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Antony, Lucille Marilyn May Kriger d’Amorim

A PHYLOGENETIC ANALYSIS OF THE RHODACAROIDEA (: ) '

The Ohio State University Ph.D. 1986

University Microfilms I nternetionel300 N. zeeb Road, Ann Arbor, Ml 48106

Copyright 1986

by

Antony, Lucille Marilyn May Kriger d’Amorim

All Rights Reserved A PHYLOGENETIC ANALYSIS OF THE RHODACAROIDEA

(ACARI; MESOSTIGMATA)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Phylosophy in the Graduate

School of the Ohio State University

By

Lucille Marilyn May Kriger d*Amorim Antony, B.Sc., M.Sc.

******

The Ohio State University

1986

Dissertation Committee: Approved by

N.F. Johnson

S.W. Fisher

D.L. Wrensch ^ idviser

Department of Entomology Copyright by

Lucille Marilyn May Kriger d'Amorim Antony

1986 To My Mother and to the Memory of My Father

ii ACKNOWLEDGMENTS

I want to thank my major Professor Dr. Donald E. Johnston, for his advice, and especially for the stimulating discussions through the course of my studies at Ohio State. Thanks to him (and others), my memories of OSU will remain indelible.

I am very grateful to the Conselho Nacional de Desenvolvimento

Cientifico e Tecnologico and to The Organization of American States for granting two-year scholarships during my graduate studies.

I want to thank the acarologists and curators of collections who

generously provided me with the material studied. Among them, special

thanks to Mr. Keith Hyatt, Miss Anne Baker, Mr. Kieran Martyn of the

British Museum of Natural History, Dr. Fausta Pegazzano of the

Istituto Sperimentale per la Zoologia Agraria, and Dr. Jo Ann Tenorio

of the Bernice Pauahi Bishop Museum, for their hospitality and for

providing access to the collections.

I thank the Instructional and Research Computer Center of The

Ohio State University, for granting invaluable computer time.

I extend my appreciation to Dr. Norman F. Johnson, for spending

a great deal of his time helping me to get acquainted with the

"mini-" and other bugs of the computer world.

iii My warmest thanks to the good friends who, directly or indirectly contributed to the accomplishment of this study.

Finally, and most specially, I thank for all the love and constant support with which my very special family have gifted me all through the years. To them, my deepest love and appreciation.

iv VITA

October 15, 1950 . . . Born - Manaus, Amazonas, Brasil

1972-1975...... Trainee student. Laboratory of Pedobiology, Instituto Nacional de Pesquisas da Amazonia (INPA), Manaus, Brasil.

1973 ...... B.Sc. - Pharmacy, Universidade Federal do Amazonas, Manaus, College of Health Sciences.

1975-1977...... Research Assistant, Laboratory of Pedobiology, INPA, Manaus.

1978-1979...... Graduate studies, Department of Biology, Georgia Southern College, Statesboro. Sponsored by The Organization of American States.

1980 ...... M.Sc. - Entomology, The Ohio State University, , Ohio, sponsored by The Organization Of American States.

1981-1983...... Graduate Teaching Associate, Department of Entomology, The Ohio State University.

1983-1985...... Doctoral Student, Department of Entomology, The Ohio State University. Sponsored by the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico, Brasil.

1985-1986...... Graduate Teaching Associate, Department of Entomology, The Ohio State University.

Publications

Antony, L.M.M.K.d'A. and Johnston, D.E., 1984. A Cladistic Analysis of the Genus Berlese (Acari; Mesostigmata). In; Griffiths, D.A. and Bowman, C.E.(eds.). Acarologv VI. v.l. Ellis Norwood Publ., Chichester, England, p.216-221. Presentations

Antony, L. M, and Johnston, D, E. Phylogenetic studies of primitive Dermanyssina: The Rhodacaroidea (Acari). Entomological Society of America, Annual Meeting, Hollywood, Florida, 8-12 December, 1985.

FIELDS OF STUDY

Major Field; Entomology

Studies in and General Biology of Oribatid , Laboratory of Pedobiology, Instituto Nacional de Pesquisas da Amazonia, Manaus, Brasil. Dr. Herbert 0. R. Schubart.

Studies in Reproductive Biology of Dermanvssus gallinae De Geer (Acari: Dermanyssidae). Department of Biology, Georgia Southern College, Statesboro, Georgia. Dr. James H. Oliver Jr., Callaway Professor.

Studies in Taxonomy of Free-living Gamasina (Acari: Mesostigmata). Laboratory of Acarology, The Ohio State University, Columbus. Dr. Donald E. Johnston, Professor.

Vi TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS...... i ü

VITA...... V

LIST OF TABLES...... ix

LIST OF FIGURES ...... x

INTRODUCTION...... 1

MATERIALS AND METHODS...... 26

CHARACTER ANALYSIS...... 34

RESULTS AND DISCUSSION...... 57

Suprafamilial relationships...... 57

Interfamilial relationships...... 71

Intrafamilial relationships...... 78

SYSTEMATIC ACCOUNTS ...... 130

Superfamily Rhodacaroidea Ouderaans ...... 130

Family Euryparasitidae n. fam...... 138

Family Ouderaans ...... 144

Family Gamasellidae n. f a m ...... 155

Family Ologaraasidae Ryke ...... 162

Subfamily Ologamasinae Ryke...... 169

Subfamily Epiphidinae Kethley...... 171

Subfamily Gamasiphinae Lee ...... 172

vii Taxa removed from the Rhodacaroidea ...... 173

Key to the families of Rhodacaroidea...... 178

Superfamily Laelaptonyssoidea Womersley ...... 182

Family Panteniphidae n. f a m ...... 188

SUMMARY...... 205

LIST OF REFERENCES...... 207

APPENDICES...... 213

A. MATERIAL EXAMINED (LISTED ALPHABETICALLY BY REPOSITORY) . .

...... 213

B. DATA MATRIX...... 256

viii LIST OF TABLES

TABLE Page

1. Family Rhodacaridae Oudemans sensu Ryke, 1962. 9

2. Superfamily Eugamasoidea Karg, 1965...... 13

3. Family Rhodacaridae Oudemans sensu Lee, 1970, 1973 .... 16

4. Family Rhodacaridae Oudemans sensu Bregetova, 1977 .... 22

5. Family Rhodacaridae Oudemans sensu Shcherbak, 1980 .... 24

6. Character Polarization ...... 49

7. Proposed classification of the Rhodacaroidea and

Laelaptonyssoidea. The arrangement of families is

phyletically sequenced ...... 126

IX LIST OF FIGURES

FIGURE Page

1. Suprafamilial and interfamilial relationships of the

Rhodacaroidea. Numbers on figure refer to stems.

Characters are listed following sequence of stems.

Character states are listed between parentheses...... 192

2. Intrafamilial relationships of the Euryparasitidae n. fam.

Refer to Figure 1 legend for explanation...... 194

3. Intrafamilial relationships of the Rhodacaridae. Refer to

Figure 1 legend for explanation...... 196

4. Intrafamilial relationships of the Gamasellidae. Refer to

Figure 1 legend for explanation...... 198

5. Intrafamilial relationships of the Ologamasinae. Refer to

Figure 1 legend for explanation...... 201

6. Intrafamilial relationships of the Epiphidinae and

Gamasiphinae. Refer to Figure 1 legend for explanation. . 203

X INTRODUCTION

The Rhodacaroidea is a large group of ecologically and morphologically diverse free-living mites of the suborder

Mesostigmata. Together with the Dermanyssoidea, Eviphidoidea,

Ascoidea and Veigaioidea they comprise the cohort Dermanyssina

(Johnston, 1982). Females of this cohort are characterized by the

presence of an accessory genital system - the spermatheca - which

consists of paired ducts that open in the region of legs III and IV.

The males possess a sperm-transfer device - the spermadactyl -

located on the movable digit of the chelicera. Their genital

aperture is placed at the anterior edge of the sternum (sternogenital

shield) and opens through an eversible ejaculatory organ by which

sperm is transferred to the chelicerae (Johnston, 1982).

The majority of species of Rhodacaroidea is primarily of

soil-dwelling, predatory habits. They inhabit forest and mineral

soils, having been collected from a wide variety of habitats such as

litter, humus of primary and secondary forests, grasslands, rotting

vegetable matter (compost, corn stalks, grass and twigs), cave

debris, under stones or bark of trees, in dung and deeper soil

layers. The last is a common habitat of Rhodacaridae sensu stricto.

with several species of having been recovered from depths

1 2 at 20-26cm in pine forest soils in California (Price, 1973). A few of its members are found associated with nests of rodents,

insectivores and birds, while others seem restricted to littoral

zones (living on sandy or rocky shores or even in crevices between

barnacle carapaces) or intertidal zones (living among algae) or

living among aquatic plants and algae of freshwater lakes. Shcherbak

(1980) points out the theoretical and practical importance of

studying rhodacarid mites as they play an important role in soil

communities "which is indirectly indicated by their widespread

distribution in soils of natural habitats and agrocoenoses, hotbeds

and greenhouses". They favor collembolans, nematodes and other mites

as prey, but small insect larvae and mole fleas may also be eaten.

This has been observed under laboratory conditions by Michael (1886),

Glynne-Williams and Hobart (1952), Karg (1961), Gressitt (1967), and

Lee (1974). Because rhodacarids use extraoral digestion (they take

their food only in liquid form), it is rather difficult to determine

their natural prey (Hurlbutt, 1968). Even though most of its genera

have been found only in the Southern Hemisphere, the group is

cosmopolitan (Lee, 1970).

However insufficiently collected and very poorly understood, the

morphological heterogeneity exhibited by the group has encouraged

taxonomic and systematic studies by several workers who have

attempted to define it. Consequently, a number of concepts of the

group have been proposed throughout the years, some very restricted

like those of Oudemans (1902, 1939) who established the subfamily

Rhodacarinae, Halbert (1915) who elevated the group to the family 3 level, and Vitzthum (1941). Broader concepts included those of

Berlese (1913), Evans (1957, 1963), Athias-Henriot (1961), Hirschmann

(1962), Ryke (1958, 1962), Karg (1965, 1971, 1977, 1979), Bregetova and Shcherbak (1977) and Shcherbak (1976, 1979, 1980, 1983).

A major effort to define the Rhodacaridae has been made by Lee

(1970), who contributed the most comprehensive systematic study of the group. Another important contribution is the monograph on the

Palearctic Rhodacaridae by Shcherbak (1980).

One of the current classifications of the Rhodacaroidea is that of Krantz (1978) who adopted Lee's concepts with the following modifications: the Rhodacaridae sensu Lee (-Rhodacaridae sensu lato ) was elevated to the superfamily level and divided into two families: the Rhodacaridae which includes the Rhodacarinae Oudemans, 1902 sensu

Lee (1970), («Rhodacaridae sensu stricto of Johnston, 1968) with the genera Rhodacarus Oudemans, 1902, Willmann, 1935,

Rhodacaropsis Willmann, 1935, Afrogamasellus Loots and Ryke, 1968; and the Ryke, 1962 («Gamasellinae Hirschmann, 1962;

Cyrtolaelapidae Berlese, 1913 sensu Johnston 1968, in part), which

includes all of Lee's non-rhodacarinae rhodacarids, namely the

Gamasiphinae Lee, 1970, Ologamasinae Ryke, 1962, Sessiluncinae Lee,

1970, Laelaptonyssinae Womersley, 1956a, and Tangaroellinae Lee,

1970.

In addition to these, Krantz expanded the limits of the

superfamily to accomodate the Evans, 1957 sensu

Lindquist (1975), who included four genera: Halbert,

1915 with the subgenera ^ (Dendrolaelaps Halbert) and ^ 4

(Dendrolaelaspis Lindquist, 1975); Dendroseius Karg, 1965;

Digamasellus Berlese, 1905; Loneoseius Chant, 1961 with the subgenera

L. (Loneoseius Chant) and ^ (Loneoseiulus Lindquist, 1975).

In the sense of Krantz (1978), the Rhodacaroidea can be defined

in the following way: tibia I with three or four ventral setae;

tibia III with seven or eight setae; genu IV with one or two ventral

setae; palptarsal claw (apotele) with two or three prongs; ventrianal

shield present; males with genital aperture placed at anterior margin

of sternal shield, and spermadactyl fused to base of cheliceral

movable digit but can be free or fused distally to digit; females

podospermal, with sperm induction pores opening on coxae, trochantera

or femora III, or on endopodal, metapodal or, rarely, on sternal

shield; sternal shield of females generally with four pairs of setae;

often, two pairs of scleronoduli present on podonotum; deutonymphs

with divided dorsal shield; protonymphal pygidial shield with seven

or eight pairs of setae, with setae S4 and S5 inserted marginally on

shield or on integument; larvae with or without pygidial shield. It

should be noted that Krantz*s concept of the Rhodacaroidea was the

one I adopted to begin this study .

