on both tarsi I and II (Meyer Smith, 1974, 1987). MALE.

The dorsal body setae are similar to those of the female and the dorsum presents four punctate areas, one on the prodorsum and three on the opisthosoma. The shaft of the aedeagus is long and ends in two short stylets (Meyer Smith, 1974, 1987). Geographical distribution South Africa (Meyer Smith, 1974, 1987). Bio-ecology

The mite has been collected on Dimorphotheca pluvialis (Linnaeus) Moench and Anthospermum dregei Sond. (Meyer Smith, 1987), and twice on the trunk of C. sinensis in South Africa (Meyer Smith, 1974, 1987). Its bio-ecology is unknown.

13.3.2.1.3 Aplonobia histricina (Berlese) (Fig. 13.6) Common name Unknown.

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B

C

E

G D

F

Fig. 13.5. Aplonobia honiballi Meyer. Female. (A) Dorsal view; (B) distal anasthomosis of peritreme; (C) pretarus I; (D) receptaculum seminis. Male. (E) Dorsal view; (F) distal anasthomosis of peritreme; (G) aedeagus (from Meyer Smith, 1974, 1987).

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B

A

D

C E

Fig. 13.6. Aplonobia histricina (Berlese). Female. (A) Dorsal view (from Jeppson et al., 1975); (B) distal anasthomosis of peritreme; (C) palp; (D) receptaculum seminis; (E) pretarsus I (from Vacante, 1985).

Diagnostic characteristics FEMALE. The body is dark red in colour and leg I is longer than the body. The dorsal body setae are lanceolate serrate and set on strong tubercles, except the prodorsal setae v2, and reach about as far as the distances between their bases. The prodorsum possesses two eyes per side arranged between the scapular setae. The peritremes end in an oval-shaped distal anasthomosis, and the receptaculum seminis is oval and smooth (Vacante, 1985). MALE. The male is smaller than the female, with leg I twice as long as the body,

and the aedeagus is long and thin (Vacante, 1985).

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Geographical distribution Australia (Berlese, 1910; Womersley, 1940); Israel (Ben-David et al., 2007); Italy (Vacante, 1985); and South Africa (Meyer and Ryke, 1959; Baker and Pritchard, 1960; Meyer Smith, 1974). Bio-ecology Aplonobia histricina has been collected on Sphaeralcea ambigua Gray (Migeon and Dorkeld, 2006), Oxalis pes-caprae Linnaeus (Baker and Pritchard, 1960; Ben-David et al., 2007), Oxalis sp. (Womersley, 1940), Pyrus communis (Baker and Pritchard, 1960) and an unknown host plant (Berlese, 1910). In Sicily (Italy), the mite is very common on Oxalis cernua Thumb. (Vacante, 1985; Vacante and Nucifora, 1986) under the canopy of citrus trees or surroundings. The species oversummers as eggs laid on the trunk and branches of citrus and fruit trees, on fences near the plants or on plastic irrigation tubes. The development of the host plant in the crops coincides with the hatching of the summer eggs and the start of winter generations. It is sometimes possible to observe immature and adult stages of the mite on the lower foliage of citrus plants, but they do not cause any damage. Similarly to B. praetiosa, the presence of the mite on citrus may induce an unjustiﬁed alarm resulting in chemical control.

13.3.3 Petrobiini Reck The claws are pad-like and the empodium uncinate (Bolland et al., 1998).

13.3.3.1 Petrobia Murray The dorsum bears not more than 15 pairs of dorsal body setae; on the venter there are three pairs of medioventral body setae. Setae h2–3 are set in a ventral position. Tarsus I has two or more sets of duplex setae. The empodium is curved distally with two rows of tenent hairs, and the true claw is devoid of hair-like processes (Bolland et al., 1998).

13.3.3.1.1 Petrobia harti (Ewing) (Fig. 13.7) Common name Unknown. Diagnostic characteristics FEMALE.

The body is rounded and dark red. Leg I is as long as the body and light. All dorsal setae are ﬁnely serrate, set on strong tubercles and are much

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A

B

C

D

Fig. 13.7. Petrobia harti (Ewing). Female. (A) Dorsal view; (B) receptaculum seminis (from Meyer Smith, 1987). Male. (C) Palp; (D) aedeagus (from Vacante, 1985).

longer than the distances between consecutive setae, with dorsal opisthosomal setae h1 about half as long as other dorsal opisthosomal setae (c1, d1, e1 and f1). The prodorsum is punctate in the central area. The peritremes end simply. The receptaculum seminis is oval to rounded and with an irregular outline (Vacante, 1985; Meyer Smith, 1987).

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Chapter 13 MALE.

The body is smaller than that of the female, lighter and with dorsal setae shorter than the distances between consecutive setae. Leg I is about three times as long as the body (Vacante, 1985; Meyer Smith, 1987). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology

Petrobia harti has been recorded on 46 spontaneous and cultivated host plants, including various species of the genus Oxalis and trees (Migeon and Dorkeld, 2006). It as been collected on Citrus grandis (Carmona, 1970), Citrus maxima (Rodrigues, 1968) and Citrus sp. (Ferragut et al., 1983; Garcia Marì et al., 1986). Reproduction is parthenogenetic arrhenotokous, and the male haploid number of chromosomes is two (Helle et al., 1970). According to Ehara (1969), the mite has been collected in different regions of the world on Oxalis and its neighbouring plants. In Sicily (Italy), the species overwinters as eggs laid on the trunk and branches of citrus trees, and hatching occurs when O. cernua develops in the vicinty; sometimes the motile forms of the mite are found in the lower foliage of citrus trees (Ciampolini et al., 1985). The same behaviour has been observed in Spain, where the species lives on O. pes-caprae in the lower canopy of citrus (Garcia Marì and del Rivero, 1982). In Israel, P. harti develops on Oxalis corniculata Linnaeus, where it lays the greatest number of eggs and lives longer than on Oxalis articulata Savigny; large densities were observed in early summer. Males represent more than 10% of the population and the eggs do not usually have a diapause (Dubitzki and Gerson, 1987). In Greece, the mite develops throughout the year on Oxalis corniculata but not on Oxalis articulata, whereas in the laboratory it may be reared on both host plants. Egg diapause depends on photoperiod (12:12), temperature (20°C), host plant and phenological stage (Koveos and Tzanakakis, 1989). Females developing with longer daylight and a temperature of 19°C lay diapausing eggs. Eggs do not develop with 70% under RH and varying conditions of light and dark, and temperatures (Koveos and Tzanakakis, 1991). The species is not injurious to citrus (Garcia Marì and del Rivero, 1982; Ciampolini et al., 1985).

13.3.3.1.2 Petrobia latens (Müller) (Fig. 13.8) Common name Brown wheat mite. Diagnostic characteristics FEMALE. The body is ovoid, about 700 μm long, dark brown or greenish in colour and the legs are pale yellow. The dorsal body setae are ﬁnely serrate, not set on tubercles and shorter than the distance between their bases.

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B

D C

Fig. 13.8. Petrobia latens (Müller). Female. (A) Dorsal view; (B) pretarsus I (from Jeppson et al., 1975); (C) distal anasthomosis of peritreme; (D) receptaculum seminis (from Vacante, 1985).

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The dorsal striae are lobates. The prodorsal setae v2 are twice as long as the other dorsal setae. The peritremes end in a large anasthomosis, about 50 long and 15 μm wide. The receptaculum seminis is smooth and tubular, about 15 μm long. Tarsi III and IV bear duplex setae (Vacante, 1985). MALE.

Unknown (Vacante, 1985).

Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The brown wheat mite has been recorded on about 116 host plants (Migeon and Dorkeld, 2006) and frequently on herbaceous plants, spontaneous and cultivated, and also on Citrus sp. in India (Prasad, 1974; Gupta and Gupta, 1994; Gupta and Chatterjee, 1997). The species reproduces by parthenogenesis thelytokous and the life cycle from egg to adult requires 8–11 days. The adult preoviposition takes 1–2 days and the adult lives for 2–3 weeks. The eggs are laid on soil or on any object in the area of plants. The female lays non-diapause and diapause eggs. The incubation of non-diapause eggs takes 7 days at 22°C, whereas diapause eggs remain unhatched for an indeﬁnite time during hot, dry weather; soil moisture and increased humidity stimulate the eggs to hatch. Summer females lay 70–90 summer eggs in 3 weeks but, in the case of winter eggs, each female lays 30 eggs at the same time (Fenton, 1951; Cox and Lieberman, 1960; Jeppson et al., 1975). In Sicily, the mite is sometimes common on grass under the canopy of citrus trees or on the lower foliage of the trees, but does not cause any damage to citrus crops. Natural enemies include coccinellid larvae and mites. The former include Chausseria sp., which occurs together with Petrobia latens in South Africa (Meyer Smith, 1981), whereas predatory mites include the bdellid B. depressa (Snetsinger, 1956) and the aceosejid Lasioseius terrestris Menon et Ghai (Menon and Ghai, 1969).

13.3.3.1.3 Petrobia tunisiae Manson (Fig. 13.9) Common name Unknown. Diagnostic characteristics FEMALE. The body is oblong, 600–700 μm long and dark green in colour. The dorsum is transversely striate. All dorsal setae are ﬁnely serrate and not set on tubercles. The prodorsal setae v2 are twice as long as the scapular setae. The opisthosomal dorsal setae f1, h1 and f2 are longer than the other opisthosomal setae. The peritremes end in a hook. The receptaculum seminis is tubular (Vacante, 1985).

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B

A

C

D

E

Fig. 13.9. Petrobia tunisiae Manson. Female. (A) Dorsal view; (B) distal end of peritreme; (C) receptaculum seminis. Male. (D) Palp; (E) aedeagus (from Vacante, 1985, partially modiﬁed).

194

Chapter 13 MALE. The male is smaller than the female and of the same colour. The aedeagus is simple, straight and thin at the distal end (Vacante, 1985).

Geographical distribution Iran (Kamali, 1990); Israel (Dubitzki, 1981; Dubitzki and Gerson, 1987; Ben-David et al., 2007); Italy (Vacante, 1985; Vacante and Nucifora, 1986); Spain (Ferragut and Escudero, 1996); and Tunisia (Manson, 1964). Bio-ecology Petrobia tunisiae lives on graminaceous plants, both spontaneous (Manson, 1964; Vacante; 1985; Dubitzki and Gerson, 1987) and cultivated as wheat and barley (Kamali, 1990), and sometimes on other weeds (Vacante, 1985). In Sicily it is common on spontaneous graminaceous plants in uncultivated lands or under the canopy of citrus trees, and occasionally on lemon leaves of the lower foliage (Vacante and Nucifora, 1986). The mite lays about 17 nondiapause eggs in the ﬁrst generation and most of these eggs enter diapause in the second generation. It is probable that the species oversummers in diapause (Dubitzki and Gerson, 1987).

13.4 TETRANYCHINAE BERLESE The female possesses one or two pairs of pseudanal setae (ps1–2) and the male four pairs of genito-pseudanal setae (g1–2, ps1–2). The empodium lacks, or if present does not bear, tenent hairs (Bolland et al., 1998).

13.4.1 Tenuipalpoidini Pritchard et Baker The opisthosomal seta f1 is set in a marginal position or absent. The empodium is claw-like or split distally. Tarsus I has two pairs of duplex setae, and tarsus II one pair (Bolland et al., 1998).

13.4.1.1 Tenuipalponychus Channabasavanna et Lakkundi The opisthosomal seta f1 is set in a marginal position. The empodium consists of a simple claw. The distal member of the duplex setae of tarsus II is a long solenidion (Bolland et al., 1998).

13.4.1.1.1 Tenuipalponychus citri Channabasavanna et Lakkundi Common name Unknown.

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Diagnostic characteristics FEMALE. The body is ovoid. The dorsal body setae are set on strong tubercles, serrate spatulate or sometimes tapering and not longer than the distances between their bases. The scapular setae sc2 are longer than the other prodorsal setae. The central area of the prodorsum has a reticulate pattern consisting of polygonal elements. The opisthosoma is feebly striate, with striae formed by discontinuous elements and transverse except between the dorsal opisthosomal setae e1, where they are longitudinal. The opisthosomal setae f2 are as long as half of f1. The peritremes end in a simple bulb (Channabasavanna and Lakkundi, 1977). MALE.

Unknown (Channabasavanna and Lakkundi, 1977).

Geographical distribution India (Channabasavanna and Lakkundi, 1977). Bio-ecology The species has been collected on C. sinensis in India (Channabasavanna and Lakkundi, 1977), and its bio-ecology is unknown.

13.4.2 Eurytetranychini Reck The empodium when present is claw-like. Tarsus I bears loosely associated setae or one pair of duplex setae; if two pairs of duplex setae are located on tarsus I, no pairs are set on tarsus II (Bolland et al., 1998). 13.4.2.1 Aponychus Rimando The opisthosoma bears ten pairs of dorsal setae. The anal region possesses one pair of pseudanal setae (ps). The empodial claw is apparently lacking (Bolland et al., 1998).

13.4.2.1.1 Aponychus chiavegatoi Feres et Flechtmann (Fig. 13.10) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid to rounded, 411 μm long and 388 μm wide, and yellow-greenish with dark green colour. The dorsum is ﬁnely granulated, with three to four pairs of circular structures on the prodorsum and three pairs of the same on the dorsal opisthosoma. The prodorsal setae are strong,

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B

Fig. 13.10. Aponychus chiavegatoi Feres et Flechtmann. Female. (A) Dorsal view; (B) seta c2 (from Feres and Flechtmann, 1988).

lanceolate serrate and set on tubercles; the scapular setae sc1 less than half as long as vertical setae v2. The eyes are located laterally to the bases of scapular setae sc1. The opisthosomal dorsal setae are strong, lanceolate serrate and set on tubercles; the dorsal opisthosomal setae c2 are very small, variable in length and displaced anteriorly on to the prodorsum; the dorsal opisthosomal setae c1, d1 and e1 decrease in length from ﬁrst to third pair (Feres and Flechtmann, 1988). MALE.

Smaller than the female, with aedeagus curved dorsal and sigmoid (Feres and Flechtmann, 1988). Geographical distribution Brazil (Feres and Flechtmann, 1988). Bio-ecology The mite has been collected on Citrus sp. in Brazil (Feres and Flechtmann, 1988). Its bio-ecology is unknown.

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13.4.2.1.2 Aponychus spinosus (Banks) (Fig. 13.11) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid. The prodorsal vertical setae v2 and scapular setae sc1 are tapering and the setae sc1 are longer and about twice as long as the scapular setae sc2; these latter setae are short, slender and sometimes blunt. The eyes are located laterally to the bases of scapular setae sc1. The dorsal opisthosomal setae tapering and set on tubercles; the setae c3 and f1 are short, slender and sometimes blunt, and the setae h1 are somewhat shorter than setae f2. Other dorsal opisthosomal setae are similar (Pritchard and Baker, 1955). MALE. Smaller than the female. The aedeagus has the distal half of the external

portion bent sharply dorsally, with the distal end curved somewhat caudally and narrowing to a blunt tip (Pritchard and Baker, 1955). Geographical distribution Brazil (Paschoal, 1969); Canada (Banks, 1909); Paraguay (Aranda and Flechtmann, 1971); the Philippines (Thewke and Enns, 1969); and the USA (Pritchard and Baker, 1955). Bio-ecology The mite has been recorded on 17 different host plants (Migeon and Dorkeld, 2006) and also on Citrus sp. (Aranda and Flechtmann, 1971). Its bio-ecology is unknown.

13.4.2.2 Eutetranychus Banks The opisthosoma possesses ten pairs of dorsal setae. The anal region bears two pairs of pseudanal setae (ps1–2). The empodial claws are apparently absent (Bolland et al., 1998).

13.4.2.2.1 Eutetranychus africanus (Tucker) (Fig. 13.12) Common name African red mite. Diagnostic characteristics FEMALE. The body is rounded, and brownish-green in colour. The dorsal setae are long, slender and set on tubercles. The prodorsum is striate and the striae

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D

C

E

Fig. 13.11. Aponychus spinosus (Banks). Female. (A) Dorsal view; (B) tarsus and tibia I; (C) pretarsus I. Male. (D) Tibia I; (E) aedeagus (from Pritchard and Baker, 1955).

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C B

E

F

D

Fig. 13.12. Eutetranychus africanus (Tucker). (A) Dorsal view of female; (B) distal portion of dorsal seta (from Jeppson et al., 1975); (C) receptaculum seminis; (D) aedeagus; (E) palptarsus of female; (F) palptarsus of male (from Meyer Smith, 1974, 1987).

possess weaker lobes. The dorsal opisthosomal setae c2, d2, e2 and f2 are long and slender and setae c1, d1, e1, f1 and h1 are lanceolate to subspatulate, varying in length from half to three-quarters the length of the other dorsal setae. The dorsal opisthosomal setae c2, d2, e2 and f2 can vary from relatively long, slender and tapering to long and subspatulate. The dorsal opisthosoma is striate and bears weakly developed lobes, and longitudinal striae between setae d1 and e1. The peritremes end simply. The recepataculum seminis is oval. Tibia II bears six setae (Meyer Smith, 1987).

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Smaller than the female, yellowish-green in colour and with dorsal body setae more slender, shorter and tapering than those of the female. The distal, bent portion of the aedeagus is shorter than the dorsal margin of the shaft, which is relatively straight (Meyer Smith, 1987). Geographical distribution Australia (Walter et al., 1995); Comoros (Bolland et al., 1998); Egypt (Attiah, 1967; Atalla and El-Atrouzy, 1971; Zaher et al., 1982); India (Manson, 1963; Gupta and Gupta, 1994); Japan (Ehara and Yogi, 1998); Madagascar (Gutierrez and Helle, 1971); Mauritius (Moutia, 1958; Baker and Pritchard, 1960); Mozambique (Meyer and Rodrigues, 1966); Myanmar (Burma) (Manson, 1963b; Baker, 1975), Papua New Guinea (Schicha and Gutierrez, 1985), the Philippines (Baker, 1975); Réunion Island (Gutierrez and Etienne, 1986); South Africa (Tucker, 1926; Baker and Pritchard, 1960; Meyer Smith, 1974, 1987); and Thailand (Uraisakul and Oates, 1999; Kongchuensin et al., 2005). Bio-ecology

The african red mite has been recorded on about 60 host plants (Migeon and Dorkeld, 2006), including C. aurantium (Gupta and Gupta, 1994), C. depressa Hayata (Ehara and Yogi, 1998), C. hystrix (Blommers and Gutierrez, 1975), C. limon (Tucker, 1926), C. paradisi (Blommers and Gutierrez, 1975), C. reticulata, C. sinensis (Tucker, 1926; Attiah, 1967), Citrus sp. (Baker and Pritchard, 1960; Gutierrez and Helle, 1971), Murraya paniculata (Linnaeus) Jack (Ehara and Yogi, 1998) and Poncirus trifoliata Linnaeus (Blommers and Gutierrez, 1975). Reproduction is parthenogenetic arrenothokous, with the female diploid and the male haploid and with two chromosomes (Bolland et al., 1981). The mite produces webbing, and its populations develop on the upper surface of the leaves; on the lower surface the females do not lay, and die after a few days. The period of preoviposition is 1–2 days and the life cycle from egg to egg takes about 15 days. At 24 ± 1°C and 85 ± 10% RH, the female lays an average of 4.2 eggs per day on C. hystrix (maximum eight), and two eggs per day on mandarin (maximum ﬁve). The time of development of egg, larva, protonymph and deutonymph is 7.2, 1.3, 1.0 and 0.7 days, respectively. The distribution of populations on the leaves seems independent of the age of the leaves (Blommers and Gutierrez, 1975). Mite dispersion is hindered by rain (Moutia, 1958; Baker and Pritchard, 1960) but it is resistant to moderate rain (Blommers and Gutierrez, 1975). Different natural enemies develop on E. africanus. The linyphiidad Hylyphantes graminicola Sundevall preys on immature and adult stages of the mite (Wipada, 1997). The insects include the coccinellid Stethorus madecassus Chazeau and some chrysopids (Blommers and Gutierrez, 1975). The phytoseiid mites include Amblyseius parasundi Blommers, Amblyseius tamatavensis Blommers, Amblyseius vazimba Blommers et Chazeau (Blommers and Gutierrez, 1975), Neoseiulus californicus, Neoseiulus longispinosus (Manita and Vatana, 2000; Kongchuensin et al., 2005), Phytoseius hongkongensis Swirski et Shechter (Charanasri and Kongchuensin, 2001), and Phytoseiulus persimilis Athias Henriot (Manita and Vatana, 2000) have also been recorded.

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Symptomatology and damage The African red mite infests various cultivated plants including C. hystrix, C. limon, C. paradisi, C. reticulata, C. sinensis and Citrus sp. (Gutierrez and Helle, 1971; Bolland et al., 1998); although it attacks all citrus, it does not prefer lemon. Immature and adult stages live exclusively on the upper surface of the leaves and produce, with feeding activity, thin whitish spots on the entire surface. The infested leaves become like grey lead and, with a high population density, may show lateral necrosis of the edges similar to the effects of mineral deﬁciency or tetranychid damage. The infested leaves may show early yellowing (Blommers and Gutierrez, 1975). Control In Madagascar, the need for control on citrus is necessary only on C. hystrix (Blommers and Gutierrez, 1975). There is no information with regard to other regions of the world.

13.4.2.2.2 Eutetranychus banksi (McGregor) (Fig. 13.13) Common name Texas citrus mite. Diagnostic characteristics FEMALE. The body is broad and robust, and tan to brownish-green in colour with dark brown to greenish spots and bars near the lateral margins. The legs are strong, with distal segments pale and the proximal brown. The dorsal setae are variable in length, commonly broader distally and not as long as the distance between their bases The prodorsum is striate longitudinally. The dorsal opisthosomal setae e1 are closer together than other dorsal setae. The opisthosomal striae are transverse, with the exception of a V pattern between dorsal setae d1 and e1. Tibia II bears six setae (Pritchard and Baker, 1955; Jeppson et al., 1975). MALE.

Smaller than the female, and similar in colour. The aedeagus is simple, abruptly turned dorsally near the distal end (Pritchard and Baker, 1955; Jeppson et al., 1975). Geographical distribution

Egypt (Zaher and El-Badry, 1961); Hawaii (Garrett and Haramoto, 1967); India (Sandhu et al., 1982); North, Central and South America (Jeppson et al., 1975; Bolland et al., 1998; Migeon and Dorkeld, 2006); Portugal (Goncalves et al., 2002); and Spain (Garcia et al., 2003, 2004). Bio-ecology The list of host plants numbers about 93 botanical species (Migeon and Dorkeld, 2006), and among arboreous cultivated plants are cited C. aurantifolia,

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C

D

Fig. 13.13. Eutetranychus banksi (McGregor). Female. (A) Dorsal view (from Jeppson et al., 1975); (B) tibia and tarsus I. Male. (C) Dorsal view; (D) aedeagus (from Pritchard and Baker, 1955).

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C. aurantium (Doreste, 1967), C. limon (Baker and Pritchard, 1962), C. paradisi (Baker and Pritchard, 1962), C. reticulata (Paschoal, 1970), C. sinensis (Baker and Pritchard, 1962; Garcia et al., 2003; Martinez et al., 2004) and Citrus spp. (Pritchard and Baker, 1955; Aranda and Flechtmann, 1971 ; Jeppson et al., 1975; Andrews and Poes, 1980; Suarez, 2004). Reproduction is parthenogenetic arrenothockous and the haploid number of chromosomes is three (Helle et al., 1970). The Texas citrus mite develops with low RH, very dry conditions and temperatures close to 27°C. At low densities, the pest commonly occurs in the southern quadrants of the canopy (Dean, 1959). On grapefruit leaves, the life cycle from egg to adult stage at 15°, 20°, 25°, 28°, 30° and 32°C and with 51–84% RH takes 29.6, 17.2, 13.1, 11.6, 11 and 9.6 days for the female and 27.7, 16.4, 12.0, 10.1, 10.8 and 8.5 days for the male, respectively. The sex ratio is female biased (<80%) at 20–30°C, but the proportion of females declines to 68% at 15°C and 61% at 32°C. The maximum net reproductive rate (Ro) is 31.26 at 28°C, followed by 26.06 and 25.29 at 30° and 25°C, respectively (Childers et al., 1991). On leaves of C. sinensis, the optimum range of development occurs from 28° to 31°C. The intrinsic rate of increase is −0.0649, 0.1723, 0.1759, 0.1973 and −0.0711, the net reproductive rate is 0.30, 8.25, 7.24, 9.21 and 0.44 and the mean generation times are 19.21, 12.79, 11.74, 11.52 and 11.64 days at 20, 25, 30, 32.5 and 35°C, respectively (Badii et al., 2003). In Texas, E. banksi is present on citrus leaves in the egg form through the year, laid along the midrib and near the lateral leaf margins, at lower densities from February to April but higher during May and June; in Mexico it develops 22–23 generations per year (Badii et al., 2003). Fungi, insects and mites limit the natural development of populations. Fungi include Neozygites ﬂoridana (Weiser et Muma) Reumadière et Keller (Neozygitaceae) (Weiser and Muma, 1966; Muma, 1969) and Hirsutella thompsonii (McCoy and Selhime, 1974). Insects include the coccinellid Stethorus atomus Casey (McMurtry et al., 1970) and Stethorus siphonulus Kapur (Raros and Haramoto, 1974); the mites include the phytoseiids Euseius mesembrinus (AbouSetta and Childers, 1989) and Galendromus helveolus (Caceres and Childers, 1991), the ascid Lasioseius scapulatus Kennet (Ramirez and Esponda, 1986) and the stigmeid Agistemus ﬂeschneri Summers (Zaher and El-Badry, 1961). Symptomatology and damage The feeding symptomatology is similar to that of the citrus red mite, Panonychus citri (Jeppson et al., 1975). The results of monthly inspections for arthropods in more than 130 citrus groves carried out in Florida in 1956–1971 showed that E. banksi numbers increased in the 16-year period (Simanton, 1976). In a citrus orchard (cultivar Rhode Red Valencia) under irrigation management, the ﬁnal mean density of feeding stipples on infested leaves were 873 per cm2 and leaf life lasted 380 days. No signiﬁcant negative relationship exists between leaf life and mite damage. The study indicated that damage by Texas citrus mites to Valencia citrus leaves promoted little or no premature leaf abscission in irrigated trees (Hall and Simms, 2003).

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Control The limits and possibilities of biological (McMurtry, 1978, 1985) and IPM strategies are well known (Knapp, 1981). With regard to chemical control, avermectins (McCoy et al., 1982; French and Villarreal, 1988); carbamates (French et al., 1985); organochlorines (French, 1975; French et al., 1985; Buergo et al., 1986; Flores et al., 1994, 1996a); organotins (Ochoa et al., 1989); pyrethroids (Ochoa et al., 1989); phenyl pyrazoles (French and Villarreal, 1988); organophosphorus (Buergo et al., 1986); petroleum oils (French, 1975; Buergo et al., 1986; French and Villarreal, 1988); pyridazinones (French and Bruno, 1996); and sulfur (Jeppson, 1989) have all been cited in the literature.

13.4.2.2.3 Eutetranychus citri Attiah (Fig. 13.14) Common name Unknown.

Fig. 13.14. Eutetranychus citri Attiah. Female. Dorsal view (partially modiﬁed from Attiah, 1967).

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Diagnostic characteristics FEMALE. The body is ovoid, 460 μm long and 280 μm wide. The dorsal body setae are spatulate and set partly on small tubercles. The prodorsum is longitudinally striate, with the scapular setae sc2 not set on tubercles. The dorsal opisthosomal setae c2, d2, e2 and f2 are slightly longer, and setae c2 and c3 are located anteriorly to dorsocentral setae c1. The dorsal opisthosomal setae c1, d1, e1, f1, h1 and setae c3 are not set on tubercles. The striae between dorsocentral setae d1 and e1 are longitudinal. The peritremes end simply. Tibia II possesses ﬁve setae (Attiah, 1967). MALE.

Unknown (Attiah, 1967).

Geographical distribution Egypt (Attiah, 1967) and India (Nassar and Ghai, 1981; Gupta and Gupta, 1994). Bio-ecology The mite has been collected on lime trees in Egypt (Attiah, 1967) and on Citrus sp. and other host plants in India (Nassar and Ghai, 1981; Gupta and Gupta, 1994). 13.4.2.2.4 Eutetranychus cratis Baker et Pritchard (Fig. 13.15) Common name Unknown. Diagnostic characteristics FEMALE. The body is 338 μm long, 255 μm wide and is greenish in colour. The dorsal body setae are nearly rod-like and serrate and set on pronounced, granulated tubercles. The prodorsum is granulate in the anterior part and ﬁnely and longitudinally striate in the posterior part, with scapular setae sc2 lateral and shortest. The dorsal opisthosomal setae are relatively longer. The setae c3, f2 and h1 are shortest; the setae c3, c2, d2, e2 and f2, f1 and h1 are set laterally. The setae d1 are wider apart than other dorsocentral setae; the setae f1 are more widely spaced than setae c1 and e1; the setae c1 are closest. The striae between the dorsal opisthosomal setae d1 and e1 form a V pattern, and the integument between setae c1, d1 and e1 presents elliptical elevations with a basketweavelike shape. The stylophore bears a pair of strong lobes ventrodistally and a dorsodistal protuberance. The peritremes end with a concave, bulbous enlargement. Tibia II possesses ﬁve setae (Baker and Pritchard, 1960). MALE.

Smaller than the female and lighter, without integumentary elevations and with the aedeagus bent abruptly dorsally and tapering (Baker and Pritchard, 1960).

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A

B

C

D

Fig. 13.15. Eutetranychus cratis Baker et Pritchard. Female. (A) Dorsal view; (B) sensory setae of palptarsus; (C) enlargements of mediodorsal hysterosomal integument. Male. (D) Aedeagus (from Baker and Pritchard, 1960).

