SCANNING ELECTRON MICROSCOPIC STUDIES ON SEVERAL OTOPHEIDOMENID (: )

VIKRAM PRASAD M.D., Ph.D.

2019

INDIRA PUBLISHING HOUSE Publisher of books in agricultural and biological Sciences P.O. Box 250456, West Bloomfi eld, MI 48325-0456, USA email: [email protected]; website: www.indirapublishinghouse.com 2

Vikram Prasad

All rights reserved by Indira Publishing House. 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 2019 Published by: Indira Publishing House PO Box 250456 West Bloomfi eld, MI 48325-0456, USA email: [email protected] website: www.indirapublishinghouse.com

Printed in the USA ISBN: 0-930337-34-4 Library of Congress Control Number: 2018915133

[Front cover photo: Atrophied fi xed digits and the movable digits surrounded by the spermatodac- tyls in the male of Prasadoseius cocytes (Prasad, 1970a); Fig. M10, page 133]. 3

3. DEDICATION Respectfully, this book is dedicated to: (1) Deceased Dr. Asher E. Treat who contributed so much information on the moth mites; (2) Dr. Glenn Walker, Professor and Scanning Electron Microscopist, Biol- ogy Department, Eastern Michigan University, Ypsilanti, Michigan, who helped in this study by taking the photographs on which this study is based; and (3) Mr. George Vieira, Fowlerville, Michigan, a young man, who patiently taught this author about InDesign, Photoshop, formatting this book, and helped in many technical problems. 4

4. ABSTRACT The results of the scanning electron microscopic (SEM) studies on the idiosoma, gnathosoma, and the legs of Prasadiseius cocytes (Prasad, 1970a), Prasadiseius donahuei (Prasad, 1970a), Prasadiseius incanus Prasad and Guanilo, in Prasad et al., 2011, and Prasadiseius kayosiekeri (Prasad, 1970b) (Acari: Otopheidomenidae: Otopheidomeninae), never published before, are presented giving numerous micro- graphs. All of these species are native to Central and South America and are obligatory ectoparasites of the hawk moths (Lepidoptera: Sphingidae). The remaining 4 species in the genus - P. achlora (Prasad, 1972), P. aporodes (Prasad, 1972), P. indicus (Prasad, 1973), and P. pholusis (Prasad, 1970b) are described from these and other tropical regions of the world. Species of the family Otopheidomenidae are most similar to those of the family Phytoseiidae but diff er in having the fi xed digit of the chelicera very much reduced, atrophied and stump-like in appearance, and generally fewer idiosomal setae. The mites of the subfamily Otopheidomeninae infest moths of the families Noctuidae and Sphingidae. They possess 11-16 pairs of minute dorsal idiosomal setae. Females have a long, tubular sperm access system of the laelapid type. The SEM sudy indicated the atrophied fi xed digit of female being diff erent than in the male. The present status of the family Otopheidomenidae is discussed. A key is included to identify the 12 species of Otopheido- meninae belonging to 2 subfamiles (Otopheidomeninae and Prasadoseiinae new subfamily). Another key is provided for the identifi cation of 7 families of Phytoseioidea (Blattisociidae, Hemipteroseiidae new family, Katydiseiidae Fain and Lukoschus, new status, Otopheidomenidae, Phytoseiidae, Podocinidae, and Treatiidae Wainstein, new status) as accepted in this work. Eickwortius termes Zhang and Orthoptero- seius sinicus Mo are transferred to Blattisociidae. Genera Nabiseius having 4 species and Treatia (= Ento- moseius) also having 4 species are included in the family Treatiidae. Twelve tables, several hundred SEM fi gures and line drawings of adults and, immature stages of Otopheidomeninae species including, some key morphological characters, and chaetotaxy of genua and tibiae I-IV, are provided.

