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Welbourn, Warren Calvin, Jr.

PHYLOGENETIC STUDIES OF TROMBIDIOID MITES

The Ohio Stale University Ph.D.

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University Microfilms International PHYLOGENETIC STUDIES OF TROMBIDIOID MITES

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Philosophy in the Graduate

School of The Ohio State University

By

Warren Calvin Welbourn Jr, B.A., M.A.

sjc >je 3jc >jc Sjt

The Ohio State University

1985

Reading Committee: Approved By

Donald E. Johnston

Susan W. Fisher

Norman F. Johnson Adviser Department of Entomology Copyright by Warren C. Welbourn Jr 1985 ACKNOWLEDGEMENTS

I wish to thank my advisor, Dr. Donald E. Johnston, for his advice, assistance and patience during this study. Very special thanks go to my wife and family for their continuing support.

Funds for my 1984 trip to the British Museum (Natural History),

London, Museum National D’Histoire Naturelle, Paris, Museum

D ’Histoire Naturelle, Geneva and Istituto Spermentale per la Zoologia

Agrara (Berlese Collection), Florence and related field work in

England and Italy was supported by a National Science Foundation

Dissertation Grant (BSR-8401206). Travel from Florence, Italy to

University "Al. I. Cuza." in Iasi, Romania was partially supported by an Ohio State University Graduate Student Alumni Research Award. My visit to the Smithsonian Institution in April, 1981 was supported by a Visiting Researcher Grant from the Smithsonian Institution.

I would like to acknowledge the assistance of Ms. Anne Baker,

Dr. Don Macfarlane, Mr. Keith Hyatt and Mr. Keiren Martin in the

Arachnid Section, of the British Museum (Natural History), London, for their assistance in 1982 and 1984 and arranging for loans of important specimens. I also want to thank Mrs. Judith Marshall

(Entomology) for her assistance in my examination of the Orthopteran collections for mites. I am especially grateful to Anne Baker and

Paul Hillyard for arranging a very successful spring 1984 collecting trip to south Wales. I also want to thank Clive and Diane Bowman for their hospitality during my two visits to London.

I want to acknowledge Dr. Yves Coineau, Director of the

Laboratory of Arthropods, Paris for his assistance while in Paris, as well as Mr. Michel Naudo for his patience and assistance as well as arranging for the loan of important specimens. I am especially grateful to Dr. Pierre Robaux for opening his laboratory to me and for the stimulating discussions on trombidioid mites, in addition to his hospitality. I am also grateful to Dr. Bernard Hauser of the

Geneva Natural History Museum for his assistance in locating specimens in the Vercammen-Grandjean collection.

I am indebted to Dr. Fausta Pegazano for her assistance in making ray visit to the Berlese Collection successful. Dr. Roberto

Nannelli is appreciated for his assistance and taking the time to take me collecting at several of Berlese's collecting localities in the mountains around Florence.

I wish to gratefully thank Dr. Libertina Solomon for providing space in her laboratory and locating trombidioid specimens in the

Feider Collection. I am especially grateful for her family's hospitality while I was in Iasi.

The 1982 field work in California and Oregon was supported by a small grant from The Ohio State University to Dr. Donald E. Johnston. I thank Dr. G. W. Krantz for opening his laboratory to us, and Dr.

David Walter for his assistance in Corvallis.

I acknowledge Drs. G. W. Wharton and G. R. Needham of the

Acarology Laboratory for their assistance, interest and support.

I am especially grateful to Dr. Edward Baker for his continuing

assistance in locating missing specimens and lost literature. I also

thank Drs. Evert E. Lindquist (Agriculture Canada) and John B.

Kethley (Field Museum Natural of History, Chicago) for their

consultation and support.

I am especially indebted to the many people who loaned and/or

donated specimens for this research. In addition to the persons mentioned above I thank Dr. B. Feldman-Muhsam (Hebrew University

Hadassah Medical School, Jerusalem), Dr. Torbjorn Kornestedt

(Naturhistoriska Rimsrauset, Stockholm), Dr. David Lee (South

Australia Museum), Drs. Ian Smith and Evert Lindquist (Agriculture

Canada, Ottawa), Dr. L. van der Hamraen (Rijksrauseura voor Natuurlijke

Historie, Leiden), Drs. R. G. Beer and G. W. Byers (Snow

Entomological Museum, University of Kansas), Dr. H. Levi (Museum of

Comparative Zoology, Harvard University), Dr. Norman Platnick

(American Museum of Natural History, New York), Dr. Gary Mullen

(Auburn University, Auburn), Dr. Barry M. OConnor (Museum of Zoology,

University of Michigan, Ann Arbor), Dr. Donald Tuttle (University of

Arizona Experiment Station, Yuma), Dr. John Moser (USDA, Forest

Service, Southern Forest Experiment Station, Pineville), Dr. John B. Kethley (Field Musuera of Natural History, Chicago), Dr. Wayne Moss

(Philadelphia Academy of Natural Sciences, Philadelphia), Dr. Harold

Denmark (Florida Department of Agriculture and Consumer Services,

Division of Entomology, Gainsville).

To the numerous other people who contributed directly or indirectly to this reseach I am very appreciative. VITA

December 19, 1946 ...... Born - Pasadena, California

1966...... A.A. - Zoology Citrus College Azusa, California

1969...... B.A. - Biology California State University, Long Beach, California

1972...... M.A. - Biology California State University Long Beach, California

1971-1978 ...... Teacher Albuquerque Public Schools Albuquerque, New Mexico

1978-1985 ...... Collection Manager Acarology Laboratory Department of Entomology The Ohio State University Columbus, Ohio

PUBLICATIONS

Loomis, R. B. and W. C. Welbourn 1969. A new species of Hannemania (Acarina, Trombiculidae) from Bufo punctatus of western North America, with comments on Hannemania hylae Ewing. Bull. So. Calif. Acad. Sci. 69:160-168.

Welbourn, W. C. and R. B. Loomis 1970. Three new species of Hannemania (Acarina, Trombiculidae) from amphibians of western Mexico. Bull. So. Calif. Acad. Sci. 69: 65-73.

Welbourn, W. C. 1972. Studies of two species of Hannemania (Acarina, Trombiculidae) on amphibians in western North America, (abstract) Masters Abstracts 10 No. 4.

Welbourn, W. C. and R. B. Loomis. 1975. Hannemania (Acarina, Trombiculidae) and their anuran hosts at Fortynine Palms Oasis, Joshua Tree National Monument, California. Bull. So. Calif. Acad. Sci. 74: 15-19.

Welbourn, W. C. 1978. Biology of Ogle Cave with a list of the cave fauna of Slaughter Canyon. NSS Bulletin 40: 27-34.

vi Welbourn, W, C. 1979. Invertebrate fauna of the earth cracks of Wupatki National Monument, Arizona. Proceedings of the First Conference on Scientific Research in the National Parks, New Orleans, LA, November 9-12,1976. Volume 1. National Park Service Trans, and Proc. Ser. No.5: 387-391.

Welbourn, W. C. 1981. Phylogenetic relationships in mites of the family Trombidiidae. (abstract) Amer. Zool. 21:1005.

Goff, M. L., R. B. Loomis, W. C. Welbourn and W. J. Wrenn. 1982. A glossary of chigger terminology (Acari; Trombiculidae). J. Med. Entomol. 19:221-238.

Welbourn, W. C. 1982. Phylogenetic relationships in mites of the superfamily Erythraeoidea. (abstract) Amer. Zool. 22:885.

Welbourn, W. C. 1983. Potential use of trombidioid and erythraeoid mites (Prostigmata: Parasitengona) as biological control agents of insect pests, pp 103-140 ’In: M.A. Hoy, G.L. Cunningham and L. Knutson eds. Biological Control of Pests by Mites, Agricultural Experimental Station Special Publication 3304, University of California, Berkeley, 185pp.

Welbourn, W. C. 1984. Phylogenetic studies on Trombidioidea. pp 135-142. In: D.A. Griffiths and C.E. Bowman eds. Acarology VI (volume 1), Ellis Horwood Limited, Chichester, England, 645pp.

Shelley, W. B., E. D. Shelley & W. C. Welbourn. 1985. Polypodium fern wreaths (Hagnaya): A new source of occupational mite dermatitis. JAMA 253:3137-3138.

GRANTS

Visiting Reseacher Grant, Smithsonian Institution April, 1981.

The Ohio State University Graduate Student Alumni Research Award January, 1984.

National Science Foundation Dissertation Research Grant: "Systematics of the Trombidioidea" March, 1984.

FIELD OF STUDY

Major Field: Entomology

Studies in Acarine Systematics. Professor Donald E. Johnston

vii TABLE OF CONTENTS

Page ACKNOWLEDGEMENTS ...... ii

VITA ...... vi

LIST OF TABLES...... '...... x

LIST OF FIGURES...... xi

INTRODUCTION ...... 1

MATERIALS AND METHODS...... 5

REVIEW OF PREVIOUS CLASSIFICATIONS ...... 10

EXTERNAL MORPHOLOGY AND DISCUSSION OFCHARACTERS ...... 15 LARVAE...... 15 Gnathosoma...... 15 Idiosoma...... 25 Leg Chaetotaxy...... 29 Specialized Setae ...... 48 POSTLARVAL INSTARS (DEUTONYMPH AND ADULT) ...... 53 Gnathosoma...... 54 Idiosoma...... 56

PHYLOGENETIC ANALYSIS...... 61 RELATIONSHIPS OF THE TROMBIDIOIDEA WITHIN THE PARASITENGONA ...... 61 RELATIONSHIPS WITHIN THE TROMBIDIOIDEA...... 72 Podothrombiidae ...... 73 Trombidiidae...... 77 Neothrombiidae...... 82 Calothrombiidae ...... 89 Eutrombidiidae...... 91 Microtrombidiidae ...... 96

KEY TO THE GENERA OF TROMBIDINA (LARVAE)...... 106

viii Page TAXONOMIC SECTION...... 123 Family Podothrombidiidae...... 124 Family Trombidiidae ...... 125 Subfamily Trombidiinae...... 126 Subfamily Allothrombinae...... 131 Family Neothrombiidae ...... 135 Subfamily Neothrombiinae...... 136 Subfamily Aethethrombidiinae...... 147 Subfamily Aegialothrombiinae...... 151 Family Calothrorabiidae...... 153 Family Eutrombidiidae ...... 156 Subfamily Eutrombidiinae...... 157 Subfamily Hoplothrombiinae...... 166 Family Microtrorabidiidae...... 169 Subfamily Feideriinae...... 171 Subfamily Microtrombidiinae ...... 179

BIBLIOGRAPHY ...... 197

APPENDIXES

A. List of Characters...... 210

B. List of Species Examined...... 227

ix LIST OF TABLES

Table Page

1. Leg Nomenclature for Larval Terrestrial Parasitengona ...... 30

2. Proposed Classification of the Parasitengona...... 63

3. Character States used on the Phylogenetic Analysis of the Parastiengona (Families and Superfamilies)...... 65

4. Proposed Classification of the Trombidioidea...... 75

5. Character States used on the Phylogenetic Analysis of the Podothrombiidae & Trombidiide ...... 78

6. Character States used on the Phylogenetic Analysis of the Neothrombiidae...... 84

7. Character States used on the Phylogenetic Analysis of the Eutrombidiidae...... 92

8. Character States used on the Phylogenetic Analysis of the Microtrorabidiidae ...... 97

9. List of Charcates used in Phylogeneitc Analysis .... 210

10. List of Speicmens Examined...... 227

x LIST OF FIGURES

Figure Page

1. Dorsal idiosoma of Trombidium holosericeum L. (Trombidiidae)...... 19

2. Subcapitular Setae of Larvae...... 24

3. Ventral view of Trombidium holosericeum L. (Trombidiidae)...... 36

4. Legs of Trombidium holosericeumL. (Trombidiidae) . . . 39

5. Lophotrix Setae from Tarsus Leg...III ...... 45

6. Representative Postlarval Dorsal Idiosoraal Setae. . . . 59

7. Cladogram of Hypothesized Relationships within the Parasitengona...... 64

8. Cladogram of Hypothesized Relationships within the Trombidioidea...... 74

9. Cladogram of Hypothesized Relationships Within the Podothrombiidae and Trombidiidae ...... 79

10. Cladogram of Hypothesized Relationships within the Neothrombiidae...... 85

11. Cladogram of Hypothesized Relationships within the Eutrombidiidae ...... 93

12. Cladogram of Hypothesized Relationships within the Microtrombidiidae ...... 98

xi INTRODUCTION

The prostigraatid cohort Parasitengona (Parasitengonae,

Trombidia, etc.) is a large assemblage of mites which share a complex pattern of postembryonic development and a partitioning of ontogeny into parasitic/dispersal and predatory phases. The Parasitengona are currently arrayed in 11 superfamilies (eight of which are Hydracarina or water mites), 45-60 families, some 520 genera and more than 7000 species (Kethley, 1982; Cook, 1974). Many of these mites are well known to non-acarologists as the red velvet mites, as the brightly colored swimmers in lakes and ponds, the obvious red engorged protelean parasites of insects and arachnids, and as the frequently pestiferous and sometimes pathogen-transmitting chiggers parasitic on terrestrial vertebrates. These properties, along with the relatively large size of the adults, have resulted in a considerable amount of anatomical (Feider, 1950; Bottazzi, 1954; Hirst, 1926a, Mather,

1954a, 1954b; Moss, 1960, 1962a; Witte, 1976, 1984), ecological

(Robaux, 1974; Michener, 1946, Newell & Tevis, 1960), and taxonomic

(Andre 1962a, b, 1958; Berlese, 1910, 1912, 1914, 1917a, b, c;

Daniel, 1955, 1956; Feider, 1977, 1979; Hirst, 1926b, 1928, 1929;

Newell, 1957, 1958, 1960; Oudemans 1909, 1910; Robaux, 1967a, b, c;

1 2

Southcott 1982; Thor and Willmann, 1947, Womersley, 1937, 1945 and others) work directed to these mites. Despite this effort, the group is poorly known and the classification of the group is unrigorously developed.

The superfamily Trombidioidea consists of an unsettled number of families (Vercammen-Grandjean, 1973; Krantz, 1978; Feider, 1979;

Welbourn, 1984). It is a diverse and ecologically important group with over 200 genera and nearly 3000 species. The larvae of some genera are parasites of economically important insect pests such as aphids (Homoptera), mosquitoes (Diptera), biting midges (Diptera) and grasshoppers (Orthoptera), while the chiggers (Trombiculidae) are well known pests and occasionally vectors of the rickettsial organism, Rickettsia tsutsugamushi. The implications for biological control of insect pests by parasitengone mites have recently been reviewed by Welbourn (1983).

Systematic work on the Trombidioidea has been hindered because of the heteromorphic post-larval instars. Taxonomic change has occurred primarily through the discovery of distinctive new taxa

(Robaux, et. al., 1976; Shiba, 1976; Mullen and Vercammen-Grandjean,

1978) or by removing smaller definable taxa from older groups

(Vercammen-Grandjean, 1980; Feider, 1952a). While most traditional groups can be recognized on the basis of combinations of plesiomorphic and apomorphic characters, the true relationships of these mites are yet to be resolved.

Nomenclatural problems associated with the Trombidioidea include the difficulty in identifying genera from old descriptions and associating larvae with these traditional genera. Many commonly encountered genera (i. e., Trombidium Fabricius. Microtrombidium

Haller, Allothrombium Berlese, Podothrombium Berlese, Eutrombidium

Verdun, Parathrombium Bruyant) were described from Europe and most are poorly defined, resulting in considerable confusion which could threaten the stability of existing classifications.

The traditional essentialist definitions of groups, and the reliance on plesiomorphic characters are further complicated by the larval/ post-larval divergence which has resulted in different taxonomies. Recent authors (Feider, 1979; Michener, 1946; Newell,

1957; Robaux, 1973; Southcott, 1982) have usually tried to integrate larval and post-larval data into their classifications. In contrast to this holomorphological approach, my work has concentrated on the comparative study of the larval instar, a position also suggested by other authors (Vercammen-Grandjean, 1969; Vercammen-Grandjean et. al., 1973; Mullen and Vercammen-Grandjean, 1978). Vercammen-

Grandjean (1969) has elaborately argued for larvae as the "repository of phylogeny". While I do not agree with much of the author's rationale, there are reasons for emphasizing the study of larvae.

First, trombidioid larvae (and parasitengone larvae in general) show similarity to those of other prostigmatid groups. Thus, it is possible to determine homologies among parasitengone and non-parasitengone larvae which permitted establishment of character polarizations. Such conclusions aided, in some instances, in understanding postlarval characters, using congruence as a criterion.

Further, the comparative study of larvae established the broad patterns (generic, familial, suprafamilial) of relationship that are obscured by the divergent array of autapomorphies of the postlarval instars. This is especially true for the water-mites (see Cook,

1974; Smith, 1976).

Because larvae are adapted for host-discovery, parasitic feeding, and dispersal, it seems possible to recognize at least some of the apomorphies of this instar as functional correlates of these habits. This is much more difficult for the postlarval instars, as we know very little about these animals. The problems presented by the postlarval parasitengones are the problems common to gaining an understanding of all free-living Acari in general. Materials and Methods

The theoretical viewpoint of phylogenetic systematics (Hennig,

1966; Wiley, 1981) was used to develop phylogenetic hypotheses of the relationships among the Trombidioidea at the generic, subfamily and family levels using characters derived from a comparative study of the external (cuticular) morphology of the larval instar, to make inferences about the evolutionary history of the Trombidioidea and

Parasitengona, compare the phylogenetic classification of the larvae with the existing postlarval classifications and develop diagnoses and a key to the trombidioid genera studied. These objectives were accomplished by comparative study of specimens borrowed from other collections, studied in some European collections and field collected as well as laboratory reared specimens.

The comparative study of the external morphology of the larval

Trombidioidea included a number of character suites not previously used in systematic studies of the Parasitengona. Appendix A presents a list of characters and character states studied. Character polarization was determined by using the out-group comparison method of Watrous and Wheeler (1981). In addition to morphological data, patterns in larval host preference and additional ecological

5 information was used to provide additional support for the

phylogenetic analysis. Welbourn (1983) listed host distributions for

34 non-chigger trombidioid genera and 77 nominate species which are,

in most cases, consistent with the character analysis.

The sister group of the Parasitengona is believed to be in the

Anystina (Adamystidae, Anystidae, Pseudochelidae, Teneriffiidae,

Caeculidae, Paratydeidae, Pomerantzidae, Pterygosomatidae). In a

thoughtful discussion, Newell (1973) proposed the Pterygosomatidae as

the "ancestral” group to the Parasitengona based on a shared pattern

of postembryonic development that involves the alternation of active

and quiescent instars (calyptostases) in the protonymph and

tritonymph (see Johnston and Wacker, 1967). While both groups share

this synapomorphy the Pterygosomatidae present a number of

autapomorphies associated with parasitism that makes it difficult to

find other characters supporting their monophyly.

The family Anystidae (subfamily Anystinae) was used as the

sister group of the Parasitengona. While the family is poorly known, with few larval forms described, the excellent description of Anystis

by Grandjean (1943) provides a good starting point. The larva of

Anystis is plesiomorphic for characters shared with the

Parasitengona. Witte (1984) suggested Anystis as the sister group of

the Parasitengona, based on reproductive behavior.

To examine the relationships of the Trombidioidea with the rest of the Parasitengona and with the Anystina, a comparative analysis, based on all the trombidioid groups, along with representatives from the Erythraoidea, Calyptostomatoidea and the less derivative water mites (Hydracarina), was undertaken.

Larval mites for study were obtained primarily by loans of determined and undetermined specimens from various institutions and individuals (Appendix B). Many specimens were field collected or reared from field collected adults. Additional species were obtained by searching berlese residues in the Acarology Laboratory collections and removing mites from the pinned Orthoptera in the British Museum

(Natural History) insect collections. Other museums visited include

United States National Museum (Smithsonian Institution), Washington;

Museum of Comparative Zoology, Harvard; Field Museum of Natural

History, Chicago, Museum National d'Histoire Naturelle, Paris, Museum d'Histoire Naturelle, Geneva, Instituto Spermentale per la Zoologia

Agraria (Berlese Collection), Florence and University "Al. I. Cuza"

(Feider Collection), Jassay, Romania.

Field collections were accomplished by extraction from leaf litter using a Berlese apparatus (MacFadyen, 1953). Leaf litter extractions were done over water to facilitate collection of live trombidioids for rearing. Adults were also hand collected under rocks, logs, bark and on trees. Field work vas conducted in Arizona

(1981), Arkansas (1979-1983), California and Oregon (1982), Kentucky

(1979-1980), Maryland (1981), New Mexico (1981), Ohio (1979-1984),

West Virginia (1981, 1983), Great Britain (Wales) (1984) and Italy

(1984).

The technique used to obtain larvae from field-collected adults was modified from the method developed by Wharton (1946; see also

Lipovsky, 1953) for rearing trorabiculid deutonymphs and adults from engorged larvae. Adults were placed in individual culture vials (4

dram glass vials with snap caps), filled one-half to two-thirds with

a mixture of plaster of Paris and activated charcoal mixture (9:1)

(Wharton, 1946). After egg deposition the female was preserved and

the eggs were allowed to develop; the larvae were preserved on

emergence (usually about two weeks). Representative prelarva were

preserved for later preparation as permanent slides. Adults can

survive several months in a culture with minimal care, but eggs are

usually deposited within the first two to four weeks. This method is

superior to rearing from engorged larvae because one can obtain up to

several hundred unengorged larvae making comparisons of individual

variation possible. In addition, the exuviae obtained from an

engorged larva is usually not suitable for detailed taxonomic work

and it is frequently difficult to rear a deutonymph to an adult.

This technique worked quite well for many Trombidiidae,

Neothrombiidae and Microtrorabidiidae but was less successful for

Tanaupodidae and Erythraeidae.

In addition to rearing larvae from eggs, some engorged larvae obtained from hosts were reared to deutonymph and adult stages. An engorged larva was placed in culture vial and observed daily. When the deutonymph emerged, usually in 10-15 days, the larval exuviae was mounted as a microscopic slide preparation. Attempts were made to get the deutonymph to feed on the eggs of a collembolan, Sinella curviseta Brook (Lipovsly, 1954). All deutonymphs were preserved upon death or after at least 30 days without feeding. After adult emergence the deutonymphal cuticle was also mounted for study. Specimens were studied using a Wild M-20 phase contrast microscope. Drawings were made using a Wild drawing tube attached to the Microscope. Standard measurements (Robaux, 1974; Goff, et. al.,

1982) were made using the Wild M-20 with the drawing tube to combine the image of the mite and the working area of an Apple Graphics

Tablet connected to an Apple lie computer. Using BASIC language programs, written specifically to measure Trombidioid mites, measurements (i.e. setal length, segment length, etc.) were made on the graphics tablet and the data were stored on a floppy disk. Basic statistics (mean, range, standard deviation, etc.) were calculated by the program and the results printed. Review of Previous Classifications

Over the years a number of classifications for the Trombidioidea

have been proposed but the familial relationships remained to be

established. A comprehensive classification of these mites by

Oudemans (1923) considered all the then known parasitengones and

contrasted the Erythraeoidea (Apobolostigmata, without Claparede’s

organ) with the remaining terrestrial and aquatic taxa

(Engonostigmata, with Claparede's organ). He then divided the latter

group into Phanerostigmata, with terrestrial or aerial (on the

surface of aquatic habitats) larvae, and Calyptostigmata, with

swimming larvae. Although there were some inconsistencies in the

placement of the genera Calyptostoma, Hydrachna, and Piersigia,

Oudeman's thoughtful attempt has, unfortunately, been ignored by

subsequent workers.

The post-Oudemans tradition has been to isolate the aquatic

parasitengones. This has resulted in the study of water mites

developing independently of other mites, especially the terrestrial

Parasitengona. Vercammen-Grandjean (1973, 1974) and Feider (1979)

removed most of the aquatic Parasitengona to the Hydrachnogona. This was also followed by Krantz (1978) and Cook (1974) who placed the

10 11 water mites in the Hydrachnidia and Hydracarina, respectively.

Robaux (1973) ignored the aquatic Parasitengona. In recent classifications of these essentially terrestrial groups,

Vercammen-Grandjean (1973) presented a nearly complete list of the terrestrial parasitengone genera and families, Feider (1979) presented a dichotomous branching scheme, and Robaux (1973) figured a polychotomous scenario.

Feider's 1979 treatment of the Trombidioidea s. .s.. was based primarily on the "respiratory" system but his confusion of the trachea-like ducts of the supracoxal gland system with tracheae requires re-evaluation of his conclusions. His classification of the Trombidia proposes a number of new higher categories (i.e., six superfamilies and six new families). Most of his classification is a dichotomous branching diagram based on the respiratory opening, uropore, palptibial chaetotaxy, crista metopica and the number of trichobothria on the prodorsum of the post-larval instars. A cladistic analysis of his classification found it to be paraphyletic.

Robaux's (1973) polychotomous tree represents a consensus of the presently accepted classifications of terrestrial parasitengones (see also Krantz, 1978). Unfortunately, too few characters were used and the three characters in his classification are distributed in such a way as to make the problem of major groupings unresolvable in terms of synapomorphies. From this, I conclude that, while the traditional groups of terrestrial Parasitengona can be recognized on the basis of combinations of plesiomorphic and apomorphic features, the phylogenetic relationships of these groups are yet to be resolved. 12

Witte (1984) in a thoughtful paper on the evolution of

reproductive mechanisms in the Parasitengona suggested the

Hydrachnidia, Erythraeoidea and Trombidioidea _s. _1* could be defined

as monophyletic based on a study of the reproductive mechanisms in 14

taxa. He tentatively placed the calyptostomatids with the

Trombidioidea.

While most of the work on the Trombidioidea has been limited to

descriptions of new taxa, several authors have attempted to provide

usable classifications (Thor, 1935a; Thor & Willmann, 1947; Feider

1952, 1955a, 1959, 1979 and Vercammen-Grandjean, 1973). Many of the current suprageneric categories were proposed by Thor (1935a, b) and elaborated on by Thor and Willmann (1947) in their catalog and classification of known species. This and most works were based exclusively on the postlarval instar. Vercammen-Grandjean*s (1973) list of trombidioid genera and suprageneric categories was without definitions or explanaton of his classification. In somes cases he added genera while omitting other genera without explanation. Feider

(1952, 1955a, 1959, and 1979) proposed a number of classifications and frequently did not follow his own classification from one year to the next. In 1984 I visited the Univeristy "Al. I. Cuza1' to study the Feider Collection, but was able to find only a few of his trombidioid specimens, casting further doubt on the value of his trombidioid classifications.

As mentioned above, nomenclatural problems associated with the

Trombidioidea have resulted in considerable confusion and could threaten the stability of existing classifications. For example, the identity of the type species of the genus Trombididum. T. holosericeum (L.), has been discussed by a number of authors (Treat,

1975, Vercammen-Grandjean, 1973, Southcott, 1961) without resolution.

The consensus is to accept Thor and Willmann's (1947) exhaustive, and somewhat speculative, synonomy of T. holosericeum. with the type species only poorly described and without a described larval instar.

Thor and Willmann (1947) indicated Atomus parasiticus DeGeer as the possible larval instar of X* holosericeum. Robaux (1974) synonymized

Atomus with Trombidium without comment while Vercammen-Grandjean et. al. (1977) proposed the synonomy of A_. parasiticus with T. holosericeum on the basis of the similarity of T. hyperi

Vercammen-Grandjean et. al. larva and deutonymph to the two aforementioned species. While I agree with Vercammen-Grandjean et. al. (1977), based on circumstantial evidence, it should be pointed out that jA. parasiticus is unrecognizable in the original description

(Southcott, 1961). Only comparative study of series of specimens including European Trombidium. with subsequent designation of a neotype, can resolve the problem. For the present, I have used larval specimens reared from adults collected in Great Britian as my working concept of the larva of Trombidium.

The best example of confusion is the genus Microtrombidium, the type species of which was originally described from an adult, T. pusillum Hermann. The larva was unknown or at least undescribed.

This has not stopped authors from speculating on what the larva might be and from ascribing larval species to the genus. There are approximately 230 species currently assigned to the genus of which 17 14 are larval forms and nine have been described since 1946, none correlated with postlarval instars. My work indicates all of these larval species are assignable to other genera (nominate and new).

For this study I have used the larva of Microtrombidium italicum

Berlese reared from field collected adults in Italy in 1984 as my working concept of Microtrombidium. Other nomenclatural problems result from the different larval and post-larval classifications where the adult of one species can be placed in one genus while its larval form, usually described separately, is placed in a different genus and sometimes different subfamily. This has occurred with the genus Valgothrombium Willmann which was originally described from an adult but with the larval forms placed in Feiderium Vercammen-

Grandjean and its subgenus Parafeiderium Vercammen- Grandjean and

Cochrane. These problems emphasize the importance of a detailed comparative study, correlating of larval and adult forms, and study of European material. EXTERNAL MORPHOLOGY AND DISCUSSION

OF CHARACTERS

The discussion of the external morphology and characters in the groups studied is divided into larval and postlarval instars. Each section is subdivided into a discussion of major groups of characters. In the following discussion I will use Trombidioidea to refer to the following families as a unit: Podothrombiidae,

Trombidiidae, Neothrombiidae, Calothrombiidae, Eutrombidiidae and

Microtrombidiidae.

LARVA

The larva is the hexapod parasitic/dispersal stage of the

Parasitengona life cycle. The morphology of the larva is designed to enable it to find a host. It is the larva which is most similar to other Acariformes, thus allowing comparisons with other parasitengone and prostigmatid mites.

Gnathosoma

The gnathosoma, a pseudotagma, is the region anterior to the

15 16 circumcapitular furrow with two pair of appendages: chelicerae and palps.

The acariform chelicerae superficially consists of three segments: the basal segment, second segment with two setae, including the fixed digit and distal segment, the movable digit (pretarsus).

In the Anystidae (and most Anystina) the fixed digit is reduced or lost and the movable digit is bladelike and the two cheliceral setae are retained. All larval Parasitengona have lost the fixed digit and cheliceral setae but retain the bladelike movable digit.

Most Trorabidiina have one subapical tooth on the cheliceral blade.

The Chyzeriidae have a large bladelike movable digit with numerous small teeth, while the Trorabiculidae have independently evolved a variety of forms (Goff, et. al., 1982).

The primitive palp of the Acariformes consists of five movable segments (trochanter, femur, genu, tibia, tarsus) (Kethley, 1982).

The coxae are fused to form the gnathobase. This primitive condition is retained in Anystidae (outgroup), the water mites (Prasad & Cook,

1972), Erythraeiodea and the families Trombellidae, Chyzeriidae,

Johnstonianidae, Trombiculidae and Neotrorabidiidae. The palpal trochanter has been lost or fused with the palpal femur in the

Calyptostomatidae and Trombidioidea. Secondary fusion of palpal segments is rare in the Parasitengona, but has occurred independently in the Tanaupodidae (Eothrombium and Polydiscia) and

Aethethrombidiinae (Neothrombiidae) where the outer surface of the palpal femur and genu has been fused.

The palpal trochanter is without setae in the Anystidae. In the 17

Parasitengona only Balaustium (Erythraeidae) has a seta on the palpal

trochanter.

The setation of the palpal femur is variable in Anystidae. The

primitive genus Anystis is hypertrichous, with up to 5 setae, and the

derived Tarsotomus has a single seta. Most parasitengone genera have

a single dorsal barbed seta. Neotrichy occurs in two related

erythraeid genera (Abrolophus and Hauptmannia) and in Eothrombium

(Tanaupodidae). In the Trombidiidae, Microtrombidiidae and

Neothrombiidae this seta is reduced or lost in most genera (see

Figure 1). In some Eutrombidiidae, Calothrombiidae and

Microtrombidiidae the seta is small and thus frequently overlooked in

descriptions.

The palpal genu of the primitive Anystis is also hypertrichous while Tarsotomus has a single barbed seta. The chaetotaxy of the

parasitengone genu is more variable. All Erythraeidae have one

barbed setae on the palpal genu, except Balaustium, Hauptmannia,

Abrolophus and some Leptus. which have two. The water mites also

have two setae on the palpal genu, while the Trombiculidae,

Trombellidae, Chyzeriidae and Johnstonianidae have a single, well developed barbed seta. In the Trombidiidae and Neothrombiidae all genera have lost this seta. In the Microtrombidiidae and

Eutrombidiidae this seta is frequently retained but is usually reduced in size. Calothrombiidae retain both setae. Podothrombium

(Podothrombiidae) is the only trombidioid which retains a well developed barbed seta on the palpal femur and genu.

The setation of the palpal tibia is consistent throughout the 18

FIGURE 1. Dorsal idiosoma of Trombidium holosericeum L.

(Trombidiidae)

AM= Anteriomedian setae; AL= Anteriolateral setae; PL=

posteriolateral setae, S= prodorsal trichobothria. AM

AL

PL

FIGURE 1. Dorsal idiosoma of Trombidium holosericeum L.

(Trombidiidae) 20

Parasitengona and Anystidae, with all genera examined having three setae. In some Eutrombidiidae and Neothrombiidae the dorsal seta is hypertrophied (i.e. spinelike) and is frequently located adjacent to the palpal tibial claw. The presence or absence of barbs are frequently important in species descriptions, but are of little importance in generic diagnosis.

The palpal tibial claw, a hypertrophied seta, is well developed in the Parasitengona. In the Anystidae the palpal tibial claw has several forms, from three small "claws'1 in Anystis to single clawlike seta with barbs in the Erythracarinae.

The palpal tibial claw in the Parasitengona is usually two-tined. In the Chyzeriinae (Chyzeriidae) most genera have well developed three-tined claws. Some genera of Trombiculidae retain the two tined claw (Eutrombicula) but other genera have developed multi-tined claws (Goff et. al., 1982). In the Calyptostomatidae the palpal tibial claw is reduced to a small stub and in Polydiscia

(Tanaupodidae) it is lost.

The chaetotaxy of the palpal tarsus has been used extensively in the classification of the Trombiculidae (Vercammen-Grandjean,

Langston and Audy, 1973; Brennan and Goff, 1977). While there appears to be consistent numbers of setae on the palpal tarsus in some groups (Chyzeriinae with 8 setae; Johnstonianidae with 7 setae and the Trombellidae with 8-9 setae) variation often makes it difficult to determine trends in many groups. In the Trombidioidea most of the setae and palpal tarsus are reduced, making it difficult to see, let alone make accurate setal counts. In the Anystidae the 21 number of setae on the palpal tarsus is variable (8-10 in Anystis and

4-6 in Tarsotomus). All Anystids and Parasitengona examined have at least one solenidion on the palpal tarsus. In the Anystidae there is a distal eupathid which is retained in the Erythraeoidea and

Calyptostomatoidea and is present in only a few non-erythraeoid

Parasitengona. It appears to be absent in the Trombidioidea, but the reduced tarsal setation makes it difficult to confirm the presence or absence of these setae. In the Chyzeriidae there is a terminal pair of short, nude setae which may be eupathidia. There are eupathidial setae in some genera of Trombellidae. No eupathidial setae were observed in the Neotrombidiidae but Lindquist and Vercammen-Grandjean

(1971) reported eupathidial setae on the palpal tarsus. The palpal tarsus*of the Johnstonianidae terminates in a spine, which appears to be non-setal in origin. This structure is absent in Tanaupodidae but there are one to four terminal eupathidia.

The subcapitulum is the ventral portion of the gnathosoma composed proximally of the fused palpal coxae and the lateral lips distally.

In the Acariformes there are up to three pair of adoral setae which are usually distinct from the subcapitular setae. Grandjean

(1943) referred to the distal two pair of setae on the subcapitulum in the larva of Anystis as "poils adoraux" and noted the third pair was added in the deutonymph. Evidence in the Parasitengona suggest this interpretation is, if not correct, at least parsimonious. I will follow Grandjean (1943) and call the distal two pairs of subcapitular setae the adoral setae (orl, or2). This leaves the 22

third or proximal pair to be subcapitular setae (scl).

The Anystidae and primitive Parasitengona have three pairs of

setae on the distal third of the subcapitulum, and have been given

various names depending on interpretation. Newell (1957) avoided

determining homologies by calling them protorostrals (protorostral 1,

protorostral 2 and protorostral 3). This was followed by Lindquist

and Vercammen-Grandjean (1971) in their revision of Neotrombidiidae;

most other authors ignore these setae.

The anterior adoral setae (orl) correspond, at least

topographically, to the galeala (Goff, et. al, 1982.) of the

Trombiculidae. All Erythraeoidea examined have three pairs of

subcapitular setae. In the Smarididae and a few Erythraeidae the

second pair of adoral setae (or2) are reduced. The difference

between the subcapitular (scl) and adoral (orl and or2) setae are not

always obvious in the Erythraeidae but in a few genera (Callidosoma,

Charletonia and Hauptmannia) the subcapitular setae are

morphologically distinct and separated from the adoral setae. An

undescribed species of Abrolophus (Erythraeidae) has a neotrichous addition of a pair of setae on the gnathobase.

The subcapitular setae (scl) within the Parasitengona have a

variety of shapes and positions. They are usually located distal to

the palpal articulation but in Trombellidae and Trombiculidae they are basal. In addition, they are separate from the adoral setae and sometimes assume the morphology similar to nearby setae on the coxal fields.

In the non-erythraeioid parasitengona, the maximum subcapitular 23 compliment of three pairs of setae (orl,or2 and scl) occurs in

Chyzeriidae, Tanaupodidae and some Trombellidae with all others losing or2 and retaining only orl and scl. In the Erythraeoidea and

Trombellidae (Trombella. Durenia), Tanaupodidae (Eothrombium),

Chyzeriidae (Chyzeria. Nothotrombicula) the or2 setae are very small or spinelike. The Trombidioidea, Johnstonianidae, Trombiculidae and

Neotrombidiidae have all lost one pair of adoral setae which is assumed to be the second pair (or2). Moser and Vercammen-Grandjean

(1979) illustrated a pair of tiny setae between the adoral and subcapitulae setae in Megophthrombium but examination of the holotype and numerous other specimens failed to reveal their presence nor were they found on any other Trombidioidea. When only two pairs of setae are present on the subcapitulum, the distal pair is assumed to correspond to the adoral setae and the proximal pair are the subcapitular setae. The scl setae take on a variety of shapes from setiform (Neothrombiidae and Calothrombiidae) to palmate (most

Microtrombidiidae and Trombidiidae) or spinelike (Eutrombidiidae and some Microtrombidiidae) (see Figure 2). These setae are often similar to the setae on the coxal fields in the Eutrombidiidae.

Primitively there is a supracoxal seta (e) located on the gnathobase. This seta is present in Erythraeoidea, but is absent in the Johnstonianidae, Trombiculidae, Neothrombiidae, Eutrombidiiae and most Microtrombidiidae, water mites and Calyptostomatidae. In the

Trombidiidae and Calothrombiidae they are located on the distal edge of the gnathobase. 24

ff 17 B

FIGURE 2. Subcapitular Setae of Larvae.

A. Podothrombium (Podothrombiidae), Neothrombium

(Neothrombiidae); B. Trombidium (Trombidiidae); C.

Cercothrombium (Microtrombidiidae); D. Campylothrombium

(Microtrombidiidae); E.Eutromidium (Eutrombididae); F.

Valgothrombium (Microtrombidiidae). 25

Idiosoma

The idiosoma.of the Acariform mites is divided into two regions: podosoma and opisthosoma, which are separated by the disjugal furrow.

In the Parasitengona the disjugal furrow is not evident but the division between the propodosoraa and opisthosoma is recognized dorsally as being between the prodorsal sclerite (scutum) and first row of idiosomal setae (C-row).

The prodorsum of the acariform mite is defined as the dorsal portion of the idiosoma anterior to the sejugal furrow. The prodorsum is recognized by the presence of the prodorsal sclerite.

Primitively this sclerite had six pairs of setae which have been given different names in almost every acariform group.

The Parasitengona prodorsum maximally has four pairs of setae

(including at least one pair of prodorsal trichobothria and three pairs of normal setae) in the primitive condition. Some neotrichous

Erythraeidae have six pairs. These fundamental setae are anteromedial (AM) which may be trichobothrial in some genera; anterolateral (AL); toward the posterior of the sclerite are the posterolaterals (PL) and a pair of trichobothria (S) (see Figure 1).

The number of prodorsal setae remains relatively constant in the

Parasitengona. In the Trombiculidae and some Johnstonianidae one or both AM setae may be lost.

