ATLAS OF OF THE WORLD (: )

Vikram Prasad, M.D., Ph.D.

2013

INDIRA PUBLISHING HOUSE Publisher of books in agricultural, biological, medical and veterinary sciences P.O. Box 250456, West Bloomfield, MI 48325-0456, USA E-mail: [email protected]; www.indirapublishinghouse.com 2

All rights reserved. Except for brief quotation in a review or reference, this book or parts thereof must not be reproduced in any form without permission in writing from the publisher.

First printing: March 2013 Published by: Indira Publishing House P.O. Box 250456, West Bloomfield, MI 48325-0456, USA Phone: (248) 661-2529 E-mail: [email protected] Website: www.indirapublishinghouse.com

Printed in the USA ISBN: 0-930337-31-X Library of Congress Control Number: 2013902148 3

DEDICATION

THIS BOOK IS LOVINGLY DEDICATED

For their constant support, to:

My grandparents Mr. Shiva Jatan Shah and Mrs. Manwa Devi; my parents Mr. Udai Narain Ram and Mrs. Yashoda Devi; my wife Indira Prasad; my daughter Nilima Prasad Chand, her husband Sunil Chand, and their daughter Sunita Chand; my son Ravi Vikram Prasad, his wife Savita Prasad, and their daughters Kavita Prasad and Anita Prasad; and my son Ravindra Vikram Prasad and his wife Mary Dimercurio Prasad.

A special dedication also to:

Dr. Harold A. Denmark, Gainesville, Florida (USA) and Dr. James A. McMurtry, Sunriver, Oregon (USA) for their enormous help to me with this book. 4

ABSTRACT

The family Phytoseiidae (Acari: Mesostigmata) is comprised of 2,800 species known to occur throughout the world with the exception of Antarctica. They are primarily predatory , many of which play an important role as biological control agents of many plant feeding mites and tiny insects. Several species of these are being mass cultured and utilized globally for this purpose by the biological control industries. Many new species are described and already known species recollected and redescribed every year from all over the world. Research work on different aspects of the Phytoseiidae is expected to increase several folds over next few decades. Thus, correct identification of these important mites will be essential. Atlases, checklists and catalogs are some important taxonomic reference books required to aid in such identification works.

This atlas follows the classification system of Chant and McMurtry (2007) and includes over 1,000 figures of different species of Phytoseiidae. As the concept of some of the genera keeps changing, for the sake of convenience of the readers, these species are arranged in alphabetical order from A to Z showing important morphological structures and current system of setal nomenclature. Errors or problems in published figures of species are pointed out. It was noted that almost 8-10% published figures had errors or problems of one kind or another which are discussed for 192 species in this book. The species reported as synonyms in literature, even if controversial, are included as they were identified originally since some may or may not be synonyms. This book complements other books, “Checklist of Phytoseiidae of the World” (Prasad, 2012) and “Phytoseiidae and Otopheidomenidae (Acari: Mesostigmata) of the World: A Self Study Guide” (Prasad, 2011). Other recent books on Phytoseiidae as “Illustrated Keys and Diagnoses for the Genera and Subgenera of the Phytoseiidae of the World (Acari: Mesostigmata)” (Chant and McMurtry, 2007), “Phytoseiidae of Greece and Cyprus (Acari: Mesostigmata)” (Papadoulis, Emmanouel and Kapaxidi, 2009), and “Phytoseiidae of North America and Hawaii (Acari: Mesostigmata)” (Denmark and Evans, 2011) are invaluable books for anyone wishing to pursue research on taxonomy of phytoseiid mites and should be included among the reference books. 5

CONTENTS

Title - 1 Publisher’s data - 2 Dedication - 3 Abstract - 4 Contents - 5 Introduction - 7 - Some important papers on Phytoseiidae and closely related families - 8 - Idiosomal chaetotaxy - 9 - Relationships of Phytoseiidae with nearest relatives in different life stages - 14 - Difficulties in identification of some dorsal idiosomal setae - 16 - Difficulties in identification of some ventral idiosomal setae - 19 - Other structures and characteristics - 20 - Problems with some published figures - 22 - Why so many genera today - 66 - New species based on pores, solenostomes, number of denticles, type of spermathecae and spermatodactyls - 70 - Our handicaps in Phytoseiidae - 71 - Variation in species - 71 - Species synonyms - 71 - Synonymy problems - 72 - Misidentification of species - 74 - Old versus new setal nomenclature of idiosoma - 75 - Where are the large collections of mites and reprints of Donald Chant - 75 - How to improve problems in publications of Phytoseiidae: Some recommendations - 75 - How to use this book - 80 - Confession - 81 Classification of Phytoseiidae - 82 Current genera of Phytoseiidae - 85 Controversial genera of Phytoseiidae - 87 New genera of Phytoseiidae - 88 Atlas (species, beginning with):