Alternative classifications of the Rhodacaridae sensu stricto

are proposed by Karg (1965, 1971) and Shcherbak (1979, 1980).

Karg (1971) includes Rhodacarus. Rhodacarellus. Protogamasellus

Karg, 1962, Dendrolaelaps sensu lato, with the subgenera Dendroseius

and Dendrolaelaps sensu stricto to form the Rhodacaridae within the

superfamily Eugamasoidea Karg, 1965.

Lindquist and Evans (1965) regard Protogamasellus as a member of 5 the subfamily Ascinae within the Voigts and Oudemans, 1905.

Karg's Eugamasoidea (an invalid name) includes another family, the Eugamasidae Hirschmann, 1962 (another invalid name) which contains three subfamilies: the Gamasellinae Hirschmann, 1962, the

Parasitinae Oudemans, 1901 sensu Evans (1957) and the Veigaiainae

Oudemans, 1901 sensu Bregetova (1961). The genera included in the

Gamasellinae Hirschmann are distributed among three subfamilies of the Ologamasidae Ryke sensu Krantz (1978).

In subsequent papers (1975, 1976a-c, 1977, 1979) Karg gradually expands his concept of the Rhodacaridae, by including the following genera: Neogamasellevans Loots and Ryke, 1967, Gamasiphoides

Womersley, 1956, Laelaptiella Womersley, 1956, Heterogamasus

Tragardh, 1907, Geogamasus Lee, 1970, Ologamasus Berlese, 1888,

Hevdeniella Richters, 1907, Allogamasellus Athias-Henriot, 1961,

Afrogamasellus Loots and Ryke, 1968 (containing the subgenera

Afrogamasellus. Podalogamasellus Karg, 1977, Foliogamasellus Karg,

1977, and Latogamasellus Karg, 1977) and Rhodacaroides Willmann, 1959

(with the subgenera Rhodacaroides. Tenacaroides Karg, 1977, and

Nodacaroides Karg, 1977), Acugamasus Lee, 1970 and Dendrolaelaps

(Dendrolaelaspis) Lindquist, 1975. He also seems to have further

modified his original concept of the family, for he excludes

Protogamasellus Karg and refers to the genera previously placed in

the Gamasellinae Hirschmann as the Rhodacaridae sensu lato.

Shcherbak (1979, 1980) divides the Rhodacaridae into three

subfamilies: Rhodacarinae Oudemans, 1902 (containing Rhodacarus

(Rhodacarus). Rhodacarus (Multidentorhodacarus) Shcherbak, 1980, 6

Rhodacaropsis. and Mediorhodacarus Shcherbak, 1976; Rhodacarellinae

Shcherbak, 1980 (containing Rhodacarellus. Afrogamasellus Loots and

Ryke, 1968, Afrodacarellus Hurlbutt, 1973 and Minirhodacarellus

Shcherbak, 1980) and Dendrolaelapinae Hirschmann, 1960 (including

Dendrolaelaps. Dendroseius. Dendrolaelaspis. Longoseius.

Multidendrolaelaps Hirschmann, 1974, Insectolaelaps Shcherbak, 1980 and Oligodentatus Shcherbak, 1980. More recently (1982), Shcherbak included Longoseiulus as a subgenus of Dendrolaelaps and added

Orientolaelaps Bregetova and Shcherbak, 1977 to the Dendrolaelapinae.

With this change, she seems to indicate that the Digamasellidae Evans sensu Lindquist (1975) should include only the genus .

Shcherbak (1980) also seems to exclude Protogamaspllns from the

Rhodacaridae, even though she does not clearly indicate this (for

example, Protogamasellus is not listed among the species excluded

from the family, nor are suggestions concerning its placement

provided).

Another instance of controversy refers to the definition and

placement of the genus Panteniphis Willmann, 1949. When first

described (from three females collected in Poland), this genus was

considered to have a single species - mirandus - and was not formally

placed into any family. In 1969, Athias-Henriot found and described

a male of Panteniphis mirandus collected in France and attributed the

genus to the Rhodacaridae sensu lato. In the same year, Genis, Loots

and Ryke described a new genus and species - Lindquistoseius

africanus - from several females collected in southern and eastern

Africa. The authors referred Lindquistoseius to the Ascidae. Lee 7

(1970) transferred Panteniphis to the Ascidae on the basis of its reduced leg setation, the presence of a two-pronged apotele and the inconspicuous enlargement of setae on the male leg II.

In 1975, Hurlbutt found and described a small series of adult males and females (collected in Tanzania, East Africa), which he considered to be conspecific with Lindquistoseius africanus. He regarded mirandus and africanus as congeneric species and reduced

Lindquistoseius to a subgenus of Panteniphis. He also discussed the relationships of the genus and concluded that Panteniphis was more closely related to the Rhodacaridae (sensu lato) than to the Ascidae.

Bregetova (1977) has referred Panteniphis to the Asceosejidae (i.e.

Ascidae, in part), while Hirschmann (1983) regards Lindquistoseius a subgenus of Panteniphis.

From the examples of classifications listed above, it can be

clearly seen that there is no general agreement between any two of

these classifications. They all reflect completely opposing points

of view, especially concerning the make-up of the Rhodacaridae sensu

stricto.

Furthermore, there is vacillation when it comes to determining

the placement of certain taxa which appear to be closely related to

the Rhodacaroidea. Among the taxa with controversial placements are;

Panteniphis. Lindquistoseius. Rhodacaroides. Protogamasellus.

Gamasellodes Athias-Henriot, 1961 and Laelaptonvssus Womersley, 1956.

The purpose of this study is to determine the internal and

external relationships of this complex group - the Rhodacaroidea -

through a phylogenetically based analysis, so that a rational (and if 8 possible, conservative) classification can be achieved.

The above mentioned concepts are examined and compared, with particular interest being given to those developed by Lee (1970),

Lindquist (1975) and Shcherbak (1979, 1980). The placement of controversial taxa is attempted on the basis of the consistency of results derived from the phylogenetic analysis.

Where necessary, diagnoses, descriptions and illustrations are provided. A summary of the major prior classifications of the

Rhodacaroidea is presented in Tables 1-5. Table 1. Family Rhodacaridae Oudemans sensu Ryke, 1962

Subfamily Rhodacarinae Oudemans,1902

Type Genus: Rhodacarus Oudemans, 1902 Type species: Rhodacarus Oudemans - containing 4 subgenera:

R. (Rhodacarus) Oudemans, 1902 Rhodacarus roseus Oudemans

R. (Rhodacaropsis) Willmann, 1935 Rhodacaropsis inexpectatus Willmann

R. (Rhodacarellus) Willmann. 1935 Rhodacarellus subterraneus Willmann

R. (Rhodacaroides) Willmann, 1959 Rhodacaroides aegyptiacus Willmann

Cyrtolaelaps Berlese, 1887 nec Berlese 1892 - containing 5 subgenera:

G. () Berlese, 1887 Gamasus mucronatus G. & R. (= Protolaelaps Tragardh, 1912) Ganestrini, 1881

G. () Berlese, 1892 Gamasus falciger G. & R. (= Laelogamasus Berlese, 1905) Ganestrini, 1881

G. (Digamasellus) Berlese, 1905 Digamasellus perpusillus Berlese. 1905 (= Dendrolaelaps Halbert, 1915)

G. () Oudemans, 1902 Gamasus emarginatus Koch, 1839

G. (Gamaselliphis) Ryke, 1961 G. (Gamaselliphis) potchefstroomensis Ryke v£> Table 1. (Continued)

Asca von Heyden, 1826 Acarus aphidioides Linnaeus, 1758 (= Ceratozercon Berlese, 1913)

Evanssellus Ryke, 1961 Evanssellus foliatus Ryke

Antennoseius Berlese, 1916 Antennoseius delicatus Berlese (= Vitzthumia Thor, 1930)

Halolaelaps Berlese & Trouessart, 1889 - containing 2 subgenera:

H. (Halolaelaps) Berl. & Trous., 1889 H. glabriusculus Berl. & Trous. (= Gamasus marinus Brady, 1875)

H. (Saprogamasellus) Goetz, iji_ H. (Saprogamasellus) strenzkei Goetz Sellnick, 1957

Saprolaelaps Leitner, 1946 Saprolaelaps subtilis Leitner

Leitneria Evans, 1957 Gamasellus (Protolaelaps) granulatus Halbert, 1923

Longoseius Chant, 1961 Longoseius cuniculus Chant

Saintdieria Oudemans, 1939 Parasitas sexclavatus Oudemans, 1902 Table 1. (Continued)

Subfamily Ologamasinae Ryke, 1962

Type Genus: Ologamasus Berlese, 1888 Type species: Ologamasus Berlese, 1888 Gamasus aberrans Berlese, 1888 (= Ologamasellus Berlese, 1914 (lapsus Hologamasus Berlese, 1892)

Antennolaelaps Womersley, 1956 Antennolaelaps affinis Womersley

Epiphis Berlese, 1916 Gamasiphis (Epiphis) rarior Berlese

Gamasiphis Berlese, 1904 Gamasiphis pulchellus Berlese (s= Heteroiphis arcuatus Tragardh, 1952) (= Hevdeniella Richters, 1907)

Gamasiphoides Womersley, 1956 G. (Gamasiphoides) propinquus Womersley

Gamasitus Womersley, 1956 Gamasitus obscurus Womerley

Hydrogamasus Berlese, 1892 Gamasus salinus Laboulbene, 1851 (= Gamasus littoralis G. & R. Ganestrini, 1881)

Laelaptiella Womersley, 1956 Laelaptiella anomala Womersley

Megaliphis Willmann, 1938 Gamasiphis (Megaliphis) giganteus Willmann Table 1. (Continued)

Micriphis Berlese, 1914 Gamasiphis gamasellus Berlese 1913

Neoeamasiphis Tragardh, 1952 Neogamasiphis hamifer Tragardh

Onchogamasus Womersley, 1956 Onchogamasus communis Womersley

Pachyselus Berlese, 1910 Pachyseius humeralis Berlese

Parasitiphis Womersley, 1956 Parasitiphis littoralis Womersley

Periphis Berlese, 1914 Iphis haemisphaericus Koch, 1839

Physallolaelaps Berlese, 1908 Physallolaelaps ampulliger Berlese

Queenslandolaelaps Womersley, 1956 Queenslandolaelaps vitzthumi Womersley

Sessiluncus Ganestrini, 1898 Gamasus heterotarsus Ganestrini, 1897

Trachygamasus Berlese, 1905 Gamasus pusillus Berlese, 1892

N> Table 2. Superfamily Eugamasoidea Karg, 1965

Family Rhodacaridae Oudemans sensu Karg, 1965, 1971

Type genus: Rhodacarus Oudemans, 1902 Type species:

Rhodacarus Oudemans, 1902 Rhodacarus roseus Oudemans Rhodacarellus Willmann, 1935 Rhodacarellus subterraneus Willmann

Dendrolaelaps Halbert. 1915 sensu lato - containing 2 subgenera:

D. (Dendrolaelaps) Halbert, 1915 Sj^ str. Dendrolaelaps oudemansi Halbert

D. (Dendroseius) Karg. 1965 Dendrolaelaps scotarius Sheals. 1958

Protogamasellus Karg. 1962 Protogamasellus primitives Karg

Family Eugamasidae Hirschmann

Subfamily Gamasellinae Hirschmann, 1962

Type genus: Gamasellus Berlese, 1892 Type species:

Cvrtolaelaps Berlese, 1887 Gamasus mucronatus G. & R. (= Protolaelaps Tragardh, 1912) Canestrini, 1881 (= Asca affinis Oudemans, 1902)

Gamasellus Berlese, 1892 Gamasellus falciger G. & R. CO Canestrini, 1882 Table 2. (Continued)

Euryparasitus Oudemans, 1901 Gamasus emarginatus Koch, 1839

Stylochirus G. & R. Canestrini, 1882 Stylochirus rovennensis G. & R. Canestrini

Hydrogamasus Berlese, 1892 Gamasus salinus Laboulbene, 1851 (= Gamasus littoralis G. & R. Canestrini, 1881)

Sessiluncus G, Canestrini, 1898 - containing 2 subgenera:

S. (Sessiluncus) G. Canestrini, 1898 Gamasus heterotarsus G. Canestrini, 1897

S. (Hydrogamasellus) Hirschmann, 1966 Hydrogamasus antarcticus Tragardh, 1907

Gamasiphis Berlese. 1903 Gamasus pulchellus Berlese. 1887

Subfamily Parasitinae Oudemans, 1901 sensu Evans, 1957

Type genus: Parasitus Latreille, 1795 Type species:

Parasitas Latreille, 1795 Acarus coleoptratorum Linnaeus. 1758 (= Acarus fucorum Degeer, 1778)

Holoparasitus Oudemans, 1936 Gamasus calcaratus Koch, 1839

Pergamasus Berlese, 1904 Acarus crassipes Linnaeus. 1758

Saprogamasus Willmann, 1949 Saprogamasus ambulacralis Willmann

Eugamasus Berlese, 1893 Gamasus magnus Kramer, 1876 Table 2. (Continued)

Poecilochirus G, & R. Canestrini, 1882 Poecilochirus carabi G. & R. Canestrini

Gamasodes Oudemans, 1939 Gamasodes spiniger Oudemans, 1936 (= Gamasus spiniger Koch, 1841)

Trachygamasus Berlese, 1906 Gamasus pusillus Berlese, 1892

Subfamily Veigaiainae Oudemans, 1939 sensu Bregetova, 1961

Type genus: Oudemans, 1905 Type species:

Veigaia Oudemans, 1905 Gamasus nemorensis Koch, 1836

Cyrthydrolaelaps Berlese, 1904 Cyrthydrolaelaps hirtus Berlese

Gamasolaelaps Berlese, 1904 Cyrtolaelaps aurantiacus Berlese, 1903

Ln Table 3, Family Rhodacaridae Oudemans sensu Lee, 1970, 1973

Subfamily Rhodacarinae Oudemans

Type Genus: Rhodacarus Oudemans, 1902 Type species:

Rhodacarus Oudemans, 1902 Rhodacarus roseus Oudemans

Rhodacarellus Willmann, 1935 Rhodacarellus subterraneus Willmann

Rhodacaropsis Willmann, 1935 Rhodacaropsis inexpectatus Willmann

Afrogamasellus Loots & Ryke, 1968 Cyrtolaelaps (Gamasellus) franzi Loots & Ryke, 1966 !

Subfamily Gamasiphinae Lee, 1970

Type Genus: Gamasiphis Berlese, 1904 Type species:

Gamasiphis Berlese, 1904 Gamasus pulchellus Berlese, 1887 (=* Micriphis Berlese, 1914) (= Heteroiphis Tragardh, 1952) (= Neoeamasiphis Tragardh, 1952)

Caliphis Lee, 1970 Caliphis calvus Lee

Euepicrius Womersley, 1942 Euepicrius filamentosus Womersley

o\ Table 3. (Continued)

Gamaselliphis Ryke, 1961 Cyrtolaelaps (Gamaselliphis) potchefstroomensis Ryke

Gamasiphoides Womersley, 1956 Gamasiphis (Gamasiphoides) propinquus Womersley

Hydrogamasus Berlese, 1892 Gamasus littoralis G, & R. Canestrini, 1881

Laelaptiella Womersley, 1956 Laelaptiella anomala Womersley

Subfamily Laelaptonyssinae Womersley, 1956

Type Genus; Laelaptonyssus Womersley, 1956 Type species:

Laelaptonyssus Womersley, 1956 Laelaptonyssus mitis Womersley

Subfamily Ologamasinae Ryke, 1962 (= Gamasellinae Hirschmann, 1962) (=» Cyrtolaelapidae Berlese sensu Johnston, 1968 - in part)

Type Genus: Ologamasus Berlese, 1888 Type species:

Tribe Ologamasini Berlese, 1888

Ologamasus Berlese, 1888 Gamasus aberrans Berlese, 1892 (= Ologamasellus Berlese, 1914) (lapsus Hologamasus Berlese, 1892 nec Berlese, 1906) Table 3. (Continued)

Cymiphis Lee, 1970 Ologamasus cymosus Lee, 1966

Geogamasus Lee, 1970 Geogamasus skoshi Lee

Heydeniella Richters, 1907 Heydeniella crozetensis Richters

Athiasella Lee, 1973 Hydrogamasus dentatus Womersley, 1942 (= Heydeniella Richters dentata - complex Lee, 1970)

Hydrogamasellus Hirschmann, 1966 Hydrogamasus antarcticus Tragardh, 1907

Neogamasellevans Loots & Ryke, 1967 Neogamasellevans preendopodalis Loots & Ryke

Parasitiphis Womersley, 1956 Parasitiphis littoralis Womersley (= Austrohydrogamasus Hirschmann, 1966)

Pyriphis Lee, 1970 Ologamasus pyrenoides Lee, 1966

Rykellus Lee, 1970 Cyrtolaelaps (Gamasellus) darglensis Ryke, 1962

00 Table 3. (Continued)

Tribe Gamasellini Hirschmann, 1962 (= Cyrtolaelaptini Berlese, 1913)

Type Genus: Gamasellus Berlese, 1892 Type species:

Gamasellus Berlese, 1892 Gamasus falciger G, & R. Canestrini, 1881

Acugamasus Lee, 1970 Digamasellus punctatus Womersley, 1942

Allogamasellus Athias-Henriot, 1961 Allogamasellus aquafortensis Athias-Henriot

Cyrtolaelaps Berlese, 1887 Gamasus mucronatus G. & R. Canestrini, 1881

Euryparasitus Oudemans, 1902 Gamasus emarginatus Koch, 1839

Evanssellus Ryke, 1961 Evanssellus foliatus Ryke

Heterogamasus Tragardh, 1907 Heterogamasus claviger Tragardh

Hiniphis Lee, 1970 Hiniphis hinnus Lee

Laelogamasus Berlese, 1905 Gamasus (Laelogamasus) simplex Berlese, 1905

Litogamasus Lee, 1970 Cyrtolaelaps setosus Kramer, 1898

VO Table 3. (Continued)

Notogainasellus Loots & Ryke, 1966 - containing 2 subgenera:

N. (Notogamasellus) Loots & Ryke, 1966 (Notogamasellus) vandenbergi Loots & Ryke

N. (Podonotogamasellus) Loots & Ryke, 1966 N. (Podonotogamasellus) magoebaensis Loots & Ryke

Periseius Womersley, 1961 - containing 2 subgenera:

P. (Periseius) Womersley. 1961 Cyrtolaelaps hammeni Wom.. 1961 (= (Periseius) littorale Womersley, 1961

P. (Psammonsella) Hag. 1965 P. (Psammonsella) nobskae Hag

Pilellus Lee, 1970 Cyrtolaelps (Gamasellus) rykei Hunter, 1967

Rhodacaroides Willmann, 1959 Rhodacaroides aegyptlacus Willmann

Solugamasus Lee, 1973 Solugamasus mustela Lee

Subfamily Tangaroellinae Lee, 1970

Type Genus: Tangaroellus Luxton, 1968 Type species:

Tangaroellus Luxton. 1968 Tangaroellus porosus Luxton Table 3. (Continued)

Subfamily Sessiluncinae Lee, 1970

Type Genus: Sessiluncus G, Canestrini, 1898 Type species:

Sessiluncus G. Canestrini, 1898 Gamasus heterotarsus G. Canestrini, 1897

Antennolaelaps Womersley, 1956 Antennolaelaps affinis Womersley

Gamasellevans Loots & Ryke, 1967 Gamasellevans epigynialis Loots & Ryke

Gamasellopsis Loots & Ryke, 1966 Gamasellopsis curtipilus Loots & Ryke

Gamasitus Womersley, 1956 Gamasitus obscurus Womersley

Onchogamasus Womersley, 1956 Onchogamasus communis Womersley

Paragamasellevans Loots & Ryke, 1968 Paragamasellevans michaeli Loots & Ryke

Queenslandolaelaps Womersley, 1956 Queenslandolaelaps vitzthumi Womersley

Stylochirus G, & R. Canestrini, 1882 Stylochirus rovennensis G. & R, Canestrini, 1882 (= Physallolaelaps Berlese, 1908) (= Periphis Berlese, 1914) (= Epiphis Berlese, 1916) (= Megaliphis Willmann, 1938)

lo Table 4, Family Rhodacaridae Oudemans sensu Bregetova, 1977

Type Genus: Rhodacarus Oudemans, 1902 Type species:

Rhodacarus Oudemans, 1902 Rhodacarus roseus Oudemans

Rhodacarellus Willmann, 1935 Rhodacarellus subterraneus Willmann

Rhodacaropsis Willmann, 1935 Rhodacaropsis inexpectatus Willmann

Protogamasellus Karg, 1962 Protogamasellus primitivus Karg

Gamasellodes Athias-Henriot, 1961 Gamasellodes vulgatior Athias-Henriot

Digamasellus Berlese, 1905 Gamasellus (Digamasellus) perpusillus Berlese, 1905 (= Cyrtolaelaps punctum Berlese, 1904)

Longoseius Chant, 1961 Longoseius cuniculus Chant

Dendrolaelaps Halbert, 1915 Dendrolaelaps oudemansi Halbert

Cyrtolaelaps Berlese, 1887 Gamasus mucronatus G. & R. (= Protolaelaps Tragardh, 1912) Canestrini, 1881

Euryparasitus Oudemans, 1901 Gamasus emarginatus Koch, 1839 (= Eurylaelaps Oudemans, 1902, nom, nud. lapsus)

Gamasellus Berlese, 1892 Gamasus falciger G. & R, (Canestrini, 1881 Table 4. (Continued)

Asca V. Heyden, 1826 Acarus aphidioides Linnaeus, 1758 (= Ceratozercon Berlese, 1913)

Leitneria Evans, 1957 Gamasellus (Protolaelaps) granulatus Halbert, 1923

Halolaelaps Berlese & Trouessart, 1889 Halolaelaps glabriusculus (= Saintdieria Oudemans, 1939) Berlese & Trouessart, 1889 (= Saprolaelaps Leitner, 1946) (= Gamasus marinus Brady, 1875)

Family Ologamasidae Ryke sensu Bregetova, 1977

Type Genus; Ologamasus Berlese, 1888 Type species:

Ologamasus Berlese, 1888 Gamasus aberrans Berlese. 1888 (= Ologamasellus Berlese, 1914) (lapsus Hologamasus Berlese, 1892)

Stylochirus G. & R. Canestrini, 1882 Stylochirus rovennensis G. & R. (= Physallolaelaps Berlese, 1908) Canestrini (= Periphis Berlese, 1914) (= Epiphis Berlese, 1916) (= Megaliphis Willmann, 1938)

Gamasiphis Berlese, 1904 Gamasus pulchellus Berlese. 1887

Sessiluncus G. Canestrini, 1898 Gamasus heterotarsus G. Canestrini, 1897

5 Table 5. Family Rhodacaridae Oudemans sensu Shcherbak, 1980

Subfamily Rhodacarinae Oudemans, 1902

Type Genus: Rhodacarus Oudemans, 1902 Type species:

Rhodacarus Oudemans, 1902 - containing 2 subgenera:

R. (Rhodacarus) Oudemans, 1902 Rhodacarus roseus Oudemans

R. (Multidentorhodacarus) Shcherbak, 1980 ^ (Multidentorhodacarus) denticulatus Berlese, 1921

Rhodacaropsis Willmann, 1935 Rhodacaropsis inexpectatus Willmann

Mediorhodacarus Shcherbak, 1976 Mediorhodacarus tetranodulosus Shcherbak

Subfamily Rhodacarellinae, Shcherbak, 1980

Type Genus: Rhodacarellus Willmann, 1935 Type species:

Rhodacarellus Willmann, 1935 Rhodacarellus subterraneus Willmann

Minirhodacarellus Shcherbak, 1980 Rhodacarellus minimus Karg, 1961

Afrogamasellus Loots & Ryke, 1968 Cyrtolaelaps (Gamasellus) franzi Loots & Ryke, 1966

Afrodacarellus Hurlbutt, 1973 Afrodacarellus femoratus Hurlbutt Table 5. (Continued)

Subfamily Dendrolaelapinae Hirschmann, 1960

Type Genus: Dendrolaelaps Halbert, 1915 Type species: sensu Hirschmann, 1960

Dendrolaelaps Halbert, 1915 - containing 2 subgenera:

D. (Dendrolaelaps) Halbert, 1915 _s^ str. Dendrolaelaps oudemansi Halbert. 1915

D. (Longoseiulus) Lindquist. 1975 Dendrolaelaps longulus Hirschmann. 1960

Dendrolaelaspis Lindquist, 1975 Digamasellus angulosus Willmann, 1936 (non Dendrolaelaps angulosus sensu Hirschmann, 1960)

Longoseius Chant, 1961 Longoseius cuniculus Chant

Dendroseius Karg, 1965 Digamasellus reticulatus Sheals, 1956

Multidendrolaelaps Hirschmann, 1974 Dendrolaelaps ulmi Hirschmann. 1960 (= Dendrolaelaps (Multidendrolaelaps) Hirschmann, 1974 part)

Orientolaelaps Bregetova & Shcherbak, 1977 Orientolaelaps eutamiasi Bregetova & Shcherbak

Insectolaelaps Shcherbak, 1980 Insectolaelaps armatus Hirschmann, 1960

Oligodentatus Shcherbak, 1980 Oligodentatus tridentatus N3 Shcherbak & Bregetova U 1 MATERIALS AND METHODS

Materials;

This study was first initiated with the sole objective of monophyletically defining the subfamily Ologamasinae, and determining its internal relationships and its relationships with the other subfamilies of the Rhodacaridae sensu Lee (1970). I was particularly interested in the Ologamasinae for three reasons: 1) its representatives display such a variety of forms, sizes and degrees of idiosomal sclerotization that, judging from their overall appearance,

they seem to share characteristics with each of the other rhodacarid

subfamilies; 2) they are found in many habitats; and, 3) the majority

of its genera is found only in the Southern Hemisphere (I plan to

devote a great deal of time studying mesostigmatid mites in

America). Soon after starting to examine specimens, I found that in

order to do an adequate study of the Ologamasinae using phylogenetic

methods, it would be necessary to include not only all subfamilies of

Rhodacaridae sensu Lee, but also include other taxa (families or

genera) considered to have some degree of relationship with the

family, so that sister-group relationships could be determined with

more confidence. For an outline of the procedures used, refer to

"Methods”.