While it is not possible to trace the progression of setae on the prodorsum, certain assumptions have been made about the origin and subsequent position of setae. First, the anterior pair of 26

trichobothria in the primitive Parasitengona (all Erythraeidae, some

Johnstonianidae, Tanaupodidae, some Chyzeriidae) are equivalent to

the AM setae of the more derived Parasitengona. This is supported in

the Johnstonianidae where the same pair of setae may be

trichobothrial in one genus and normally developed in another closely

related genus, representing at least topographical homology. In the

Neotrombidiidae, Trombellidae, Trombiculidae and Trombidioidea the

bothridial bases of the AM setae have been lost.

The Trombidioidea are very conservative in modifications of the

prodorsal setae with no losses and only occasional modifications in setal morphology and position. In one species of Trombidium the PL setae have similar morphology to modified coxal field setae. Changes in relative position of setae have occurred in a few genera and in some suprageneric groups. In Dinothrombium (Trombidiidae) the distance between the PL setae is less than the distance between the trichobothrial bases (SB); this is a very useful autapomorphy for the genus. In the Neothrombiinae (Neothrombiidae) the trichobothria have migrated to the posterior end of the prodorsal sclerite displacing the PL setae anteriorly.

The morphology of the prodorsal trichobothria is relatively conservative, being usually filiform, in most groups, with or without setules. In a few groups the prodorsal trichobothria exhibit varying degrees of expansion (Trombiculidae) while others (Trombidiidae,

Neothrombiidae and Eutrombidiidae) expanded trichobothria have only occasionally independently evolved.

The shape of the prodorsal sclerite is relatively constant in 27

most groups with the Johnstonianidae, Trombellidae and

Neotrombidiidae having triangular prodorsal sclerites while the

Chyzeriidae and Trombiculidae have rectangular prodorsal sclerites.

The Trombidioidea have rounded or rectangular sclerites which cover

most of the prodorsum. In many genera in the Microtrombidiidae and

Eurtrorabidiidae the prodorsal sclerite extends over the anterior end

of the prodorsura to form what have been called anterolateral

shoulders (=stolascutum of Moser and Vercammen-Grandjean, 1979). In

Eothrombium (Tanaupodidae) the anterior portion of the sclerite with

the AM setae is separated from the rest of the sclerite, a primitive

character shared with Anystis and some water mites.

The setae on the idiosoma posterior to the disjugal furrow has

never been comparatively studied and generally given in descriptions

as total number of setae with little attempt to identify setal rows.

The dorsal idiosomal chaetotaxy presents a difficult problem in

analysis because of the large number of variations which occur. By

comparing the position of setae in parasitengone larvae with other acariform mites and looking at the location of idiosomal lyrifissures

(cupules), present in the primitive Parasitengona (most erythraeids,

Tanaupodidae, water mites and some trombellids), it is possible to homologize at least some dorsal idiosomal setae.

The Anystidae and primitive Parasitengona possess the dorsal

proprioceptors (cupules) in the cuticle. They are also visible in

some of the postlarval erythraeids. In the Anystidae, Trombellidae,

Tanaupodidae, Johnstonianidae the cupules ia, im, ip, and ih are

present. Cupules are lost in the Trombiculidae, Neotrombidiidae and 28

Trombidioidea.

The opisthosoma of the primitive larval Acariformes consists of

six parallel rows of setal pairs 4-2-2-3-3-3 designated C, D, E, F, H

and PS respectively. In Anystis the number of setae appears to be to

be 2-2-2-2-2-3; while other anystids are hypertrichous they still

retain the original setae which are usually longer than the added

setae.

The Erythraeoidea and Calyptostomtoidea species examined are

hypertrichous in all rows with the rows becoming obscured

posteriorly. In Tanaupodidae the basic number of setal pairs is

2-2-2-2-2 with the PS series having one or two pairs of setae. This

same pattern appears in the lower water mites. The Chyzeriidae,

Trombellidae and Johnstonianidae tend to have three pair of setae in

each row (C, D, E, F, and H). The Trombiculidae are generally hypertrichous dorsally. In the Trombidioidea secondary reductions in

this pattern can occur in rows E and F (see Figure 1). Rows H and PS appear to have one pair of setae each, which are frequently longer

than the other dorsal idiosomal setae.

In the Trombidioidea there is at least one additional dorsal

sclerite (scutellura) which is found mid-dorsally in the C-row bearing

the first pair of setae (cl). In Eutrombidiidae and

Microtrombidiidae the first pair of setae in the D-row are on large

circular sclerites. In one genus in the Manriquiini

(Microtrombidiidae) the sclerites surrounding the H and PS setae have fused to form what has been called the pygosomal plate (Goff, et. al., 1982). Rows tend to be obscured at the H and PS level. 29

LEG CHAETOTAXY

The acariform leg consists of six articulating segments beyond

the coxa (trochanter, femur, genu, tibia, tarsus, pretarsus). The

coxae are fused to the venter of idiosoma. The pretarsus, distal to

the tarsus, is a separate segment and primitively consists of paired

claws and padlike empodium.

While the legs are obviously for locomotion, they are also the

primary structures used to locate and contact host or prey, requiring

a wide array of tactile and cheraoreceptor setae. The arrangement,

number and morphology of the setae (tactile and chemosensory) have

certain patterns in position and number which are usually fixed for a

given taxon. Some setal patterns can be given as formulas which are

useful in systematic analysis (Grandjean, 1935). In the Acariformes

formulas have been used extensively in the Oribatida, but have been

applied only to the Smarididae and Erythraeidae by Grandjean (1947a,

b).

Over the years different workers (Southcott, 1961, Newell, 1957,

Wharton, et. al., 1951, Robaux, 1967b, 1974, Grandjean, 1947) have

developed different terminologies for the leg setae in most

parasitengone groups. Recently, there has been an effort to

integrate some of these terminologies (Goff, et. al., 1982). Table 1

summarizes the different leg setae terminologies for the terrestrial

Parasitengona. I will follow the terminology as applied by Robaux

(1967b, 1974) and Grandjean (1935, 1947). 30

Table 1. Leg Nomenclature for Larval Terrestrial Parasitengona.

Trad) icul idea Troriildlldae (Brennan 1 Goff, Looais, (hobauB, Grandjean, Erythraeoidea Johnstonianidae Varcaraen-Grandjean, Mullen I Vercaaaen (Southcott. 1961) (Newell. 19S7) ■harton) Grandjean)

taispeclalited setae scobalse nonal setae nonal, ordinary, branched n (branched) scobofaairalae, setae, barbed setae

Specialized > c u h t

tolanidia" spiaofaaorala solanldlon Jt Sj faaorala 9 theta

Genu solenidia splnoganiala solanidion 3. S 3 gaiuala • sigaa

atcroseta sastigiala vestigial sets aicroapur, nicrogenuala k

Tibia solenidia splnotlblala solanidion S. tiblala ¥> pal proximal; solanidion 4 , S 4 digital

atcroseta vestlglala vestigial seta adcroapur, adcrotlbiala k

eofanion teu CDOfMnsls a a p s i i o n seta of solanidion 4> S 4

l W a solenoldsls solanidion 3 , S, tarsala, I, II u onega (solanotarsalt) (Tarsus I) soleAidion ,, Sj (Tersus 11)

■ l C I D M U t a i l u s (fassls) faaultm 1 , 2 faaulus I, II a, faaulus aicrospur, aicrotarsala

eupathidia

A. distal "sansills alongside distivantral cipathid pretarsala f seta pedicle" of tarsi or "eupathid at 0.92 pv*'

1 . proximal spinotarsala dorsal eipathid of subtemlnala f seta tarsi or "eipathid at O.nd"

aapenion sets

A. c n ^ a n a l a (?) coapanlon seta of S,. S 2 . S 4 1

*. coeipanlon seta of psrasifeterminala f teta dorsal eupathid

modified sctse on D, liphotrix T s n u s III N, V, scopa

long whiplike sets sens i 11m bothridia aastisetae usually nude (trioobothriala, asstala bothrisls)

other modified vdified scobslee e c u ("taetalae") 31

In the Acariformes all tactile (normal setae) including

mastiseta, trichobothria, microsetae and famuli) and presumed

chemoreceptors (eupathida) contain optically active substance, called

actinopilin, which causes the seta to become birefringent when viewed

under polarized light (Grandjean, 1935). Solenidia, which are

strictly chemoreceptors, are not optically active.

Tactile setae (normal setae including mastisetae and

trichobothria), regardless of the external morphology, consist of

cuticular elements, sensory neuron and sheath cells (Mclver, 1975).

The sensory neuron (dendrite) is attached to the base of the seta and

when the seta moves the neuron is stimulated. Haupt and Coineau

(1975) found that, despite the external morphological appearance,

there was no difference between trichobothria and normal setae in

Caeculidae other than the amount and type of movement which was dictated by the shape of the setal base.

Famului and eupathidia are modified normal setae with a hollow core, usually a blunt tip and actinopilin. The function of these setae are unknown. These are considered modified setae and will be covered separately under Specialized Setae.

Our understanding of the modern leg chaetotaxy in the

Acariformes is based on Grandjean's (1940) concept of verticils

(reviewed by Norton, 1977 and Andre, 1979). This assumes the legs of the Acari are composed of numerous small annuli, each supplied with a set of setae (verticil) and the modern chaetotome of a leg is the result of additions and/or losses to the basic number of setae in the primitive verticils. Verticils in the Parasitengona can be 32 recognized on distal femur, genu and tibia. The verticils are more difficult to see on the tarsus because of setal losses and realignment of setae.

There is some question as to how many setae are in the primitive verticil. Van der Hammen (1980) suggests the primitive verticil number as 8 using the notation: d, Id", Id’, 1", 1', lv", lv', and v and Grandjean describes the basic verticil as five setae (v",v',1",1" and d). Based on my examination of the anystoids and the

Parasitengona I will follow Grandjean. From the basic verticil of five setae most chaetotaxies can be derived and designations can be given to individual setae.

Setae are identified by their morphology and position, often without full understanding of their function or origin. The legs of the larval Parasitengona provide a rich array of setal types, patterns and morphologies which provide numerous characters. In this work tactile setae will be called normal setae and the specialized setae will be referred to by their respective names (famulus, solenidia, eupathidia). Leg trichobothria in the Parasitengona are found only on members of the family Smarididae and will not be discussed.

Most abundant are the normal or tactile setae. These can be of several forms: barbed, nude (without barbs), mastisetae, and various other shapes (i.e. lophotrix and scopa). With few exceptionsthese setae have been ignored by most workers. Papers by

Vercammen-Grandjean and those by Pierre Robaux at least provide counts of setae on each leg segment. The positions of many of these 33 setae are constant and an attempt has been made to develop formulas and point out patterns.

The coxa is the basal segment of the leg and in the Acariformes it is immobile (i.e., fused with the ventral idiosoma). Generally, the coxal field and adjacent intercoxal area have been mislabeled as the coxa and sternum, respectively (by the chigger workers). In this work I will use the terms coxal field and intercoxal area.

The setae in the coxal field and intercoxal area are identified by their position relative to the adjacent coxal field and on the basis of a maximum of three pair of setae for each coxal field and intercoxal area. The setae in coxal field leg I and adjacent intercoxal area would be designated 1A for the seta closest to the saggital line and IB and 1C occupying the outermost position. It is assumed the outermost setae are lost first, which is supported by observations in different parasitengone genera.

The number of setae in the Trombidioidea is relatively constant within each of the families, but some setae have been modified in certain groups. Primitively, the Trombidioidea have two setae on coxal field leg I. These result from seta 1A migrating onto the coxal field from the intercoxal area and seta 1C being lost from the coxal field. Setae on coxal field I are designated 1A and IB with 1A derived from the intercoxal area. In one genus, Milliotrombidium

(Eutrombidiidae), seta 1A is located on the edge of the coxal field and in some specimens appears to be in the intercoxal area. In the family Johnstonianidae seta 1A is located on a small sclerite attached to the inner side of the coxal field I. 34

Seta IB is derived from the original coxal field setae. This is

' supported by the observation that intercoxal setae IB, 2B and 3B

frequently take on the same form as, for example, in the

Eutrombidiinae (Eutrombidiidae) these setae are stout and bifid while

setae 1A and 3A are setiform.

On coxal field leg II seta 2A is apparently lost leaving seta

2B. In the Trombidiidae, Calothrombiidae and the Neothrombiinae

(Neothrombiidae) a synapomorphic second seta (designated 2C) appears

independently in each group and can take various positions on the

coxal field (see Figure 3). In the Aethethrombidiinae and one genus

in the Neothrombiinae ( both Neothrombiidae) there is a tendency for

coxal field leg II to be neotrichous.

Coxal field leg III exhibits little variation with seta 3A

present in all but a few genera. Seta 3B usually takes the same form

as setae IB and 2B.

All Parasitengona, except Calyptostomatidae and some

Erythraeoidea, have only a single branched seta on the trochanter

legs I, II and III. Southcott (1982) mentioned this as a possible

character in the Trombellidae, but it provides no useful information

for classification purposes.

The presence of a subdivided femur (basifemur and telofemur) is

primitive. The femur of all active postlarval instars of

Parasitengona is subdivided while the femur is divided in only some

larvae. The femur is divided in all larval Erythraeoidea, the water

mites and the Calyptostomatidae (Calyptostomatoidea). In the

remaining Parasitengona only the Chyzeriidae have a divided femur 35

FIGURE 3. Ventral view of Trombidium holosericeum L. (Trombidiidae)

orl= adoral setae; scl= subcapitular setae; e= gnathosomal

supracoxal setae; el= supracoxal setae at leg I; 1A, IB

setae on coxal field leg I; 2B, 2C setae on coxal field leg

II •c

1ft

u

tc

SB

FIGURE 3. Ventral view of Trombidium holosericeum L. (Trombidiidae) 37

in all species. The distribution of divided femora is variable in

the Neotrombidiidae, Johnstonianidae and Trombiculidae where some

taxa have all femora divided, others have only leg I divided and

others only legs I and II. Some authors (Southcott, 1982; Brennan

and Goff, 1977) have used the divided femur (plesioraorphic) as an

important character in determining group placement, when it is the

fusion of the femur that is derived and is not really very useful

because it has occurred independently in many trombidioid groups.

Femora of legs I, II and III are fused in all larvae in the

Trombidioidea.

In Anystis the maximum femoral chaetome (combining the basifemur and telofemur) is 9-8-8 for legs I, II, and III, respectively. From this arrangement all femoral chaetomes of the Parasitengona can be derived. In Anystis the maximum setal complement for the basifemur is 4 setae (d, 1’, 1", bv) and 5 setae on the telofemur (d, 1’, 1", v', v") on leg I. The setae of the basifemur comprise a single verticil and this usually remains distinct when the femur is fused.

In the Erythraeoidea there can be reduction of the basifemur to only one or two setae while the telofemur component remains unchanged. In the Chyzeriidae and many Johnstonianidae the femur remains divided, but there has been a loss of the "d" and a ventral seta on the basifemur leaving only the "bv" seta, which is also present in all other Parasitengona.

The water mites and Tanaupoidea have femoral chaetomes of

10-10-10 and 9-9-9 respectively. When compared with the anystids the femur must be considered neotrichous, but when compared with most 38

FIGURE 4. Legs of Trombidium holoserlceum L. (Trombidiidae)

k- microseta tibia leg I and genu legs I and II; (j) -

solenidia tarsus legs I and II; (J ■ solenidia genu legs I,

II and III; i p ■ solenidia tibia lege I and II; € ■

famulus tarsus legs I and II; N ■ lophotrix; V « scopa;

« eupathid 39

FIGURE A. Legs of Trombidium holosericeum L. (Trombidiidae) 40

Erythraeina the setal count represents a reduction in setae. The

Trombidioidea have1 a maximum of 6-5-4 on legs I, II and III, respectively. In most of the Trombidioidea the basiventral seta "bv" is the only remnant of the basifemur verticil (see Figure 4). Seta

"bv" varies in form from a short nude seta to a long and multibranched, but is usually distinct from the other setae on the femur. In some Microtrombidiidae the dorsal seta (d) is reduced, without barbs and has migrated to the extreme distodorsal end of the segment.

The chaetotaxy of the femur leg I in Neothrombiidae,

Calothrombiidae, Eutrorabidiidae and Microtrombidiidae is identical with six setae ( d, 1’, 1", v 1, v", bv), while Trorabidiidae and

Podothrombiidae have five setae (d, l1, v ’,v", bv) on the femur of leg I. Leg II differs between the six families. In the

Trombidiidae, Podothrombiidae and Neothrombiidae setae 1" and v" are lost leaving d,l,,v', and bv. The Microtrombidiidae,

Calothrombiidae, Eutrombidiidae have a different arrangement with leg

II having 5 setae ( d, 1’, v ’, v", and bv). All have the same chaetotaxy for femur leg III. The chaetome of the telofemur remains unchanged with five setae in the other Parasitengona.

The genu in the Trombidioidea is short compared with the

Erythraeidae and Anystidae and as a result it has a reduced chaetome.

In the Erythraeoidea the genu contain two or three verticils of four setae each (1', 1", v', v", ml', ml", mv', mv", pi', pi", pv1, pv") in the Callidosoraatinae and 8-8-8 (1', 1", v', v", pi1, pi", pv', pv") in the Erythraeinae, while in the Anystidae the genus Anystis 41

has two verticils of five setae each (d, 1’, 1", v 1, v", d2, pi'.pr'.pv*, pv").

The Chyzeriidae, Trombellidae, Neotrombidiidae and Trombidioidea

all have a constant number of four setae (1’, 1", v ’, v") on the genu

of leg I. On the genua of legs II and III there have been

independent losses of one or two setae in several families. In the

Neothrombiidae (except in the neotrichous genus Acridothrombium).

Calothrombiidae, Eutrombidiidae and Microtrombidiidae the loss of two

ventral setae is a synaporaorphy uniting the four families. The

Trombiculidae and Trombidiidae have independently lost a ventral

seta.

The chaetotaxy of the tibia presents a problem because the

number of setae is variable within the Parasitengona, ranging from 18

in some Erythraeidae to 5 or 6 in most Trombidina. Within the

Anystidae the range is from 8 to 18. The lower water mites retain 10

setae (1', 1", v f, v", v2, plf, pi", pv', pv", d) although these

positions do not correspond to the verticils of other Parasitengona,

because of what appears to be an extra ventral seta.

In the Tanaupodidae (Eothrombium) the "pv" seta is lost from the proximal verticil and the dorsal seta "d" from the proximal verticil migrates anteriorly. The other posibility is the seta (d) is lost posteriorly and then gained de novo in the anterior verticil.

Regardless of the mechanism the result is a total of 9 setae (l1, 1", v 1, v", d, pi1, pi", pv', pv") on the tibia of leg I, II and III.

In the remaining terrestrial Parasitengona (Trombidioidea,

Chyzeriidae, Trombellidae and Neotrombidiidae) there has been a 42 reduction in the tibial I chaetome of one to four setae. In all genera examined a posterior ventral seta is missing, leaving only three setae on the venter of the tibia. These setae are the two anterior ventral setae "(v)" and one posteroventral "pv" seta. The tibia I chaetome of the Trombidiidae is more reduced with five setae present, resulting from the loss of the remaining "pv" seta (1*, 1", v \ v", d).

The tibiae of legs II and III in all Trombidioidea all have five setae (d, 1’, 1”, v', v"). It is not possible to be sure of the exact origin of each ventral seta because of movement in the setae but since it is the pv seta which appears to be lost on leg I it seems likely that the same seta is lost on legs II and III also. In some Eutrombidiinae (Eutrombidiidae) the pv seta is lost, again leaving only two ventral setae.

The Trombiculidae and some Neotrombidiidae still have eight setae (l1, 1", v', v", pv, pi', pi", d) on tibia leg I resulting from the addition or retention of one or both of the "pi" series or even a "pv". The Johnstonianidae, Chyzeriidae and Ralphaudyana have d,

1', 1", v', v", and pv. Tibia legs II and III each have six setae.

The chaetotaxy of the tarsus is the most difficult of all the leg segments to study because of the number of setae and variation in the number of setae within species. In this study it was not possible to assign signatures to most of the setae on the tarsus because of the variation in the location of setae and lack of good specimens in all groups. Even though setal signatures were not assigned to the tarsal setae in the Trombidioidea there appeared to 43

be a relatively constant range on setae (18-14-13) which varied two

or three setae within species and even on individuals of the same

species.

In the Microtrombidiidae, Eutrorabidiidae, Calothrombiidae and

Trombidiidae, two distal normal setae on tarsus III have been

modified into structures of unknown function, resulting in reduction

to the pretarsus and, in some genera, a modification of the tarsus

and occasionally the tibia. These two setae have been called the

lophotrix ("a long, strong, dichotomus, and abundantly branch

ciliated plume") and scopa ("a thick little brush") (Mullen and

Vercammen-Gradjean, 1978) and given the signature "D" and "N"

respectively (Robaux, 1974, 1977).

The lophotrix is a distoventral normal seta that has displaced

the pretarsus at the terminus of tarsus leg III while being modified

into a variety of forms. This migration and modification is best

developed in the Microtrombidiidae and Eutrombidiidae but has

occurred independently in the Trombidiidae and Calothrombiidae but

not in the Neothrombiidae or Podothrombiidae. The original

definition by Mullen and Vercaramen-Grandjean (1978) referred to a

long dichotomous seta ('Y' shape) in Megophthrombium. I will expand the definition of Mullen and Vercammen-Grandjean (1978) to cover all

the variations found in Trombidiidae, Calothrombiidae, Eutrombidiidae and Microtrombidiidae.

Primitively, the lophotrix was setiform and located distoventrally. In the less derived taxa (Trombidiidae and

Calothrombiidae) the lophotrix is still in the ventral position. In 44

FIGURE 5. Lophotrix Setae from Tarsus Leg III.

A. Manriquia. Pygothrombidium (Microtrombidiidae); B.

Tricohotrombidium (Microtrombidiidae); C. Ettmuelleria

(Microtrombidiidae); D. Microtrombidium

(Microtrombidiidae); E. Dictotrombidium (Calothrombiidae);

F. Hoplothrombium (Eutrombidiidae); G. Eutrombidium

(Eutrombidiidae); H. Campylothrombium (Microtrombidiidae);

I. Platythrombidium (Microtrombidiidae) t v H

FIGURE 5. Lophotrix Setae from Tarsus Leg 46 the Microtrombidiidae there are three basic forms for the lophotrix: one has one or two long basal barbs (most of the Cercothrombiinae and

Feideriinae); the second has three to five or more long barbs and includes most of the Eutrombidiinae (Eutrombidiidae); and the third form is more like an enlarged seta and may have one long distal barb

(this includes most of the Microtrombidiinae) (see Figure 5). In the

Trombidioidea when the lophotrix and scopa are not developed tarsus

III has the normal compliment of 13 setae.

Tarsus III may be further modified to the condition where the segment appears to be bifurcate, with the reduced pretarsus on one tip and the lophotrix on the other (i.e. Hoplothrombium.

Cryptothrombidium and Milliotrombidium). This condition has occurred twice independently in the Eutrombidiidae with the Milliotrombidium

(Eutrombidiinae) and in the Hoplothrombiinae (Eutrombidiidae).

The second seta, scopa, is located on the paraxial side of tarsus leg III just proximal to the modified paraxial pretarsal claw and is also a modified normal seta. I will use the definiton of

Mullen and Vercammen-Grandjean (1978). Usually when the lophotrix is present the scopa is also present. The scopa in Milliotrombidium is slightly thickened and Atopothrombidium townsendi (Dumbelton), while in the closely related Hoplothrombiinae it is a normal seta. The scopa is bifurcate in Calothrombiidae and not developed in

Neothrombiidae.

The pretarsus of the Acariformes is the terminal segment of each leg and bears paired claws (olf, ol") and a padlike empodium (oc).

Primitively, the parasitengone pretarsus has paired claws and a 47 clawlike erapodiura. Since the empodiura and claws are quite similar, it is usually assumed that if only a single pair or structures are present then the empodiura is lost and if only a single clawlike structure is present then the paired claws are lost. One must be careful in making this assumption because some Microtrombidiidae have lost the antaxial claw leaving the paraxial claw and clawlike erapodiura although with careful examination it is possible to determine which structure is absent and there will frequently be a minute portion of the base of the antiaxial claw still in the pretarsus.

In the Parasitengona the paired claws and clawlike empodium are present in most groups although they may be reduced in some genera.

In the Trombiculoidea some genera have lost the paired claws or the empodium.

In most groups there are at least some species which have modified or reduced the paraxial claw usually in association with the modification of two normal setae (lophotrix and scopa).

Atractothrombium and Feiderium (Microtrombidiidae) have independently lost the erapodiura. No trorabidioids have lost paired claws. The loss of the empodium occurs only on pretarsus I and II. Pretarsus III may be reduced or modified independently of pretarsus I and II.

Pretarsus III shows a reduction of the paraxial (ocr) and a lateral movement of the claw. Mullen and Vercammen-Grandjean, 1978 called this modified claw a smilum because it appeared to be similar to "small pruning knife". In some groups, i.e. Microtrombidiidae and

Eutrombidiidae, this claw becomes thickenend to two or three times 48 the width of a normal claw and less than one-half the length of the antaxial claw. In the Trombidiidae, a similar reduction on the length and rotation to the paraxial occurs in some genera but it has become only slightly thickened and may be lost altogether.

SPECIALIZED SETAE

Solenidia are thin walled, hollow, specialized setae found on the palp tarsus and legs. They are characterized by the lack of actinopilin and may have an annulate appearance caused by numerous fenestra and helicoid ridges (Norton, 1977). They occur in a variety of shapes that can include baculiform, ceratiform, piliform, or claviform and are believed to be chemoreceptors. The notations for solenidia are greek letters (Grandjean, 1935; van der Haramen, 1980;

Goff, et al, 1982). Solenidia on the tarsus are omega, those on the tibia phi, those on the genu sigma, and those on the femur are theta.

In the Parasitengona, solenidia can be present on the femur, genu, tibia of legs I, II or III and tarsus of legs I and II. Solenidia on the femur are found scattered through a number of genera in the

Trombellidae, Johnstonianidae, Neotrombidiidae and some

Trombiculidae. The abbreviation for the solenidiataxy is fsol=

I(0-2-2-l), 11(0-1-2-1), 111(0-1-0-0) where I(0-2-2-l) means leg I femur is without solenidia, genu leg I has two solendia, tibia leg I has two solenidia and tarsus leg I has 1 solenidia.

The presence of femoral solenidia, theta, is apomorphic, as they are absent in the Erythraeidae and Anystidae. The Trombellidae, 49

Johnstonianidae, Neotrombidiidae and some Trombididae are the only

families which regularly have femoral solenidia. The Trombellidae

tend to be hypertrichous for theta ranging from 4-2-2 to 10-10-8. In

the Johnstonianidae the number ranges from 2-1-1 to 1-0-0 with the

Neotrombidiidae with 3-3-3 to 2-2-2. It appears that most

Trorabiculidae have secondarily lost theta except in some groups like

Hannemania (Leuweenhoekiinae), which have up to 10 theta.

In the Trombidioidea the solenidiotaxy of the genu varies with each leg. Leg I usually has two setiform sigma (s3 of Newell, 1957).

The length and position of these solenidia are used in species descriptions. Legs II and III usually have a single setiform sigma, but in some groups there can be multiple solenidia. The

Allothrombiinae (Trombidiidae) usually have two sigma on legs II and

III. Those Microtrombidiidae which have three sigma on genu I usually have two sigma on legs II and III. The number of sigma on the genu appears to have significance only at the generic level, because each family has independently modified the numbers of sigma either by addition or reduction.

In the Chyzeriini (Chyzeriidae) have lost sigma on all legs while the Pteridopiini have only a single sigma on each leg.

The solenidia on the tibia are remarkably constant throughout the Parasitengona. The Trombidioidea usually has two phi: one proximal and usually piliform and the other (distal) is a smaller version of omega on tarsus leg I, usually ceratiform or occasionally baculiform. The position of these two setae varies within families and their relative positions can be used in species descriptions. On leg II there are also two phi similar in structure to tibia I. One

is proximal and the other distal. These two solenidia exhibit little

variation in position throughout the terestrial Parasitengona and are

not used taxonomically. In the Tanaupodidae and some Erythraeoidea

there may be a companion seta associated with phi. On leg III there is a single setiform phi near the proximal end of the tibia, except in the Trombidioidea where it is lost. Some water mites have independently lost the phi on leg III.

The tarsi of legs I and II each have a single solenidion (omega) located on the proximal half of the segment. The omega is different from the leg solenidion in being baculiform or occasionally ceratiforra. The length and position of omega is used in species descriptions. The relative positon of omega and the famulus

(epsilon) is used as a family character for the Microtrombidiidae.

While the omega on tarsus I and II are similar in appearance and shape, omega II is generally smaller and narrower (Newell, 1957). In the Trombidioidea this distinction is less evident than in the

Trombellidae, Chyzeriidae and Johnstonianidae.

The famulus (epsilon) is a small, hollow seta with actinopilin, found on the dorsal surface of the tarsus of legs I and II of all

Parasitengona, except possibly Calyptostomatidae. The term famulus, coined by Grandjean in 1935, has only recently been applied to the terrestrial Parasitengona (Newell, 1957 and Goff, et. al., 1982) to replace the chigger terra microtarsala. Robaux (1974) and Goff, et. al. (1982) used the notation "e" for the famulus. Since epsilon was used by Grandjean (1935) and "e" is used to refer to the supracoxal 51

seta (van der Haramen, 1980) I will follow van der Hammen (1980) here.

The famulus on tarsus leg I is usually distal to omega I in most

Parasitengona. In the Microtrombidiidae the famulus is proximal to

omega I resulting in a synapomorphy distinguishing the family. The

famulus on tarsus II is relatively constant in its position, being

approximately at the same level as omega II. In the johnstonianid

genera Diplothrombium and Centrothrombium the famulus on leg I is

elongated.

Eupathids (zeta) are modified normal setae with a hollow center, and actinopilin. These setae are spiniform and occur on tarsus of

legs I, II and occasionally III.

The Trombellioidea have the eupathidial formula (2-1-0) (except

Neotrombidiidae with 1-0-0) and the erythraeoids have a eupathidial

formula of 2-1-1 or occasionally 3-2-2. In the Trombidioidea the

Trombidiidae and Neothrombiide have only two eupathidia on tarsus leg

I (2-0-0) and Calothrombiidae, Eutrombidiidae and Microtrombiidae have 2-1-0.

On tarsus I of the trombidioids there are two eupathidia. One is placed in the distodorsal half of the segment (called subterminala in chiggers). The second is located distoventrally or laterally (the pretarsala in Goff et. al., 1982). The distodorsal eupathidium may have a closely associated normal seta which is called the parasubterminala by chigger workers. This seta is called a companion seta by Newell, 1957 and is present in the Tanaupodidae, Chyzeriidae,

Trombellidae, Trombiculidae and Johnstonianidae. Companion setae are absent in Neotrombidiidae and the Trombidioidea. 52

In the Johnstonianidae, and independently in the genus

Podothrombium (Podothrombiidae), some species have a multiplication of eupathidia on the ventral surface of tarsus leg I.

The microseta (vestigial seta of Newell, 1957; vestigiala of

Southcott, 1961; microgenuala and microtibiala of chigger workers) is a small spinose seta, of unknown function, on the genu of legs I, II, tibia leg I and rarely genu leg III. The seta has been given the notation k (Robaux, 1974, Goff, et. al., 1982) to distinguish it from the famulus on tarsus of legs I and II. On the genu this seta is located distodorsally in about the same position on all genera studied. The genus Parathrombium (Trombidiidae) is the only parasitengone to have a k seta on genu leg III. In the family

Neothrombiidae the microseta on genu leg II has been lost. In the

Microtrombidiidae some genera have independently lost this seta. In the Johnstonianidae all genera except Johnstoniana have lost microsetae.

The microseta on tibia leg I is universally present at approximately the same position in all terrestrial Parasitengona. In some Erythraeidae the microseta is baculiform or ceratiform, where as it is setiform in other Parasitengona.

Occasionally, some of the normal (usually distolateral) setae

(n) on the legs become modified into long whiplike setae without barbs or with only basal barbs. Newell (1957) called them bothridial setae while chigger workers call them mastisetae (i.e. mastifemorala, mastitarsala etc). While these setae are modified normal setae they do occasionally have larger bases which appear to be bothridial but 53 are not the same as those found on the legs of larval Smarididae. In

the Trombidioidea these setae are best developed in the Trombidiidae where they are found on tarsus I as a pair of mastisetae flanking omega I in Trombidium.

Postlarval Instars

(Deutonymphs and Adults)

Most generic and many species descriptions are based on the active postlarval instars, especially older descriptions. Their size and hirsute idiosoma and legs make them difficult to study often resulting in limited descriptions. The characters used in descriptions emphasize the dorsal idiosomal setae, tarsus leg I and palp (usually only the palpal tarsus and tibia). While in many cases this is sufficent at the generic level, it is generally not sufficient for species recognition. In fact, we really do not have a good understanding of the intra and interspecific variation in the postlarval Parasitengona because most species and genera are known from only a few species and usually few specimens. It was not my intention to provide an exhaustive and comprehensive description of the postlarval instars but to review some of the aspects of postlarval morphology to find additional characters useful in phylogenetc analysis. 54

Gnathosoma

The gnathosoma, like the larva, is a pseudotagma with two pair of appendages: palps and chelicerae. Unlike the larva, the palps and subcapitulum are usually densely setose, making comparison and identification of individual setae impossible. Only the palpal tibia and tarsus are used taxonomically.

The cheliceral blade provides few characters useful in phylogentic analysis because of little variation. In the Erythraeina

(Erythraeoidea and Calyptostomatoidea) the cheliceral blade is stylet-like while the rest of the Parasitengona retain the curved cheliceral blade. In the Microtrombidiidae some genera have reduced the cheliceral blade to a stylet-like form without increasing its length or modifying the cheliceral base as in the Erythraeina.

Feider (1955) illustrated differences in length of cheliceral bases but data are available on only a few trorabidioid taxa.

The palpal tibia, like other palpal segments is usually densely setose, but unlike other segments there are a number of taxonomically important spinose setae in the Trorabidiina, which first appear in the

Tanaupodidae. At the distal end of the tibia is the tibial claw, which has not been used taxonomically. On the dorsal surface immediately behind the tibial claw is a second hypertrophied seta, called the accessory claw. The accessory claw first appears in

Eothrombium (Tanaupodidae) and is best developed in the more derived groups (i.e., Microtrombidiidae and Eutrombidiidae) and is secondarily lost, reduced or relocated in the trombiculoids. In the 55

Johnstonianidae this seta is usually in a ventral position between

the palpal tibial claw and articulation of the palpal tarsus. In the

Trombidiidae, Trombiculidae and Neotrombidiidae the seta has been reduced or lost. In addition, the size of the accessory claw can vary in relation to the tibial claw, but it is not known if this

variation is of phylogenetic significance because of limited sample size.

Other parts of the palpal tibia have varying numbers and arrangement of spines. The primitive arrangement being a random pattern with a few spines in the less derived terrestrial

Parasitengona (i.e., Tanaupodidae, Chyzeriidae and Trombiculoidea).

Podothrombiidae is the only family in the Trombidioidea to retain the primitive randoin pattern of spines while its sister group, the

Trombidiidae, has lost all spines on the palpal tibia. In the more derived Eutrombidiidae and Microtrombidiidae, the palpal tibial spines are arranged in one or two rows, with the first row starting immediately posterior to the accessory claw and the second row beginning about midway along the dorsum of the tibia. In some forms both rows are united into one long row as is found in Valgothrombium

(Microtrobidiidae)

In addition to the dorsal rows of setae, some Microtrombidiidae and most Eutrombidiidae have additional large ventrally directed spines located near the palpal tarsus articulation. The number of spines appears to be variable among species in the same genus but their significance in generic determination is unknown.

The palp tarsus has three types of setae: normal setae, 56 eupathidia and solenidia (omega). Some authors (Robaux, 1967) believe the number of terminal eupathidia is an important taxonomic character and make an effort to count and illustrate the terminal eupathidia. I found this character to be unreliable and of questionable value due to variation in number of setae and difficulty in observing eupathidia. Other characters of the palpal tarsus include general shape and size relative to the palpal tibia.

Idiosoma

The idiosoma of most Trombidioidea is densely covered with setae, while those usually associated with wet habitats (i.e.,

Johnstonianidae, Calyptostoraatidae, and Hydracarina) have only widely distributed simple setae. It is the dorsum of the idiosoma which has provided previous workers with a rich and frequently confusing array of characters used in present adult classifications.

The crista metopica (crista (L)=ridge; metopa (L)= space between) is widely used in descriptions and is synonomous with what is now called the prodorsal sclerite. Berlese (1912) published a figure of the different forms of the crista metopica which has been widely used and was reproduced by Thor and Willmann in 1947. If one carefully examines the prodorsal area of the trorabidioid adult or deutonymph, one finds that the "crista metopica" is actually composed of two elements: 1) an internal heavily sclerotized ridge projecting out from and/or connecting the prodorsal trichobothria; and 2) a sclerotized area which is expressed as a surface sclerite. I will 57 retain the term crista metopica, but redefine it to include only the internal, heavily sclerotized area associated with the prodorsal trichobothria and the term prodorsal sclerite will refer to the surface sclerite. In those groups which have two pair of prodorsal trichobothria the crista metopica connects both bothridial bases. In many groups (most Microtrombidiidae, some Johnstonianidae and

Trorabiculidae) the prodorsal sclerite and the internal crista metopica coincide. In the Trombidiidae, Podothrombiidae,

Calothrombiidae and Neothrombiidae there has been considerable additional prodorsal sclerotization which provides useful synapomorphic characters.

The Trombidiiae are characterized by a derived prodorsal sclerite which extends laterally and anteriorly on either side of the crista metopica. The amount of sclerotization varies among the genera in the family. In the Neothrombiidae the prodorsal sclerite is also well developed (Neothrombium and Acridothrombium) and extends laterally to include the ocular plate, while in the cave adapted genus Ceuthothrombium the prodorsal sclerite is secondarily reduced.

In the Microtrombidiidae the prodorsal sclerite is limited to the area of the crista metopica.

The shape of the dorsal idiosoraal setae has been used extensively in taxonomic descriptions, especially in the

Microtrombidiidae where the greatest variation occurs. From the primitive simple nude or barbed seta a wide array of forms have evolved (see Figure 6). Many groups retain the primitive setal form with some reducing setation, especially in those forms living in wet 58

FIGURE 6. Representative Postlarval Dorsal Idiosomal Setae

A. Allothrombium (Trombidiidae); B. Wilmannellia

(Microtrombidiidae); C. Manriquia (Microtrombidiidae); D.

Trombidium (Trombidiidae); E. Campylothrombium

(Microtrombidiidae); F. Microtrombidium

(Microtrombidiidae); G. Acridothrombium (Neothrombiidae) 59

FIGURE 6. Representative Postlarval Dorsal Idiosomal Setae habitats. The Trombidiidae and Eutrombidiidae retain the basic setiform shape but have added barbs producing a thicker setae. The

Neothrombiidae, at least in the three postlarval forms known, have modified dorsal idiosomal setae where the basal barbs have become spines as in Acridothrombium. In the Microtrombidiidae idiosomal setae range from the setiform to slightly inflated setae of

Microtrombidium to the bizzare setae in Hiothrombidium Woraersley.

The idiosomal setae in the Microtrombidiidae appear to have evolved in two groups. The first group includes the setiform shape and those types only slightly expanded or otherwise modified. The second group

(tribe Cercothrombiinae) include a wide array of inflated and globose setae. Associated with the inflated setae in the Cercothrombiinae is the presence of brushlike setae on the palpal genu, femur and basal leg segments.

Other idiosomal characters include the autapomorphic presence of a pygidial plate in Eutrombidiidae, the setose fingerlike projections in Chyzeriidae and the circular sclerites in the Trombellidae. In the genus Valgothrombium the cuticle in the anterior prodorsal region along either side of the crista metopica is grooved.