A - 89 B - 177 C - 229 D - 329 E - 370 F - 410 G - 446 H - 476 I - 522 J - 554 K - 570 L - 603 M- 662 N - 762 O - 817 P - 842 Q - 937 R - 943 S - 985 T - 1076 6

U - 1131 V - 1140 W- 1160 Y - 1177 Z - 1185

Species list of figures - 1188 Acknowledgements - 1220 References - 1221 Index - 1293

7

INTRODUCTION Species of the family Phytoseiidae are considered to have evolved from Gamasina or Mesostigmata with holotrichous setation with a single postanal seta and 66 pairs of dorsal and ventral idiosomal setae (Lindquist and Evans, 1965) which have lost many of these setae during different evolutionary adaptations to various environmental conditions. Of these 66 pairs, 23 pairs of setae have been suppressed and 43 pairs are known to occur in Phytoseiidae but no known phytoseiid species has this full complement (Chant, 1993c; Chant and McMurtry, 1994, 2007). Of these 43 pairs of idiosomal setae, only 22 pairs (dorsal = 12 pairs and ventral = 10 pairs) occur as fixed setae and 21 pairs (dorsal = 16 pairs and ventral = 5 pairs) occur as variable setae on all phytoseiids (Chant and McMurtry, 2007, Figs. 1, 2). The most setose phytoseiid species that is known is Australiseiulus angophorae (Schicha), 1981a (subfamily Typhlodrominae), with 38 pairs of dorsal and ventral setae on the female idiosoma, and the most hypotrichous species is Afroseiulus robertsi (Baker), 1990 (subfamily ), with 25 pairs of dorsal and ventral setae on the female idiosoma.

The evolution in the Phytoseiidae has been marked by simplification of many characteristics and suppression of many setae on the idiosoma and legs, but also by modification to the tectum and corniculi, and reduction in the hypostomal dentition and idiosomal sclerotization. This setal suppression has resulted in extensive paedomorphosis (retention of juvenile or larval traits into later life including the adult) leading to loss or suppression of adult ancestral setal forms and adapting to more hypotrichous phytoseiids now that resemble the setal complement of immature holotrichous blattisociines (Chant, 1993c). This suppression of idiosomal setae appears to have been discontinuous over a period of time (Chant, 1993a).

Mites of the family Phytoseiidae (Acari: Mesostigmata) are predators of other mites and certain kinds of insects and nematodes. They are well known as biological control agents of plant feeding mites and some insects. Several species, for this reason, are being commercially utilized for this purpose in several countries. Among these, the well-known predators are: Amblydromalus limonicus (Garman and McGregor), 1956 (Amblyseiinae), andersoni (Chant), 1957c (Amblyseiinae), Amblyseius swirskii Athias-Henriot, 1962 (Amblyseiinae), occidentalis (Nesbitt), 1951 (Typhlodrominae), degenerans (Berlese), 1889 (Amblyseiinae), californicus (McGregor), 1954 (Amblyseiinae), (Oudemans), 1930b (Amblyseiinae), Neoseiulus fallacis (Garman), 1948 (Amblyseiinae), longipes Evans, 1958c (Amblyseiinae), Phytoseiulus persimilis Athias-Henriot, 1957a (Amblyseiinae) and Transeius montdorensis (Schicha), 1957b (Amblyseiinae).

At present, about 2800 phytoseiid species are known in the world. New species, especially from tropical and neotropical countries, are being described at a rate of about 40 per year (Prasad, 2012). It is expected that hundreds of new species will be discovered over the next several decades. It is also expected that several phytoseiid species, other than those listed above, would be utilized in the biological control. Thus, knowledge of these mites, taxonomic or in biological control and applied research, would be expanding gradually requiring knowledge of trained acarologists in various lines of research, and for correct identification of these mites.

The researchers of the plant and soil associated mites require various tools for the correct identification of the phytoseiid mites. For example, descriptions, figures, keys, and information on the distribution, hosts, habitat, etc. Some of this information may be available in comprehensive taxonomic publications such as Atlases, Bibliographies, Catalogs, Checklists, Monographs, etc. Each of these provides only certain specific information and does not replace the importance of the other. Thus, various reference books as tools are necessary to achieve the goal of identification and writing about these mites.