26 27

All of the material exeunined in this study was from museums and private collections. Listed below are the persons who kindly made specimens available, followed by their institutions:

Dr. T. K. Crosby and Dr. Graeme W. Ramsay, Entomology Division,

D.S.I.R., Auckland (DSIR); Dr. Valerie , Queensland Museum,

Queensland (QM); Dr. Robert Domrow, Queensland Institute of Medical

Research, Brisbane (QIMR); Dr. H. Fechter, Zoologische Staatssammlung

München, Munich (ZSM); Dr. Keith S. Hyatt, Ms. Anne Baker and Mr.

Kieran P. Martyn, British Museum (Natural History), London (BMNH);

Dr. John Kethley, Field Museum of Natural History (FMNH); Dr. Donald

E. Johnston, Acarology Laboratory, The Ohio State University,

Columbus (ALOSU); Dr. Wolfgang Karg, Institut fur

Pflanzenschutzforschung, Kleinmachnow, Berlin (KARG); Dr. David C.

Lee, South Australian Museum, Adelaide (SAM); Dr. Evert E. Lindquist,

Biosystematics Research Institute, Canadian National Collection,

Ottawa (CNC); Dr. Sandor Mahunka, Zoologische Abteilung des

Ungarischen Naturwissenchaftlichen Museums, Budapest (UNMB); Dr.

Henri Naudo, Museum National d'Histoire Naturelle, Paris (MNHN); Dr.

Fausta Pegazzano, Istituto Sperimentale per la Zoologia Agraria,

Florence (ISZAF); Dr. F. Puylaert, Musee Royal de l’Afrique Centrale,

Tervuren (MRAC); Dr. Karel Samsinak, Institute of Parasitology,

Czechoslovakian Academy of Sciences, Prague (CAS); Dr. Jo Ann

Tenorio, Bernice Pauahi Bishop Museum, Honolulu (BPBM).

Efforts to borrow specimens from the Potchefstroom University

Collection in South Africa (PUSA) were unsuccessful. As a

consequence of this, I was unable to examine the following taxa: 28

Acugamasus natalensls species complex of Lee, 1970; Gamaselliphis

Ryke, 1961a (all species except for a specimen labelled "nr. potchefstroomensis" from the Hurlbutt Collection, deposited at

ALOSU), Gamasellevans Loots and Ryke, 1967; Gamasellopsis Loots and

Ryke, 1966; Notogamasellus (Notogamasellus) Loots and Ryke, 1966;

Notogamasellus (Podonotogamasellus) Loots and Ryke, 1966;

Paragamasellevans Loots and Ryke, 1968 and Rvkellus Lee, 1970. For these, it was necessary to resort to literature-based data.

It was not possible to collect all data necessary to characterize some of the genera and species examined, for several reasons: the specimens available for study were poorly mounted (and I had no permission to remount them) or, broken into several pieces or, some body part was missing or, one of the sexes was not known, so that a complete account of morphological characteristics could not be made. These taxa included the genera Gamasitus Womersley, 1956a,

Laelogamasus Berlese, 1905, Onchogamasus Womersley, 1956, and the following species: Afrogamasellus citri Loots. 1969 sensu Hurlbutt

(1973); Afrogamasellus squamosus Karg, 1977; Afrogamasellus furculatus Karg, 1979: Neogamasellevans berlesei Womersley. 1956b;

Rhodacaroides minvaspis Lee, 1973 and Rhodacarus costai Sheals, 1962.

A total of 1232 specimens representing 293 species and 60 genera was examined; when necessary (and permission was granted), a number

of preparations was remounted.

Methods :

Specimens were examined and illustrated with the aid of a Wild

Heerbrug compound microscope with phase contrast microscopy, equipped 29 with camera lucida, 15x wide field oculars and lOx, 20x, 40x bright field, 40x phase contrast, lOOx bright field and lOOx phase contrast objectives, providing a range between 150 - 1500x. Measurements were made with a 1mm scale (100 divisions of 0.01mm), using the Wild

Heerbrug M20. Dissecting and mounting were accomplished with an

Olympus stereoscopic microscope equipped with lOx oculars and 0.7x -

4x objectives, providing a range between 7x-40x.

Slides containing specimens to be remounted were soaked in distilled water for 3 to 4 hours (to loosen mites from under the coverslip), then transferred to a 1:1 mixture of 70% ethyl alcohol and Nesbitt's fluid for 24-48 hours, at room temperature (Nesbitt's effectively macerates internal tissues with little or no harm to the exoskeleton). After clearing, specimens were dissected and mounted in Berlese's medium. Preparations were allowed to dry completely

(three to five weeks at 50 degrees Celsius) before being sealed with

Glyptal, a non-soluble protective paint used to prevent moisture uptake by aqueous media such as Berlese's.

The terminology of morphological structures was that used by

Evans and Till (1965) and Lee (1970) in part; idiosomal chaetotaxy, that of Lindquist and Evans (1965); palpal chaetotaxy, that of Evans

(1964); leg chaetotaxy followed Evans (1963, 1969); and finally, adenotaxy was that of Athias-Henriot (1969a).

Selection of material:

The material utilized for outgroup comparisons was extracted from the Digamasellidae Evans, 1957 (sensu Lindquist, 1975), the

Veigaiaidae Oudemans, 1939 and the Ascidae Voigts & Oudemans, for the 30 following reasons:

The current placement of the Ologamasidae Ryke sensu Krantz,

1978, is in the superfamily Rhodacaroidea which also includes the

Rhodacaridae sensu Johnston. 1968 (= Rhodacaridae sensu stricto) and the Digamasellidae sensu Lindquist.

Lindquist (1975) pointed out the close relationships of the

Digamasellidae with the Rhodacaridae sensu stricto and of both families with the Cyrtoloaelapidae sensu Johnston, 1968 (=

Ologamasidae sensu Krantz, 1978, in part).

To Shcherbak (1979, 1980, 1982, 1983), the Rhodacaridae sensu stricto should include all of the nominal genera placed in the

Digamasellidae sensu Lindquist (1975) except for the genus

Digamasellus.

Since the Digamasellidae sensu Lindquist includes taxa regarded as more derived than taxa of the Rhodacaridae and Ologamasidae, I selected two of its genera perceived as more "generalized" and which illustrate different conditions concerning the development of scleronoduli (a very common feature in rhodacarids and digamasellids): Dendroseius which, according to Lindquist (1975), resembles the ancestral stock and possesses the "normal set" of four scleronoduli on the podonotal area of the divided dorsal shield; and

Digamasellus. the only representative of the family which has secondarily lost all four scleronoduli and autapomorphically developed enlarged anal valves. Also examined (for the sake of general comparisons within the Digamasellidae) were representatives of Dendrolaelaps and Longoseius (ALOSU collections). 31

 Parasitidae-Velgalaidae relationship to the Rhodacaridae sensu latu (= Rhodacaroidea) has been suggested by Lee (1970) and Lindquist

(1975). Because Parasitidae are devoid of a truly functional spermadactyl and are not podospermal (whereas all Dermanyssina are),

I did not include them as a potential outgroup. From the

Veigaiaidae, Veigaia Oudemans, 1905 and Berlese, 1903 were included.

Some of the taxa with controversial taxonomic placement have been referred to another dermanyssine family - the Ascidae - by several workers. Examples include Gamasellodes Athias-Henriot, 1961 and Protogamasellus Karg (Lindquist & Evans, 1965); Panteniphis

Willmann (Lee, 1970; Bregetova, 1977) and Lindquistoseius Genis,

Loots & Ryke, 1969. Therefore, two genera from the subfamily

Ascinae, Lasioseius Berlese, 1916 (tribe Blattisocini) and

Proctolaelaps Berlese, 1923 (tribe Melicharini) were chosen as the most generalized representatives to be compared with Gamasellodes and

Protogamasellus (placed in the tribe Ascini by Lindquist & Evans,

1965), and with Panteniphis and Lindquistoseius as well.

Character selection;

The characters selected for this study were based upon examination of many cuticular structures used by by Loots (1967), Lee

(1970) and Lindquist (1975), which appeared to exhibit the least variation up to the intrageneric level and therefore, potentially valuable in the elucidation of generic relationships. Some of these structures included: the site of the spermathecal opening, the female and male cheliceral features, the degree of fusion of shields, the 32 development of scleronoduli, and the chaetotactic pattern of body dorsum, venter and legs. In addition, characters not previously utilized in comparative studies of the Rhodacaridae sensu lato were examined and included. Some of these were: the structure of the podonotal glands gd4 and ventral glands gv2; the placement of the male genital setae (st5); the presence, number and form of the preendopodal shields; and the relative positions of ventral setae Jvl and Zvl. Structures subject to interspecific variation were also used in estimating sister species relationships.

Analytical methods:

With the objective of producing a classification that would reflect genealogical affinities and starting with the assumption that the groups under study represented monophyletic lineages (that is, they shared recency of common ancestry), my efforts concentrated into testing these hypotheses through the use of phylogenetic sysyematics in the sense of Hennig (1965, 1966). A series of independent homologous characters held in common by some or all members of the studied groups was identified and their direction of change

(character polarity) was postulated, on the basis of a combination of three criteria:

Outgroup comparisons: the character state most often found among members of the group under study and among members of the presumably related group (outgroup), was considered primitive (plesiomorphic).

Character correlation: when plesiomorphy or apomorphy of a

character state could not postulated by outgroup comparison, its

polarity was inferred by correlation with the transformation series 33 of other character states already determined by outgroup comparison.

Commonality: where a character was found to occur only in the group under study and not in the presumably related group (outgroup) and where the the direction of change could not be postulated by correlation, the most frequent state was considered plesiomorphic as its "commonness" was unlikely due to convergence (the commonality principle of Schaeffer et al., 1972). •

Analysis of the polarized character states was performed by using Farris' computerized algorithm "Wagner 78". The reason for the choice of this particular algorithm stems from the fact that it operates under the principle of parsimony, combined with phylogenetic methods and therefore, produces trees based upon shared derived character states (synapomorphies) with the smallest number of character transformations. Because it was not possible to obtain all data for several species (in cases where males were not known, for example), it was necessary to analyze various combinations of data matrices. From these, a total of 105 computer-generated cladograms was produced with each carefully evaluated regarding the distribution of character states, total homoplasy, deviation ratio and tree length. From the most parsimonious trees, six consensus cladograms were constructed to illustrate suprafamilial and familial relationships. CHARACTER ANALYSIS

The direction of change of some of the characters revealed most useful in the elucidation of familial relationships is briefly discussed below. Throughout the text, the concepts of Rhodacaridae and Digamasellidae are respectively, those of Lee (1970, 1973 with the inclusion of the genus Afrodacarellus Hurlbutt, 1973), and

Lindquist (1975). Terms referring to primitive (least derived, plesiomorphic) and derived (most derived, apomorphic) conditions, are used interchangeably. Character numbers are given between parentheses

(and see Table 6).