The shape of tarsus leg I is frequently figured in taxonomic descriptions to show the inflated or elongated condition of the tarsus. There seems to be considerable variation in the shape of the tarsus but the Microtrombidiidae generally have an inflated tarsus leg I. Other characters include the autapomorphic appearance of aliform setae subtending the pretarsal claws in Allothrombium

(Trorabididiae) and the highly modified sete on leg IV in

Coccothrombium (Microtrombidiidae). PHYLOGENETIC ANALYSIS

Relationships of the Trombidioidea within

the Parasitengona

Using the Anystina, specifically the Anystidae, as an outgroup, the cohort Parasitengona can be defined by four synapomorphies: 1) a postlarval lifecycle involving alternating active (deutonymph and adult) and calyptostatic (protonymph and tritonymph) instars

(Johnston and Wacker, 1967); 2) heteromorphic postlarval instars; 3) larva without peritreraes (18); 4) larva without cheliceral setae

(17). The alternating active and calyptostatic postlarval cycle is not autapomorphic for the Parasitengona but is shared with the

Pterygosomatidae (Anystoidea) a highly derived group parasitic on lizards, cockroaches and reduviids whose phylogenetic relationships are unknown.

The Parasitengona is a large and diverse group, with over 50 per cent of all Prostigmata (Kethley, 1982), in which the relationships among some groups are obscured by numerous autapomorphies and the successful invasion and subsequent radiation in new habitats by two groups, the water mites in fresh water and the chiggers on vertebrates.

61 62

Starting with the least derived Parasitengona I will discuss each group, leading to a redefinition of the Trombidioidea as well as defining several new hypothesized monophyletic groups using shared derived characters. A proposed new classification of the higher categories of the Parasitengona is given in Table 2. Figure 7 is a cladograra of the hypothesized relationships within the Parasitengona using 23 characters (Table 3).

The Erythraeoidea (Erythraeidae and Smarididae), with more than

34 genera and 200 nominant species, are considered the least derived

Parasitengona by the plesiomorphic characters they retain (two pair of prodorsal trichobothria, neotrichy on the legs) and their overall similarity to the Anystidae, especially the Erythracarinae. The erythraeoids are defined by four synapomorphies: 1) presence of stylet-like cheliceral blades in the active postlarval instars

(120c); 2) neotrichous dorsal idiosoma in the larva (31c, 32c, 33c,

34d); 3) loss of Claparede’s organ (58); 4) absence of an anal opening (60). The systeraatics of the Erythraeoidea was comprehensively reviewed by Southcott (1961), but currently is in need of revision.

The Calyptostomatoidea is a small monophyletic group of mites represented by a single genus, with five nominate species, found in moist and wet habitats world wide. The phylogenetic position of the calyptostomatids have remained unclear. Robaux (1974) ignored them and Vercammen-Grandjean (1973) just listed them in his classification. Feider (1955) placed the origin of the calyptostomatids close to the erythraeoids in his "evolutionary 63

Table 2. Proposed Phylogenetic Hypothesis for the Parasitengona

Cohort Anystina Cohort Parasitengona

Subcohort Erythraeina sf. Erythraeoidea f. Erythraeidae Robineau-Desvoidy, 1828 f. Smarididae Vitzthum, 1929

sf. Calyptostomatoidea f. Calyptostomatidae Oudemans, 1923

Subcohort Hydracarina

Subcohort Trombidiina sf. Tanaupoidea f. Tanaupodidae Thor, 1935

sf. Chyzerioidea f. Chyzeriidae Womersley, 1954

sf. Trombiculoidea f. Johnstonianidae Thor, 1935 f. Trombiculidae Ewing, 1929 f. Neotrombidiidae Feider, 1959 f. Trombellidae Thor, 1935

sf. Trorabidioidea f. Podothrombiidae Thor, 1935 f. Trombidiidae Leach, 1815 f. Neothrombiidae Feider, 1955 f. Calothrombiidae Oudemans, 1940 f. Eutrombidiidae Thor, 1935 f. Microtrombidiidae Thor, 1935 MICROTROWBtOtrDAE

trombidioidea It. >0. II II. TROM8ICULOIDE A

CHYZERIOIDEA

TANAUPODIOIDEA

ERYTHRAEINA

ANYSTINA PARASITENGONA

FIGURE 7. Cladograin of Hypothesized Relationships within the

Parasitengona. Table 3. Character States used on the Phylogenetic Analysis of the Parastiengona

(Families and Superfamilies).

Tax* Characters 1 13 19 31 32 33 34 35 36 48 58 59 76 85 87 94 96 97 104 105 106

Anystidae a a a a a a a a a a a 7 a a a a a a a a a Erythrarldae a a a c c d d a a a b b a a b b a a a a a Saarldldae a a a c c d d a a b b a a b b a a a a a Calyptostooatldae ? a b c c d d a a 7 a a? a a a a a a a a a Hydracarina b b a a a a a a a b a a b a a a b a a a a Tanaupodoidea a a a a a a a a a a a a b b b b a b a a a Ghyaerloidea a a a/b b b b b a a a a b b a b a a a a a Troabellldae a b a b b b b b b b a b b b a b a a b a b Neotroabldildae b b a b/c b/c b/d b/d b b b a abb a b a a b a b Troablculldae b b b c c d d b b b a b b/k b a b b a a a a Johoatonlanldae b b a/b b b b b b b b a b b b a b a a a b a Troabidloldea b a/b b b b b/c b/c b b b a b b/c b a a b a a/b a/b a/b

O' Ul 66 tree". Witte (1984) tenatively placed them closer to the

Johnstonianidae and traditional trombidioids based on his comparative study of reproductive structures. While the calyptostomatids share two synapomorphies with the erythraeoids (stylet-like cheliceral blades and neotrichous larval dorsal idiosoma), they can be defined by five additional synapomorphies: 1) one pair of prodorsal trichobothria in the larval (19) and postlarval (121) instars; 2) prodorsal sclerite reduced in larval (29c) and postlarval instars

(122b); 3) stellate cuticular pattern on the idiosoma in the larval

(47c) and postlarval (130) instars; 4) linear palps in the postlarval instars (125); 5) retractile gnathosoma in the postlarval instar

(126). I was unable to find any larval synapomorphies supporting relationships with the other trombidioids. Traditionally, the calyptostomatids have been placed with the troraibidoids based on the presence of Claparede's organ (a plesiomorphy) and the presence of one pair of prodorsal trichobothria (an apomorphy which has reoccurred throughout the Parasitengona) with the assumption that the stylet-like chelicerae were independently derived. I suggest the

Erythraeoidea and Calyptostomatoidea are sister groups in the monophyletic Erythraeina, based on the shared derived stylet-like cheliceral blades and neotrichous larval idiosoma. I reject the idea that the stylet-like cheliceral blades were independently derived in the calyptostomatids. It could be also argued that the cosmopolitan distribution and few species suggest the calyptostomatids are a relictual group with an ancient origin and is the only member of an early derived erythraeoid group. 67

The remaining parasitengones are separated by the presence of four or less setae on the genua legs I, II and III (76, 77b). The

Water Mites or subcohort Hydracarina is a complex group of 11 superfamilies, with over 50 families and thousands of species. The origin of the Hydracarina has been discussed widely in the literature

(Cook, 1974; Smith, 1976; Newell, 1957; Mitchell, 1957; Barr, 1972) without resolution. Barr (1972) suggested, based on his study of the water mite ejaculatory complex, the water mites were monophyletic

(except possibly Hydrovolvzia). Within the terrestrial

Parasitengona, the Johnstonianidae have been suggested as the possible sistergroup of the Hydracarina (Newell, 1957; Smith, 1976).

While it was not within the scope of this study to determine the origin of the water mites, my comparative study suggests the most likely sistergroup for the water mites is in the Tanaupodoidea which is also hypothesized as the primitive trombidioid sister group. This is based on examination of larval "lower water mites" which have aerial larvae and have not been modified for swimming. The

Hydracarina are characterized by three synapomorphies: 1) loss of one pair of adoral setae (1); 2) loss of supracoxal setae on gnathosoma

(13); 3) loss of supracoxal setae above leg I (48). Only with the discovery, rearing of additional taxa, and careful comparative study of more taxa will a better understanding of the relationship of the water mites and their sistergroup be recognized.

The subcohort Trombidiina (Tanaupodoidea, Chyzerioidea,

Trombiculoidea and Trombidioidea) are characterized by three synapomorphies: 1) one pv setae on tibia leg I (85); 2) dorsal 68 eupathid with companion seta (94b); 3) accessory claw on palpal tibia in the postlarval instars(115).

The Tanaupodoidea (based on the Eothrombium and Polydisica) retain a number of primitive larval characters (reduced idiosomal chaetotaxy, anal sclerites with setae; supracoxal setae e and el; two pairs of prodorsal trichobothria) but are also distinguished by three synapomorphies; 1) pregenital tubercle (127) in the postlarval forms;

2) reduction of palpal tibial claw in the larva (10e,f); 3) companion setae for omega on tarsus leg I (88). Thor 1935a recognized

Tanaupodus as different when he errected the subfamily Tanaupodinae for it and six genera (Eothrombium Berlese, Rhinothormbium Berlese,

Typhlothrombium Berlese, Tanaupodaster Vitzhum, Calothrombium

Berlese, and Neotanaupodus Garman). Feider (1959) elevated the tanaupoids to the family Tanaupodidae and superfamily Tanaupodoidea at the same time he elevated Calothrombium to a family. Subsequent workers ignored his classification and continued to place the tanaupodids with the trorabellids (Vercammen-Grandjean, 1973, Krantz,

1978, Kethley, 1982). Vercaramen-Gandjean (1973) removed

Typhlothrombium and Tanupodaster to the Speleothrorabiinae

(Trombellidae) leaving only Rhinothrombium, Eothrombium. Tanaupodus and Paratanaupodus Andre in the group, but made no reference to

Neotanaupodus. Newell's (1957) excellent study of the

Johnstonianidae did not include a comparison with Tanaupodus and

Eothrombium, resulting in his classification being paraphyletic and this has been followed by other workers (Vercammen-Grandjean, 1973;

Robaux, 1974). In spring 1984 I had the opportunity to study 69

Berlese's types of Eothrombium and Rhinothromium and to compare them with postlarval specimens of Newell's genus Lassenia and found them to be congeneric. Robaux (1967d) speculated that Lassenia and

Eothrombium were synonoraous. Since Newell (1957) had previously described the larva of Lassenia it was possible to determine larval relationships in the Tanaupodidae based on my synonomy of the two genera.

The Chyzeriidae is another group which retains a number of primitive characters (anal sclerite with setae, supracoxal setae e and el, unmodified pretarsal claw and ernpodium) while possessing several autapomorphies (reduced solenidiataxy (78b, c,79e), modified cheliceral blade (16c) and the setose projection in the postlarval instars (128)) as well as synapomorphies uniting them with the rest of the trombidioids (dorsal idiosoraal setation neotrichous with the addition of an extra pair of setae in row C, D, E, F, and H (31b,

32b, 33b, 34b, 35b), the apparent loss of setae from the PS series

(36b) and loss of companion setae with phi and omega on leg I

(87,88)). This group has been included in the Trombellidae by many authors (Thor and Willmann,1947; Vercammen-Grandjean, 1973; Feider,

1979; Southcott, 1982). Vercammen-Grandjean (1973) and

Vercammen-Grandjean et. al. (1974) recognized the chyzeriids as a distinct group but did not define the group. Southcott (1982) included the chyzeriids in the Trombellidae on the basis of one pair of prodorsal trichobothria and the mistaken observation that the crista raetopica was absent. He also stated there were no larval characters to reliably separate the group. My comparative study 70

suggests that the Chyzeriidae is distinct from the Trombiculoidea

because of two synapomorphies shared by Trombiculoidea not found in the Chyzeriidae.

Vercammen-Grandjean et. al. (1974) included the genus

Ralphaudyana in the chyzerids without an adequate definition.

Southcott (1982) suggested it was more like Lassenia (=Eothrombium); my study indicates that Ralphaudyana is not a chyzeriid nor is it a tanaupoid but should be placed in the Trombiculoidea although its relationships remain uncertain.

The superfamily Trombiculoidea, which is the oldest available valid family group name despite the numerous family group names proposed by Feider (1955, 1959, 1979), contains four families

(Johnstonianidae, Trorabiculidae, Neotrombidiidae and Trombellidae) and two unplaced genera, Ralphaudyana and Audyana. The group is defined by at least three larval synapomorphies, the presence of theta on the femur of at least one leg (73, 74, 75) and the absence of supracoxal setae (13, 48). Because of the apparent heterogeneous nature of the group I could find no postlarval characters which are consistent within the group. The hypothesized monophyletic

Trombiculoidea must be considered weak and the possibility of paraphyly exists. Only when the larval and postlarval instars of the remaining genera, especially the three subfamilies in the

Trombellidae, are correlated with larval and postlarval instarscan the question of monophyly of the Trombiculoidea be looked at in detail.

Newell (1957) provided us with an excellent revison of the Johnstonianidae which has been the basis for speculation on the relationships of the Parasitengona. Unfortunately his study was limited to western North America. The Johnstonianidae can be characterized by two larval synapomorphies: the presence of only paired claws (105, 107) on all legs (empodium absent) and an autapomorphic spine on the palpal tarsus (12) and a postlarval synaporaorphy with the accessory claw on the palpal tibia on the lateral or ventral surface (116). These mites are frequently found near water which has lead to speculation that they were the sister group of the Hydracarina.

The Trombiculidae (chiggers), with nearly 2000 nominate species, are the only parasitengones to regularly parasitize vertebrates

(excluding fishes), this has lead to a wide diversification and an obscuring of phylogenetic relationships. Some authors recognize two chigger families, the Trombiculidae and Leeuwenhoekiidae (Krantz,

1978; Kethley, 1982; Vercammen-Grandjean, 1973). I did not attempt to fully examine the relationships of the chiggers, but ray study suggested there may well be two families. The current classification is in need of careful revision. I will follow Goff et. al. (1982) in this work.

The remaining two families (Neotrombidiidae and Trombellidae) are united by the synapomorphic absence of the dorsal eupathid on tarsus leg I (93) and absence of pretarsal claws on legs I and II

(104). The Neotrombidiidae is characterized by two synapomorphies:

1) absence of paired claws on leg III (106); 2) reticulate patten on at least one coxal field (129). Lindquist and Vercammen-Grandjean 72

(1971) and Singer (1971a, b) independently revised the two genera and respective species. Robaux (1977) described a third larval genus from California. The postlarval forms in this family are known only for the type species and are characterized by the autapomorphic trifurcate dorsal idiosomal setae.

The larvae of Trombellidae are characterized by the presence of multiple theta (4+) on femur leg I and the postlarval circular structures on the dorsal idiosoma of adults (131). This family is known from 15 genera but only three have correlated larval and postlarval instars.

As stated above the Trombiculoidea and Trombidioidae are sister groups. The superfamily Trombidioidea is characterized by seven larval synapomorphies: 1) presence of at least one additional dorsal sclerite (scutellum) with at least one pair of setae posterior to the prodorsal sclerite (38); 2) loss of phi on tibia leg III (86); 3) femur leg III with four setae (67d); 4) fsp=6-6-6 (62,63,64); 5) genu legs II and III usually with less than four setae (77c, d); 6) absence of companion setae with the dorsal eupathid on tarsal leg I

(94); 7) without dorsal eupathid and companion seta on tarsus leg II

(96,97).

Phylogenetic Relationships within the Trombidioidea

The Trombidioidea is a monophyletic group consisting of six families (Podothrombiidae, Trombidiidae, Microtombidiidae,

Calothrombiidae, Eutrombidiidae and Neothrombiidae) with the shared 73

synapomorphies described above. The cladogram in Figure 8 presents the hypothesized phylogenetic relationships at the familial and subfamilial level. Table 4 presents a sequenced listing of the hypothesized phylogenetic relationships for the trombidioid families and subfamilies based on studied taxa. Within the Trombidioidea there are two monophyletic family groups. One group, the families

Podothrombiidae Thor, 1935 and Trombidiidae Leach, 1815 form an unnamed monophyletic group characterized by five synapomorphies: 1) five setae on femur leg I (65c); 2) femur leg II with four

(occasionally five) setae (66d); 3) three setae in genu legs II and

III (occasionally four setae) (67c); 4) five setae on tibia leg I

(82f); 5) subterminal eupathid on tarsus leg II absent (99). Table 5 lists the characters and character states used in the phylogenetic analysis of the Podothrombiidae and Trombidiidae. Figure 9 is a cladogram of the hypothesized relationships of the Trombidiidae and

Podothrombiidae. The remainder of the trombidioid families

(Neothrombiidae, Calothrombiidae, Eutrombidiidae and

Microtrombidiidae) are united by a single unique synapomorphy, two setae on genu legs II and III (77d).

Podothrombiidae

The family Podothrombiidae Thor, 1935 was erected for the single nominante genus Podothrombium Berlese, 1910. Feider (1959) elevated it to family and superfamily status. Vercammen-Grandjean (1973), without reference to Feider’s earlier work, also elevated NEOTHROMBIINAE HOPCOTHROMBRNAE FEIOERIIMAE MICROTROMBIDIINAE AEQIALOTHROMBIDIINAE CALOTHROMBINAE EUTROMBMMNAE AETHETHROMBIDIINAE PODOTHROMBIIDAE ALLOTHROM8BNAE TROMBWNNAC M 01 o a

NEOTHROMBIIOAE 75

Table 4. Proposed Phylogenetic Classification for the Trombidioidea

Trombidioidea f. Podothrombiidae Thor, 1935 Podothrombium Berlese, 1910

f. Trombidiidae Leach, 1815

Trombidiinae Leach, 1815 Oligothrombidium NEW GENUS Trombidium Fabricius, 1775 Parathrombium Bruyant, 1910

Allothrombiinae Thor, 1935 Nanothrombidium NEW GENUS Allothrombium Berlese, 1903 Dinothrombium Oudemans, 1910 f. Neothrombiidae Feider, 1959

Neothrombiinae Feider, 1959 Adelothrombidium NEW GENUS Kermathrombidium NEW GENUS Neothrombium Oudemans, 1909 Acridothrombium NEW GENUS Ceuthothrombium Robaux, Webb & Campbell, 1976

Aethethrombidiinae NEW SUBFAMILY Aethethrombidium NEW GENUS Megalothrombium NEW GENUS

Aegialothrombidiinae NEW SUBFAMILY Aegialothrombidium NEW GENUS f. Calothrombiidae Oudemans, 1940 Calothrombium Berlese, 1918 (adult) Dectothrombidium NEW GENUS f. Eutrombidiidae Thor, 1935

Eutrombidiinae Thor, 1935 Eutrombidiini Thor, 1935 Eutrombidium Verdun, 1909 Scutodesmus NEW GENUS Table 4. (continued)

Milliotrombiini NEW TRIBE Atopothrombidium NEW GENUS Milliotrombidium Shiba, 1976 Hoplothrombiinae NEW SUBFAMILY Hoplothrombium Ewing, 1925 Cryptothrombidium NEW GENUS f. Microtrorabidiidae Thor, 1935

Feideriinae Vercammen-Grandjean & Cochrane, 1974 Manriquiini Welbourn, 1984 Pygothrombium NEW GENUS Manriquia Boshell & Kerr, 1942 Reticulatrombidium Shiba, 1976 Feideriini Vercammen-Grandjean & Cochrane, 1974 Valgothrombium Willmann, 1940 Gonothrombium Feider, 1948

Microtrombidiinae Thor, 1935 Microtrombidiini Thor, 1935 Microtrombidium Haller, 1882 Platytrombidium Thor, 1936 Ettmulleria Oudemans, 1911 Atractothrombium Feider, 1951 Achelothrombidium NEW GENUS Cercothrombiini NEW TRIBE Diathrombium Moser & Vercammen-Grandjean, 1979 Campylothrombium Krausse, 1912 Lithothrombidium NEW GENUS Trichotrombidium Kobulej, 1950 Willmannella Feider, 1951 Cercothrombium Methlagl, 1928 77

Podothrombiinae to the family level. The Podothrombiidae can be recognized by four synapomorphies: 1) reduction of palpal tibial claw to a single tine (lOe); 2) dorsal idiosomal setae neotrichous (31c,

32c, 33d, 34d); 3) scutellum reduced to include only the basal sclerites of cl of setae (41b); 4) tarsus leg I with extra ventral eupathidia (95). Podothrombium is unique in the Trombidioidea in the retention of primitive characters of the Trombiculioidea (well developed setae on the palpal femur and genu and the primitive random arrangement of palpal tibial spines on the postlarval instars). In addition, the dorsal idiosomal setae of the postlarval instars are nude or have few barbs. The prodorsal sclerite, of the postlarval instars, has extensive anterolateral sclerotization and the larval chaetotaxy suggests close affiliation with the Trombidiidae (figure

9). Welbourn (1984) suggested the podothrombiids as the possible sister group the rest of the the Trombidioidea.

Little is known of the biology of Podothrombium other than the primary hosts are Aphidae (Homoptera) (Welbourn, 1983). The genus is quite large with 37 species listed from Europe (Robaux, 1967a) and only one named from North America.

Trombidiidae

The sistergroup to the Podothrombiidae is the Trombidiidae

Leach, 1815. This family can be distinguished by three larval and two postlarval synapomorphies: 1) Palpal genu without setae (7d); 2) secondary loss of one pair of dorsal idiosomal setae from row E Table 5. Character States used in the Phylogenetic Analysis of the Podothrombiidae and Trombidiidae

Taxa Characters

5 6 7 10 22 24 26 33 37 41 50 52 79 95 101 102

Podothrombiidae Podothrombium dacea aadbbaaabaa

Trombidiidae Trombidilnae OloKOthrombidium d b d b b b a c a a b a a a a a Parathrombium d b d b b a a c a a b c a a a a Trombidium d b d b a a a c a a b b a a b b Allothrombiinae Nanothrombidium a — d b a a a c a a b b a a a a Dinothrombium e — d b a a b c a a b a b a b b Allothrombium e — d b a a a c a b d a b a b b FIGURE 9. Cladogram ofHypothesized Relationships the withinFIGURE PODOTHROMBIUM 32c, 3c 33 , c 2 3 . c 1 3 d, 10* 0 ,1 6 0 , 4d 3 PODOTHROMBIIDAE Podothrombiidaeand Trombidiidae. 0. 0, 102 101, 70b. 06c, 77c, 01 00 0 021, , ,d c 7 7 . ,d c 6 0 , c 6 6 26 LOHO BIINAE ALLOTHROM 2 4b 2 62b 6* TROMBIDIIDAE d 33c. 11d 116c 17d, 1 , b 0 6 . c 3 3 7d. ROMBI IAE IINA ID B M O TR 6d. • c 2 6 22b 2b 6 80

(33c); 3) coxal field leg II with two setae (50b); 4) palpal tibia of postlarval instars without spines (117d); 5) palpal tibia of postlarval instars without acccessory claw (115c).

The family, as defined by the above synapomorphies, includes seven genera and more than 50 species worldwide. Based on the chaetotaxy of the the palpal femur the Trombidiidae are divided into two subfamilies, Trombidiinae and Allothrombiinae.

The Trombidiinae as currently recognized consists of seven genera (Vercammen-Grandjean, 1973; Robaux, 1967a). Feider (1950) proposed the genus Teresothrombium based on variation of the dorsal idiosomal setae in the postlarval instars. Examination of larvae of

Teresothrombium from Feider's collection indicate this genus is congeneric with Trombidium. My phylogenetic hypothesis of relationships in the Trombidiinae is based on examination of

Trombidium, Parathrombium and Oligothrombidium.

Within the Trombidioidea, Oligothrombidium is distinguished by the autapomorphic presence of expanded trichobothria (24b). The genus is known from two free-living larval specimens, one each from

Tennessee and Alabama. In addition, the arrangement of barbs on the dorsal idiosomal setae and barbs on the palpal femoral seta can also be used to distinguish this genus.

The two sister genera, Trombidium and Parathrombium are defined by the synapomorphic presence of an AM seta without barbs (22b).

Both are very similar morphologically and differ in the morphology of seta la on coxal field leg I and host preference. Welbourn (1983) listed three genera of Hemiptera (Miridae and Pyrrhocoridae) and two 81 species of Hymenoptera (Pompilidae and Formici'dae) as hosts for

Parathrombium. Trombidium. on the other hand, are primarily parasites of Lepidoptera, Coleoptera and to lesser extent Diptera,

Hemiptera and various other arthropods (Araneae, Hymenoptera,

Homoptera and others) (Welbourn, 1983).

The subfamily Allothrombiinae is defined by the absence of setae on the palpal femur (5e) and genu. Feider (1979) listed seven genera in what I recognize as a subfamily. Larvae of the following nominate genera were examined: Allothrombium Berlese, 1903, Dinothrombium

Oudemans, 1910, Isothrombium Andre, 1949, Angelothrombium Newell and

Tevis, 1960 and Aphithrombium Childers and Vercammen-Grandjean, 1981.

Based on comparison of Isothrombium and Angelothrombium with reared specimens of I), tinctorum L. from Newell's collection (now part of the USNM) both Isothrombum and Angelothrombium are congeneric with

Dinothrombium. Tevis and Newell 1962 stated that Angelothrombium was a junior synonym of Dinothrombium but both Vercammen-Grandjean (1973) and Feider (1979) listed Angelothrombium in their classifications while ignoring Dinothrombium.

Dinothrombium is distinguished by the position of the PL setae in relation to the prodorsal trichobothridia bases, where in

Dinothrombium the distance between the PL setae is less than the distance between the prodorsal trichobothrial bases (26). Hosts for

Dinothrombium are Orthoptera and Tevis and Newell (1962) summarized the biology of I), pandorae from the deserts of California.

Allothrombium is distinguished by having the scutellum much narrower than the the width of the prodorsal sclerite. Both genera are united 82

by the presence of second sigma on genu legs II and III. Welbourn

(1983) listed 25 host species with 21 from Homoptera, mostly Aphidae.

A new genus, Nanothrombidium was discovered parasitizing

Thysanoptera in Italy; this differs from other Allothrombiinae in not

having the additional sigma on genu legs II and III but can be

distinguished by the presence of seta la on coxal field leg I without

barbs (52b).

Childers and Vercammen-Grandjean (1981) named a new genus,

Aphithrombium. closely related to Allothrombium but differing in only

the absence of the paraxial claw in pretarsus leg III. Examination

of the paratypes and the observed variation of the paraxial claw in

Allothrombium and other Trombidiidae indicate this synapomorphy is not sufficient to separate this species as a new genus.

Neothrombiidae

The monophyletic group, Neothrombiidae, known only from one genus prior to 1976 has been placed in various familial groups at different levels, under different names by different authors (Feider,

1955, 1959; Vercammen-Grandjean, 1973; Robaux et. al, 1976). This family is defined by six larval synapomorphies: 1) femur leg II with four setae (66d); 2) microseta k absent on genu leg II (81); 3) palpal genu without seta (7d); A) tarsus legs II without eupathidia

(99); 5) empodium legs I, II and III with distally expanded tip

(118); 6) prodorsal sclerite and scutellum striate (28b, 39b). While many of the larval characters have evolved at least once in other 83

Trombidioidea this combination of synapomophies is unique to the

Neothrombiidae. The use of adult characters is of questionable value because postlarval instars are known only for three of the terminal taxa (Neothombium Oudemans. 1910, Ceuthothrombium Robaux. et, al.,

1976 and Acridothrombium) in the subfamily Neothrombiinae. The discovery and rearing of additional postlarval instars will provide a good test of the hypothesized relationships.

Phylogenetic analysis of the family used 23 characters (Table 6) resulting in three hypothesized monophyletic and very different subfamilies arising from an unresolved trichotomy (Figure 10). The

Aethethrombidiinae exhibit a number of autapomorphies (palpal femur-genu fused (9), chelicera needle-like (15), palpal tibial claw entire (10e)), in addition, both members of this monophyletic group have neotrichous coxae (49c, 50c, 51c). The two included genera are each known from single specimens taken from litter samples in the neotropics (Brasil and Jamaica).

The subfamily Aegalothrombldiinae represented by the genus

Aegalothrombidium, with two species and four specimens, exhibits a number of autapomorphies: 1) reduction of number of solenidia on genu

(78b) and tibia (85) of leg I; 2) addition of a dorsal seta on tibia leg I (82d); 3) PL setae between eyes (off prodorsal sclerite) (25).

Both species are known only from the coast of Mexico, Baja California

Sur and Guatemala as parasites of Phaleria (Coleoptera,

Tenebrionidae). Table 6. Character States used on the Phylogenetic Analysis of the Neothrombiidae.

Taxa Charactera 10 15 22 23 24 25 28 29 39 40 45 49 50 51 57 77

Naothroablinae Neothroablua a a a b b a d Acrldothroablua d a a b c a b Outhothroablua d a a b b a d keraathroablua 7 a a b b a d Adelothroabldlua d a c a b a d

Aethothroabllnae Mggalothroablaa a a a Ae thothroabIdlua a a a

Aegalothroabllnae Aegalothroabldlu 1 s s m HI ! o % * * I *

M e tie ••e

lie t4c lie

4»e 40c 41b

II

AETHETHROMilDJINAE NEOTHROMBIMAE 0, 10*. 14 II, 101 40c. tOc. tie

NEOTHROMBIIDAE

n , * • . h i, n

M l , II, 110

FIGURE 10. Cladogram of Hypothesized Relationships within the

Neothrombiidae. 86

The subfamily Neothrombiinae is represented by five closely

related genera (only two known prior to my study) and at least 9

species. The subfamily is defined by the two larval and three

postlarval synapomorphies: 1) coxal field leg II with two setae

(50a); 2) prodorsal sclerite rectangular in shape (29); 3) prodorsal

sclerite expanded in the postlarval instars; 4) idiosomal setae in

postlarval instars with basal spines; 5) accessory claw in postlarval

instars reduced. Most genera in the Neothrombiinae are similar in

morphology and are orthopteran parasites as larvae. Kermathrombidium

known only from Nasutitermes nests in Panama has the only

non-orthopteran host. The distribution of the Neothrombiinae, as

currently known, is Holarctic, Central America and Africa.

Post-larval instars have been described for only Neothrombium but

they are known for Acridothrombium and Ceuthothrombium.

The genus Neothrombium Oudemans, 1909 was proposed for

Allotrombidium neglectum Bruyant. 1909 described from a European

Gryllotalpa gryllotalpa L. Five species of Gryllotalpa were examined

at the British Museum (Natural History) and only two species, G..

gryllotalpa L. and £. unispina Saussure had mites attached between

the coxae. Feider (1948) proposed Mycterothrombidium for the adult

of what is now known as Neothrombium and in 1955 synonomized it with

Neothrombium. The subfamily Neothrombiinae was proposed by Feider in

1959. The type species was redescribed by Robaux (1972) based on

specimens from the Oudemans Collection. In 1976 Robaux, Webb and

Campbell described a second genus, Ceuthothrombium cavatum from

between the coxae and on the neck of cave crickets (Ceuthophilus) from caves in Texas, New Mexico and northern Mexico. At the sam6

time the authors redefined the subfamily Neothrombiinae, Ignoring the

previous work of Feider. Webb, et. al. (1977) reported rearing the

undescribed deutonymphs and discussed the possible cave adaption of

Ceuthothrombium. Ceuthothrombium larvae have adapted to the cave

environment by loss of pigment and reduction of the prodorsal eyes to

a single eye (occasionally absent). Preliminary work with the

postlarval instars indicate they are restricted to the cave, although

not yet recovered from cave soils, by reduction in pigment and

absence of eyes. This genus probably entered the cave environment

via their host, Ceuthophilus. which leaves the cave nightly to feed.

The discovery of a widespread genus, Acridothrombidium, in the

southwestern United States provides a sister group for the

cave-adapted Ceuthothrombium.

Acridothrombium is a widespread parasite of acridid grasshoppers

in the western United States with at least two undescribed species in

California, New Mexico, Texas and Arizona. All species occupy

similar sites of attachment around the coxae, but are also found in

the cervical region of the host. Based on studies of limited

collections it appears that A^. chiricahuaie new species, is

widespread in the arid regions of Arizona, New Mexico and west Texas

and another undescribed species is limited to more mesic conditions

found in higher elevations. Unfortunately, host information is not

available for most specimens, making host comparisons impossible.

More extensive collections and host identifications are needed to

determine species distributions and relationships. 88

A second new genus, Kermathrombidium, was discovered on the head

of worker termites (sterile females) of three species of Naautitermes

in Panama. Infestation rate in individual nests was less than 1 per

cent (B. Thorne personal communication). A third new genus,

Adelothrombidium. is known from a few specimens, representing two

species, from South African gryllacridid, Libanasidus vittatus, differs from the other Neothrombiinae in that the prodorsal trichobothria are anterior to the PL setae. I am aware of one additional undescribed genus of Neothrombiinae from Africa, but the specimens were not available for this work.

The post-larval instars of Neothrombium were described by

Willmann (1950) and Feider (1950,1955), but were not available for this study. The postlarval instars of Acridothrombium are similar to

Neothrombium in color (red), setal shape, shape of the prodorsal sclerite and palp to Neothrombium. Ceuthothrombium, as suggested by

Webb et. al. (1977) is adapted for cave life by loss of pigment, loss of eyes, reduction on the prodorsal sclerite to width of the crista metopica and the loss of basal spines on the dorsal idiosomal setae.

The remainder of the Trombidioidea (Calothrombiidae,

Eutrombidiidae and Microtrombidiidae) are characterized by four larval and two postlarval synapomorphies: 1) the presence of a buccal ring on the anterior end of gnathosoma (14); 2) femur leg II with five setae (66c); 3) scopa on tarsus leg III well developed

(occasionally secondarily lost) (101); 4) lophotrix well developed

(occasionally secondarily lost) (102); 5) palpal tibia of postlarval instars with ventral spines (118); 6) dorsal spines on palpal tibia 89

of postlarval instars in one or two rows (117b,c). In addition, the

dorsal idiosomal setae of postlarval instars range from setiform to

the highly derived setae of Cercothrombiini (Microtrombidiidae).

Calothrombiidae

The genus Calothrombium was proposed for an adult trombidioid mite from "nidi Dociostaurus maroccansus" from Foggia, Italy by

Berlese, 1918 as a new name for a Parathrombium paolii Berlese, 1918 because the name Parathrombium Bruyant, 1910 was preoccupied. The placement of this genus has been a case history of recent workers basing their classification on the errors of earlier workers. Thor

(1935) placed the genus in Microtrombidiinae, but Thor and Willmann

(1947) placed it in the Tanaupodinae. While Womersley (1942) recognized Thor’s (1935) placement, subsequent workers (Feider, 1959;

Vercammen-Grandjean, 1973) continued to place it with the tanaupoids

(Feider, 1959) and Trombellidae (Vercammen-Grandjean, 1973) and in

1979 Feider elevated it to a superfamily. In 1984 I examined the type of Calothrombium paolii (Berlese, 1918) in the Berlese

Collection housed at the Istituto Sperimentale per la Zoologia

Agraria, Florence, Italy. Based on my comparison I will retain the family status proposed by Feider and concur, in general, with the relationship with the Microtrombidiidae implied by Thor's 1935 classification. The family can be defined by three postlarval synapomorphies: 1) prodorsal sclerite wide (122); 2) idiosomal setae bifurcate (124e); 3) two rows of 8-11 spines with both starting at or 90 near the palpal tibial claw (117c). Phylogenetically this group falls between the Neothrombiidae and the Eutrombidiidae.

The larva of Calothrombium is unknown, but Paoli (1937) described a mite Metathrombium danuium from Steropleurus cavannae.

Dociostarus maraccanus. Oedaleus decorus. Prionotropis hystrix appula, Dectius albifrons and D. loudoni all from southern Italy and suggested it was the larva of Calothrombium. I was not able to locate Paoli's type, but while examining Orthoptera at the British

Museum of Natural History two specimens, each representing a new species, of a new genus (Dectothrombidium) were found on J). albifrons from Spain and Morocco. Welbourn (1983) placed M. danuium in the genus Trombidium based on the chaetotaxy of the coxal field and

Paoli’s description, but rexamination of Paoli's description and figures based on the new material indicates the species should be transferred to Dectothrombidium.

Dectothrombidium can be defined on the basis of four synapomorphies: 1) bifurcate scopa on tarsus leg III (lOlf); 2) two setae in coxal field leg II (50b); 3) lophotrix on tarsus leg III pilose (103j); 4) two idiosomal scutella, one each in dorsal setal rows C and D (42b). It is very likely that Dectothrombidium is the larva of Calothrombium but until larval and postlarval instars are reared the names should remain separate.

The Eutrombidiidae and Microtrombidiidae are united by a single synapomorphy involving the modification of the subcapitular setae

(scl) into a variety of nonsetiform shapes (rounded, spinlike, minute and palmate (4b, c, e)). In addition, the scopa and lophotrix are 91 usually well developed.

Eutrombidiidae

The Eutrombidiidae was first proposed as a subfamily by Thor in

1935 primarily on the basis of a posterior opisthosoraal sclerite in the postlarval instar. Feider (1959) elevated the group to family level, but subsequent workers have followed Thor (1935a).

The larval Eutrombidiinae are characterized by five synapomorphies: 1) subcapitular setae (scl) hypertrophied (4c,d); 2)

AM seta nude (22b); 3-5) setae lb, 2a and 3b on coxal fields legs I and III respectively thick, short and usually bifid (53, 54, 55). In some species of Hoplothrombium the setae on all coxal fields are setiforra distally while retaining a thick basal portion. The modification of setae lb on coxae legs I and III and seta la on coxa leg II into stout bifid setae are autapomorphic in this group.

My comparative study of the family was based on examination of

17 characters (Table 7) and indicates there are six larval genera in two subfamilies which show a diverse array of forms including apparent convergences in the morphology of tarsus leg III. Figure 11 is a cladograra of the hypothesized relationships within the

Eutrombidiidae. The type genus Eutrombidium Verdun, 1909, proposed for Trombidium trigonum Hermann, 1804, appears to be the most primitive eutrombidioid as well as the most widespread and least specialized of all the taxa studied. The lophotrix in the

Eutrombidiinae primitively consists of three large basal branches and Table 7. Character States used on the Phylogenetic Analysis of the Eutrombidiidae.

Taxa Characters

4 24 41 42 45 48 54 56 66 78 79 80 81 101 103

Eutroabldllnae Eutroabldlua c a a a a b b a c a a a a b e Scutodeaaua c a c a a b a a c a a a a b e AtODOthroabldlua d a a a a a b a d b e a b d g Mllllothroabidiua d b a a a a b b d b a/e a b c b

Hoplothroablinae CiTDtothrobldiua d a b a c a b b c b a b b d f HoDlothroablua d a a c/d a a b/a b c a a a b d f 93

64b 41c 70c 20b 42c.

103a 66b 4 6 c

61b 76b

101c

103h

EUTROMBIDIINI MILLIOTROMBIDIINI

4 c . 46 4d, 20c, 30d

HOPLOTROMBIDIINAE

EUTROMBIDIINAE 4d, 66b, 61. 60a.b

61b, 101d, 1031 103a

EUTROMBIDIIDAE

4c,d. 22b, 63b,e

64b,c, 66b,c

FIGURE 11. Cladogram of Hypothesized Relationships within the

Eutrombidiidae. 94

short setules on the distal half of the seta. This form has been modified within the Eutromibidiinae with Scutodesmus having one long branch, one medium length branch and the remainder of the seta with

short setules. In the Milliothrombiini, for example,

Atopothrombidium has the lophotrix reduced to a long, thick single clawlike setae and in its sister genus, Milliotrombidium. the lophotrix similar to Atopothrombidium except bifurcate in M.

japanensis while M. milliopdoium Shiba retains the three branches with barbs.

The Eutrombidinae are cosmopolitan in distribution with nearly all larvae parasites of Orthoptera. Welbourn (1983) listed 48 orthopteran species as hosts for three species of Eutrombidium. Of these, 39 hosts were in the family Acridiae, 6 Tettigoniidae, two

Gryllidae and one mantid. Hosts are unknown for Scutodesmus.

Little is known of the habits of Milliotrombidium as one species

(M. milliopodium) is known from a millipede in Malaysia (Shiba, 1976) and a second species (M. japanensis) is known from Rhaphidophoridae of the genus Diestrammena in Japan. It is possible the millipede host is accidental. Atopothrombidium is known only from the "cave weta" in New Zealand.

The systematics of Eutrombidium at the species level is chaotic with most records in Europe, North America, and India listed as E. trigonum (Riziv & Aziz, 1974; Kamal, et. al., 1976, Severin, 1944).

Preliminary work with the North American and African Eutrombidium indicates considerable larval diversity with many additional species remaining to be identified. Despite the systematic chaos the biology 95

of E. locuBtarum (Walsh) in North America was well documented by

Severin (1944).

The genus Hoplothrombium Ewing was placed in the Trombidiidae by

Vercammen-Grandjean (1973) baed on the atypical setae on coxa leg II.