Key words - Blattisociidae, chelicera, Hemipteroseiidae new family, Katydiseiidae Fain and Lukoschus, new status, , mites, Otopheidomenidae, Otopheidomeninae, parasites, Phytosei- idae, Podocinidae, Prasadseiinae new subfamily, Treatiidae Wainstein, new status. 5 5. CONTENTS 1. Title - 1 2. Publisher’s data - 2 3. Dedication - 3 4. Abstract - 4 5. Contents - 5 6. Introduction - 7 7. Material and methods - 12 8. - 14 (A) Genus Prasadiseius Wainstein, 1972 - 14 1. Prasadiseius cocytes (Prasad, 1970a) - 15 2. Prasadiseius donahuei (Prasad, 1970a) - 15 3. Prasadiseius incanus Prasad and Guanilo, 2011 - 16 4. Prasadiseius kayosiekeri (Prasad, 1970b) - 16 (B) Key for the identifi cation of 4 species of Prasadiseius - 17 9. Results - 18 1. Prasadiseius cocytes (Prasad, 1970a) - 18 (A) Male # 1 (dorsal) - 18 (a) Idiosoma (dorsal) - 18 (b) Gnathosoma (dorsal) - 18 (c) Legs I-IV, left and right (dorsal) - 19 (B) Female # 2 (ventral) - 25 (a) Idiosoma (ventral) - 25 (b) Gnathosoma (ventral) - 25 (c) Legs I-IV, right (ventral) - 26 2. Prasadiseius donahuei (Prasad, 1970a) - 29 (A) Females (dorsal) - 29 (a) Idiosoma (dorsal) - 29 (b) Gnathosoma (dorsal) - 30 (c) Legs I, II, and IV, right (dorsal) - 31 (B) Females (ventral) - 32 (a) Idiosoma (ventral) - 32 (b) Gnathosoma (ventral) - 32 (C) Males # 1 and # 2 (dorsal and ventral) - 32 (a) Idiosoma (ventral) - 32 (b) Gnathosoma (dorsal and ventral) - 33 (c) Leg II, Femur (ventral) - 33 3. Prasadiseius incanus Prasad and Guanilo, 2011 - 33 (A) Females n-female, females # 1, 2, 3 (dorsal) - 33, 38, 42, 45 (a) Idiosoma (dorsal) - 34, 38 (b) Gnathosoma (dorsal) - 34, 39, 42, 45 (c) Legs I-IV, left and right (dorsal) - 35, 40, 43 (B) Females # 4 and # 5 (ventral) - 45, 49 (a) Idiosoma (ventral) - 45, 49 (b) Gnathosoma (ventral) - 46, 50 (c) Legs I-IV, left and right (ventral) - 47, 50 (C) Male # 1 (ventral) - 52 (a) Idiosoma (ventral) - 52 (b) Gnathosoma (ventral) - 52 (c) Legs I-IV, left and right (ventral) - 53 (D) Male # 2 (ventral) - 56 (a) Idiosoma (ventral) - 56 (b) Gnathosoma (ventral) - 57 (c) Legs I-IV, left and right (ventral) - 57 6

4. Prasadiseius kayosiekeri (Prasad, 1970b) - 58 (A) Female (dorsal) - 58 (a) Idiosoma (dorsal) - 58 (b) Gnathosoma (dorsal) - 58 (c) Legs I-IV, left and right (dorsal and ventral) - 59 (B) Females (ventral) - 60 (a) Idiosoma (ventral) - 60 (b) Gnathosoma (ventral) - 60 10. Discussion - 61 11. Acknowledgements - 96 12. References - 97 13. Tables 1-12 - 100 14. Abbreviations and terms - 115 15. SEM Figures: 1. Prasadiseius cocytes (Prasad, 1970a) - 123, 197 Males [Figs. M1-M73] - 124 Females [Figs. F1-F58] - 198 2. Prasadiseius donahuei (Prasad, 1970a) - 256, 306 Females [Figs. F1-F49] - 257 Males [Figs. M1-M10] - 307 3. Prasadiseius incanus Prasad and Guanilo, 2011 - 317, 494 Females [Figs. F1-F175] - 318 Males [Figs. M1-M67] - 495 4. Prasadiseius kayosiekeri (Prasad, 1970b) - 562 Females [Figs. F1-F28] - 563 16. Line Drawings: 1. Prasadiseius cocytes (Prasad, 1970a) - 591 Male [Figs. L1-L12] - 592 Female [Figs. L13-L23] - 604 2. Prasadiseius donahuei (Prasad, 1970a) - 616 Females [Figs. L24-L39] - 617 Males [Figs. L40 and L41] - 633 3. Prasadiseius incanus Prasad and Guanilo, 2011 - 636 Female [Figs. L42-L80] - 637 Males [Figs. L81-L98] - 676 4. Prasadiseius kayosiekeri (Prasad, 1970b) - 695 Females [Figs. L99-L107] - 696 17. Addendum (Some additional SEM Figures) - 705 18. Some key fi gures - 741 19. Conclusions - 762 20. Index - 771 7 6. INTRODUCTION Mites of the families Blattisociidae, Otopheidomenidae, Phytoseiidae, and Podocinidae belong to superfamily Phytoseioidea of the cohort Gamasina and order Mesostigmata. Detail characteristics of these and related superfamilies are given by Lindquist, Krantz, and Walter (2009). The number of setae on diff er- ent segments of the palp of a typical mesostigmatic are as follow (from trochanter to tarsus; palp coxa is fused with gnathosoma): larva - 0, 4, 5, 12, 11; protonymph - 1, 4, 5, 12, 15; deutonymph and adult - 2, 5, 6, 14, 15. Thus, in comparison to a larva, 1 seta is added on the trochanter and 4 setae added on the tarsus in the protonymph. Also, in comparison to a protonymph, 1 seta is added on the trochanter, 1 seta added on the femur, 1 seta added on the genu and 2 setae added on the tibia in the deutonymph, a pattern retained in the adult. The superfamily Phytoseioidea is most closely related to superfamily Ascoidea. Thus, some features are common to both the superfamilies.