Postanal seta (1);

A long postanal seta (at least one and a half times as long as the adanal setae) is primitive; in the opinion of Bregetova (1974), it represents "an archaic character, a modified telson" which is still observed "in some larvae of Mesostigmata including

Eurvparasitus emarginatus Koch, 1839 and many Parasitidae". In some adults of the presumably related groups (the outgroups), this primitive state is found (Gamasellodes. Protogamasellus). and several

taxa of the Rhodacaridae exhibit this condition as well:

Tangaroellus. Rhodacarus. Rhodacaropsis. Rhodacarellus. Litogamasus

Lee, 1970, Parasitiphis Womersley, 1956, Hvdrogamasus Berlese, 1892,

34 35

Periseius Womersley, 1961 and Rhodacaroides; however, the apomorphic trend for a gradually shortened postanal seta is observed throughout the family. In the Digamasellidae, Laelaptonyssinae and the incertae sedis genera Panteniphis and Lindquistoseius. the most derived condition is observed (where the adanals are longer than the postanal seta).

Scleronoduli (5):

Scleronoduli are subsurface (apodemal) cuticular structures

(usually four in number), shaped like bulges (nodules) that project into the body cavity, and to which longitudinal intertergal muscles are attached; they are located on the podonotal shield within a dorsal hexagonal area comprising setae j5, z5 and j6; from them, several muscle bundles radiate running posteriad to attach to the anterior margin of the opisthonotal shield. The association between these structures and the muscle bundles, makes up a special muscular apparatus which provides flexibility to the opisthosoma.

Shcherbak and Akimov (1974) studied this muscular apparatus in three genera: Rhodacarus. (with 3 scleronoduli), Rhodacarellus and

Dendrolaelaps (each with 4 scleronoduli) and found that, regardless of whether there are 3 or 4 scleronoduli present, the total number of muscle bundles involved is the same (twelve) and that the regrouping of muscles is determined by the number of scleronoduli and is directly connected with the different types of opisthosomatic movement observed in the different genera. It seems very likely that scleronoduli have evolved to fit certain living conditions - perhaps the mobility of the opisthosoma is particularly useful as the 36

moves about the narrow pore spaces of deeper soil layers or the bark

of trees, for instance.

The ocurrence of scleronoduli seems restricted to certain

groups; it is commonly found among Rhodacarinae and Digamasellidae,

but other taxa of Rhodacaridae also possess them: Solugamasus Lee,

1970; Paragamasellevans Loots & Ryke, 1968; several species of the

genus Gamasellus Berlese, 1892; Hydrogamasellus racovitzai

Trouessart, 1903 and Hiniphis hinnus Lee. 1970 are some of them;

also, Protogamasellus mica Athias-Henriot 1961 (currently placed in

the Ascidae), possesses 3 scleronoduli on its dorsal hexagonal area

in the same way that Rhodacarus and Rhodacaropsis do.

For the polarization of this character, I regarded the absence

of scleronoduli as plesiomorphic (on the basis of outgroup comparison); the direction of change followed the commonality

principle, with the gradual decrease from four to two scleronoduli representing a higher degree of apomorphy.

Palp claw (6):

A palp claw with three well developed tines is predominantly

found in Rhodacaridae and ; in the rhodacarid genera

Heterogamasus and Evanssellus however, the basal tine is very

reduced, almost vestigial, whereas a two-tined palp claw is found

throughout the Digamasellidae, the Ascidae, and in Panteniphis and

Lindquistoseius. I regard the gradual loss of the basalmost tine as

apomorphic, on the basis of outgroup comparison.

Female metasternal setae (7):

The metasternal setae (st4) are commonly enclosed on metasternal 37 platelets in the females of Veigaiidae, Ascidae and several other groups; this condition is plesiomorphic. The tendency towards the fusion of these platelets with the sternal shield is apomorphic, regardless of whether the fusion attained forms a completely well

sclerotized sternal shield or, a shield with punctate sclerotization at its posterior margin; also apomorphic is the condition where the

platelets are lost and the metasternal setae lie on the soft cuticle

immediately posterior to the sternal shield.

Preendopodal shields (14);

Preendopodals are placed anteriorly to the sternal shield; they

are usually small platelets produced into a weakly or well

sclerotized pair and, in this form, are commonly found in the

Rhodacaridae and related groups. I regard this state as the least

derived one in a transition series from which two lineages

originated. On the one hand, this pair of plates becomes incorporated

to the sternal shield or is lost; on the other hand the plates may

become fragmented into two, three or more pairs.

Female peritrematal shields (17):

Two states are observed relative to this character: the shields

are separated from the exopodals around leg IV or, they are connected

(fused) to them. The first condition is the most often found in the

outgroups - Veigaiidae, Ascidae, Digamasellidae - and several

rhodacarid taxa: Afrogamasellus. Paragamasellevans. Rhodacaropsis.

Rhodacarus. Rhodacarellus. Tangaroellus. Neogamasellevans.

Antennolaelaps Womersley, 1956, Sessiluncus Canestrini, 1898, Epiphis

Berlese, 1916 and Laelaptonyssinae. However, there is a strong trend 38 toward the fusion between peritrematals and exopodals IV throughout the Rhodacaridae; the incertae sedis Panteniphis and Lindquistoseius exhibit this state as well. On the basis of outgroup comparison, the first condition (lack of fusion of the shields) is considered primitive.

Podonotal setae zl (22);

Relative to setae jl, this pair of podonotal setae is most often produced as a much shorter one. This state is commonplace in

Veigaiidae, Ascidae, some Digamasellidae and most Rhodacarinae genera; therefore, 1 regard it as the least derived state in the character transformation series. Apparently, two distinct lineages seem to have originated from the primitive state: the first one (and the main trend among rhodacarids) in which zl evolved from subequal in length to longer than jl, and the second one in which zl was lost.

The latter state seems to occur only rarely as in some Gamasellopsis species.

Sternal setae stl (24):

The first sternal setae are placed near the anterior margin of the sternal shield in both males and females of Veigaiidae, Ascidae and the majority of Rhodacaridae (this is also quite common in other

Dermanyssina). 1 regard this condition as the plesiomorphic one.

There are two main trends in the evolution of this character. In

one, the first sternal setae are removed from the sternal shield and are placed on the soft cuticle between the sternal and preendopodal

shields or, they are directly placed on the preendopodal shields. The

first state is found in Lindquistoseius (incertae sedis) and although 39 rare, is less derived than the latter, which occurs in Afrogamasellus

(Rhodacarinae) and a few members of the Ascidae. In the other trend, setae stl are placed on an area of weak sclerotization of the sternal shield; of the taxa exhibiting this state, some are members of the

Rhodacarinae (Afrodacarellus. Rhodacarus. Rhodacarellus),

Gamasellinae (Notogamasellus), Ascidae (Gamasellodes. some

Protogamasellus species), and most members of Digamasellidae. It is interesting that this character state - and a few others relating to areas of weak sclerotization in different shields - is observed in species that tend to inhabit confined spaces such as deeper soil layers or spaces under bark. It is very likely that at least some of these characters may be correlated and represent adaptations that enable these species to move more freely in these confined spaces.

Podonotal region (25);

The anterior half of the dorsum of the idiosoma is referred to as podonotum (the case where an entire dorsal shield is present) or podonotal shield (where the dorsal shield is subdivided ). Since both entire and divided dorsal shields occur in Rhodacaridae, for practical purposes this anterior half will be referred to as podonotal region. There are four longitudinal rows of setae referred to as j, z, s and r series and the holotrichous chaetotaxy for this region is 24 pairs of setae, (according to the modified setal nomenclature for the idiosoma of Gamasina developed by Lindquist and

Evans in 1965, using the Ascidae as a model); I regard holotrichy as the plesiomorphic condition. The Veigaiidae, Digamasellidae and the majority of Rhodacaridae share an oligotrichous condition where there 40 are between 20 to 23 pairs of podonotal setae. There are a few exceptions, however: Tangaroellus porosus. Laelaptonvssus mitis and the genus Panteniphis exhibit further oligotrichy with 16 to 19 pairs of setae, while this oligotrichy has evolved to a paedomorphic condition in Laelaptonvssus chinensis Samsinak, 1964 which has between 13 to 15 pairs of podonotals (the larval chaetotactic complement is 10 pairs; that of the protonymph is 15 pairs).

On the other hand, the primitive condition is still observed in

Cvrtolaelaps Berlese, 1887 whereas Notogamasellus Loots & Ryke, 1966,

Pillelus Lee, 1970 and Euepicrius Womersley, 1942 developed hypertrichy with more than 25 pairs of podonotals present.

Opisthonotal region (26):

This dorsal posterior half of the idiosoma has a holotrichous condition of 28 pairs of setae, usually distributed as 5J, 5Z, 5S, 7R and 6UR, according to Lindquist and Evans. Using the criterion of character correlation I regarded holotrichy as plesiomorphic. As in character 25, it appears that two distinct lineages evolved from holotrichy: one towards hypertrichy and the other towards oligotrichy, with further development of paedomorphic suppression of setae where there are between 11 to 15 pairs of opisthonotals (here again, the larval complement is 10 pairs and the protonymphal complement is 15 pairs).

Arthrodial process of female chelicera (27):

A membranous arthrodial region, located at the base of the movable digit of the chelicera, enables the movable digit to articulate ventrally to the fixed digit. This is accomplished with 41 the aid of two condyles developed on the limb of the fixed digit, which engage acetabula on the movable digit (Evans and Till, 1965).

This arthrodial membrane is usually produced into setiform processes collectively referred to as "corona" and this condition is observed in most Rhodacaridae and Veigaiidae; I consider this state plesiomorphic. The evolution of this character produced two trends: the corona and setiform processes are progressively reduced (as seen in many Ascidae and Digamasellidae) and eventually, the entire arthrodial process was lost (as it occurs in Laelaptonyssinae) or, the setiform processes became elongate, developing what is called an

"arthrodial brush". The arthrodial brush is typically found in the

Rhodacarinae genus Afrogamasellus: in Latogamasellus (which Karg regards as a subgenus of Afrogamasellus); in the genus Afrodacarellus referred to the Rhodacaridae sensu stricto by Hurlbutt and in the genus Lindquistoseius.

Pretarsus I (30):

Like the pretarsi of the other legs, the pretarsus of leg I bears two claws and bilobed pulvilli but in addition, is usually set on a long peduncle in Veigaiidae, Ascidae and several members of

Rhodacaridae; based on outgroup comparison, this state is considered

plesiomorphic. There is a unidirectional trend concerning the

transformation of this character: first, the pretarsal peduncle is

shortened (this is the state most commonly observed among

Rhodacaridae); then, the peduncle is eventually lost, giving the

pretarsus the "effect" of sessile claws (this state is found in

several Digamasellidae and Rhodacaridae taxa). Ultimately, the 42 pretarsus is lost; this condition occurs in Rhodacarus

(Rhodacarinae); Euepicrius (Gamasiphinae); Evanssellus Ryke. 1961,

Notogamasellus and Solugamasus (Gamasellini).

Submarginal (UR) series (38):

The fifth longitudinal series of setae placed ventrolaterally on the opisthosoma is referred to as "submarginals" following the setal nomenclature of Lindquist and Evans. The authors noted that these setae are "regularly paired, and positioned and do not constitute hypertrichy" contrary to what Hirschmann (1957) thought when he attributed "Rx" signatures to them. According to Lindquist and Evans, the full complement of submarginals is 6 pairs, but there may be 0-15 pairs in the Ascidae. In Veigaiidae and some Rhodacaridae

(Cvrtolaelaps. Pyriphis Lee. 1970), there are 7 pairs and species of the Gamasellus discutatus complex of Lee, 1970 may have 6, 7 or 9 pairs of submarginals. Also, 4 or more pairs are found in Caliphis

Lee, 1970 (Gamasiphinae), Epiphis. Eurvparasitus Oudemans, 1902 and

Pilellus (Gamasellini). On the other hand, submarginals are absent in

Digamasellidae, Panteniphis. Lindquistoseius and many Rhodacaridae taxa.

The direction of change for this character was relatively simple

to ascertain, since there is a definite trend towards the loss of

submarginals. However, it was difficult to establish limits (states)

relating the number of UR setae with the degree of aporaorphy. After

examination of all species it was possible to notice three patterns:

the first and the most primitive, was that in which species tended to

possess at least 4 pairs (and up to 9 pairs); the second was found in 43 species which possessed at least 1 pair, but never more than 3; the third and most derived, was seen in species which completely lost the

UR series.