Other than bibliographical works on Phytoseiidae by Kostiainen and Hoy (1996), most comprehensive works on these mites over the last decade have been by Moraes, McMurtry, Denmark and Campos (2004); Chant and McMurtry (2007); Papadoulis, Emmanouel and Kapaxidi (2009); Denmark and Evans (2011) and Prasad (2012). Of these, only the “Catalog” by Moraes, McMurtry, Denmark and Campos (2004) and “Checklist” by Prasad (2012) include all phytoseiid species known in the world but do not include figures. Atlases on insects and mites, by tradition, are important reference books that include illustrations of published species which are very useful not only in the correct identification of species but also to compare the closely related species which may turn out to be synonym or misidentification. As such works require years of dedicated work, except for “Atlas of the Swedish water mites” by Lundblad (1962) and “Illustrations of the mites and of Japan” edited by Shôzô Ehara (1980), no Atlas of mites, especially on Phytoseiidae of the world has been published. 96

G F ST1 B ST2 JV 1 L - Shaped JV2 ZV2 ST3 ST4 J V 4 E ST5

ZV1 PE > j1 JV1 A MP2 j1 j3 JV2 ZV2 r3 j 4 z2 JV3 z4 s4 z5 JV4 R1 j6 Z1

StIV S2 J2 StiIV

S4 C SgeIV

S5 Vesicular Z4

Z5 J5 D

acaridophagus, (Collyer), 1964b – Acarologia 38 (3), 1997, p 225 (after Yoshida-Shaul and Chant, 1997) [A-D, female: A. dorsal shield, B. ventral shields, C. spermatheca, D. leg IV, E. peritremal shield; F-G, male: F. spermatodactyl, G. ventrianal shield] - AMBLYSEIINAE. 97

PE j1 A j1 B ST1 j3 D ST2 z2 Vesicular j 4 ST3 z4 r3 j5 ST4 z5 s4

j6 ST5 MP1 PD + C R1 ZV1 FDCH Z1 = 3 JV1 MDCH J2 MP2 = 2 ZV2 ZV3 S2 JV2 JV4

Z4 S4

JV5

S5 Z5 J5 G

E Beaked F JV1

ZV2 SgeIV JV2

JV5

StIV

accessus, Neoseiulus (Ueckermann and Loots), 1988 – IJA 32 (3), 2006, p 243 (after Zannou, Moraes, Ueckermann, Oliveira, Yaninek and Hanna, 2006) [A-E, female: A. dorsal shield, B. ventral shields, C. chelic- era, D. spermatheca, E. leg IV; F-G, male: F. spermatodactyl, G. ventrianal shield] - AMBLYSEIINAE. 112

A

PE j1

j1 j3 D z2 z3 j 4 z4 r3 j5 s4 z5

j6 s6 R1 StIV S2 J2

S4 Z4 B ST1

ST2 S5 Z5 J5 ST3 ST4? C MP1 ST5

ZV1

MP2 JV1 E PD + ZV2 Fundibular JV2 ZV3 MDCH FDCH = 2 JV3 = 5-6 JV4

JV5

alba, Amblydromella Denmark and Evans, 2011 – Denmark and Ev- ans (2011), p 311 (after Denmark and Evans, 2011) [female: A. dorsum, B. ventral shields, C. spermatheca, D. peritremal shield, E. chelicera] - TYPHLODROMINAE. 238

j1 A PE j1 B z2 ST1 j3 z3 j 4 z4 ST2 s4 r3 j5 z5 ST3 j6 s6

ST4

ST5 MP2 JV1 ZV1 ZV2 JV2 ZV3

Z4 JV5 Z5 J5

E PD + D C

MDCH Fundibular FDCH = 1 = 3 SgeIV

StiIV StIV

canadensis, Phytoseius Chant, 1965b – Papadoulis, Emmanouel and Kapaxidi (2009), p 107 (after Papadoulis, Emmanouel and Kapaxidi, 2009) [female: A. dorsal shield, B. ventral shields, C. chelicera, D. spermathecae, E. leg IV] - PHYTOSEIINAE. 1319

Z

{zamoranus}, {Swirskiseius} Denmark and Evans – 66, 1185

{zuluagai}, {} Denmark and Muma – 66, 1186

{zwoelferi}, {Neoseiulus} (Dosse) – 1187

***** 1320

BOOKS OF THE AUTHOR

Atlas of Phytoseiidae of the World (Acari: Mesostigmata), in color, by V. Prasad, 2013, 1319 pp, (hard cover).

Checklist of Phytoseiidae of the World (Acari: Mesostigmata), by V. Prasad, 2012, 1063 pp, (hard cover).

Phytoseiidae and Otopheidomenidae (Acari: Mesostigmata) of the World: A Self Study guide, in color, by V. Prasad, 2011, 208 pp, (hard cover).

Memories of Acarology in Color (Part 1. Honoring Acarologists), in color, by V. Prasad, 2003, 200 pp, (hard cover).

Mites: A Bibliography (1926-1966), by V. Prasad, 1988, 230 pp, (hard cover).

A Decade of Acarology, by V. Prasad, 1987, 183 pp, (hard cover).

History of Acarology, V. Prasad (Ed.), 1982, 472 pp, (hard cover).

The Taxonomy of Water Mite Larvae, by V. Prasad and D. R. Cook, 1972, 326 pp, (hard cover).

A Catalogue of Mites of India, by V. Prasad, 1974, 320 pp, (hard cover).

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