In the adult mites of the superfamily Phytoseioidea, the femur of leg I usually has 11-12 setae (in- cluding 4-5 dorsal, 3 ventral and 4 lateral), the tibia of leg I with 7-11 setae (4-6 dorsal, 2-3 ventral and 1-2 anterolateral), and the tibia of leg III with 7-8 and, rarely, with 9 setae. The palp tarsus usually has a 2-tined palpal apotele in Phytoseiidae. In addition, the “Phytoseiid type sperm access system” is present in which sperm induction pores (= solenostomes) in the females are present at the base of coxa III on the left and right that often lead to a sclerotized spermathecal calyx, with an associated sac-like vesicle, atrium, and mi- nor duct (such mites are called “podospermic” but having the “Phytoseiid type” sperm access system). Lar- vae are with or without a pygidial shield, protonymphs have a pygidial shield and with 5-8 pairs of setae, and deutonymphs have a dorsal shield covering almost the entire idiosoma with lateral incisions or divided as in the adults. In comparison to the above, mites of family Ameroseiidae representing the superfamily Ascoidea, possess the “Laelapid type sperm access system” in which sperm induction pores in the females are also present on each lateral side at the bases of coxae III but lead to a single unpaired central sper- mathecal sac. This is similar to some otopheidomenid mites (such mites with secondary sperm induction pores are called “podospermic” in comparison to “tocospermous taxa” that have “Direct vaginal or other sperm access systems”). In podospermic mites like Dendrolaelaps representing the family Digamasellidae, superfamily Rhodacaroidea (Wiśniewski and Hirschmann, 1991), Hattena (representing the family Amero- seiidae, superfamily Ascoidea (Faraji and Cornejo, 2006), Prasadiseius (representing the family Oto- pheidomenidae, superfamily Phytoseioidea (Prasad, 1970a, b), and Tropicoseius (representing the family Melicharidae, superfamily Ascoidea (Wiese and Fain, 1996) typically possess a long tubular sperm access system including the “Tubulus annulatus” as seen in the “Laelapid type” sperm access system (Krantz and Walter, 2009). Adult females of the ameroseiid genera Afrocypholaelaps, Hattena and Neocypholaelaps are phoretic on bees, butterfl ies, wasps and fl ower-visiting fl ies and scarab beetles in the tropics and subtropics and feed on nectar of fl owers that are visited also by sphingid moths. Prasad (1968a) reported 4 females of Proctolaelaps lobatus De Leon, 1963 (representing family Melicharidae, superfamily Ascoidea) from the proboscis of a noctuid moth collected at Manoa, Oahu Island, Hawaii, that have a tubular spermatheca of the Laelapid type but without having single central spermathecal sac. This is similar to Noctuiseius treati Prasad, 1968, infesting Noctuidae and in all species of Prasadiseius infesting Sphingidae.

In some families of the superfamily Phytoseioidea (Blattisociidae, Otopheidomenidae, Phyto- seiidae, and Podocinidae) and the superfamily Ascoidea (Ameroseiidae, Ascidae, and Melicharidae), the sperm access system is either of “Laelapid type” or of “Phytoseiid type”. For example, as mentioned above, some species of Ameroseiidae, e.g. Neocypholaelaps lindquisti Prasad, 1968a, has a Laelapid type with a single, central spermathecal sac in which both left and right tubuli annulati enter at the anterior end of the sac (Prasad, 1968a: 130, Fig. 1e; Treat, 1975: 118, Fig. 56e). But, some species of Melicharidae, e.g. Proctolaelap lobatus stated above have a Laelapid type without showing a single central spermathecal sac but showing tubuli annulati with a tiny enlarged sac at the end (Prasad, 1968a: 131, Fig. 2F; Treat, 1975: 138, Fig. 71f). At present, the current classifi cation system of Phytoseioidea is not based on the sperm ac- cess systems but based on other morphological and host preference systems. 8 Species of Otopheidomenidae comprise a distinct group of phytoseioid mites that are known to parasitize insects of the orders Hemiptera, Isoptera, Lepidoptera and Orthoptera in temperate and tropi- cal regions throughout the world (Treat, 1975; Prasad, 1976; Fain and Lukoschus, 1983; Syed and Goff , 1983; Halliday, 1994; Zhang, 1995; Mo, 1996). Presently, the family is subdivided into 3 subfamilies based on host preference and morphological characteristics: (1) Katydiseiinae Fain and Lukoschus,1983, which infest katydids (Orthoptera - Pseudophyllidae and Tettigoniidae) and termites (Isoptera - Termitidae); (2) Otopheidomeninae Treat, 1955, which infest noctuid and sphingid moths (Lepidoptera - Noctuidae and Sphingidae); and (3) Treatiinae Wainstein, 1972, which infest various true bugs (Hemiptera - Lygaei- dae, Miridae, Nabidae, and Pyrrhocoridae). The subfamily Otopheidomeninae is comprised of 3 genera: Noctuiseius Prasad, 1968 and Otopheidomenis Treat, 1955, which infest noctuid moths and Prasadiseius Wainstein, 1972, which infest sphingid moths. Nabiseius rivnayae Amitai and Swirski, 1980, is known from Caryophyllales (fl owering plants), Noctuiseius batoridgi Prasad, 1987, from unidentifi ed moth in Malaise trap collection, Treatia ghaiguptaorum (= Treatia indica) Zhang, 1995, from malformed mango infl orescence, and Treatia phytoseioides (Baker and Johnston, 1959) from unknown Hemiptera.