Geographical distribution Congo (Gutierrez and Bonato, 1994), Congo (RDC, ex-Zaïre) (Baker and Pritichard, 1960) and Nigeria (Matthysse, 1980). Bio-ecology Eutetranychus cratis has been collected on Bixa sp., Macaranga sp. and orange leaves in Zaïre (Baker and Pritichard, 1960), on Trema orientalis Blume and Manihot esclulenta Crantz in Nigeria (Matthysse, 1980) and Congo (Gutierrez and Bonato, 1994). Immature and adult stages live isolated on the upper leaf surface and the females lay eggs along the veins (Gutierrez and Bonato, 1994). On the various host plants, the feeding activity of the mite produces a

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thin, light stippling on the upper leaf surface. The symptomatology on citrus is unknown. With regard to natural enemies, Gutierrez and Bonato (1994) have reported mites and predaceous insects in Congo. 13.4.2.2.5 Eutetranychus eliei Gutierrez et Helle (Fig. 13.16) Common name Unknown. Diagnostic characteristics FEMALE. The body is about 415 μm long and 250 μm wide. The dorsal body setae are spatulate and set on small tubercles. The prodorsum is longitudinally

A

B

C

F D

E

Fig. 13.16. Eutetranychus eliei Gutierrez et Helle. Female. (A) Distal end of pertitreme; (B) palptarsus; (C) associated setae on tarsus I; (D) associated setae on tarsus II. Male. (E) aedeagus; (F) palptarsus (from Meyer Smith, 1974).

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striate with striae strongly lobed. The dorsal opisthosomal setae are of different length. The dorsal opisthosomal setae c1, d1 and e1 are shortest and c1 a third of the length of setae c2. The opisthosoma is striate and the striae between setae e1 and f1 form a V pattern. The peritremes end simply. Tibia II possesses six setae (Gutierrez and Helle, 1971). MALE.

The male is small, 330 μm long and 160 μm wide. The aedeagus has the distal part forming an angle of less than 90° to the axis of shaft; the dorsal margin of the shaft is concave and twice as long as the distal, bent portion (Gutierrez and Helle, 1971). Geographical distribution Madagascar (Gutierrez and Helle, 1971). Bio-ecology

The species has been collected on Artocarpus incisa Linnaeus, Plumeria alba Linnaeus and Citrus sp. in Madagascar (Gutierrez and Helle, 1971).

13.4.2.2.6 Eutetranychus orientalis (Klein)1 (Fig. 13.17) Common name Oriental red mite, citrus brown mite, lowveld citrus mite. Diagnostic characteristics FEMALE.

The body is ovoid, about 400 μm long, from green–brown to dark green in colour and with dark inner blotches. The dorsal body setae are set on tubercles and vary in length and shape. The prodorsum is striate longitudinally and the striae are tuberculate. The dorsal opisthosomal striae between setae d1 and e1 vary from longitudinal to V-shaped. The dorsal opisthosomal setae c2, d2, e2 and f2 are long and lanceolate, subspatulate or broadly spatulate; the setae c1, d1, e1, f1 and h1 are short and spatulate, lanceolate or subspatulate. The setae c1 are more or less in line with setae c2 and c3. The setae c1 and f1 form a square. The receptaculum seminis is pointed distally. Tibia II bears six seta (Baker and Pritchard, 1960; Meyer Smith, 1987).

1According

to Meyer Smith (1987), Eutetranychus monodi André, 1954, Eutetranychus anneckei Meyer, 1974 and Eutetranychus sudanicus Elbadry, 1971, recorded for citrus and other plants in different regions of the world, are regarded as conspeciﬁc with E. orientalis.

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A

C B

D

E F

Fig. 13.17. Eutetranychus orientalis (Klein). Female. (A) Dorsal view; (B) palptarsus; (C) associated setae on tarsus I; (D) associated setae on tarsus II; (E) receptaculm seminis. Male. (F) Aedeagus (from Meyer Smith, 1974, 1987).

MALE. Smaller than the female, 360 μm long and with dorsal body setae shorter

than those of the female and more lanceolate or subspatulate. The aedeagus has the bent portion longer than the dorsal margin of the shaft (Baker and Pritchard, 1960; Meyer Smith, 1987). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006).

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Bio-ecology The list of host plants numbers over 217 botanical species (Migeon and Dorkeld, 2006), including C. aurantium (Nassar and Ghai, 1981; Zaher et al., 1982); C. grandis (Meyer Smith, 1974); C. jambhiri; C. karma Raf.; C. limon (Baker and Pritchard, 1960; Chaudhri et al., 1974; Nassar and Ghai, 1981; Gupta and Gupta, 1994; Garcia et al., 2003); C. medica (Gupta and Gupta, 1994); C. paradisi (Baker and Pritchard, 1960; Gupta and Gupta, 1994; Ben-David et al., 2007); C. reticulata (Prasad, 1974; Gupta and Gupta, 1994; Ben-David et al., 2007); C. sinensis (Rodrigues, 1968; Chaudhri et al., 1974; Gupta and Gupta, 1994; Walter et al., 1995); Citrus sp. (Baker and Pritchard, 1960; Rambier, 1965; Chaudhri et al., 1974; Nassar and Ghai, 1981; Walter et al., 1995; Gupta and Chatterjee, 1997; Kreiter et al., 2002; Garcia et al., 2003); and Citrofortunella × microcarpa (Bunge) Wijnands (Ehara, 2004). Although the oriental red mite is fundamentally a pest of citrus, it can attack other cultivated plants such as cotton, squash, ﬁg, frangipani, pear, grapevines, quince, walnut and Euphorbia (Jeppson et al., 1975). Reproduction is parthenogenetic arrenothokous, with a haploid condition of three chromosomes, and a male:female ratio of 1:3.25 (Helle et al., 1970). During the summer, the mite is found mostly along the midrib of the upper leaf surface, but in the winter or at high density also on both leaf surfaces. The developmental threshold is 11°C. Above 26°C, reproduction becomes slower; at 23°C the time of pre-oviposition is 12 days, at 20–22°C from 2.5 to 3.0 days and at 14–15°C from 4 to 8 days. Egg development rate decreases with low RH. The optimum conditions for mite development are 21°C and 59–70% RH. The species develops with temperatures ranging from 18° to 30°C and 35–75% RH. Beyond these limits development decreases or stops (Bodenheimer, 1951). The longevity of fertilized females is lower (6.9 days) than that of nonfertilized females (9.8 days) (Angsumarn and Kosol, 1988). In summer, the adult has a life span of about 12 days, in spring and autumn 14–18 days and in winter more than 21 days (Bodenheimer, 1951). In summer, the period of pre-oviposition, oviposition and postoviposition is 1.0, 10.4 and 2.4 days, respectively, and 2.4, 12.2 and 3 days, respectively, in winter. In summer, the egg, larva and protonymph develop in 4.3, 2.9 and 1.7 days, respectively, and in winter in 5.7, 3.7 and 2.4 days, respectively (Siddig and El-Badry, 1971). At 20°C, egg development, immature stage development and adult longevity last 8.06, 20.75 and 21.88 days, respectively, while at 30°C they are 3.57, 10.0 and 7.36 days, respectively (Atwa et al., 1987a). In colder months a generation occurs in 14.5 days, and in 11.2 days in summer, and in 12 months the species can produce up to 27 generations (Siddig and El-Badry, 1971). Net reproductive rate (Ro) is 18.85, cohort generation time (Tc) is 17.03 days, the innate capacity for increase (Rc) is 0.17 and the ﬁnite rate of increase is 1.19 mites per day (Angsumarn and Kosol, 1988). In Egypt, on seedlings of six kinds of citrus, the sour orange and citron were found to be the most favourable hosts, as they were severely infested,

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followed by sweet orange, Morrakishi orange, Navel orange and mandarin (Rasmy and Zaher, 1983). The structure and physiology of host varieties show a positive correlation between rate of population development on leaves, and both the density of oil glands and the quantity of tannic acid in the leaves, and a negative correlation with the thickness of the cuticle (Mohamed, 1965). Protein concentration in the leaves is positively correlated with mite infestation, and lemon (cultivar Baramasi) contains phenols and free amino acids, which both seem to be negatively correlated with mite infestation (Jyotika and Mandeep, 2003). A positive correlation between temperature and day length and the population development of E. orientalis on C. aurantifolia, C. reticulata, C. limon and C. paradisi has been observed, whereas RH, rainfall and the Phytoseiid, Amblyseius delhiensis, are negatively correlated with a population increase on all citrus varieties except C. reticulata (Neena and Sadana, 1998). In coastal regions where humidy is moderate, populations develop during the summer, whereas in regions with low humidity populations do not increase in spring and summer, but may develop later and cause serious damage in autumn (Jeppson et al., 1975; Jeppson, 1989). In India, two population peaks occur per year, in late April and in September, and no mites have been found since the end of November until the third week of January (Bhumannavar and Singh, 1986). With temperatures of between 21° and 27°C and RH of 30–40%, a peak has been observed in May–June and a decrease in July–August following intense rainfall (Dhooria and Butani, 1984). In South Africa, the damage is more evident in autumn, especially in dry conditions. Infestations appear between February and March and the peak is reached between March and April (Meyer Smith, 1998). The factors limiting the development of populations include entomophatogenic fungi, predatory mites and insects. Pathogens include several fungi such as the Ascomycota Verticillium lecanii, the Clavicipitaceae H. thompsonii and Metarligium anisopliae, and the Exobasidiomycetidae Meira argovae, Meira geulakonigii, and Acaromyces ingoldii (Table 13.4). Predatory mites include cheyletids, stigmaeids, tydeids, cunaxidae, smarididae (Table 13.4) and phytoseiids (Table 13.5). Various species of insects have been collected in different regions of the world (Kazimi, 1980; Dhooria and Butani, 1984; Meyer Smith, 1998), including coccinellids, sthaphylinids, anthocorid, chrysopids and eolothripids (Table 13.4).

Symptomatology and damage The oriental red mite prefers sour lemon to stock to sweet lemon, mandarin and orange (Mohamed, 1965), and lays a higher number of eggs on lemon leaves than on orange and mandarin (Rasmy, 1977). Feeding on the upper leaf surface and fruit causes stippling, similar to that of the citrus red mite, P. citri. With high levels of attack, the trees become silver–grey and the leaves stop development and may fall, and the shoots die back. Bare trees represent a serious problem in the nursery or in young orchards. The combined effect of

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Table 13.4. Eutetranychus orientalis (Klein) natural enemies (fungi, insects and mites). Families and species Ascomycota Verticillium lecanii (Zimmermann) Viégas Clavicipitaceae Hirsutella thompsonii Fisher Metaharrizium anisopliae (Metschnikoff) Sorokin Exobasidiomycetidae Acaromyces ingoldii Boekhout et al. Meira argovae Boekhout et al. Meira geulakonigii Boekhout et al. Aeolothripidae Scolothrips indicus Priesner Scolothrips longicornis Priesner Scolothrips sp. Anthocoridae Orius spp. Chrysopidae Chrysopa spp. Coccinellidae Stethorus pauperculus (Weise) Staphylinidae Oligota (Somatium) oviformis (Casey) Oligota spp. Cheyletidae Acaropsellina sollers (Rohdendrof) Acaropsis aegyptiaca Wafa et Soliman Cheletogenes ornatus (Canestrini et Fanzago) Chiapacheylus macrocorneus Zaher et Soliman Eutogenes punctata Zaher et Soliman Cunaxidae Cunaxa capreolus (Berlese) Smaridiidae Smaris biscutatus Meyer et Rike Stigmaeidae Agistemus buntex Chaudhri Agistemus exsertus Gonzalez Rodriguez Tydeidae Tydeus californicus (Banks) Tydeus kochi Oudmans Tydeus munsteri Meyer et Ryke Pronemtus ubiquitus (McGregor)

Reference(s)

Sewify and Mabrouk, 1990; El-Hady, 2004 McCoy and Selhime, 1974; Gerson et al., 1979; Dhooria and Butani, 1984 El-Hady, 2004

Paz et al., 2009a Paz et al., 2009a Paz et al., 2009a Lal, 1977; Dhooria, 1983 Heikal et al., 1990 Meyer Smith, 1998 Meyer Smith and Schwartz, 1998a Meyer Smith, 1998 Banu and Channabasavanna, 1972 Banu and Channabasavanna, 1972 Meyer Smith and Schwartz, 1998a Nassar and Kandeel, 1986 Wafa et al., 1970 Rizk et al., 1983; Rezk and Gadelhak, 1996 Zaher and Soliman, 1971 Hassan, 1989 Zaher et al., 1975 Meyer Smith, 1998 Khan and Muhammad, 2005 Rizk et al., 1983; Rezk and Gadelhak, 1996 Rizk et al., 1983 Rasmy, 1970b Meyer Smith and Schwartz, 1998a Rasmy, 1969, 1970b

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Table 13.5. Eutetranychus orientalis (Klein) natural enemies (phytoseiid mites). Species

Reference(s)

Amblyseius deleoni Muma et Denmark Amblyseius zaheri Yousef et El-Borolossy Cydnoseius negevi (Swirski et Amitai) Euseius alstoniae (Gupta) Euseius bwende (Pritchard et Baker) Euseius citri (van der Merwe et Ryke) Euseius delhiensis (Narayanan et Kaur) Euseius ﬁnlandicus (Oudemans) Euseius gossipi (El-Badry) Euseius hibisci (Chant) Euseius olivi (Nasr et Abou-Awad)

Rasmy et al., 2000 Rasmy et al., 2003 Abou-Awad et al., 1989 Dhooria, 1983 Meyer Smith, 1998 Meyer Smith, 1998 Neena, 2001 Sharma and Sadana, 1987 El-Enany et al., 1993 Swirski et al., 1967, 1970 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Shih et al., 1993 Swirski et al., 1967, 1970 Tanigoshi et al., 1990; Abd El-Samad et al., 1996 Swirski and Dorzia, 1969 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Ibrahim et al., 2005 Angsumarn, 1994 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Abou Awad et al., 1998b; Momen and El-Borolossy, 1999 Nawar et al., 2001a

Euseius ovalis (Evans) Euseius rubini (Swirski et Amitai ) Euseius scutalis (Athias Henriot) Galendromus occidentalis (Nesbitt) Neoseiulus barkeri Hughes Neoseiulus californicus (McGregor) Neoseiulus longispinosus (Evans) Paraseiulus talbii (Athias Henriot) Proprioseiopsis badryi (Yousef et El-Borolossy) Phytoseius plumifer (Canestrini et Fanzago) Proprioseiopsis asetus (Chant) Proprioseiopsis cabonus (Schicha et Elshaﬁe) Proprioseiopis lindquisti (Schuster et Pritchard) Typhlodromips swirskii (Athias Henriot) Typhlodromus athiasae Porath et Swirski Typhlodromus balanites El-Badry Typhlodromus citri Hassan et Romeih Typhlodromus mangiferus Zaher et El-Borolossy Typhlodromus praeacutus van der Merwe Typhlodromus pyri Scheuten Typhlodromus transvaalensis (Nesbitt)

Fouly, 1997 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Hoda et al., 1987 Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999; Nawar et al., 2001b Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999 Romeih et al., 2005 El-Halim, 2004 Meyer Smith and Schwartz, 1998a Rasmy, 1970b Abou-Awad et al., 1998b; Momen and El-Borolossy, 1999

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insufﬁcient water and low infestation produces as much defoliation and twig dieback as a high density of attack (Jeppson et al., 1975; Jeppson, 1989). Control Hot, dry environmental conditions and/or irrational programmes of control can stimulate outbreaks of mite populations. In Egypt, if citrus orchards are not treated with chemicals constantly, E. orientalis is no longer a problem (Rasmy, 1969). If sprays are applied against other citrus pests, it is necessary to add acaricides to control phytophagous mites (Rasmy, 1970a). In South Africa, in orchards where pyrethroids have been sprayed against other pests, attacks are more severe and an average of three or more females/leaf justiﬁes chemical control (Meyer Smith, 1998). In Egypt, one spray application of acaricide applied as populations of mite start to increase in size is sufﬁcient to maintain control throughout the year (Wafa et al., 1969). BIOLOGICAL CONTROL. In spite of the numerous natural enemies well known throughout the world, biological control alone is insufﬁcient and chemical control is usually neccessary. Recently, Paz et al. (2009a) have evaluated the action of the Exobasidiomycetidae M. argovae, M. geulakonigii and A. ingoldii in the laboratory, observing that after 14 days M. argovae and M. geulakonigii produced 83.6% and 87.8% mortality, whereas A. ingoldii resulted in mortality of 83.9% and 79.2% after 7 and 14 days, respectively. CHEMICAL CONTROL.

Chemicals include organochlorines (El Kady et al., 1977; Dhooria, 1983; Dhooria and Butani, 1984; El-Zemaity et al., 1984; Deshpande et al., 1988; Iskander et al., 1993; Helal et al., 1995; Rezk and Gadelhak, 1996; Marquez et al., 2006); petroleum oils (Jeppson et al., 1975; Iskander et al., 1993; Ibrahim, 1994; Osman et al., 1994; Osman, 1997); sulfur (Bodenheimer, 1951; Klapperich, 1957; Jeppson et al., 1975; Singh et al., 1986; Deshpande et al., 1988; Jeppson, 1989; Meyer Smith, 1998); or other acaricides effective in controlling other tetranychid mites.

INTEGRATED PEST MANAGEMENT. The above-mentioned aspects clearly indicate the rational use of insecticides and, among these, the appropriate distribution of petroleum oils and selective acaricides helps to limit the development of mite populations and indirectly to realize a preliminary status of IPM. In this context, various citrus cultivars have been investigated to evaluate their resistance to mite attacks (Sadana and Kanta, 1972; Dhooria and Sandhu, 1973; Dhooria, 1982; Singh et al., 1983; Bhumannavar et al., 1988).

13.4.2.2.7 Eutetranychus pantopus (Berlese) (Fig. 13.18) Common name Unknown.

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A

B

Fig. 13.18. Eutetranychus pantopus (Berlese). Female. (A) Dorsal view; (B) aedeagus (partially modiﬁed from Sayed, 1946).

Diagnostic characteristics FEMALE. The body is oval–round, deep brown in colour, 400 μm long and 283 μm wide. The dorsal body setae are set on small tubercles, relatively long and acutely tapering. The prodorsum is striate longitudinally. The dorsal opisthosomal setae are as long as the distances between their bases or longer. The dorsal opisthosomal setae d2, e2 and f2 seem longest and the seta h1 shorter. The integument is striate transversally between setae c1 and forming a V pattern between setae d1 and e1, and transversally between setae f1. The peritremes end simply (Sayed, 1946).

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Chapter 13 MALE.

The male is triangular in shape and light brown in colour, 383 μm long, with aedeagus curved upwards (Sayed, 1946). Geographical distribution Australia (Berlese, 1910; Walter et al., 1995), Egypt and Sudan (Sayed, 1946). Bio-ecology Eutetranychus pantopus has been collected on Albizia lebbeck Linnaeus, Azadirachta indica (Linnaeus), leaves and fruit of Citrus sp., Ficus sp. and Ricinus communis Linnaeus in Egypt and Sudan (Sayed, 1946), and on Ficus sp. (Berlese, 1910) and citrus (Walter et al., 1995) in Australia. Its bio-ecology is unknown.

13.4.2.2.8 Eutetranychus pyri Attiah (Fig. 13.19) Common name Unknown. Diagnostic characteristics FEMALE. The body is oval–round, 450 μm long and 310 μm wide. The prodorsum presents striae of tortuous lines forming crescent shapes, with three

Fig. 13.19. Eutetranychus pyri Attiah. Female. Dorsal view (partially modiﬁed from Attiah, 1967).

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pairs of setae, rod-like and serrate and set on strong tubercles, except the scapular setae sc2. The dorsal opisthosomal setae are rod-shaped and serrate, around of same length and set on strong tubercles, except the setae c3. The dorsal opisthosomal setae e1 form a square with setae f1. The striae between setae d1 and e1 are longitudinal. The peritremes end in a simple enlargement. The tibia II bears ﬁve setae (Attiah, 1967). MALE.

Unknown (Attiah, 1967).

Geographical distribution Egypt (Attiah, 1967; Zaher et al., 1982). Bio-ecology Eutetranychus pyri has been collected in Egypt on P. communis, R. communis, Cynodon dactylon Linnaeus, C. aurantium, Solanum melongena Linnaeus and Dalbergia sisso Roxburgh ex De Candolle (Attiah, 1967; Zaher et al., 1982). The bio-ecology of the mite is unknown.

13.4.2.3 Meyernychus Mitrofanov The opisthosoma possesses nine pairs of dorsal setae, and the setae c2 are lacking. The anal region bears two pairs of pseudanal setae (ps1–2). The empodial claws are apparently absent (Bolland et al., 1998).

13.4.2.3.1 Meyernychus emeticae (Meyer) (Fig. 13.20) Common name Angolan citrus mite. Diagnostic characteristics FEMALE. The body is ovoid, 550 μm long and about 400 μm wide and red– black coloured. The prodorsum bears three pairs of short setae, from subspatulate to spatulate, and the striae are distinctly lobed and medially interrupted. The dorsal opisthosomal setae vary from subspatulate to spatulate, and some are set on small tubercles. The dorsal opisthosomal setae e1 and f1 form a rectangle and the striae between the setae d1 and e1 are longitudinal. The peritremes are bilobed distally. The receptaculum seminis is depicted. The tibia II bears six setae. The solenidion and the proximal tactile seta of associated setae both on tarsi I and II are of about equal length. The species may be easily confused with P. citri, which differs in that the dorsal setae are markedly shorter (Meyer Smith, 1974, 1987).

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A

B

D

C

Fig. 13.20. Meyernychus emeticae (Meyer). Female. (A) Dorsal view; (B) receptaculum seminis; (C) distal end of peritreme; (D) palptarsus (from Meyer Smith, 1974, 1987).

MALE. The male is smaller than the female and pale orange (Meyer Smith, 1981).

Geographical distribution Angola (Meyer Smith, 1974) and South Africa (Meyer Smith, 1974, 1987). Bio-ecology The angolan citrus mite is recorded on several host plants, including C. limon (Meyer Smith, 1974), C. reticulata (Meyer Smith, 1987), C. sinensis (Meyer

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Smith, 1987) and C. nobilis Lour. Its bio-ecology is unknown. The mite lives on the upper surface of the leaves and the feeding activity of immature and adult stages produce grey spots that may become brown and lead to death of the leaf. No control measures are required (Meyer Smith, 1981).

13.4.3 Tetranychini Reck The opisthosomal seta f1 is set in the normal dorsal position. The empodium is claw-like or split distally. The tarsus I has two pairs of duplex setae and the tarsus II one pair (Bolland et al., 1998).

13.4.3.1 Acanthonychus Wang The dorsal body setae are set on tubercles. The anal region bears one pair of pseudanal setae (ps), and two pairs of setae h2–3. The empodium is split distally (Bolland et al., 1998).

13.4.3.1.1 Acanthonychus jiangfengensis Wang (Fig. 13.21) Common name Unknown. Diagnostic characteristics FEMALE. The body is green coloured, 416 μm long and 273 μm wide. The idiosoma is striate transversally, also on tubercles. The dorsal propodosomal setae, except the second pair, are set on small, circular tubercles. The dorsal opisthosomal setae, except c3, f1, f2 and h1, are set on prominent and conical tubercles. All dorsal setae are prominent, and deeply serrate. The opisthosoma bears a single pair of nude pseudanal setae ps and two pairs of h2–h3, set on small tubercles and with pubescence. The distal eupathidium of the palptarsus is about one and a half times as long as broad at base. The empodium is divided into three pairs of hairs in the middle (Wang, 1982). MALE. The aedeagus is slender, long, tapering and curved dorsally (Wang, 1982).

Geographical distribution China (Wang, 1982). Bio-ecology The mite has been collected on C. sinensis (Wang, 1982). Its bio-ecology is unknown.

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B

Fig. 13.21. Acanthonychus jiangfengensis Wang. Female. (A) Dorsal view; (B) tarsal appendage (from Meyer Smith, 1987).

13.4.3.2 Eotetranychus Oudemans The dorsal body setae are as long or longer than the distances between their bases. The opisthosoma is transversely striate. The anal region bears two pairs of pseudanal setae (ps1–2) and two pairs of setae h1–2. The empodium is split near the middle into three pairs of hairs (Bolland et al., 1998).

13.4.3.2.1 Eotetranychus cendanai Rimando (Fig. 13.22) Common name Citrus yellow mite. Diagnostic characteristics FEMALE. The body is 390 μm long, 240 μm wide and pale yellowish-green in colour and with silvery setae. The dorsum of the idiosoma possesses longitudinal striae between the dorsal opisthosomal setae e1. The dorsal opisthosomal setae are slender, tapering and pubescent. The dorsal opisthosomal

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A B

D

C

Fig. 13.22. Eotetranychus cendanai Rimando. Female. (A) Tibia and tarsus I. Male. (B) Palptarsus; (C) aedeagus; (D) tibia and tarsus I (from Manson, 1963b).

setae c1, d1, e1 and f1 are about as long as or longer than longitudinal intervals between them. The peritremes end in a simple bulb. The tibiae I and II have eight and ﬁve tactile setae, respectively; the tarsus I has four tactile setae proximal to the duplex setae. The genital ﬂap and the area anterior to the ﬂap are striated transversally (Ehara, 1969; Jeppson et al., 1975). MALE. The body is 270 μm long and 270 μm wide. The aedeagus turns upward

at almost a right angle to the shaft, is strongly sigmoid and tapers distally (Ehara, 1969; Jeppson et al., 1975). Geographical distribution Cambodia (Manson, 1963b; Baker, 1975); China (Wang, 1980); the Philippines (Rimando, 1962a); Taiwan (Ehara, 1969); and Thailand (Manson, 1963b; Baker, 1975). Bio-ecology The citrus yellow mite is recorded on several host plants, including C. aurantifolia (Baker, 1975), C. aurantium (Rimando, 1962a), C. grandis (Ehara, 1969), C. medica, C. sinensis (Baker, 1975), C. hystrix (Baker, 1975) and Citrus sp. (Baker, 1975; Ehara and Wongsiri, 1975) in the Philippines and South-east

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Asia. In Thailand, the mite has been collected on six different citrus species (C. hystrix, C. aurantifolia, C. reticulata, C. maxima, C. madurensis and Murraya paniculata) (Thongtab et al., 2002). On C. medica and C. mitis, the ratio of males to females in the offspring of mated females averages 1:3, and from unmated females 1:0 (Barrion and Corpuz Raros, 1975). The fertilized females produce offspring with a male:female sex ratio of 1:1.02–1:2, and unfertilized females generate only male offspring (Thongtab et al., 2002). The mite feeds on the upper surface of the leaves, produces little webbing and frequently causes damage to young trees. At 28 ± 1°C and 58 ± 5% RH on lime, leech lime and Shogun orange, the egg, larva, protonymph and deutonymph stages ranged from 4.9 to 5.8, 1.9 to 3.0, 1.9 to 2.6 and 1.8 to 2.7 days, respectively (Thongtab et al., 2002). On C. medica and C. mitis, the duration of the egg, larval, protonymphal, deutonymphal and adult stages averaged 5–6, 3.5–4.5, 3.0–3.5, 2.5–3.5 and 19.0–20.5 days, respectively, in males, and 5–6, 4.0–4.5, 3, 4 and 15.5–18.5 days in females (Barrion and Corpuz Raros, 1975). The life cycle required 11.2–14.0 days, and the fertilized female laid 6.30–26.38 eggs (0.96–2.43 eggs/day) (Thongtab et al., 2002). Mated and unmated females laid 22–23 (11–13) eggs on C. medica and 24–25 (10–18) on C. mitis, and 46.25% (41.59%) and 55.11% (45.46%) of the eggs hatched (Barrion and Corpuz Raros, 1975). The female longevity lasted from 6.55 to 10.52 days; longevity of unfertilized females lasted 7.52–11.62 days and 14.35–17.73 eggs (1.40–1.97 eggs/day) are laid per female (Thongtab et al., 2002). The mortality averages were 51.45% on C. medica and 65% on Citrofortunella mitis (Barrion and Corpuz Raros, 1975). The net reproductive rate of increase (Ro) is 14.1 times, the cohort generation time (Tc) is 17.37 days, the intrinsic rate of increase (rm) is 0.156 individuals/day and ﬁnite rate of increase (λ) is 1.165. Highest mortality occurs during the larval stage, followed by the nymphal and egg stages (Thongtab et al., 2002). The natural enemies limiting the development of populations include the linyphiidae H. graminicola (Wipada, 1997) and the phytoseiid mite N. longispinosus (Thongtab et al., 2001; Kongchuensin et al., 2005). In the Philippines, appeared to be a potential citrus pest and has damaged young trees previously treated with insecticides (Rimando, 1962a). Control seems unjustiﬁed.

13.4.3.2.2 Eotetranychus kankitus Ehara (Fig. 13.23) Common name Citrus yellow mite. Diagnostic characteristics FEMALE. The body is ovoid. The tibiae I and II bear nine and eight tactile setae, respectively, the tarsi I and II ﬁve and three tactile setae proximal to the

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Fig. 13.23. 1975).

223

Eotetranychus kankitus Ehara. Male. Aedeagus (from Jeppson et al.,

duplex setae, respectively. The peritremes are slightly curved distally but not hooked. The genital ﬂap is striate transversally on the posterior part and longitudinally on the anterior central part; the area anterior to the genital ﬂap is longitudinally striate (Ehara, 1955; Jeppson et al., 1975). MALE. The empodium I is split distally into three pairs of ventrally direct hairs.