Femur _! (39):

According to Evans (1963), the most widely distributed chaetotactic pattern for femur I of free-living Gamasina is 13 setae

(including 4 ventrals); the Veigaiidae, Digamasellidae and most

Rhodacaridae (among other families) exhibit this plesiomorphic pattern. In the Ascidae, Lindquistoseius. Panteniphis. and

Tangaroellus porosus Luxton, 1968 (Tangaroellinae), this number is reduced to 12 setae, while in the Laelaptonyssinae there is further reduction to 11 setae, this being uniquely derived within

Rhodacaridae. The gradual loss of setae is regarded as derived.

Genu I_ (42) :

The full complement for genu I is 13 setae and this is found

(among other groups) in Veigaiidae, Ascidae and most Rhodacaridae, except for Laelaptonvssus chinensis and Gamasellopsis Loots & Ryke,

1966 which possess 12 setae. This derived state (12 setae) is observed in Digamasellidae and Panteniphis as well, while

Lindquistoseius has 11 setae.

Genu III (43):

This segment bears 9 setae (including 2 ventrals) in Veigaiidae,

Rhodacaridae and Ascidae, with a few exceptions in the latter two families; I regard this complement as plesiomorphic, even though the full set for this segment is 10 setae (which occur in families not directly related to the groups here studied). Apomorphic conditions 4 4 concern the reduction to 8 setae (loss of 1 ventral) observed in some

Ascidae, some Digamasellidae, Gamasellopsis. Lindquistoseius.

Panteniphis and Laelaptonvssus chinensis. Another apomorphic state refers to an apparent neotrichy, where an extra seta was added to the genua of Heterogamasus. Notogamasellus and Periseius. all members of the Gamasellini.

Genu IV (44):

The genua of leg IV bear a maximum of 10 setae, including 2 ventrals. This is the plesiomorphic pattern which is present in

Veigaiidae and most Rhodacaridae; the exceptions, Sessiluncus.

Gamasiphis Berlese, 1904, Gamasiphoides. Hvdrogamasus. Gamasitus and

Gamasellopsis have lost 1 or 2 setae on this segment, a derived condition exhibited by the Ascidae as well. In the Digamasellidae there are 7 setae, while in Panteniphis and Lindquistoseius there is a maximum of 6 setae, a state rarely found among free-living Gamasina except for some species of a non-related genus of Macrochelidae

(Macrocheles Latreille. 1829).

Tibia I (45):

The full complement of tibia I is 14 setae, which includes 4 ventrals. This pattern is observed in Veigaiidae and all

Rhodacaridae, except for the Laelaptonyssinae which possess 9 (L. mitis) or 11 (L. chinensis) setae. In the Ascidae there are 13 setae, whereas in the Digamasellidae there is further reduction to 12.

Panteniphis also possesses 12 setae and Lindquistoseius has 11 setae.

Taking into account the criteria of outgroup comparison and character correlation, the progressive loss of setae is apomorphic. 45

Tibia IV (47);

Evans (1963) observed six types of chaetotactic patterns for this segment, the number of setae ranging from 6 to 10. On the basis of outgroup comparison, character correlation and commonality principle, I regard as plesiomorphic the condition where 10 setae are present.

The full set of tibial setae is found in Veigaiidae, most Ascidae and most Rhodacaridae. Exceptions to the Ascidae are Protogamasellus species and to the Rhodacaridae, the genera Hvdrogamasus and

Gamasellopsis (each of these with 9 setae), and Laelaptonyssinae which possess 7 or 8 setae. Digamasellidae also have 7 setae, while

Panteniphis and Lindquistoseius uniquely share 6 setae.

Dorsum of the idiosoma (49):

The dorsum of the idiosoma is commonly divided into two shields

(a podonotal and an opisthonotal) in Veigaiidae, Digamasellidae, several Ascidae and many Rhodacaridae taxa (Rhodacarinae,

Gamasellini). Based on outgroup comparison, I regard this state as plesiomorphic. A number of different conditions can be seen

throughout the Rhodacaridae, but they are probably all derived from the basic pattern of divided shields. For example, certain taxa are

characterized by divided shields in both sexes but occasionaly,

females of certain species may have a holonotal shield; this is the

case for Geogamasus. In some genera the shield is usually entire in

both sexes, but the male may have it divided; this is the case for

some species of Athiasella Lee, 1973, Hvdrogamasellus. Gamasellus

discutatus complex and Lindquistoseius. The shields tend to 46 eventually coalesce and evidence for this is provided by taxa which have entire shields in both sexes, but a suture or, at least a line of demarcation, across the shield can still be seen, indicating the site of fusion; this condition occurs in Rykellus Lee, 1970,

Gamasiphoides and Hiniphis hinnus. A holonotal shield is the most derived state and it is found in several genera of Rhodacaridae, particularly in the Ologamasini, Gamasiphinae and Sessiluncinae.

Female genital shield (54);

A slightly longer than wide genital shield is the state most

commonly found among females of Veigaiidae, many Digamasellidae and

Rhodacaridae taxa; by outgroup comparison, this was considered the

plesiomorphic condition, with two lineages evolving from this basic

shape. In one, the genital shield became very elongate, at least one

and a half times as long as wide (perhaps as a correlation with a

narrow body). This seems to be the case for Longoseius and

Longoseiulus (Digamasellidae); Protogamasellus and Gamasellodes

(Ascidae); Afrodacarellus. Litogamasus. Solugamasus and

Paragamasellevans (Rhodacaridae). In the other lineage, evolution

followed the opposite direction, with the genital shield becoming as

wide as long or even wider than long; this is the main trend among

members of the Rhodacaridae, especially the Ologamasinae,

Gamasiphinae and Sessiluncinae. Panteniphis and Lindquistoseius also

exhibit this character state.

Opisthogastric setae Zvl (55):

The common position for setae Zvl is at a level parallel to or

slightly above that of Jvl. This is observed in Veigaiidae, 4 7

Digamasellidae, several members of Ascidae and Rhodacaridae. Here also, two lineages evolved from the plesiomorphic condition. On the one hand, Zvl migrated upwards reaching a position well above Jvl, nearly vertical to it; on the other hand, Zvl migrated downwards to a level below Jvl; ultimately, Zvl was lost, a condition observed only in Laelaptonvssus mitis Womersley, 1956, Panteniphis and

Lindquistoseius. It should be noticed that these taxa also exhibit overall hypotrichy.

Female exopodal shields III (59):

The exopodal plates around the acetabula of legs III are rudimentarily developed, not completely encasing the circumaxial edges of these acetabula in females of Veigaiidae, Digamasellidae, several Ascidae and most Rhodacaridae. There seems to be a unidirectional trend towards the gradual increase in the size of these shields so that, they eventually become united forming an entire sclerite around the acetabula. This is considered the most derived state and is observed in some Sessiluncinae, Gamasiphinae, one Ologamasinae genus (Rvkellus). some Ascidae and in Panteniphis and Lindquistoseius.

Site of the spermathecal opening (61);

Among podospermal Gamasina, the most primitive site for the opening to the spermathecae is in the region of acetabula IV. This is found in Veigaiidae, many Rhodacaridae genera, some Digamasellidae and Ascidae. It appears that there is a tendency for the site of the accessory opening to migrate anteriad, first to the basal segments of legs IV or III (coxae), then to a more distal position on leg III 48

(trochanter or femur). A spermathecal tube opening in the region of acetabulum IV is common among Rhodacarinae, Gamasiphinae and

Gamasellini. The spermathecal opening on coxa IV is the second most common site and it occurs in Ologamasini, some Sessiluncinae, in

Tangaroellus. some Digamasellidae and Panteniphis. The acetabulum, coxa, trochanter and femur of leg III are less common sites for the spermathecal opening; the last condition is found in many

Digamasellidae, in the genus Rhodacarus. and in Lindquistoseius. On the basis on the criteria of outgroup comparison and commonality principle, I regard the site for the spermathecal opening associated with the acetabulum of leg IV as the plesiomorphic state, followed by coxa IV, coxa III, trochanter III and femur III.

The direction of change of the sixty- one characters utilized in this study is listed on Table 6. State ”0” indicates the plesiomorphic condition followed by sequential increase in apomorphy.

Negative values in the transition series of a character indicate evolution of distinct lineages relative to that character. 49

Table 6. Character Polarization

Character ...... Description Character state

1. Postanal seta: At least one and a half times as long as the adanal setae. . . 0 Only slightly longer than adanals...... 1 Shorter than adanals ...... 2

2. Spermadactyl (length of): Shorter than or, subequal in length to the movable digit of chelicera...... 0 Less than one and a half times longer than movable digit . . . 1 At least one and a half times longer than movable digit. . . . 2

3. Spermadactyl (degree of fusion): Fused to movable digit throughout most of its length...... 0 Freer than fused to movable digit...... 1

4. Spermadactyl (shape of): Generally straight throughout its length ...... 0 Recurved at basal or median level of movable digit before projecting freely or curving at its tip...... 1 Recurved at distal level of movable digit before projecting freely ...... 2 Curving around movable digit...... -1

5. Scleronoduli: Absent from the dorsal hexagonal area (dha) of the podonotal shield ...... 0 Present on dha, four (04) in number...... 1 Present, three (03) in number (with median pair fused) .... 2 Present, two (02) in number...... 3

6. Palp claw: With three tines ...... 0 With basal tine reduced in size...... 1 With two tines...... 2

7. Female metasternal setae: Placed on metasternal plates ...... 0 Placed on well sclerotized sternal shield...... 1 Placed on stippled cuticle of the sternal shield (area of weak sclerotization)...... 2 Placed on the integument, posteriorly to the sternal shield. .-1 50

Table 6. (Continued)

8. Male genital setae (st5): Placed on sternogenital shield ...... 0 Placed on separate platelets (or on endopodals IV) ...... 1 Placed outside the sternogenital shield, on the integument . . 2 Placed posteriad, on ventrianal shield ...... 3 Absent...... -1

9. Posterior margin of female genital shield: Extending posteriorly, well beyond acetabulum IV ...... 0 Not extending well beyond acetabulum IV...... 1

10. Adanal setae: Placed at level of posterior margin of the anal valves .... 0 Placed at mid-level of anal valves ...... 1 Placed at level anterior or above anal valves...... 2

11. Female ventri-anal shield: Not fused to exopodal IV shields ...... 0 Fused to exopodal IV shields...... 1

12. Male sternogenital shield: Not fused to ventri-anal shield...... 0 Fused to ventri-anal shield...... 1

13. Podonotal setae jl: Always simple (at most, inconspicuously pilose)...... 0 Usually simple, although may be barbed at times...... 1 Always barbed or strongly pilose ...... 2 Pilose and spatulate ...... 3 Modified into a strong spine...... -1 Modified into a wide,leaf-shaped process...... -2

14. Preendopodal shields: Present, as a weakly or well-sclerotized pair...... 0 Produced into two (02) pairs...... 1 Produced into three (03) or more pairs...... 2 Incorporated (fused) to the sternal shield...... -1 Absent...... -2

15. Female ventri-anal shield: Not fused to notai shield...... 0 Fused to notai shield...... 1

16. Female peritrematal shield: Not fused to ventrianal shield ...... 0 Fused to ventrianal shield ...... 1 51

Table 6. (Continued)

17. Female peritrematal shields: Not fused to exopodal IV shields ...... 0 Fused to exopodal IV shields...... 1

18. Female sternal shield: Entirely well defined, regardless of whether or not it is well sclerotized...... 0 Weakly defined (with an area of granulate cuticle at its anterior or posterior margins, or both ...... 1

19. Ventral gland gv2: Present, as three or more pores...... 0 Present, as one or two pores at m o s t ...... 1 Absent or not discernible...... 2

20. Anal valves: Not enlarged...... 0 Enlarged ...... 1

21. Dorsal gland gd4: Not enlarged (present as a small, pore-size structure) .... 0 Slightly larger than a pore...... 1 Enlarged (produced into a conspicuously developed structure) . 2

22. Podonotal setae zl: At least one and a half times shorter than setae jl...... 0 Only slightly shorter than jl...... 1 Longer than j l ...... 2 Absent...... -1

23. Opisthonotal setae Z5: At least one and a half times as long as setae J5...... 0 Slightly longer or, subequal in lenght to J5 ...... 1

24. Sternal setae stl: Placed on sternal shield ...... 0 Placed between pre-endopodals and sternal shield ...... 1 Placed on pre-endopodals...... 2 Placed on area of granulate sclerotization (stippled cuticle) of sternal shield...... -1

25. Podonotal region of the idiosoma: Hypertrichous, with 25 or more pairs of setae...... -1 Holotrichous, with 24 pairs of setae ...... 0 Oligotrichous, with 20 to 23 pairs of setae...... 1 Oligotrichous, with 16 to 19 pairs of setae...... 2 Paedomorphic, with 13 to 15 pairs of setae...... 3 52