The family Otopheidomenidae is considered to be closely related to the Phytoseiidae (Krantz and Khot, 1962; Evans, 1963; Chant, 1965; Chant and Yoshida-Shaul, 1992; Chant and McMurtry, 2007) but comprises species that have adapted to parasitize various insects. Unlike the chelicerae of phytoseiids, which have well developed sickle-shaped fi xed and movable digits, the otopheidomenids have a reduced (atrophied) fi xed digit but well developed sickle-shaped or narrowly elongate movable digit. In addition, there is a reduction in the number of setae on the dorsal idiosoma in Otopheidomenidae. Because of these unique features and their close relation to phytoseiids, otopheidomenids have received more attention since the family was established by Treat in 1955 leading to description of several new species (Krantz and Khot, 1962; Evans, 1963; Chant, 1965; Chant and Lindquist, 1965; Treat, 1965, 1975; Prasad, 1968a, b, 1970a, b, 1972, 1973, 1975 (1976), 1987, 2011a, b, c, d, e, 2012; Wainstein, 1972; Fain and Lukoschus, 1983; Syed and Goff , 1983; Halliday, 1994; Prasad, Guanilo, Grados, and Prasad, 2011; Negm and Alatawi, 2013).

Treat (1955) established the family Otopheidomenidae with type genus Otopheidomenis and type species Otopheidomenis zalelestes collected from diff erent species of noctuid moths (Lepidoptera: Noctui- dae) of the genus Zale. He considered that this gamasine species belonged to the superfamily Parasitoidea Banks. He gave the following diagnostic characteristics for the family: (1) Palp tarsus with 2-tined spe- cial seta (referring to palpal apotele); (2) Tritosternum vestigial or absent in both sexes and all stages; (3) Sternal shield with 3 pairs of sternal setae (metasternal setae absent); (4) Peritremes of female extending beyond anterior border of coxae II; and (5) Ectoparasitic on insects. Note here that he did not include ab- sence or reduction of the fi xed cheliceral digit as a diagnostic feature for this family although he mentioned on page 556 that this and the vestigial tritosternum in association with the long peritreme distinguished Otopheidomenidae from any previously described family. Further, he gave the following diagnostic char- acteristics for the genus Otopheidomenis: (1) Fixed digit of female chelicera absent or greatly reduced; (2) Movable digit of female chelicera denticulate; (3) Dorsal shield deeply notched near middle of lateral mar- gins; and (4) Eggs laid in or near external tympanic recess of noctuid moths. Note that he never mentioned about the anal shield with anus being in terminal position. He did not give diagnostic features of the new species but described it in suffi cient details to identify this species.

Treat (1965) recognized problem of family Otopheidomenidae when he discovered several new species of Hemipteroseius from pyrrhocorid bugs (Hemiptera: Pyrrhocoridae, red cotton bugs) that had several diff erent characteristics than given above in his family diagnosies and yet placed them in the fam- ily Otopheidomenidae. He found species of this genus having a 2-tined palpal apotele, reduced fi xed digit with a variable length of the peritreme, but with a well developed tritosternum and 4 pairs of sternal setae (including metasternal setae). Also, he noted that the dorsal shield in diff erent species was either entire, par- tially incised bilaterally, or divided into 2 separate shields, had a total of 16 pairs of dorsal idiosomal setae in which several variations were present (e.g. pair of j6 were absent, right Z4 was absent, and pair of s6 were on integument in H. antilleus Treat, 1965, when s6 were present on dorsal shield in H. ageneius Treat, 1965, and H. sabbaticus Treat, 1965). In addition, the anus in some species was in a terminal position (as in

H. antilleus and H. parvulus) while in a ventral position in other species (as in H. sabbaticus)while. Thus, only 2 9 characters (reduced fi xed digit of chelicera and parasites of insects) were consistent for the family.