Examination of other Hoplothrombium species indicate it is phylogenetically closer to the Eutrombidiidae.

The Hoplothrombiinae, with two genera Hoplothrombium and

Cryptothrombidium NEW GENUS, are characterized by the bifurcation of tarsus leg III (110), absence of paraxial claw on tarsus leg III

(108c), secondary loss of a developed scopa (lOle), presence of a preanal sclerite (61), a multibranched lophotrix (103f) and coxal field leg III with extra sclerotized area (56). Despite the similarity in appearance of tarsus leg III in the genus

Milliotrombidium and the Hoplothrombiinae, it appears that the divided tarsus was independently derived in both groups.

Milliotrombidium has several synapomorphic characters which suggest a closer relationship with Atopothrombidium and the Eutrombidiinae than

Hoplothrombiinae (no preanal sclerite, femur leg II with four setae, lophotrix with 1 to 3 branches and paraxial claw present). In addition to sharing the general morphology of tarsus leg III with the

Hoplothrombiinae, the coxal field of leg III in Milliotrombidium has additional sclerotization, although not as extensive, as in the

Hoplothrombiinae. The function of this sclerotization is unknown but it may be necessary for additional muscle attachment needed to move the enlarged leg and tarsus III. Members of the genus Hoplothrombium are parasites of Coleoptera. 96

In the Hoplothrombiinae the genus Cryptothrombidium NEW GENUS, known from the cave crickets, Hadenoecus, in Kentucky and Tennessee, appears to have moved into the cave habitat as suggested by the loss of eyes (45c) and reduction of the scutellum (41b) to a narrow sclerite (similar to Ceuthothrombium in Neothrombiidae).

Microtrombidiidae

The Microtrombidiidae, with more than 50 nominate genera is the largest and most diverse trombiuioid family. While the hypothesized subfamily and tribal groups are monophyletic the placement of genera will be subject to further analysis and testing as more larval forms become available. The microtrombidiids have been placed at several familial ranks (Vercammen-Grandjean, 1973; Feider 1955, 1979,

Welbourn, 1984). Until recently the group was loosely defined on postlarval characters (Welbourn, 1984). Here the group is defined on the basis of a single unique larval synapomorphy, the position of famulus on tarsus leg I proximal to omega I (91). While the postlarval instars are still difficult to define at the family level, the subfamilies are more easily defined. The family is divided into two monophyletic subfamilies Microtrombidiinae Thor, 1935 and

Feideriinae Vercammen-Grandjean and Cochrane, 1974 (not

Vercammen-Grandjean, 1973, see Welbourn, 1984). Figure 12 presents a cladogram of the hypothesized phylogenetic relationships in the

Microtrombidiidae. Table 8 lists the characters used in my analysis of the Microtrombidiidae. Table 8. Character States used on the Phylogenetic Analysis of the Microtrombidiidae

Taxa Characters 4 22 28 39 44 47 52 68 69 78 79 81 101 102 105 108

Nanriaula b a a a a b a a a d b a b a d a Retlculatroabldlua b a a a a b a a a d b a a a d a Preoathroablua b a a a a a a a a a a a b a d a Valaothroablua b a a a a a a b b a a b d c b Goaothroablua b a b b a a a b b a a b d c _ b Achelothroabldliia e b e b a a a b b a a a d c - a Atractothroabldl.ua e b b b a a a b b a a b d c b Ettauelleria a b b b b a a c a a a b a It a PlatTtroabldlua e b b b b a a b b a a a b a b a Mlcrotroabldlua e b a a b a a c b a a a b a d a Diathroablua e b a a b a a 9 • a a a b a d a CaaDTlothroablua e b b b a b b b a a a a b a d a Trlcotroabldlua e b c c b a a b b a a b a d a Llthothroabidiua e b b b a a b b a a a b b a d a Villaannella a b b a b c a b b a a b a d a Cercothroablua a a a a a a a a a a a a b a d a 4b 7 8 d 108 37 4 0 7 9 9 1 c

102c

6 2 b

81b, 105

28 b

28b. 86.89

MICROTROtftSDHNI CERCOTHMOMBI1MI

103b losa

MCBOTROHBIOUNAE

4b

MICROTROMBIDIIDAE

FIGURE 12. Cladogram of Hypothesized Relationships within the

Microtrombidiidae. 99

The subfamily Microtrombidiinae Thor, 1935a, b, with more than

40 nominate genera, is defined on the basis of two larval synapomorphies, the AM seta on the prodorsal sclerite is without barbs (22b) and scl setae palmate (4e). The subfamily is subdivided into two tribes each based on a larval and a postlarval character.

The Microtrombidiini are defined by a larval lophotrix without long basal branches (103d). The lophotrix is more setiform and secondarily lost in some microtrombidine genera. The dorsal idiosomal setae in the postlarval instars are usually simple to inflated while retaining their setiform appearance. The

Cercothrombiini are defined by a lophotrix which has a long basal branch and 2-3 shorter branches (103d) and postlarval instars with dorsal idiosomal setae that are globose or other complex nonsetiform shapes. The palpal femur and genu and the basal leg segments in postlarval instars have numerous brushlike setae.

The Microtrombidiini recognized in this analysis includes five monophyletic genera (Ettmulleria Oudemans, 1911, Platvtrombidium

Thor, 1936, Microtrombidium Haller, 1882, Atractothrombium Feider,

1952 and Achelothrombidium). This group is difficult to define on the basis of synapomorphies because of generic variation and small sample size. The members of this group are cosmopolitan and parasitize a variety of insects with Diptera being the most common hosts.

The genus Microtrombidium Haller, 1882, like many trombidioid genera, was described from the adult. While many authors have speculated on the larva of Microtrombidium (Vercammen-Grandjean, 1973; Robaux 1974, 1973; Feider, 1950) no one has collected or reared the type species (T. pusillum Hermann, 1804). The original description of M. pusillum (Hermann,1804) is unrecognizable beyond

Trombidioidea, and the oldest specimens originally identified as M. pusillum are in the Berlese Collection. These 1884 specimens, although renamed M. italicum Berlese, 1912, were used to provide a diagnosis for the postlarval Microtrombidium. Collecting near

Florence, Italy during May 1984 revealed adult M. italicum from which larvae were reared. Comparison with other specimens of

Microtrombidium £. j3. in the the Berlese and Andre Collections support my conclusion to define Microtrombidium based on M. italicum and its reared larvae. The larva of M. italicum is congeneric with the larval genus Megophthrombium Mullen and Vercammen-Grandjean,

1978. Based on these tenative conclusions Microtrombidium is defined by the following synapomorphies; 1) lophotrix without long basal branches, with one long distal branch; 2) dorsal seta on femur leg I and II nude and located on anterior end of segment; 3) idiosomal setae in postlarval instars setiform; 4) dorsal palpal tibia with two short rows of 2-4 spines; 5) palpal tibia without large ventral spines. Nothing is known of the biology of Microtrombidium other than that the larva of one species (=Megophthrombium) is a parasite of Culicidae (Diptera) in North America.

Vercammen-Grandjean (1973) placed Atractothrombium in the

Feideriinae on the basis of missing empodium on pretarsus legs I and

II and the missing microseta k on genu leg II. Despite these two shared synapomorpies with the Feideriini, Atractothrombium possess a 101

palmate scl setae, nude AM seta and a large ventral spine on the

palpal tibia of postlarval instars.

The tribe Cercothrombiini, as stated above, is defined on the

morphology of the larval lophotrix, the morphology of the dorsal

idiosomal setae and the brushlike palpal setae in the postlarval

instars. The five larval genera included in this groups are very

similar and differ in the presence or absence of striations on the

prodorsal sclerite and scutellum, shape of setae on the coxal fields and size of idiosomal setae in rows D, E, F and H.

The genus Diathrombium Moser and Vercammen-Grandjean, 1979 is known only from Dentroctonus frontalis Zimmerman (Coleoptera,

Scolytidae) and is distinguished by the reduction of the scl setae to a few barbs. The larva of Campylothrombium Krausse, 1916 was described by Robaux (1974) and is distinguished by the striations on the prodorsal sclerite. The host is unknown and the genus is known primarily from Europe with additional species from tropical Africa and southeast Asia which could represent different genera.

Lithothrombidium is known from a species originally described by

Banks, 1916 from California. The species was recollected and reared in 1982. Nothing is known of the biology of this mite other than the adults collected in 1982 were associated with ants.

Cercothrombium Methlagl, 1928 was described from free-living larvae in Austria and remained uncollected and unknown until 1984 when it was reared from field collected Camerothrombium from Great

Britain which deposited eggs from which larval Cercothrombium emerged. Cercothrombium Methlagl, 1928 predates Camerothrombum Thor, 102

1936. Nothing is known of the hosts or biology of these mites.

Trichotrombidium Kobulei, 1950 was named for T. muscarum Riley, 1878 parasitizing Musca domestica worldwide, especially in tropical regions. Willmannella Feider, 1951 originally named from the adult

(M. (Enemothrombium) racovitzai Feider, 1951) with the larva reared later. A species described by Robaux (1975) referred to

Compsothrombium johnstoni is transferred to Willmannella based on examination of type of Compsothrombium Andre and comparison with larval Willmanella from the Feider collection making it the first record for the genus in North America. Nothing is known of the hosts or habits of this mite.

The subfamily Feideriinae Vercammen-Grandjean and Cochrane, 1974

(not Vercammen-Grandjean, 1973; see Welbourn, 1984) was proposed by

Vercammen-Grandjean (1973) to include the the invalid new genus,

Feierium and three other genera (Gonothrombium, Platytrombidium and

Atractothrombium). Platytrombidium and Atractothrombium have been transferred to the Microtrombidiini. The subfamily is redefined on the basis of a single larval synapomorphy, the reduction of the scl setae to minute spines (4b). In addition the AM setae are barbed.

The subfamily consists of two monophyletic groups: Feideriini and

Manriquiini Welbourn, 1984.

The Feideriini Vercammen-Grandjean and Cochrane, 1974 are characterized by four synapomorphies: 1) microseta k absent on genu leg II (81); 2) pretarsus leg III without lophotrix (102); 3) pretarsus leg III without scopa (lOle); 4) pretarsus legs I and II without empodium (105). There are two larval genera, one with a correlated adult, included in the tribe. The genus Feiderium was

named and the type species (M. tarnavense Feider, 1949) designated by

Vercammen-Grandjean (1973) and but neither were described nor a

diagnosis provided. In 1974 Vercammen-Grandjean and Cochrane

described a new species which they placed in a new subgenus

(Parafeiderium) on the basis of a missing claw on tarus leg III.

Feiderium. made available by Vercammen-Grandjean and Cochrane, 1974

is a parasite of Ceratopogonidae (Diptera) and has been suspected as

being the larva of Valgothrombium Willmann, 1950. While no larva has

been reared in Europe providing direct evidence that Valothrombum is

the senior synonym of Feiderium, evidence from specimens from Panama

and North America indicate they are congeneric. Michener, 1946

described M. fluminis from reared larvae and field collected adults

from Panama. A comparison of M. fluminis with the type of

Valgothrombium (Ottonia valga George, 1909) from the British Museum

(Natural History) show the adults to be congeneric and the larvae to

be congeneric with Feiderium. Additional evidence comes from several

field collected adult trombidioids from Oregon during the spring,

1982 and placed together. After several egg masses were deposited,

the adults were removed and identified. When the eggs hatched, all

specimens save one were Atractothrombium as were all the adult but

one adult identified as Valgothrombium. Based primarily on

Michener's work in Panama and the additional data from Oregon I am

placing Feiderium in synonomy with Valgothrombium Willmann, 1940.

Parafeiderium remains as a subgenus in Valgothrombium. Larval

Valgothrombium is defined by the reduced barbs on the coxal field 104 setae and the shape of the prodorsal sclerite and scutellum. Larvae are known to parasitize Ceratopogonidae (Diptera) over a wide range

(Holarctic, Neotropical and Africa) and are probably cosmopolitan.

Only one larval Valgothrombium is known from Europe; a species found free-living and described by Willmann (1958) as Allothrombium ignotum is here placed in Valgothrombium. A second larva, that of

Microtrombidium spasiscutum Robaux, 1974 from Argentina, is also a member of the genus Valgothrombium and only the second known

Nortropical species. The genus Valgothrombium is defined by the following synapomorphies: 1) setae on coxal fields short with one or two long barbs; 2) prodorsal sclerite with three posterior lobes; 3) scutellum with medial anterior projection; 4) postlarval instars with prominant striations on anterior end of prodorsal sclerite; 5) dorsal idiosomal setae inflated.

The second genus Gonothrombium was described by Feider (1948) and known from two nominate species, G. oudemansianum and G. bimaculatum, from eastern Europe. A third species .G. bequaerti

(Fuller and Wharton, 1951) was hidden under several misidentifications and wrong names until Vercammen-Grandjean (1973) placed it in Gonothrombium. Nothing is known of the biology of

Gonothrombium. Gonothrombium is characterized by the synapomorphy of a multisetose scutellum (40).

The tribe, Manriquiini Welbourn, 1984, originally defined on the basis of few taxa, is redefined on the basis of additional specimens and a new genus. The synapomorphies defining this group are the presence of a scopa (101b), the lophotrix with one very long basal 105

branch and the dorsal idiosoma with reticulate pattern or secondary

scleritization posteriorly. See Robaux (1967b, c) for a

redescription of the adult.

Manriquia Boshell and Kerr and Reticulatothrombium Shiba are

closely related genera with Reticulatothrombium in Malaysia (and

undescribed species in Africa) and Manriquia in the Neotropics. They are united by the following synapomorphies: genu legs II and III each with two solenidia (sigma); genu leg I with two or three (2-3) solenidia; dorsal opisthosoma with distinct reticulate pattern; all known specimens have been either collected in litter samples or in the case of the type species, reared from eggs. Nothing is known of the habits or hosts of these mites.

The genus Pygothrombium was also found in litter samples in

Malaysia and the neotropics as well as a malaise trap in New Guinea.

This genus is characterized by the autapomorphicpresence of a pygidal plate bearing two pairs of setae. Nothing is known of the biology or hosts of these mites. KEY TO THE GENERA OF TROMBIDIINA

(LARVAE)

1. Dorsum with a prodorsal sclerite (scutum) and at least one

(1) other sclerite (scutellum) bearing at least one (1) pair

of setae; tibia leg III without phi; parasitic on arthropods

Superfamily Trombidioidea ...... 2

— Dorsum with prodorsal sclerite but lacking scutellum; tibia

leg III with at least one (1) phi; parasitic on vertebrates

or arthropods ...... 39

2 (1). Coxal field II each with at least two setae...... 3

— Coxal field II each with one (1) seta...... 18

3 (2). Femur leg II with four setae; idiosoma with only two

sclerites (prodorsal and scutellum) ...... 5

— Femur leg II with five setae; idiosoma with more than two

sclerites ...... A

106 107

4 (3). Tarsus leg III bifurcate; lophotrix large and-multibranched;

idiosoma with four plates; scopa absent; host: Coleoptera;

Nearctic ...... Family Eutrombidiidae ...... Subfamily

Hoplothrombiinae (in part) ......

...... Hoplothrombium Ewing, 1925

Tarsus leg III normal; lophotrix setiform; idiosoma with two

plates; scopa bifurcate; host: Orthoptera; Southern Europe

and Northern Africa . . . Family Calothrombiidae ......

...... Dectothrombidium NEW GENUS

5 (3). Genu leg II and III each with a single sigma; palpal femur

usually with one (1) dorsal seta; palpgenu without setae . .

...... 7

Genu leg II and III each with two (2) sigma; palp femur and

genu without setae .... Family Trombidiidae (in part) . .

Subfamily Allothrombiinae ...... 6

6 (5). Distance between prodorsal trichobothria (SW) less than

distance between the posterior lateral setae (PL) on scutum

(SW

Homoptera and Orthoptera; cosmopolitan ......

...... Allothrombium Berlese, 1903

Distance between prodorsal trichobothria (SW) greater than

the distance between the PL on the scutum (SW>PW); femur leg 108

II with four (4) branched setae; host: Orthoptera;

cosmopolitan ...... Dinothrombium (Oudemans, 1910)

7 (5), Tibia and femur leg I each with five (5) branched setae;

genu legs II and III each with three (3) branched setae;

microseta (k) present on genu leg II ......

Family Trombididae .... Subfamily Trombidiinae...... 8

— Tibia and femur leg I each with six (6) branchedsetae; genu

legs II and III each usually with two (2) branched setae

(except Acridothrombium with 4); microseta (k) absent on

genu leg II . . . . Family Neothrombiidae...... 11

8 (6). Seta la on coxal field I setiform; microseta (k) on genu leg

III absent ...... 9

— Seta la on coxal field I a highly modified comb-like seta;

microseta (k) present on genu leg III; hosts: Homoptera,

Hemiptera, and Hymenoptera; Holarctic, Ethiopian, Australian

...... Parathrombium (Bryant, 1910)

9 (8). Seta la on coxal field leg I nude; prodorsal trichobothria

flagelliform; seta on palpal femur nude or absent . . . 10

— Seta la on coxal field leg I branched; prodorsal

trichobothria expanded; seta on palpal femur branched; host

unknown; eastern North America

...... Oligothrombidium NEW GENUS 109

10 (9). Palpal femur with single nude seta; anterior end of

prodorsal sclerite nearly straight; subcapitular setae (scl)

short with heavy setules; hosts: Diptera, Lepidoptera,

Homoptera, Coleoptera; cosmopolitan ......

...... Trombidium Fabricius, 1775

— Palpal femur without setae; prodorsal sclerite with anterior

extension tapering distally; subcapitular setae (scl) long

and setiform; hosts: Thysanoptera; Italy ......

...... Nanothrombidium NEW GENUS

11 (7). Palpal tibial claw bifurcate; palpal genu femur separate;

coxal field leg I with two (2) setae...... 12

— Palpal tibial claw entire; palpal genu and femur fused along

outer surface; coxal field leg I multisetose (6-11 setae) .

.... Subfamily Aethethrombidiinae ...... 17

12 (11). PL setae on prodorsal sclerite; genu and tibia leg I each

with at least two solenidia ......

. . . Subfamily Neothrombiinae ...... 13

— PL setae off prodorsal sclerite; genu and tibia leg I each

with only one solenidion; host: Phaleria (Coleoptera); Baja

California, Guatemala .... Subfamily Aegalothrombidiinae

...... Aegalothrombidium NEW GENUS 110

13 (12). Prodorsal trichobothria placed posterior to PL setae on

prodorsal sclerite...... 14

— Prodorsal trichobothria placed anterior to PL setae on

prodorsal sclerite; host; Orthoptera; South Africa ......

...... Adelothrombidium NEW GENUS

14 (13). Scutellum with two setae; eyes absent or reduced to a single

eye on each side of prodorsum; prodorsal sclerite and

scutellum without striations...... 15

— Scutellum multisetose (6-13); two pair of eyes with ocular

plates; prodorsal sclerite and scutellum with striations . .

16

15 (14). One pair of eyes 1/1 (occasionally absent) without ocular

plates; anteromedial (AM) setae branched; prodorsal

trichobothria flagelliform; host; cave dwelling crickets of

the genus Ceuthophilus; southwestern US and northern Mexico

...... Ceuthothrombium Robaux, et al, 1976

— Eyes absent; anteromedial (AM) setae forked; prodorsal

trichobothria expanded; host: termites; Panama......

...... Kermathrombidium NEW GENUS

16 (14). Coxa leg II each with at least four (4) setae; genu leg II

and III with four (4) branched setae; host Orthoptera Ill

(Acridldae); western US and Mexico......

...... Acridothrombium NEW GENUS

— Coxa leg II each with two (2) setae; genu leg II and III with

two (2) branched setae; host: Orthoptera (Gryllotalpidae);

Palearctic, Africa ...... Neothrombium Oudemans, 1910

17 (11). Tibia legs II and III each with six (6) setae; genu legs I,

II and III with numerous sigma (7-22-28); tarsus leg I, II

and III neotrichous; host unknown; Jamaica ......

...... Megalothombium. NEW GENUS

— Tibia legs II and III each with five (5) setae; genu legs I,

II and III with normal number of sigma (2-1-1); tarsus leg

I, II and III with 18-14-13 setae respectively; host

unknown; Brasil...... Aethothrombidium NEW GENUS

18 (2). Genu leg II and III each with two (2) branched setae; femur

leg I with six (6) branched setae; tarsus leg I with maximum

of one ventral eupathid; scutellum not usually reduced;

claws on pretarsus III may be modified (asymmetrical) . . 19

— Genu legs II and III each with three (3) or four (4)

branched setae; femur leg I with five (5) branched setae;

ventral surface of tarsus leg I may have five (5) or more

eupathids; scutellum reduced to small platelet with two

setae; claws on pretarsus III symmetrical; hosts: Homoptera;

Holarctic .... Family Podothrombiidae ...... 112

Podothromblum (Berlese, 1910)

19 (18). Setae lb and 3b on coxal fields legs I and III respectively

stout and usually bifid; tarsus leg III greatly modified

(asymmetrical); famulus (e) on tarsus I usually distal to

omega I ...... Family Eutrombidiidae...... 20

— Setae on coxa legs I, II, or III setiform; tarsus leg III

may or may not be modified; famulus (e) tarsus legI usually

proximal to omega I . . . . Family Microtrombidiidae. . . 24

20 (19). Tarsus leg III entire; tarsus leg III with empodium, normal

claw and modified claw...... 21

— Tarsus leg III bifurcate; tarsus leg III with only empodium

and modified c l a w ...... 23

21 (20). Seta 2a coxal field leg II thick and bifid; all setae on

tibia leg I setiform; prodorsal sclerite and scutellum not

covering entire dorsal idiosoma ...... 22

— Seta 2a coxal field leg II spinelike; some setae on tibia

leg I modified into comblike setea; prodorsal sclerite and

scutellum in unengorged specimens completely cover dorsum

pushing setae onto venter; host unknown; Africa ......

...... Scutodesmus NEW GENUS

22 (21). Femur leg II with five (5) branched setae; one sigma on 113

genus legs II and III; subcapitular setae (scl) spiniform;

host: Orthoptera; cosmopolitan ......

...... , ...... Eutrombidium Verdun, 1909

— Femur leg II with four (4) branched setae; without sigma on

genu legs II and III; subcapitular setae (scl) trilobed;

host: Orthoptera; New Zealand ......

...... Atopothrombidium NEW GENUS

23 (20). Femur leg II with five (5) branched setae; scopa absent;

with preanal sclerite; one sigma on genu leg III; eyes

absent; prodorsal trichobothria flageliform; host

Orthoptera; known only from caves in Kentucky and Tennessee

...... CryptothrombidiumNEW GENUS

— Femur leg II with four (4) branched setae; scopa nude;

without preanal sclerite; without sigma on genu leg III;

eyes present; prodorsal trichobothria slightly expanded;

hosts: millipede, Orthoptera; Malaysia & Japan ......

...... Milliotrombium Shiba, 1976

24 (19). Posterior idiosoma not sclerotized ...... 25

— Posterior idiosoma with pygosomal plate; host unknown;

Malaysia & Brasil...... Pygothrombium NEW GENUS

25 (24). Dorsal opisthosoma with conspicuous reticulate pattern

between setal bases; genu legs II and III each with at least 114

two (2) sigma: pretarsus legs I and II each with paired

claws and clawlike empodium...... 26

— Dorsal opisthosoma without reticulate pattern between setal

bases; genu legs II and III each usually with one (1) sigma:

pretarsus legs I and II each with paired claws but may be

lacking clawlike empodium ...... 27

26 (25). Genu leg I with three (3) sigma: host: unknown; Neotropical

...... Manriquia Boshell and Kerr, 1942

— Genu leg I with two (2) sigma: host unknown; Ethiopian and

Oriental...... Reticulatrombidium Shiba, 1976

27 (25). Pretarsus legs I and II each with paired claws but lacking

empodium...... 28

— Pretatsus legs I and II each with paired claws and clawlike

empodium . 30

28 (26). Scutellum with only two setae...... 29

— Scutellum with numerous setae (10+); prodorsal sclerite and

scutellum with longitudinal striations; host unknown;

Palearctic ...... Gonothrombium Feider, 1948

29 (28). Prodorsal sclerite with three rounded posterior projections

with middle projection aligning with an anterior projection

on the scutellum; scutellum with broad groove down middle; 115

prodorsal sclerite and scutellum without prominent

longitudinal striations; host; Diptera; cosmopolitan? . . .

...... Valgothrombium Willmann, 1940

— Prodorsal sclerite without posterior projections; scutellum

triangular in outline; scutum and scutellum with distinct

longitudinal striations; tarsus leg III normal

(symmetrical); hosts: Diptera; Holarctic ......

...... Atractothrombium Feider, 1952

30 (29). Pretarsus leg III with antaxial claw present, usually

modified; scopa and lophotrix well developed ...... 31

— Pretarsus leg III with antaxial claw absent; without scopa

and lophotrix; host unknown; Panama ......

...... Achelothrombidium NEW GENUS

31 (30). Prodorsal sclelrite and/or scutellum with longitudinal

striations; subcapitular setae (scl) palmate distally . . 32

— Prodorsal sclerite and scutellum without longitudinal

striations; subcapitular setae spiniform or setiform. . . 37

32 (31). Prodorsal sclerite with longitudinal striations over entire

surface ...... 33

— Scutum with longitudinal striations limited to the lateral

margins; host; Diptera (primarily Musca domestica);

cosmopolitan ...... Trichotrombidium Koubulei, 1950 116

33 (32). Longitudinal striations limited to prodorsal sclerite;

scutellum punctate ...... 34

— Longitudinal striations present on prodorsal sclerite and

scutellum...... 35

34 (33). Basal sclerites ofthe first pair of dorsal setae posterior

to the scutellum (dl) fused; idiosomal setae in rows E and F

more than half (1/2) length of SL setae; host unknown;

Palearctic ...... Cercothrombium Methlagl, 1928

— Basal sclerites of the first pair of dorsal setae posterior

to the scutellum separate; idiosomal setae in rows E and F

less than half (1/2) length of SL setae; host unknown;

Holarctic ...... Willmannella Feider, 1952

35 (33). Seta la on coxa I branched...... 36

— Seta la on coxa I nude; host unknown; Holarctic ......

...... Campylothrombium Krausse, 1916

36 (35). Famulus on tarsus leg I proximal to omega; lophotrix without

long basal branches ...... Platytrombidium Thor, 1936

— Famulus on tarsus leg I distal to omega; lophotrix with at

least one long basal branch ......

...... Lithothrombidium NEW GENUS 117

37 (32). Famulus tarsus leg I proximal to omega I ...... 38

— Famulus tarsus leg I distal to omega I; genu legs II and III

each with two (2) sigma ......

...... Ettmuelleria Oudemans, 1911

38 (37). Prodorsal eyes unequal in size with anterior being much

larger than the posterior; femur legs I and II with distal

dorsal nude seta (d); host: Diptera; Holarctic......

...... Microtrombidium Haller, 1882

=Megophthrombium Mullen & Vercammen-Grandjean, 1978

— Eyes on prodorsum equal in size or nearly so; femur legs I

and II without distal dorsal nude seta (d); hosts: Diptera

and Coleoptera; cosmopolitan?? ......

...... Diathrombium Moser & Vercammen-Grandjean, 1979

39 (1). Femur leg I, II and III without theta (present in some

Trombiculidae); hosts arthropods or vertebrates ...... 40

— Femur leg II and III with at least one theta; hosts

arthropods...... 48

40 (39). Anal sclerite absent...... 46

— Anal sclerite present ...... 41

41 (40). Coxal formula 2-1-2 (fcx=2,l,2) ......

. . . Family Tanaupodidae...... 42

» 118

— Coxal formula 2—1—1 (fcx=*2,l,l). . Family Chyzeriidae . . 43

42 (41). Anal sclerite with only one (1) pair of setae; host unknown;

Holarctic ...... Subfamily Polydiscinae......

...... Polydiscia Methlagl, 1928

— Anal sclerite with two (2) pairs of setae; host diptera;

Nearctic ...... Subfamily Tanaupodinae......

...... Eothrombium Berlese, 1902

43 (40). Genu leg I, II and III without sigma...... 45

— Genu leg I, II and III with at least one (1) sigma . . . 44

44 (43). Palp femur with two (2) setae; tibia legs I, II and III with

six (6) branched setae; femur leg II with six (6) branched

setae; genu legs II, III with three (3) sigma; host:

Orthoptera; Japan ......

...... Ralphaudyana Vercammen-Grandjean, et. al., 1974

— Palp femur with one (1) seta; tibia legs I and II with eight

(8) branched setae; femur leg II with seven (7) branched

setae; genu legs II, III with three (3) sigma; host:

Orthoptera?; Ethiopian ......

...... Pteridopus Newell and Vercammen-Grandjean, 1964

45 (44) Scutum with median anterior projection (naso); host:

Orthoptera; New Zealand...... 119

...... Nothotrombicula Dumbleton, 1947

— Prodorsal scutum without median anterior projection (naso);

host: Orthoptera; southern hemisphere......

Chyzeria Canestrini, 1897

46 (39). Pretarsus legs I and II with paired claws and clawlike

empodium; most species are vertebrate parasites...... 47

— Paired claws present without clawlike empodium on pretarsus

I and II; host: scorpion; Malaysia ...... Family

Trombellidae (in part) Audyana Woraersley, 1954

47 (46). Genu leg II with three (3) branched setae; scutum with one

(1) anteriomedian seta (AM) (may be absent in some genera);

host: vertebrates; cosmopolitan......

...... Family Trombiculidae

(see Brennan and Goff, 1977 for a key to the New World

genera)

— Genu leg II with four (4) branched setae; scutum with two

(2) anteriomedian setae (AM); host: vertebrates;

cosmopolitan...... Family Leeuwenhoekiidae

(see Brennan and Goff, 1977 for a key to the New World

genera)

48 (39). Femur legs II and III eachwith a single theta or none. . 49

— Femur legs II and III each with at least two (2) theta . 52 120

49 (48). Femur legs II and III with one theta; prodorsal

trichobothria flageliform or globose ...... 50

Femur legs II and III without theta; prodorsal trichobothria

expanded midway along length; host unknown; Holarctic . . .

. Family Johnstonianidae ...... Subfamily

Charadracarinae . Charadracarus Newell, 1960

50 (49). Microseta (k) absent on genu legs I and II ...... 51

Microseta (k) present on genu legs I and II; host: Diptera;

Holarctic...... Johnstoniana George, 1909

51 (50). Prodorsal sclerite with four (4) pairs of setae (including 1

pair of flagelliform trichobothria); femur leg I with two

(2) theta; hosts; aquatic Coleoptera; Holarctic ......

Diplothrombium Berlese, 1909

Prodorsal sclerite with two (2) pairs of setae (including 1

pair of globose trichobothria); femur leg I with one (1)

theta; host: Diptera; holarctic ......

...... Centrotrombidium Kramer, 1896

52 (48). Femur leg I with seven (7) branched setae; coxal field legs

I, II and III with distinct reticulate pattern ......

...... Family Neotrombidiidae .... 53

Femur leg I with six (6) branched setae; coxal field legs I, 121

II and III without reticulate pattern 54

53 (52). Femur legs I to III each with three (3) theta; leg femora

undivided; parasite of streblid flies; Neotropical . . . . .

...... Monunguis Wharton, 1938

— Femur legs I to III each with two (2) theta; femur leg I

divided; host: Coleoptera; cosmopolitan......

...... Neotrombidium Leonardi, 1901

54 (52). Femur legs II and III each with six (6) branched setae; genu

leg I with four (4) sigma; two pairs of prodorsal eyes

present .... Family Trombellidae...... 55

— Femur legs II and III each with five (5) branched setae;

genu leg I with three (3) sigma; one pair of prodorsal eyes

present; host unknown; California. . Family Neotrombidiidae

(in part)...... Taraxithrombium Robaux, 1977

55 (54). Tibia leg II and III each with six (6) branched setae;

microseta k absent from genu legs I and II; leg III with

paired claws but without empodium...... 56

— Tibia leg II and III each with seven (7) branched setae;

microseta k present on genu legs I and II; leg III without

paired claws but with clawlike empodium; host unknown;

cosmopolitan ...... Trombella Berlese, 1888 122

56 (55). Tibia leg I with seven (7) branched setae; femur leg I with

four (4) theta; genu leg I with six (6) sigma: host:

Orthoptera; Maldive Islands ......

...... Womersleya Radford, 1950

— Tibia leg I with sex (6) branched setae; femur leg I with

seven (7) theta; genu leg I with four (4) sigma; host:

Diptera; cosmopolitan ......

...... Durenia Vercammen-Grandjean, 1985 TAXONOMIC SECTION

This section contains diagnoses and descriptions of new taxa included in this study. Diagnoses are provided for all Trorabidioidea studied and are arranged to follow the proposed classification.

Referred speices under each genus represent species actually examined. Complete locality data for all species examined can be found in Appendix B.

123 124

Superfamily Trombidioidea Banks, 1894

Diagnosos (LARVA): fsp= 6-6-6; fst= 0-0-2; fnTr= 1-1-1; fsol=

I(0—2/3—2—1), 11(0-1/2-2-1), 111(0-1/2-0-0); with dorsal and

subterminal eupathidia on tarsus leg I; dorsal eupathid absent on leg

II; legs without companion setae; femur leg III with four (4) setae;

genu legs II and III with two (2) or three (3) setae (occasionally

four); v/ith at least one additional sclerite (scutellum) immedately

posterior to prodorsal sclertite (usually in C-row) bearing at least

one pair of setae.

Family Podothrombiidae Thor, 1935

Podothrombiinae Thor, 1935, p. 137

Ocypetinae Vitzthum, 1941; Thor & Willmann, 1947, p. 515; Baker and

Wharton, 1952, p. 249

Podothrombiodea Feider, 1959, p.454; Feider, 1979, p. 422.

Podothrombiidae Feider, 1959, p. 454; Vercammen-Grandjean, 1973,

p. 112; Feider, 1979, p. 422.

Type Genus: Podothrombium Berlese, 1910

Podothrombium Berlese, 1910

Podothrombium Berlese, 1910, Redia 6:354; Type species Trombidium

I 125

filipes Koch, 1837.

Ocypete Leach, 1814, p. 434; Thor & Willmann, 1947, p. 508 ; Baker

and Wharton, 1952, p. 249

Diagnosis (LARVA): fcx= 2-1-1; fnFe= 5-5-4 or 5-4-4; fnG= 4-3-3

(occasionally 4-4-4); fsol= I(0-2-2-l) 11(0-1-2-1) 111(0-1-0-0)

fk= 1(1-1) 11(1-0) 111(0-0); palpal femur and genu each with a single

seta; scutellum reduced; tarsus leg I many have multiple eupathidia.

For a recent description and figures see Robaux and Schiess, 1982.

(POSTLARVAL): Dorsal idiosomal setae setiform; palpal tibia without

accessory claw; palpal tibia with or without scattered spines;

prodorsal sclerite wide anteriorly.

Referred Species:

Podothrombium montanum Berlese, 1910

P.. shellhammeri Robaux, 1977

_P. piriformis Robaux and Schiess, 1982

P_. verecundum Berlese, 1910

Family Trombidiidae Leach, 1815

Sercothrombidiidae Feider, 1959, p. 546

Type Genus: Trombidium Fabricius, 1775 126

Diagnosis (LARVA): fcx= 2-2-1; fnFe= 5-5-4 or 5-4-4; fnG= 4-3-3

(occasionally 4-4-4); fsol= I(0-2-2-l), 11(0-1-2-1), 111(0-1-0-0)

(except in most Allothrombiinae with I(0-2-2-l), II (0-2-2-1),

111(0-2-0-0)); with two eupathidia on tarsus leg I; fk= 1(1-1)

11(1-0) 111(0-0) (except Parathrombium with 1(1-1) 11(1-0) 111(1-0)); supracoxal setae e and el present; (POSTLARVAL): Dorsal idiosomal setae setiform; palpal tibia without accessory claw or spines; prodorsal sclerite wide anteriorly.

Referred Genera:

Trombidiinae

Trombidium Fabricius, 1775

Parathrombium Bruyant, 1910

Oligothrombidium NEW GENUS

Allothrombiinae

Allothrombium Berlese, 1903

Dinothrombium Oudemans, 1910

Nanothrombidium NEW GENUS

Subfamily Trombidiinae Thor, 1935

Type genus: Trombidium Fabricius, 1775

Diagnosis (LARVA): AM seta nude (except Oligothrombidium); coxal 127

field seta la nude or modified; palpal genu without seta; palpal

femur with one seta, all three setae on palpal tiba nude; paraxial

claw on pretarsus leg III reduced or absent (except

Oligothrombidium); dorsal idiosomal setal row E with two pairs setae.

(POSTALRVAL): Palpal tibia without accessory claw or other spines;

palpal tibia densely covered with setiform setae.

Trombidium Fabricius, 1775

Trombidium Fabricius, 1775, Syst. Ent. p.430; Type species: Acarus holosericeus Linneaus, 1758

Atomus Latreille, 1795, }ag. enc. 4:18; Type species: Acarus parasiticus DeGeer, 1778

Sericothrombium Berlese, 1910, Redia 6:365; Type species: Acarus holosericeus L., 1758

Teresothrombium Feider, 1950, Acad. R.P.R. Filiala Iasi, 1:846-855;

Type speices: Sericothrombium carpaticum Feider, 1948. NEW SYN0N0MY

Diagnosis (LARVA): Coxal field seta la nude; tarsus leg I with one pair of mastisetae; subcapitular setae (scl) with heavy setules.

(POSTLARVAL): Dorsal idiosomal setae setiform with expanded tips; 128

Referred Speices:

Trombidium auroraense Vercammen-Grndjean, Van Driesche & Gyrisco,

1977; X* brevimanum (Berlese, 1910); T. cancelai Robaux. 1967; T. dacicum Feider, 1948; T. holosericeum (L., 1758) (=Atomus parasiticus

DeGeer, 1778); £. heterotrichum (Berlese, 1910); T. hyperi

Vercammen-Grndjean, Van Driesche & Gyrisco, 1977; T. mediterraneum

(Berlese, 1910); T. rhopalicus (Vercammen-Grandjean & Popp, 1967);

T.susteri (Feider, 1956) NEW COMBINATION; T. tenuicualtum (Berlese,

1917); T. teres (Andre, 1929) NEW COMBINATION

Parathrombium Bruyant, 1910

Parathrombium Bruyant, 1910, Zool. Anz. 35:347;Type species: P_. egregium Bruyant, 1910.

Diagnosis (LARVA): Coxal field seta la a modified feather-like structure; subcapitular setae (scl) setiform. For descriptions and figures of larval species, see Newell, 1958 and Robaux, 1969.

(POSTLARVAL): Dorsal idiosomal setae setiform; tarsus all legs without aliform setae. For descriptions of postlarval instars see

Newell, 1958.

Referred Species:

Parathrombium bidactylus Newell, 1958

P. megalochirum (Berlese, 1910) 129

£. quadriseta Newell, 1958

OliRothrombidium NEW GENUS

Type Species: OliRothrombidium mulleni NEW SPECIES

Diagnosis (Larva): Prodorsal trichobothria expanded; seta on palpal femur barbed; pretarsus leg with normally developed claws and empodium.

Holotype from Alabama, Lee Co., Auburn, lO-v-1978, D. Reed, DJR

79-0510-2d, ex moss (UAA). One paratype from Tennessee, Shelby Co.,

Maeeman-Shelby Forest State Park, 7-vi-1976, E. R. Hobeke, BMOC

#76-0804-5, ex hardwood litter.

Description of species. Larva. Idiosoma. Holotype unengorged 443 long, 135 wide; eyes 2/2, with ocular plate, anterior eye larger.

Dorsal idiosomal setae 3—3—2—3—1—1; cupules absent; supracoxal seta el present. One pair of branched intercoxal setae between coxae III; two pair of preanal setae.

Prodorsal Sclerite. Punctate without striations, anterior margin rounded, posterior margin convex; AM > PL > AL; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria expanded with setules.

Prodorsal sclerite measurements of holotype with variation of paratype in parenthesis: AM 49 (48), AA 32 (38), MA 69 (70), AW 61

(79), AL 41 (49), PW 92 (103), PL 45 (44), AP - (41), SB 71 (79), S

47 (51), PSB - (36), ASB 115 (106), SD - (142). Scutellum: HS -

(42), LSS 87 (79), SL 40 (39), SS 34 (-) .