The aedeagus is slightly downcurved, tapering distally and slightly sigmoid (Ehara, 1955; Jeppson et al., 1975). Geographical distribution China (Wang, 1980); India (Manson, 1963b); and Japan (Ehara, 1955). Bio-ecology The list of host plants of citrus yellow mite includes 15 different botanical species (Migeon and Dorkeld, 2006), including Citrus sp. in Japan (Ehara, 1955), India (Gupta and Gupta, 1994) and China (Wang, 1980; Chen and Li, 1985a; Li, 1990; Yang, 1998; Zhou et al., 1999) and C. reticulata in India (Manson, 1963b; Prasad, 1974), where the mite infests leaves, ﬂowers, shoots and fruitlets, but young leaves are usually attacked severely (Yang, 1998). The haploid number of chromosomes is four (Zou et al., 2002). The highest mean number of eggs (87.96 eggs/female), with a daily oviposition rate of 2.06 eggs per female per day, occurs at 20°C, whereas the highest daily oviposition rate (3.73 eggs per female per day) has been observed at 35°C, with a total of 55 eggs per female (Zhou et al., 1999). At 15°C, the lowest net reproductive rate of increase (R0) (16.64) and the longest generation time (T) (48.62 days) induced a lowest intrinsic rate of increase (rm) (0.058) and longest dt (11.95 days), whereas the highest intrinsic rate of increase (0.223) and a ﬁnite rate of increase (λ) (1.2493) were obtained at 35°C, and the population can double in only 3.11 days (Zhou et al., 1999). In spring, the dispersal pattern of the mite is of the gregarious translocation type. There is a signiﬁcant negative correlation between the population densities on the autumn and spring tips before the peak of mite population, but a negative one after the peak when numbers decline rapidly (Chen and Li, 1985a). The warm winter and infrequent rainfall registered in China, omitting orchard winter cleaning, not spraying chemicals in the early spring

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and outbreaks of mite populations, are related to irrational use of chemicals (Yang, 1998). Among the natural enemies are recorded the phytoseiid mite E. nicholsi (Zhi et al., 1994, 1998) and the stigmaeid Amblyseius exsertus (Wang, 1981b). Symptomatology and damage In Japan, the mite causes injury to citrus similar to that of the six-spotted spider mite, Euseius sexmaculatus (Ehara, 1964b). The injury is related to leaf age, mite abundance and soil moisture. Leaves infested earlier were able to compensate for damage more strongly than those infested later (Chen and Li, 1985b). Severe infestations on citrus trees cause leaves, ﬂowers and fruit to fall prematurely, and withering of branches (Chen, 1999). According to Li (1990), E. kankitus in China is thermophilic, producing shoot damage on citrus during warm, dry weather as the shoots elongated. Control Sampling methods for estimating the population density of E. kankitus in China has been investigated by Wang (1985), who has compared the methods of the insect density estimation technique and the sequential sampling technique, determining the practicality and effectiveness of the latter method by random sampling computer simulation. Spraying with pyrazoles in early April is quite effective (Yang, 1998), also with organotins and pyridazinones (Chen, 1999). 13.4.3.2.3 Eotetranychus lewisi (McGregor) (Fig. 13.24) Common name Lewis spider mite. Diagnostic characteristics FEMALE. The body is ovoid, about 360 μm long and pale greenish-amber in colour, with blackish spots along the lateral margin. The genital ﬂap and the area anterior to the ﬂap are both striate transversally. The peritremes are hooked distally. The tibiae I and II bear nine and eight tactile setae, respectively, and the tarsus I has ﬁve tactile setae proximal to the duplex setae (McGregor, 1950; Jeppson et al., 1975). MALE.

Smaller than the female, and mustard yellow in colour. The tibia I has nine tactile setae. The aedeagus gradually tapers distally and forms a broad, sigmoid ventral bend (McGregor, 1950; Jeppson et al., 1975). Geographical distribution Bolivia (Gonzalez, 1989; Flechtmann, 1996); Chile (Gonzalez, 1989); Colombia (Urueta, 1975); Costa Rica (Baker and Pritchard, 1962); El Slavador (Berry, 1959;

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A

B

Fig. 13.24. Eotetranychus lewisi (McGregor). Female. (A) Tibia and tarsus I (from Pritchard and Baker, 1955). Male. (B) Aedeagus (from Meyer Smith, 1987).

Andrews and Poes, 1980); Guatemala (Ochoa et al., 1991); Hawaii (Garrett and Haramoto, 1967); Honduras (Baker and Pritchard, 1962); Libya (Damiano, 1964); Madeira Island (Carmona, 1992); Mexico (Tuttle et al., 1974), Nicaragua (Baker and Pritchard, 1962); Panama (Ochoa et al., 1991); Peru (Bolland et al., 1998); South Africa (Meyer Smith, 1987); Taiwan (Ho and Shih, 2004); and the USA (McGregor, 1943). Bio-ecology The Lewis spider mite has been collected on about 66 spontaneous and cultivated plants (Bolland et al., 1998). It is recorded on C. limon and C. sinensis exclusive of the desert areas of California (McGregor, 1943, 1950, 1956; Jeppson et al., 1975; Jeppson, 1989), and in Israel (Jeppson, 1989). The mite reproduces by parthenogenesis arrhenotokous and the haploid number of chromosomes is two (Helle et al., 1981). The female lays two to three eggs per day in depressions of the citrus fruit surface. The egg has a short stalk, but no threads extend from the top of the stalk to the host. The life cycle from egg to adult lasts 12 days for the male and 14.5 days for the female. Each female lays two to three eggs per day (Stiller, in Jeppson et al., 1975). The natural enemies include the phytoseiid mites E. hibisci and Typhlodromalus limonicus (Garman et McGregor) (McMurtry, 1985). Recently, in

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Taiwan, N. longispinosus, E. ovalis, Paraseiulus minutus Athias Henriot, Ph. rugatus Tseng, some cecidomyiids, Scolothrips spp., Oligota sp., Lasioseius sp., stigmaeids and Orius sp. (Ho, 2006) have been collected on poinsettia. Symptomatology and damage The Lewis spider mite is injurious only for citrus fruit, its feeding resulting in a pale stippling of the rind. Commonly no injury is produced on the leaves, and heavy infestations cause silvering on lemon and either a silvering or russeting of oranges. The abundant webbing collects dust, making detection of heavy infestations easy (McGregor, 1956; Jeppson et al., 1975). Control The harvesting of the citrus fruit may remove most of the populations (Jeppson et al., 1975). Injurious populations may be controlled by treatments with sulfur or other acaricides (Jeppson, 1989).

13.4.3.2.4 Eotetranychus limonae Karuppuchamy et Mohanasundaram Common name Unkown. Diagnostic characteristics FEMALE.

The body is 405 μm long, 260 μm wide and yellowish-orange in colour. The dorsal setae are long, serrate and slightly longer than the distance between their bases; the dorsal opisthosomal setae h1 and f2 are shorter. The dorsal opisthosomal setae c1 are set on weak tubercles and d1, e1 and f1 on strong tubercles. The other dorsal setae lack tubercles. The prodorsum is longitudinally striate; the hysterosoma is transversally striate, except on the area between the dorsal setae e1 where the striae are longitudinal, and on the area between the dorsal setae f1 where the striae form a more or less diamond-shaped pattern. The striae are ornate with triangular lobes. The peritremes end in a bulb. The genital ﬂap and the anterior area are striate transversally. The tibia I and II bears eight and six tactile setae, respectively. On tarsus I, the duplex setae are distal and approximate (Karuppuchamy and Mohanasundaram, 1988). MALE. The body is 300 μm long and 145 μm wide. The aedeagus is bent dorsally and again bent posteriorly and tapers gradually, forming a sigmoid pattern (Karuppuchamy and Mohanasundaram, 1988).

Geographical distribution India (Karuppuchamy and Mohanasundaram, 1988).

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Bio-ecology The mite has been collected on C. limon in India (Karuppuchamy and Mohanasundaram, 1988), and its bio-ecology is unknown.

13.4.3.2.5 Eotetranychus limoni Blommers et Gutierrez (Fig. 13.25) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid, from 325 to 380 μm long and yellow coloured. The palptarsal eupathidial spinneret is about twice as long as broad. The genital ﬂap is transversely striate and the area anterior to the genital ﬂap is longitudinally striate. The empodium I possesses three pairs of hairs. The tibia I has eight tactile setae, one bothridial seta and one solenidion; tarsus I with 12 tactile setae, three eupathidia and three solenidia. Tibia II bears eight tactile setae and tarsus II 11 tactile setae, three eupathidia and two solenidia. Tarsus I has ﬁve tactile setae proximal to the duplex setae. The peritremes end hooked (Blommers and Gutierrez, 1975). MALE. Smaller than the female. The peritreme is similar to those of the female.

The tibia I has eight tactile setae, one bothridial seta and four solenidia; tarsus I has 12 tactile setae, three eupathidia and ﬁve solenidia. The duplex setae of the tarsus I possess a short, proximal tactile seta and a long, distal solenidion. Tibia II bears eight tactile setae, and tarsus II 11 tactile setae, three eupathidia and two solenidia, and duplex setae similar to those of tarsus I. The aedeagus is sigmoid, wider in the proximal part and tapering in the end, ﬁrst bent downward and then upward (Blommers and Gutierrez, 1975). Geographical distribution Madagascar (Blommers and Gutierrez, 1975). Bio-ecology The mite has been collected on C. aurantium, C. limon, C. sinensis, C. nobilis and C. paradisi in Madagascar. Immature and adult stages commonly feed on the lower surfaces of the leaves, and the females produce mostly silk and spin webs. The distribution of the mite populations on the plants seems independent of the age of the leaves (Blommers and Gutierrez, 1975). At 23 ± 2°C and 75 ± 10% RH, the egg, larva, protonymph and deutonymph develop in 5.9, 1.9, 2.2 and 2.1 days, respectively. The life cycle requires 16.2 days, pre-ovideposition 2–3 days and each female lays three eggs per day (Blommers and Gutierrez, 1975).

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A

B

D

E

C

F

Fig. 13.25. Eotetranychus limoni Blommers et Gutierrez. Female. (A) Genito-anal region; (B) palptarsus; (C) distal end of peritreme; (D) tibia and tarsus I. Male. (E) Palptarsus; (F) aedeagus (from Blommers and Gutierrez, 1975).

The symptomatology of feeding is similar to other Eotetranychus spp. living on citrus. The leaves show patchy light yellow and some swelling of the surface (Blommers and Gutierrez, 1975). Several phytoseiid mites develop on E. limoni in Madagascar, the most important being A. parasundi (Blommers and Gutierrez, 1975). Among

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recorded insects are the coccinellid S. madecassus (Blommers and Gutierrez, 1975). Control is unjustiﬁed.

13.4.3.2.6 Eotetranychus mandensis Manson (Fig. 13.26) Common name Unknown. Diagnostic characteristics FEMALE.

The body is ovoid and about 270 μm long. The palptarsal eupathidial spinneret is about twice as long as broad. The peritremes end in a simple bulb. The dorsal body setae are slender and tapering, and the dorsal opisthosomal are about as long as or longer than the distances between their bases. The dorsal striae between the dorsal opisthosomal setae e1 are longitudinal. The genital ﬂap and the area anterior to the genital ﬂap are both transversally striate. The tibia I bears nine tactile setae and one sensory seta; tarsus I with 13 or 14 tactile setae and one sensory seta, as well as the duplex setae. Tibia II with six tactile setae, rarely ﬁve; tarsus II with 12 tactile setae, one sensory seta and one duplex seta (Manson, 1963b). MALE.

Smaller than the female and about 205 μm long. The palptarsal eupathidial spinneret is absent. The aedeagus is long, slender and tapering, with distal portion directly posterior (Manson, 1963b). Geographical distribution India (Manson, 1963b; Gupta and Gupta, 1994). Bio-ecology The species has been collected on Citrus sp. in India (Manson, 1963b; Gupta and Gupta, 1994).

13.4.3.2.7 Eotetranychus pamelae Manson (Fig. 13.27) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid and about 385 μm long. The palptarsal eupathidial spinneret is about twice as long as broad. The peritreme ends simply and bent at a right angle. The dorsal body setae are long and tapering, longer than the distances between their bases. The dorsal striae are transverse between the dorsal opisthosomal setae e1. The genital ﬂap and the area

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A

B

C

Fig. 13.26. Eotetranychus mandensis Manson. Female. (A) Tibia and tarsus I. Male. (B) Tibia and tarsus I; (C) aedeagus (from Manson,1963b).

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A B

C

D

E

Fig. 13.27. Eotetranychus pamelae Manson. Female. (A) Tibia and tarsus I; (B) distal end of peritreme. Male. (C) Palptarsus; (D) aedeagus; (E) tibia and tarsus I (from Manson, 1963b).

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anterior to the genital ﬂap are striate longitudinally, whereas the area posterior to the genital ﬂap has transverse striae. Tibia I bears nine tactile setae and one sensory seta; tarsus I bears 14 tactile setae and one sensory seta, as well as the two duplex setae. Tibia II has eight tactile setae; tarsus II possesses 13 tactile setae and one sensory seta, as well as one duplex seta. The empodial spur is prominent on all legs (Manson, 1963b). MALE.

Smaller than the female and 269 μm long. The palptarsal eupathidial spinneret is longer than broad, gradually narrowing from base to apex and rather sharply pointed. The peritremes end somewhat variably, but usually sharply bent at an angle of about 90°. The aedeagus is short, directly posterior and ending bluntly (Manson, 1963b). Geographical distribution India (Manson, 1963b; Gupta and Gupta, 1994). Bio-ecology The mite has been collected on Citrus sp. in India (Manson, 1963b; Gupta and Gupta, 1994). Its bio-ecology is unknown.

13.4.3.2.8 Eotetranychus sexmaculatus (Riley) (Fig. 13.28) Common name Six-spotted spider mite. Diagnostic characteristics FEMALE. The body is ovoid, about 300 μm long and lemon yellow in colour, commonly with blackish spots organized in three blotches along each side of the body, and sometimes these spots are inconspicuous or absent. Tibiae I and II bear nine and eight tactile setae, respectively. Tarsus I possesses ﬁve tactile setae proximal to duplex setae. The peritreme is hooked distally. The genital ﬂap is longitudinally striate on the anterior central part and on the area anterior to the ﬂap (Jeppson et al., 1975). MALE.

Smaller than the female and pale. The tibiae I and II are similar to those of the female. The aedeagus curves dorsally near the middle of the shaft, and is distally direct caudoventrally with the tip deﬂected (Jeppson et al., 1975). Geographical distribution Australia (Froggatt, 1921); China (Ma et al., 1979; Wang, 1980; Wang et al., 1985; Yang, 1987); Formosa (Pritchard and Baker, 1955); Hainan Island (Ma et al., 1979); Hawai (Garrett and Haramoto, 1967); Korea (Rep. South) (Lee, 1989); India (Gupta and Gupta, 1976, 1994); Iraq (Mohamed and El-Haidari,

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A

B

Fig. 13.28. Eotetranychus sexmaculatus (Riley). Female. (A) Tibia and tarsus I. Male. (B) Aedeagus (from Pritchard and Baker, 1955).

1965, 1968); Japan (Ehara, 1956); Korea (Lee, 1989); New Zealand (Manson, 1983); Okinawa Island (Ehara, 1975b); Taiwan (Pritchard and Baker, 1955; Ehara, 1969); and the USA (Riley, 1890; Pritchard and Baker, 1955). Bio-ecology The list of host plants of the six-spotted mite numbers 43 spontaneous and cultivated species (Migeon and Dorkeld, 2006), including C. grandis, C. limon, C. sinensis (Pritchard and Baker, 1955) and Citrus sp. (Riley, 1890; McGregor, 1950; Pritchard and Baker, 1955; McGregor, 1956; Tuttle and Baker, 1964; Gupta and Gupta, 1994). In Florida, E. sexmaculatus has been a pest of oranges since at least 1885, and in California it attacks citrus only in coastal areas (McGregor, 1956; Jeppson, 1989). The mite feeds on the lower surfaces of the leaves, near the midrib or the principal veins, and only at high density does it also infest the fruit. It produces abundant web, which covers their populations, protecting them from predators and offering support for the eggs. The female:male ratio in the progeny of mated females is 2.5:1. Mated females reproduce arrhenotokously, and fecundity reduces as the female’s age increases. The number of females in the progeny of newly emerged and mated females is relatively high, and decreases with increased age of females (Yang, 1987).

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The pre-oviposition period is 2–3 days; each female lays 25–40 eggs in 10–20 days (Jeppson et al., 1975). The number of eggs laid by mated females (35.16 ± 6.90) is greater that than laid by unmated ones (20.95 ± 3.85) (Yang, 1987), and egg development in summer lasts from 5 to 8 days and the developmental stages each last 2–3 days (Jeppson et al., 1975). In southern California, the six-spotted spider mite is restricted to the coastal areas, and although at times it can be a serious pest, it does not usually cause severe damage (McGregor, 1956). Population dynamics is adversely inﬂuenced by dry weather conditions, and the populations reach injurious levels only in the more humid coastal regions. The dry winds reduce the development of mite populations, with a gradual increase during winter and a rapid one in spring and summer (Jeppson et al., 1975; Jeppson, 1989). The factors limiting the development of populations include several natural enemies, including pathogens, predatory insects and mites. Among the pathogens recorded are Entomophthora sp. (Entomophthoraceae) (Muma, 1969), and H. thompsonii (Clavicipitaceae) (McCoy and Selhime, 1974). The insects include the coccinellid Stethorus utilis Horn, the aeolothripid Scolothrips sexmaculatus (Pergande), the chrysopid Chrysopa lateralis Guérin and the coniopterygid Coniopteryx vicina Hagen (McMurtry et al., 1970). The commonest predatory mites are the stigmaeids and phytoseiids. Among the ﬁrst recorded were A. exsertus (Wang, 1981b; Tna, 1986) and A. terminalis (Wang, 1981b). Recently recorded are A. herbicolus (Chang and Chen, 1984), Amblyseius maai Tseng (Chang and Chen, 1984), Chelasius ﬂoridanus (Muma) (Muma, 1970), E. hibisci (Tanigoshi et al., 1981; McMurtry, 1985), E. mesembrinus (McCoy and Rakha, 1985; Abou-Setta and Childers, 1989), E. ovalis (Chang and Chen, 1984), G. helveolus (Caceres and Childers, 1991), G. occidentalis (Solow and Sherman, 1997), Neoseiulus collegae (De Leon) (Mizell and Schiffhauer, 1991), Neoseiulus taiwanicus (Ehara) (Chang and Chen, 1984), N. womersleyi (Osakabe, 2002), T. limonicus (McMurtry, 1985) and Typhlodromips okinawanus (Ehara) (Chang and Chen, 1984). Symptomatology and damage The six-spotted spider mite feeds on the undersides of leaves and normally does not inhabit the fruit except during severe infestations. Leaves near the ground are attacked ﬁrst, and only after some time may the leaves more than about 1 m from the ground be injured (Jeppson et al., 1975). Feeding of pests partly or completely evacuates the spongy mesophyll cells during feeding from the lower surface of the leaves. There is some recovery several weeks following injury, which appeared to be related to physical damage or carbohydrate depletion but not to toxins or growth stimulants (Albrigo et al., 1983). Feeding on the lower surfaces of the leaves produces yellow depressions on the undersides of the leaves, covered by webbing. The areas of the upper surfaces corresponding to the sides of the lower surfaces where the mite colonies live become raised and yellow or yellow–white, with a smooth, thin surface. With the increase of the infestation, the yellowish areas converge and the leaves become entirely yellow, distorted or

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misshapen and drop prematurely (McGregor, 1956; Jeppson et al., 1975; Jeppson, 1989). Control The webbing covering the mite colonies probably protects the pest from predators, and the increase in prey numbers represents a source of food for predators. The action of phytoseiid mites, together with unfavourable weather, reduces the mite to one of occasional importance on citrus in California and Florida (Jeppson et al., 1975). As regards chemical control, it is possible to employ the same acaricides used against the citrus red mite. Petroleum oils (Jeppson et al., 1975) or sulfur can both reduce populations (Jeppson, 1989).

13.4.3.2.9 Eotetranychus yumensis (McGregor) (Fig. 13.29) Common name Yuma spider mite. Diagnostic characteristics FEMALE. The body is ovoid, about 320 μm long and 180 μm wide, varying in colour from straw colour to dark pink. The dorsal body setae are ﬁnely setose, each reaching well beyond the base of seta, just behind and not set on the tubercles. Tibiae I and II bear nine and eight tactile setae, respectively. Tarsus I possesses four tactile setae proximal to the duplex setae. The end of the peritremes is strongly hooked. The genital ﬂap is striate transversally, with longitudinal striae in the area anterior to the ﬂap (McGregor, 1950; Jeppson et al., 1975). MALE. Smaller than the female and pale. The aedeagus forms a shallow, nearly

sigmoid dorsal curve, with the distal end gradually tapering to a caudally direct point (McGregor, 1950; Jeppson et al., 1975). Geographical distribution Mexico (Tuttle et al., 1974) and the USA (McGregor, 1934, 1956). Bio-ecology The Yuma spider mite is recorded in over 20 host plants (Bolland et al., 1998), and among these are reported C. limon (McGregor, 1950; Pritchard and Baker, 1955; Tuttle and Baker, 1964); C. paradisi (McGregor, 1950; Tuttle and Baker, 1964); C. reticulata (McGregor, 1950; Pritchard and Baker, 1955; Tuttle and Baker, 1964); C. sinensis (McGregor, 1950; Tuttle and Baker, 1964); and Citrus sp. (McGregor, 1934, 1950, 1956; Elmer, 1965, 1969; Jeppson et al., 1975). On citrus the mite infests leaves, fruit and twigs, and the development of its

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Fig. 13.29. Eotetranychus yumensis (McGregor). Aedeagus (from Pritchard and Baker, 1955).

populations seems restricted to desert areas. It feeds chieﬂy on the under surface of the leaves and produces quantities of webbing to which dust adheres, making it easy to detect dense mite colonies (McGregor, 1956; Jeppson et al., 1975). Reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is two (Helle et al., 1981). Relatively high temperatures are necessary for the development of E. yumensis. The mite does not lay eggs at constant temperatures of 10°C, and at 15.5°C the eggs do not hatch if the RH is below 50%. At temperatures between 21°C and 38°C complete development occurs, but at 43.5°C the eggs do not hatch. This adaptation probably conﬁnes the species to the hot desert areas (Elmer, 1969; Jeppson et al., 1975). The populations of Yuma spider mite increase in October and November, remain high throughout the winter and decrease in spring or early summer. During the summer, the species aestivate as female stages under the bark of the trunk and limbs or in cracks and crevices, and it is possible to ﬁnd a few eggs, but not immature stages (Jeppson et al., 1975). Predatory insects and mites are commonly associated with populations of Yuma spider mite, similarly to other Eotetranychus species (McMurtry, 1985).

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Symptomatology and damage The mite feeds on leaves, fruit and green twigs of citrus and produces a silvering of mature fruit. The increase of populations accentuates the development of dieback, normally caused by low RH, wind and other factors that either increase leaf transpiration or otherwise lead to the leaves to developing a moisture deﬁciency. High levels of infestation produce dry leaves and fruit drops, while young twigs and even whole limbs become devoid of leaves and die (Jeppson et al., 1975). Control Chemical control may be entrusted to the use of sulfur during the autumn, winter or early spring, and treatment should be applied during cool weather periods (Elmer, 1965; Jeppson, 1989). Alternatively, it is possible to employ speciﬁc acaricides.

13.4.3.3 Mixonychus Meyer et Ryke The opisthosoma bears ten pairs of stout dorsal setae, with a reticulate pattern. The empodial claw is strong, much longer than the pads of the true claws, and without proximoventral hairs (Bolland et al., 1998).

13.4.3.3.1 Mixonychus ganjuis Qian, Yan et Ma Common name Unknown. Diagnostic characteristics FEMALE. The body is oval, and including the gnathosoma is 519 μm long, 256 μm wide and orange in colour. The gnathosoma is slender, reaching the distal end of genu I. The prodorsum has mostly irregular longitudinal and transverse striae, the mediolateral area of the metapodosoma presents dorsal reticulations and the posterolateral area of the opisthosoma with dorsal and ventral reticulations. The dorsal setae are 13 pairs, all strongly setiform, pubescent, set on prominent tubercles, each slightly enlarged near the distal end. The stylophore is rounded anteriorly. Tarsus I bears two pairs of duplex setae, and tarsus II one single pair of duplex setae. The empodial claw is without proximoventral hairs. The species is similar to Mixonychus acaciae Ryke et Meyer, 1960 in having the empodial claw without proximoventral hairs, and dorsal integument with reticulate pattern, but Mixonychus ganjuis is distinct, with the stylophore rounded anteriorly (Qian et al., 1980). MALE.

Unknown (Qian et al., 1980).

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Geographical distribution China (Qian et al., 1980). Bio-ecology The species has been collected on C. reticulata and Amelanchier spp. in the Sichuan province of China (Qian et al., 1980).

13.4.3.3.2 Mixonychus ziolanensis (Lo et Ho) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid, 428 μm long, 247 μm wide and orange in colour. The dorsal body setae are strongly serrate and rod-like, most slightly shorter than the distances between them and set on tubercles. The prodorsal vertical setae v2 are slightly shorter than the scapular setae sc1, and the prodorsal central area possesses reticulate elements and, laterodorsally, complete and incomplete reticulate elements. The dorsal opisthosomal setae f2 are two-thirds as long as setae f1. The dorsocentral area mostly has transverse striae, except for longitudinal striae between the setae e1 and f1. The peritremes end in a simple bulb and a short hook (Lo and Ho, 1989). MALE.

Smaller than the female, with dorsal setae and striae similar to those of the female. The shaft of the aedeagus is broad and curving dorsally to form a knob, with an acuminate anterior projection and a longer, upturned posterior projection; the knob is less than the length of dorsal margin of shaft and forms an acute angle with the shaft axis (Lo and Ho, 1989). Geographical distribution Taiwan (Lo and Ho, 1989). Bio-ecology The species has been collected on C. sinensis and C. nobilis in Taiwan (Lo and Ho, 1989).

13.4.3.4 Oligonychus Berlese The opisthosoma possesses ten pairs of dorsal setae. The anal region has two pairs of pseudanal setae (ps1–2) and one pair of setae h. The empodium is claw-like and has proximoventral hairs. All the legs or most of them bear

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empodial claws as long as or longer than the proximoventral hairs. The duplex setae of tarsus I are distal and adjacent (Bolland et al., 1998). 13.4.3.4.1 Oligonychus biharensis (Hirst) (Fig. 13.30) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid, with dorsal body setae longer than the distances between them and dorsal striae transverse. The peritremes are retrorse distally. The terminal sensillum of the palptarsus is short and broad. Tibia I has nine tactile setae and one solenidion. Tarsus I possesses four tactile setae and one solenidion proximal to duplex setae (Pritchard and Baker, 1955; Meyer Smith, 1974). MALE.

The terminal sensillum of the palptarsus is about three and a half times as long as broad, and the tibial claw is dentate. Tibia I bears nine tactile setae and four solenidia, and tibia II seven tactile setae. The shaft of the aedeagus curves dorsally and forms at the tip a large knob longer than the dorsal margin; the anterior projection is small and acute, and the posterior projection is long and curves downwards; the axis of the knob is parallel to the axis of the shaft, and the dorsal margin of the knob is convex (Pritchard and Baker, 1955; Meyer Smith, 1974). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology Oligomychus biharensis is tropical and subtropical and has been collected on 69 host plants, including Citrus sp. (Migeon and Dorkeld, 2006). The reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is two (Helle and Gutierrez, 1983). According to Saitô (1985), the mite belongs to the Wn-u subtype, serving the web production to shelter it from adverse climatic factors and predators. Among these are recorded some aeolothripid (Chazeau, 1985) and phytoseiid mites, such as E. ovalis (Sabelis, 1985) and N. longispinosus (Kongchuensin et al., 2005).

13.4.3.4.2 Oligonychus coffeae (Nietner) (Fig. 13.31) Common name Tea red spider mite, red tea mite.

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C

B

Fig. 13.30. Oligonychus biharensis (Hirst). (A) Tibia and tarsus I of female; (B) aedeagus (from Baker and Pritchard, 1960); (C) pretarus I of female (from Pritchard and Baker, 1955).

Diagnostic characteristics FEMALE .

The body is ovoid, with the propodosoma bright red and the hysteroma wine coloured. The peritremes end in a bulb. The dorsal body setae are serrate and longer than the distances between consecutive setae. The opisthosoma is striate, mostly transversally, but sometimes U-shaped or irregular between the third pair of dorsal setae e1. Some-

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A

B

C

D

E

Fig. 13.31. Oligonychus coffeae (Nietner). Female. (A) Tibia and tarsus I; (B) pretarsus I (from Pritchard and Baker, 1955); (C) palptarsus. Male. (D) Palptarsus; (E) aedeagus (from Meyer Smith, 1974).

times the striae between the dorsal setae f1 are irregular to irregular longitudinal. The striae anterior to genital flap are longitudinal to transverse. Tibia I bears seven tactile setae. Tarsus I possesses three tactile setae and one solenidion well proximal to duplex setae, and one tactile setae occurs opposite duplex setae on ventral side. Tarsus II has two tactile setae well proximal to duplex setae and one tactile setae ventral to duplex setae (Pritchard and Baker, 1955; Meyer Smith, 1974; Jeppson et al., 1975). MALE.

Smaller than the female and lighter in colour. The palptarsus has a tiny terminal sensillum. Tarsus I bears three tactile setae and two solenidia well proximal to duplex setae; the empodia have ﬁve pairs of proximoventral hairs. Distally, the aedeagus bends ventrally at a right angle to the shaft axis and gradually narrows to a slender end (Pritchard and Baker, 1955; Meyer Smith, 1974; Jeppson et al., 1975). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006).