During 1965-2015, several new species were described in the family Otopheidomenidae from dif- ferent insect hosts and non-insect habitats (Bermuda grass - Negm and Alatawi, 2013; Hemiptera - Costa, 1968; Amitai and Swirski, 1980; Ramsay, 1973; Halliday, 1994; Menon et al., 2011; Isoptera - Zhang, 1995; Lepidoptera - Prasad, 1968a - Noctuiseius treati; 1970a - cocytes and donahuei; 1970b, 1970c - kayosiekeri and pholusis; 1972 - achlora and aporodes; 1973 - indicus; 1987 - Noctuiseius batoridgi; Syed and Goff , 1983; Prasad et al., 2011; Orthoptera - Fain and Lukoschus, 1983; Mo, 1996; mal formed mango infl orescence - Ghai and Gupta, 1984). In these, the following characteristics are noted: (1) Only Eickwor- tius termes Zhang described from Isoptera has 21 pairs of dorsal idiosomal setae (16 pairs on dorsal shield and 5 pairs on dorsal integument) while all others have 11-18 pairs of dorsal idiosomal setae (9-17 pairs on dorsal shield and 1-6 pairs on dorsal integument). (2) Dorsal shield entire, incised, or divided into 2 sepa- rate shields. (3) Tectum smooth. (4) Fixed digit of chelicera reduced. (5) Palpal apotele absent or present. (6) Capitular setae absent or present on ventral gnathosoma. (7) Tritosternum absent, present, or reduced. (8) Sternal shield with 3 pairs, ST1-ST3, or fewer setae and others on integument. (9) Metasternal setae ST4 absent or present. (10) Genital shield seta ST5 absent or present. (11) Ventral opisthosomal setae from 1-10 pairs. (12) Anal shield without preanal setae or ventrianal shield with from 1-3 pairs of preanal setae. (13) Anus terminal or ventral. (14) Sperm induction system in female being laelapid type or phytoseiid type. (15) Legs I-IV with or without macrosetae. Considering the original defi nition of Otopheidomenidae, it is evident that only 3 characters hold true for all the known species (except Eickwortius termes) which are: (1) Hypotrichous condition of the dorsal idiosoma, (2) Reduced fi xed digit of chelicera and (3) Associ- ated with insects.

The mites of the Otopheidomenidae are most closely related to Phytoseiidae on the basis of DNA studies and being more hypotrichous than the mites of Phytoseiidae, and thus both are considered sister groups sharing a recent common ancestor, with the otopheidomenids generally being more morphologically regressive than the free-living phytoseiids (Chant and McMurtry, 2007). On the other hand, these families are also closely related to Blattisociidae of the same superfamily Phytoseioidea but being generally more hypotrichous than the Blattisociidae. This superfamily Phytoseioidea is related to the superfamily Ascoidea that include the families Ameroseiidae, Ascidae, and Melicharidae. Several species belonging to both of these superfamilies, their families, and genera are associated with or are free-living and visit fl owers and feed on pollen. Some possess characteristics of Otopheidomenidae including the sperm induc- tion system of the female, reduced fi xed digit and reduced tritosternum, etc. However, they have greater number of dorsal idiosomal setae than in the Otopheidomenidae or the Phytoseiidae. The tritosternum is also reduced in Hattena rhizophorae Faraji and Cornejo, 2006 (family Ameroseiidae) and occasionally in other taxa of the cohort Gamasina (e.g. in the superfamily Epicrioidea, but here the mode of sperm transfer is “tocospermous” in which the male uses the primary genital opening of the female for sperm transfer, and in the family Spinturnicidae in the superfamily Dermanyssoidea, which are ectoparasites of bats). The fi xed digit of the chelicera is reduced also in Tropicoseius chocoensis (Wiese and Fain, 1996) (family Meli- charidae) and in Laelaps hilaris Koch (family Laelapidae). Thus, some of the characteristics mentioned by Treat (1955) as characteristics of Otopheidomenidae, are present also in some other families of Gamasina, and may just represent the morphological and functional adaptations of these mites. Also, some of the mites of Ascoidea and Phytoseioidea feed on pollen present in fl owers that are also visited by moths of the families Noctuidae and Sphingidae, which harbor some mites of the Otopheidomenidae, where these moths feed on the fl ower nectar.