Table 6. (Continued)

26. Opisthonotal region of the idiosoma: Hypertrichous, with 29 or more pairs of setae...... -1 Holotrichous, with 28 pairs of setae ...... 0 Oligotrichous, with 23 to 27 pairs of setae...... 1 Oligotrichous, with 20 to 22 pairs of setae...... 2 Oligotrichous, with 16 to 19 pairs of setae...... 3 Paedomorphic, with 13 to 15 pairs of setae...... 4

27. Arthrodial process of female chelicerae: Produced into a conspicuous corona ...... 0 Corona with reduced fringes...... 1 Arthrodial process absent...... 2 Process produced into a conspicuous brush ...... -1

28. Podonotal area: Entire ...... 0 With a partial or complete line of incision...... 1

29. Dorsal setae of the idiosoma: Always simple (setiform or at most, inconspicuously pilose). . 0 Predominantly simple, except for a few complex setae distributed throughout the podonotum and opisthonotum...... 1 Predominantly complex (strongly pilose, barbed, spatulate or pilose-spatulate...... 2

30. Pretarsus I : Present, set on a long peduncle...... 0 Present, set on a shortened peduncule...... 1 Present, but not pedunculate (sessile) ...... 2 Absent ...... 3

31. Number of opisthogastric setae: Ten pairs (holotrichous condition) ...... 0 Eight or nine pairs...... 1 Seven pairs...... 2 Six pairs...... 3 Five pairs...... 4 Four pairs (paedomorphic condition)...... 5 Eleven or more pairs...... -1

32. Female tectum: With central tine undivided...... 0 With central tine bifid or trifid...... 1 53

Table 6, (Continued)

33. Length of peritreme: Normal, i.e. reaching to level of dorsal setae zl or jl. . . . 0 Reduced, reaching to anterior level of acetabulum of leg II. . 1 Reduced, reaching to median level of acetabulum of leg II. . . 2 Extremely reduced, so that only the stigma is present...... 3

34. Female chelicerae; With predominantly small teeth ...... 0 With small and large teeth combined...... 1 With predominantly large teeth ...... 2 Cheliceral dentition reduced, i.e. a maximum of two (small) teeth present...... -1

35. Female podonotal setae placed on soft cuticle (pairs): Three to six pairs...... 0 Three pairs, always...... 1 Usually two pairs...... 2 Usually one p a i r ...... 3 Usually none, but occasionally one pair...... 4 None, always...... 3

36. Female opisthonotal setae placed on soft cuticle (pairs): Usually eight or more pairs...... 0 Usually three to six pairs ...... 1 Two pairs...... 2 Usually one p a i r ...... 3 Usually none, but occasionally one pair...... 4 None, always...... 3

37. Female opisthogastric setae (excluding the circumanals) placed on soft cuticle: At least four pairs...... 0 Three pairs...... 1 Usually two pairs...... 2 Usually one p a i r ...... 3 Usually none, but occasionally one pair...... 4 None, always...... 3

38. Sub-marginal series (UR series); Present, as four or more pairs...... 0 Present, from one to three pairs...... 1 Absent ...... 2

39. Femur I: With 13 setae...... 0 With 12 setae...... 1 With 11 setae...... 2 54

Table 6. (Continued)

40. Femur III; With 6 setae...... 0 With 4 setae...... 1 With 7-8 setae (Neotrichy)...... -1

41. Tarsus IV : With setae pl4 present...... 0 Lacking pl4...... 1

42. Genu I: With 13 setae...... 0 With 12 setae...... 1 With 11 setae...... 2

43. Genu III: With 9 setae...... 0 With 8 setae...... 1 With 7 setae...... 2 With 9-10 setae (neotrichy)...... -1

44. Genu IV: With 10 setae...... 0 With 8 or 9 setae...... 1 With 7 setae...... 2 With 6 setae...... 3

45. Tibia I: With 14 setae...... 0 With 13 setae...... 1 With 12 setae...... 2 With 11 setae...... 3 With 9 setae...... 4

46. Tibia III: With 8 setae...... 0 With 7 setae...... 1 With 6 setae...... 2 With 9 setae (neotrichy)...... -1

47. Tibia IV: With 10 setae...... 0 With 9 setae...... 1 With 8 setae...... 2 With 7 setae...... 3 With 6 setae...... 4 55

Table 6. (Continued)

48. Tarsus IV: With a total of 18 setae...... 0 With 17 setae...... 1 With 16 setae...... 2

49. Dorsum of the idiosoma: Divided on both sexes...... 0 Usually divided on both sexes, but female may have it entire. .1 Usually entire on both sexes, but male may have divided. . . . 2 Entire on both sexes, but with an apparent line of demarcation between the podonotal and opisthonotal regions ...... 3 Entire, with no apparent line of demarcation between the podonotal and opisthonotal regions ...... 4

50. Seta pv2 on tarsus IV: Simple (setiform or inconspicuously pilose)...... 0 Complex (bifid)...... 1 Complex (strongly pilose or with several lateral prongs). . . -1

51. Female tarsus I: With no solenidion half as long as the length of tarsus. . . . 0 With a solenidion half as long as the length of tarsus .... 1

52. Opisthonotal setae J5: Placed at level anterior to Z 5 ...... 0 Placed at level subequal to Z 5 ...... 1 Placed at level posterior to Z5...... 2

53. Femur IV: With 6 setae...... 0 With 7 setae (neotrichy) ...... 2

54. Female genital shield: Slightly longer than wide...... 0 At least one and a half times as long as wide...... 1 As wide as long or wider than l o n g ...... -1

55. Opisthogastric setae Zvl: Placed at a level parallel to or slightly above Jvl...... 0 Placed at level below Jvl...... 1 Absent ...... 2 Placed at level well above Jvl, nearly vertically to it . . . -1 56

Table 6. (Continued)

56. Movable digit of female chelicerae: Usually with 3 teeth...... 0 With 4-8 teeth...... 1 With 9 or more teeth (up to 19)...... 2 With 1-3 teeth at most (rarely edentate)...... -1

57. Fixed digit of female chelicerae: With 3-5 teeth at m o s t ...... 0 With 6-12 teeth...... 1 With 13 teeth or more (up to 1 9 ) ...... 2 With 0-2 teeth...... -1

58. Female sterno-metasternal shield: Entirely separated from endopodal shields IV ...... 0 Partially fused to endopodals IV ...... 1 Fused to endopodals I V ...... 2

59. Female exopodal shields III: Not completely encasing paraxial edges of acetabula III. . . . 0 Not coalesced (split)...... 1 Coalesced (entire) ...... 2

60. Seta all on palp genu: Simple (spiniform or slightly lanceolate)...... 0 Spatulate...... 1 Bifid (with one lateral prong) ...... 2 Trifid (with two lateral prongs) ...... 3 With 5 or more prongs (up to 1 5 ) ...... 4 Finely pilose...... -1

61. Site of spermathecal opening: Associated with acetabula IV ...... 0 On coxae IV...... 1 On coxae or acetabula III...... 2 On trochantera I I I ...... 3 On femora III...... 4 RESULTS AND DISCUSSION

Hypotheses about suprafamilial and interfamilial relationships

(Fig. 1), and intrafamilial relationships (Figs. 2-6) are presented below. Numbers on cladograms indicate stems. Refer to Fig. 1 legend for explanation.

Suprafamilial relationships :

The results of my analysis indicate that the Laelaptonyssoidea

Womersley, 1956 (new status) is the sister group of the Rhodacaroidea

Oudemans, 1902. The two lineages share eight synapomorphies as follows: postanal seta shorter than adanal setae (1); male

spermadactyl freer than fused to movable digit of the chelicera (3);

female metasternal setae placed on the sternal shield (7); adanal

setae placed anteriorly or above level of the anal valves (10);

podonotal setae zl shorter than setae jl (22); podonotal area of the

dorsal shield with a maximum of 23 pairs of setae (25); one pair of

podonotal setae commonly placed on soft cuticle (35) and a maximum of

three pairs of opisthogastric setae placed on soft cuticle (37).

The Laelaptonyssoidea includes the Digamasellidae Evans, 1957

sensu Lindquist, 1975 (= Dendrolaelapinae Hirschmann sensu Shcherbak,

1980 in part), the Panteniphididae n. fam., and the Laelaptonyssidae

Womersley, 1956. The Rhodacaroidea as recognized in this study,

57 58

Includes four families: 1) Rhodacaridae Oudemans 1902 (= Rhodacarinae sensu Lee, 1970), 2) Euryparasitidae n. fam. (= Gamasellini

Hirschmann sensu Lee. 1970 in part), 3) Gamasellidae new status (=

Ologamasini Lee, 1970 in part; Gamasellini Hirschmann sensu Lee, 1970 in part) and 4) Ologamasidae Ryke, 1962 (= Ologamasinae Ryke sensu

Lee, 1970 in part; Gamasiphinae Lee, 1970 in part; Sessiluncinae Lee,

1970 in part), which includes three subfamilies - Ologamasinae Ryke,

1962, Gamasiphinae Lee, 1970 and Epiphidinae Kethley, 1983.

In 1975, Lindquist applied phylogenetic systematica to his studies on the Digamasellidae Evans and concluded that the

Rhodacaridae Oudemans sensu stricto appeared to be its sister group.

According to his findings, the two families comprised a monophyletic group which was equivalent to the Rhodacaridae sensu Karg, (1965,

1971) "excluding the ascid genus Protogamasellus". and he listed the synapomorphies shared by both: the presence of "sclerotic subsurface nodules" (scleronoduli) on the podonotal shield, between setae z5; sternal shield of female weakly sclerotized anteriad of the first pair of sternal pores; male spermadactyl recurved basally before projecting freely anteriad. Among other character states shared by both groups which he felt "unable to judge if apomorphic or plesiomorphic", were: tectum basically triramous; submarginal (UR) series absent; male ventrianal shield not fused to the sternogenital shield, but commonly united posteriorly and laterally with the posterior dorsal shield. Interpreted as parallelisms were: podonotal setae "j2 commonly displaced anteriorly, in transverse row between jl and zl on vertex, leg IV tending to lose pl3 on basitarsus and female' 59 spermatheca with duct commonly opening on leg III".

Lindquist regarded "the group ancestral to the Cyrtolaelapidae"

Csensu Johnston, 1968), as the sister group of the Rhodacaridae and

Digamasellidae, defined by the following synapomorphies; "female

sternal shield with four pairs of setae, due to incorporation of the metasternal plates; female genital shield rounded anteriorly,

truncate posteriorly and separated from a conspicuous ventrianal

shield". Also pointed out were the plesiomorphies that the ancestor

had in common with the "parasitid-veigaiid stock": tibia I with 14

setae, including 4 ventrals; male leg II with large ventral apophyses

especially on the femur (these are modified anteroventral setae);

deutonymph with divided shields whether or not they are separated or

fused in the adult; deutonymph is the phoretic instar in associations

with other .

Lindquist concluded that the Rhodacaridae were more similar to

the ancestral stock than the Digamasellidae, based upon a series of

character states which are found plesiomorphically in the

Rhodacaridae and "other related families (Cyrtolaelapidae,

Parasitidae)". Such character states are: palp claw 3-tined (2-tined

in digamasellids); postanal seta longer than adanal setae (adanals

are longer than the postanal seta in digamasellids - "a derived,

neotenous condition"); deutosternal denticles in 7 rows, none of them

widened (digamasellids with 5, rarely 6 rows of denticles of which

the fifth is usually widened); leg setation of genua and tibiae I in

particular, with the complete set (digamasellids exhibit "neotenous

suppression of some deutonymphal setae"); the 3 tines of the tectal 60 process originate at the same level from the surface of the tectum

(in digamasellids, the median tine occupies a more ventral position than the lateral tines - "a derived condition expressed after the larval instar”); male genital setae (st5) remain on the sternogenital shield - except for the genera Rhodacaropsis and Afrodacarellus -

..."this condition is derived and similar to that of digamasellid males, which have the genital setae on small plates, nearly always separated medially from one another as well as from the sternogenital shield".

My findings partly concur with those of Lindquist and I agree with his general conclusions regarding the Rhodacaridae as a less derived group compared to the Digamasellidae. However, I have reservations concerning his assessment of sister group relationships between the two families. I do not agree that they are sister groups; they represent early derivatives within two very closely related lineages - one which gave origin to the Laelaptonyssoidea and the other from which the Rhodacaroidea originated. Several of the character states listed by Lindquist as shared synapomorphies are not unique to the Digamasellidae and Rhodacaridae. In fact, they are derived independently within other groups of Rhodacaroidea, and are discussed below.