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Bio-ecology The species is pantotropical and recorded on about 125 host plants (Migeon and Dorkeld, 2006); it was collected also on Citrus sp. in South Africa (Baker and Pritchard, 1960; Meyer Smith, 1974), India (Gupta and Gupta, 1994) and Taiwan (Ehara, 1969). In tropical and subtropical areas, the mite is considered the most serious of all tea and coffee pests and also attacks cotton (Jeppson et al., 1975; Meyer Smith, 1981), but is not injurious to citrus. Its reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is three (Helle et al., 1970). The tea red spider mite feeds on the upper surface of the leaves of the host plant and produces little webbing. At 22°C, the females lay 40–50 eggs during their life span. In these conditions, the life spans of larva, protonymph and deutonymph are 8, 40 and 2–3 days, respectively. The development from egg to adult requires 14–15 days. The temperature and humidity limit the development of mite populations. Under constant temperatures of 34°C and 17% RH, the eggs fail to hatch; at 33°C and RH below 72%, the eggs do not hatch, whereas with RH just above 72% some hatching occurs. The optimal conditions for development are 20–30°C and 49–94% RH. In these conditions the mite may complete 22 generations in a year on tea. In north-east India, the mite can be found throughout the year. The increase of populations occurs in early March and the higher densities are observed in late March and early April. Injury to the leaves is severe during May and June or until the monsoon rains. The attacks are renewed after the rainy season, but never become as injurious as during the pre-monsoon period. In cool weather, the densities of the populations are lower and damage is uncommon. On tea, pruning removes many of the old leaves and the shoots, and thus a large part of the mite populations (Das, 1959, 1960; Hu and Wang, 1965; Das and Das, 1967). Coccinellids prey on eggs and other biological stages of O. coffeae (Das, 1959). Among these are recorded Stethorus comoriensis Chazeau (Chazeau, 1971), S. madecassus (Blommers and Gutierrez, 1975) and Stethorus gilvifrons (Mulsant) (Banerjee and Cranham, 1985). Other predatory insects include chrysopids (Meyer Smith, 1981; Banerjee and Cranham, 1985) and staphylinids (Banerjee and Cranham, 1985). Also included are some phytoseiids such as Amblyseius parasundi (Blommers and Gutierrez, 1975), N. longispinosus (Kongchuensin et al., 2005)) and other secondary species, and some stigmaeid mites (Banerjee and Cranham, 1985).

13.4.3.4.3 Oligonychus gossypii (Zacher) (Fig. 13.32) Common name Unknown.

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A

B

Fig. 13.32. Oligonychus gossypii (Zacher). Female. (A) Tibia and tarsus I (from Baker and Pritchard, 1960). Male. (B) Aedeagus (from Meyer Smith, 1974).

Diagnostic characteristics FEMALE.

The body is ovoid, and reddish in basic colour. The dorsal body setae are serrate and longer than the distances between their bases. The peritremes are strongly retrorse distally. The opisthosoma is longitudinally striate between the dorsal setae e1 and f1 and with a diamond-shaped ﬁgure. The tibia I bears nine tactile setae and tibia II seven tactile setae. Tarsus I has four tactile setae proximal to the duplex setae, and two ventral setae proximal to the duplex setae (Meyer Smith, 1974; Jeppson et al., 1975).

MALE.

Smaller than the female. The peritremes are similar to those of the female. The terminal sensillum of the palptarsus is about four times as long as broad. The aedeagus narrows distally and curves dorsally at about a right angle; distally, it presents a large knob with a small anterior projection and a long, undulate posterior projection, and the tip is direct ventrally. Tarsus I has four tactile setae and three solenidia proximal to the duplex setae. The empodia possess a claw and three proximoventral hairs (Meyer Smith, 1974; Jeppson et al., 1975).

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Geographical distribution Angola (Carmona, 1967); Benin (Carmona, 1967); Brazil (Flechtmann and Baker, 1970); Cameroun (Meyer Smith and Bolland, 1984); Central Africa Rep. (Gutierrez, 1987); Colombia (Urueta, 1975); Congo (Gutierrez and Etienne, 1981); Congo (RDC, ex-Zaire) (Baker and Pritchard, 1960); Costa Rica (Byrne et al., 1983); Ecuador (McGregor, 1955); Ethiopia (Matthysse, 1980); Guinea-Bissau (Hirst, 1926); Honduras (Baker and Pritchard, 1962); Kenya (Nyiira, 1982); Madagascar (Helle et al., 1970); Nigeria (Baker and Pritchard, 1960); Sao Tome and Principe (Hirst, 1926); Senegal (Gutierrez and Etienne, 1981); Sierra Leone (Hirst, 1926); Tanzania (Nyiira, 1982); Togo (Zacher, 1921); Uganda (Nyiira, 1982); Venezuela (Byrne et al., 1983); and Zaire (Baker and Pritchard, 1960). Bio-ecology The species is common in African and South American equatorial areas, recorded in over 48 host plants (Bolland et al., 1998; Migeon and Dorkeld, 2006) and on citrus in Zaïre (Baker and Pritchard, 1960) and Colombia (Urueta, 1975). It produces much webbing and lives on both leaf surfaces. It is a pest of cotton (Jeppsn et al., 1975; Meyer Smith, 1987). Reproduction is characterized by parthenogenesis arrhenotokous and the haploid number of chromosomes is two (Helle et al., 1970).

13.4.3.4.4 Oligonychus peruvianus (McGregor) (Fig. 13.33) Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid and yellowish green in colour. The dorsal body setae are short, strongly lanceolate, nearly nude and broadest near the base. The peritremes end simply. The opisthosoma presents dorsal longitudinal striae between the dorsal setae e1. The tibia I has nine tactile setae and one solenidion. The tarsus I bears four tactile setae and one solenidion proximal to the duplex setae, and one single ventral tactile seta. The tibia II has seven tactile setae. The empodium possesses three pairs of proximoventral hairs (McGregor, 1950; Pritchard and Baker, 1955). MALE.

Smaller than the female. The tibia I bears nine tactile setae and two solenidia, and tibia II seven setae. Tarsus I has four tactile setae and three solenidia proximal to the duplex setae, and one single ventral tactile seta. The aedeagus bends ventrally, and the distal part is short (McGregor, 1950; Pritchard and Baker, 1955).

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C

B

Fig. 13.33. Oligonychus peruvianus (McGregor). (A) Dorsal view of female; (B) aedeagus (from Jeppson et al., 1975); (C) tibia and tarsus I of female (from Pritchard and Baker, 1955).

Geographical distribution Colombia (Thewke and Enns, 1969); Costa Rica (Thewke and Enns, 1969); Ecuador (McGregor, 1955); Guatemala (Baker and Pritchard, 1962); Mexico (Estebanes Gonzalez and Baker, 1968); Peru (McGregor, 1917); Trinidad and Tobago (Hirst, 1922); USA (Pritchard and Baker, 1955); and Venezuela (Doreste, 1982). Bio-ecology The species is recorded on several host plants, including C. aurantium (Doreste, 1967) and Citrus sp. in Central America (Anon., 1973). It feeds in restricted colonies on the undersides of the leaves of the host plants under

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small, compact webs, resembling the egg covering of certain true spiders (McGregor, 1950; Baker and Pritchard, 1953b; Estebanes Gonzalez and Baker, 1968). Its bio-ecology is unknown.

13.4.3.5 Panonychus Yokoyama The empodial claw is as long as or longer than the proximoventral hairs, which are inserted at right angles to the claw (Bolland et al., 1998).

13.4.3.5.1 Panonychus citri (McGregor) (Fig. 13.34) Common name Citrus red mite. Diagnostic characteristics FEMALE. The length of the body ranges from 350 to 400 μm and the colour is deep red to purplish. The legs I are shorter than body. All dorsal setae are set on strong tubercles of the same colour as the body, and longer than the distance between their bases. The prodorsal vertical setae v2 are shorter and scapular setae sc1 longer. The opisthosoma is striate transversally and the striae bearing lobes conical and narrow; ventrally, the lobes are broad and round. The dorsal opisthosomal setae f2 and h1 are of equal length and both as long as about one third of setae f1. The peritremes end in a simple bulb. Tarsus I bears three tactile setae and one solenidion proximal to duplex setae (Vacante, 1985). MALE.

Smaller than the female, and light in colour, sometimes orange. The aedeagus is sigmoid, with broad base and distal end pointed and turned upward (Vacante, 1985). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology

The citrus red mite is recorded on 111 host plants (Migeon and Dorkeld, 2006) and is very common on citrus, where it has been reported on C. mitis (Blanco) Ingram et Moore (Vierbergen, 1994); C. aurantifolia (Doreste, 1967); C. aurantium (Rodrigues, 1968; Gupta and Gupta, 1994); C. grandis (Gotoh et al., 2003); Citrus hassaku Tanaka; C. junos Siebold ex Tanaka (Gotoh et al., 2003); C. limon (McGregor, 1950); C. medica (Gotoh et al., 2003); C. mitis (Bowman and Bartlett, 1978); Citrus natsudaidai Hayata, C. paradisi (McGregor, 1950); C. reticulata (McGregor, 1950); C. sinensis (McGregor, 1950); Citrus sudachi Hort. ex Shirai; Citrus tamurana Hort. ex Tanaka (Hyuganatsu); C. unshiu (Gotoh et al., 2003);

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A

B

C

E

D

Fig. 13.34. Panonychus citri (McGregor). (A) Dorsal view of female; (B) receptaculum seminis (from Meyer Smith, 1987); (C) distal end of peritreme of female; (D) aedeagus; (E) pretarsus I of female (from Vacante, 1985).

Citrus sp. (McGregor, 1916; Rambier, 1958; Baker and Pritchard, 1960; Baker and Pritchard, 1962; Ferragut et al., 1983; Garcia Marì et al., 1987; Gupta and Gupta, 1994; Vierbergen, 1994; Gupta and Chatterjee, 1997; Kreiter et al., 2002; Gotoh et al., 2003; Suarez, 2004); Fortunella spp. (Vierbergen, 1994; Gotoh et al., 2003); and Poncirus trifoliata (Ehara, 1955). The species reproduces through parthenogenesis arrhenotokous and the virgin females lay eggs hatching into only males, whereas mated females produce both sexes. The karyotype is n = 3, 2n = 6 or n = 4, 2n = 8. The chromosomes are holokinetic, with mixoploidy in the karyotypes (Zou et al., 2002). The mite presents diapausing and non-diapausing forms characterized by morphological differences (Okasabe and Saito, 1991), and the diapause occurs

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through winter eggs (Shinkaji, 1979; Takafuji and Morimoto, 1983). The ratio of females to males on leaves and on fruit of lemon is 2.7:1 and 3.1:1, respectively (Chiavegato, 1988). Panonychus citri develops on leaves, twigs and mature or green immature fruit, and prefers light green maturing growth, feeding commonly in greatest numbers on the upper surface of the young and middle-aged leaves. It has been observed that 60% of populations live in the exterior canopy and 80% on the upper side of the leaves. The mite prefers the green to the yellow fruit, and in winter in California on mature fruit orange, the densities of populations are higher than on the leaves (Mico et al., 1992). On lemon, the species develops more or less constantly, where new growth and new fruits are available (Jeppson et al., 1975). In Korea, it has been observed that on citrus in autumn the density of populations of citrus red mite is higher on fruit than on leaves according to fruit maturing level (Song et al., 2003), but laboratory experiments have shown that lemon leaves are more suitable than fruits for the development of the pest (Chiavegato, 1988). The age of the leaf and level of injury of the feeding activity inﬂuence the mite populations. The number of eggs laid on young and middle-aged leaves is greater than on old leaves, but the feeding injury inﬂuences oviposition (Henderson and Holloway, 1942) and cannot represent a regulative factor in the density of populations. Furthermore, a direct relationship exists between the density of populations and the nitrogen levels of the host plant (Schwartz, 1984). At 26°C the life cycle is completed in 14 days, and at 10°C it is ﬁve times longer; furthermore, at 10°C it is nine times longer than at 27°C. The range of development with high temperatures is balanced by a life cycle longer in colder conditions (English and Turnipseed, 1941; Shinkaji, 1954). On lemon and Valencia orange fruit at 23.9°C and 56% RH, the eggs hatch in 7–9 days, while the larval, protonymphal and deutonymphal stages last 2, 2 and 1–3 days, respectively. The males hatch before the females and oviposition reaches its maximum within 7–8 days after the females’ birth. In these conditions, the life cycle from egg to egg takes 17 days. The longevity of male and female is 22 and 27 days, respectively (Keetch, 1968). At 25 ± 1°C, 58 ± 5% RH and light to dark 24:0 with leaves and fruits of lemon, 28.9 and 13.0 days are necessary for the female life span, and 9.6 and 10.9 days for the male life span, respectively. The number of eggs per female is 77.2 and 37.7 on leaves and in fruits, respectively (Chiavegato, 1988). The development threshold for the complete cycle is 10.82°C and the thermal constant is 151.61 degree-days (Delrio and Monagheddu, 1986). The instantaneous rates of reproduction are 0.1723 at 30°C and 0.1688 at 25°C, and no reproduction occurs at 15°C (Muraoka and Shiyomi, 1986). At 25°C, the intrinsic rate of increase (rm) is 0.209/day and the net reproduction rate (R0) is 46.61 (Gotoh et al., 2003). The temperature represents the major factor affecting mite development (Keetch, 1968). Temperatures of 40.5°C, or several days of hot dry weather (5% RH and 32°C) together with strong wind, frequently gie rise to a high mortality. Similar conditions may be observed during the summer in California, but they can occur also in the winter. Extended periods of high RH are also

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unfavourable for the development of mite populations. The susceptibility to extreme conditions of both temperature and RH limits the mite distribution and affects the seasonal trends of the populations (Jeppson et al., 1975). Study of the direct inﬂuence of irrigation on citrus red mite development has shown than the direct effects of irrigation on yield are greater than the physiological effects on the trees and on mites’ development (Hare et al., 1989). Direct and interacting effects of four seasons of uncontrolled citrus red mite populations with 36 other combinations of grove management practices have been examined by Hare et al. (1992), to investigate the existence of the long-term deleterious consequences of not controlling mite populations on Navel orange. Effectiveness of the acaricide, densities of mites/leaf, differential acaricide application, fertilization level and differential irrigation cause yield reductions and/or produce reduction in fruit size. Total yield and average fruit size did not differ consistently because of any other interactions between effective P. citri treatments and other cultural practices. Marketing conditions continues to favour large fruit; thus, increases in average fruit size on trees where citrus red mite populations are not effectively suppressed continue to be at least a compensating, if not beneﬁcial, consequence of holding acaricide applications. The dynamics of populations vary from season to season, depending on climatic conditions, chemical control of different citrus pests and other factors. Sometimes the mite seems to disappear for some years. McGregor (1956) reported that, in the interior areas of Florida and California, the period of heaviest occurrence of the citrus red mite is in autumn and early winter. Jeppson (1978) indicates that two peaks of populations occur, in spring or early summer and in autumn or early winter. In South Africa, the maximum number of mites has been observed during spring and autumn (Smith, 1953; Stofberg, 1959; Brodrick, 1965), or in some countries from November to January (Van Rooyen, 1966). In Spain, the seasonal trend of the mite populations has shown an increase during August–September, followed by a drop or maintenance in the population levels until winter (Garcia Marì et al., 1983). In Italy, the mite populations increase commonly in spring and autumn, with a pause or lowering of reproduction in summer and winter, with some countries where in environmental factors permit, mite development can occur in those seasons (Vacante, 1995, 2009a). Normally, the citrus red mite does not infest crops uniformly, with damage varying from tree to tree. The inﬂuence of dust covering the tree alongside orchard roads on predators as well known (Van Rooyen, 1966). Furthermore, the use of different insecticides for scale insects or other pests control can stimulate, directly (Schwartz, 1978; Jones and Parrella, 1984a) or indirectly (Keetch, 1968; Grout and Richards, 1992), citrus red mite development for the elimination of predacious mites. The species is dispersed by wind, and by human movement labour and farm implements, and by harvesting or various machines. Natural enemies include pathogens, predatory insects and mites. As regards pathogens, in various regions of the world a non-inclusion virus and various species of fungi are recorded (Table 13.6).

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Table 13.6. Panonychus citri (McGregor): natural enemies (viruses, fungi, insects and mites). Natural enemies

Reference(s)

Viruses Non-inclusion virus

Munger et al., 1959, 1979; Rath, 1991

Fungi Clavicipitaceae Beauveria bassiana (Balsamo) Vuillemin Hirsutella thompsonii Fisher Entomophthoraceae Entomophthora sp. Exobasidiomycetidae Acaromyces ingoldii Boekhout et al. Meira geulakonigii Boekhout et al. Meira argovae Boekhout et al. Insects Coccinellidae Stethorus spp. Coniopterygidae Conwentzia barrette (Banks) Coniopteryx vicina Hagen Semidalis fuelleborni Enderlein Coniopteryx (Holoconiopterix) turneri Kimmins Conwentzia sp. Semidalis meridionalis Kimmins Conwentzia capensis Tjeder Conwentzia psociformis (Curtis) Endomychidae Saula japonica Gorham Staphylinidae Oligota spp. Oligota pygmea Solier Reduviidae Ploearia culiciformis (de Gen.) Mites Bdellidae Bdellodes longirrostris (Hermann) Phytoseiidae Amblyseius eharai Amitai et Swirski Amblyseius herbicolus (Chant) Euseius hibisci (Chant) Euseius addoensis (van der Merwe et Ryke) Euseius citri (van der Merwe et Ryke)

Shi and Feng, 2006 Rath, 1991 Fisher, 1951 Paz et al., 2009a Sztejnberg et al. , 2004; Paz et al., 2009a Paz et al., 2009a

McMurtry et al., 1970; McMurtry, 1977b, 1985; Garcia Marì et al., 1983; Vacante, 1995 McMurtry et al., 1970 McMurtry et al., 1970 Van Rooyen, 1966 Van Rooyen, 1966 Van Rooyen, 1966 Meyer Smith and Schwartz, 1998 Meyer Smith and Schwartz, 1998 Garcia Marì et al., 1983; Vacante, 1995 Nakao et al., 1977 McMurtry et al., 1970; McMurtry, 1977b, 1985; Garcia Marì et al., 1983 Gonzalez, 1969 Garcia Marì et al., 1983

Garcia Marì et al., 1983 Tanaka and Kashio, 1977; Beattie, 1978 Tanaka and Kashio, 1977; Beattie, 1978 McMurtry, 1969 Meyer Smith, 1981 Meyer Smith, 1981 (Continued)

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Table 13.6. (continued) Natural enemies Euseius stipulatus (Athias Henriot)

Stigmaeidae Zetzellia sp. Agistemus cyprio (González) Agistemus exsertus Gonzalez Rodriguez Agistemus fanari Dosse Agistemus terminalis (Quayle)

Reference(s) Vacante, 1986, 1995; Vacante and Nucifora, 1986; Vacante et al., 1989; Ferragut et al., 1988, 1992; Abad et al., 2006 Garcia Marì et al., 1983 Garcia Marì et al., 1983 Yue and Childers, 1994 Dosse, 1967 Inoue and Tanaka, 1983

In the USA many species of insects and mites are recorded (Quayle, 1938; DeBach et al., 1950; Fleschner and Ricker, 1953a, b; Ebeling, 1959; McMurtry, 1969, 1977b, 1985; McMurtry et al., 1970, 1979; Yue and Childers, 1994b; Goldarazena et al., 2004; Villanueva et al., 2004, 2005). The insects include coccinellids, staphylinids, endomychids, chrysopids, coniopterygids and reduviids (Table 13.6). The most important predatory mites belong to the Phytoseiidae family (Table 13.6), and it is very difﬁcult to present a summary of this aspect. According to McMurtry (1977b), 21 species of the genus Amblyseius are reported on citrus worldwide, and the species of the ﬁnlandicus group are frequently associated with citrus and are general feeders on pollen. The species commonest are presented in Table 13.6 and according, to McMurtry (1992), they are generalists. In the humid areas Amblyseius species of the largoensis group are common. In addition to phytoseiids, some stigmaeid and bdellid mites are recorded. More detailed information can be found in Gerson et al. (2003). Symptomatology and damage The feeding symptomatology of the red spider mite may be confused with that of the oriental citrus mite, E. orientalis. The feeding from the upper surface of the leaves results in the removal of all cytoplasmic contents except for some starch grains (Albrigo et al., 1983). This produces on the leaves, mostly on the upper surface, little, empty spaces appearing as stippling, more or less numerous according to density of the attack, consisting of light-coloured spots with a grey or silvery appearance. The green, immature fruit becomes pale and mature fruit turns a pale straw yellow. The fruit damaged while still green will ripen and colour, but if the mite infestation occurs on mature fruit, the markings are more permanent. Severe levels of attack before fruit maturity may cause fruit fall (Stofberg, 1959; Keetch, 1968; Jeppson et al., 1975).

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The inﬂuence on the quality of the fruit is commonly of minor importance (Jeppson et al., 1975). The damage is severe when a serious density of attack is associated with a high transpiration rate owing to a low RH and wind, or a deﬁciency in leaf moisture owing to drought, poor root system or other inﬂuences that limit the capability of supplying adequate moisture to the leaves. The combined action of the mite attack and weather may produce heavy leaf and fruit drop, twig dieback and even death of large limbs (Jeppson et al., 1975). The twigs without leaves are bare and show that the leaf petioles are still attached and commonly die back as a result of the defoliation. The defoliation rate during winter may reach levels of 13.5, 20.6, 53.1 and 72.6% at densities of 1, 3, 6 and >10 mites/leaf, respectively (Choi and Kim, 1998). Severe infestation conditions negatively affect the yield in subsequent years. The use of one or more pesticides on Tahiti lime trees can negatively impact on the populations of citrus red mite and their associated predators, to the detriment of yield. The magnitude of impact is dependent upon the treatment regime and application date. Spider mite numbers increase because of reduced predator populations and use of these pesticides, especially in combination, is to be discouraged (Childers and Abou-Setta, 1999). Like other spider mites, P. citri is responsible for worker allergies (Burches et al., 1996). Control The methodology of sampling of the mite populations in citrus has been examined by various researchers. In the Mediterranean region the fortnightly counting of mobile forms present on four developed leaves of the newest vegetation has been suggested, sampled at random from 10% of the plants, keeping the threshold within the limits of 3 mites/leaf of new vegetation, or 20–40% of leaves infested (Cavalloro and Prota, 1983). Jones and Parrela (1984b) and Zalom et al. (1986) have examined the method of presence– absence and Rodriguez and Ramos (1998) have calculated on Marsh grapefruit seedlings that the proportion of infested leaves shows a strong relationship to the average density of mites for any phase. The values of R2 are 0.88, 0.85 and 0.85, respectively, for females, juveniles and the total population. Using the threshold value of 0.30 females/leaf and the results obtained, it is possible to apply a presence–absence sampling plan. The method provides a rapid estimation of the citrus red mite population based on 16% of infested leaves in a 30-leaf sample. Hare and Phillips (1992) did not ﬁnd a signiﬁcant relationship between the variation in density and duration of citrus red mite populations peaking at >19 adult females/leaf and variation in total yield, size or grade of lemon fruit. They have observed no effects of high mite populations in the ﬁrst year upon crop yield, grade or size in the second year, and have adopted a conventional treatment threshold for P. citri on lemon of two adult female mites/leaf. The data do not support the premise that growers who observe that threshold are achieving an economic return. Song et al. (2003) have produced a sequential sampling plan for adults of the citrus red mite, developing

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a decision-making citrus red mite population level based on the different action threshold levels (2.0, 2.5 and 3.0 mites/leaf), with 0.25 precision. The maximum number of trees and required number of trees sampled on ﬁxed sample size plans on 2.0, 2.5 and 3.0 thresholds with 0.25 precision level are 19, 16 and 15 and their critical T values are 554, 609 and 659, respectively. Nevertheless, the monitoring of populations is difﬁcult because variables such as climate, mite location and chemical control can affect the economic threshold. McMurtry (1985) report that successful monitoring of mite populations is based more on experience than on the use of economic thresholds, and that more citrus red mites can be tolerated in California during spring and summer than during the autumn, when the dry winds may cause stress to the trees. In South Africa, an average of 2–3 adult females/leaf represents the economic threshold, but the use of chemicals must be related to climatic conditions. Integrated control is the obvious solution to the citrus red mite problem. Panonychus citri very seldom reaches pest proportions in unsprayed orchards or orchards under integrated control (Meyer Smith and Schwartz, 1998e). In this context the selective acaricides must be employed. BIOLOGICAL CONTROL.

As regards biological control, practical applications and other control strategies become necessary. Recently, Paz et al. (2009a), evaluating in the laboratory the role of the pathogenic fungi M. geulakonigii, M. argovae and A. ingoldii (Exobasidiomycetidae), observed that after 14 days M. geulakonigii and A. ingoldii assured 86.7% and 87.5% of mortality, respectively, whereas M. argovae in the same conditions produced only 72.7% of mortality. The results encourage us to believe that a radical turning point in the control of the mite (and also of other herbivorous mites) is possible, but at this moment we do not know the practical implications of usage of these pathogens.

CHEMICAL CONTROL. Among acaricides, acid benzhydroxamic derivates (Benfatto and Lanza, 1980; Lanza et al., 1980; Vacante et al., 1980; Vacante, 1995); amidines (Benfatto and Lanza, 1980; Lanza et al., 1980; Garcia Marì et al., 1983; Vacante, 1995; Meyer Smith and Schwartz, 1998); avermectins (McCoy et al., 1982; French and Villarreal, 1988; Benfatto, 1994; Peña et al., 1999); organochlorines (Lanza et al., 1980; Vacante et al., 1980; Garcia Marì et al., 1983, 1988; French et al., 1985); organosulfurs (Benfatto and Lanza, 1980; Lanza et al., 1980; Vacante et al., 1980; Garcia Marì et al., 1983, 1988); organotins (Benfatto and Lanza, 1980; Lanza et al., 1980; Garcia Marì et al., 1983; Vacante, 1995; Meyer Smith and Schwartz, 1998e); petroleum oils (Jeppson et al., 1975; Benfatto and Lanza, 1980; Lanza et al., 1980; Garcia Marì et al., 1983; Ohkubo, 1983; French and Villarreal, 1988; Benfatto, 1994; Herron et al., 1995; Vacante, 1995; Walker and Aitken, 1996; Meyer Smith and Schwartz, 1998e; Nohara et al., 2000); phenyl pyrazoles (Yamada et al., 1987; French and Villarreal, 1988); pyridazinones (Benfatto, 1994; Peña et al., 1999; Shi and Feng, 2006); pyrazoles (Benfatto, 1994); quinazolines (Longhurst et al., 1992), tetrazines (Santaballa et al., 1994); and tetronic acid derivates (Izquierdo et al., 2002) are recorded, alone or mixed.

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Chapter 13 INTEGRATED PEST MANAGEMENT.

IPM represents the best strategy of control. It is necessary to evaluate the variation of susceptibility to different acaricides (Gotoh et al., 2004), and commonly the use of petroleum oils once or twice per year is required to control the problem appropriately. Alternatively, it is possible to utilige organotins. In all cases, mite control is strictly related to general control of adversities and horticultural practices. In the Mediterranean region, the phytoseiid mite E. stipulatus provides excellent control of citrus red mite populations, but it is necessary to preserve its presence in the citrus groves using selective acaricides or petroleum oils (Vacante, 1986).

13.4.3.5.2 Panonychus elongatus Manson (Fig. 13. 35) Common name Unknown. Diagnostic characteristics FEMALE. The body is 410 μm long and bright red in colour. The anterior margin

of the stylophore is rounded. The peritremes end in a bulb-like swelling. The dorsal body setae are long, slender, tapering, ﬁnely pubescent and most are set on prominent tubercles. The podosomal venter bears four pairs of midventral setae. The coxae I and II each bear two setae; the coxae III and IV each with one seta. Two pairs of anal setae, two pairs of pseudanal setae and two pairs of genital setae. The species is very similar to P. citri, but the body setae of female of this latter tetranychid, except for the prodorsal setae v2, are all slightly smaller than the corresponding body setae of P. elongatus (Manson, 1963b). MALE. The body is 273 μm long and bright red in colour. The terminal sensillum

of the palptarsus is about twice as long as broad. The peritremes end in a simple bulb. Tarsus I bears three tactile setae and three sensory setae proximal to duplex setae. The tibia I possesses seven tactile setae and four sensory setae. Tarsus II has three tactile and one sensory seta proximal to duplex setae. Tibia II presents ﬁve tactile setae. Tarsus III has nine tactile and one sensory setae. Tibia III has ﬁve tactile setae. The tarsus and tibia IV are similar to III. The distal part of the aedeagus is elongate, slender, tapering, directed posteriorly, similar to that of P. citri, but distinctly longer (Manson, 1963b). Geographical distribution Australia (Manson, 1968); China (Ma and Yuan, 1980); Korea (Han, 1970); Myanmar (Burma) (Manson, 1963b); Papua New Guinea (Schicha and Gutierrez, 1985); Taiwan (Tseng, 1990); and Thailand (Manson, 1963b; Baker, 1975).

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B C

D

Fig. 13.35. Panonychus elongatus Manson. Male. (A) Distal end of peritreme; (B) palptarsus; (C) aedeagus; (D) tibia and tarsus I (from Manson, 1963b).

Bio-ecology Panonychus elongatus is recorded on 22 host plants (Migeon and Dorkeld, 2006), including C. limon (Davies, 1968), C. reticulata (Davies, 1968; Ma and Yuan, 1980), C. sinensis (Davies, 1968), Citrus sp. (Manson, 1963b, 1968; Davies, 1968) and P. trifoliata (Davies, 1968). Its bio-ecology is unknown. The natural enemies include several phytoseiid mites, recorded for Papua New Guinea (A. largoensis, A. tamatavensis, N. longispinosus, Phytoseius hawaiiensis Prasad, Ph. hongkongensis, Ph. rubiginosae Schicha, Proprioseiopsis peltatus (van der Merwe) and Typhlodromips deleoni) (Schicha and Gutierrez, 1985), and for Taiwan (E. ovalis) (Shih et al., 1993).

13.4.3.5.3 Panonychus ulmi (Koch) (Fig. 13.36) Common name European red mite.

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Chapter 13 A

f1

B

C

h1

f2

Fig. 13.36. Panonychus ulmi (Koch). Female. (A) Dorsal opisthosomal setae h1, f2 and f1 (from Vacante, 1985); (B) receptaculum seminis; (C) pretarsus I (from Meyer Smith, 1987).