Gnathosoma. Palpal setal formula B-0-NNN2-4NB (palpal trochanter

absent); palp tibial claw two pronged; adoral setae (orl) nude,

subcapitular setae (scl) branched; palpal supracoxal seta (e) not

observed; cheliceral blade with one ventral tooth.

Legs. Femora on legs I to III undivided, five segments beyond the

coxa; pretarsus legs I to III with paired claws and clawike empodium.

Measurements of holotype with those of paratype in parentheses.

Positions of specialized setae of the holotype are given as a ratio

of the segment length variation of paratypes given in parentheses.

IP = 555 (564). Leg I 216 (226); coxa 2B; trochanter IB; femur 5B,

bv seta barbed, d seta barbed, genu 4B, two sigma 23 (23) at 0.23

(0.23) and 0.60 (0.50) respectively, k 4 (4) at 0.82 (0.77); tibia

5B, two £hi 19 (23) and 21 (23) at 0.49 (0.44) and 0.82 (0.85)

respectively, k 5 (3) at 0.80 (0.78); tarsus 17B, omega 26 (21) at

0.67 (0.67), famulus 2 (2) at 0.62 (0.58), two eupathidia 13 (12) and

10 (10) at 0.93 (0.92) and 0.96 (0.97) respectively. Leg II. 160

(157); coxa 2B; trochanter IB; femur 5B, bv seta barbed, d seta

barbed; genu 3B, sigma 21 (22), at 0.41 (0.41); tibia 5B, two phi 16

(17) and 13 (17) at 0.28 (0.31) and 0.79 (0.70) respectively; tarsus

15B, omega 13 (14) at 0.49 (0.47), famulus 2 (2) at 0.49 (0.44). Leg

III. 178 (181); coxa IB; trochanter IB; femur 5B, bv seta barbed, d seta barbed; genu 3B, sigma 19 (21) at 0.42 (0.45); tibia 5B, tarsus 131

14B.

Subfamily Allothrombiinae Thor, 1935

Type Genus: Allothrombium Berlese, 1903

Diagnosis (LARVA): Genu legs I, II and III each with two sigma

(except Nanothrombidium); palpal femur and genu each without setae; coxal field seta la barbed; at least one seta on palpal tibia barbed.

(POSTLARVAL): Dorsal idiosomal setae setiform.

Allothrombium Berlese, 1903

Allothrombium Berlese.1903. Redia 1:351-352; Type species: Trombidium fuliginosum Hermann, 1804.

Diagnosis (LARVA): Subcapitular setae (scl) setiform; the distance between the prodorsal trichobothrial bases less than the distance between the bases of the posterolateral setae; scutellum reduced.

(POSTLARVAL): Tarsus legs I, II, III and IV each with aliform setae subtenting paired pretarsal claws. 132

Referred Species:

Allothrombium brevitarsuum (Berlese, 1888); _A. canaanense Feider,

1977; A^ crassicomum Berlese, 1910; h_. fuliginosum (Hermann, 1804);

_A. lerouxi Moss, 1960; _A. recki Feider, 1967; A., (Aphithrombium) mali

(Childers and Vercammen-Grandjean, 1980) NEW COMBINATION.

Dinothrombium Ouderaans, 1910

Dinothrombium Oudemans, 1910, Ent. Ber. Nederl. Ver. 3:48; Type

species: Acarus tinctorius Linneaus, 1767

Isothrombium Andre, 1949, Bull. Mus. Nat. Hist. Nat. Paris, 2e Serie

21:354-357; Type species: Isothrombium oparbellae Andre, 1949. NEW

SYN0N0MY

Angelothrombium Newell & Tevis, 1960, Ann. Ent. Soc. America 53:

293-304; Type species: Angelothrombum pandorae Newell & Tevis, 1960.

Diagnosis (LARVA): Subcapitular setae (scl) short and thick; the distance between the prodorsal trichobothrial bases more than the distance between the bases of the posterolateral setae; scutellum not reduced.

(POSTLARVAL): Tarsus legs I, II, III and IV each without aliform setae subtending paired pretarsal claws; idiosomal setae setiform; 133

Referred Species:

D. tinctorum (L., 1767); I), magnificum (Le Conte, 1852) (=T. superbum

Banks, 1910) NEW SYN0N0MY; D. oparbellae Andre, 1949, NEW

COMBINATION; D. pandorae (Newell & Tevis, 1960).

Nanothrombidium NEW GENUS

Type Species: Nanothrombidium nannelli NEW SPECIES

Diagnosis (LARVA): Coxal field seta la nude; genu legs II and III with one sigma; pretarsus leg III unmodified;

Postlarval instars unknown.

Nanothrombidium nannelli NEW SPECIES

Type Data: Holotype Italy 16 km W Rome ex sweepings of annual vegetation along Via Casia. Paratype ex thrips (Thysanoptera) from same locality. 13 June 1984; W.C. Welbourn and S.C. Clement colls.; original number WCW840613-5.

Description of species. Larva. Idiosoma. Holotype partially engorged

289 long, 135 wide; eyes 2/2 subequal, ocular plate (23 x 12).

Dorsal idiosomal setal rows 3—3—2—3—1—1; cupules absent; supracoxal seta Ie present. One pair of branched intercoxal setae between coxae

III; four preanal setae; 134

Prodorsal Sclerite. Lightly punctate without striations, anterior margin extended into broad naso, slightly narrower and with

horizontal striations at level of AM setae, posterior margin broadly rounded; PL > AM > AL; SB < PW; trichobothridial bases anterior to

PL setal bases; trichobothria flagellate with setules.

Prodorsal Sclerite measurements of holotype with variations of paratype in parenthesis: AM 28 (25), AA 29, MA 30 (28), AW 50, AL 23

(23), PW 57 (56), PL 36, AP 25 (28), S 44 (42), PSB 33 (36), ASB 72

(70), SD 105 (106). Scutellum: HS 37 (40), LSS 67; SL 37 (29); SS 22

(18).

Gnathosoma. Palpal setal formula 0-0-NNB2-wB3N (palpal trochanter absent); palp tibial claw 2 pronged; adoral setae (orl) setae nude

(4), subcapitular setae barbed (27); palpal supracoxal seta (e) 6; cheliceral blade small with one ventral tooth.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawike empodium.

Measurements of holotype with variation of paratype in parentheses.

Positions of specialized setae of the holotype are given as a ratio of the segment length with variation of paratype are given in parentheses. Seta bv on femur all legs long with 2-4 barbs. IP =448

(438). Leg I 157 (151); coxa NB; trochanter IB; femur 5B; genu 4B, two sigma 17 (15) at 0.36 and 0.57 respectively, k 2 at 0.83; tibia

5B, 2 phi 22 and 12 (11) at 0.86 (0.89) and 0.88 (0.91) respectively, k 2 at 0.86; tarsus 17B, omega 13 at 0.53), famulus 1 at 0.50, two eupthidia 20 (19) and 11 at 0.68 and subterminally respectively. Leg

II. 138 (137); coxa 2B; trochanter IB; femur 5B; genu 3B, sigma 14 135 at 0.47, k 3 at 0.76; tibia 5B, phi 13 and 12 (10) at 0.84 (0.80) and

0.85 (0.90) respectively; tarsus 14B, omega 14 at 0.44, famulus 1 at

0.44. Leg III. 153 (150); coxa IB; trochanter IB; femur 5B; genu 3B, sigma 14 at 0.55; tibia 4B; tarsus 13B.

Family Neothrombiidae Feider, 1959

Type Genus: Neothrombium Oudemans, 1909 (larva)

Family Diagnosis (larva): fsp= 6-6-6; fnTr= 1-1-1; fnFe= 6-4-4; fnG=

4-2-2 (except Acridothrombium with 4-4-4); fnTi= 6-5-5 (except

Megalothrombium with 6-6-6 and Aegialothrombidium with 7-5-5); fk=

1(1-1) 11(0-0) 111(0-0); fZ= 2-0-0; famulus I distal to omega I; pretarsal claws claw like; empodium with distal end expanded; scl setiform; supracoxal seta Ie present.

Referred subfamilies and genera:

Neothrombiinae Feider, 1959

Neothrombium Oudemans, 1909

Ceuthothrombium Robaux, Webb & Campbell, 1976

Acridothrombium NEW GENUS

Kermathrombidium NEW GENUS

Adelothrombidium NEW GENUS

Aethethrombidiinae NEW SUBFAMILY

Aethethrombidium NEW GENUSA 136

Megalothrombium NEW GENUS

Aegialothrombidiinae NEW SUBFAMILY

Aegialothrombidium NEW GENUS

Subfamily Neothrorabiinae Feider, 1959

Type Genus: Neothrombium Oudemans, 1909

=sMyceterothrombidiinae Feider, 1948 p. 92

Subfamily Diagnosis (LARVA): Prodorsal sclerite rectangular usually with the prodorsal trichobothria placed posterior to PL setae; dorsal

eupathid on tarsus I set on pedicile.

(POSTLARVAL): Prodorsal sclerite wide, wider than crista metopica, in

some genera the eyes are incorporated into the prodorsal sclerite, idiosoraal setae armed and short, palpal tibia with scattered spines, not arranged in rows; tarsus leg I not expanded.

Neothrombium Oudemans, 1909

Neothrombium Oudemans 1909, Ent. Ber. Nederl. Ver. 3:17-20;Type speices Allothrombium neglectum Bruyant, 1909 (larva).

Myceterothrombium Feider, 1945, Bull. R. Acad. Sect. Scienti.

27:528-531; Type speices: Myceterothrombium neglectum (postlarval 137

instars).

Diagnosis (LARVA): fsol- I(0-2-2-l) 11(0-1-2-1) 111(0-1-0-0); fPp=

0-0-NNN2- ANB; AM, AL, and PL barbed; eyes 2/2 on platelets; prodorsal sclerite and scutellum punctate and with striations; scutellura with 6 setae; palpal femur without seta; legs with longitudinal.striations; bv seta on legs II and III short and nude.

For a complete redescription and figures see Robaux, 1973.

(POSTLARVAL): For a description of the adult see Willmann, 1950 and

Feider, 1955.

Hosts: Gryllotalpa gryllotalpa and £. unispina (Orthoptera,

Gryllotalpidae)

Distribution: Europe, North Africa and Middle East.

Referred Species:

Neothrombium neglectum

Ceuthothrombium Robaux, Webb & Campbell, 1976

Ceuthothrombium Robaux, Webb & Campbell, 1976, Ann. Speleol.

31:213-218; Type species: Ceuthothrombium cavaticum Robaux, Webb &

Campbell, 1976 (No holotype or paratypes designated in the description). 138

Diagnosis (LARVA): fsol=-I(0-2-2-l) 11(0-1-2-1) 111(0-1-0-0); fPp=0-N-BNN2-4NB; AM, AL and PL barbed; eyes 1/1 (occasionally absent), without ocular plates; prodorsal sclerite and scutellum punctate without striations; scutellum narrow with 2 setae, prodorsal trichobothria with few distal setules, bv setae on all legs barbed, legs without striations.

(POSTLARVAL): Prodorsal sclerite about as wide as crista metopica; idiosomal setae with 3 to 6 pair of large, alternate barbs; eyes absent; accessory spine on palpal tibia as long as palpal tibial claw; palpal tarsus with 3-4 terminal euapthidia and one omega.

Host: Ceuthophlus carlsbadensis Caudell, C_. conicaudus Hubbell, C. longipes Caudell, C. paucispinosus Rehm, £. pima Hubbell, C. spp.

(Orthoptera, Rhaphidophoridae)

Distribution: southwestern USA and Mexico

Referred Speices:

C. cavaticum Robuax, Webb, & Campbell, 1976.

Acridothrombium NEW GENUS

Type Species: Acridothrombium chiricahuae NEW SPECIES 139

Generic Diagnosis (LARVA): fnG*= 4-4-4; tarsus legs I, II and III neotrochous (25-22-20); fPp= N-0-NNN2-6N or N-0-BBB2-5NB; AM, AL and

PL barbed; eyes 2/2 on striated ocular plates; prodorsal sclerite and scutellum punctate with striations; scutellum with 8-13 setae, with one pair longer than the rest; legs with faint striations.

(POSTLARVAL): Prodorsal sclerite wider than crista metopica, widest in middle with eyes incorporated into the prodorsal sclerite, idiosomal setae with three large spines basally and setiform distally; palpal tibia with 6 to 10 scattered spines, palpal tarsus with 4-5 terminal eupathidia.

Acridothrombium chiricahuae NEW SPECIES

Holotype and 12 paratypes reared from eggs deposited by female collected under a rock 2.5 km ESE Portal, Arizona, Cochise Co.,

3-VII-1981; W.C.Welbourn, culture 247; 11 paratypes reared from eggs deposited by 5 other females collected under rocks at same locality

(cultures 248, 254, 255 and 257); 6 paratypes removed from neck and coxae of Xanthippus c. pantherinus (WCW810803-15); 23 paratypes from five specimens of Trimerotropis £. pallidipennis; 5 paratypes from two juvenile Brachystola magna; 18 paratypes from residue in vials with above mentioned hosts.

Description of species. Larva. Idiosoma. Holotype unengorged 274 long, 157 wide, 19 unengorged paratpyes averaged 257 (233-288) long by 175 (140-212) wide; eyes 2/2, with ocular plate (33 x 21), 140 anterior larger (13), posterior eye smaller (11). Dorsum hypertrichous obscuring setal rows; cupules absent; supracoxal seta el present. One pair of branched intercoxal setae between coxae III; twenty preanal setae.

Prodorsal sclerite. Punctate with longitudinal striations, shape roughly rectangular with anterior margin slightly concave, posterior margin straight; AM > AL > PL; SB < PW; trichobothridaial bases slightly posterior to PL setal bases; trichobothria flagellate with distal setules.

Scutal measurements of holotype with mean, range and number of paratypes in parenthesis; AM 35 (35, 30-40,45), AA 36 (39, 32-46,

47), MA 30 (39, 32-46, 47), AW 48 (54, 48-59, 47), AL 29 (26, 23-30,

43), PW 61 (61, 57-66, 47), PL 22 (21, 18-25,47), AP 28 (28, 24-34,

47), SB 32 (34, 31-38, 47), S - (68, 59-76, 47), PSB 19 (19, 16-24,

47), ASB 65 (66, 62-74, 46), SD 84 (86, 79-93, 46). Scutellum; HS 45

(46, 36-53, 47), LSS 127 (127, 118-136, 47), with 8 to 12 setae

(14-20) and one pair of SL setae 25 (25, 20-33, 46), SS 24 (26,

19-34, 44).

Gnathosoma. Palpal setal formula N-0-NNN2-w6N (palpal trochanter absent); palp tibial claw with two pronged; orl setae branched, scl setae branched; palpal supracoxal seta absent; cheliceral blade with one ventral tooth; gnathobase punctae with striations.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawlike empodium; empodium with expanded tip. Measurements are of the holotype with mean, range and number of paratypes in parenthesis. 141

Positions of specialized setae of the holotype are given as a ratio

of the segent length mean, range and number of paratypes in

parenthesis. The bv setae on femur all legs with 1-3 small barbs.

IP 750 (755, 725-784, 38). Leg I 253 (254, 240-268, 39); coxa 2B;

trochanter IB; femur 6B; genu 4B, two sigma 25 (25, 22-29, 46) and 24

(25, 22-26, 46) at 0.29 (0.30, 0.20-0.40, 46) and 0.60 (0.60,

0.49-0.72, 47) respectively, k - (3, 3-4, 23) at - (0.70, 0.64-0.77,

33); tibia 6B, two £hi 16 (17, 13-20, 47) and 13 (14, 13-17, 47) at

0.33 (0.36, 0.26-0.42, 47) and 0.77 (0.81, 0.75-0.86, 46) respectively, k - (3, 2-4, 16) at - (0.68, 0.61-0.79, 23); tarsus 25B

(range in paratypes 22 27), omega 16 (16, 13-18, 45) at 0.42 (0.42,

0.36-0.47, 47), famulus 2 (3, 2-4, 37) at 0.60 (0.61, 0.55-0.69, 44), two eupthidia 18 (18, 14-21, 44) and 10 (10, 8-11, 43) at 0.83 (0.83,

0.80-0.87, 46) and 0.92 (0.93, 0.91-0.97, 47) respectively. Leg II.

226 (231, 220-245, 40); coxa 6-7B; trochanter IB; femur 4B; genu 4B, sigma 27 (25, 21-28, 46), at 0.45 (0.45, 0.37-0.53,47); tibia 5B, two phi 16 (15, 11-18, 46) and 12 (13, 11-15, 47) at 0.39 (0.38,

0.32-0.46, 47) and 0.77 (0.77, 0.72-0.84, 46) respectively; tarsus

22B (range in paratypes 19-25), omega 17 (17, 15-18, 47) at 0.32

(0.35, 0.31-0.38, 46), famulus 2 (2, 2-3, 23) at 0.33 (0.35,

0.29-0.41, 28). Leg III. 271 (272, 253-288, 45); coxa 1-2B; trochanter IB; femur 4B; genu 4B, sigma 35 (34, 31-38, 46) at 0.37

(0.43, 0.32-0.54, 46); tibia 5B, tarsus 20B (range in paratypes

18-22). 142

Kermathrombidium NEW GENUS

Type Species: Kermathrombidium panamensis NEW SPEICES

Diagnosis (LARVA): fsol« I(0-2-l-l) 11(0-1-2-1) 111(0-1-0-0); fPp=

0-0-NBB2-3B3N; AM bifurcate; AM, AL and PL barbed; eyes absent; prodorsal trichobothria expanded; prodorsal sclerite and scutellum punctate without striations; scutellum with 2 setae. Postlarval stages unknown.

Kermathrombidium panamensis NEW SPECIES

Holotype and 7 paratypes ex corniger (Termitidae), Panama, C.Z.,

Frijoles, 19-V-1980, B. Thorne coll.

Description of species. Larva. Idiosoma. Holotype engorged; due to the condition of the idiosoma in all specimens it was not possible to make measurements or determine setal rows; eyes absent; cupules absent; supracoxal seta el present; one pair of branched intercoxal setae between coxae III.

Prodorsal Sclerite. Punctate without longitudinal striations, anterior margin concave, posterior margin convex; PL > AL > AM; SB <

PW; trichobothridial bases posterior to PL setal bases; trichobothria expanded with setules. Scutellum rectangular with one pair of setae (SL) near anterior margin.

Prodorsal sclerite measurements of holotype with mean, range and 143

number of paratypes in parenthesis: AM 16 (16, 15-18, 7), AA 18 (20,

18-21, 7), MA 10 (11, 10-11, 7), AW 28 (29, 28-30, 7), AL 23 (21,

18-23, 7), PW 33 (33, 32-34, 7), PL 26 (23, 21-25, 7), AP 12 (12,

11-13, 7), SB 26 (25, 22-26, 7), S 27 (26, 25-28, 5), PSB 15 (16,

14-18, 7), ASB 31 (29, 28-30, 7), SD 46 (45, 44-47, 7). Scutellum:

HS 22 (22, 20-23, 6), LSS 37 (39, 36-42, 6), SL 32 (29, 26-31, 5), SS

11 (11, 10-12, 6).

Gnathosoma. Palpal setal formula 0-0-BBN2-3B2N (palpal trochanter

absent); palp tibial claw two pronged; orl setae nude, scl setae

branched; palpal supracoxal seta absent; cheliceral blade not

observed.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawike empodium; emposium with expanded tip. Measurements are of the holotype with mean, range and number of paratypes in parenthesis. Positions of specialized setae of the holotype are given as a ratio of the segment length with mean, range and number of paratypes in parenthesis. Seta bv on the femur legs I, II and III long with 1-3 barbs. IP= 299

(301, 290-310). Leg I 102 (104, 98-108, 7); coxa 2B; trochanter IB; femur 6B; genu 4B, two sigma 11 (11, 9-12, 7) and 10 (10, 9-11, 7) at

0.27 (0.24, 0.20-0.23, 7) and 0.30 (0.31, 0.29-0.39, 7) respectively, k 5 (4, 3-5, 7) at 0.80 (0.79, 0.70-0.87, 7); tibia 6B, two phi 12

(11, 9-12, 7) and 13 (13, 12-14, 7) at 0.21 (0.25, 0.16-0.38, 7) and

0.68 (0.68, 0.63-0.67, 7) respectively; tarsus 18B, omega 16 (15,

14-16, 7) at 0.36 (0.36, 0.31-0.38, 7), famulus - (2, 1-2, 5) at 0.56

(0.55, 0.50-0.61, 7), two eupthidia 11 (11, 8-12, 6) and 7 (8, 6-9, 144

7) at 0.82 (0.80, 0.75-0.82, 6) and 0.84 (0.83, 0.79-0.87, 5) respectively. Leg II 90 (94, 87-101, 7); coxa 2B; trochanter IB; femur 4B; genu 2B, sigma 10 (10, 8-11, 7) at 0.48 (0.40, 0.35-0.46,

7); tibia 5B, two £hi 9 (11, 11-12, 7) and 10 (11, 9-12, 7) at 0.21

(0.25, 0.19-0.36, 7) and 0.60 (0.62, 0.54-0.76, 7) respectively; tarsus 13B, omega 15 (14, 13-15, 7) at 0.31 (0.33, 0.31-0.34, 7), famulus 2 (2, 2-3, 5) at 0.32 (0.34, 0.31-0.37, 6). Leg III 106

(102, 88-106, 6); coxa IB; trochnter IB; femur 4B; genu 2B, sigma 12

(11, 10-12, 6) at 0.46 (0.46, 0.44-0.54, 7); tibia 5B, tarsus 11B.

Adelothrombidium NEW GENUS

Type Species: Adelothrombidium africana NEW SPEICES

Diagnosis (LARVA): Scutellum with two setae; prodorsal sclerite and scutellum punctate and without striations; prodorsal trichobothria anterior to PL setae. Postlarval stages unknown.

Adelothrombidium africana NEW SPECIES

Holotype and 10 paratypes ex Libanasidus vittatus (Orthoptera,

Gryllacridiae, Heniciinae), South Africa, Transvaal, Malta Forest,

Trichardtsdaal, 5-iii-1984, Otte, Cale & Tom #145, BM0C #84-0628-2.

Description of species. Larva. Idiosoma. Holotype engorged and not 145

measured; eyes 2/2, with ocular plate (30 x 15), anterior eye larger

(11), posterior eye 8. Dorsal chaetotaxy. Dorsal idiosoma

hypertrichous; cupules absent; supracoxal seta el present. Ventral

Chaetotaxy. One pair of branched intercoxal setae between coxae III;

four preanal setae; with a total of fourteen ventral setae posterior

to coxal field leg III.

Prodorsal Sclerite. Punctate without striations, anterior and

posterior margins convex; AL > AM > PL; SB < PW; trichobothridial

bases anterior to PL setal bases; trichobothria flagellate with few

distal setules.

Prodorsal sclerite measurements of holotype with mean, range and

number of paratypes in parentheses: AM 39 (37, 32-41, 9), AA - (53,

50-58, 5), MA 28 (31, 28-33, 3), AW 59 (59, 55-61, 10), AL 40 (42,

40-45, 8), PW 84 (83, 80-87, 10), PL 23 (22, 20-23, 10), AP 54 (55,

51-58, 10), SB 59 (61, 58-64, 10), S 57 (58, 51-63, 6), PSB 20 (22,

20-27, 5), ASB - (105, 1), SD - (130, 1). Scutellum: HS 32 (31,

28-35, 10), LSS 87 (85, 81-90, 10), SL 32 (30, 28-32, 7), SS 34 (35,

32-39, 10).

Gnathosoma. Palpal setal formula N-0-BBN2-4NB (palpal trochanter

absent); palp tibial claw two pronged; orl setae nude, scl setae

branched; palpal supracoxal seta absent; cheliceral blade with two

teeth.

Legs. Femora on legs I to III undivided, five segments beyond the

coxa; pretarsus legs I to III with paired claws and clawlike

empodium; distal tip of empodium expanded. Measurements of holotype with mean, range and number of paratypes in parenthesis. Positions of specialized setae of the holotype are given as a ratio of the segment length with mean, range and number of paratypes are given in parentheses. Seta bv on femur legs I and II long and barbed, bv on femur leg III short and without barbs. IP = 587 (573, 564-583, 9).

Leg I 204 (202, 196-210, 9); coxa 2B; trochanter IB; femur 6B, bv seta branched; genu 4B, two sigma 17 (17, 15-18, 9) and 18 (18,

16-19, 8) at 0.26 (0.23, 0.19-0.26, 10) and 0.44 (0.42, 0.37-0.51,

10) respectively, k 3 (3, 3-4, 9) at 0.81 (0.78, 0.72-0.84, 9); tibia

6B, two £hi 17 (16, 15-18, 9) and 17 (17, 16-19, 10) at 0.35 (0.32,

0.28-0.38, 10) and 0.77 (0.77, 0.70-0.81, 10) respectively, k 4 (3,

2-4, 10) at 0.73 (0.71, 0.67-0.75, 6); tarsus 18B, omega 23 (21,

17-23, 10) at 0.37 (0.39, 0.37-0.41, 10), famulus 2 (3, 2-4, 10) at

0.50 (0.50, 0.48-0.53, 10), two eupthidia 24 (22, 20-24, 9) and 13

(12, 10-13, 6) at 0.84 (0.84, 0.83-0.85, 9) and 0.92 (0.91,

0.88-0.95, 8) respectively. Leg II. 174 (171, 161-175, 10); coxa

2B; trochanter IB; femur 4B, bv seta branched; genu 2B, sigma 25 (25,

22-29, 10), at 0.32(0.29, 0.25-0.38, 10); tibia 5B, two £hi 15 (14,

12-16, 10) and 15 (16, 14-17, 10) at 0.36 (0.37, 0.35-0.40, 10) and

0.69 (0.74, 0.68-0.74,10) respectively; tarsus 14B, omega 17 (17,

15-19, 10) at 0.34 (0.34, 0.33-0.36, 10), famulus 1 (2, 1-3, 6) at

0.30 (0.30, 0.27-0.31, 9). Leg III. 200 (199, 195-206, 9); coxa IB; trochanter IB; femur 4B, bv seta branched; genu 2B, sigma 19 (20,

19-22, 9) at 0.50 (0.44, 0.33-0.53, 9); tibia 5B, tarsus 13B. 147

Aethethrombidiinae NEW SUBFAMILY

Type species: Aethethrombidium braziliensis NEW SPECIES

Diagnosis (LARVA): Coxae on all legs multisetose; palp femur and genu fused on antaxial surface; palp claw with single prong; PL posterior to prodorsal trichobothria; scutellum with two setae; cheliceral blade elongate. Postlarval stages unknown.

Aethethrombidium NEW GENUS

Type Species: Aethethrombidium brazilensis NEW SPECIES

Generic Diagnosis (LARVAL): fcx= 5/6B-6B-3/5B; fst= 0-0-0; fsol=

I(0-2-2-l) 11(0-1-2-1) 111(0-1-0-0); fPp= N-0-NNS1-4NB; one seta on palp tibia spiniforra; AM and PL Barbed; AL nude. Postlarval stages unknown.

Aethethrombidium brazilensis NEW SPECIES

Holotype: Brasil, Amazonas, Manaus; ex miscellaneous litter at INPA;

31-111-1978; D. E. Johnston; original number 31-111-78; Acarology

Laboratory Accession number AL-1070.

Description of species. Larva. Idiosoma. Holotype appears to be partially engorged 426 long, 280 wide; eyes 2/2, with ocular plate 148

(42 x 27), anterior eye larger (17), posterior eye 9. Due to the condition of the holotype it was not possible to determine dorsal idiosomal chaetotaxy; cupules absent; supracoxal seta el present.

One pair of intercoxal setae between coxal field III; six pairs of preanal setae.

Prodorsal Sclerite. The prodorsal sclerite is roughly circular with anterior margin extending over prodorsum; punctate with longitudinal striations, anteriomargin rounded, posterior margin nearly straight with rounded corners; AM > PL = AL; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules.

Prodorsal sclerite measurements of holotype: AM 59, AA 53, MA 64, AW

127, AL 41, PW 129, PL 41, AP 61, SB 132, S 103, PSB 43, ASB 106, SD

148. Scutellum: HS 53, LSS 141, SL 63, SS 35.

Gnathosoma. Palpal setal formula N-0-NBS1-2B3N; two setae on palpal tarsus very long, one nude and other branched; palpal trochanter absent and femur genu fused on external surface; palp tibial claw with single prong; adoral setae (orl) nude, subcapitular setae (scl) branched (83); palpal supracoxal seta absent; cheliceral blade partially obscured, but appeared to be elongate.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawlike empodium. Measurements of holotype with positions of specialized setae given as a ratio of the segment length. Setae bv on femur all legs barbed. IP = 986. Leg I 319; coxa 5-6B; trochanter IB; femur

6B; genu 4B, two sigma 30 and 29 at 0.17 and 0.57 respectively, k 6 at 0.59; tibia 6B, two phi 22 and 30 at 0.83 and 0.91 respectively, k 149

7 at 0.75; tarsus 18B, omega 22 at 0.12, famulus 7 at 0.35, two

eupthidia 39 and 13 at 0.74 and 0.98 respectively. Leg II. 306; coxa

6B; trochanter IB; femur 4B; genu 2B, sigma 40 at 0.31; tibia 5B, phi

34 and 17 at 0.25 and 0.78 respectively; tarsus 13B, omega 17 at

0.27, famulus 4 at 0.23. Leg III. 360; coxa 3-5B; trochanter IB;

femur 4B; genu 2B, sigma 41 at 0.29; tibia 5B, tarsus 12B.

Megalothrombium NEW GENUS

Type Species: Megalothrombium jamaicensis NEW SPECIES

Generic Diagnosisis (LARVA): cx= 10B-13B-15B; fst= 0-0-4; fnTi=

6-6-6; fnTa= 50+-50+-50+; fsol= I(0-7-2-l) 11(0-22-2-1)

111(0-24-0-0); fPp= N-0-NNNl-4NBw; AM and AL nude; PL barbed; scutum and scutellum striate and punctate; idiosoma hypertrichous.

Postlarval stages unknown.

Megalothrombium jamaicensis NEW SPEICES

Holotype from Jamaica; St. Thomas Par., Corn Pass Gap; ll-V-1950; H.

B. Mills coll.; 111. Nat. Hist. Survey Acc. No. 49627, KU collection.

Description of species. Larva. Idiosoma. Holotype unengorged 589 long, 366 wide; eyes 2/2, with ocular plate (40 x 22), anterior eye

26, posterior eye 14. Dorsal idiosoma hypertrichous; cupules absent; 150 supracoxal seta el present. Five pairs of branched intercoxalsetae between coxae III; more than fifty preanal setae.

Prodorsal Sclerite. Punctate with longitudinal striations, anterior margin rounded (convex), posterior margin slightly convex with rounded corners; AL > AM > PL; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules.

Prodorsal Sclerite measurements of holotype: AM 70, AA 67, MA 144,

AW 150, AL 86, PL 65, AP 67, SB 111, S 144, PSB 74, ASB 208, SD 282.

Scutellum: HS 74, LSS 192, SL 96, SS 31.

Gnathosoma. Palpal setal formula N-0-NNN1-3NB; plapal trochanter absent and femur-genu fused on external side; palp tibial claw entire; adoral setae (orl) nude (26), subcapitular setae (scl) branched (69); palpal supracoxal setae absent; cheliceral blade elongate terminating in a sharp point.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawlike empodium with expanded tip. Measurements of holotype with positions of specialized setae of the holotype given as a ratio of the segment length. Seta bv not differentiated from other femoral setae on all legs. IP = 2207. Leg I 721; coxa 9—1IB; trochanter IB; femur 6B; genu 4B, seven sigma 32-36 along entire dorsal surface, k 10 at 0.72; tibia 6B, two phi 40 at 0.70 and 0.90 respectively, k 11 at 0.90; tarsus 100+B, omega 62 at 0.66, famulus 10 at 0.75, two eupthidia 39 at 0.88 and 0.94. Leg II. 662; coxa 12-13B; trochanter IB; femur

4B; genu 2B, twenty-two dorsal sigma 39-46; tibia 6B, two phi 36 and

34 at 0.37 and 0.80; tarsus 50+B, omega 33 at 0.54, famulus 5 at 151

0.40. Leg III. 809; coxa 13-14B; trochanter IB; femur 4B; genu 2B,

twenty-four dorsal sigma 39-43; tibia 6B, tarsus 50+B.

Aegialothrombidiinae NEW SUBFAMILY

Type Species: Aegialothrombidium roexicana NEW SPECIES

Generic Diagnosis: fnTi= 7-5-5; fsol= I(1-2-1) 11(1-1-1) 111(1-0-0);

fPp= 0-0-NNN2-4BN or 0-0-NNN2-5N; AM nude, AL and PL barbed; eyes 2/2

without ocular plate; PL seta between eyes; prodorsal sclerite and

scutellum with striations and punctaions; prodorsal trichobothria

distally bifurcate; genu leg III without sigma; tibia leg II with one

phi; coxal field leg II with one seta. Postlarval instars unknown

Aegialothrombidium NEW GENUS

Type Species: Aegialothrombidium mexicana NEW SPECIES

Aegialothrombidium mexiciana NEW SPECIES

Holotype and two paratypes from under elytra of £ pilifera le Conte,

Mexico, Baja California Sur, Puerticetos, L. E. Watrous.

Description of species (Larva). Idiosoma. Holotype unengorged 220 long, 99 wide; eyes 2/2 without ocular plate, anterior eye smaller 152

(7), posterior eye larger (8). Dorsal setal rows 3-3-3-3-1-1; cupules absent; supracoxal seta el (10) under coxal field leg I.

Thirty branched setae posterior to coxal field leg III; one pair of branched intercoxal setae between coxal fields leg III.

Prodorsal Sclerite. Punctate with longitudinal striations, anterior margin slightly convex with anterolateral edge of scutum extending over the prodorsum, posterior margin straight with sharp corners; AM

> AL > PL; SB < PW; PL setae located between eyes; trichobothria flagellate, forked distally, AM setae nude.

Prodorsal Sclerite measurements of holotype with mean of two paratypes in parentheses: AM 14 (15), AA 21 (21), MA 45 (47), AW 49

(48), AL 25 (24), PL 24 (24), SB 44 (45), S 32 (34), PSB 17 (15), ASB

67 (72), SD 83 (87). Scutellum: HS 39 (42), LSS 53 (54), anterior SL

26 (27), posterior SL 27 (26), anterior SS 44 (41), posterior SS 32

(33).

Gnathosoma. Palpal setal formula 0-N-NNS2-5N (palpal trochanter absent); palp tibial claw two pronged; adoral (orl) setae nude (7), subcapitular (scl) setae branched (15); cheliceral blade very small with one ventral tooth, gnatobase punctate with striations.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III with paired claws and clawike empodium with distally expadned tip. Measurements of holotype with mean of two paratypes in parentheses. Positions of specialized setae of the holotype are given as a ratio of the segment length with mean of two paratypes in parentheses. IP <= 389 (416). Leg I 143 (154); coxa 2B; trochanter IB; femur 6B, bv seta branched; genu 4B, one sigma 16 (18) at 0.38 (0.33), k 5 (5) at 0.79 (0.70); tibia 7B, two phi 9 (14) and

11 (11) at 0.88 (0.78) and 0.88 (0.82) respectively, k 4 (4) at 0.85

(0.78); tarsus 18B, omega 15 (16) at 0.53 (0.55), famulus - (2) at

0.66 (0.63), two eupthidia 17 (16) and 8 (10) at 0.60 (0.64) and0.89

(0.86) respectively. Leg II 117 (121); coxa IB; trochanter IB; femur 4B, bv seta branched; genu 2B, sigma 32 (35), at 0.22 (0.26); tibia 5B, one phi 14 (14) at 0.74 (0.71); tarsus 15B, omega 15 (15) at 0.37 (0.43), famulus 2 (3) at 0.35 (0.41). Leg III 130 (140); coxa IB; trochanter IB; femur 4B, bv seta branched; genu 2B; tibia

5B, tarsus 13B.

Family CAL0THR0MBIIDAE Oudemans, 1940

Type Genus: Calothrombium Berlese, 1918

Diagnosis (LARVA): Dorsal idiosoma with two scutella at rows C and D each with one pair of setae (cl and dl respectively); scopa bifurcate; lophotrix pilose, without any long branches or barbs.

(POSTLARVAL): Prodorsal sclerite with lateral scleritization, widest at prodorsal trichobothria; without eyes; idiosomal setae bifurcate and barbed; palp tibia with two parallel rows of 8-11 spines; with three larve ventral spines on palpal tibia. 154

Referred Genera:

Calothrombium Berlese, 1918 (adult)

Dectothrombidium NEW GENUS (larva)

Dectothrombidium NEW GENUS

Type Species Dectothrombidium marshalli NEW SPECIES

Diagnosis (LARVA): Dorsal idiosoma with two scutella at rows C and D

each with one pair of setae (cl and dl respectively); scopa

bifurcate; lophotrix pilose, without any long branches or barbs.

Postlarval instars unknown.

Referred Species:

D. marshalli NEW SPECIES

JD. danium Paoli, 1937 (not examined) NEW COMBINATION

Dectothrombidium marshalli NEW SPECIES

Holotype ex Decticus albifrons F. (Orthoptera, Tettigoniidae), in the

British Museum (Natural History) collectons; specimen label with

"Spaino 1876"

Description of species. Larva. Idiosoma. Holotype engorged (not measured); eyes 2/2, with ocular plates (41 x 15), anterior eye 19, posterior eye 16. Dorsal idiosomal setae 3—3—3—2—1—1; cupules 155 absent; supracoxal seta el present. One pair of branched intercoxal setae between coxae III; nine preanal setae.

Prodorsal Sclerite. Punctate with striations, anterior margin convex with a rounded point in the middle, posterior margin undulating; AM >

AL = PL; SB < PW; AM barbed; trichobothridial bases anterior to PL setal bases; trichobothria flagellate with without setules.

Prodorsal sclerite measurements of holotype: AM 41, AA 69, MA 62, AW

102, AL 31, PW 97, PL 32, AP 53, SB 82, S 83, PSB 43, ASB 96, SD 140.

Scutellum: HS 43, LSS 98, SL 36, SS 59.

Gnathosoma. Palpal setal formula N-N-BNS2-2B4N (plapal trochanter absent); palp tibial claw two pronged; adoral setae (orl) nude, subcapitular setae (scl) spiniform; palpal supracoxal seta (e) present; cheliceral blade broken; buccal ring present.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to III withpaired claws and clawike empodium.

Measurements of holotype. Positions of specialized setae of the holotype are given as a ratio of the segment length. IP = 758. Leg

I 273; coxa 3B; trochanter IB; femur 6B, bv seta barbed, d seta barbed; genu 4B, two sigma 17 and 18 between 0.29 and 0.31, k 2 at

0.82; tibia 6B, two phi 18 and 19 at 0.23 and 0.90 respectively, k 7 at 0.88; tarsus 18B, omega 20 at 0.44, famulus 3 at 0.52, two eupthidia 30 and 19 at 0.75 and 0.87 respectively. Leg II. 240; coxa 2B; trochanter IB; femur 5B, bv seta barbed, d seta barbed; genu

2B, sigma 22 at 0.23, k 3 at 0.77; tibia 5B, two phi 17 and 13 at

0.26 and 0.84 respectively; tarsus 14B, omega 17 at 0.33, famulusnot observed, one eupathid 19 at 0.90. Leg III. 245; coxa IB; 156 trochanter IB; femur 4B, bv seta barbed, d seta barbed; genu 2B, sigma 21 at 0.22; tibia 5B, tarsus 11B, scopa bifurcate, lophotrix thickened seta.

Family EUTROMBIDIIDAE Thor, 1935

Type Genus: Eutrombidium Verdun, 1909

Subfamily Diagnosis (LARVA): Setae lb, 3b and usually 2a on coxal fields legs I, III and II respectively thickened and bifid (except some species of Hoplothrombium where all coxal field setae are setiform); subcapitular setae (scl) hypertrophied; lophotrix present; scopa usually developed; AM nude; famulus on tarsus leg I distal to omega I.

(POSTLARVAL): Dorsal idiosomal setae setiform; posterior idiosoma with pygosomal plate; palpal tibia with two rows of dorsal spines each with approximately 7-10 spines; palpal tibia with 1 to 4 ventral spines.

