Preface Among the Acari eriophyoid mites- the Eriophyoidea or Tetrapodili- are second only" to the spider mites (Tetranychidae) in their economic importance as plant pests throughout the world. They surpass all other groups of phy- tophagous mites in the extent of their morphological and biological special- ization for obligate phytophagy. Moreover, they surpass these other groups in the degree to which they are specialized on their host plants, making them superior in their potential as weed control agents. Despite the reductive structural simplicity evident among even the most primitive extant members of the Eriophyoidea, the more derived subgroups have augmented their body plan secondarily in a variety of ways, either for !iving in closely confined spaces like sheaths, buds, erinea and galls or moreso for living, freely exposed, on plant surfaces. Behavioral and life cycle modifi- cations correlated with these structural changes reflect the adaptation and evolution of this lineage into a disparity of highly host-specific forms that, to date, have bewildered any meaningful classification of them. At the time of the last compilation of world knowledge about eriophyoid mites in the book Mites Injurious ot Economic Plants by Jeppson, Keifer and Baker (1975), some 1800 species in 511 genera were known. During only the 20 years since then, approximately 1000 more species and nearly 511 more genera have been described, giving testimony to how poorly known and taxon-rich this group is. Yet, major regions of the world remain virtually untouched in surveying for these mites, such that Amrine and Stasny (1994), in their new comprehensive Catalog of the Eriophyoidea of the World, estimated that not more than 5 percent of the world species of Eriophyoidea have been described! Just as the number of described taxa of Eriophyoidea has doubled during the last two decades, so has our knowledge of the biology, ecology and importance of these mites expanded. At the same time, the actual and potential economic importance of eriophyoids continues to grow worldwide, and their success in colonizing new regions makes them an ongoing quarantine threat in many parts of the world (a new case in point, as this is written, is the note in Florida Entomologist by Pefia and Denmark (1996) on the recently confirmed presence in Florida of Tegolophus perseaflorae Keifer, a neotropical pest of avocado). Thus, this book is timely in compiling and synthesizing information that is now available on these behaviorally fascinating, economically important mites. We realize that such a book, containing updated knowledge on nearly 3000 species, will perforce be incomplete and overly generalized in some areas. However, we have asked the best specialists available concerned with the bi- ology, ecology and control of eriophyoid mites, as well as some generalists in iv Preface acarology, to join us in contributing to the compilation of this book. The book gives much attention to fundamental aspects of eriophyoid anatomy, behav- ior, ecology and even systematics, as bases for understanding the ways of life of these mites and their effects on host plants; in turn, this will lead to develop- ing the most appropriate means of regulating these mites as detrimental or beneficial organisms. Previous general accounts of eriophyoid mites have been primarily from the perspectives of single authors - notably Nalepa during the first third, and Keifer during the middle third, of this century- followed by more regional perspectives from subsequent specialists as, for example, Boczek in Poland, Shevchenko in Russia, Mohanasundaram in India, Manson in New Zealand, and Smith Meyer in South Africa. The present book is unique in being not only the first compilation of knowledge on Eriophyoidea by a multiplicity of au- thors (47, including some of those just named), but also in the international as- pect of its contributors (from 41 countries) and in many of them being hands-on specialists in the biology, behavior and economic importance of these mites. The book is also unique in its perspective of treating eriophyoid mites as a lin- eage - no matter how specialized - of acariform mites, such that standard terms and notation for structures common to other such mites are applied to them as well. i i i The disparity of interests and also linguistic backgrounds among authors has led to quite different, often refreshing, approaches to the subject of their chapters, such that uniformity in content and presentation has not been possi- ble or even encouraged by the editors. Considerable latitude has also been given to authors on the subject matter presented in sections that are of a paral- lel nature, e.g., eriophyoid pests of citrus, of apple and pear, etc. With an eye to the future, however, we have strongly encouraged authors to consider the needs of further research in the conclusions of each of their sections. Such a multi-authored book will have some unavoidable overlap of content, and even some discord, in various sections. On balance, we view this as advantageous in cross-referring and stimulating readers to other sections of the book. Our book is generally organized in four parts. Part I deals with aspects of eriophyoid mites themselves, including: external anatomy, systematics (including the first illustrated key limited to genera with species of economic importance), and nomenclatural problems (Chapter 1.1); internal anatomy and physiology (Chapter 1.2); morphogenesis and cytogenetics (Chapter 1.3); biol- ogy, ecology and general accounts of eriophyoids associated with primitive vascular plants (Chapter 1.4); evolution and phylogeny (Chapter 1.5); and field and laboratory techniques for their scientific study (Chapter 1.6). Part II treats the natural enemies of eriophyoid mites, including: predatory phytoseiid mites, potentially the most effective biological control agents of phytophagous mites (Chapter 2.1); predatory stigmaeid mites, long in need of ecaferP vii greater scrutiny as auxiliary biological control agents (Chapter 2.2); and other predatory arthropods (Chapter 2.3) and pathogens (Chapter 2.4). Part III begins with an account of the nature of damage by eriophyoids and its assessment (Chapter 3.1), followed by a series of 14 sections that treat erio- phyoid pest problems and their control in major world agro-ecosystems (Chap- ter 3.2). This part continues with presentations on host plant resistance (Chap- ter 3.3), pesticide resistance in eriophyoids and their associates (Chapter 3.4), and an extensive review of chemical control (Chapter 3.5). Part IV deals with eriophyoid mites as beneficial organisms, and includes accounts of various species in the biological control of weeds (Chapter 4.1). The effects and potential impact of the presence of eriophyoid mites as competitors of other phytophagous mites and as alternative prey for the natural enemies of other phytophagous mites are also considered (Chapter 4.2). We are grateful to the contributing authors, not only for their individualis- tic experience and knowledge as put forward in their presentations, but also for valuable input by some of them as reviewers for various sections. Permission to reuse Fig. 1.1.2.50 was given by DSIR Plant Protection, Auckland, New Zea- land; figures used with permission from other sources are acknowledged in ap- propriate captions. Special thanks go to Barry Flahey (Agriculture & Agri- Food Canada, Ottawa) for timely artistic support in Chapters 1.1.1 and 1.5.1, to Alice Boerrigter and Hans Bolland for their enormous support in creating a reference collection of literature on eriophyoid mites, to Simon van Mechelen for producing hundreds of glossy prints, and to Lia Out who was instrumental in constructing the indices and in giving the book its final touch. We hope that this book meets the needs for an up-to-date compilation of the basic and applied knowledge on eriophyoid mites and their control that is otherwise scattered in a variety of languages and literature throughout the world. In doing so, it also presents new views intended to stimulate interest in eriophyoids and their enemies, and it points to areas where further research is needed. The contents are intended for students, teachers, researchers, extension workers and other clients in the areas of acarology and plant protection. They are also intended for readers having broader interests in ecology and evolu- tionary biology who may find eriophyoids to be rewarding experimental ani- mals for formulating and testing biological concepts that may provide new in- sights about general biological phenomena. We further hope that the book stimulates readers to critically test the views presented and aimed ultimately toward environmentally safe, sustainable and economically efficient means of regulating detrimental and beneficial eriophyoid mites. Evert E. Lindquist Maurice W. Sabelis Jan Bruin ShdnPargotohP no front cover era yb courtesy fo .E.W Frost and .M.P Ridland ,)tfel( .N.G dleifdlO )elddim( .E.W Styer .)thgir( ehT first photo ni this ecaferp si yb ysetruoc fo .M.C.D .nosnaM VIII ... Scale This page: Acaricalus eac~poxeli on Ilex acapo leaf (photo by W.E. Styer). Opposite page, top: Abacarus hystrix on bal, point pen; middle: Parasitus sp. (Mesostigmata: Parasitidae) plus three specimens of Abacarus hystrix (asterisks) on perennial ryegrass; bottom: Aceria sp. in leaf grooves of wheat (photos by W.E. Frost and P.M. Ridland). r,~ Life forms .... ,:: ,i, 48(cid:1)~ This page: Acalitus aeniregaf in erineum on sugaF ailofidnarg (photo by W.E. Styer). Oppo- site page, top: suracabA hystrix on perennial ryegrass (photo by W.E. Frost and P.M. -di~T land); middle: wax-secreting setporemirT ;sfmrofiaoryela bottom: Cymeda acidnalaez (pho- tos by D.C.M. Manson). % ~x egamaD (cid:12)9 ~ ~, ..i,, " 3.)," Top: coconuts (Photo by D. Moore and F.W. Howard); middle: tulip bulbs (photo supplied by C.G.M. Conijn); bottom: apples (photo by M.A. .)koorbretsaE egamaD ~176176 XIII Top: wheat grain with Aceria allehcisot (Photo by W.E. Frost and P.M. Ridland); middle: lucerne (photo by P.M. Ridland); bottom: pear leaves with blister galls (photo by M.A. Easterbrook). XV Contributors to this Volume .G ALBERTI Zoologisches Institut und Museum, Universit/it Greifswald, Johann-Sebastian- Bachstr. 11/12, D-17489 Greifswald, Germany J.W. AMRINE, Jr. Division of Plant and Soil Sciences, West Virginia University, P.O.Box 6108, Morgantown, WV 26506-6108, USA .J BOCZEK Department of Applied Entomology, Warsaw Agricultural University, 02-766 Warszawa, ul. Nowoursynowska 166, Poland .R BRONNER Laboratoire de C6cidologie, Institut de Botanique, Universit6 Louis Pasteur, 82 rue Goethe, 67083 Strasbourg Cedex, France .J BRUIN Section Population Biology, Institute of Systematics and Population Biology, University of Amsterdam, Kruislaan 320, 1098 MS Amsterdam, The Netherlands M. CASTAGNOLI Istituto Sperimentale per la Zoologia Agraria, Via Lanciola, Cascine del Riccio, 50125 Firenze, Italy G.P. CHANNABASAVANNA Department of Entomology, University of Agricultural Sciences, Rajajinagar, Bangalore 065 010, India C.C. CHILDERS Citrus Research and Education Center, University of Florida, 007 Experiment Station Road, Lake Alfred, FL 33850-2299, USA D.R. CLEMENTS Department of Biology, Trinity Western University, 7600 Glover Road, Langley, British Columbia V3A 6H4, Canada C.G.M. CONIJN Bulb Research Centre, Vennenstraat ,22 2160 BA Lisse, The Netherlands