Diagnostic characteristics FEMALE. The body is reddish in colour, and the dorsal body setae are prominent and set on prominent tubercles, white at the bases. The dorsal opisthosomal setae h1 and f2 are one third and two-thirds as long as the dorsal opisthosomal setae f1, respectively. The genital plate possesses transverse striae and the anterior region longitudinal striae. The peritremes end in a simple bulb. The species is similar to P. citri (Vacante, 1985). MALE. Smaller than the female, light in colour, sometimes orange. The aedeagus is sigmoid, with broad base and distal end pointed and turned upward (Vacante, 1985).

Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The European red mite is recorded on 144 host plants (Migeon and Dorkeld, 2006), including C. aurantifolia, C. aurantium, C. grandis and Citrus sp. (Zhou et al., 1999). It is a serious pest of deciduous fruit and is very common in orchards. Its presence on citrus is sporadic and not injurious, arising from infested deciduous fruit trees, vineyards or other host plants from which the motile forms can be transported by wind, man or other means. Misidentiﬁcation in ﬁeld with P. citri or P. eleongatus is also possible. Reproduction is characterized by

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parthenogenesis arrhenotokous and the haploid condition is n = 3 (Helle and Bolland, 1967). The sex ratio is 63% females and 37% males (Jeppson et al., 1975). The mite overwinters as eggs, laid in groups, starting in August and extending to October or November, depending on the climate, on bark areas at the base of the buds, spurs, in bud wounds, and points that mark the beginning of new growth. Hatching takes place when a total of 195 degreedays have accumulated. The ﬁrst hatch occurs when the earliest blossom buds show pink (Jeppson et al., 1975); the immature stages live fundamentally on the undersurfaces of the leaves, whereas the adult can live on both surfaces. The summer eggs are laid on the lower leaf surface from spring to September or October. The life cycle requires 3–4 weeks, and ﬁve to six generations per year occur in Canada and as many as nine to ten in Virginia, USA (Parent and Beaulieu, 1957). At 23.5°C, the egg develops in 5 days and at 13°C in 20 days. The development from egg to adult requires 4 days at the higher temperature and 19–22 days at the lower temperature. The average life span of the female takes about 19 days and oviposition produces 10–90 eggs. At 25 ± 1°C the intrinsic rate of increase (rm) is 0.180/day and the ﬁnite rate of increase (λ) is 1.197 (Rabbinge, 1976). The species is fundamentally spread by transporting the overwintering eggs to new plantings on nursery stock. The dynamics of population is inﬂuenced by weather, cultural practices, predators, chemical treatments and host plant (Jeppson et al., 1975). Pathogens, predatory mites and insects develop at different biological stages of the European red mite. For further information, one may consult various works (Jeppson et al., 1975; Chazeau, 1985; Gerson, 1985b; Helle and Sabelis, 1985; Santos and Laing, 1985; van de Vrie, 1985; van der Geest, 1985; Gerson et al., 2003).

13.4.3.6 Schizotetranychus Trägårdh The opisthosoma possesses ten pairs of dorsal setae. The empodium is clawlike and split into two claw-like structures (Bolland et al., 1998).

13.4.3.6.1 Schizotetranychus baltazari Rimando (Fig. 13.37) Common name Citrus green mite. Diagnostic characteristics FEMALE. The body is 294 μm long, 238 μm wide, somewhat ﬂattened, yellow– green with dark spots along each side, with integumentary striae very smooth, and short legs. The peritremes end in simple bulbs. The palptarsus has the distal eupathidium twice as long as wide at base. The prodorsal striae

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A

B

C

D

Fig. 13.37. Schizotetranychus baltazari Rimando. Female. (A) Dorsal view; (B) tibia and tarsus I. Male. (C) Palptarsus; (D) aedeagus (from Manson,1963b).

are longitudinal and continue posteriorly to dorsal opisthosomal setae c1. The dorsal body setae are broad proximally and narrow distally, and about half as long as the distances between their bases. The propodosomal setae are equal in length, the length of the ﬁrst and second pairs nearly equal to the distances between their bases. The dorsal opisthosomal setae c3 are longest of all dorsal setae. The dorsal opisthosomal setae c1, d1, e1, c2, d2, e2 and f1 are similar, shorter than propodosomal setae. The setae f1 are well spaced. The setae f2 are longer than f1 and slightly shorter than h1 setae. The tibia I bears six tactile setae, and tarsus I two tactile setae approximate to duplex setae (Rimando, 1962a). MALE.

The body is 252 μm long. The distal eupathidium of the palptarsus is absent or rudimentary. The peritremes end distally with a simple bulb. Tibia

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I bears eight tactile setae, and tarsus I three proximal tactile setae; tibia II with ﬁve tactile setae, and tarsus II with one proximal tactile seta. The aedeagus is bent dorsally at right angles to the main shaft, and with a short distal sigmoid curve (Rimando, 1962a). Geographical distribution China (Wang, 1981a); Hong Kong (Manson, 1963b); India (Manson, 1963b; Gupta and Gupta, 1994); Indonesia (Ehara, 2004); Myanmar (Burma) (Manson, 1963b); the Philippines (Rimando, 1962a); Taiwan (Lo and Hsia, 1968); and Thailand (Ehara and Wongsiri, 1975). Bio-ecology Schizotetranychus baltazari is recorded for Dioscorea sp., H. koenigii, C. grandis, C. madurensis Loureiro, C. medica, C. sinensis (Gupta and Gupta, 1994) and Citrus sp. (Prasad, 1974). It feeds on leaves and fruit of citrus, and all biological stages are present throughout the year, but the quiescent period continues in the cooler season. The female lays close to the principal vein on both surfaces of the leaves and spins webbing. The development of egg, larva, protonymph and deutonymph occurs in 3.4, 6.6, 5.4 and 5.2 days, respectively, and the duration of preoviposition is 3.2 days. The life cycle takes about 23.8 days. The time of oviposition is 20 days and a female lays an average of 31.6 eggs (Lo and Hsia, 1968). The mite infests leaves and fruit of citrus, producing large, discoloured or grey spots on both sides of the leaves and on the fruit (Jeppson et al., 1975).

13.4.3.6.2 Schizotetranychus hindustanicus (Hirst) (Fig. 13.38) Common name Unknown. Diagnostic characteristics FEMALE.

Unknown (Jeppson et al., 1975).

MALE.

The dorsal body setae are about two-thirds as long as the distances between their bases; the dorsal opisthosomal setae f1 are much further apart than the other dorsal opisthosomal setae. The aedeagus has the distal part turned dorsally to form a sigmoid curve, slightly hooked at the tip and with the axis of the knob almost parallel with the shaft (Jeppson et al., 1975). Geographical distribution India (Hirst, 1924; Gupta and Gupta, 1994).

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Fig. 13.38. Schizotetranychus hindustanicus (Hirst). (A) Aedeagus (from Jeppson et al., 1975).

Bio-ecology The species is recorded for citrus and some other host plants in India (Hirst, 1924; Gupta and Gupta, 1994). Its bio-ecology is unknown.

13.4.3.6.3 Schizotetranychus lechrius Rimando Common name Unknown. Diagnostic characteristics FEMALE.

The body is 350 μm long and 252 μm wide. The peritremes end distally in a simple bulb. The palptarsus possesses a distal eupathidium, slightly more than twice as long as wide at base. Tibia I bears nine tactile setae and one sensory seta, and tarsus I ﬁve tactile setae and one sensory seta proximal to duplex setae. Tibia II possesses ﬁve tactile setae, and tarsus II four tactile setae and one sensory seta proximal to duplex setae. The empodia are claw-like, with a pair of dorsal appendant hairs on each digit (Rimando, 1962a).

MALE.

The body is 256 μm long. The peritremes are similar to those of the female. The palptarsus is devoid of distal eupathidium. The dorsal setae are slender, slightly broad at near base, pubescent, slightly longer than the distance between their bases. Tibia I bears nine tactile setae and four sensory setae, and tarsus I ﬁve tactile setae and three sensory setae proximal to duplex setae. The tibia II has six tactile setae, and tarsus II three tactile setae and one sensory seta proximal to the duplex setae. The empodium I possesses a pair of claw-like appendices, and empodia II–IV each with a pair of dorsal hairs on each division. The aedeagus is bent dorsally, with a knob characterized by a deﬁnite proximal angle and a slender, tapering posterior angle oblique dorsally (Rimando, 1962a). Geographical distribution Indonesia (Bolland et al., 1998); the Philippines (Rimando, 1962a); and Taiwan (Lo, 1969).

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Bio-ecology The mite has been collected on Colocasia esculentum (Linnaeus) Schott (Rimando, 1962a) and other host plants, and also on Citrus spp. (Bolland et al., 1998). Its bio-ecology is unknown.

13.4.3.6.4 Schizotetranychus spiculus Baker et Pritchard (Fig. 13.39) Common name Unknown. Diagnostic characteristics FEMALE. The body is 382 μm long and 240 μm wide. The peritremes end hooked. The palptarsus bears a distal eupathidium one and one half times as long as broad and slightly edentate at apex. The stylophore is edentate anteriorly. The prodorsal setae v2 and sc1 and the dorsal opisthosomal setae are awl-shaped and nearly nude, with c1, d1 and e1 slightly less than one half as long as distances between them longitudinally. The prodorsal setae sc2 and the opisthosomal setae c3, f2 and h1 are longer than the oblate setae, slender and serrate. The opisthosomal setae f1 are set apart than the setae c1, d1 and e1. The prodorsum is ﬁnely striate with longitudinal striae with small, elongate lobes. The opisthosoma is transversely striate between dorsocentral setae c1 and d1, forming between the dorsocentral setae d1 and e1 a V-pattern. Tarsus I bears two tactile setae and one solenidion proximal to duplex setae. Tibia I possesses seven tactile setae and one solenidion, and tibia II ﬁve tactile setae (Baker and Pritchard, 1960). MALE.

Unknown (Baker and Pritchard, 1960).

Geographical distribution India (Karuppuchamy and Mohanasundaram, 1987); Kenya (Baker and Pritchard, 1960); and Zaire (Baker and Pritchard, 1960). Bio-ecology The mite has been collected on Citrus sp. (Baker and Pritchard, 1960) and on Murraya koenigii (Karuppuchamy and Mohanasundaram, 1987). Its bioecology has not been investigated.

13.4.3.6.5 Schizotetranychus youngi Tseng Common name Unknown.

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A C

B

D

Fig. 13.39. Schizotetranychus spiculus Baker et Pritchard. Female. (A) Dorsal view; (B) distal segment of palp; (C) dorsal seta; (D) tibia and tarsus I (from Baker and Pritchard, 1960).

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Diagnostic characteristics FEMALE. The peritremes end simply and are curved distally. The distal eupathidium of the palptarsus is longer than wide at the base. The dorsal body setae are slightly serrate, with dorsal opisthosomal c1, d1 and e1 fusiform, broad at mid-length and narrow and tapering distally; the opishosomal setae h1 not tapering and the setae h2–3 are slender and tapering. Tibia I bears seven tactile setae and one little sensory seta, and tarsus I seven tactile setae and three solenidia, of which those forming the duplex setae set around the middle of the article. The tibia II has ﬁve tactile setae, and the tarsus II eight tactile setae, one single solenidion and a duplex seta set at the middle of article (Tseng, 1975). MALE.

Tibia I possesses seven tactile setae and three solenidia, and tarsus I nine tactile setae, one solenidion, set proximally, and two duplex setae set distally. Tibia II bears ﬁve tactile setae, and tarsus II eight tactile setae, one little solenidion set proximally and a pair of duplex setae in the distal part. The aedeagus is broad at the base, gradually narrowing and forming a large curve turned ventrally; the distal end is truncate, with the lower angle acute and the upper obtuse (Tseng, 1975). Geographical distribution Taiwan (Tseng, 1975). Bio-ecology The mite has been collected on C. medica (var. sarcodactilis) and C. paradisi (Tseng, 1975). Its bio-ecology is unknown.

13.4.3.7 Tetranychus Dufour The peritreme is recurved distally. The anal region possesses one pair of setae h. The empodium is split distally, commonly into three pairs of hairs, and the empodial spur is commonly visible. The duplex setae of tarsus I are well separated (Bolland et al., 1998).

13.4.3.7.1.Tetranychus desertorum Banks (Fig. 13.40) Common name Desert spider mite. Diagnostic characteristics FEMALE. The body is about 490 μm long, reddish in colour and with dorsal setae ﬁnely serrate and longer than the distances between their bases. The

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A

C

D E

Fig. 13.40. Tetranychus desertorum Banks. Female. (A) Tibia and tarsus I; (B); pretarsus; (C) palptarsus. Male. (D) Palptarsus (from Pritchard and Baker, 1955); (E) aedeagus (from Jeppson et al., 1975).

peritremes end in a simple hook. The dorsal opisthosomal striae form a broad triangle between the setae e1 and f1, and between these setae being longitudinal. The tactile setae on tarsus I are in line with the posterior set of duplex setae (Pritchard and Baker, 1955). MALE.

Smaller than the female. The aedeagus has the dorsal margin sigmoid, the anterior angulation small and acute, and posterior angulation also acute and curved ventrally to a variable extent; the width of the knob is not more than one quarter as long as the dorsal margin of the shaft (Pritchard and Baker, 1955).

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Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The desert spider mite is recorded on over 201 host plants (Migeon and Dorkeld, 2006), including Citrus sp. It is a pest of various cultivated plants and severely injures cotton and other tropical and subtropical crops (Jeppson et al., 1975), and its presence on citrus is occasional. Reproduction is characterized by parthenogenesis arrhenotokous and the haploid number of chromosomes is three (Helle et al., 1981). The mite belongs to the subtype CW-u of Satô (1985), constructing a highly complicated and irregular web on the leaf surface. The development of the egg occurs in summer in 2 days, and in 4–5 days in the winter. Each developmental stage requires 1.0–1.5 days in summer and 1.6–3.0 days in winter. The mite will not survive at temperature below 10°C, and the optimum conditions for increase in populations are 30°C and 85–90% RH. The intrinsic rate of increase (rm) at 30°C is 0.46 individuals/female/day at 25–30% RH and 0.36 individuals/female/day at 85–90% RH (Nickel, 1960). Temperature and humidity probably limit the distribution of the species in the warmer climates, and arid conditions appear to be preventing the species from becoming a major cotton pest in California and Arizona. During the winter the pest occurs on native host plants and by mid-March is abundant in localized patches; by the end of April it attacks cotton, where it develops and occurs abundantly in July. In autumn the mite moves to its winter hosts. The spring rains repress its development. Dispersion is through both wind and crawling. Control of the winter host favours mite control (Iglinsky and Rainwater, 1954; Hightower and Martin, 1956; Jeppson et al., 1975). In Japan on goldenrod (Solidago altissima Linnaeus,) the seasonal population trend of the desert spider mite is characterized by an exponential increase followed by a strong decline, with three peaks of density throughout the year. In this country among its natural enemies are recorded the phytoseiid mites Phytoseius nipponicus Ehara, Ph. capitatus Ehara, Amblyseius deleoni Muma et Denmark, the stigmaeid mite Agistemus exsertus (González Rodriguez) and the cecidomiid Feltiella sp. (Takafuji, 1980).

13.4.3.7.2 Tetranychus ﬁjiensis Hirst (Fig. 13.41) Common name Unknown. Diagnostic characteristics FEMALE. The body is 350 μm long and orange–red in colour. The peritremes are strongly U-shaped distally. The opisthosomal dorsal striae form a

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B

C D

Fig. 13.41. Tetranychus ﬁjiensis Hirst. Female. (A) Tibia and tarsus I. Male. (B) Palptarsus; (C) pretarsus I (from Pritchard and Baker, 1955); (D) aedeagus (from Jeppson et al., 1975).

diamond-shaped pattern between the setae e1 and f1. The empodia have only two pairs of proximal hairs, the dorsal pair being smaller than the ventral pair, and mediodorsal spur strong, about one half as long as the longer pair of proximal hairs. Tarsus I is attenuate, with tactile setae in line with the duplex setae, which are widely spaced (Pritchard and Baker, 1955). MALE. The aedeagus is long, slender, tapering and curved dorsally, without an

enlarged knob (Pritchard and Baker, 1955). Geographical distribution Australia (Flechtmann and Knihinicki, 2002); Carolina Islands (Gutierrez, 1980); China (Ehara and Tho, 1988); Fiji (Hirst, 1924); Hainan Island (Ehara and Tho, 1988); India (Hirst, 1924; Manson, 1963b; Gupta and Gupta, 1994); Kiribati (Gutierrez, 1977); Malaysia (Ehara and Tho, 1988); Marianas Northern (Gutierrez, 1980); Marshall Islands (Gutierrez, 1980); Micronesia Federated States (Gutierrez, 1980); New Caledonia (Bolland et al., 1981);

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Papua New Guinea (Schicha and Gutierrez, 1985); the Philippines (Baker, 1975); Seychelles (Gutierrez, 1974); Sri Lanka (Anomym, 1977); Taiwan (Tseng, 1990); and Thailand (Manson, 1963b; Baker, 1975). Bio-ecology Tetranychus ﬁjiensis occurs on 25 different host plants in Asia, Australia and Polynesia (Migeon and Dorkeld, 2006), including C. aurantium (Prasad, 1974; Gupta and Gupta, 1994), C. medica, C. microcarpa (Yusof and Zhang, 2003), C. paradisi, C. reticulata (Baker, 1975) and Citrus sp. (Ehara and Wongsiri, 1975). The reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is four (Bolland et al., 1981). The unmated females produce only males and the mated females both males and females, at a ratio of 1:8 (Vatana et al., 2001). The mite feeds on the lower surface of the leaves, where the different stages occur in webs spun by the adults (Daniel, 1979). The uninseminated females lay fewer eggs, but live longer than inseminated females. The 50% survival rate of inseminated females is attained 14.5 days before that of uninseminated females. The fecundity of inseminated females is roughly twice as great in comparison with uninseminated females. This behaviour is an adaptation to the climate of the area of origin (Bonato and Gutierrez, 1999). In India on leaves of betelnut (Areca catechu Linnaeus) the mite lives throughout the year, with maximum populations in summer and minimum in the rainy season. The egg stage lasts 4.0–4.5 days. The larval period lasts 43–68 h, including a quiescent period of 11–20 h. The protonymphal and deutonymphal periods are 47–52 and 47–72 h, respectively (including quiescent periods of 4–19 and 13–23 h). The immature stages last from 9.9 to 12.1 days (Daniel, 1978). On the passion fruit leaf, the developmental period from egg to adult of male and female is 10.7 and 12.6 days, respectively. Each unmated female could lay on average 15.1 eggs throughout her adult life, whereas the mated female could lay on average 20 eggs (Vatana et al., 2001). The complex of natural enemies is represented by various predators such as coccinellids, cecidomyiids and chrysopids, recorded on citrus (Daniel, 1979), on various crops (Chazeau, 1985) or experimentally investigated such as the coccinellid Delphastus catalinae (Horn) (Jing et al., 2003). Several phytoseiid mites has been studied in different regions of the world, such as Amblyseius channabasavannai Gupta et Daniel (Daniel, 1979); Amblyseius cinctus Corpuz et Rimando (Vatana et al., 2001); A. deleoni (Schicha and Gutierrez, 1985); A. largoensis (Schicha and Gutierrez, 1985); A. tamatavensis (Schicha and Gutierrez, 1985); N. longispinosus (Schicha and Gutierrez, 1985; Kongchuensin et al., 2005); E. nicholsi (Vatana et al., 2001); Euseius ovaloides (Blommers) (Schicha and Gutierrez, 1985); Ph. peltatus (Schicha and Gutierrez, 1985); Ph. hawaiiensis (Schicha and Gutierrez, 1985); Ph. hongkongensis (Schicha and Gutierrez, 1985); Ph. rubiginosae (Schicha and Gutierrez, 1985); and Papuaseius dominiquae (Schicha et Gutierrez) (Schicha and Gutierrez, 1985).

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Symptomatology and damage The mite feeds on the lower surface of the leaves, and on betelnut (A. catechu) it caused yellow speckles that later coalesced to give the whole leaf a yellow colour (Daniel, 1978). The symptomatology of the attack on citrus is unknown and on this crop control is unjustiﬁed.

13.4.3.7.3 Tetranychus gloveri Banks (Fig. 13.42) Common name Unknown. Diagnostic characteristics FEMALE. The body is carmine in colour, and with dorsal setae ﬁnely serrate, long and slender. The opisthosomal dorsal striae form a diamond-shaped

A

B

D

C

E

Fig. 13.42. Various kinds of aedeagus. Tetranychus gloveri Banks (from Jeppson et al., 1975); (B) Tetranychus kanzawai Kishida (from Meyer Smith, 1987); (C) Tetranychus paraguayensis Aranda (from Aranda and Flechtmann, 1971); (D) Tetranychus turkestani (Ugarov et Nikolski) (from Jeppson et al., 1975); (E) Tetranychus urticae Koch (from Vacante, 1985).

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pattern between the dorsal setae e1 and f1. The empodia are rayed and with a strong dorsal spur. Tarsus I with tactile setae proximal to the posterior duplex setae. The fertilized eggs are red (Jeppson et al., 1975). MALE.

Smaller than the female and lighter. Empodia I and II are claw-like and each with a strong dorsal spur; other empodia end in ventrally directed hairs, each having a dorsal spur. The knob of the aedeagus is indented dorsally (Jeppson et al., 1975). Geographical distribution

American Samoa (Gutierrez, 1978); Australia (Womersley, 1942; Halliday, 2000); Bermuda (Nyiira, 1982); Brazil (Bondar, 1930); Colombia (Urueta, 1975); Costa Rica (Ochoa et al., 1991); Cuba (Livshits and Salinas Crochea, 1968); French Polynesia (Navajas et al., 1996); French West Indies (Gutierrez and Hetienne, 1988); Greece (Papaioannou-Souliotis et al., 1994); Guadeloupe (Gutierrez and Etienne, 1988; Flechtmann and Etienne, 2006); Guam Island (McGregor, 1950); Hawaii (Garrett and Haramoto, 1967); Honduras (Ochoa et al., 1991); Les Saintes (Flechtmann and Etienne, 2006); Marianas Northern (Gutierrez, 1980); Mexico (Estebanes Gonzalez and Baker, 1968); Panama (Baker and Pritchard, 1962); Paraguay, Peru, Puerto Rico (Banks, 1917); Samoa (American) (Gutierrez, 1978); Suriname (Bolland et al., 1998); Trinidad and Tobago (Byrne et al., 1983); the USA (Banks, 1900); and Venezuela (Doreste, 1967). Bio-ecology The mite is recorded on 113 host plants (Migeon and Dorkeld, 2006), among these numerous herbaceous cultivated plants and Citrus sp. Its bio-ecology is unknown.

13.4.3.7.4 Tetranychus kanzawai Kishida (Fig. 13.42) Common name Kanzawa spider mite, cassava red spider mite. Diagnostic characteristics FEMALE. The body is carmine in colour, with dorsal body setae long and slender. The opisthosomal dorsal striae form a diamond-shaped pattern between the dorsal setae e1 and f1. The lobes of the dorsal striae are taller than broad. Tibiae I and II with nine and seven tactile setae, respectively. Tarsus I with four tactile setae proximal to the posterior duplex setae (Pritchard and Baker, 1955; Jeppson et al., 1975). MALE.

Smaller than the female, and lighter. Tibiae I and II and tarsus I are similar to those of the female. The aedeagus has the anterior part of the knob

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rounded and the posterior acutely angled (Pritchard and Baker, 1955; Jeppson et al., 1975). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The Kanzawa spider mite is recorded on about 160 host plants (Migeon and Dorkeld, 2006), including C. grandis, C. limon (Estebanes Gonzalez and Baker, 1968), Citrus sp. (Ehara, 1960), M. paniculata, Orixa japonica Thunb. and Zanthoxylum piperitum DC (Osakabe et al., 2002). It is a pest of tea and other crops (Banerjee and Cranham, 1985). The reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is three (Helle and Bolland, 1967). The proportion of females ranges from 0.76 to 0.83, and the sex ratio is determined by the genotype of the mothers, appearing to be controlled by several genes, without the involvement of cytoplasm (Takafuji and Ishii, 1989). The males probably use odours to determine the mating status of females. These abilities of males may play an important role in gaining access to virgin females. Alternatively, the behaviour of adult females varies with their mating status. Virgin females are more gregarious and remain on infested kidney leaves for a longer time than mated females. This behaviour is likely to increase the mating opportunities of virgin females (Oku et al., 2005). The Kanzawa spider mite is infected by endosymbiont Wolbachia, a group of maternally inherited bacteria infecting arthropods, responsible in the spider mite for cytoplasmatic incompatibility (Breeuwer, 1997), but not found in the Japanese populations of T. kanzawai (Gotoh et al., 2004). The tetranychid belongs to the subtype CW-u, constructing a complex and irregular web on the leaf surface (Satô, 1985). The mite lives on the lower surface of the leaves. At 15°, 20°, 25° and 30°C the egg development takes 15.1, 8.9, 4.4 and 2.3 days, the larval period 6.4, 3.5, 1.9 and 1.2 days, respectively; protonymphal period 5.6, 2.8, 1.7 and 1.0 days and deutonymphal period 7.5, 4.0, 2.1 and 1.3 days, respectively. The corresponding adult female life spans take 33.3, 16.8, 15.5 and 13.4 days, and the average numbers of eggs laid per female are 37.5, 59.4, 100.6 and 103.3, respectively. The developmental threshold temperatures for the egg, protonymphal and deutonymphal stages are 13.9°, 12.6° and 12.6°C, respectively, and the corresponding temperature sums for development 39.2, 21.4 and 18.2 degree-days (Kim et al., 1993). On tea at 15°, 20°, 25° and 30°C, the intrinsic rates of increase (rm) are 0.0443, 0.1079, 0.2149 and 0.2764, respectively. The net reproductive rate (R0) ranges from 10.85 female offspring/female at 15°C to 31.05 at 30°C, and the ﬁnite rate of increase (λ) from 1.0453 to 1.3183. The mean generation time (T) ranges from 12.4 days at 30° to 53.9 days at 15°C (Tsai et al., 1989). At 15°, 20°, 25° and 30°C, the intrinsic rates of increase (rm) are 0.062, 0.134, 0.252 and 0.371, respectively (Kim et al., 1993). According to Gotoh and Gomi (2003),

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the intrinsic rates of natural increase (rm) vary on different host plants, from 0.187/day to 0.283/day. The factors limiting the development of populations include predatory insects and mites. The former include the staphylinid Oligota kashmirica beneﬁce Naomi, the cecidomyiid Feltiella sp., the aeolothripids S. sexmaculatus (McMurtry et al., 1970) and S. takahashii Priesner (Morishita, 2000), and the anthocorids Anthocoris sp. and Orius sp. (McMurtry et al., 1970). Among the mites are recorded the phytoseiids Amblyseius eharai (Morishita, 2000), N. californicus (Takada and Kashio, 2004), N. fallacis (Ho, 1990), N. longispinosus (Nakagawa, 1984; Nagatomo et al., 1991; Kongchuensin et al., 2005), N. womersleyi (Kim and Paik, 1996a), Euseius sojaensis (Ehara) (Osakabe et al., 1987; Morishita, 2000) and Phytoseiulus persimilis (Zhang et al., 1996). Symptomatology and damage The mite is a pest on tea and other crops (Banerjee and Cranham, 1985) and a minor pest of citrus, where the symptomatology and damage are unclear. Control On citrus, the need for control is infrequent. Nevertheless, in other crops various acaricides are employed, such as avermectins (Kim and Paik, 1996b), diphenil carbinols (Wang and Liu, 1993), diphenyl ether (Wang and Liu, 1993), organophosphates (Ozawa, 1995), sulfur-bridged compounds (Wang and Liu, 1993; Tatara et al., 1999) and other active ingredients, such as the synthetic pyrethroids (Ozawa, 1995). Resistance to various acaricides such as pyrazole, fenpyroximate and pyridaben is known (Goka, 1998).

13.4.3.7.5 Tetranychus lambi Pritchard et Baker (Fig. 13.43) Common name Unknown. Diagnostic characteristics FEMALE. The body is carmine in colour. The distal sensillum of the palptarsus is about twice as long as wide. The peritremes end hooked. The dorsal body setae are long and slender. The opisthosomal dorsal striae form a diamond-shaped pattern between the dorsal setae e1 and f1. Ventrally, the area anterior to the genital ﬂap, as well as the anterior part of the ﬂap, with longitudinal striae. Tarsus I with proximal duplex setae distal to four tactile setae. The mediodorsal spur is absent on each empodium (Pritchard and Baker, 1955). MALE.