Referred Genera:

Subfamily Eutrombidiinae Thor, 1935

Tribe Eutrombiini

Eutrombidium Verdun, 1909

Scutodesmus New Genus

Tribe Milliotrombiini NEW TRIBE 157

Atopothrombidium New Genus

Milliotrombidium Shiba, 1976

Subfamily Hoplothrombiinae NEW SUBFAMILY

Hoplothrombium Ewing, 1925

Cryptothrombidium NEW GENUS

Subfamily Eutrombidiinae Thor, 1935

Type Genus: Eutrombidium Verdun, 1909

Diagnosis (LARVA): Subcapitular setae (scl) setae spiniform or rounded; tarsus legs III entire; scopa well developed; paraxial claw on pretarsus leg III modified.

Tribe Eutrombidiini

Type Genus: Eutrombidium Verdun, 1909

Diagnosis (LARVA): Subcapitular setae (scl) setae spiniform; supracoxal seta el absent; femur leg II with five setae.

Eutrombidium Verdun, 1909

Eutrombidium Verdun, 1909, Soc. Biol. 67: 244; Type species: 158

Trombidium trigonum Hermann, 1804

Generic Diagnosis (LARVA): Coxal field seta 2a thickened and bifid; genu legs II and III each with one sigma; palpal femur and genu each with one dorsal seta. For a description and figures of larval

Eutrombidium trigonum Hermann see Robaux, 1974. While Robaux (1974) presents a good description he did omit the small setae on the palpal femur and genu.

(POSTLARVAL): Dorsal idiosomal setae setiform; posterior idiosoma with pygosomal plate; palpal tibia with two rows of dorsal spines with 9 and 7 respectively; palpal tibia with 1 to 4 ventral spines.

Referred Species:

Eutrombidium sp.

E.* feldmanmusae Feider, 1977

E_. trigonium (Hermann, 1804)

Scutodesmus NEW GENUS

Type Species: Scutodesmus araneivarum NEW SPECIES

Generic Diagnosis (LARVA): coxal field seta 2a setiform, thickened only at base, not bifid; palpal femur without seta; prodorsal sclerite and scutellum cover entire dorsal idiosoma; scl seta spiniform. Postlarval instars unknown. 159

Scutodesmus araneivarum NEW SPECIES

Holotype from South Africa, Natal, World's View Hilton, ex forest

leaf litter, iii-1974, B. H. Lamoral. AL-1114.

Description of species. Larva. Idiosoma. Holotype unengorged but

distorted during mounting and not measured; eyes 2/2, with ocular

plates, anterior eye larger smaller. Prodorsal sclerite and

scutellum occupy the entire dorsal idiosoma displacing the normal

dorsal idiosomal setae ventrally and distorting setae patterns.

Cupules absent; supracoxal seta el absent. One pair of nude

intercoxal setae between coxae III; two preanal setae.

Prodorsal Sclerite. Punctate with without striations, covering

entire prodorsum, anterior margin broadly rounded, posterior margin

slightly convex; AM < AL = PL; SB < PW; trichobothridial bases

anterior to PL setal bases; trichobothria flagellate without setules.

Scutal measurements of holotype: AM 14, AA 67, MA 93, AW 182, AL 26,

PW 230, PL 26, AP 44, SB 204), S 66, PSB 20, ASB 107, SD 127.

Scutellum: HS 133, LSS 211, SL 30), SS 76.

Gnathosoma. Palpal setal formula 0-N-SNN2-7NB (palpal trochanter

absent); palp tibial claw two pronged; adoral setae (orl) nude,

subcapitular setae (scl) hypertrophied; palpal supracoxal seta

absent; cheliceral blade not observed, surrounded by buccal cone.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw short, thick and rotated nearly opposite

paraxial claw, paraxial claw and empodium normally developed.

Measurements of holotype with positions of specialized setae of the

holotype given as a ratio of the segment length. IP = 648. Leg I

227); coxa 2B; trochanter IB; femur 6B, bv seta long with 1-2 barbs,

d seta barbed; genu 4B, two sigma 19 and 22 at 0.22 and 0.34

respectively, k 3 at 0.70; tibia 6B, two phi 17 at 0.29 and 0.81

respectively, k 4 at 0.77; tarsus 18B, omega 20 at 0.38, famulus 4 at

0.47, two eupthidia 40 and 17 at 0.67 and 0.92 respectively. Leg II.

219; coxa IB; trochanter IB; fenrar 5B, bv seta long and nude, d seta

barbed; genu 2B, sigma 21 at 0.28, k 3 at 0.69; tibia 5B, two phi 16

and 11 at 0.34 and 0.80 respectively; tarsus 14B, omega 19 at 0.43,

famulus 3 at 0.40, eupathid 20 at 0.93. Leg III. 202; coxa IB;

trochanter IB; femur 4B, bv seta long with one barb, d seta barbed;

genu 2B, sigma 22 at 0.32; tibia 5B, tarsus 11B, scopa well developed with 5-6 barbs, lophotrix with two long basal branched and 6-8 small

barbs.

Milliotrombidiini NEW TRIBE

Type genus: Milliotrombidium Shiba, 1976

Diagnosis (LARVA): Subcapitular setae (scl) rounded (not spinelike); coxal field seta 2a thickened and usually bifid; fnF= 6-4-4; prodorsal sclerite with crescent shaped sculptures. Postlarval 161 instars unknown

Atopothrombidium NEW GENUS

Type Species: Ettmulleria townsendi Dumbelton, 1962, N.Z.J. Sci.

5:8-12.

Generic Diagnosis (LARVA): Subcapitular setae (scl) trilobed; coxal field seta 2a thickened and bifid; genu legs II and III without sigma; palpal femur with single dorsal seta; prodorsal sclerite with faint crescent shaped sculptures. Postlarval instars unknown.

Atopothrombidium townsendi (Durableton, 1962) NEW COMBINATION

Holotype and four paratypes from New Zealand, Unites Creek, Roding

River, ex cave weta (Orthoptera, Rhaphidophoridae) in limestone cave,

J. I. Townsend collector, 6-viii-1960. Types deposited in Entomology

Division, D.S.I.R. Dumbleton, 1962. New Zealand J. Science 5:8-12.

This description based on holotype and two paratypes as well as four additional specimens from New Zealand.

Description of species. Larva. Idiosoma. Holotype engorged but not measured; eyes not observed. Dorsal idiosomal rows with

3-3-3-3-2-1-1; cupules absent; supracoxal seta el present. One pair of branched intercoxal setae between coxae III; two pair of preanal setae.

Prodorsal Sclerite. Punctate without striations, with faint

crescent shaped sculptures, anterior margin broadly rounded,

posterior margin slightly convex; PL > AL > AM; SB < PW;

trichobothridial bases anterior to PL setal bases; trichobothria

flagellate without setules.

Prodorsal sclerite measurements of holotype with mean of two

paratypes in parentheses: AM 37 (45), AA 59 (61), MA 118 (123), AW

177 (180), AL 64 (71), PW 186 (198), PL 73 (81), AP 59 (57), SB 135

(142), S 129 (162), PSB 30 (31), ASB 146 (167), SD 176 (198).

Scutellum: HS 66 (77), LSS 194 (206), SL 63 (77), SS 85 (89) .

Gnathosoma. Palpal setal formula N-N-NNN2-6N (palpal trochanter

absent); palp tibial claw two pronged; adoral setae (orl) nude,

subcapitular setae (scl) thick with three lobes; palpal supracoxal

seta absent; cheliceral blade with one ventral tooth surrounded by

buccal cone.

Legs. Femora on legs I to III undivided, five segments beyond the

coxa; pretarsus legs I to II with paired claws and clawlike empodium;

leg III with antiaxial claw short, thick and rotated, paraxial claw

and empodium longer the other legs but otherwise normally developed.

Measurements of holotype with positions of specialized setae of the

holotype given as a ratio of the segment length. IP =757. Leg I

273; coxa 2B; trochanter IB; femur 6B, bv seta nude, d seta barbed;

genu 4B, one sigma 39 at 0.50, k - (7) at 0.85; tibia 6B, two phi 37 and 34 at 0.47 and 0.90 respectively, k 9 at 0.87; tarsus 19B, omega

29 at 0.22, famulus 3 at 0.42, two eupthidia 42 and 20 at 0.73 and 163

0.99 respectively. Leg II. 249; coxa IB; trochanter IB; femur 5B,

bv seta nude, d seta barbed; genu 2B, k 8 at 0.68; tibia 5B, two phi

27 at 0.41 and 0.81 respectively; tarsus 14B, omega 29 at 0.36,

famulus 4 at 0.23, eupathid 24 at 0.83. Leg III. 234; coxa IB;

trochanter IB; femur 4B, bv seta with few basal barbs, d seta barbed;

genu 2B; tibia 5B, tarsus 11B, scopa without barbs, tapered at tip,

lophotrix reduced to single large prong.

Milliotrombidium Shiba, 1976

Milliotrombidium Shiba, 1976, Nature & Life in S.E. Asia 7:209-212;

Type species: Milliotrombidium milliopodium Shiba, 1976.

Generic Diagnosis (LARVA) (based on a new species (M. japanensis)

described below because type species was not available): Prodorsal

trichobothria expanded; prodorsal sclerite with obvious crescent

shaped sculptures; scopa with or without setules; setae on coxal

fields II and III thickened but may or may not be bifid; without

preanal sclerite; tarsus leg III bifurcate; genu leg without sigma.

Postlarval stages unknown

Milliotrombidium japanensis NEW SPECIES

Holotype and thirty-seven paratypes from Japan, Shikoku, Ichizuchi 164

Mountains National Park, ex Tachycines robustus Ander from mixed warm temperate forest, 18-25-VIII-1980, S.Peck coll., AL-2514, original number WCW810416-12.

Description of species. Larva. Idiosoma. Holotype partially engorged 358 long, 172 wide, paratypes ranged from 284 x 164

(unengorged) to 550 x 252 (engorged); eyes 2/2, with ocular plate

(39 x 15), anterior eye 11; posterior eye 17. Dorsal setae

3-3-3-3-2-3; seta C2 long (67), setae in rows D, E, and F 42-52, H setae 54-67; cupules absent; supracoxal seta (Ie) present. One pair of branched intercoxal setae between coxae III; six preanal setae.

Coxal field leg III rectangular due to increased sclerotization medially.

Prodorsal Sclerite. Lightly punctate with crescent shaped scultures, without striations, anterior margin rounded, posterior margin straight; AL > AM > PL; trichobothridial bases anterior to PL setal bases; trichobothria gradually expanded distally with small setules.

A single scutellum with one pair of setae.

Prodorsal Sclerite measurements of holotype with mean, range and number of paratypes in parentheses: AM 32 (34,23-40, 36),AA 43 (44,

39-49, 36), MA 82 (81, 78-86, 37), AW 103 (105, 99-111, 36), AL 56 (

52, 47-56, 35), PW 122 (121, 116-131, 34), PL 17 (16, 12-20, 37), AP

34 (32, 25-37, 37), SB 95 (95, 90-101, 34), S 89 (88, 83-95, 28), PSB

20 (21, 17-23, 37), ASB 108 (110, 101-119, 37), SD 128 (131,

121-139,35). Scutellum: HS 41 (42, 38-47, 36), LSS 120 (119,

112-126, 34), SL 46 (46, 41-53, 34), SS 39 (41, 35-48, 34). 165

Gnathosoma. Palpal setal formula N-N-NNS2-5N (palpal trochanter absent); dorsal seta on palpal tibia spinelike, palp tibial claw 2 pronged; adoral (orl) setae nude (9), subcapitular (scl) setae clavate, nude with obvious striations (11); palpal coxa without setae; palpal supracoxal seta absent; cheliceral blade not observed.

Legs. Femora on legs I to III undivided (five segments beyond the coxa); pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw thick, short and rotated opposite other claw and empodium, paraxial claw and empodium on leg III normally developed, but longer tha legs I and II. Measurements of holotype with mean, range and number of paratypes in parentheses. Positions of specialized setae of the holotype are given as a ratio of the segment length with mean, range and number of paratypes are given in parentheses. IP = 421 (417, 395-429,34). Leg I 167 (169,

159-180,36); coxa with 2 setae, one nude (N) and other modified into short stout bifid seta (F); trochanter IB; femur 6B; genu 4B, sigma

19 (20, 16-23,36) at 0.57 (0.55, 0.41-0.64, 36), k 6 (7, 5-9, 32) at

0.73 (0.76, 0.67-0.89, 34); tibia 6B, two £hi 19 (18, 14-21, 35) and

23 (23, 20-36, 36) at 0.41 (0.41, 0.34-0.49, 37) and 0.84 (0.83,

0.70-0.94, 37) respectively, k 7 (8, 6-9, 32) at 0.81 (0.82,

0.76-0.93, 36); tarsus 18B (range in paratypes 18-21), omega 21 (22,

19-24, 36) at 0.50 (0.51, 0.47-0.58, 37), famulus 7 (7, 5-9, 23) at

0.59 (0.59, 0.54-0.64, 28), two eupthidia 28 (28, 23-32, 36) and 18

(18, 15-20, 27) at 0.72 (0.72, 0.54-0.74, 36) and 0.82 (0.82,

0.78-0.88, 27) respectively. Leg II. 134 (132, 126-138, 36); coxa

IF; trochanter IB; femur 4B; genu 2B, sigma 16 (15, 14-17, 37), at 166

0.51 (0.48, 0.42-0.58, 37); tibia 5B, two £hi 14 (15, 14-17,37) and

14 (15, 13-18, 36) at 0.47 (0.43, 0.37-0.53, 37) and 0.74 (0.74,

0.69-0.82, 37) respectively; tarsus 14B (range in paratypes 15-14), omega 19 (18, 13-23,37) at 0.24 (0.34, 0.29-0.39, 36), famulus 4 (3,

2-5, 25) at 0.13 (0.19, 0.13-0.23, 31). Leg III. 119 (115,

106-124,36); coxa IF; trochanter IB; femur 4B, bv branched; genu 2B; tibia 5B, tarsus 11B (range in paratypes 11). Tarsus bifurcate with pretarsus on one side and lophotrix on opposite. Scopa reduced to thick seta without setules. Lophotrix thick bifurcate seta without setules.

Hoplothrombiinae NEW SUBFAMILY

Type Genus: Hoplothrombium Ewing, 1925

Tribe Diagnosis (LARVA): Tarus leg III bifurcate; tarsus leg III with only one claw; scopa reduced; tibia leg II with two ventral setae; preanal sclerite present; coxal field leg III rectangular.

Postlarval stages unknown.

Hoplothrombium Ewing, 1925

Hoplothrombium Ewing, 1925, Amer. J. Trop. Med. 5:251-265; Type species: Hoplothrombium quinquescutatum Ewing, 1925, ex "beetle mite 167

taken from the stomach of a toad (Bufo americanus Holbrook)”

collected at Hudson Bay (U.S. Biological Survey No. 585). Holotype in

U. S. N. M. Washington, D.C. no.893.

Generic Diagnosis (LARVA): fcx= 2-1-1 or 2-2-2; coxal field setae lb,

2a and 3b thick and bifid or setiform with thick bases; idiosoma with

three or four scutella in rows C, D, E and H; scopa not developed;

preanal sclerite present; subcapitular setae (scl) thick and

spinelike. Vercaramen-Grandjean (1967) redescribed and illustrated

the type species. Postlarval instars unknown.

Hosts: Cicindelide, Carabidae (Coleoptera)

Distribution: Holarctic, Neotropical & Africa.

Referred Species:

H. quinquescutatum Ewing, 1925

II. cicindelae (Floch and Abonnenc, 1941) NEW COMBINATION

Cryptothrombidium NEW GENUS

Type Species: Cryptpothrombidium howarthi NEW SPECIES

Generic Diagnosis (LARVA): With only one scutellum in setal row C;

palpal femur with single nude seta; genu leg I with only one sigma; 168

without eyes; preanal sclerite small; microseta k absent from genu

all legs. Postlarval instars unknown

Cryptpothrombidium howarthi NEW SPECIES

Holotype and 2 paratypes from Tennessee, Warren Co., McMinnville,

Higganbottom Entrance to Cumberland Caverns, deep zone, ex

Hadenoecus, (Orthoptera, Rhaphidophoridae) 24-vii-1981, F. G.

Howarth.

Description of species. Larva. Idiosoma. Holotype engorged but not maeasured; eyes absent. Dorsal idiosomal setal rows with 3-3-3-3-1-1 pairs; cupules absent; supracoxal seta el present. One pair of nude intercoxal setae between coxae III; twelve preanal setae; preanal sclerite small.

Prodorsal Sclerite. Punctate with striations, anterior margin convex, posterior margin slightly convex; AM > AL > PL; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules; PW/SD = 0.77; W/L = 0.80.

Prdorsal sclerite measurements of holotype with mean two paratypes in parentheses: AM 63 (64), AA 71 (70), MA 73 (72), AW 109 (110), AL 59

(67), PW 128 (130), PL 49 (45), AP 46 (46), SB 89 (90), S 130 (142),

PSB 29 (31), ASB 142 (146), SD 169 (172). Scutellum: HS 24 (25), LSS

113 (114), SL 55 (50), SS 37 (37).

Gnathosoma. Palpal setal formula N-N-NNN2-7Nv/ (palpal trochanter absent); palp tibial claw two pronged; adoral (orl) setae nude, 169 subcapitular (scl) setae spiniform; palpal supracoxal seta absent; cheliceral blade not observed, with buccal cone; gnathobase lightly punctate.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw thick, short and rotated, paraxial claw absent, empodium leg III normally developed but longer than empodia on other legs. Measurements of holotype with mean of two paratypes in parentheses. Positions of specialized setae of the holotype are given as a ratio of the segment length; variation of paratypes is given in parentheses. IP = 500 (501). Leg I 204 (206); coxa 2B; trochanter IB; femur 6B, bv seta nude, d seta barbed; genu 4B, one sigma 10 (10) at 0.57 (0.53); tibia 6B, two phi (19 (18) and 22 (20) at 0.56 (0.53) and 0.82 (0.83) respectively, k - (11) at - (0.60); tarsus 18B, omega 27 (27) at 0.48 (0.49), famulus 9 (10) at 0.57

(0.57), two eupthidia 33 (33) and - (19) at 0.76 (0.74) and - (0.86) respectively. Leg II. 130 (133); coxa IB; trochanter IB; femur 5B, bv seta nude, d seta barbed; genu 2B, sigma 9 (9) at 0.50 (0.55); tibia 5B, two phi 12 (10) and 15 (13) at 0.42 (0.51) and 0.79 (0.78) respectively; tarsus 14B, omega 27 (25) at 0.34 (36), famulus 3 (4) at 0.29 (29). Leg III. 204 (206); coxa IB; trochanter IB; femur 4B, bv seta nude, d seta barbed; genu 2B, sigma 9 (9) at 0.39 (0.37); tibia 5B, tarsus 11B, scopa not developed, lophotrix with many branches fused except at tips giving appearance of brushes. 170

Family MICROTROMBIDIIAE Thor, 1935

Type Genus: Microtrombidium Haller, 1882

Family Diagnosis (Larva): famulus tarsus leg I proximal to omega I; subcapitular setae (scl) modified (minute, spinelike or palmate); lophotrix with one large basal spine or none; lophotrix may be lost in some genera.

(POSTLARVAL): Prodorsal sclerite narrow; 1-2 rows of palpal tibial spines; accessory claw well developed; idiosomal setae variable from setiforra to expanded and globose; eyes present.

Referred Genera:

Subfamily Feideriinae Vercammen-Grandjean, 1973

Tribe Manriquiini Welbourn, 1984

Manriquia Boshell & Kerr, 1942

Reticulatrombium Shiba, 1976

Pygothrombium NEW GENUS

Tribe Feideriini Vercammen-Grandjean, 1973

Valgothrombium Willmann, 1940

Gonothrombium Feider, 1948

Subfamily Microtrombidiinae Thor, 1935

Tribe Microtrombidiini

Microtrombidium Haller, 1882

Platytrombidium Thor, 1936 171

Ettmulleria Oudemans, 1911

Atractothrombium Feider, 1951

Achelothrombidium NEW GENUS

Tribe Cercothrombiini NEW TRIBE

Diathrombium Moser & Vercammen-Grandjean, 1979

Campylothrombium Krausse, 1912

Lithothrombidium NEW GENUS

Trichotrombidium Kobulej, 1950

Willmannella Feider, 1951

Cercothrombium Methlagl, 1928

Subfamily FEIDERIINAE Vercammen-Grandjean & Cochrane, 1974

Type Genus: Feiderium Vercammen-Grandjean & Cochrane, 1974

Subfamily Diagosis (Larva): subcapitular seta (scl) reduced, spinaforra; fcx= BB-B-B; AM setae branched.

(POSTLARVAL): Dorsal idiosoma with two types of setae: most setae are short, setiform to slightly inflated, some sete long and bladelike with distal barbs; palpa tibia with two rows of 6 and 10 spines respectively; prodorsal sclerite not much wider than crista metopica. 172

Tribe Feideriini

Type Genus: Feiderium Vercammen-Grandjean & Cochrane, 1974

Tribe Diagnosis (Larva): pretarsus legs I and II without empodium; microseta k absent on genu leg II.

Valgothrombium Willmann, 1940

Valgothrombium Willmann, 1940, Zool. Anz. 131:250; Type species:

Ottonia valga George, 1909.

Feiderium Vercammen-Grandjean, 1973, Acarologia 15:113, Type species:

Microtrombidium tirnavense Feider, 1949. NEW SYN0N0MY

F. (Parafeiderium) Vercammen-Grandjean & Cochrane, 1974, J. Kansas

Ent. Soc. 47:66-72, Type species: JF. (Parafeiderium) culicoidium. NEW

SYN0N0MY

Diagnosis (Larva): prodorsal sclerite with three rounded posterior projections with the middle projection aligning with an anterior projection on the scutellum; scutellum with prorainant grove in the middle; prodorsal sclerite and scutellum without prominant longitudinal striations.

(POSTLARVAL) prominant striations on anterior end of prodorsal 173 sclerite; palpal tibia with one long row of dorsal spines; dorsal idiosomal setae expanded.

Hosts: Ceratopogonidae (Diptera)

Distribution: Holarctic, Neotropical

Referred Species:

Valgothrombium valRa (George, 1909)

V_. culicoidium (Vercammen-Grandjean & Cochrane, 1974) NEW COMBINATION

V. confusum (Berlese, 1910) NEW COMBINATION

V. fluminis (Michener, 1946) NEW COMBINATION

V. homocomum (Berlese, 1918) NEW COMBINATION

Gonothrombium Feider, 1948

Gonothrombium Feider, 1948, Ann. Scienti. Universite Jassy 31:205;

Type species: Microtrombidium bimaculatum Feider, 1948.

Blankaartia Oudemans, 1911, Zool. Jahrb. Suppl. 14:118; Type speices:

Trombidium niloticum Tragardh, 1905.

Pseuoblankaartia Fuller & Wharton, 1951, Psyche 58:87; Type species:

Pseudoblankaartia bequaerti Fuller & Wharton, 1951 (name applied to larva associated with postlarval Trombidium niloticum Tragardh, 1905. 174

Diagnosis (LARVA): Scutellum with at least 10 setae; prodorsal

sclerite and scutellum striate; idiosoma neotrichous.

(POSTLARVAL): Dorsal idiosomal setae expanded; palpal tibia with

ventral spine.

Hosts: unknown

Distribution: Europe

Referred Speices:

Gonothrombium bimaculatum (Feider. 1948)

£. oudemansianum (Feider, 1948)

£. bequaerti (Fuller & Wharton, 1951)

Tribe Maniquiini Welbourn, 1984

Type Genus: Manriquia Boshell & Kerr, 1942

Diagnosis (LARVA): Scopa present; lophotrix with one very long basal branch; dorsi idiosoma with reticulation or secondary scleritization.

(POSTLARVAL): Dorsal idiosoma with two distinct types of setae; palpal tibia with one long row of dorsal spines.

Manriquia Boshell & Kerr, 1942 175

Manriquia Boshell & Kerr, 1942, Rev. Acad. Colombia Cienc. Bogota

5:17; Type species: Manriqlii bequaerti Boshsell and Kerr, 1942.

Diagnosis (LARVA): Genu legs II and III each with two sigma; dorsal opisthosoma with distinct reticulate pattern; For description and figures of species see Robaux (1977).

(POSTLARVAL): Dorsal idiosoma with two distinct types of setae; palpal tibia with one long row of dorsal spines. See Robaux (1967) for a redescription of the adult.

Hosts: unknown

Distribution: Neotropical

Referred species:

Manriqui bequaerti Boshsell and Kerr, 1942; M_. boshelli

Michener,1946; M. kethleyi (Robaux, 1977) NEW COMBINATION; M. panaminsis Michener,1946.

Reticulatrombidium Shiba, 1976

Type species: Reticulatrombidium serratum Shiba, 1976, Nature and

Life in Southeast Asia 7:212; Holotype from Malaysia, Negeri

Senbilan, Pasoh Forest Reserve, east of plot 1; 4-iii-1971; collector

Shiba. 176

Generic Diagnosis (LARVA): fsol I (0-2-2-1) II (0-2-2-1)

111(0-2-0-0); fz_= 2-1-0; fpP= 0-0-NNS2-5Nw_; scl setae reduced to small spines; dorsal and posterioventral idiosoma with reticulate pattern; dorsal setae heavily setulate; prodorsum wider than long; pertarsus leg III without paraxial claw; lophotrix and scopa absent.

See Shiba (1976) for descripion and figures of ji. serratum.

Postlarval instars unknown.

Hosts: unknown

Distribution: Asia, Africa

Referred Species:

R^. serratum Shiba, 1976

Pygothrombium NEW GENUS

Type Species: Pygothrombium NEW SPECIES

Generic Diagnosis (larva): Posterior idiosoma with sclerotized plate

(pygosmal plate); subcapitular setae (scl) reduced, rounded; setae on coxal plate leg I nude; lophotrix and scopa present. Postlarval instars unknown. 177

Hosts: uknown

Distribution: Papua New Guinea, Brasil

Pygothrombium bakeri NEW SPECIES

Holotype and 2 paratypes from Papua New Guinea, Morobe Province, Buso

Forest, ex malaise trap, September 1979, J. H. Martin, original number 2559.

Description of species. Larva.

Idiosoma. Holotype engorged; due to condition of idiosoma the holotype and paratyes could not be measured; eyes 2/2, ocular plates

(89 x 26) with anterior projection, anterior eye 11, posterior eye 8.

Due to the condition of the type series it was not possible to determine setal rows; pygosomal plate with two pair of setae, possible the H and PS series; cupules absent; supracoxal seta absent.

One pair of nude intercoxal setae between coxae III; one pair of preanal setae.

Prodorsal Sclerite. Punctate without striations, anterior margin convex, posterior margin straight; AL > AM > PL; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules.

Prodorsal sclerite measurements of holotype with mean of two paratypes in parentheses: AM 69 (71), AA 70 (82), MA 89 (88), AW 162

(166), AL 116 (119), PW 182 (188), PL 53 (50), AP 30 (36), SB 146 (153), S - (121), PSB 19 (23), ASB 117 (127), SD 136). Scutellum: SL

119 (110), HS 87 (85), SS 131 (143), LSS 182 (193).

Gnathosoma. Palpal setal formula N-N-NSS2-6Nw (palpal trochanter absent); palp tibial claw two pronged; adoral setae (orl) nude, subcapitular setae (scl) reduced and rounded; palpal supracoxal seta absent; cheliceral blade with a single tooth and surrounded by buccal ring.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw short, thick and rotated opposite other claw, paraxial claw and empodium longer than conterparts on legs I and II but otherwise normally developed. Measurements of holotype.

Positions of specialized setae of the holotype are given as a ratio of the segment length. Seta bv nude on femur leg I and with 1-3 barbs on femur legs II and III. IP = 866. Leg I 307; coxa 2B; trochanter IB; femur 6B; genu 4B, two sigma 40 both at 0.27, k 2 at

0.71; tibia 6B, two phi 40 and 27 at 0.35 and 0.91 respectively, k 7 at 0.65; tarsus 18B, omega 33 at 0.44, famulus 14 at 0.21, two eupthidia 26 and 18 both at 0.88. Leg II. 244; coxa IB; trochanter

IB; femur 5B; genu 4B, sigma 25 at 0.23, k 4 at 0.55; tibia 5B, two phi 22 and 20 at 0.42 and 0.55 respectively; tarsus 14B, omega 22 at

0.46, famulus 3 at 0.37, eupathid 16 at 0.88. Leg III. 316; coxa

IB; trochanter IB; femur 4B; genu 2B, sigma 27 at 0.25; tibia 5B, tarsus 11B, scopa with 4-5 barbs, lophotrix with one long basal branch. 179

Subfamily Microtrorabidiinae Thor, 1935

Type Genus: Microtrombidium Haller, 1882

Subfamily Diagnosis (larva): AM nude; subcapitular setae (scl) palmate (may be secondarily reduced in some genera); d seta femur legs I, II and occasionally III short, usually nude and located at the anterior end of the tarsus.

(POSTLARVAL): Dorsal idiosomal setae setaform or slightly inflated.

Tribe Microtrombidiini

Type genus: Microtrombidium Haller, 1882

Diagnosis (larva): lophotrix without larva basal spine, may be setaform; (POSTLARVAL): Dorsal idiosomal setae setaform or slightly inflated (not divided into two parts; palp femur, genu and basal segments of the legs without brushlike setae; palpal tibia with large ventral spine, may be located anteriorly in some genera (absent in

Platytrombidium and reduced in Microtrombidium); spines on dorsal palpal tibia in two short rows. 180

Microtrombidium Haller, 1882

Microtrombidium Haller, 1882, Jahr. Wurtonb. 38:322; Type species:

Trombidium pusillum Hermann, 1804

Megophthrombidium Mullen & Grandjen, 1978, Intl. J. Acarology

4:39-45; Type species: Megophthrombidium welleslyi Mullen &

Vercammen-Grandjean, 1978. NEW SYN0N0MY

Diagnosis (LARVA): Lophotrix with long branch located distally giving the seta a "Y" shape; prodorsal sclerite and scutellum without striations; anterior eye very large, with ocular plate fused to prodorsal sclerite; scl reduced.

(POSTLARVAL): Dorsal idiosomal setae setaform; palpal tibia with two short rows of 2 to 4 spines; tarsus leg I expanded.

Hosts: Culicidae (Diptera)

Distribution: Cosmopolitan

Referred Species:

Microtrombidium columbianum Berlese, 1910; M. gracile (Moser &

Vercammen-Grandjean, 1979) NEW COMBINATION; M. italicum Berlese,

1912; M. l'abanicum Berlese, 1910; M. leptochirum Berlese, 1912; M. welleslyi (Mullen and Vercammen-Grandjean, 1978) NEW COMBINATION; M. spp. Platytrombidium Thor. 1936

Platytrombidium Thor. 1936, Zool. Anz. 114:32; Type species:

Trombidium vagabundum Berlese, 1903.

Cylindrothrombium Feider, 1956, Ann. Stiint. Univeristy "Al. I.

Cuza", Iasi Sect.II, Stiint. Nat. 1:86-87; Type species: Trombidium fasciatum Koch, 1836. NEW SYN0N0MY

Diagnosis (LARVA): Lophotrix without large distal branch; prodorsal sclerite and scutellum striate.

(POSTLARVAL): Dorsal idiosomal setae expanded;

Hosts: Anthomyziidae, Chloropidae, Drosophilidae, Muscidae,

Tephrididae (Diptera)

Distribution: Europe

Referred Species:

P. vagabundum Berlese, 1884

P. fasciatum (Koch, 1836) NEW COMBINATION

Ettmuelleria Oudemans, 1911

Ettmuelleria Oudemans, 1911, Ent. Ber. 3:57; Type species: Trombidium 182 sucidium Tragardh, 1910.

Diagnosis: famulus tarsus leg I distal to omega I; genu legs I, II and III each with two sigma; prodorsal scperite and scutellum without striations; bv setae with few setules. Postlarval instars unknown

Hosts: unknown

Distribution: Europe

Referred Species:

Ettmulleria sucidum (Tragardh, 1910)

Atractothrombium Feider, 1951

Atractothrombium Feider, 1951, Bull. Stiint. Sect. Stiint. Biol.,

Agron., Geol. et Geog. 4:609-610; Type species: Microtrombidium

(Enemothrombium) fusiconum Berlese, 1910.

Atractothrombodies Feider, 1975, Rev. Roum. Biol.- Biol. Anim.

20:223-227, Type species: Atractothromboides danubialis

NEW SYN0N0MY

Diagnosis (LARVA): Lophotrix absent; scopa absent; genu leg II without microseta k; pretarsus legs I and II without empodium; prodorsal sclerite and scutellum striate. (POSTLARVAL): Dorsal idiosomal setae inflated; palpal tibia with

large ventral spine;

Hosts: Ceratopogonide (Diptera)

Distribution: Holarctic, Neotropical

Referred Species:

danubialis (Feider, 1975)

A_. dictyostracum Vercammen-Grandjean & Cochrane, 1974

_A. fusiconum (Berlese, 1910)

A.. heterocomum (Berlese, 1918) NEW COMBINATION

A. littorale (Michener, 1946) NEW COMBINATION

A. sylvaticum (Koch, 1836) NEW COMBINATION

A. trispinum (Berlese, 1910) NEW COMBINATION

Achelothrombidium NEW GENUS

Type Species: Microtrombidium pistae Mitchener, 1946.

Diagnosis (LARVA): Pretarsus leg III without paraxiial claw; parodorsal sclerite and scutellum without striations;

(POSTALARVAL): Dorsal idiosomal setae setiform; palpal tibia with short thick ventral spine between palpal tibial claw and palpal tarsus articultion. 184

Hosts: Culicidae (Diptera)

Distribution: Panama, Guatemala

Referred Species:

A. pistiae (Michener, 1946) NEW COMBINATION

A. maculatum (Michener, 1946) NEW COMBINATION

Achelothraombidium pistiae (Mitchener, 1946) NEW COMBINATION

Panama, Juan Mina, Canal Zone, 18-vi-1945; "hatched from eggs laid on

Pistia plants"; C. D. Michener coll.

Description of species. Larva. Idiosoma. Holotype unengorged; eyes

2/2, with ocular plate, anterior eye larger smaller. Dorsal setal rows 3-3-3-2-1-1. Cupules absent; supracoxal seta (el) absent. One pair of branched intercoxal setae between coxae III; two preanal setae.

Prodorsal Sclerite. Punctate without longitudinal striations, anterior margin rounded, posterior margin straight to convex ; PL >

AM > AL; SB > PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate with setules.

Prodorsal sclerite measurement of 12 larvae with range in parenthesis: AM 42 ( 35-43), AA 41 (35-46), MA 46 (42-48), AW 68

(64-72), AL 37 (34-40), PW 82 (77-85), PL 53 (50-57), AP 47 (44-50),

SB 57 (54-60), S 83 (79-87), PSB 24 (22-29), ASB 82 (75-88), SD 107 185

(100-112). Scutellum: HS 34 (30-39), LSS 77 (74-80), SL 86 (81-92),

SS 40 (38-45) .

Gnathosoma. Palpal setal formula 0-0-NNN2-5N (plapal trochanter absent); palp tibial claw two pronged; adotal setae (orl) nude , subcapitular setae (scl) short and nude; palpal supracoxal seta (e) absent; cheliceral blade with one tooth; buccal cone present.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw absent, paraxial claw and empodium leg

III normally developed. Measurements of 12 larvae with range in parenthesis. Positions of specialized setae of the holotype are given as a ratio of the segment length with range in parenthesis.

IP= 850 (826-869). Leg I 285 (276-290); coxa 2B; trochanter IB; femur 6B, bv seta branched, d seta short and nude at distal end of segment; genu 4B, two sigma 47 (40-55) and 54 (46-60) at 0.10

(0.10-0.15), k 8 (7-9) at 0.79 (0.75-0.82); tibia 6B, two phi 35

(32-37) and 36 (32-38) at 0.23 (0.22-0.25 ) and 0.33 (0.27-0.39) respectively, k 7 (6-8) at 0.85 (0.82-0.87); tarsus 18B, omega 33

(28-37) at 0.15 (0.14-0.15), famulus 12 (9-13) at 0.06 (0.06-0.07), two eupthidia 30 (29-30) and 16 (15-17) at 0.77 (0.76-0.78) and 0.95

(0.93-0.97) respectively. Leg II. 267 (259-273); coxa IB; trochanter IB; femur 5B, bv seta branched, d seta short and nude at distal end of segment; genu 2B, sigma 51 (42-59), at 0.18

(0.13-0.20); k 7 (6-9) at 0.78 (0.75-0.84); tibia 5B, two phi 32

(30-34) and 33 (32-33) at 0.16 (0.14-0.19) and 0.83 (0.81-0.86) respectively; tarsus 14B, omega 35 (34-36) at 0.45 (0.43-0.47), 186 famulus 4 (3-5) at 0.43 (0.42-0.43), eupathid 16 (14-17) at 0.93

(0.91-0.95). Leg III. 297 (290-307); coxa IB; trochanter IB; femur

4B, bv seta branched, d seta branched; genu 2B, sigma 50 (50-51) at

0.17 (0.13-0.18); tibia 5B, tarsus 11B, scopa and lophotrix absent.

Cercothrombiinae NEW TRIBE

Type genus: Cercothrombium Methlagl, 1928

Diagnosis (Larva): Lophotrix with at least one nude basal branch and

2-3 midlength branches.

(POSTLARVAL): Idiosomal setae variously expanded, usually globular; palpal femur and tibia, and at least basal segment of legs with brushlike setae; prodorsal sclerite limited to crista metopica; palpal tibia with two rows of spines and 0-3 spines opposite palpal tarsal articulation.

Cercothrombium Methlagl, 1928

Cercothrombium Methlagl, 1928, Denk. Ak. Wien 101:228-231; Type species: Cercothrombium loricatum Methlagl, 1928

Camerotrombidium Sig Thor, 1935, Zool. Anz. 114:31-32, type speices:

Trombidium pexatum Koch, 1837. NEW SYN0N0MY 187

Diagnosis (LARVA): Basal plates of setae dl fused; coxal field* seta la nude; coxal field seta lb with long barbs; prodorsal sclerite and scutellum with striations; bv setae on all femur legs I, II and III with barbs.

(POSTLARVAL): Dorsal idiosoraa with two types of setae, expanded globose and short irregular setae; palpal tibia with two rows of 6-8 spines each; two to four large ventral spines.

Hosts: unknown

Distribution: Palearctic, Southeast Asia

Referred Species:

C. curtulum (Berlese, 1910) NEW COMBINATION; £. diversum (Berlese,

1910); C. pextum (Koch, 1837) NEW COMBINATION; C. rasum (Berlese,

1910); C^. sanguineum (Koch, 1837) NEW COMBMBINATION; £. sigthori Thor and Willmann, 1947 NEW COMBINATION; C. vesiculosum (Thor, 1900) NEW

COMBINATION.

Cercothrombium hillyardi NEW SPECIES

Holotype and 15 paratypes reared from eggs deposited by female collected under rock in mixed woodland, Great Britan, Wales, Powys,

Glaisfer nr. Llagynide, elev. 420 m, 22-iv-1984, W. C. Welbourn, A. 188

Baker and P. Hillyard.

Description of species. Larva. Idiosoma. Holotype unengorged; eyes

2/2, with ocular plate (55 x 24), anterior eye 15, posterior eye 7.

Dorsal idosomal rows 3-3-3-3-1-1; basal sclerite of setal pair cl fused; PS setae very long; cupules absent; supracoxal seta (el) present. One pair of branched intercoxal setae between coxae III; two pairs of preanal setae.

Prodorsal Sclerite. Punctate with longitudinal striations, anterior margin rounded with overlap onto ventral idiosoma, posterior margin convex; PL > AL > AM; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules.

Prodorsal sclerite measurements of holotype with mean, range and number of paratypes in parentheses: AM 46 (44, 35-50, 15), AA 82 (81,

77-89, 8), MA 82 (83, 77-89, 15), AW 155 (154, 146-159, 11), AL 72

(71, 67-74, 15), PW 123 (119, 128-117, 10), PL 86 (87, 84-93, 15), AP

74 (73, 68-80, 15), SB 127 (130, 123-135, 12), S 108 (102, 91-109,

15), PSB 46 (46, 43-53, 13), ASB 167 (160, 150-175, 15), SD 210 (208,

194-217, 16). Scutellum: HS 33 (35, 33-40, 13), SL 88 (85, 82-89,

15), LSS 178 (173, 153-179,8), SS 53 (51, 48-57, 15).