The 30 species known in 10 genera of family Otopheidomenidae are as follows: (1) Eickwor- tius - termes Zhang, 1995; (2) Entomoseius - dysderci (Evans, 1963); (3) Hemipteroseius - adleri Costa, 1968; ageneius Treat, 1965; antilleus Treat, 1965; indicus (Krantz and Khot, 1962); parvulus Treat, 1965; sabbaticus Treat, 1965; vikrami Menon, in Menon, Joshi, Hussain and Ramamurthy, 2011, junior synonym of indicus (Krantz and Khot, 1962) per Prasad (2017); womersleyi Evans, 1963; (4) Katydiseius - nadcha- trami Fain and Lukoschus, 1983; (5) Nabiseius - arabicus Negm and Alatawi, 2013; duplicisetus Chant and Lindquist, 1965; melinae Halliday, 1994; rivnayae Amitai and Swirski, 1980; (6) Noctuiseius - batoridgi Prasad, 1987; treati Prasad, 1968b; (7) Orthopteroseius - sinicus Mo, 1996; (8) Otopheidomenis - ascala- 10 phae Syed and Goff , 1983; zalelestes Treat, 1955; (9) Prasadiseius - achlora (Prasad, 1972); aporodes (Prasad, 1972); cocytes (Prasad, 1970a); donahuei (Prasad, 1970a); indicus (Prasad, 1973); incanus Prasad and Guanilo, in Prasad, Guanilo, Grados and Prasad, 2011; kayosiekeri (Prasad, 1970b); pholusis (Prasad, 1970b); and (10) Treatia - dieuches Ramsay, 1973; ghaiguptaorum Zhang, 1995 (new name for indica Ghai and Gupta, 1984); and phytoseioides (Baker and Johnston, 1959).

Systematic position of some of the above species included in Otopheidomenidae is controversial as they possess characteristics of Ascoidea or Phytoseioidea, including several pairs of “R” setae and “S” setae, but are placed in Otopheidomenidae due to comparative hypotrichy, some minor features, and as- sociation with insects. Thus, they do not share all the diagnostic features Treat (1955) gave for the family Otopheidomenidae. Seeman (2012) described a new hypotrichous mite (Krantzoseius walteri) in the family Blattisociidae from Australian burrowing carabid beetles that has a total of 20 pairs of dorsal idiosomal setae and the phytoseiid type sperm access system that could have been placed in Otopheidomenidae. He mentioned the diffi culty of this mite and why he placed it in family Blattisociidae. He also mentioned that Otopheidomenidae is likely not a phytoseioid taxon.

The species of the genus Prasadiseius Wainstein, 1972 (Otopheidomeninae) are known to only parasitize hawk moths (Lepidoptera: Sphingidae) and have been found primarily in the Neotropical region with two species from other tropical regions of the world (Prasad, 1976, 2011). Recently, Prasadiseius cocytes was found to be widely distributed in several neotropical countries: Brazil, Ecuador, Guatemala and Peru (Prasad, 2011) and in Colombia and Mexico (Prasad, 2013). Also, Prasadiseius donahuei was found to be present in Cuba, Mexico and the state of Arizona in USA (Prasad, 2013). All stages of the mite are present dorsally mainly in the rectangular area of the tympanum, the metathorax, and fi rst abdominal tergite of the moths; occasionally, they are found on the proboscis, around the eyes, and on the dorsum or venter of the head, thorax, abdomen and wings.

The only paper based on scanning electron microscopic studies of Otopheidomenidae was published on Prasadiseius kayosiekeri (Prasad, 1970b) by Prasad and Walker (2011). That SEM study is expanded to include other species of this subfamily from recently collected specimens and include not only the idiosoma but the gnathosoma and legs as well, the results of which are presented in this book. It also in- cludes details of males and females as seen in SEM studies never done before. In addition to morphological details as seen in SEM studies, reconstruction of the gnathosoma including the tectum, corniculi and some other structures in schematic diagrams are also provided.

It should be noted that some species of the predatory mite genus Veigaia that have the “Laelapid type sperm access system” in which sperm induction pores are present in between the bases of coxae III-IV in the females, may have bisexual reproduction or may have only unisexual population where only females occur (Di Palma, Alberti, Nuzzaci, and Krantz, 2006). It is interesting to know that unisexual Veigaia have a wider geographic distribution (Cuellar, 1977) than the bisexual species and also have a more northern distribution (Hurlbutt, 1979, 1984). Do the otopheidomenids have only bisexual reproduction or unisexual populations also? Why was the infestation rate by the male mites much higher than that of the female mites, of some moth hosts in some countries than other countries as shown by Prasad (2013b)? Neither these nor the ultrastructure of these mites have been studied in the Otopheidomenidae and will need to be done in fu- ture. It is hoped that this major work will draw the attention of other morphologists and anatomists improv- ing this work further in the future.