Smaller than the female. The sensillum of the palptarsus is about four times as long as broad. Empodium I consisting of two pairs of nearly united, triﬁd plates. Tarsi I and II are devoid of mediodorsal spurs. The aedeagus has

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B A

E

C D

F

Fig. 13.43. Tetranychus lambi Pritchard et Baker. (A) Tibia and tarsus I of female; (B) palptarsus of male; (C) palptarsus of female; (D) distal end of peritreme; (E) pretarsus I of male (from Pritchard and Baker, 1955); (F) aedeagus (from Jeppson et al., 1975).

the axis of its knob parallel to axis of shaft, slender, scarcely widened dorsoventrally, with the anterior angulation acute and pronounced, and caudal angulation very slender and acute (Pritchard and Baker, 1955). Geographical distribution Australia (Davis, 1968); Cook Islands (Gutierrez, 1980); Fiji (Swaine, 1971); French Polynesia (Gutierrez, 1980); Iran (Khalil Manesh, 1973); New Caledonia

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(Gutierrez, 1977); New Zealand (Pritchard and Baker, 1955); Papua New Guinea (Schicha and Gutierrez, 1985); Samoa (American) (Gutierrez, 1977); Samoa (Western) (Bolland et al., 1998); Taiwan (Baker, 1975); Tasmania (Miller, 1966); Tonga (Gutierrez, 1980); Vanuatu (Gutierrez, 1977); Wallis and Futuna (Gutierrez, 1981); and Western Samoa (Bolland et al., 1998). Bio-ecology The mite is recorded on about 61 host plants and on different cultivated plants (Migeon and Dorkeld, 2006). As regards citrus, it has been collected on P. trifoliata (Davis, 1968) and on Citrus sp. (Gerson, 2003). The reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is three (Bolland et al., 1981). Natural enemies include, in various crops, coccinellids, Staphylinidae, Cecidomyiidae and Aelothripidae (Chazeau, 1985). Symptomatology and damage On strawberry, the injury appears as an abnormal and irregular purple colour on the upper leaf surface, commonly close to the principal veins. Severe densities of attack produce a silvering appearance on the entire undersurface, the leaf edges roll, growth regresses and the fruit ripens prematurely (Davis and Heather, 1962). On citrus the damage is unknown, and control is unjustiﬁed. 13.4.3.7.6 Tetranychus ludeni Zacher (Fig. 13.44) Common name Red spider mite, red-legged spider mite. Diagnostic characteristics FEMALE. The body is carmine in colour and the legs reddish. The dorsal body setae are serrate, long and slender. The opisthosomal dorsal striae between the dorsal setae e1 and f1 are longitudinal and forming a diamond-shaped pattern. Tarsus I has the proximal pairs of duplex setae on the same line with four tactile setae; the empodia have a small mediodorsal spur (Pritchard and Baker, 1955; Jeppson et al., 1975). MALE. Smaller than the female. The mediodorsal spur of empodium I is about one third the length of two proximoventral spurs. The empodia II–IV have three pairs of proximoventral hairs and a small mediodorsal spur. The aedeagus has the distal knob very small, without posterior projection, and anterior projection small and acuminate (Pritchard and Baker, 1955; Jeppson et al., 1975).

Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006).

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B

C

E

D

F

G

Fig. 13.44. Tetranychus ludeni Zacher. (A) Dorsal view of female; (B) distal end of peritreme of female; (C) palptarsus of female; (D) palptarsus of male (from Ehara and Masaki, 1989); (E) tibia and tarsus I of female; (F) pretarsus I of female; (G) aedeagus (from Pritchard and Baker, 1955).

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Bio-ecology The red spider mite is recorded on about 312 host plants, spontaneous and cultivated (Migeon and Dorkeld, 2006). In the warmer regions of the world, the species occurs throughout the year and attacks many agricultural crops, such as cotton, fruit trees, vegetables and ornamentals, in both ﬁeld and greenhouse (Jeppson et al., 1975), and has been collected on C. aurantium (Davis, 1968), C. limon (Meyer Smith, 1974), C. paradisi, Citrus sp. (Gutierrez and Schicha, 1983; Gupta and Gupta, 1994; Gupta and Chatterjee, 1997) and P. trifoliata (Davis, 1968). The reproduction is parthenogenetic arrhenotokous and the haploid number of chromosomes is three (Helle et al., 1970). The male to female ratio is 1:2.7 (Puttaswamy and Channabasavanna, 1979a). The pest initially feeds on underleaf surfaces or in the young, curled leaf, and as population increases distributes over all leaf surfaces and produces webbing over the entire plant (Coates, 1974). Field studies carried out in India on French bean plants with temperatures ranging from 19.3° to 28.4°C and 53% to 88% RH have shown that the development of males and females takes 11.96 and 12.48 days, respectively. Mated and unmated females have pre-oviposition periods of 1.54 and 1.43 days, respectively, and oviposition periods of 22.83 and 27.41 days; they laid a total of 165.9 and 132.0 eggs, and lived for 27.98 and 32.14 days, respectively. The adult male life span takes 25.20 days (Puttaswamy and Channabasavanna, 1979a). On Phaseolus vulgaris Linnaeus at 26.34 ± 3.92°C and 69.44 ± 19.44% RH, the mean duration of the different developmental stages for egg, larva, protonymph and deutonymph is 4.68, 1.75, 1.31 and 1.85 days, respectively. The total life cycle is 9.98 and 9.25 days for females and males, respectively (Morros and Aponte, 1994). On cotton at 20°, 23°, 25°, 28° and 30°C, 70 ± 10% RH, and a 12 h:12 h (L:D) photoperiod, the development time from egg to adult ranges from 20.77 days at 20°C to 8.50 days at 30°C for males, and from 18.83 days at 20°C to 7.75 days at 30°C, for females (da Silva, 2002). On Ph. vulgaris the reproduction rate (R0) is 77.42, the mean generation time (T) is 19.63 days, the intrinsic rate of increase (rm) is 0.2526 individuals/ female/day and the ﬁnite rate of natural increase (λ) is 1.2874 individuals/ female/day (Morros and Aponte, 1994). On cotton, the values of thermal constant are 138.34 degree-days for females and 130.91 degree-days for males. At 30°C the highest values of the intrinsic rate of increase (rm) (0.418), number of eggs/females/day (3.47), fecundity (61.29), net reproductive rate (48.00), and minimum value to the mean generation time (9.27) are recorded (da Silva, 2002). All stages are found throughout the year, and many overlapping generations per year are possible (Jeppson et al., 1975). Wind was found the most important factor affecting the distribution of T. ludeni (Puttaswamy and Channabasavanna, 1979b). Natural enemies include pathogens and several predatory species. Among the ﬁrst reported is the fungus dematiaceaed Cladosporium sp. (Godse and Patil, 1977). The predatory insects number aeolothripids, coccinellids,

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staphylinids and chrysopids. Several phytoseiids, and some stigmaeid and cunaxid mites have been investigated in various regions of the world on various crops (Table 13.7). Symptomatology and damage Attacked plants become yellowish, wilt and drop rapidly, normally during dry conditions. The leaves ﬁrst turn yellow, but soon necrotic patches encompass the leaves (Jeppson et al., 1975). Mite feeding on leaves of French bean causes white stippling, which resulted in drying and leaf loss (Puttaswamy and Channabasavanna, 1979a). Damage from the red spider mite to the citrus is fundamentally unknown and judged of minor importance. Control On various crops different acaricides are employed, such as avermectins (Kumar et al., 2003); formamidines (Chiavegato et al., 1975); organochlorines (Duncombe, 1972; Chiavegato et al., 1975; Nangia and Channabasavanna, 1983; Jagadish and Channabasavanna, 1983; Nangia et al., 1990; Kumar and Sharma, 1993; Kumar et al., 2003); organosulphurs (Nangia et al., 1990; Kumar and Sharma, 1993); organotins (Brun et al., 1983; Peter and David, 1988); or sulfur (Nangia and Channabasavanna, 1983; Nangia et al., 1990; Kumar and Sharma, 1993; Kumar et al., 2003), but on citrus the need does not arise. 13.4.3.7.7 Tetranychus mexicanus (McGregor) (Fig. 13.45) Common name Unknown. Diagnostic characteristics FEMALE.

The body is carmine in colour, with dorsal setae long, ﬁnely serrate and slender. The opisthosoma possesses a diamond-shaped pattern between the dorsal setae e1 and f1, and the striae are longitudinal between each of these pairs of setae. The empodium has a large mediodorsal spur, and the proximal pair of duplex setae on tarsus I are placed distal to the proximal tactile setae (Pritchard and Baker, 1955).

MALE.

Smaller than the female. The empodium possesses a large mediodorsal spur and the proximal pair of duplex setae on tarsus I are placed distal to the proximal tactile setae. The aedeagus has the axis of the knob parallel to that of the shaft, the anterior angulation short and acutely angulate and posterior longer and also acutely angular (Pritchard and Baker, 1955). Geographical distribution Argentina (Pritchard and Baker, 1955); Brazil (Paschoal and Reis, 1968); China (Cheng, 1994); Colombia (Urueta, 1975); Costa Rica (Ochoa et al., 1991); Cuba

Tetranychidae Donnadieu Table 13.7.

277

Tetranychus ludeni Zacher: natural enemies (fungi, insects and mites).

Natural enemies

Reference(s)

Fungi Dematiaceae Cladosporium sp.

Godse and Patil, 1977

Insects Aeolothripidae Scolothrips indicus Priesner Chrysopidae Chrysoperla carnea Stephens Coccinellidae Stethorus pauperculus Weis Scymnus coccivora Ayyar Staphylinidae Oligota oviformis Casey Mites Cunaxidae Cunaxa setirostris Hermann Phytoseiidae Amblyseius multidentatus (Chant et al.) Euseius alstoniae (Gupta) Euseius delhiensis (Narayan et Kaur) Euseius ﬁnlandicus (Oudemans) Euseius neococcineae (Gupta) Euseius rhododendronis (Gupta) Neoseiulus californicus (McGregor) Neoseiulus fallacis (Garman) Neoseiulus idaeus Denmark et Muma Neoseiulus indicus (Narayan et Kaur) Neoseiulus longispinosus (Evans) Phytoseius jujuba Gupta Phytoseius minutus Narayanan et al. Phytoseius mixtus Chaudhri Phytoseius roseus Gupta Phytoseiulus persimilis Athias Henriot Typhlodromips tetranychivorus Gupta

Typhlodromus homalii (Gupta) Stigmaeidae Agistemus industani Gonzalez Rodriguez

Reddy and Jagadish, 1977 Reddy, 2002 Ansari and Pawar, 1992 Ansari and Pawar, 1992 Ansari and Pawar, 1992

Arbabi and Singh, 2000 Kumar and Sharma, 1993; Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Krishnamoorthy, 1983 Escudero et al., 2005 Biasi and Santos, 1988; Putatunda and Anupam Tagore, 1997 Escudero et al., 2005 Putatunda and Anupam Tagore, 1997 Ansari and Pawar, 1992; Abhilash and Sudharma, 2003 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Escudero et al., 2005 Puttaswamy and Channabasavanna, 1979c; Nangia and Channabasavanna, 1983; Ansari and Pawar, 1992; Kumar and Sharma, 1993 Putatunda and Anupam Tagore, 1997 Arbabi and Singh, 2002

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A

C

Fig. 13.45. Tetranychus mexicanus (McGregor). Female. (A) Tibia and tarsus I; (B) pretarsus I. Male. (C) Aedeagus (from Pritchard and Baker, 1955).

(Livshits and Salinas Croche, 1968); El Salvador (Andrews and Poes, 1980); Guadeloupe (Flechtmann et al., 1999; Flechtmann and Etienne, 2006); Honduras (Ochoa et al., 1991); Les Saintes (Flechtmann and Etienne, 2006); Mexico (Beer and Lang, 1958); Nicaragua (Ochoa et al., 1991); Paraguay (Aranda, 1969); Peru (Bolland et al., 1998); USA (McGregor, 1950); Uruguay (Bernal and Piñeiro, 1982); and Venezuela (Dominguez Gil and McPheron, 1992; Quiros de Gonzalez, 2000). Bio-ecology Tetranychus mexicanus has been collected on 105 spontaneous and cultivated plants (Migeon and Dorkeld, 2006), including C. aurantifolia, C. latifolia (Quiros de Gonzalez, 2000), C. limon (Pritchard and Baker, 1955), C. paradisi, C. reticulata, C. sinensis (McGregor, 1950), Citrus sp. (Pritchard and Baker, 1955; Aranda and Flechtman, 1971; Suarez, 2004), Fortunella japonica (Thunb.) Swingle (Fernadez, 1972), Zanthoxylum stipitatum Huang and P. trifoliata (Feres et al., 2005) and Esenbeckia leiocarpa Engl. (Paschoal and Reis, 1968). Its bio-ecology is unknown. The natural enemies include some predatory mites, such as the stigmaeid Agistemus ﬂoridanus Gonzalez and the phytoseiids Euseius concordis (Chant),

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Neoseiulus anonymus (Chant et Baker) (Ferla and de Moraes, 2003) and Phytoseiulus macropilis (Banks) (Jaime de Herrero, 1984). Symptomatology and damage The mite is sometimes injurious to citrus in Texas, Mexico and Argentina (Jeppson, 1989), but the attacks are of minor importance. Control The employment in various crops of several acaricides such as avermectins (Nakayama et al., 1987), dinitrophenol derivates (Nakayama et al., 1987), organosulphurs (Nakayama et al., 1987; Hernandez et al., 1988), organotins (Nakayama et al., 1987), pyrethroids (Nakayama et al., 1987) and sulfur (Hernandez et al., 1988) has been reported, but their use on citrus is not justiﬁed.

13.4.3.7.8 Tetranychus neocaledonicus André (Fig. 13.46) Common name Vegetable mite. Diagnostic characteristics FEMALE.

The body is bright red in colour. The peritremes are hooked distally. The terminal sensillum of the palptarsus is about one-and-three-quarters as long as broad. The dorsal body setae are ﬁnely serrate, slender and longer than distances between their bases. Dorsal opisthosomal striae form a diamond-shaped pattern between the dorsal setae e1 and f1, and the striae between these setae are longitudinal. The lobes on the dorsal striae have different sizes and shapes, mostly semicircular, with basal spots. Medioventral striae bearing lobes low and wide, sometimes hardly more than an occasional incision on the striae. The tibia I and tarsus I each with one solenidion. Tarsus I with four tactile setae proximal to the duplex setae (Meyer Smith, 1974, 1987).

MALE.

Smaller than the female. The terminal sensillum of the palptarsus more than twice as long as broad. Tarsus I with four tactile setae and two solenidia proximal to duplex setae. Empodium I with small dorsal spur, about one quarter the length of the proximoventral spur; empodium II split into three pairs of proximoventral hairs and devoid of obvious mediodorsal spur. The shaft of the aedeagus bends dorsally at nearly a right angle; anterior and posterior projections of knob rounded and separated by a small, dorsal indentation (Meyer Smith, 1974, 1987). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006).

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A

C B

F

D

E

Fig. 13.46. Tetranychus neocaledonicus André. Female. (A) Dorsal view (from Attiah, 1969); (B) palptarsus. Male. (C) Palptarsus; (D) pretarsus I; (E) pretarsus II; (F) aedeagus (from Meyer Smith, 1987).

Bio-ecology The vegetable mite is pantotropical, collected about on 440 spontaneous and cultivated plants in warmer regions of the world (Migeon and Dorkeld, 2006), including C. aurantifolia; C. aurantium (Attiah, 1969); C. grandis; C. limon; (Prasad, 1974; Nassar and Ghai, 1981; Gupta and Gupta, 1994); C. sinensis

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(Attiah, 1969); Citrus sp. (Gutierrez and Schicha, 1983; Gupta and Chatterjee, 1997); Evodia madagascariensis Baker; Feronia limonia (Linnaeus) Swingle; Galipea sp.; P. trifoliata; and Zanthoxylum tsihanimposa Perrier (Migeon and Dorkeld, 2006). In tropical and subtropical areas it is a serious pest of vegetable and ﬁeld crops (Goldsmid, 1962; Jeppson et al., 1975), and in Egypt has been found on leaves and fruit of mandarin and orange (Attiah, 1969; Soliman et al., 1973). The 50% survival rate of inseminated females is attained 15.5 days before that of uninseminated females, and the fecundity of inseminated females is 1.5 times as great in comparison with uninseminated females (Bonato and Gutierrez, 1999). The mite feeds on lower surfaces of the leaves and on fruit, and produces webbing. The eggs are laid in the web, commonly on the lower surfaces of the leaves. On cardamom, the female completes its life cycle in 10.47 days and the male in 10.01 days. The egg, larval, deutonymphal and protonymphal stages average 4.66, 1.71, 1.47 and 2.63 days, respectively, for the female, and 4.65, 1.60, 1.41 and 2.35 days, respectively, for the male. Mated females each lay an average of 126.42 eggs during their oviposition period of 20.50 days, whereas unmated females lay an average of 87.00 eggs over 23.50 days. The life span of mated females is 24.00 days, of unmated females 26.50 days and of males 12.25 days (Puttaswamy and Reddy, 1980). On Amaranthus viridis (Linnaeus) at 23–26°C and 74–81% RH the developmental period from egg to adult lasts 10.19 days. Females lay 147.42 eggs and the life span is 27.69 days (Puttaswamy and Channabasavanna, 1981). In general, there is a negative relationship between population size and rainfall level, and the responses of the mite to the different levels of rainfall vary according to the food plant (Sharma and Pande, 1983). The species does not survive at temperatures beyond 37°C, and between 30° and 37°C it fails to complete its life cycle. With increase in temperature above 20°C, development is faster but mortality of all stages in the life cycle increases. Overall, 30°C is the optimal temperature (Pande and Sharma, 1986). At 25 ± 1°C on cotton, the net reproduction (R0) is 57.2, the intrinsic rate of increase (rm) is 0.260 individuals/female/day and the ﬁnite rate of increase (λ) is 1.297 individuals/female/day (Gutierrez, 1976). The vegetable mite overwinters as fertilized females on weeds, and their activity begins when the weather warms in the spring. In this period, populations increase and infest the cultivated plants, reaching the highest densities in May to mid-July and decreasing during late July and August. After that, populations increase again in September and October, commonly on weeds. The females move to winter crops and continue their activity until December and February. Successively, the females move to winter crops or weed hosts to overwinter (Jeppson et al., 1975). The destruction of overwintering or summer hosts decreases infestation levels, leaving the land free of plants between crop plantings (Rahman and Sapra, 1945; Khot and Patil, 1956). The natural enemies number pathogens, predatory insects and mites. The ﬁrst are represented by several fungi. The insects include coccinellids, staphylinids and chrysopids, and the mites several phytoseiids and several Anystidae (Table 13.8).

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Table 13.8. Tetranychus neocaledonicus André: natural enemies (fungi, insects and mites). Natural enemies

Reference(s)

Fungi Ascomycota Verticillium lecanii (Zimmermann) Viégas Sreenivas et al., 2005 Clavicipiteae Beauveria bassiana (Balsamo) Vuillemin Sreenivas et al., 2005 Metarhizium anisopliae (Metschnikoff) Sorokin Hanchinal and Manjunatha, 2000; Sreenivas et al., 2005 Insects Chrysopidae Chrysoperla carnea Stephens Coccinellidae Stethorus madecassus Chazeau Stethorus pauperculus (Weise) Stethorus proximus Chazeau Rosolia sp. Chilocorus sp. Menochilus sexmaculatus (Fabricius) Brumus suturalis Fabricius Illeis cincta (Fabricius) Staphylinidae Oligota ﬂavicornis (Boisduval et Lacordaire) Oligota pallidicornis Cameron Mites Anystidae Anystis sp. Phytoseiidae Amblyseius deleoni Muma et Denmark Amblyseius largoensis (Muma) Amblyseius multidentatus (Chant et al.) Amblyseius tamatavensis Blommers Amblyseius vazimba Blommers et Chazeau Euseius alstoniae (Gupta) Euseius concordis (Chant) Euseius delhiensis (Narayan et Kaur) Euseius ﬁnlandicus (Oudemans) Euseius neococcineae(Gupta) Euseius ovaloides (Blommers) Galendromus pilosus (Chant) Neoseiulus bibens (Bloomers) Neoseiulus fallacis (Garman) Neoseiulus indicus (Narayan et Kaur) Neoseiulus longispinosus (Evans) Neoseiulus masiaka (Blommers et Chazeau) Papuaseius dominiquae (Schicha et Gutierrez)

Sharanabasava and Manjunatha, 2001 Gutierrez and Chazeau, 1972; Chazeau, 1974 Puttaswamy and Channabasavanna, 1976 Chazeau, 1979 Pande and Sharma, 1984 Pande and Sharma, 1984 Pande and Sharma, 1984 Pande and Sharma, 1984 Pande and Sharma, 1984 Gutierrez, 1976 Gutierrez, 1976

Singh and Singh, 1999 Schicha and Gutierrez, 1985 Schicha and Gutierrez, 1985 Putatunda and Anupam Tagore, 1997 Schicha and Gutierrez, 1985 Blommers and Chazeau, 1974 Putatunda and Anupam Tagore, 1997 Jagadish et al., 1994 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Schicha and Gutierrez, 1985 Aguilar and Salas, 1989 Blommers, 1976 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Schicha and Gutierrez, 1985 Blommers and Chazeau, 1974 Schicha and Gutierrez, 1985 (Continued)

Tetranychidae Donnadieu Table 13.8.

283

(continued)

Natural enemies Phytoseius hawaiiensis Prasad Phytoseius hongkongensis Swirski et Shechter Phytoseius jujuba Gupta Phytoseius minutus Narayanan et al. Phytoseius mixtus Chaudhri Phytoseius roseus Gupta Phytoseius rubiginosae Schicha Proprioseiopsis peltatus (van der Merwe) Typhlodromus homalii (Gupta)

Reference(s) Schicha and Gutierrez, 1985 Schicha and Gutierrez, 1985 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Putatunda and Anupam Tagore, 1997 Schicha and Gutierrez, 1985 Schicha and Gutierrez, 1985 Putatunda and Anupam Tagore, 1997

Symptomatology and damage The mite feeds on the leaves, producing white spots that gradually coalesce. The leaves lose their green colour, gradually wilt, dry and drop. The infestation adversely affects growth, ﬂowering and fruiting. The damaged areas of some plants may turn red. The pest produces a web, which forms a thick sheath that covers the canopy (Jeppson et al., 1975). In Egypt, T. neocaledonicus infests leaves and fruit of mandarin and orange, causing yellowish blotches in fruit (Attiah, 1969), but it is considered a minor pest for citrus crops. Control Generally, in various crops mite control is entrusted to several acaricides, such as diphenyl carbinols (Sidhu and Singh, 1971; Gupta et al., 1972; Krishnaiah and Tandon, 1975; Dhooria and Mann, 1980; Singh et al., 1981, 1988; Peter et al., 1987; Pillai and Jolly, 1986), organotins (Krishnaiah and Tandon, 1975; Singh et al., 1981; Peter et al., 1987; Yadav et al., 1987), organophosphates (Peter et al., 1987), sulfur-bridged (Krishnaiah and Tandon, 1975; Dhooria and Mann, 1980; Singh et al., 1981; Zhang et al., 1990); and sulfur as lime sulfur (Zhang et al., 1990) or wettable sulfur (Anand and Chauhan, 1983; Yadav et al., 1987). Nevertheless, the chemical control is not justiﬁed on citrus.

13.4.3.7.9 Tetranychus paciﬁcus McGregor (Fig. 13.47) Common name Paciﬁc spider mite. Diagnostic characteristics FEMALE. The body is yellow coloured, with three irregular dark spots on each side of the hysterosoma. The dorsal body setae are ﬁnely serrate, long and

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A

B

Fig. 13.47. Tetranychus paciﬁcus McGregor. Female. (A) Dorsal view (from Pritchard and Baker, 1955). Male. (B) Aedeagus (from Jeppson et al. 1975).

slender. The opisthosoma bears transverse striae between the dorsal setae e1 and f1. The peritremes end simply. The tactile setae of tarsus I are proximal to the duplex setae. The empodia are devoid of dorsal spurs (Pritchard and Baker, 1955; Jeppson et al., 1975). MALE.

Smaller than the female. Empodium I is claw-like; with a prominent dorsal spur, and other empodia are similar to those of the female. The aedeagus has a distal knob with a long posterior angulation, its tip reaching well beyond the level of the caudal end of the bend of the neck, and forming an obtuse angle (Pritchard and Baker, 1955; Jeppson et al., 1975). Geographical distribution Canada (Baker and Pritchard, 1953); Mexico (Beer and Laing, 1958); and the USA (McGregor, 1919). Bio-ecology The paciﬁc spider mite has been collected on 50 host plants (Migeon and Dorkeld, 2006). It is a serious pest of deciduous trees, cotton, grapes and other crops (Jeppson et al., 1975). It was found on orange and lemon trees in Los Angeles County, and on grapefruit in Riverside County (McGregor,

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1950, 1956), and has produced serious damage in San Joaquin Valley in California, but is rare on this crop and was probably dispersed from nearby food plants. The infestations occurred under clean crops indicating that the mite has become established as a pest of citrus (Jeppson, 1989). The species possesses a haploid number of three chromosomes and reproduces by parthenogenesis arrhenotokous (Helle and Bolland, 1967). It feeds on both leaf surfaces and produces webbing, which may cover the entire plant. Each female lays 50–100 eggs in 2–4 weeks, and the life cycle from egg to adult lasts 10–14 days (Jeppson et al., 1975). On cotton cotyledons, the average development periods ranges from 25.8 to 29.0 days at 15.5°C and from 6.1 to 6.7 days at 29.4°C for the females, and a slightly shorter time for the males. The total progeny at 23.8°C averaged 78.9 eggs/ female and the corresponding number at 29.4°C is 68.3 eggs/female. The percentage of female offspring is 57. The females lived for a mean of 12.71 days at 23.8°C, and 8.91 days at 29.4°C (Carey and Bradley, 1982). On cotton at 29°C, the life span of adult females is 8.68 days and the fecundity is 57 eggs/female. The immature survivorship is 89.7%, and 78.7% of the offspring are females. The intrinsic rate of increase (rm) is 0.305 individuals/female/day, the net reproductive rate (R0) is 40.12, and the ﬁnite rate of increase (λ) is 1.35 individuals/female/day, the population doubling time 2.28 days, and the mean generation time (T) is 12.12 days (Bai and Carey, 1988). It is a warm weather mite, and high populations occur only during summer and early fall (Jeppson, 1989). The populations develop throughout the year in greenhouses and in warmer areas. In temperate climates, the species migrates in late summer to the bark and crevices of the host plants, where it overwinters and remains until March. It lives in protective winter places, migrates on different hosts and re-emerges to deposit eggs. The ﬁrst populations on trees are observed at the tops or the tips of the branches, and may escape notice until the infestations become severe and leaf drop occurs (Jeppson et al., 1975). Among the natural enemies are recorded pathogens, insects and mites. The ﬁrst include some fungi, the insects eolothripids and coccinellids, for example, and phytoseiids and bdellid mites (Table 13.9). Symptomatology and damage The feeding activity causes a stippling on the leaves as it removes chlorophyll from the plant cell. The serious injury caused by low levels of populations on pear, citrus and other plants suggests the presence of a toxin. The symptoms on the leaves begin at the tops of the trees, which turn brown and die as though they had been scorched by ﬁre. The attacked plants ﬁrst become stippled, and when the mite population increases, they are covered by webbing (Jeppson et al., 1975). On citrus, the mite populations do not tend to colonize and live on the entire undersurface of the leaves and on fruit. Injury is similar to that of citrus red mite, but leaf drop and dead twig incidence rise from lower populations (Jeppson, 1989).

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Table 13.9.

Tetranychus paciﬁcus McGregor: natural enemies (fungi, insects and mites).

Natural enemies

Reference(s)

Fungi Entomophthoraceae Entomophthora sp.

Steinhaus and March, 1962

Insects Eolothripidae Franklinothrips sp.. Scolothrips sexmaculatus (Pergande) Coccinellidae Stethorus picipes Casey Mites Bdellidae Bdella longicornis (Linnaeus) Phytoseiidae Amblyseius andersoni (Chant) Euseius citrifolius Denmark et Muma Euseius hibisci (Chant) Euseius scutalis (Athias Henriot) Euseius stipulatus (Athias Henriot) Euseius tularensis Congdon Galendromus annectens (De Leon) Galendromus occidentalis (Nesbitt)

Iphiseius degenerans (Berlese) Metaseiulus pini (Chant) Metaseiulus validus (Chant) Phytoseiulus longipes Evans Phytoseiulus persimilis Athias Henriot Typhlodromus kettanehi Dosse Typhlodromus persianus McMurtry

Hoddle et al., 2000 Laminman, 1935; Rice and Jones, 1972 McMurtry et al., 1974

Sorensen et al., 1983 Amano and Chant, 1977 de Moraes and McMurtry, 1981 Zhimo and McMurtry, 1990 Bounfour and McMurtry, 1987 McMurtry, 1977a; Zhimo and McMurtry, 1990; Flechtmann and McMurtry, 1992 Zhimo and McMurtry, 1990 Badii et al., 1990 Croft and McMurtry, 1972; Rice and Jones, 1972; Hoy et al., 1982; Flechtmann and McMurtry, 1992 Takafuji and Chant, 1976; Flechtmann and McMurtry, 1992 Charlet and McMurtry, 1977 Charlet and McMurtry, 1977 Badii and McMurtry, 1983 McMurtry and Scriven, 1975; Takafuji and Chant, 1976; Amano and Chant, 1977 McMurtry, 1977c McMurtry, 1977c

Control The occurrence of this mite on citrus in USA is rather rare and of minor importance (McGregor, 1956). On other crops, the control of tetranychids leads to many problems, with populations developing resistance to several acaricides (Jeppson et al., 1975). On cotton and other crops, avermectins (Hoy and Conley, 1987; Leigh et al., 1990); phenyl pyrazoles (Hoy and Ouyang, 1986); organochlorines (Dennehy and Granett, 1982; Leigh et al., 1990); organosulphurs (Stafford and Kido, 1969; Hoy et al., 1982; Leigh et al., 1990); organotins (Rice and Jones, 1972; Hoy et al., 1982); and tetrazines (Hoy and Ouyang, 1986) have been employed.

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13.4.3.7.10 Tetranychus paraguayensis Aranda (Fig. 13.42) Common name Unknown. Diagnostic characteristics FEMALE. The distal eupathidium of the palptarsus is prominent and about 1.5 times as long as broad. The peritremes end strongly recurved. The dorsal integument is striate and the striae form a diamond-shaped pattern between the dorsal opisthosomal setae e1 and f1. The other morphological characteristics are those of the genus (Aranda, 1969; Aranda and Flechtmann, 1971). MALE.

The terminal eupathidium of the palptarsus is slender and long, about 2.5 times as long as broad. The aedeagus has a knob with anterior and posterior projections and upper side slightly sigmoid; the anterior projection is small and the posterior greater, pointed and with the tip slightly turned down (Aranda, 1969; Aranda and Flechtmann, 1971). Geographical distribution Paraguay (Aranda, 1969; Aranda and Flechtman, 1971). Bio-ecology The mite is recorded on Carica papaya Linnaeus and Citrus sp. in Paraguay (Aranda, 1969; Aranda and Flechtman, 1971). Its bio-ecology is unknown.