Gnathosoma. Palpal setal formula N-N-NNN2-6N (palpal trochanter absent); palp tibial claw recurved with two prongs; adoral setae

(orl) nude, subcapitular setae (scl) palmate; palpal supracoxal seta absent; cheliceral blade with one ventral tooth and tubercel midway along length; buccal cone present.

Legs. Femora on legs I to III undivided, five segments beyond the 189 coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw short, thick and rotated opposit other claw, paraxial claw and empodium longer than ones on legs I and II otherwise normally developed. Measurements of holotype with mean, range and number of paratypes in parentheses. Positions of specialized setae of the holotype given as a ratio of the segment length variation of paratypes are given in parentheses. IP =697

(694, 675-709, 15) . Leg I 258 (254, 248-260, 25); coxa NB; trochanter IB; femur 6B, bv seta barbed, d seta barbed; genu 4B, two sigma 26-28 (29, 25-33, 15) between 0.15 and 0.30, k 6 (5, 4-7, 15) at 0.72 (0.69, 0.63-0.73, 15); tibia 6B, two phi - (26, 17-29, 15) and 18 (20. 17-23, 15) at 0.23 (0.23, 0.20-0.26, 15) and 0.51 (0.52,

0.49-0.55, 15) respectively, k 5 (6, 4-8, 11) at 0.87 (0.84,

0.79-0.89, 15); tarsus 18B, omega 23 (24, 22-26, 15) at 0.19 (0.22,

0.19-0.25, 15), famulus 5 (6, 5-8, 11) at 0.11 (0.13, 0.10-0.15, 15), two eupthidia 35 (36, 32-38, 15) and 17 (18, 16-19, 14) at 0.72

(0.72, 0.70-0.77, 15) and 0.91 (0.92, 0.89-0.94, 15) respectively.

Leg II. 207 (208, 203-212, 15); coxa IB; trochanter IB; femur 5B, bv seta barbed, d seta barbed; genu 2B, sigma 29 (31, 27-33, 15) at

0.26 (0.24, 0.21-0.30, 15), k 5 (5, 4-7, 14) at 0.60 (0.65,

0.59-0.74, 15); tibia 5B, two £hi 22 (24, 21-27, 15) and 15 (17,

14-19, 15) at 0.25 (0.24, 0.18-0.25, 15) and 0.43 (0.42, 0.37-0.50,

15) respectively; tarsus 14B, omega 21 (22, 20-23, 15) at 0.45 (0.43,

0.39-0.45, 15, famulus 2 (3, 2-3, 14) at 0.45 (0.44, 0.38-0.49, 15), eupathid 16 (18, 16-19, 15) at 0.92 (0.91, 0.89-0.97, 15). Leg III.

232 (232, 222-241, 15); coxa IB; trochanter IB; femur 4B, bv seta 190 barbed, d seta barbed; genu 2B, sigma 29 (30, 27-33, 15) at 0.26

(0.28, 0.23-0.33, 15); tibia 5B, tarsus 11B, scopa with four dorsal barbs, lophotrix three long barbs, basal longest.

Campylothrombium Krausse, 1916

Campylothrombium Krausse, 1916, Zool. Anz. 47:181; Type species:

Microtrombidium (Campylothrombium) langhofferi Krausse, 1916

Diagnosis (LARVA): Coxal field seta la nude; coxal field seta lb setiform; d seta on femur legs I and II nude

(POSTLARVAL): Dorsal idiosomal setae in two distict segments with distal being smaller than basal; palpal tibia with two rows of 8-14 setae; palpal tibia with one large spine opposite palpal tarsal articulation; accessory claw half length of palpal tibial claw.

Hosts: unknown

Distribution: Europe

Referred Species:

C. barbarum (Lucus, 1847); CL boreale (Berlese, 1910); _C. densipapillum (Berlese, 1910); £. langhofferi Krausse, 1916. 191

Trichotrombidium Kobulej, 1950

Trlchotrombidium Kobulej, 1950, Acta. Vet. Hung. 7:85; Type species:

Trombidium muscarum Riley, 1878.

Philogothrombium Feider, 1952, Bull. Stiinit. Sect. Stiinie Biol.,

Agron., Grol. si Geog. 4:619-620.

Diagnosis: bv seta on all femora barbed; setae in coxal fields legs

I, II and III branched; prodorsal sclerite and scutellum with marginal striations; d setae femora legs I and II nude

(POSTLARVAL): Dorsal idiosoma with two types of setae, expanded globose and shore irregular setae; palpal tibia with two rows of 6 and 4 spines respectively; one large spine opposite palpal tarsal articulation; palpal tibial accessory claw nearly as long as tibia claw.

Hosts: Calliphoridae, Muscidae, Sarcophagidae (Diptera)

Distribution: Cosmopolitan

Referred Species:

Trichotrombidium muscarum Riley, 1878

(=Trombidium hemistriatum Womersley, 1942) NEW SYN0N0MY 192

Diathrombiunr Moser & Vercammen-Grandjean, 1979

Diathrombium Moser & Vercammen-Grandjean, 1979, Int. J. Acarology

5:22; Type species: Diathrombium diaphane

Moser & Vercammen-Grandjean, 1979

Diagnosis (LARVA): coxal field seta la bifurcate; lophotrix seyiform; subcapitular setae (sci) reduced to few setules. For description see

Moser and Vercammen-Grandjean, 1979

Postlarval forms unknown

Host: Dentroctomus frontalis Zimmerman, 1868 (Coleoptera, Scolytidae) and Diptera

Distribution: North America

Referred Species:

Diathrombium diaphane Moser & Vercammen-Grandjean, 1979

Willmannella Feider, 1951

Willmannella Feider, 1951, Bull. Stiinte. Biol., Agron., Geol. si

Geog. 4:625-626; Type species: Ottonia phyllophora Canestrlni, 1897.

Diagnosis (LARVA): Coxal field seta la nude; coxal field seta lb 193

barbed; dorsal idiosoraal setae in rows E and F short (less than half

the length of SL setae on scutellum); bv setae on femor legs I, II and III with 1-2 setules; prodorsal sclerite with striations; scutellum with reticulate pattern.

(POSTLARVAL): Dorsal idiosomal setae of two types, one with expanded distal portion and other with disc shaped top; palpal tibia with two rows of 10-13 and 20-23 spines respectively; palpal tibia without ventral spines; accessory claw subequal to tibial claw.

Host: unknown

Distribution: Holarctic

Referred Species:

Willmannella. johnstoni (Robaux, 1977) NEW COMBINATION

W. racovitzia (Feider, 1948);.

Lithothrombidium NEW GENUS

Type Species: Trombidium perscabrum Banks, 1916

Diagnosis (LARVA): Famulus on tarsus leg I and II distal to omega I and II respectively; d on femur legs I and II nude; bv seta on femur legs I, II and III with 1-3 setules. 194

Host: unknown

Distribution: known only from California.

Description of larval Lithothrombidium perscabrum (Banks, 1916) reared from female collected under rock associated with ants in San

Gabriel Canyon, NE of Azusa, Los Angeles Co., California, 9-iv-1982,

W. C. Welbourn.

Idiosoma. Holotype unengorged not measured; eyes 2/2, with ocular plate (29 x 14), anterior eye 11, posterior eye 6. Dorsal idiosomal setal rows 3-3-3-3-1-1; PS seta long; basal sclerites of dl pair separate; cupules absent; supracoxal seta absent. One pair of branched intercoxal setae between coxae III; two pairs of preanal setae.

Prodorsal Sclerite. Punctate with longitudinal striations, anterior margin rounded, with lateral overlap, posterior margin convex; AM >

PL > AL; SB < PW; trichobothridial bases anterior to PL setal bases; trichobothria flagellate without setules; scutellum with striations.

Prodorsal sclerite mean measurements of holotype with range of 22 reared larvae and number of specimens in parentheses: AM 46 (42-52,

15), AA 56 (55-58, 3), MA 63 (57-66, 18), AW 97 (95-98, 4), AL 27

(23-34, 16), PW 105 (103-109, 10), PL 31 (28-33, 18), AP 46 (43-48,

18), SB 85 (79-91, 14), S 68 (62-73, 17), PSB 26 (23-31, 18), ASB 124

(116-137, 13), SD 150 (144-163, 13). Scutellum: HS 50 (46-54, 18),

LSS 129 (122-135, 7), SL 33 (29-35, 18), SS 60 (54-67, 18). 195

Gnathosoma. Palpal setal formula N-N-NNN2-6N (palpal trochanter absent); palp tibial claw recurved with two prongs; adoral setae

(orl) nude, subcapitular setae (scl) palmate; palpal supracoxal seta absent; cheliceral blade with one ventral tooth and tubercle in internal side; buccal cone present.

Legs. Femora on legs I to III undivided, five segments beyond the coxa; pretarsus legs I to II with paired claws and clawike empodium; leg III with antiaxial claw short, thick, rotated opposite other claw, paraxial claw and empodium longer than claws and empodium on other legs. Mean measurements of 22 reared larvae with range and number of specimens in parentheses. Positions of specialized setae of the holotype given as a ratio of the segment length; range of specimens in parentheses. IP = 627 (604-641, 15). Leg I 231

(217-336, 15); coxa 2B; trochanter IB; femur 6B, bv seta 2-4 barbs , d seta nude and near distal end of femur; genu 4B, two sigma 49-53

(45-55, 15) between 0.10 and 0.19 (0.06-0.25, 15), k 4 (2-5, 12) at

0.81 (0.75-0.92, 13); tibia 6B, two £hi 29 (28-31, 15) and 16 (15-18,

15) at 0.47 (0.41-0.51, 15) and 0.73 (0.69-0.77, 15) respectively, k

4 (3-5, 10) at 0.89 (0.81-0.92, 14); tarsus 18B (17-19,22), omega 21

(20-23, 15) at 0.36 (0.33-0.40, 15), famulus 4 (3-5, 10) at 0.43

(0.38-0.49, 14), two eupthidia 30 (28-32, 14) and 13 (12-14, 15) at

0.72 (0.68-0.76, 15) and 0.95 (0.93-0.98, 15) respectively. Leg II.

196 (190-203, 15); coxa IB; trochanter IB; femur 5B, bv seta few barbs, d seta nude and near distal end of femur; genu 2B, sigma 52

(42-57, 15) at 0.11 (0.08-0.16, 15), k 4 (3-5, 10) at 0.76

(0.69-0.81, 15); tibia 5B, two £hi 29 (27-31, 15) and 12 (10-14, 15) at 0.17 (0.15-0.19, 15) and 0.51 (0.45-0.57, 15) respectively; tarsus

14B (14-15, 21), omega 22 (20-23, 15) at 0.41 (0.37-0.43, 15), famulus 2 (1-3, 11) at 0.46 (0.41-0.49, 15), eupathid 13 (12-14, 15) at 0.95 (0.91-0.97, 15). Leg III. 200 (195-208, 15); coxa IB; trochanter IB; femur 4B, bv seta with few barbs, d seta barbed; genu

2B, sigma 53 (51-55, 14) at 0.16 (0.14-0.19, 15); tibia 5B, tarsus

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Table 9. List of Characters used in Phylogenetic Analysis

210 Table 9. List of Characters Used in Phylogenetic Analysis

Primitive Derived

Number of adofal setae a) two pairs b) one pair

Number of phi on tibia leg I a) two b) one

Position of subcapitular setae (scl) a) distal b) proximal

Shape of subcapitular setae (scl) a) setiform b) minute

c) spinelike

d) rounded

e) palmate

Number of setae on palp femur a) 4f b) 3

c) 2

d) 1

e) 0

Seta on palpal femur a) setiform b) reduced Table 9 (continued)

7. Number of setae on palpal genu a) 3 b) 2

c) 1

d) 0

8. Seta on palpal genu a) setiform b) reduced

9. Palp femur-gend articulation a) separate b) fused

10. Number of tines on palpal tibial claw a) entire b) bifurcate

c) trifurcate

d) multifurcate

e) entire (reversal)

f) absent

11. Zeta on palp tarsus a) present b) absent

12. "Spine" on distal end of palp tarsus a) absent b) present

13. Supracoxal setae (e) a) present b) absent

14. Buccal ring a) absent b) present

15. Cheliceral shape a) blade like b) needle like 212 16. Teeth on chelicerae a) 1 b) 2

c) many Table 9 (continued)

17. Setae on chelicerae a) present b) absent

18. Peritreme a) present b) absent

19. Prodorsal trichobothria a) two pairs b) one pair

20. Number of setae on prodorsal sderite

(excluding trichobothria) a) 6 b) less then

c) more then (neotrichous)

21. AM seta a) present b) absent

22. AM seta a) branched b) nude

c) bifurcate

23. Position of prodorsal trichobothria

relative to PL setae a) anterior b) posterior

24. Prodorsal tichobothria a) seta form b) expanded

c) bifurcate

25. Position of PL relative to

prodorsal scelrite Table 9 (continued)

26. SB relative to PW a) SBPW

27. Naso on prodorsal sderite a) present b) absent

28. Ornamentation of prodorsal sderite a) none b) striate

c) striate on margin only

d) crescent sculptures

e) reversal

29. Shape of prodorsal sderite a) rounded b) rectangular

c) irregular

d) ventral or absent

30. Cupules a) present by absent

31. Number of setae in C-row a) two pairs b) three pairs

c) neotrlchous

32. Number of setae in D-rov a) two pairs b) three pairs

c) neotrlchous

33. Number of setae in B-row a) two pairs b) three pairs

c) two pairs 214

d) neotrlchous Table 9 (continued)

34. Number of setae in F-row a) two pairs b) three pairs

c) two pairs

d) neotrlchous

35. Number of setae in H-row a) 2+ pairs b) one pair

36. Number of setae in PS-row a) 2+ pairs+ b) one pair

37. Pygosomal plate on dorsal idiosoma a) absent b) present

38. Scutellum a) absent b) present

39. Ornamentation of scutellum a) none b) striations

c) striations on margin

d) crescent shaped scultures

e) reticulate

40. Setae on scutellum a) 2 b) multiple setse (4+) Table 9 (continued)

41. Scutellum size a) normal b) reduced

c) very large

42. Additional dorsal idiosomal plates a) none b) D row

c) D & E rows

d) D, E & H rows

43. Basal setal plates of c2 setae a) small b) large

44. Basal setal plates of dl setae a) normal b) large

c) fused

45. Number of eyes a) 2 pair b) 1 pair

c) none

46. Ocular sderite a) present b) absent

47. Cuticle of dorsal opistosoma a) striate b) reticulate

c) stellate

48. Supracoxal setae (el) a) present b) absent

49. Coxal field leg I a) two setae b) one seta

c) multisetose Table 9 (continued)

50. Coxal field leg II a) one seta b) two setae

c) multisetose

51. Coxa leg III a) one seta b) two setae

c) multisetose

52. Seta la coxal* field I a) setiform b) nude

c) featherlike

53. Seta lb coxal field I a) setiform b) short and thick

c) bifid

54. Seta 2a coxal fields II a) setiform b) short and thick

c) bifid

55. Seta 3b coxal field III a) setiform b) short and thick

c) bifid

56. Coxal field leg III a) rounded b) rectangular

57. Intercoxal setae a) 0-0-2 b) none

c) 0-0-3+

58. Claparede's organ a) present b) absent Table 9 (continued)

59. Anal sderite with setae a) present b) absent

60. Anus a) present b) absent

61 • Preanal s d e r i t e a) absent b) present

62. Femur leg I a) divided b) entire

63. Femur leg II a) divided b) entire

64. Femur leg III a) divided b) entire

65. Number of setae on femur leg I a) 6+ b) 6

c) 5

66. Number of setae on femur leg II a) 6t b) 6

c) 5

d) 4

67. Number of setae on femur leg III a) 6f b) 6

c) 5

d) 4

68. Dorsal seta femur legs I &II a) barbed b) nude

c) reversal Table 9 (continued)

69. Dorsal seta femur legs I &II a) normal b) anterior

70. Seta bv leg I a) present b) absent

71. Seta bv leg II a) present b) absent

72. Seta bv leg III a) present b) absent

73. Theta femur ldg I a) absent b) present

74. Theta femur leg II a) absent b) present

75. Theta femur leg III a) absent b) present

76. Number of setae on genu leg I a) 5+ b) 4

77. Number of setae on genua legs II & III a) 5+ b) 4

c) 3

d) 2

78. Number of sigma on leg I a) 2 b) 1

c) 0

d) 3+

e) multiple (4+) Table 9 (continued)

79. Number of sigma on legs II & III a) 1 b) 2

c) 3

d) 4+

e) 0

80. Microseta k on 'genu leg I a) present b) absent

81. Microseta k on genu leg II a) present b) absent

82. Number of setae on tibia leg I a) 10* b) 9

c) 8

d) 7

e) 6

f) 5

83. Number of setae on tibia leg II a) 10* b) 9

c) 7

d) 6

e) 5 220 Table 9 (continued) 00 84. Number of setae on tibia leg III a) lOf b)

c) 6

d) 7

e) 5

85. "pv" seta on tibia leg I a) two b) one

86. Phi tarsus leg III a) present b) absent

87. Companion setae with phi I a) absent b) present

88. Companion setae with omega I a) absent b) present

89. Famulus tarsus leg I a) present b) absent

90. Famulus leg I a) long b) minute

91. Position famulus leg I relative

to omega a) distal b) proximal

c) distal (reversal)

92. Position famulus leg II relative

to omega a) proximal b) distal 221

93. Dorsal eupathid tarsus leg I a) present b) absent Table 9 (continued)

94. Dorsal eupathid tarsus leg I

vith companion seta a) absent b) present

c) reversal

95. Extra ventral eupathidia a) present b) present

96. Dorsal eupathid tarsus leg II a) present b) absent

97. Dorsal eupathid tarsus leg II

with companion seta a) absent b) present

98. Subterminal eupathid tarsus leg I a) present b) absent

99. Subtermlnal eupathid tarsus leg II a) present b) absent

100. Subterminal eupathid tarsus leg III a) present b) absent

101. Scopa on tarsus leg III a) absent b) normal

c) reduced

d) absent (reversal)

e) bifurcate

102. Lophotrix on tarsus leg III a) absent b) present 222 c) absent (reversal) Table 9 (continued)

103. Form of lophotrix a) seta form b) large seta without

long branches

c) one long distal branch

d) one long basal branch

e) 1-3 long basal branches

f) multiple branches (5+)

g) long spinelike

h) long, spinelike & # bifurcate

i) pilose

j) branches fused,

tips brushlike

k) setiform (reversal)

NJro (_o Table 9 (continued)

104. Pretarsus legs I, II a) paired claws b) without paired claws

105. Pretarsus legs I, II a) clawlike empodium b) without empodium

106. Pretarsus leg III a) paired claws b) without paired daws

107. Pretarsus leg III a) clawlike empodium b) without empodium

108. Paraxial d a w leg III a) normal b) short, thick, rotated

109. Paraxial claw of leg III a) normal b) absent

110. Tarsus leg III a) normal b) modified

111. famulus leg I a) minute b) long

112. Omega leg II a) setiform b) distally expanded

113. Striation on legs a) absent b) present

114. Tip of empodium a) normal b) expanded tip

POSTLARVAL CHARACTERS

115. Accessory claw a) absent b) present

c) absent (reversal) NJ N 3 Table 9 (continued)

116. Placement of accessory claw a) dorsal b) ventral

117. Arrangement of palp tibial spines a) random b) one row

c) two rows

d) absent

118. Large ventral seines a) absent b) present

119. Brushlike sete on palp femur and genu a) absent b) present

120. Prodorsal trichobothria a) two pair b) one pair

121. Cheliceral blade a) bladelike b) styletlike

c) needlelike

122. Prodorsal s d e r i t e a) present b) with anterior extension

c) including eyes

d) wide

e) reduced or absent

123. Aliform setae legs I to IV a) absent b) present

124. Idiosomal setae 225 125. Shape of palps a) raptoral b) linnear Table 9 (continued)

126. Retractile gnathosoma a) absent b) retractile

c) chelicerae only

127. Pregenital tuberide a) absent b) presant

128. Idiosomal fingerlike structures a) absent b) present

129. Coxal field a) without reticulation b) reticulate

130. Idiosomal cuticle a) smooth or striate b) stellate

131. Dorsal idiosoma a) without special b) with circular sderites

structures 226 APPENDIX B

Table 10. List of Species Examined

The following list of species and slide label data is arranged by

superfamily and family according to the proposed new classification.

Within each family the genera and species are arranged alphabetically. Individual localities are listed chronologically, except for types which are listed first. The number of specimens examined is given, in parentheses, after the slide label data for each speices. An "L" indicates larvae and "P11 indicates postlarval instars (deutonymphs and adults). Following the number of specimens is an abbreviation (also in parentheses) indicating the source of the specimens. A key to the abbreviations is provided at the end of the list.

227 228

Table 10. List of Species Examined

Subcohort Erythraeina

SF. Erythraeoidea

F. Erythraeidae Robineau-Desvoidy, 1828

Abrolophus sp. AL-2660 Oregon, Lincoln Co., Ona Beach State Park, on barrier rocks, l-IV-82, W.C. Welbourn, (4P, 80L reared) (OSU).

Augustsonella tuberculata (Auguston, 1940) California, Marin Co., Pt. Reyes National Seashore, 21-vii-1979, W.C. Welbourn, (3P) (OSU).

Augustsonella sp. Arizona, Yuma Co., Yuma, ex Cressa truxillensis H.B.K.. vi-5-81, D.M. Tuttle, (4P) (TUTTLE).

Balaustium spp. AL-2664 California, Los Angeles Co., San Gabriel Cyn., NE of Azusa, ex oak litter, 9-IV-82, W.C. Welbourn, (8P, 2L) (OSU). AL-2955 Italy, Rome, USDA Laboratory, 12-vi-1984, W.C. Welbourn, (20P, 8L reared) (OSU).

Caeculisoma spp. AL-1114 South Africa, Natal, World’s View Hilton, ex forest leaf litter, iii-1974, B.H. Lamoral, (IP) (OSU). AL-2450 Mississippi, 24-iii-81, R.L. Brown, (2P) (OSU).

Callidosoma metzi. Sharma, Drooz, & Treat, 1983 West Virginia, Jefferson Co., 10 km E Kearneysville, VI-1983, (3L) (OSU).

Charletonia sp. on Camnula pellucida, Cederville, Calif., Sept. 3, 1952, H.H. Keifer, 52-1-51, Lot 52-9947, (1L) (OSU). AL-1411 Utah, Locomotive Spr., ex fly, 25-vii-1969, G.F. Knowlton, (1L) (OSU).

Erythraeoides serratus (Womersley, 1946) Delair, S.A. 28-9-35, Baltaurs, N.S.W., 35-5-34, (IP, IP paratype) (SAM).

Erythraeus styriacus Turk, 1981 AL-2778 Austria, Steiermark (IP, 1L) (OSU). Table 10 (continued) 229

Erythraeus sp. AL-2534 New Mexico, Eddy Co., Lincoln Nat. For., Cottonwood Lookout, ex under rocks, 31-vii-1981, W.C. Welbourn, (2P, 10L reared) (OSU).

Erythrites reginea (Hirst, 1928) Parklands Adelaide, S.A., 3-12-27. on water trough, Nat. Pk. Bera, S.A. , 18-11-49, (P holotype, 2P syntypes, 2L) (SAM).

E. urrbrae (Wormersley, 1934) on Caladendron, Burnley vie., 10-10-45, on Jassid, Delaide, 1937, (2L) (SAM).

Erythraxus sp. Colombia, S.N. de Santa Marta, San Pedro, San Javier, 1563m, iii-29-1975, J.A. Kochalka, (2P) (OSU).

Hauptmannia sp. AL-2510 Maryland, Garrett Co., 0.5 mi. S Hwy 48 on New Germany Road, ex litter around rotten log, 10 April 1981, W.C. Welbourn, (4L) (OSU). AL-2512 West Virginia, Preston Co., Coopers Rock St. Forest, 0.25 mi S Hwy 48, ex moss and bark on log, 10-iv-1981, W.C. Welbourn, (2L) (OSU).

Leptus spp. Maine, Piscataquis Co., Baxter State Park, ex beating sprice, 18-vii-1984, W.C.Welbourn, (10P, 100L reared) (OSU).

Mypongia sp. AL-2723 R.S.A., Transvaal, bot. garden of Potchefstroom Univ., ex moist litter in grass (Cynodon dactylon), 28-V-1982, D.E. Johnston and P. Theron, (19P, 10L) (OSU).

Paraphanolohus metcalfei Smiley, 1968 On Saccharosydne saccharivora West., Br. Honduras, J.R. Metcalf, Lot 67-2949, (L holotype and 1L paratype) (USNM).

Paraphanolohus sp. AL-1034 Brasil, Sao Paulo, ESALQ, Paracicaba, ex under dry dung in horse pasture, l-iii-1978, D.E. Johnston, (IP, 15L reared) (OSU).

Rainbowia celeripes (Rainbow, 1906) Heidelberg, Victoria, Oct. 1959, E. Coghill, (2P) (SAM).

Tepoztlana sp. Texas, Jack Co., 3 mi NW Jacksboro on US 281, 1050’, lO-v-1972, (IP) (OSU). Texas, Pecos Co., 23 mi SW Ft. Stockton, 4-vii-1979, (IP) (OSU). Table 10 (continued) 230

F. Smarididae Vitzthum, 1929

Calorema aztaka Southcott. 1963 Orchid plants, Maiz, SLP, Mexico at Laredo, Feb. 4, 1954, Lot54-1358, Fouts coll., (IP) (OSU).

Fessonia sp. AL-2519 Arkansas, Stone Co., 3.5 mi NE Fiftysix, ex litter, base of dead tree, 25-IV-1981, W.C. Welbourn, (IP, 18L reared) (OSU).

Smaris lanceolate Southcott, 1963 on orchid plants, Orizaba, V.C. Mex. at Laredo, Aug. 21, 1945, Lot45-15902, (IP) (OSU).

Trichosmaris sp. AL-2534 New Mexico, Eddy Co., Lincoln Nat. For., Cottonwood Lookout, ex under rocks, 31-vii-1981, W.C. Welbourn, (IP, 18L reared) (OSU).

SF. Calyptostomatoidea Feider, 1955

F. Calyptostomatidae Oudemans, 1923

Calyptostoma spp. Ohio, Hocking Co., Rock House, ex mosquito, (3L) (OSU). on cranefly, Cuernavaca, Mor. Mex., Oct. 16, 1944, N.L.H. Krauss coll., Lot45-1651, (1L) (OSU). C. velutinus (Muller, 1776) ex Limnophila nemoralis, Allagauer Alpen Rappensee, 2000m (Ml 115), 27-6-1-7-1976, (1L) (OSU). Elkwater Lake, Cypress Hills Prov. Pk. Alberta, ex wet algal mat amidst sedges by lake, 7-vii-1978, E.E. Lidquist, (1L) (CNC).

Subcohort Hydracarina

Hydrachnoidea

F. Persigiidae

Persigia limnophila Protz, 1896 R2-63, East Marsh ESGR, Mich. (7L) (OSU).

F. Limnocharidae

Limnochares americana Lundblad, 1941 Barton Pond, Washtenaw Co., Michigan, 29 May 1951, D.R. Cook, DC55-51, (1L) (OSU). Table 10 (continued) 231

Hydryphantoidea

F. Hydryphantoidae

Hydryphantes ruber (Geer, 1778) Lot 162-14, -17 Washtenau Co. Mi., Portage Lake, 14 mi north-west of Ann Arbor, vi-21-51, R.D. Mitchell, (7L reared) (OSU).

Trichothyas musicola Mitchell, 1953 II., 10 mi N.W. of Kankakee, Rock Creek Canyon, in seepage on canyon wall, vii-29-1950, Lot 144, R.D. Mitchell and H. Dybas, (5L) (OSU).

Thyas stolli Koenike, 1895 Lot 162-11, -12, -15, -16 Washtenau Co. Mi., Portage Lake, 14 mi north-west of Ann Arbor, vi-21-51, R.D. Mitchell (4L reared) (OSU). Lot 161-R1, Livingston Co., Mich. George Reserv., 2 mi W of Pinckney, vi-11-51, (8L) (OSU).

F. Thermacaridae

Thermacarus nevadensis Marshall, 1928 Oregon, Harney Co., 5 mi S White Horse Ranch, ex hot spring, Aug. 3, 1979, G.T. Baker, (2L) (OSU).

F. Hydrodromidae

Hydrodroma despiciens (Muller, 1776) Head Lake, Berry Co., Mich., 30 June 1967, R.D. Cook, 18-2, (1L) (FMNH). '

Superfaraily Tanaupodioidea Thor, 1935

F. Tanaupodidae

Eothrombium echinatum Berlese, 1910 151/12 Beragun, (lP) (BERLESE). 175/22 ex Palacia (Pila) (Musca), (IP) (BERLESE). 191/7 ex Firenze (Humus), (IP) (BERLESE). 191/8 Firenze- Siardieso R. Nov. 1917, (IP) (BERLESE). 251/23 Cetta-di-Cashello ex Musco, (IP) (BERLESE). 251/24 Zuchuro, (IP) (BERLESE). CXIV/1 No data, (IP) (BERLESE).

Eothrombium leptotarsum Berlese, 1910 251/25 (CXIIl/50) tipico ex Umbria, (IP) (BERLESE). Table 10 (continued) 232

Eothrombium lasseni (Newell, 1957) NEW COMB. 1073(HC)-01. Hat L. drainage, Lassen Park, Calif., Under rocks, etc. along stream, Aug. 5, 1955, I.M. Newell, (1L) (USNM). 1079-13. Origin of King's Cr. Lassen Park, Calif., Aug.6, 1955, I. M. Newell, (1L) (USNM). 1089-01. E. Fork Hat Cr. Lassen Park, Calif., On fly at 7250' El., Aug. 7, 1955, I.M.Newell, (1L) (USNM). 1090-13. E. Fork Hat Cr. Lassen Park, Calif., Along steam at 7250’ El., Aug. 7, 1955, I.M. Newell, (1L) (USNM).

Eothrombium nemoricola (Berlese, 1886) NEW COMB. 45/27 ex musco, (IP) (BERLESE). 49/39, /40 ex Firenze, (2P) (BERLESE). 113/24, /25, /26, /28 ex Bevagna-Umbria, (4P) (BERLESE). 113/33 ex Vallombria, (IP) (BERLESE).

Eothrombium scutelata (Newell, 1957) NEW COMB. 818-07. Spencer's Butte, Eugene, Oregon, On shrubs, July 1, 1952, I.M. Newell, (1L holotype) (USNM).

Eothrombium siculum Berlese, 1910 CXIII/48, /49 No data, (2P) (BERLESE).

Eothrombium spp. 1020-07. Salt Cr. Falls, Ore Under rocks on wet talus slope, July 12, 1954, I.M. Newell, (IP) (USNM). 1092(HC)-01. East Fork Hat Creek Lassen Park, Calif., On small fly at 7250'elev., Aug. 7, 1955, I.M. Newell, (1L) (USNM). 1459-01. Shasta Springs, Shasta Co., Calif., June 26, 1959, I.M. Newell, (1L) (USNM).

Polydiscia squamata Methlagl, 1928 AL-2933 Italy, 1 km W. Vallombrosa, sweeping vegetation, 20-V-1984, W.C. Welbourn, (1L) (OSU).

Tanaupodus passimpilosus Berlese, 1910 116/5 Tipico Bevagua-Umbria (IP) (BERLESE) 93/48 ex niti Dociostaurus moricenus Paoli- inverm 1918 Foggia, (IP) (BERLESE).

Superfamily Chyzerioidea

F. Chyzeriidae

Chyzeria hirsti Womersley, 1934 ACB 21 9.12.1938, Glen Osmond, South Australia, 5-vi-1938, R.V. Southoctt, (IP) (OSU). ACB468L1 (ACB 408L1) South Australia, 3-ix-1950, R.V. Southcott, (1L reared) (OSU). Table 10 (continued) 233

Chyzeria sp. BMOC #84-0628-2, -3 South Africa, Transvaal, Malta Forest, Trichardtsdaal, ex Libanasidus vittatus, 5-iii-1984, Otte, Cale & Tom #145, (6L) (UMMZ).

Nothotrombicula deinacridae Dumbleton, 1947 Parasitic on Giant Weta, Mt. Peel, N.Z., coll. M. Laird, 29-ii-42, (L holotype, 1L paratype) (NZ).

Pteridopus auditor Newell SVercammen-Grandjean, 1964 5128-05, -06, -07, -09, -10 Hot Springs, Sabi Valley. S. Rhodesia, on cricket, 12 Feb. 1961, I.M. Newell, (13L paratypes on 5 slides) (USNM).

Pteridopus pseudhannemania Newell &Vercammen-Grandjean, 1964 Dundo (Angola), ex forest soil in Furi, Dr. Barros Machado, 1-1948, (L holotype) (USNM).

SF. TR0MBICUL0IDEA

F. Johnstonianidae

Centrotrombidium dichotomoicoxala Vercammen- Grandjean & Cochrane, 1974 Etna, Tompkins Co., NE of Ithaca, New York, USA, polluted seepage area - cow pastaure, ex Culicoides crepuscularis, A. Cochrane coll., (12L) (OSU).

Centrotrombidium distans Newell, 1957 893PP-01, -02, -04. Tule Lake, California, Mud flat by slough, Sept. 4, 1952, I.M. Newell, (3L paratypes) (USNM). 893-ff-12. Tule Lake, Califrnia, Mud flat by slough, Sept. 4, 1952, I. M. Newell, (1L paratype) (USNM).

Centrotrombidium misellum (Berlese, 1918) 191/4, /5 tipico ex Messico - Guajauato A. Duges, (IP) (BERLESE).

Centrotrombidium spp. 1653(HC)-01, -02, -04. HC Rusk Co., Wise., (T35N, R7W, S22), On Culicoides crepuscularis Mall, in light trap, July 3, 1959, David W. Lupton, (6L) (USNM).

Charadracarus delitescens Newell, 1960 1178-29. Angeles Nat. For., L.A. Co., Calif., Engelmann Oak (sic-21B), April 7, 1957, I.M. Newell, (IP) (USNM). 1185-04, -32, -34 Angeles N.F. L. A. Co., Calif. (sic-21B), Engelman Oak Litter, June 10, 1957, I. M. Newell, (3L) (USNM). Table 10 (continued) 234

Charadracarus spp. 1431-01, -11. Hwy.79, ca.4.8 mi S. of Julian, San Diego Co., Calif., Under live oaks, Apr. 24, 1959, I.M. Newell, (2P) (USNM). 2121-28, -48. ca. 2 mi. W of Arrowhead Village, San Bernardino Mts., California, Oak and Fir litter, 29 June 1965, I.M. Newell, (2P) (USNM).

Diplothrombium cascadeuse Newell. 1957 816-01, -02. Salt Creek Falls, Oegon, Grass and flowerws, July 7, 1952, I.M. Newell (1L) (USNM). 816-04. Salt Creek Falls, Oregon, Grass and flowers, July 7, 1952, I.M. Newell, (IP paratype) (USNM).

Diplothrombium longipalpe (Berlese. 1887) 101/14, /15 tipico ex Vallombrosa, (IP) (BERLESE). 116/1 tipico ex Vallombrosa, (IP) (BERLESE). 149/48 Vallombrosa, (IP) (BERLESE). 197/18 Palacia (Pisa) ex musco, (IP) (BERLESE). 220/37 Vallombrosa ex musco 1921, (IP) (BERLESE).

Diplothrombium moldavicum Feider. 1959 Magura-Odobasti, Romania, ex Dicranota bimaculata. 19-V-1952, no. 372, Z. Feider, (1L) (FEIDER).

Diplothrombium monoense Newell, 1957 605-01, -03. Mono L., Calif., On beetle pupa, under wet stone, June 1, 1952, I.M. Newell, (2L) (USNM). 729-77, -101. Mono L. Calif., Seepage water, May 30, 1952, I.M. Newell, (2P) (USNM).

Diplothrombium spp. KEH #36, #39. Light trap, Whiple Dam, Huntingdon Co., Pa., 5 Jul 1948, D.M.& K.E. Hyland, (2L) (USNM). 736-08. Mono L., Calif., Under rocks, June .1, 1952, I.M. Newell, (1L) (USNM).

Johnstoniana latiscuta Newell 1957 1080-02. Origin of King's Creek, Lassen Park, Calif., Aug. 6, 1955, I.M. Newell, (IP paratype) (USNM). 1452-HC-01. Shasta Springs, Shasta Co., Calif., Larva attached to adults ., June 26, 1959, I.M. Newell, (1L) (USNM).

Johnstoniana maxima Feider, 1958 Dorna, Suceava, sous pierres, 18-vi-1950, Nr. 468, (1L paratype) (FEIDER).

Johnstonaiana ventripilosa Feider. 1958 pe Tipulidae, Timisuldejos, Brasov, 25-vi-1955, Nr. 601, (1L paratype) (FEIDER). Table 10 (continued) 235

F. Trombiculidae Ewing, 1944

Acomatacarus arizonensis Ewing. 1942 A.P.24595, ex Phrynosoma c. blainvilli, California, Walker Pass, 5-30-1941, (2L) (OSU).

Apolonia tigipioensis Tores & Baraga, 1938 Brasil, Pernambuco, ex quills of new feathers, chicken, May 1940, Sylvia Torres, (7L) (OSU).

Eutrombicula splendens Ewing, 1913 OSU, ex culture, (5P, 25L) (OSU).

Euschoengastia radfordi Brennan & Jones, 1954 #29444, ex Peromyscus maniculatus. Hastings Reservation, Monterey Co., April 7, 1951, (1L) (OSU). #29457, ex Peromyscus boylii. Hastings Reservation, Monterey Co., April 14, 1951, (1L) (OSU).

Hannemania hegeneri Hyland, 1956 Ex Rana pipiens, Marshall Co., Okla., E.O. Morris, (2L) (OSU). Ex Acris crepitans. Marshall Co., Okla., E.O. Morris, (1L) (OSU).

Pseudoschoengastia farneri Lipovsky, 1951 #38045, Peromyscus leucopus. Mareiga Refuge, Cameron Co., Texas, June 5, 1962, (1L) (OSU). #38194, Liomys sp., Mareiga Refuge, Cameron Co., Texas, Sept. 5, 1962, (1L) (OSU).

Sasacarus furmani (Hoffmann, 1954) RML 44401, Tylomys watsoni. Panama, C.Z. , Pina, 7 Feb. 1962, C.M. Keeman, (1L) (OSU). RML 44965, Proechimys semispinosus. Panama, C.Z. , France Field, 13 April 1962, C.M. Keeman, (1L) (OSU).

Sauracarella whartoni Lawrence, 1949 From Pseudocoroylus subriridis Herschel. Cape, N.M. 4830, (6L) (OSU).

Whartonia nudosetosa (Wharton, 1938) #45384 ex Carollia perspicillata, Taraana Caves, Trinidad, 9 Jan. 1954, Hummelinck, (1L) (OSU).

Xenodontacarus plumosus (Greenberg, 1951) 2 mi S Aetna, KS, v-6-1956, off Peromyscus. D.S. Narayan, (3L) (OSU). Table 10 (continued) 236

F. Neotrombidiidae Vercammen-Grandjean, 1973

Discotrombidium villasiense Felder, 1977 Cuba, Cueva de Colon, Las Villas, guano, 24-iv-1969, Nr. 1856d, (IP) (FEIDER).

Monunguis streblida Wharton, 1938 Trinidad, La Fontaine Cave, Marne Coco Road, ex streblid flies, 23-viii-1958, Diego Martin, (4L) (CNC). 1/2 mi W Bells, Dominica, from bat flies, Trichobium on bat Myotis nigricans. 30-iii-1966, J.K. Jones Jr., (12L) (CNC).

Neotrombidium anuroporum Lindquist & Vercammen-Grandjean, 1971 Tobago Is., Panama, ex Chalcolepidius silbermanni Chevrolet. 1 June, (2L paratypes) (CNC).

Neotrombidium bengalense Lindquist & Vercammen-Grandjean, 1971 India, West Bengal, Gonsaipur Thana Deganga, District 24, ex Batocera numitor Newm., 18 June 1967, (L holotype, 2L paratypes) (CNC).

Neotrombidium tricuspidum Borland, 1956 ex hickory log, gate 7, Duke Forest, 21 June 1947, G.W. Wharton, (1L) (OSU).