The 4 species of family Otopheidomenidae included for SEM studies in this book represent subfamily Otopheidomeninae. The species are: (1) Prasadiseius cocytes (Prasad, 1970a), (2) Prasadiseius donahuei (Prasad, 1970a), (3) Prasadiseius incanus Prasad and Guanilo, 2011, and (4) Prasadiseius kay- osiekeri (Prasad, 1970b). Some of these species were recently collected in a survey, and the SEM study was completed. The objectives of this study were: (1) To describe the morphological structures, (2) To identify and describe characteristics not reported before, and (3) Come to some conclusion about the systematic position of these mites. 11 For the results, each species is discussed separately giving SEM photos of the dorsal idiosoma and the setae, ventral idiosoma and associated structures with the setae, dorsal gnathosoma and associated structures followed by ventral gnathosoma and associated structures, and left and right legs I-IV. Internal structures of female sperm access system are not included as these were not seen in this study. As setae on the leg segments are arranged in whorls (usually in pairs into four sections of the leg segment: al = antero- lateral, ad/av = anterodorsal/anteroventral, pd/pv = posterodorsal/posteroventral, and pl = posterolateral) and legs are photographed in fl attened condition, some of the dorsal or ventral setae may not appear in the same position. Also, some of these setae may have apparently moved to anterior or posterior position on the segment depending on the orientation of the legs. Readers are suggested to refer Evans (1963a) who has discussed the chaetotaxy of the legs of motile stages (larva, protonymph, deutonymph, female and male adults) in the free-living Gamasina (Mesostigmata) in detail. In this work, he gave the chaetotactic formula [al, ad/av + pd/pv, pl] very nicely and in detail as seen in diff erent families of Gamasina. He called unpaired setae appearing during ontogeny on the dorsal and ventral surfaces of tarsi II-IV as mediodorsal (md) and medioventral (mv), respectively. This system and notations, never given before in the Otopheidomenidae (except for a few), are used here.

Articles and verbs are avoided in description of the species as much as possible. If a mite is seen dorsally, its left is considered left (evident on left of the examiner) and right is considered right (evident on right of the examiner) but if this mite is seen ventrally, its left is considered left and right is considered right. All setae on the idiosoma, the gnathosoma and the legs are smooth unless stated otherwise. Most of the leg segments are rotated anteriorly toward the gnathosoma or posteriorly toward the opisthosoma or the anal shield. In such cases when the mite is in dorsal view, these are indicated on the fi gures, keeping anterodorsal and posterodorsal setae in mind, but when the mite is in ventral view then the anteroventral and the posteroventral setae, along with ventral hook (= condyle), was kept in mind. As one specimen may not show all the characters, details of more specimens are included as available, which have provided ad- ditional information.

Several tables are included for the benefi t of the readers, who may like to know more details about all or some species of the Otopheidomenidae, other than the 4 species included in the present work. Many line drawings are also given to give the readers more clarity. For the same reason, a lot of historical and other details of this family along with the related families are also included. It should be noted that the 4 species included in the present study are rare mites, having very low infestation rates on their hosts (usu- ally about 1-3%) including immature stages much more than the adults (usually males much less than the females) that have next developing stages inside the body indicating accelerated developments and having short life stages of the immature stages in these obligatory ectoparasitic mites. Finding of suitable speci- mens for SEM preparations was limiting as well. In spite of all diffi culties, this work was completed after 8 years of hard work and presented with the hope that this book will be helpful to the readers. In a work of this kind, errors are possible. If noted, readers are requested to inform the author. I feel as I have included too many SEM photographs but I had no choice but to take these in diff erent magnifi cations for better understanding of the characteristics. When this work was complete, several photographs of eggs and other stages (including motile immature stages) were added at the end for better understanding of these mites. It should be noted that this author has placed several videos of these otopheidomenid mites (including the live stages) on “Youtube” (under Vikram Prasad M.D.) that are worth watching. Opinions expressed in this book, including current defi nition of Otopheidomenidae and creation of several new taxa, are entirely of this author and are done to facilitate the future work on these mites. 592 R T TR L A I E P T TI E - F FE D G GE F FD R R-PAL H D S C -P = tibia of palp, TR-P = TR-P = tibia of palp, -P - - R R-SD R R-CH E H T TE C - D L L-CH S D - F FD L L-SD

H

C H S

- d structures in dorsal view (scale bar = 10 μm) [BS E E C I H

R F FE -

T TI G GE C CHS , R

S , R BBS, R-CH S xed digit, FE-P = femur of palp, GE-P = genu of palp, GN = femur of palp, GE-P xed digit, FE-P T TR L M MS, R-CH A P -