13.4.3.7.11 Tetranychus salasi Baker et Pritchard Common name Unknown. Diagnostic characteristics FEMALE.

The dorsal body setae are long and slender, and the dorsal striae possess rounded lobes. Tarsus I bears one sensory seta and four tactile setae proximal to the posterior duplex setae, and tibia I one sensory seta and nine tactile setae. The empodia possess a strong dorsomedial spur, about as half long as the length of the proximoventral hairs (Baker and Pritchard, 1962). MALE. Smaller than the female. Tarsus I has three sensory setae and four tactile setae proximal to the duplex setae; empodium I has the proximoventral hairs free, not claw-like, with a strong dorsomedial spur, about as half as long as the length of the proximoventral hairs. Tibia I bears three sensory and nine tactile setae. The aedeagus has the axis of the knob forming an acute angle with the axis of the shaft; the knob presents a little anterior

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angulation and an acute and large posterior angulation (Baker and Pritchard, 1962). Geographical distribution Costa Rica and Nicaragua (Baker and Pritchard, 1962). Bio-ecology The mite is recorded for Ceiba pentandra (Linnaeus) Gaertn., Chamaedorea sp., Croton niveus Jacques, Croton sp. and Citrus sp. in Costa Rica and Nicaragua (Baker and Pritchard, 1962). Its bio-ecology is unknown.

13.4.3.7.12 Tetranychus taiwanicus Ehara Common name Unknown. Diagnostic characteristics FEMALE. The body is ovoid, 450 μm long and reddish in colour. The peritremes are U-shaped distally. The terminal sensillum of the palptarsus is slightly longer than wide, with a terminal pit. The dorsal body setae are slender, pubescent and longer than intervals between longitudinal bases. The dorsal striae form a diamond-shaped pattern between the dorsal setae e1 and f1. The genital ﬂap has transverse striae; the area immediately anterior to the ﬂap has longitudinal striae. Each empodium possesses a strong mediodorsal spur and two pairs of proximoventral hairs. Tarsus I has a separate set of duplex setae, and one sensory and four tactile setae. Tibia I bears nine tactile and one sensory setae (Ehara, 1969). MALE.

The body is 320 μm long. The terminal sensillum of palptarsus is about twice as long as wide. Each empodium has two pairs of proximoventral hairs and a strong mediodorsal spur; tarsus I bears four tactile and three sensory setae, and tibia I nine tactile and four sensory setae. The aedeagus is broadly bent dorsally to form a distal, narrow part of subequal width, with caudal end truncate (Ehara, 1969). Geographical distribution China (Wang, 1981c); Hainan Island (Ma et al., 1979); Taiwan (Ehara, 1969); and Thailand (Ehara and Wongsiri, 1975). Bio-ecology Tetranychus taiwanicus has been collected on P. odoratissimus Linnaeus in Taiwan (Ehara, 1969), and on C. reticulata and Citrus sp. (Ehara and Wongsiri, 1975). The natural enemies number some phytoseiid mites, such as

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N. longispinosus (Kongchuensin et al., 2005). On citrus, damage from T. taiwanicus is secondary (Gerson, 2003) and control is unjustiﬁed.

13.4.3.7.13 Tetranychus tumidus Banks (Fig. 13.48) Common name Tumid spider mite.

A

B

C

Fig. 13.48. Tetranychus tumidus Banks. Female. (A) Tibia and tarsus I (from Pritchard and Baker, 1955); (B) pretarsus I. Male. (C) Aedeagus (from Jeppson et al., 1975).

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Diagnostic characteristics FEMALE. The body is light green to yellow in colour, marked on the back by a pair of large, U-shaped, sooty spots. The dorsal body setae are slender and longer than the intervals between longitudinal bases. The peritremes are simple and hooked distally. The dorsal opisthosomal striae form a diamondshaped pattern between the dorsal setae e1 and f1, the striae being longitudinal in the interval between each of these pairs of setae. The tactile setae of tarsus I are proximal to the duplex setae. The empodia have large spurs (Pritchard and Baker, 1955; Jeppson et al., 1975). MALE.

Smaller than the female. Empodium I is claw-like and the other empodia end in three pairs of ventrally directed hairs. The aedeagus has a broadly rounded anterior angulation and a short, acute angulation posteriorly. The knob is at right angles to the neck, and the axis of the knob is parallel to that of the shaft (Pritchard and Baker, 1955; Jeppson et al., 1975). Geographical distribution

Colombia (Urueta, 1975); Cuba (Boudreaux, 1979); Greece (Hatzinikolis, 1986b); Guam (Baker and Pritchard, 1962); Panama (Baker and Pritchard, 1962); Puerto Rico (Baker and Pritchard, 1962); Thailand (Kongchuensin et al., 2005); and the USA (Banks, 1900). Bio-ecology The tumid spider mite has been collected on 44 host plants (Migeon and Dolkerd, 2006) and occurs sometimes on citrus in south-eastern USA (Jeppson, 1989). It possesses a haploid number of six chromosomes and reproduces through parthenogenesis arrhenotkous (Helle et al., 1970, 1983). Adults and immature stages feed on leaves of host plants. The developmental periods of immature stages range from 39.6 days at 15°C to 7.4 days at 30°C. The mite fails to develop beyond the larval stage at 10°C. The lower temperature developmental thresholds for egg, larva, protonymph, deutonymph and the combined immature stages are estimated at 11.1°, 12.9°, 12.1°, 11.1° and 11.9°C, respectively. The upper temperature thresholds for development of immature stages are 25.9–35.9°C. The percentages of survivorship of immature stages vary from 56.4% to 93.7% within 15–35°C. The average longevity of adult females ranges from 48.7 days at 15°C to 7.9 days at 35°C. The average oviposition per female life span varies from 86.02 to 61.75 eggs within the temperature range of 20–30°C. However, oviposition is greatly reduced to 19.44 and 20.54 eggs at 15°C and 35°C, respectively (Liu and Tsai, 1998). The minimum development threshold for the incubation period is 14°C with a thermal constant of 35 ± 6.4 degree-days. Corresponding temperatures for the larval stages are 13.7°C with a thermal constant of 26.3 ± 5.2 degree-days; for nymphal stages, 12.9°C with a thermal constant of 50.1 ± 5.2 degree-days; and for the whole cycle from egg to adult stages, the minimum

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threshold was 13.9°C with a thermal constant of 112.6 ± 8.5 degree-days (Perez Alvarez et al., 1997). At 30°C the intrinsic rate of increase (rm) and the net reproduction (R0) present the largest values, respectively of 0.289 individuals/female/day and 59.67. At 15° and 35°C the lower values of net reproduction (R0) of 12.74 and 12.99, respectively, have been observed. The lowest values of intrinsic rate of increase (rm) (0.042) is recorded at 15°C. The mean generation time (T) of the population ranges from 60.7 days at 15°C to 11.7 days at 35°C. The optimal temperature for the development of the populations is 30°C (Liu and Tsai, 1998). The natural enemies number the phytoseiid mites N. longispinosus (Kongchuensin et al., 2005). Symptomatology and damage On citrus, the mite causes damage to fresh ﬂush leaves similar to that of the six-spotted spider mite (Jeppson, 1989). Control On Chamaedorea elegans Martius several acaricides such as organochlorines (Reinert and Neel, 1977), organosulphurs (Reinert and Neel, 1977) and organotins (Reinert and Neel, 1977) have been employed, but on citrus chemical control is not justiﬁed, because the attacks are of minor importance.

13.4.3.7.14 Tetranychus turkestani (Ugarov et Nikolski) (Fig. 13.42) Common name Strawberry spider mite. Diagnostic characteristics FEMALE. The body is variable in colour, from amber, green, brownish or almost black in the summer females to bright orange in overwintering females; present on each side is a broad spot, behind the eyes and extending beyond the middle of the body, and one black spot for each side towards the end of the opisthosoma. The dorsal body setae are slender and longer than the intervals between longitudinal bases. The opisthosomal dorsal striae between the dorsal setae e1 and f1 are longitudinal, and transverse striae between the latter two pairs of setae form a diamond-shaped pattern. The dorsal lobes of the striae are semicircular and sometimes triangular; several broad, ventral lobes are present between the third and fourth pairs of ventral setae. On tarsus I the tactile setae are proximal to duplex setae. The empodia end in three pairs of ventrally directed hairs and there is no dorsal spur (Pritchard and Baker, 1955; Meyer Smith, 1987). MALE.

Smaller than the female. Empodium I is claw-like and other empodia are similar to those of the female. The aedeagus has a distal knob moderately

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enlarged, about a quarter as long as the dorsal margin of the shaft. Anterior projection broad and rounded, and posterior angulation small and acute, the axis of the knob forming an angle with that of the shaft (Pritchard and Baker, 1955; Meyer Smith, 1987). Geographic distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology The strawberry spider mite has a wide host range of about 207 host plants (Migeon and Dorkeld, 2006), including low-growing forage, vegetable and fruit crops, C. sinensis (Tuttle and Baker, 1968) and Citrus sp. (Jeppson et al., 1975; Mijuskovic, 1981; Kreiter et al., 2002); in southern California, the species occurs rarely, appearing as a result of straggling from other plants growing under or near the trees, but it is not of any economic importance on citrus (McGregor, 1956), whereas it is a serious pest of cotton in different region of the world (Leigh, 1985). The haploid number of chromosomes is three and the reproduction is parthenogenetic arrhenotokous (Helle et al., 1970). It feeds mainly on the lower surface of the leaves and produces webbing, which may cover leaves and stems together. High daily temperatures and low rainfall favour its development (Canerday and Arant, 1964). The species overwinters as bright orange females, which appear at the end of September, whereas the summer forms disappear by the end of October. The overwintering females shelter in litter, under bark scales and in the soil (Mellot and Connell, 1965). The mite hosts the endosymbiont Wolbachia bacteria, responsible for cytoplasmatic incompatibility (Breeuwer, 1997). The pre-oviposition ranges from 1 to 6 days. The female lays an average of 7.4 eggs per day during midsummer, but later in the summer and autumn each female lays two to three eggs per day. The female longevity is about 8 days in the summer and 33 days in autumn (Cagle, 1956). On cotton cotyledons, the average development periods range from 25.8 to 29.0 days at 15.5°C and from 6.1 to 6.7 days at 29.4°C for the females, and a slightly shorter time for the males. The total progeny at 23.8°C averages 84.6, and at 29.4°C is 73.5 eggs/female. The female offspring is 55%. The females live for means of 12.46 days at 23.8°C, and 8.79 days at 29.4°C. At 25 ± 1°C on cotton cotyledons, the intrinsic rate of increase (rm) is 0.203 individuals/female/day, the net reproduction (R0) is 46.8 and the ﬁnite rate of increase (λ) is 1.225 individuals/female/day (Carey and Bradley, 1982). The mite develops from eight to 16 generations per year (Cagle, 1956). Natural enemies include the coccinellid Stethorus gilvifrons, which preys voraciously on different growth stages of T. turkestani (Ahmed and Ahmed, 1989; Afshari, 1999), and in Pakistan have been investigated in laboratory for their functional and numerical responses (Sohrabi and Shishehbor, 2007).

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Symptomatology and damage The strawberry spider mite feeds mainly on the lower surface of the leaves, and the damage shows on the upper surface as dead areas at the point of feeding on the leaf. High infestations produce leaves drop and the plants can die, and the webbing mats leaves and stems together (Jeppson et al., 1975). The mite injects a local toxin into the plants during the feeding process (Simons, 1964). On citrus the symptomatology of the attack and the damage are unknown, and the control is unjustiﬁed.

13.4.3.7.15 Tetranychus urticae Koch (Figs 2.1, 2.2, 2.4, 2.5, 2.6, and 13.42) Common name Two-spotted spider mite. Diagnostic characteristics FEMALE. The body is variable in colour; the overwintering forms are yellowishorange and the summer forms are similar in colour to strawberry spider mite but with one conspicuous black spot on each side. The peritremes end simply. The dorsal body setae are ﬁnely serrate, slender and longer than the intervals between longitudinal bases. The striae between the dorsal setae e1 and f1 form a diamond shape. The dorsal lobes on the striae are triangular to semicircular (Vacante, 1985). MALE.

Smaller than the female. Empodium I has a prominent mediodorsal spur and two proximoventral spurs. The aedeagus has a small knob, with the axis of the knob parallel to axis of the shaft and the anterior and posterior angulations of the knob small and similar (Vacante, 1985). Geographical distribution Worldwide distribution (Migeon and Dorkeld, 2006). Bio-ecology

The species is recorded about on 1054 botanical species, spontaneous and cultivated (Migeon and Dorkeld, 2006), mostly including ornamental plants and the important agricultural crops such as C. aurantifolia (Doreste, 1967); C. aurantium (Hatzinikolis, 1969; Zaher et al., 1982); C. decumana (Hatzinikolis, 1969); C. grandis (Meyer Smith, 1974); C. limon (Womersley, 1940; McGregor, 1950; Hatzinikolis, 1969; Nucifora, 1981); C. medica (Hatzinikolis, 1969); C. paradisi (Cuenod, 1956; Hatzinikolis, 1969); C. reticulata (Hatzinikolis, 1969; Meyer Smith, 1974), C. sinensis (Cuenod, 1956; Hatzinikolis, 1969; Gupta and Gupta, 1994; Martinez et al., 2004); and Citrus sp. (Ehara, 1956; Rambier, 1958; Gupta and Gupta, 1994; Kreiter et al., 2002; Suarez, 2004).

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The haploid number of chromosomes is three and the the reproduction is parthenogenetic arrhenotokous (Helle and Bolland, 1967). The sex ratio is three females: one male. Tetranychus urticae red and green forms are well known, and the red form has overlapping niches with T. turkestani in which reproductive interactions between the two species are likely to occur naturally, and a post-zygotic reproductive barrier exists between T. turkestani and T. urticae red form (BenDavid et al., 2009). The mite is infected by endosymbiont Wolbachia bacteria (Tsagkarokou et al., 1996; Gotoh et al., 2004), responsible for cytoplasmatic incompatibility (Breeuwer, 1997) in European populations, but not in Japan (Gotoh et al., 2004). The two-spotted spider mite commonly lives on young leaves, feeding on the upper leaf surface, but at greater densities it invades all plant surfaces, blooms and fruit. Large populations invade also the older leaves, and the webbing production may cover the entire plant. According to Satô (1985), the mite belongs to the subtype CW-u, characterized by a complex construction web on the leaf surface. On rose leaves at 20°C the pre-oviposition period is 1.7 days, and the development of egg, larva, protonymph and deutonymph stages requires 6.7, 2.8, 2.3 and 3.1 days, respectively. The development of a generation at 15°, 20° and 30°C requires 36.3, 16.6 and 7.3 days, respectively (Sabelis, 1981). On bean leaves at 20°C the female lays 37.9 eggs, with a daily average of 2.4 eggs, and with higher temperatures up to 200 eggs (Laing, 1969). On cotton cotyledons the average development periods range from 25.8 to 29.0 days at 15.5°C and from 6.1 to 6.7 days at 29.4°C for the females, and a slightly shorter time for the males. The fecundity at 23.8°C averages 103.3 eggs/female and at 29.4°C averages 64.3 eggs/female. Females comprise 74% of offspring, living for means of 14.71 days at 23.8°C, and for 9.71 days at 29.4°C (Carey and Bradley, 1982). The intrinsic rate of increase is 0.0067 for each degree, and the thermal threshold is 12°C (Sabelis, 1981). At 25 ± 1°C on cotton cotyledons, the net reproduction (R0) is 74.8, the intrinsic rate of increase (rm) is 0.219 individuals/female/day and ﬁnite rate of increase (λ) is 1.245 individuals/female/ day (Carey and Bradley, 1982). The species develops in glasshouses and in the ﬁeld, and their populations are constantly active in the warmer regions of the world, while overwintering as diapause females in the temperate climates. Shortened periods of light, decreased temperatures and unfavourable food supplies stimulate the appearance of diapause females, which stop feeding and laying eggs, leave the plants, change in colour and overwinter in various protected sites (Cagle, 1949; Gasser, 1951). In warmer climates the populations continue to reproduce on their host plants (Jeppson et al., 1975). In relatively mild and cooler climates, the species overwinters on citrus in a diapause form, and although the diapause forms commonly leave the plant host, in San Joaquin Valley in California they are found on the citrus trees. Normally, the populations develop on weeds or cover crops, and when these plants dry or are cut, they move to citrus where they cause damage to leaves on lower limbs (Jeppson,

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1989). Similar behaviour has been registered on citrus in southern Italy, where the species mainly infests the lemon and, in certain conditions, other citrus crops (Vacante, 1995). The female lays the eggs on the underside of leaves, mainly along the midrib, and with severe levels of attack the eggs are laid on both leaf surfaces and on fruit. On mandarin trees, high salt concentrations negatively affect the twospotted spider mite, but the rm values obtained at the lowest concentrations are signiﬁcantly higher than those of the control, and this may contribute to the observed outbreaks of the mite (Ansaloni et al., 2006). On lemon in Sicily (Italy), it has been observed that the two-spotted spider mite is also active in winter and presents three peaks – in July, November and February, respectively (Vacante, 1986). On clementines in Spain, the populations of two-spotted spider mite show three peaks during winter, late spring and autumn. Apparently, population increases in the trees and green cover are related to a drop in humidity, whereas ﬂuctuations in temperature have less effect on the seasonal trend (Pascual and Ferragut, 2003). Workers’ allergies to T. urticae are recorded (Burches et al., 1996). The natural enemies include pathogens, predatory insects and mites. Among the former are recorded some fungi. The insects include conyopterigids and coccinellids, and the mites several species of phytoseiids and cheyletids (Table 13.10). Symptomatology and damage On herbaceous plants, the symptomatology and damage are similar to those of the strawberry spider mite. McGregor (1956) reported in California that the two-spotted spider mite was an incidental pest of citrus trees, their presence originating from nearby alternate hosts by dispersion, and their resulting damage on citrus of minor importance, with the exception of the nurseries repeatedly treated with insecticides. Successively, in the same region (San Joaquin Valley), the mite increased its damage on citrus, similarly to that of the six-spotted spider mite, E. sexmaculatus, its feeding activity on the undersides of the young leaves producing chlorotic areas visible on the upper surface, with severe injury possibly resulting in leaf drop (Jeppson, 1989). In semitropical areas of the world, populations of the two-spotted spider mites infest the young leaves and green or mature fruit of all citrus species. Commonly, they develop on limited portions of the leaves. The leaves buckle at the site invaded by the colonies, and the upper surface becomes raised and yellow ochre. On orange, lemon and other citrus fruit, the feeding activity of the mite produces a blackish area around the navel end of fruit, and increases in populations may involve the whole fruit (Lewis et al., 1951; Dosse, 1964; Di Martino, 1985; Vacante, 1995, 2009a). In the lemon fruit of southern Italy, the problem is very common and normally involves only the blossom end of the lemons (Di Martino, 1985; Vacante, 1995). At present, the pest status of the species on citrus worldwide is considered of minor importance.

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Table 13.10. Tetranychus urticae Koch: natural enemies (fungi, insects and mites). Natural enemies

Reference(s)

Fungi Exobasidiomycetidae Acaromyces ingoldii Boekhout et al. Hirsutella thompsonii Fisher Meira argovae Boekhout et al. Meira geulakonigii Boekhout et al. Neozygitaceae Neozygites ﬂoridana Weiser et Muma

Paz et al., 2009a Aghajanzadeh et al., 2006 Paz et al., 2009a Paz et al., 2009a

Insects Coccinellidae Clitostethus arcuatus (Rossi) Stethorus punctillum Weise Stethorus utilis Horn Conyopterigidae Conwentzia psociformis (Curtis)

Liotta, 1981 Abad et al., 2006 Mora Morin, 1991 Ripolles and Melia, 1980; Abad et al., 2006

Mites Cheyletidae Acaropsellina sollers (Rohdendrof)

Nassar and Kandeel, 1986

Buergo et al., 1986

Phytoseiidae Amblyseius zaheri Yousef et El-Borolossy Euseius mesembrinus (Dean) Euseius stipulatus (Athias Henriot)

Rasmy et al., 2003 Abou-Setta and Childers, 1987 Vacante, 1986; Ferragut et al., 1987; Grafton Cardwell et al., 1997 Neoseiulus californicus (McGregor) Buergo et al., 1986; Grafton Cardwell et al., 1997; Ibrahim et al., 2005; Abad et al., 2006 Neoseiulus cydnodactylon (Shehata et Zaher) El-Banhawy et al., 2000 Neoseiulus longispinosus (Evans) Kongchuensin et al., 2005 Neoseilus picanus (Ragusa) Ragusa et al., 2000 Galendromus helveolus (Chant) Caceres and Childers, 1991 Galendromus occidentalis (Nesbitt) McMurtry, 1985; Grafton Cardwell et al., 1997 Phytoseiulus longipes Evans Grafton Cardwell et al., 1997 Phytoseiulus persimilis Athias Henriot Buergo et al., 1986; Abad et al., 2006 Phytoseius plumifer (Canestrini et Fanzago) Nawar et al., 2001a Proprioseiopsis badryi (Yousef et El-Borolossy) Abou-Awad et al., 1998a Typhlodromus athiasae Porath et Swirski Nawar et al., 2001b Typhlodromus exhilaratus Ragusa Ragusa, 1981 Typhlodromus phialatus Athias Henriot Ferragut et al., 1987

Control Guidelines for the control of the two-spotted spider mite in the citrus groves of the Mediterranean region suggest examination of four shoots per plant (newest vegetation) and in situ of 20 fruit per tree on 10% of plants. The

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threshold consists of 2% of fruit infested and/or 8–10% of leaves occupied by active colonies of the mite (Cavalloro and Prota, 1983). Successively, the need to rationalize control has been suggested for clementine in Spain, because the mite shows an aggregated pattern of distribution. The sampling method comprises random choice of both the primary units (trees in orchard) and secondary units (leaves or fruits of each tree). The sampling plans combine the observation of symptomatic leaves at a 0.25 m2 diameter and pest population density (Martinez Ferrer et al., 2006). Other methods, such as photographic experience with ‘false colour’ and infrared ﬁlm for localization of some unhealthy conditions in a citrus grove, provide a less practical approach (Spicciarelli and Arpaia, 1991). BIOLOGICAL CONTROL.

The natural enemies alone do not assure a constant biological control and pose a need to integrate their action with selective acaricides (da Silva and de Oliveira, 2006). In bioassay on tomato leaﬂets and in an experimental greenhouse on tomato plants, Chandler et al. (2005) have evaluated the action of isolates of Beauveria bassiana, H. thompsonii, Metarhizium anisopliae and V. lecanii, and concluded that B. bassiana assures the best control in reducing the number of adults, nymphs and eggs of the two-spotted spider mite by 97–98%. Moreover, we do not have any information on citrus. Recently, Paz et al. (2009a) essayed in the laboratory the actions of the fungi M. geulakonigii, M. argovae and A. ingoldii and observed that after 14 days, M. geulakonigii and A. ingoldii produce 81.9% and 90% mortality respectively, whereas M. argovae in the same conditions assures only 46.1% of mortality. At this moment, we do not have practical indications of employment of these pathogens.

CHEMICAL CONTROL. Also in this case it is desirable to have knowledge of the presence of resistant strains in the ﬁeld (Garcia Marì, 2005). Several chemicals have been positively examined, such as avermectins (Aucejo et al., 2003; Conti et al., 2005); organochlorines (Osman and Rasmy, 1976; Di Martino, 1985; Buergo et al., 1986; Garcia Marì et al., 1988; Vacante, 1995; Aucejo et al., 2003); quinazolines (Aucejo et al., 2003); organotins (Vacante, 1995; Aucejo et al., 2003); organosulphurs (Di Martino, 1985; Garcia Marì et al., 1988; Vacante, 1995; Aucejo et al., 2003); petroleum oils (Nucifora, 1981; Di Martino, 1985; Buergo et al., 1986; Vacante, 1986, 1995; Aucejo et al., 2003; Conti et al., 2005); pyrazoles (Aucejo et al., 2003); phenyl pyrazoles (Aucejo et al., 2003); pyridazinones (Aucejo et al., 2003); sulfur (Buergo et al., 1986; Conti et al., 2005); and tetronic acid derivates (Izquierdo et al, 2002), alone or variably mixed in combination. INTEGRATED PEST MANAGEMENT.

IPM represents a better solution for control of mite attacks on citrus. The strategy demands the use of selective acaricides and insecticides, and particularly of petroleum oils (Vacante, 1986, 1995, 2009a).

14

Conclusions

The book has examined different groups of mites characterized by typical ecological adaptations and various levels of harmfulness. Many featured species, although reported in the literature for citrus and in more than one case for other host plants, are unknown as regards the bio-ecology and pest status. Other species are more speciﬁcally investigated. To some extent, the use of modern research approaches (biochemical, biological, genetic, statistical, etc.) has revealed important aspects of their bio-ecology that signiﬁcantly expand the framework of knowledge and help better to plan their control. This examination cannot be regarded as exhaustive and invites specialists to ﬁll the inevitable gaps found in the work, suggesting modiﬁcations and/or updates that may mprove it. From this point of view, any study of a systematic nature, both biological and ecological, can only help to resolve better the applied issues of the problem. Let us brieﬂy summarize the most important aspects in order to frame this complex topic in a context of greater rationality.

14.1 SYSTEMATICS The existence in the different taxa of several systematic problems has been highlighted, including the need for a radical revision of the group, as in the case of the tenuipalpidae, taking care to complement the traditional morphological analysis of individuals with that of molecular tools (Navajas and Boursot, 2003; Ben-David et al., 2007, 2009), in order to better understand the biological and genetic structure of species, identifying where possible those adjustments that facilitate control, as in the case of clones of populations of Brevipalpus phoenicis already reported (Weeks et al., 2000), adapted to different environmental conditions and host plants (Groot et al., 2005) and variously capable of transmitting viral diseases (Rodrigues et al., 2003). 298

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14.2 BIO-ECOLOGY The bio-ecology of many species is unknown. This deﬁciency is mainly for three reasons: (i) the difﬁculty of carrying out studies on the subject in certain regions of the world; (ii) lack of adequate facilities and/or expertise, the sporadic ﬁndings of which do not allow any intervention; and (iii) the inconsistent pest status of many species, which is not a priority for the attention of specialists. Many species are available only to study in a traditional museum, with a small number of specimens mounted on glass slides, sufﬁcient for classic morphological analysis, but absolutely useless for any other type of scientiﬁc approach. Aggravating the situation is the propensity of many specialists for the study of issues of a systematic or wildlife nature rather than studies of bio-ecology and/or control. This calls for a greater importance to be attached to the study of bio-ecology of various species, including those designated with either minor or unknown pest status. In this context, there is a need to study the biological mechanisms within the species, such as the role of endosymbiont bacteria (Breeuwer, 1997; Gotoh et al., 2004), the relationship between host plants and pests, and biotic and abiotic factors involved in population dynamics. More information on the ecology of different species would demarcate with greater certainty the range of distribution of various entities and build predictive models of risk of their acclimation following an accidental introduction.

14.3 PEST STATUS Many species are formally insigniﬁcant in terms of damage, and their presence in no way constitutes problems in citrus and consequently does not require control. However, the reporting of species marked by an unknown pest status is not generally without risk, which could in part be related to either inadequate assessment of the problem in the ﬁeld or the intervention of subsequent, unforeseen factors. From this point of view, it is necessary to evaluate the actual pest status of each insufﬁciently known entity, to build in different areas of the world a reliable picture of the risk associated with their presence and avoid irrational choices that can stimulate the development, as in the case in Egypt of Brevipalpus californicus (but it is not the only case), where the employment of phosphorus compounds may be causing an increase in mite populations (Attiah and Wahba, 1973); or of Eutetrancychus orientalis in the same country, where if citrus orchards are not treated with chemicals constantly the mite is no longer a problem (Rasmy, 1969), and if no sprays are used it does not become serious. If sprays are applied against other citrus pests, it is necessary to add acaricides to control phytophagous mites (Rasmy, 1970a). The need arises, within certain limits, even for better-known pest species – as in the case of the citrus bud mite, Aceria sheldoni, where ﬁeld evaluations of its harmfulness on lemon in Italy (Vacante and Nucifora, 1984) and California (Hare et al., 1999) have shown modest damage. A further examination

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conducted on lemon by Vacante et al. (2007) in southern Italy suggested the hypothesis of a mutual symbiosis between the eryophioid and lemon, where the mite adjusted the ﬂowering of the host plant, which offers protection and nutritional resources for the citrus bud mite, and only in special cases (nursery, young lemon plants, Valencia orange, etc.) did it become harmful. Indeed, in Italy, only 1–2% of lemon fruit appears deformed by the mite despite its presence on 100% of the buds. This contrasts with the suggestions of various Italian researchers who have recklessly set empirical thresholds for the control of the mite on lemon equal to about 30% of infested buds (Areddia et al., 2000; Barbagallo, 2000) or between 30% and 50% (Asero et al., 1994), allowing a paradoxically unnecessary consumption of pesticides. Another interesting example, already mentioned in the text, covers the North and Central American populations of Brevipalpus phoenicis (Weeks et al., 2000), consisting of several host-specialized clones instead of one generalist form, each adapted to different environments and host plants (Groot et al., 2005) and capable of transmitting leprosis (Rodrigues et al., 2003), while the Mediterranean populations are incapable of transmitting this disease.