Neotrombidium spp. New Jersey, Burlington Co., nr. Atsion, ex Lichenophanes armiger. 23 July 1980, W.L. Brown et. al., (20L) (OSU). AL-2710 Kenya, Amboseli Nat. Pk., Amboseli Serena Lodge at light, ex elateridae, 20-iv-1982, D.E. Johnston, (19L) (OSU).

Traxithrombium gordoni Robaux, 1977 USPR135 California, Big Basin Redwoods State Park (Santa Cruz Co.), 10-vi-1974, P. Robaux, (1L "type") (MHNNP).

F. Trombellidae Feider, 1955

Audyana thompsoni Womersley, 1954 on Heterometrus longimanus. Ulu Bomback forest Reserve, nr. Pahang road, Selangor, Malaysia, 14 April 1948, (1L, IP) (SAM).

Durenia bakavuensis Vercammen-Grandjean, 1955 Bakavu, Congo Belgan, ex Culex pipiens, (5L) (OSU).

Nothrothrombidium otiorum (Berlese, 1902) Babadag, 28-V-1958, foret, Nr. 595a (1L) (FEIDER). Table 10 (continued) 237

Ralphaudvana amamiensis Vercammen-Grandjean. N. Kumada, I.M. Newell, P. Robaux & H. Suzuki, 1974 AL-2514 Japan, Shikoku, Ichizuchi Mts. Nat. Pk., ex Tachycines robustus Ander, from mixed warm temperate forest, 18-25 August 1980, S. Peck, (45L) (OSU).

Trombella glandulosa Berlese, 1887 45/43 tipico Owina Sottern, (IP) (BERLEE). 123/13 tipico, (only leg I and palpus) (BERLESE).

Trombella nothroides Berlese, 1888 8/48 Matto Grosso tipico, (IP) (BERLESE). 34/40 tipico Matto Grosso, (2P) (BERLESE). 45/42 tipico Paraguay (label illegible), (IP) (BERLESE).

Trombella. sp. Nigeria, Western Prov., Univ. Coll. Ibaden, ex Gryllotalpa debilis Gerstaeker, BM1965-474, 17-xii-1960, (1L) (BMNH).

Typhlothrombium hystricinum Berlese. 1910 31/16 tipico (label illegible) (only palp on slide) (BERLESE). 109/50 (label illegible) (only palp and leg I on slide) (BERLESE).

Womersleyia minuta Radford. 1950 Maldive Is., gran Is., Addu Attol, 20 December 1944, (2L) (USNM, SAM).

SF. TR0MBIDI0IDEA

F. Podothrombiidae Thor, 1935

Podothrombium montanum Berlese, 1910 Vallombrosa, (IP) (BERLESE).

Podothrombium piriformis Robaux & Schiess, 1982 Mont La Schera (pare National Suisse), 2500m, 1978, (2L) (MNHNP).

Podothrombium shellhammeri Robaux. 1977 Calif., Santa Cruz Co., Big Basin Redwoods St. Park, 1974, (1L) (MNHNP).

Podothrombium verecundum Berlese, 1910 115/15, /16 Columbia, (2P) (BERLESE).

Podothrombium spp. Canada, Ontario, feeding on aphid, Cinera lasiocampa on Aibes balsuma, (6L) (OSU). Table 10 (continued) 238

P.Q.: Quebec, June 1975, J. McNiel, ex aphids, (6L) (CNC). AL-1625 Utah, Ephrim Cyn., ex fir detritus, 10,000' elev., 15 Aug. 1975, G.F. Knowlton, (IP, 2L) (OSU).

F. Trombidiidae Leach, 1815

Allothrombium brevitarsum (Berlese, 1888) 8/43 tipico Matto Grosso, (IP) (BERLESE). 34/41 tipico Rio Apa, (IP) (BERLESE).

Allothrombium canaanense Feider. 1977 Ex Perionosthenus galericulatus, Jerusalem, 1-5-1967, (1L) (HUMS).

Allothrombium crassiconum Berlese, 1910 111/41, /42, /43, /44 tipico Sta. Cruz, (2P) (BERLESE). 170/38, /39, /40, /41 tipico La Plata (Bruch), (3P) (BERLESE).

Allothrombium fuliginosum Berlese, 1903 numerous postlarval specimens were studied in the BERLESE, MNHNP and BMNH collections. Firenze, Italy, 2-X-1962, (15P) (OSU). Italy, 2km E Tosi (nr Vallombrosa), ex leaf litter, 20-V-1984, W.C. Welbourn, cultures 1984-26, -27, -28, (5P, 35L reared) (OSU).

Allolthrombium lerouxi Moss, 1960 Ottawa, Ont., ex culture, (8L) (ANSP). under apple trees, Cistereien Father's Apple orchard, Rougemont, P.Q., 21 April 1960, W.W. Moss, (P holotype, 2P paratypes) (CNC).

Allothrombium recki Feider. 1967 sur Takecales taiwaneanus, 30-V-1965, Nr. 1845, (1L paratype) (FEIDER).

Allolthrombium (Aphithrombium) mali Childers and Vercammen-Grandjean, 1980 NEW COMB. Horticulture farm, New Franklin, Howard Co., Mo., 7 June 1973, (1L paratype) (OSU).

Allothrombium spp. Tenn., Lake Co., on aphids in cotton, Lot53-5624, 19 May 1953, A.P. Morris coll., No. 6, (3L) (USNM). USSR Krasnada, ex aphid in alfalfa, 1 June 1965, 65-17813, (3L) (USNM). AL-2508 Arkansas Stone Co., ca 1 mi ENE Fiftysix, at small spring nr Mirror Lake, ex tree, 27-iii-1981, (IP, 20L) (OSU). Table 10 (continued) 239

Dinothrombium tinctorum L.. 1767 5039C Nyanyadai, Sabi V., S. Rhodesia, 16 Oct. 1960 + 23 Nov. 1960, I.M. Newell, (50L reared) (USNM).

I), magnificum (Le Conte, 1852) =D. superbum (Banks, 1910) NEW SYN0N0MY Texas, (3P) (MCZ). Tucson, Arizona, (3P) (MCZ). Arizona, (10P) (MCZ). 1764 San Simion Valley, Arizona - New Mexico, 10 July 1963, I.M. Newell, (31L reared) (USNM). Arizona, Cochise Co., ex grasshoper, 3-viii-1981, W.C. Welbourn, (1L) ()SU).

I), pandorae Newell and Tevis, 1960 1263-01 to -16 Rancho Mirage, Riv. Co., Calif., in sand, June 2, 1958, I.M. Newell, (30L on 16 slides "type series") (USNM).

I). oparbellae Andre, 1949 (=Isothrombium Andre. 1949) NEW SYN0N0MY Type, Register: no. 513, (1L "type") (MNHNP).

Nanothrombidium nannelli NEW GENUS AND NEW SPECIES Italy, 16km NW Rome, ex sweeping annual vegetation along Via Casia, 13-vi-1984, W.C. Welbourn & S.C. Clement, (L holotype and 1L paratype) (OSU).

Oligothrombium mulleni NEW GENUS AND NEW SPEICES Al., Auburn, 19 May 1979, D. Reed, ex moss, DJR 79-0510-2d, (L holotypw) (AU). Tenn., Shelby Co., Maeeman-Shelby Forest State Park, 7-vi-1976, E.R. Hobek, BM0C 76-0804-5, ex hardwood forest, (1L paratype) (CU).

Paratrombium bidactylus Newell, 1958 1036B-11 Sherwood Cr., Mason Co., Wash., sandy mud in grass, July 27, 1954, I.M. Newell, (P holotype) (USNM). 1036B-02, -03, -07, -10, -19 Sherwood Cr., Mason Co., Wash., sandy mud in grass, July 27, 1954, I.M. Newell, (14L reared paratypes on 6 slides) (USNM).

Parathrombium quadriseta Newell, 1958 9196-10 Riverside Calif., (Camp Evans), muddy bank of stream, Apr. 23, 1955, (P holotype) (USNM). 9196-01, -02, -05 Riverside Calif., (Camp Evans), muddy bank of stream, Apr. 23, 1955, (12L reared paratypes on 3 slides) (USNM).

Parathrombium megalochirum (Berlese, 1910) larva, eleve, July 1965, P. Robaux, (5L) (MNHNP). Table 10 (continued) 240

Trombidium auroraense Vercammen-Grandjean, van Driesche and Gyrisco, 1977 N.Y. Tompkins Co., Texas Hollow. 29 July 1972, L.L. Peckuman, ex. lampyrid adult, GRM 72-730-A, (1) (OSU).

AL-2918 Michigan Dickenson Co., T43N, R3W, S along border of sec. 7., mixed hardwood forest, May 1984, N. Sefera, (3P, 25L reared) (OSU).

Trombidium brevimanum (Berlese, 1910) 110/11 Norvegia strand!, (IP) (BERLESE). 110/16 Norvegia Leangerund (strand), (IP) (BERLESE). 110/35, /36 Kuhaarodeut (Germania), (2P) (BERLESE).

Trombidium cancela Robaux, 1967 P. Robaux, (no data) (IP) (MNHNP).

Trombidium dacicum Feider, 1948 Odorhei, Romania, ex elevage, Nr. 259-a, Z. Feider, (1L) (FEIDER).

Trombidium heterotrichum (Berlese, 1910) 110/30, /31, /32, /33, /34 Norvegia tipico, (5P) (BERLESE).

Trombidium holosericeum (L.. 1785) numerous postlarval specimens were studied in the BERLESE, MNHNP and BMNH collections. AL-2924 G.B. Wales, Gwent nr. Abergavenny, Querqus - Fagus woodland, 750", ex under rocks, 22-iv-1984, culture 1984-8, (IP, 63L reared) (OSU). G.B. Wales, Powys nr Crickhowell, elev. 800-1000', under rocks along wall, 22-iv-1984, culture 1984-10, (3P, 30L reared).

Trombidium hyperi Vercammen-Grandjean. van Driesche and Gyrisco, 1977 N.Y., Cayauga Co., alfalfa field nr Aurora, ex Hypera postica, (L holotype) (USNM). Ohio, ex Hypera postica. orignal numbers M20179, M21279, M19779, M19479, (8P reared, 65L) (OSU).

Trombidium mediterraneum (Berlese, 1010) 110/22, /23 tipico Corfu - Thor, (2P) (BERLESE).

Trombidium rhopalicus Vercammen-Grandjean & Popp 1967 NEW COMB. Ottershausen/Oberbayern, Juli 1966, Rhopalicus tutela Walker, leg Wichmann, (P1254a), (L holotype) (ZS).

Trombidium susteri (Feider, 1955) NEW COMB. Feliceni, Harghita, viii-1959, sur Phyllotreta atra Nr. 557, (2L) (FEIDER). Table 10 (continued) 241

Trombidium tenuiclavatum (Berlese. 1910) 166/7 Allando & Jeannel Africa Oriental tipico, (IP) (BERLESE).

T. teres (Andre, 1929) NEW COMB, label illegible, (1L) (MNHNP).

F. Neothrombiidae Feider, 1959

Acridothrombium chiricahuae NEW GENUS AND NEW SPECIES Arizona, Cochise Co., 2.5 km ESE Portal, ex under a rock, 3-viii-1981; W.C. Welbourn, culture 247, (L holotype and 12L paratypes reared from eggs deposited by female) (OSU). Arizona, Cochise Co., 2.5 km ESE Portal, 3-viii-1981; W.C. Welbourn, cultures 248-254,255 and 257, (11L paratypes reared from eggs deposited by several females) (OSU). WCW810803-15 Arizona, Cochise Co., 2.5 km ESE Portal, 3-viii-1981; W.C. Welbourn, ex from neck and coxae of Xanthippus c. pantherinus, (6L paratypes) (OSU). WCW810803-15 Arizona, Cochise Co., 2.5 km ESE Portal, 3-viii-1981; W.C. Welbourn, ex five specimens of Trimerotropis j). pallidipennis, (23L paratypes) (OSU). WCW810803-15 Arizona, Cochise Co., 2.5 km ESE Portal, 3-viii-1981; W.C. Welbourn, ex two juvenile Brchystola magna, (5L, paratypes) (OSU). WCW810803-15 Arizona, Cochise Co., 2.5 km ESE Portal, 3-viii-1981; W.C. Welbourn, ex residue in vials with above mentioned hosts, (18L paratypes) (OSU). Arizona, Cochise Co., 25 km S junction 1-40 and Potal Road, between San Simion and Portal, ex T. £. pallidipennus, 3-viii-1981, W.C. Welbourn, (5L) (OSU). Arizona, Cochise Co., 5 km E Portal, ex grasshoppers, 12-vi-1958, W.J. and J. W. Gertsch, (2L) (USNM). Arizona, Gila Co., 44 km N Globe, 3-viii-1981, ex P^. aztecus, (3L) (OSU).

Adelothrombidium africana NEW GENUS AND NEW SPECIES BMOC #84-0628-2 South Africa, Transvaal, Malta Forest, Trichardtsdaal, ex Libanasidus vittatus, 5-iii-1984, Otte, Cale & Tom #145, (L holotype and 10L paratypes) (UMMZ).

Aegialothrombidium mexicana NEW GENUS AND NEW SPECIES Mexico, Baja California Sur, Puerticetos, ex under elytra of Phaleria pilifera Le Conte; L.E. Watrous, (L holotype and 2L paratypes) (OSU).

Aegialothrombidium sp. Guatemala, Champerico, under elytra of Phaleria quatemalensis Champion, (1L) (BMNH). Table 10 (continued) 242

Aethethrombidium brazilensis NEW GENUS AND NEW SPECIES AL-1070 Brasil, Amazonas, Manaus; ex miscellaneous litter INPA; 31-111-1978; D. E. Johnston; original number 31-111-78, (L holotype) (OSU).

Ceuthothrombium cavaticum Robaux. Webb, & Campbell, 1976 New Mexico, Eddy Co., Sentinal Cave, ix-1972, C. sp., W.C.Welbourn, (3L) (OSU). New Mexico, Eddy Co., Carlsbad Caverns National Park, Lake Cave, ex £. sp., 27-ix-1973, W.C. Welbourn, (2L) (OSU). New Mexico, Lincoln Co., Lobo Cave, 18-xi-1973, ex C_. sp., W.C. Welbourn, (17L) (OSU). New Mexico, Sandoval Co., Ojo Del Diablo, 25-viii-1973, ex _C. sp., W.C. Welbourn, (1L) (OSU). New Mexico, Eddy Co., Jurnigan Cave, 26-iv-1975, C. sp., W.C. Welbourn, (12L) (OSU). Arizona, Coconino Co., Wupatki National Monument,, Dangling Flake Crack, ex C. sp., 31-i-1976, W.C. Welbourn, (2L) (OSU). Texas, Sutton Co.,Harrison Cave, 12-iv-1976, ex C. sp., W.R. Elliott and J. Reddell, (1L) (OSU). New Mexico, Eddy Co., Carlsbad Caverns National Park, Spider Cave, ex CL sp., 27-vii-1981, W.C. Welbourn, (2L) (OSU). New Mexico, Eddy Co., Carlsbad Caverns National Park, ex C. sp., 27-ix-1981, R. Kerbo, (2L) (OSU). Texas, Culberson Co., Border Cave, 27-vii-1981, ex sp., W.C. Welbourn, (2L) (OSU). Arizona, Cochise Co., Happy Jack Cave, 4-X-1981, ex Ceuthophilus paucispinosus, B.& D. Buecher, (1L) (OSU). Arizona, Gila Co., Crinoid Cave, 18-X-1981, Ceuthophilus pima, B. & D. Buecher, (4L) (OSU).

Kermathrombidium panamensis NEW GENUS AND NEW SPECIES Panama, C.Z., Frijoles, 19 May 1980, ex nest of Nasutitermes coriger, B. Thorne, (L holotype, 7L paratypes) (OSU).

Megalothrombium jamaicens NEW GENUS AND NEW SPECIES Jamaica; St. Thomas Par., Corn Pass Gap; ll-v-1950; H. B. Mills coll.; 111. Nat. Hist. Survey Acc. No. 49627, (L holotype) (KU).

Neothrombium neglectum (Bryant, 1909) Algeria, Biskra, ex Gryllotaipa gryllotalpa L. BM1922-422, (4L) (BMNH). Austria, ex Gryllotalpa gryllotalpa L. BM1964-638, (6L) (BMNH). Berthat Basrah, 23-xii-1978-105), ex Gryllotalpa unispina Saussure, BM1948-506, (11L) (BMNH). Cyprus, Larnak, 2-vii-1931, ex Gryllotalpa gryllotalpa L., (1L) (BMNH). Egypt ex Gryllotalpa gryllotalpa L.. (1L) (BMNH). Table 10 (continued) 243

Iran ex Gryllotalpa gryllotalpa L. BM93-199, (2L) (BMNH). Iran, 3-iii-1920, ex Gryllotalpa gryllotalpa L. BM no. 1957, (2L) (BMNH). Iraq, Bagdad, summer 1923, ex Gryllotalpa gryllotalpa L. BM1923-569, (3L) (BMNH). Italy, Rome, ex Gryllotalpa gryllotalpa L. BM1927-68, (2L) (BMNH). Kuait, iv-1937, ex Gryllotalpa gryllotalpa L., (2L) (BMNH). Palestine, 2-X-1926, ex Gryllotalpa gryllotalpa L. BM1927-346, (1L) (BMNH). Switzerland, Engadine, ex Gryllotalpa gryllotalpa L. BM1924-199, (4L) (BMNH). Transcaspia Buiram Ali, 17-V-1921, ex Gryllotalpa unispina Saussure, BM1925-540, (1L) (BMNH). Turkey ex Gryllotalpa gryllotalpa L., (1L) (BMNH). Upper Egypt, Meir District (N of Assiant), 21-111-1912, ex Gryllotalpa gryllotalpa L. BM1921-269, (1L) (BMNH).

F. Calothrombiidae

Calothrombium paolii (Berlese, 1918) 190/44 tipico ex nidi Deciostarus 1917 - Paoli Foggia - sotterra, (IP) (BERLESE). 190/45 juv. ex nidi Deciostarus maroccanuus Foggia 1918, (IP) (BERLESE). 190/46 cotipico pasti ex Foggia - Deciostarus ooteche - sotterra, (IP) (BERLESE). 190/47 cotipi - pasti ex ooteche di Deciostarus maroccanuus Foggia Paoli, (IP) (BERLESE).

Dectothrombidium marshalli NEW GENUS AND NEW SPECIES "Spaino 1876", ex Decticus albifrons F., (L holotype) (BMNH)

F. Eutrombidiidae Thor, 1935

Atopothrombidium townsendi (Dumbleton, 1962) NEW COMB. New Zealand, Unites Creek, Roding River, ex cave weta (Orthoptera, Rhaphidophoridae) in limestone cave, 6-viii-1960, J.I. Townsend, (L holotype, 2L paratypes) (NZ). New Zealand, Dawson’s Falls, ex Macropathus, M. Buchler, i-1955, (3L) (NZ).

Cryptpothrombidium howarthi NEW GENUS AND NEW SPECIES Tennessee, Warren Co., McMinnville, Higganbottom Entrance to Cumberland Caverns, deep zone, ex Hadenoecus (Orthoptera, Rhapidophoridae) 24-vii-1981, F. G. Howarth, (L holotype and 2L paratypes) (BPBM). Tennennessee, Grundy Co., Montegle, 17-iii-1931, J. M. Valentine, ex Hadenoecus subterraneus (Orthoptera, Rhapidophoridae), (2L) (USNM). Table 10 (continued) 244

Eutrombidium feldmanmusae Feider, 1977 Israel, Jerusalem ex Rivetina betica, 15-3-67, (1L) (HUMS).

Eutrombidium trigonium (Hermann, 1804) (label illegible) (4L on two slides) (0C).

Eutrombidium sp. Texas, Dallas Co., Dallas, ex cricket, H. P. Wood, 25-viii-1919, (12L) (USNM). Ohio, Ross Co., Chillicothe, ex eggs, A.E. Miller, 26-vi-1923, (5L) (OSU). Nicaragua, El Salto Palacaquina, ex Sch. parvanensis, F. A. estrada, 23-X-1958, Lot 2-21642, (10L) (USNM). Utah, Locomotive Springs, ex grasshopper, G.F. Knowlton, 25-viii-1969, (1L) (OSU). Kansas, Johnson Co., Overland Park, ex grasshopper, 13-ix-1984, (1L) (OSU).

Hoplothrombium cicindelae (Floch and Abonnenc, 1942) NEW COMB. French Guiana, Cayenne, ex Cicindela cayennese F., (type series on one slide with no holotype designated), (16L) (MNHNP). Brasil, Pirelli Rubber Plantation (Iribaca), just off Belem-Castanhal Hwy nr. Marituba, P. F. Darlington and W. L. Brown, viii-1962, ex leaf litter, humus and rotten wood from virgin rain forest, (1L) (OSU). Equador, Napo Pro., Laguna Taracoa, South side rio Napo, 240’, 25-vi-1980, J. P. & K. E. Donahue, ex Cincindellidae, (4L) (OSU). Equador, Napo Prov., Primarera, North bank Rio Napo; O'25" S, 76'46" W, 26-vi-1980, J. P. & K. E. Donahue, ex cicindellidae, (3L) (OSU).

Hoplothrombium quinquescutatum Ewing, 1925 ex beetle mite taken from stomach of a toad (Bufo americanus) collected at Hudson Bay (US Geol. Survey No.585), USNM No. 893, (L holotype) (USNM).

Hoplothrombium sp. Oregon, Salt Creek Falls, under rocks on wet tallus slope, I. M. Newell, 12-vii-1954, (1L) (USNM). Zimbabwe, Matopos Hills, marshy ground near stream, I. M. Newell, 28-iv-1961, (1L) (USNM). Ontario, Thunder Bay District, associated with Formica oveas, 21-V-1970, F. E. Elsey, (1L) (CNC). Mexico, Tepostlan, M. A. Bastida, l-x-1974, (1L) (OSU).

Milliotrombidium japanensis NEW SPECIES AL-2514 Japen, Shikoku, Ichizuchi Mountains National Park, ex Tachycines robustus Ander from mixed warm temperate forest, 18-25-viii-1980, S. Peck coll., original number WCW810416-12, (L holotype and 37L paratypes) (OSU). Table 10 (continued) 245

Scutodesmus araneivarum NEW GENUS AND NEW SPECIES AL-1114 South Africa, Natal, World’s View Hilton, ex forest leaf litter, iii-1974, B.H. Lamoral, (L holotype) (OSU),

F. Microtrorabidiidae Thor, 1935

Achelothrombidium maculatum (Michener 1946) NEW COMB. ex. Mansonsonia sp., (1L) (AMNH). Juan Mina, C.Z. Panama, VI-10-45, (3P) (AMNH). Juan Mina, C.Z. Panama, hatched from eggs of M. maculatum. VI-19 to VI-23-45, (5L) (AMNH).

Achelothrombium pistiae (Michener, 1946) NEW COMB. Juan Mina, C.Z. Panama, VII-2-45 on Pistia stratiotes, (4P) (AMNH). Juan Mina, C.Z. Panama, June 18,1945. hatched from eggs laid on Pistia plants, (10L) (AMNH).

Atractothrombium danubialis (Feider, 1975) (=Atractothrombodies Feider, 1975) NEW SYN0N0MY Dubova, Nehedint Fruzar, lO-v-1968, Nr. 1102, larva, holotypus, (L holotype) (FEIDER).

Atractothrombium dictyostracum Vercammen-Grandjean & Cochrane, 1974 Treman State Park, Tompkins Co., New York, U.S.A., wooded area, stream, A. Cochrane, ex Culicoides stellifer, (8L paratypes) (OSU).

Atractothrombium fusicomum (Berlese, 1910) NEW COMB. 102/37, /38, /39, /40 Tipico Prussia Kuhlgatr!, (4P) (BERLESE).

Atractothrombium heterocomum (Berlese, 1918) NEW COMB. 189/2, /3, /4 Messico - Guanajuato A. Duges Tipico, (2P) (BERLESE).

Atractothrombium littorale Michener. 1946 NEW COMB. Old Panama, ex female: 27-xi-45, (10L) (AMNH) Old Panama, xi-27-45, intertidal, (7P) (AMNH).

Atractothrombium sylvaticum (Koch, 1835) NEW COMB. 110/45, /46, /47, /48, /49, /50 ex Norvegia (Stand), (5P) (BERLESE).

Atractothrombium trispinum (Berlese, 1910) NEW COMB. 110/43, /44 Tipico Ambrugo (Strand), (2P) (BERLESE). Table 10 (continued) 246

Atractothrombium spp. AL-2506 Arkansas, Stone Co., ca 1 mi ENE Fiftysix small spr. nr Mirror Lake, ex moss, 27-iii-1982, W.C. Welbourn, (2P, 38L reared) (OSU). AL-2656 Oregon, Benton Co., N side of Mary's Pk. 3.3 mi S US Hwy 20, Woods Cr. Rd., ex moss on rocks in stream, 30-iii-1982, W.C. Welbourn & D.E. Johnston, (3P, 21L reared) (OSU).

Campylothrombium boreale (Berlese, 1910) 111/8, /9, /10 tipico Amburgo (Strand), (3P) (BERLESE). 111/11 Ambrurgo (Strand), (IP) (BERLESE). 111/12, /13, /14 Hamburg (Strand), (3P) (BRELESE).

Campylothrombium densipapillum (Berlese. 1910) 111/6, /I tipico (label illegible) (2P) (BERLESE). 111/15, 16 Portici erbe, (2P) (BERLESE).

Campylothrombium langhofferi Karusse. 1916 France, Fond de Torel, v-1968, P. Robaux. (IP, 3L reared) (OSU).

Cercothrombium curtulum (Berlese, 1910) NEW COMB. 131/14, /15 tipico Samarang Java, (IP) (BERLESE).

Cercothrombium diversum (Berlese, 1910) NEW COMB. 111/4, /15 Nouviea (Strand!), (2P) (BERLESE). 223-6 N. Caledonia, (IP) (BERLESE).

Cercothrombium hillyardi NEW SPECIES Great Britain, Wales, Powys, Glisfer nr. Llagynide, elev. 420 m, 22-iv-1984, under rock in mixed woodland, W.C.Welbourn, A. Baker and P. Hillyard, (L holotype, 15L paratypes reared, 3P) (OSU).

Cercothrombium pexatum (Koch, 1837) NEW COMB. 111/1, /2, /3 tipico Norvegia, (3P) (BERLESE).

Cercothrombium rasum (Berlese, 1910) NEW COMB. 102/41 tipico Prussai, (IP) (BERLESE).

Cercothrombium sanguineum (Koch, 1837) NEW COMB. 45/36 tipico Vallombrosa, (IP) (BERLESE). 111/17 Marburg, Germania (Strand), (IP) (BERLESE). 135/43 ex Sardegna Krausse, (IP) (BERLESE).

Compsothrombium tegularum Andre, 1957 1493-6, Reg. no. 861, (3P speicmens on 6 slides) (MNHNP). 60.07.02.02, Reg no. 805 J.P. Adam, (2P) (MNHNP).

Dendrotrombidium perligerum (Berlese, 1888) 21/47 tipico Paraguay Rotto, (IP) (BERLESE). 124/26 tipico Sud Americ, (IP) (BERLESE). Table 10 (continued) 247

Diathrombium diaphnae Moser and Vercammen-Grandjean, - 1979 Parasite Dend. frontais, coll fontalure sample #1-72, Rapides Parish, La., 8 May 1975, trap 8, #17,264, (L holotype) (USNM). Louisiana, Rapides Parish, Pineville, ex water frontalure baited bucket trap, 8-vi-1979 (#228039); 10-vi-1979 (#22529); 18-vi-1979 (#22768, 22779, 22793), (5L) (OSU).

Echinothrombium spinosum (Canestrini, 1885) (=0xithrombium Feider, 1952) NEW SYN0N0MY 111/18, /19, /20 Firenze, (3P) (BERLESE).

Enemothrombium bifoliosum (Canestrini, 1884) 162/38, /39 Falterous, (2P) (BERLESE). CXIII/45 no data, (IP) (BERLESE).

Ettmulleria sucidum Oudemans, 1910 "Ettmulleria sucidum Tragdh. larva”, from Tragardh collection, Sweden, (label illegible) (2L on one slide) (NR).

Eutrichothrombium eutrichum (Berlese, 1912) 37/15, /20 tipico, (label illegible) (2P) (BERLESE). 110/37 tipico, (label illegible) (IP) (BERLESE).

Gonothrombium bimaculatum Feider. 1955 Odorheium - secuiese, Harghita, 9-vii-1938, nr. 193, paratype, (1L paratype) (FEIDER).

Gonothrombium oudemansianum Feider, 1955 Feliceni, Harghita, 22-vii-1953, eleve, Nr. 555, paratype, (1L paratype) (FEIDER).

Gonothrombium bequaerti (Fuller and Wharton, 1951) Gebel Ahmed Aga, White Nile, Sudan, Egypt, 20-3-01, I. Tragard, (1L holotype) (NR).

Laminothrombium plicifrons (Berlese, 1917) 162/48 cotipia La Plata Bruch, (IP) (BERLESE).

Lithothrombidium perscabrum (Banks, 1916) NEW COMB. P. Canyon, Jan. 1913, M. Shaw, type Bks, (IP holotype) (MCZ). AL-2663 California, Los Angeles Co., San Gabriel Cyn., NE of Azusa, ex under rocks with ants, 9-iv-982, W.C. Welbourn, culture 367, (2P, 22L reared) (OSU).

Manriquia bequaerti Boshell and Kerr, 1942 Bolivia, San Rafail, ex humus, USNM cotype #53013, (2P) (USNM). Colombia, Muzo Boyeas, 5-xi-1937, "hatched from eggs” USNM cotype #53013 (1L) (USNM). Old Panama, eggs laid before V-12-45, hatched V-30-45, Michener, (5L) (AMNH). Table 10 (continued) 248

Manriquia boshelli Michener, 1946 Santa Rosa, Colon Prov., Panama, ix-10-45, C.D. Michener, (5P paratype) (AMNH).

Manriquia kethleyi (Robaux, 1977) NEW COMB. FM(HD) #72-148 Colombia, Meta, 23 km NW Villavicencio, Quebrada, Susumako, 1,000 m elev., 5-iii-1972, S. & J. Peck coll., (L holotype) (FMNH).

Manriquia panamensis Michener, 1946 Santa Rosa Colon. Prov., Panama, 1-25-46, Michener, (2P) (AMNH). Old Panama. VI-2-45, reared from jar of adult M. panamensis, C.D. Michener, (8L) (AMNH).

Manriquia sp. AL-408 Brasil, Amazonas, 20-50 km from Manaus, ex miscellaneous material, ix-1962, W. L. Brown, (1L) (OSU). AL-1070 Brasil, Amazonas, Manaus, ex miscellaneous letter at INPA, 31-iii-1978, D.E. Johnston, (3L) (OSU).

Microtrombidium italicuin Berlese, 1912 109-21, /22 tipico Bevagna-Umbria, (2P) (BERLESE). 109/23, /25 Corfu - Thor, (2P) (BERLESE). 109/28, /29, /30, /31 Palermo, (4P) (BERLESE). Italy, Vallomborsa, ex litter, May 1984, (IP, 3L reared) (OSU).

Microtrombidium gracile (Mullen & Vervcaramen-Grandjean, 1978) =Megophthrombium Mullen & Vervcamraen-Grandjean, 1978 NEW SYN0N0MY Parasite head of Dend. frontalis, frontalure sample #1-88, Rapides Parish, LA., 3 July 1975, trap 8, (L holotype) (USNM). ex.water frontalure baited bucket trap 2a (ground level) 7 mi. N.E. Pineville, 8 June 1979, 22865, 22862, 22871, (3L) (OSU). ex water frontalure baited bucket trap 2a (ground level) 7 mi. NE Pineville, La., 28 June 1979, 22862, (1L) (AU).

Microtrombidium jabanicum Berlese, 1910 37/24 tipico, (label illegible) (IP) (BERLESE). 109/3, /4 tipico, (label illegible) (2P) (BERLESE). 124/22 tipico, (label illegible) (IP) (BERLESE).

Microtrombidium columbianum Berlese, 1910 110/5, /6 tipica Columbia, (2P) (BERLESE).

Microtrombidium leptochirum Berlese, 1912 109/18, /19, /20 tipico Sta. Cruz, (label illegible) (3L) (BERLESE). Table 10 (continued) 249

Microtrombidium spp. AL-2474, -2476, -2477, -2478, -2482, -2490, -2495. Arkansas, Stone Co., 3.5 mi NE Fifty-six, ex litter at base of dead tree, 28-IV-1980. W.C. Welbourn, (8P, 15L reared) (OSU).

AL-2486. Ohio, Hocking Co., OSU. Barnaby Center, ex tree hole, culture #110, lO-v-1980, D.E. Jonston, (IP, 13L reared) (OSU).

Platytrombidium vagabundum (Berlese, 1884) NEW COMB. 45/33 (label illegible) (IP) (BERLESE). 45/34 Firenze, (3P) (BERLESE). 110/2 Veneto, (2P) (BERLESE). 111/23, /24 tipico Nola, (2P) (BERLESE). 123/43 Mt. Pellegrino 1911 Palermo, (IP) (BERLESE).

Platytrombidium fasciatum (Koch,1836) NEW COMB. larves, elevage, 08-09-1966, Longhjuneau 91, (3L reared) (MNHNP). SP 452(i), Franz, (IP) (MNHNP).

Pygothrombium bakeri iN'ew Species Papua New Guinea, Morobe Province, Buso Forest, ex malaise trap, September 1979, J. H. Martin, original numbr 2559,(L holotype and 2L partypes) (BMNH).

Reticulatrombidium serratum Shiba, 1976 AL-1869 Malaysia, Pahang, Betang, berlese of rotten log and fungus, 28-iv-1977, L,E. Watrous, (3L) OSU AL-1971 Malaysia, Pahang, 19 miles NE Juala Lamput, berlese of debris along stream, 29-iv-1977, L.E. Watrous, (2L) OSU.

Trichotrombidium muscarum (Riley, 1878) (=Phlogothrombium Feider, 1952) NEW SYN0N0MY Jerusalem, 15-ix-63, A. Barkia, (7L) (OSU). Banglore, ex Musca domestica. Oct. 76, (25L) (OSU). AL-1031 Brasil, S.P., ESAL0 Piracicaba, horse stable, 26-ii-78, ex Musca domestica, (6L) (OSU). AL-1032 Brasil, S.P., ESAL0 Piracicaba, horse stable, 26-ii-78, (4P) (OSU).

Valgothrombium confusum (Berlese, 1910) NEW COMBINATION 162/35, /36, /37 Vallombrosa, (3P) (BERLESE). 166/4 Vallombrosa, (IP) (BERLESE).

Valgothrombium fluminis (Michener, 1946) NEW COMB. Tocumen, Panama Prov., vi-1-45 at margin of drying stream, C.D. Michener, (8P paratypes) (AMNH). Tocumen, Panama Prov., iii-25-45, among very moist dead leaves at edge of water in intermittent stream, C.D. Michener, (IP paratype) (AMNH). Table 10 (continued) 250

Tocumen, Panama Prov., ex female: iii-25-45, C.D, Michener, (7L) (AMNH).

Valgothrombium homocomum (Berlese, 1918) NEW COMB. 189/5, /6 tipico Messico Guanajnato A. Duges, (IP) (BERLESE).

Valgothrombium valgum (George, 1909) Type, Scotland, 1940.3.12.13, (P holotype) (BMNH).

Valagothrombium (Parafeiderium) culicoidium (Vercammen-Grandjean & Cochrane, 1974) NEW COMB. Malloryville, Tompkins County, New York, U.S.A., sphagnum bog, A. Cochrane, ex Culicoides furensoides, (8L paratypes) (OSU).

Valgothrombium spp. Maryland, Prince Georges Co., Patuxent Wlf. Res. Ctr., 17 May 1976, Wm.L. Grogan, Jr., ex. Isohelea sp. (Ceretopogonidae), GRM-78-0307-la, (1L) (AU). Maryland, Prince Georges Co., Patuxent Wlf. Res. Ctr., June 1976, W.L. Grogan, Jr., ex. Culicoides sp. (Ceratopogonidae), GRM-77-0702-13a, (1L) (AU). Maryland, Prince Georges Co., Patuxent Wlf. Res. Ctr., June 1976, W.L. Grogan, Jr., ex. Echinohelea lanei (Ceratopogonidae), GRM-77-0702-4a, (1L) (AU). Maryland, Prince George Co., Patuxent Wlf. Res. Ctr., July 1976, W.L. Grogan, Jr., ex. Bezzia flavitarsus (Ceratopogonidae), GRM-77-0702-1, (1L) (AU). Maryland, Prince Georges Co., Patuxent Res. Ctr., July 1976, W.L. Grogan, Jr. ex. Dasyhelea grisea (Ceratopogonidae), GRM-77-0702-9, (1L) (AU). AL: Lee Co., Chewacla St. Pk. Auburn, 5 Oct. 76, J.I. Glick, ex. Africhopogon, CSP-76-1005-Alla, (1L) (AU). AL: Lee Co., Auburn, Chewacla St. Pk., 1 June 1977, J.I. Glick, ex. Echinohelea lanei, CSP-77-0601~Ala, (1L) (AU). Al:Lee Co., Loachapoka, 8-11 June 1977, J.I. Glick, ex. Monohelea stonei, N.J. light trap, LCP-77-0608-ll-la, (1L) (AU). AL. Lee Co., Auburn, Chewacla St. Pk., 6 July 1977, J.I. Glick, ex. Culicoides stellifer, CSP-77-0706-Ala, (1L) (AU). AL., Geneva, Slocorab, 21 May 1979, G.R. Mullen, ex. Dasyhelea scutelleta (Ceratopogonidae), GRM-78-0608-2c, (1L) (AU).

Willmannella johnstoni (Robaux, 1977) NEW COMB. West Virginia, Hampshire Co., N of Yellow Springs, 23-vi-1961, T. Savage, (IP, 2L) (OSU).

Willmannella racovitzae (Feider, 1948) Romania, Babaday, Tulcea, 3-vii-1955, Nr. 523 and 604, (2L) (FEIDER). Table 10 (continued) 251

The Academy of Natural Sciences of Philadelphia ANSP 19th and The Parkway Philadelphia, PA 19103

American Museum of Natural History AMNH Department of Entomology Central Park West at 79th St. New York, NY 10024

Berlese Collection BERLESE Dr. Fausta Pegazanno Instituto Spermentale per la Zoologica Agraria Firenze, Italy

Bernice P. Bishop Museum BPBM P.O. Box 19000-A Honolulu, Hawaii 96819

British Musuem (Natural History) BMNH Cromwell Road London, England SW7 5BD

Canadian National Collection CNC Biosystematics Research Institute Ottawa, Canada K1A 0C6

Cornell University CU Department of Entomology Ithaca, NY 14850

Feider Collection FEIDER Dr. Libertina Solomon Laboratorul de Zoologie Universitate "Al. I. Cuza" Iasi, Romania

Dr. B. Feldman-Muhsara HUMS Department of Medical Entoraolgy The Hebrew University-Hadassah Mecical School P.O.B. 1172 Jerusalem, Israel

Field Musuem of Natural History FMNH Chicago, IL 60605

Dr. Gary R. Mullen AU Department of Zoology-Entomology Auburn University Auburn, AL 38830 Table 10 (continued) 252

Museum of Comparative Zoology MCZ Harvard University Cambridge, MA 02138

Museum Natural D'Histoire Naturelle de Paris MNHNP Laboratorie De Zoologie 61 Rue Buffon Paris, France 75005

Museum of Zoology UMMZ University of Michigan Ann Arbor, MI 48109

Naturhistoriska Riksmuseet NR Entomologiska Avdelingen S-104 05 Stockholm 50 Sweeden

Ouderaans Collection 0C Dr. L. Van Der Hammen Rijksrauseum voor Natuurlijke Histoire Raamsteg 2 Leiden, Netherlands

Dr. Graeme Ramsay NZ D.S.I.R. Entomology Division Mt. Albert Reseach Center Private Bag Auckland, New Zealand

Smithsonian Institution USNM Washington, D.C. 20560

Snow Entomological Museum KU Department of Entomology University of Kansas Lawrence, KA 66045

South Australia Musuem SAM Arachnid Section North Terrace Adelaid, S.A. 5000 Australia

Tuttle Collection TUTTLE Dr. Donald M. Tuttle 6425 W 8th St. Yuma, Arizona 85364 Table 10 (continued)

Zoologische Staatssarnmlung Maria - Work - Strabe lb D-8000 Munich, West Germany