L L-PAL , A H M C M O

- - S

R R-CH L O A H S T chelicera, PA, PAL = palp, SD = spermatodactyl, TE = tectum, TI TE = tectum, = palp, SD spermatodactyl, PAL chelicera, PA, A R N O E seta, DS = dorsal shield, FD fi T TE G GNATHOSOMA, - M D DORSAL H C - L L-CH L A P - R R-PAL (Prasad, 1970a) [VP10-36, male # 1] - Gnathosoma with associate N S G GN D DS D Prasadiseius cocytes S SD L A T Fig. L1. D P N F FD - I INT L L-PAL gnathosoma, INT = integument, M male, MS middle segment of gnathosoma, INT trochanter of palp]. = basal segment of chelicera, CH CHS cheliceral 593 r ? ? ? ??? D S - ? R R-SD ? ? ??? D S - R R-SD xed digit (FD) of chelicera and spermatodactyl (SD) ed fi

, D ed structure of spermatodactyle (???) in dorsal view (scale ba A S - M L L-SD M O

- S D

L S O - A H L L-SD S T A R N O GNATHOSOMA, G GNATHOSOMA, - M D DORSAL

, n S e D H k F C - ral seta (CHS), and an unidentifi o - r M M R R-FD R-CHS, R R-CHS, b broken R R D A ARM A ARM H S C - - R R-SD R R-CH D F - R R-FD D P PD (Prasad, 1970a) [VP10-36, male # 1] - Enlarged views of atrophi (Prasad, 1970a) [VP10-36, male # 1] - Enlarged Prasadiseius cocytes D D F F H - - Fig. L2. M S D C L L-FD L L-FD - R H S L L-CH - A ARM C L L-SD - L L-CHS showing arthrodial membrane (ARM), pilus dentilis (PD), chelice = 10 μm). 742 s (Prasad, n o i e t s c D o e j n M i o - P. cocytes P. r p L L-MD s spinose p projections

,

h , p 7 s 1 e m

6 t , A y l 3

y , n - a ctions. Right - c r o 0 H o t t D xed digit (FD) and well developed 1 c n C o

(left) - Note much enlarged distal end M - r P . e - P. cocytes, P P. P Protonymph, v ventral, A17 R R-CH, V VP10-36 R R-MD s n o l end of atrophied fi i e t s c o e j n i o

R r p , 8 s spinose p projections s

O , M u C

1 4 - , n

l 3 L L-COR a # - a

c r 0 t e n l 1 i n

a P . e P. incanus, P P. m male # 1, v ventral, M8 V VP10-34 . Middle, male # 1, ventral: Left corniculus with spinose proje owing spinose projections. ly placed denticles posterior to pointed and hooked tip. Inset 1 2 D 3 4 M MD p i t tip Prasad and Guanilo, 2011 [VP10-34] - Left, larva, dorsal: Dista D F FD

D , l M MD a

, s r 2 s o 3 u 4 d n A

3

Prasadiseius incanus , a , - c a 0 H v n 1 i r C

- P D a . P. incanus, P P. L Larva, dorsal, A32 R R-CH, V VP10-34 F FD Fig. A36. xed digit showing shrivelled or collapsed region (dark arrows) A36 of fi entire movable digit (MD) of chelicera in a larva with 4 distal 1970a) [VP10-36] - Protonymph, ventral: Left movable digits sh 761 A L C CLA L , I I I T L - R R R-LIII, P PRT L D ctyli (PARD), and basally wide ctyli (PARD), to positioning of the specimens. (Prasad, 1970a) [VP10-36, male # 1, with distally expanded pretarsal folds 5 6 P. cocytes P. s , e M 1 6 t ht,

L , 3 y l # - U c

a 0 P PUL o e s l 1 c r

a P o . P. cocytes P P. m male # 1, d dorsal, M65 V VP10-36 X E P A - R R-APEX A L C CLA et-like concavities for holding the substrate. Bifurcate parada aradactyli and tiny claws may not be seen in all specimens due S stally wide dorsal (D) and narrowly elongate lateral (L) stalks , D I T L L R - O L L-LI, P PRT F X

D E T P R (Prasad, 1970a) [VP09-9, Tiedt, female # 5, left leg I] and Rig Tiedt, (Prasad, 1970a) [VP09-9, PRT FOLDS P PRT A - D D-APEX , t d , e i Prasadiseius donahuei i S e T D 5 u

, L h # 0 O a 9 2 F e - n

l F 9 o T a . 0 d R ) X m g

P i . PRT FOLDS P PRT D e E Fig. A55. Left, Fig. P. donahuei, P P. f female # 5 Tiedt, F20, F Fig. V VP09-9 R P A A A L - P C CLA L L-APEX ( (PARD) A55

and centrally located membranous pulvillus having numerous pock claws (CLA) with hooked and pointed tips also seen. Note that p right leg III] - Distal pretarsus (PRT) in dorsal view with di right leg III] - Distal pretarsus (PRT)