14.4 NATURAL ENEMIES The natural enemies of mites injurious to citrus include insects, mites and pathogens. Under natural conditions, the indigenous biodiversity hinders (in varying degrees depending on the species) the development of populations of harmful mites. Moreover, you can enrich the complex of natural enemies by means of classical biological control, based on introduction and acclimation of beneﬁcials introduced from other regions of the world. In several cases, the information provided is not exhaustive; complete information included natural enemies found only in association with other injurious mites but without the certainty of a interspeciﬁc interaction. The choice was basically dictated by the need to make a not too rigid selection that would have eliminated a priori or made less obvious assumptions and/or hypotheses for work worthy of attention. In any case, it is possible to integrate the picture given with recent contributions on the subject of other specialists (Gerson et al., 2003). Most of the bibliographic contributions show a priority interest in the phytoseiid mites, witnessed by many studies on systematic, bio-ecology or biological control programmes and/or IPM. In the context of systematic, there is a considerable increase in the described species, from about 1500 in 1986 (Moraes et al., 1986) to about 2250 in 2004, of which only 81 were collected for the ﬁrst time on citrus (Moraes et al., 2004 ); moreover, the bibliographic records indicate that their number is greatest on citrus. However, most taxonomic studies are based on traditional morphological examination, and hopes both for injurious and beneﬁcial species increased the use of molecular tools needed to identify races, biotypes and cryptic species. The remaining predators, such as cheiletids, stigmaeids etc., do not play a role similar to that of phytoseiids and their contribution, although it is

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sometimes relegated to peculiar ecological conditions and not easily generalizable. In the context of natural enemies playing an important role in containing various injurious mites, several pathogens such as bacteria and entomopathogenic fungi have been cited. The former are more extensively investigated and could also allow short-term interest in the technical aspects (Paz et al., 2009a). The implementation of the method needs to identify and/or select those strains most suitable for investigation of their bio-ecology and industrial production. This is similar to that of pesticides, and the use of pathogens leads to a lower psychological impact on the farmer than a common biological means (packs containing the predatory mites or various entomophagous organisms). The framework established conﬁrms the importance of natural enemies in control, and calls for knowledge in the ﬁeld, in agreement with a modern ecological view of citrus groves, understood as an agro-ecosystem with a good resilience and a low resistance (Vacante, 2009b). In this particular environment, the concept of ‘disturbance’ (Pickett and White, 1985) assumes particular importance and also involves the disciplines of landscape ecology and ‘habitat management’ (Landis et al., 2000), making the research of Altieri and Letourneau (1982) and Gravena et al. (1993) on the inﬂuence on citrus of green cover on predatory and phytophagous mites in citrus very relevant.

14.5 MEANS OF CONTROL Regarding the use of chemical, the rapid evolution of the market for pesticides following the withdrawal of many substances and introduction to the market of new molecules has suggested that we focus on information on the main chemical families of acaricides, to avoid providing technical information easily overtaken by events. Actually, the possibilities for chemical control by existing compounds seem limited and, as regards the future, new technologies may be established through the introduction of new molecules, improving existing products – such as some isomers of known molecules – and looking for new solutions, such as activator chemistry in the control of plant diseases (Carroll, 2000). This strategy is proﬁtable in terms of control of the various pests, but we cannot always be sure with regard to the impact of new substances on the environment. In this instance, the experience with derivatives of tetronic acid on pollinators (http://www.koppert.nl/side_ effects.htlm, 2007) and phytoseiids is signiﬁcant (Gravena et al., 2004; Rodrigues and Torres, 2007). As an alternative to chemical control, biological agents and different natural means are proposed. In the context of biological agents, the more concrete prospects can be derived, as well as broader and wiser use of natural enemies present in ﬁeld, from extensive entomopathogenic fungi and some bacterial derivatives. The prospect for natural substances seems less encouraging. Indeed, as regards bioinsecticides and also bioacaricides, in 1998 the USA global

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end-user sales were estimated at US$85–120 million – corresponding to around 0.3–0.4% of total pesticide sales (Meneley, 2000). Obviously, the sales of acaricides are less. In this context, it is difﬁcult to forecast the types of chemicals and means of securing more sales, although it is certain that the study of biochemical and genetic mechanisms involved in the biology of different species and in relation to the host plant can offer valuable insights.

14.6 HORTICULTURAL PRACTICES Several biotic and abiotic factors and horticultural practices (Jeppson et al., 1975; McCoy, 1977; Zamorra and Nasca, 1985; Smith and Papacek, 1991) can inﬂuence the population density of injurious mites. In this case, for example, the role of fertilization is known (Puttaswamy and Channabasavanna, 1982; Jackson and Hunter, 1983). In B. phoenicis, horticultural practices intended to limit the spread of leprosis and the Tenuipalpids include: (i) employing resistant varieties, disease- and/or mite-free plants; (ii) removing or pruning infected trees or parts; (iii) avoiding hedge windbreak plants that are heavily infested or plants that are alternate hosts for the mites; and (iv) removing weeds and other susceptible plants from the area (Maia and Oliveira, 2004). In B. californicus, pruning and successive periods of dry weather with light rainfall may have favoured their development (Baptist and Ranawerra, 1955), and in Oligonychus coffeae the pruning of tea removes many of the old leaves and the shoots, and thus a large proportion of the mite population (Das, 1959, 1960; Hu and Wang, 1965; Das and Das, 1967). In general, it was found that the proper management of horticultural practices hampers the development of pest mites (and other pests) in citrus. However, despite the experiences highlighted in this case, the bibliography lacks matter that constitutes a call for research. This point of view seems to suggest the need for appropriate research and/or intensifying that in progress.

14.7 PREVENTION The control of injurious mites may involve several aspects and may require different methodological and technical approaches. In this context, prevention plays a key role, usually making it easier to eradicate or control a new plant problem (pests, diseases, weeds). In this case, the information presented appropriately directs the activities of those responsible for prevention, in terms of directing them towards involvement of successive levels of intervention specialists and/or operatives. This provides a character of unity in a context of time where no updated information about the subject is available, also assigning motivation to the treatment of those species described as having unknown pest status. Moreover, the work tends to address the discrepancies between the risk associated with the actual introduction of injurious species and that provided by the legislation in force. To meet this need, the

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EPPO (European and Mediterranean Plant Protection Organization) annually sends to the member countries an Alert List of species (mainly selected on the basis of the literature) that may pose a serious risk. These species are not automatically included in the quarantine lists, the latter operation requiring a long process, including the approval of the member states, which does not allow any rapid action. European Union countries have a quarantine legislation (Directive 2000/29/EC), by which the Plant Protection Services offer a number of preventive measures and also inspection, including the correct identiﬁcation of species, the speed of operation and implementation of preventive measures. In the case of mites, these are unfortunately quite inadequate. Indeed, in the face of the many species at risk of introduction, for example, in the year 2005 the EPPO List A1 contained only the eryophiid Aculops fuchsiae Keifer and the tetranychid Oligonychus perditus Pritchard et Baker, while the A2 List hosted the tetranychid E. orientalis. Equally inadequate are the guidelines in Annex II Community that prohibit the presence of E. lewisi on plant parts of Citrus, Fortunella and Poncirus and their hybrids, but admitted it on the fruits, whereas E. orientalis is not allowed on the green parts of Citrus, Fortunella and Poncirus but admitted on the fruits (Part A, Sec. II). However, whereas E. orientalis lives almost exclusively on leaves and fruits, its discovery is accidental and not the rule, and in the case of E. lewisi, the reverse is true, making the species substantially carpophagous. This case highlights blatant disinformation, which is compounded by the fact that the legislation deals with only the above two species in the face of dozens taken into consideration in the text. This regulatory gap and organization results in an increased risk if it is true that, for example, in Italy from 1945 until the present 152 species of insect pests and ten species of mites have been accidentally introduced (Pellizari and Vacante, 2005; Pellizzari et al., 2005). In the year 2008, the EPPO A1 List made no reference to these species, while the A2 List reported E. orientalis and the tomato red spider mite, Tetranychus evansi Baker et Pritchard. The Alert List of 2009 (last updated February) shows only the red palm mite, Raoiella indica Hirst, as being injurious to palms (EPPO, 2009). The complexity of the topic does not allow any simpliﬁcation and standards and operational choices must be set, supported by more appropriate information technology to manage with greater rigour the movement of plant material and/or nurseries worldwide.

14.8 INTEGRATED PEST MANAGEMENT As regards control, it is more productive to report the presence of injurious mites and their natural enemies, and every technical choice, from the horticultural to those strictly phytoiatric, on account of the impact that can determine the populations of pests and beneﬁcials. It is better to present each choice, for both ecotoxicological and economic reasons, as part of IPM strategies. It can be concluded, according to Hoy (2000), regarding the biological control of insects, that in the case of injurious mites the potential solutions for

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implementation of classical biological control involve the application of molecular tools, the knowledge of the effects on non-target species and global cooperation. Moreover, augmentative biological control requires knowledge of release rates, timing and monitoring methods, and on the quality and purity of natural enemies. An important role is played by educational challenges. This involves various aspects of research and, according to the aspects variously discussed in the book, it is necessary to know the systematics and the bionomy of the various species, both injurious and beneﬁcial. In this respect the efforts of specialists can also be directed towards the application of modern study technologies and, according to Vacante and Gutierrez (1997), the principal research areas involved during recent years by various researchers have involved the use of information application and mathematical models, population dynamics, the behaviour of prey and predators, population genetics and also molecular biology.

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Index

Page numbers in bold type refer to ﬁgures and tables. Acanthonychus jiangfengensis 49, 219, 220 Acaricides avermectins 82, 98 carbamates 34 dithiocarbamates 73, 75 petroleum oils 66, 98 sulfur compounds 75, 87, 98, 111–112 tetronic acid derivatives 76, 98, 301 Aceria 43 Aceria sheldoni (citrus bud mite) 43, 62, 63–66 biochemical effect on host 30, 65 bud damage, symptoms and effects 29, 64–66 chemical control 66, 87, 299–300 pest status and climate 31, 63–64 Aceriini, tribe identiﬁcation 61 Aculops 44 Aculops pelekassi (pink citrus rust mite, Japanese citrus mite) 44, 60, 70–76, 71 cycle of host attack 28, 73, 74 effect on fruit yield and sugar concentration 30, 74 epidermal cell damage, fruit 29, 74–75 Aculops suzhouensis 44, 76

Aculus advens 44, 76–78, 77 Aedeagus 16, 17, 23, 176, 268 African red mite (Eutetranychus africanus) 51, 197, 199, 199–201 Alternate host plants, control 150, 281, 295, 302 Angolan citrus mite (Meyernychus emeticae) 49, 217–219, 218 Anthocoptini, tribe identiﬁcation 70 Aplonobia 48 Aplonobia citri 50, 183–185, 184 Aplonobia histricina 50, 185, 187, 187–188 Aplonobia honiballi 50, 185, 186 Aponychus 48 Aponychus chiavegatoi 50, 195–196, 196 Aponychus spinosus 50, 197, 198 Avermectins 82, 98

Bacteria endosymbionts and cytoplasm incompatibility (Wolbachia sp) 145, 270, 292, 294 feminizing (Cardinium sp) 121, 142, 145 pathogenic to mites 35, 301

371

372

Index Biological control bacteria 35, 301 fungi 96–97, 253, 301 insect predators 35, 97 predatory (beneﬁcial) mites 34, 35, 300 see also Natural enemies Brevipalpinae, subfamily identiﬁcation 117 Brevipalpus 46 Brevipalpus amicus 46, 118, 118–119 Brevipalpus californicus (citrus ﬂat mite) 47, 119–123, 120, 299, 302 Brevipalpus chilensis (grape ﬂat mite) 46, 123–125, 124 Brevipalpus cucurbitae 47, 125–127, 126 Brevipalpus cuneatus 47, 127–129, 128 Brevipalpus deleoni 47, 129, 129–130 Brevipalpus dosis 47, 130–132, 131 Brevipalpus jambhiri 47, 132 Brevipalpus jordani 47, 133, 134 Brevipalpus karachiensis 47, 134–136, 135 Brevipalpus lewisi (citrus ﬂat mite, scab/ bunch mite) 47, 136, 136–139 Brevipalpus mcgregori 46, 139, 139–140 Brevipalpus obovatus (ornamental ﬂat mite, privet mite) 46, 140–143, 141 Brevipalpus phoenicis (leprosis mite, red and black/reddish black ﬂat mite, red crevice mite, scarlet mite) 47, 115, 116, 143–151, 144 control measures, by horticultural practices 150, 302 morphology, compared with Brevipalpus jordani 133 natural enemies 147, 150 population clone variations 148, 298, 300 symptoms, compared with Aculops pelekassi 75 see also Leprosis (viral disease, CiLV) Brevipalpus phoenicoides 47, 133n, 144, 151 Brevipalpus rugulosus 47, 152, 152–153 Brevipalpus tinsukiaensis 47, 153–154 Broad mite see Polyphagotarsonemus latus Brown citrus rust mite (Tegolophus australis) 45, 79–82, 80

Brown mite (Bryobia rubrioculus) 50, 181–183, 182 Brown wheat mite (Petrobia latens) 50, 190–192, 191 Bryobia 48 Bryobia graminum 50, 177–179, 178 Bryobia praetiosa (clover mite) 50, 179–181, 180 Bryobia rubrioculus (brown mite) 50, 181–183, 182 Bryobiinae, subfamily identiﬁcation 177 Bryobiini, tribe identiﬁcation 177 Bud mites see Aceria sheldoni; Phytoptus ﬁcivorus Bunch mite (Brevipalpus lewisi) 47, 136, 136–139

Calacarini, tribe identiﬁcation 83 Calacarus citrifolii (citrus grey mite, citrus blotch mite) 43, 84, 84–87 toxicity of saliva 30, 86 wax secretion (symptom of attack) 28, 85 California damage assessment 32, 253 harmful mite species 31 Carbamates 34 Cassava red spider mite (Tetranychus kanzawai) 53, 268, 269–271 Cecidophyinae, subfamily identiﬁcation 67 Chaetotaxy 12, 117, 164 Chelicerae (mouthparts) modes of action 26–27 morphology 15, 15, 19, 25, 26 Chemical control market-related factors 10, 95 new product research 34, 301–302 of other citrus pests 214, 299 side effects of use 34–35, 97, 150–151, 301 treatments used against mites 33 see also Acaricides China, effect of temperature on infestation 31 Circaces citri 42–43, 67–68 Citrus blotch mite see Calacarus citrifolii Citrus brown mite see Eutetranychus orientalis Citrus bud mite see Aceria sheldoni

Index

373 Citrus crops global production 3–4, 7 pest and disease damage 9, 25 species cultivated 4–7 see also Crop protection; Market destinations Citrus ﬂat mite see Brevipalpus californicus; Brevipalpus lewisi Citrus green mite (Schizotetranychus baltazari) 52, 257–259, 258 Citrus grey mite see Calacarus citrifolii Citrus red mite see Panonychus citri Citrus rust mite see Phyllocoptruta oleivora Citrus silver mite see Polyphagotarsonemus latus Citrus yellow mite see Eotetranychus cendanai; Eotetranychus kankitus Classiﬁcation see Systematics Clearing, mite specimens 22 Clementine (Citrus reticulata) 7, 8 Climate, inﬂuence on mite populations 10, 31, 63–64, 91, 294–295 Clover mite (Bryobia praetiosa) 50, 179–181, 180 Collection techniques from ground 21 from plants 20–21, 95 Colomerini, tribe identiﬁcation 67 Control of mite pests see Biological control; Chemical control; Integrated pest management (IPM) Cosella ﬂeschneri 42, 68–70, 69 Crop protection assessment of risk 32, 95–96, 299–300 costs of mite control 10, 32, 36, 148 mite population monitoring 32, 33, 203, 252–253 mite-resistant crop varieties 150, 302 pest introduction alerts 32, 150, 302–303

Damage biochemical effects 29, 30, 65 to embryonic tissue 29, 64–65 mechanical (cellular) 27, 28–29

virus/fungal pathogen transmission 30, 148, 149 see also Symptoms Desert spider mite (Tetranychus desertorum) 53, 263–265, 264 Diptilomiopidae characteristics and taxonomy 41, 101 pest status and distribution 59, 101 Diptilomiopidae, subfamily identiﬁcation 102 Diptilomiopus assamica 45, 102–103, 103 Dithiocarbamates 73, 75

Ecology of citrus grove habitat 249, 301 habitat/trophic range of mites 11, 28 knowledge deﬁciency 299 Economic injury level (EIL) 32, 35, 95 Endosymbiont bacteria see Bacteria Environment climate, inﬂuence on mite populations 10, 31, 63–64, 91, 294–295 contamination by acaricides 34, 301 effect of water supply on mite damage 28, 30, 249, 252 Eotetranychus 49 Eotetranychus cendanai (citrus yellow mite) 28, 51, 220–222, 221 Eotetranychus kankitus (citrus yellow mite) 31, 51, 222–224, 223 Eotetranychus lewisi (Lewis spider mite) 28, 51, 224–226, 225 transport/inspection regulations 303 Eotetranychus limonae 51, 226–227 Eotetranychus limoni 51, 227–229, 228 Eotetranychus mandensis 51, 229, 230 Eotetranychus pamelae 51, 229, 231, 232 Eotetranychus sexmaculatus (six-spotted spider mite) 51, 232–235, 233 habitat and feeding preferences 28, 29, 234 Eotetranychus yumensis (Yuma spider mite) 28, 51, 235–237, 236 Epidermis, host plant, mite damage to 29, 74–75, 93–94 Eradication, of ﬁrst outbreaks 32

374

Index Eriophyidae biological control 35 collection technique 21 diagnostic characters 41, 58–61 feeding mechanisms 26, 29 pest status and distribution 58, 59, 303 taxonomy 61 Eriophyinae, subfamily identiﬁcation 61 European and Mediterranean Plant Protection Organization (EPPO) 302–303 European red mite (Panonychus ulmi) 52, 255–257, 256 Eurytetranychini, tribe identiﬁcation 195 Eutetranychus 48 Eutetranychus africanus (African red mite) 51, 197, 199, 199–201 Eutetranychus banksi (Texas citrus mite) 31, 51, 201–204, 202 density of feeding punctures 27, 203 Eutetranychus citri 51, 204, 204–205 Eutetranychus cratis 50, 205–207, 206 Eutetranychus eliei 51, 207, 207–208 Eutetranychus orientalis (Oriental red mite, citrus brown mite, lowveld citrus mite) 51, 208–214, 209 chemical control issues 214, 299 as factor in host defoliation 30, 211 feeding preferences 28, 210, 211 natural enemies 211, 212–213, 214 transport/inspection regulations 303 Eutetranychus pantopus 51, 214–216, 215 Eutetranychus pyri 51, 216, 216–217

False spider mites see Tenuipalpidae Feeding consequences, for host plant 27–30 structures 15, 15–16, 25–27 Fertilizer application 302 Flat mites see Brevipalpus spp. Florida chemical control treatments 10 harmful mite species 31 Fruit (fresh), production 7–8, 10 Fungi citrus pathogens, combined with mite attack 149, 167 entomopathogenic 95–97, 253, 301

Galls, bud 122, 149 Gnathosoma (capitulum) 14–16, 15 Grape ﬂat mite (Brevipalpus chilensis) 46, 123–125, 124 Grapefruit (Citrus paradisi) 6, 8

Halo scab 143, 149 Hedges (windbreaks) 150, 302 Heinze’s PVA medium 23 Hoyer’s medium 23 Hystrichonychini, tribe identiﬁcation 183

Identiﬁcation internet information resources 26 key to citrus mite taxa 41–54 microscopic examination 21–23 Idiosoma 16 India, citrus yield factors 10 Insects, predatory 35, 97 Integrated pest management (IPM) alternate host plant control 150, 281, 295, 302 deﬁnition, requirements and challenges 36–37, 303–304 use of resistant varieties 150, 302 Introductions, accidental 32, 302–303

Japanese citrus mite see Aculops pelekassi Juice production chemical pest control 10 major markets and producers 8–9 sugar content 30, 74

Kanzawa spider mite (Tetranychus kanzawai) 53, 268, 269–271

Lactophenol (clearing/mounting medium) 22, 23 Legs 12, 16–17, 18 Lemon (Citrus limon) 5, 8 relationship with Aceria sheldoni (citrus bud mite) 299–300 Leprosis (viral disease, CiLV) 10, 30, 31–32, 148, 300 Leprosis mite see Brevipalpus phoenicis

Index

375 Lewis spider mite see Eotetranychus lewisi Life cycle developmental stages 11 reproductive behaviour 16, 270, 294 Lime (Citrus aurantifolia) 5–6, 8 Lowveld citrus mite see Eutetranychus orientalis

Maceration 22 Mandarin (Citrus nobilis) 6–7, 8 Market destinations essential oils 9 fresh fruit production 7–8, 10 processing for juice 8–9, 10 Mediterranean region chemical control treatments 10, 32 citrus varieties cultivated 5, 6, 7 Meyernychus emeticae (Angolan citrus mite) 49, 217–219, 218 Microscopic examination 22–23 Mixonychus 49 Mixonychus ganjuis 52, 237–238 Mixonychus ziolanensis 52, 238 Monitoring, of mite populations see Population sampling Morphology appendages 15–16, 16–17 body regions and features 11–12, 12 chaetotaxy 12, 117, 164 embryonic development 14, 14, 16 Mounts, of prepared specimens 22–23 Mouthparts see Chelicerae

Natural enemies management, for biological control 35, 37, 301, 303–304 types of organisms 35, 300–301 see also under individual mite species New brown citrus rust mite (Tegolophus brunneus) 45, 82–83, 83 Nothopodinae, subfamily identiﬁcation 68 Nothopodini, tribe identiﬁcation 68

Oligonychus 50 Oligonychus biharensis 54, 239, 240 Oligonychus coffeae (tea red spider mite, red tea mite) 53, 239–242, 241

infestation, effect of tea pruning 242, 302 Oligonychus gossypii 54, 242–244, 243 Oligonychus peruvianus 53, 244–246, 245 Orange (Citrus sinensis) 4–5, 8 Oriental red mite see Eutetranychus orientalis Ornamental ﬂat mite (Brevipalpus obovatus) 46, 140–143, 141 Ornamented mite (Tuckerella knorri) 48, 166, 166–168 Oudeman’s ﬂuid 21

Paciﬁc spider mite see Tetranychus paciﬁcus Palps 16, 19, 26–27 Panonychus 49 Panonychus citri (citrus red mite) 31, 52, 246–254, 247 damage assessment 32, 252–253 feeding preferences 28, 248 long-term effect of infestation on host growth 30, 249, 252 natural enemies 250–251 Panonychus elongatus 52, 254–255, 255 Panonychus ulmi (European red mite) 52, 255–257, 256 Paratetra murrayae 44, 78–79 Peacock spider mite (Tuckerella pavoniformis) 48, 165, 166, 170–171 Pentamerismus tauricus 46, 154–155 Peritremes 13, 14 Permanent mounts 22–23 Pesticide resistance 34, 150–151 Petrobia 48 Petrobia harti 50, 188–190, 189 Petrobia latens (brown wheat mite) 50, 190–192, 191 Petrobia tunisiae 50, 192–194, 193 Petrobiini, tribe identiﬁcation 188 Petroleum oils 66, 98 Phyllocoptinae, subfamily identiﬁcation 70 Phyllocoptini, tribe identiﬁcation 87 Phyllocoptruta 43 Phyllocoptruta citri 44, 87–89, 88 Phyllocoptruta oleivora (citrus rust mite) 44, 81, 89–98, 90, 110

376

Index Phyllocoptruta oleivora continued cell and biochemical damage to fruit 29, 94 chemical control 33, 34, 82, 97–98 comparison with Aculops pelekassi 72, 73 damage and symptoms of leaf attack 30, 93–94 natural enemies 92, 93 pest status and climate 31, 91 population monitoring 32, 95–96 Phyllocoptruta paracitri 44, 98–100, 99 Phytoptidae, characteristics, status and taxonomy 41, 55, 59 Phytoptinae, subfamily identiﬁcation 56 Phytoptus ﬁcivorus 42, 56–57 Phytoseiidae 34, 35, 300 Pink citrus rust mite see Aculops pelekassi Plant protection measures see Crop protection Polyphagotarsonemus latus (citrus silver mite, broad/tropical/white mite, yellow tea mite) 45, 108–112 attack on young fruit 28, 109–110 female 105, 108 male 106, 108 symptoms, compared with Aculops pelekassi 74–75 Pomelo (Citrus grandis) 6, 8 Population sampling 20, 32, 33, 95–96, 252–253 Predatory mites 34, 35, 300 Preservation, of mite specimens 21–22 Privet mite (Brevipalpus obovatus) 46, 140–143, 141 Prostigmata (order) 18–19 Pruning, effect on mite populations 242, 302 Pseudotarsonemoidinae, subfamily identiﬁcation 107 Pseudotarsonemoidini, tribe identiﬁcation 107

Quarantine legislation 150, 303

Rearing 24 Red and black ﬂat mite see Brevipalpus phoenicis Red crevice mite see Brevipalpus phoenicis

Red-legged spider mite see Tetranychus ludeni Red spider mite see Tetranychus ludeni Red tea mite see Oligonychus coffeae Reddish black ﬂat mite see Brevipalpus phoenicis Reproduction 16, 270, 294 Resin-based media 23 Respiratory structures 13, 14 Russeting 27–28, 29, 30, 94, 110 Rust mites see Aculops pelekassi; Phyllocoptruta oleivora; Tegolophus spp.

Sampling protocols see Population sampling Scab mite (Brevipalpus lewisi) 47, 136, 136–139 Scarlet mite see Brevipalpus phoenicis Schizotetranychus 49 Schizotetranychus baltazari (citrus green mite) 52, 257–259, 258 Schizotetranychus hindustanicus 52, 259–260, 260 Schizotetranychus lechrius 52, 260–261 Schizotetranychus spiculus 52, 261, 262 Schizotetranychus youngi 52, 263 Sensory organs 13, 14, 16 Setae 12, 13, 14, 16 see also Chaetotaxy Six-spotted spider mite see Eotetranychus sexmaculatus Spider mites see Tetranychidae Strawberry spider mite (Tetranychus turkestani) 53, 268, 291–293, 294 Stylophores 15, 25–26 Sulfur compounds 75, 87, 98, 111–112 Symptoms bud galls 122, 149 fruit distortion and drop 27, 65 growth/yield reduction 30, 74, 252 halo scab 143, 149 internet information resources 26 leaf and shoot damage 27, 30, 86, 94, 211 russeting 27–28, 29, 30, 94, 110 webbing 27, 113, 172, 226, 294 see also Damage Systematics citrus mite identiﬁcation key 41–54

Index

377 comparison of mites and insects 11 subdivisions of Acari 17–19, 298 taxonomic position of Acari 11, 17

Tangerine 6–7, 8 Tarsonemidae diagnostic characters 42, 104–107 feeding mechanisms 27 pest status and distribution 59, 104 taxonomy 107 Tea red spider mite see Oligonychus coffeae Tegolophus 45 Tegolophus australis (brown citrus rust mite) 45, 79–82, 80 Tegolophus brunneus (new brown citrus rust mite) 45, 82–83, 83 Temporary mounts 22 Tenuipalpidae (false spider mites) diagnostic characters 42, 113–117 pest status and distribution 113, 114 taxonomy 117, 298 Tenuipalpinae, subfamily identiﬁcation 155 Tenuipalpoidini, tribe identiﬁcation 194 Tenuipalponychus citri 49, 194–195 Tenuipalpus 45 Tenuipalpus caudatus 45, 155–156 Tenuipalpus emeticae 46, 156–158, 157 Tenuipalpus mustus 46, 158 Tenuipalpus orilloi 46, 159, 160 Tenuipalpus sanblasensis 46, 157, 159–161 Tetranychidae (spider mites) biological control 35 diagnostic characters 23, 42, 172–173, 176–177 feeding mechanisms 25–26, 27 pest status and distribution 172, 173, 174–175, 303 taxonomy 177 web (silk) production 27, 172, 226, 233, 294 Tetranychinae, subfamily identiﬁcation 194 Tetranychini, tribe identiﬁcation 219 Tetranychus 50 Tetranychus desertorum (desert spider mite) 53, 263–265, 264 Tetranychus ﬁjiensis 52, 265–268, 266

Tetranychus gloveri 53, 268, 268–269 Tetranychus kanzawai (Kanzawa spider mite, cassava red spider mite) 53, 268, 269–271 Tetranychus lambi 53, 271–273, 272 Tetranychus ludeni (red/red-legged spider mite) 53, 273–276, 274 natural enemies 277 Tetranychus mexicanus 53, 276–279, 278 Tetranychus neocaledonicus (vegetable mite) 53, 279–283, 280 natural enemies 282–283 Tetranychus paciﬁcus (Paciﬁc spider mite) 53, 283–286, 284 natural enemies 286 Tetranychus paraguayensis 53, 268, 287 Tetranychus salasi 53, 287–288 Tetranychus taiwanicus 52, 288–289 Tetranychus tumidus (tumid spider mite) 53, 289, 289–291 Tetranychus turkestani (strawberry spider mite) 53, 268, 291–293, 294 Tetranychus urticae (two-spotted spider mite) 53, 268, 293–297 feeding preferences 28, 29, 294, 295 increase, following carbamate treatment 34 natural enemies 296 Tetronic acid derivatives 76, 98, 301 Texas citrus mite see Eutetranychus banksi Treatment action thresholds 32, 95–96, 252–253 Tropical mite see Polyphagotarsonemus latus Tuckerella 47 Tuckerella knorri (ornamented mite) 48, 166, 166–168 Tuckerella nilotica 48, 165, 168, 168–169 Tuckerella ornata 48, 164, 168, 169–170 Tuckerella pavoniformis (peacock spider mite) 48, 165, 166, 170–171 Tuckerellidae diagnostic characters 42, 163–165 pest status and distribution 114, 163 taxonomy 165 Tumid spider mite (Tetranychus tumidus) 53, 289, 289–291 Two-spotted spider mite see Tetranychus urticae

378

Index Ultratenuipalpus gonianensis 45, 161–162, 162

Vegetable mite see Tetranychus neocaledonicus Virus transmission 30, 148

Weed control, in citrus crops 150, 281, 295, 302 White mite see Polyphagotarsonemus latus

Whiteﬂy, as mite dispersal agents 109

Yellow tea mite see Polyphagotarsonemus latus Yield geographical variation 7 reduction due to mite attack 30, 74, 252 Yuma spider mite (Eotetranychus yumensis) 28, 51, 235–237, 236