For example, the presence of scleronoduli is common in

Rhodacaridae sensu stricto and Digamasellidae. However, scleronoduli are also present in several taxa of Gamasellidae: Hiniphis hinnus; two species of the Gamasellus pyriformis complex (G. pvriformis

Berlese, 1916a and G. muscosus Hurlbutt. 1978); one species of the 61

Gamasellus discutatus complex (G. discutatus Lee. 1966); one species

of the Gamasellus falciger complex (G. concinnus Womersley. 1942);

all three species of the Gamasellus tragardhi complex (G. cophinus

Lee, 1973, G. grossi Lee. 1973 and G. tragardhi Womersley. 1942); in

Rhodacaroidea (Nodacaroides) coniunctus Karg, 1977 and Rhodacaroides

(Nodacaroides) crinitus Karg, 1979; and, in two species of

Ologamasidae: Hvdrogamasellus racovitzai ; the deutonymph of

Hvdrogamasellus striatus Sheals, 1962 (I was unable to examine the

adult). In addition, Protogamasellus mica (an ascid species)

possesses three scleronoduli on its podonotal shield (just as in

Rhodacarus). whereas Digamasellus (the type-genus of the

Digamasellidae) and Latogamasellus (Rhodacaridae) have secondarily

lost their scleronoduli. The highly derived digamasellid genus

Longoseius may or may not have scleronoduli on its podonotal shield.

Regarding the anterior weak sclerotization of the female sternal

shield, this derived condition is commonly found in digamasellids but

it is observed in less than half of the known rhodacarid genera

recognized by me (5 out of 12 genera).

A recurved male spermadactyl in the sense of Lindquist (..."with

an initial posteroventral bend before projecting freely

anteriad..."), is widely distributed throughout the Rhodacaroidea (29

genera within 3 families) and is also observed in the Panteniphididae

and the Laelaptonyssidae. Therefore, it is not a synapomorphy of

Digamasellidae and Rhodacaridae but rather, a convergence of the

Laelaptonyssoidea with several genera of the Rhodacaroidea. However,

a condition observed in some taxa of Rhodacaroidea and in the 62

Laelaptonyssidae (but not in the Digamasellidae and Panteniphididae), is a strongly recurved spermadactyl, projecting posteriad, seen in the following taxa: Rhodacarus. Rhodacarellus. Rhodacaropsis.

Solugamasus. Afrodacarellus concavus Hurlbutt, 1973, ^ femoratus

Hurlbutt, 1973, A. pocsi Hurlbutt. 1973, A. minutus Hurlbutt. 1973

(Rhodacaridae); Pyrvphis (Gamasellidae); Geogamasus. Sessiluncus and

Hvdrogamasus (Ologamasidae).

Complete loss of the submarginal (UR) series of setae is an apomorphic condition; it is a trend observed throughout the

Rhodacaroidea (except for the Euryparasitidae and Epiphidinae), as well as in the Panteniphididae, Laelaptonyssidae and Digamasellidae.

A "basically triramous" tectum is relatively common in

Rhodacaridae, Digamasellidae, other rhodacaroid families,

Panteniphididae, Laelaptonyssidae, Ascidae and Veigaiidae; given its wide distribution, I regard this condition as a plesiomorphic one. A derived condition of the triramous tectum is a bifid and/or more elaborate central tine.

I regard as plesiomorphic the condition where the male ventrianal shield is not fused to the sternogenital shield. The fusion of these two shields is an apomorphic trend observed in

Heterogamasus (Euryparasitidae), Rvkellus (Ologamasidae), Pyriphis.

Hiniphis hinnus. Hiniphis bipala. Periseius (Periseius), Periseius

(Psammonsella). Pillelus and Evanssellus (Gamasellidae) and

Panteniphididae.

On the parallelisms (they are actually convergences) pointed out by Lindquist, the tendency towards loss of seta pl3 on basitarsus IV 63 is the only valid one; the anterior displacement of podonotal setae j2 is commonly seen in two ascid genera (Protoeamasellus and

Gamasellodes) and the opening of the female spermatheca on leg III is also a common feature of ascids. On the other hand, rhodacarid genera such as Afrogamasellus. Afrodacarellus and Latogamasellus do not have setae j2 displaced anteriorly, and Rhodacarus and Latogamasellus are the only rhodacarids with spermathecal ducts opening on leg III.

It is understandable that the character states discussed here could be interpreted as synapomorphies of the Rhodacaridae sensu stricto and Digamasellidae, if only a limited analysis were to be done, that is, if the taxa to be studied included only the two groups, one outgroup, and a hypothetical ancestral group such as the

"parasitid-veigaiid like” stock. However, if a broader study were to be undertaken, the conclusions then reached might not be the same as those of Lindquist or, they might corroborate his analysis with valuable information resulting in an improved definition of the

Rhodacaroidea. It was precisely with the search for additional evidence in mind that I decided to include in this study taxa not

traditionally thought of as belonging to the Rhodacaridae sensu

stricto as well as other potential outgroups such as the Ascidae, and

taxa of disputed familial placement CGamasellodes. Protogamasellus.

Panteniphis. Lindquistoseius).

The resulting inadequacy of several studies on the Rhodacaridae

stems mainly from the fact that some workers form a preconceived

notion of what should be a typical Rhodacaridae and, therefore, do

not even consider studying other taxa that do not accommodate to that 64 notion. Shcherbak, for example, formed her concept of the family

(Table 5) on the basis of Oudemans* very restricted interpretation of the Rhodacaridae (which was derived from the monotypic genus

Rhodacarus) and, except for a few changes (such as the exclusion of

Protogamasellus from this family), has remained relatively faithful

to it (1976, 1979, 1980, 1982, 1983).

She feels that, since the early 50*s, the family Rhodacaridae has lost its clear cut interpretation to a considerable degree, due

to the inclusion of what she calls "random genera" by many authors

such as Evans and Ryke.

She strongly opposes Lee's concept of the Rhodacaridae (1970 -

refer to Table 3), which includes forty five genera and excludes

thirteen genera "assigned to the family in the past by other

researchers". Lee's interpretation is as follows:

"Sclerotization very variable, but always separate podonotal and opisthonotal shields on deutonymph and a posteriorly truncated female genital shield separated from a conspicuous ventro-anal shield. Female metasternal seta, st4, usually on a sterno-metasternal shield. Leg chaetotaxy usually as for the "Rhodacarus - group" genera referred to by Evans (1963). Apotele usually three-pronged, never with associated hyaline flap, and if two-pronged then there are four ventral setae on tibia I. Male with distally free spermadactyl, presternal genital orifice and seta on femur II larger than that of the female and usually considerably modified to a conspicuous spur."

According to Shcherbak, Lee's diagnosis is highly extensive and

completely deviates from the original description of the family. And

to illustrate that, she makes the following comments: 1) the body

sclerotization cannot vary in the same family to such an extent, that

its representatives would have either completely separated dorsal

plates or a single dorsal plate; 2) a posteriorly straight genital 65 plate is not a characteristic feature in numerous species of

Rhodacaridae; 3) a distally free spermadactyl and the modification of one of the setae on the male femur II into a spur, are characteristic of numerous gamasid families.

I subscribe in part to the objections of Shcherbak, in particular to the highly variable degree of body sclerotization within the same family. However, she overlooks important diagnostic characteristics such as the position of st4, the pattern of leg chaetotaxy and the three-pronged palp claw, while failing to recognize that Ouderaan's original concept included features which were/are not solely characteristic of the family.

The characteristics considered of the utmost importance to her interpretation of the Rhodacaridae, are; "three or four well developed scleronoduli "constantly observed on the carapace"

(podonotal shield), and two dorsal plates (shields) on adults and deutonymphs. Her 1979 diagnosis also includes: female with four pairs of setae on large sternal plate; small genital plate with one pair of setae; ventrianal plate of various forms and sizes; male with two large ventral plates - a sternogenital with the genital opening placed at its anterior margin, and a ventrianal plate; spermadactyl basally fused with movable digit, distally always free.

It seems ironical that, while being critical of Lee's choice of features regarding the characterization of the Rhodacaridae and of the great variation in body sclerotization, Shcherbak adopts several of the same features and allows a similarly broad range regarding body shapes and the degree of sclerotization of certain ventral 66 shields in the "Description" section which follows her diagnosis.

References to the various body shapes are made (from elongated, i.e.. four or more times as long as wide, to spherical-pentagonal with an angular or bilobed posterior end), as well to the broad range of

shapes and dimensions of the female ventrianal shield. Additional

features included by Shcherbak are: large male ventrianal plate

distinctly separated from the large sternogenital plate; ventrianal

plate often posteriorly and laterally fused with the opisthonotal plate; adults usually with eight pairs of opisthogastric setae

(excluding post-anal seta) on the ventrianal plate or off it; male

sternogenital plate with four or five pairs of setae (if four pairs,

then fifth pair is placed on soft cuticle or on discrete triangular platelets); anal opening small in immatures and adults; tectum with

three processes of equal or different length; female chelicerae with well developed teeth ( movable digit with one large tooth); spermadactyl of various lengths, fused basally with the movable digit, its fused part mostly "S" shaped; male leg II bears large spine on femur, genu with highly enlarged spine-like setae and many species with other segments containing these enlarged setae as well; hypostomal groove with six to eight furrows and euanal setae present

in protonymphs and larvae.

Furthermore, on the basis of her diagnosis alone, one would have

to include with the Rhodacaridae sensu stricto, several taxa not recognized by her as members of that family. These taxa are:

Paragamasellevans: Litogamasus: Solugamasus; Gamasellus concinhus;

Gamasellus cophinus; Gamasellus discutatus; Gamasellus falciger; 67

Gamasellus grossi ; Gamasellus muscosus; Gamasellus ovriformis;

Gamasellus tragardhi; Rhodacaroides (Nodacaroides) coniunctus;

Rhodacaroides (Nodacaroides) crinitus; Rhodacaroides minvaspis;

Rhodacarus costai; Hiniphis hinnus. Excluded, would be:

Hvdrogamasellus racovitzai (for having an entire dorsal shield) and

Protogamasellus (for not bearing st4 on the sternal shield).

Clearly, Shcherbaks' diagnosis is not sufficient to characterize the family and, if used together with her description, would still have to include Paragamasellevans. Solugamasus. R. costai. R. minvaspis. and ^ (Nodacaroides) coniunctus. Moreover, her overall concept of the Rhodacaridae (which includes 13 genera and is primarily based on the Palearctic fauna) is actually considerably more inclusive than that of Lee, which includes 45 genera distributed throughout the major zoogeographical zones of the world and therefore, takes into account a higher degree of morphological diversity.

It should be pointed out that some of the diagnostic features listed by Lee (1970) and Shcherbak (1979) are not only characteristic of the Rhodacaridae (=Rhodacaroidea in this study), but also of the

Dermanyssina and although they are synapomorphies at some higher level, they represent symplesiomorphies at the familial level. These are: male genital aperture placed at the anterior margin of the sternal shield; male leg II with some setae modified into enlarged spurs particularly on femur, genu and tibia; female genital shield with one pair of setae. Plesiomorphies within the family include: two dorsal shields in deutonymphs and adults; palp claw three-tined; 68

tibia I with four ventral setae.

At any rate, neither Shcherbak's nor Lee's concepts of the taxon

Rhodacaridae can be monophyletically defined. Lee's interpretation of

the Rhodacaridae is partly corroborated by my results except for his

inclusion of the Laelaptonyssinae and Tangaroellinae and for the

internal organization of some groups.

The relationships between the Rhodacaroidea and

Laelaptonyssoidea are presented below, followed by detailed

discussions of the relationships within each of these superfamilies.

The two lineages which seem to have arisen from a veigaiid-like

ancestor (the Veigaiidae, a group of free-living predators commonly

found in ground habitats, emerged as their likely sister group in the

vast majority of the computer-generated cladograms), early undertook

different evolutionary pathways. The Laelaptonyssoidea evolved a

number of uniquely derived features especially concerning their

pattern of leg chaetotaxy: genu I with 12 setae (42); genu III with 8

setae (43); genu IV with 7 setae (44); tibia I with a maximum of 12

setae (45) and tibia IV with a maximum of 7 setae (47). They also

achieved several apomorphic (although not unique) conditions such as

complete loss of the submarginal (UR) series from the opisthonotal

area of the dorsal shields (38), a bifid palpal claw (6), a basally

recurved spermadactyl (4), an anteriorly weakly sclerotized female

sternal shield (18), a reduction in the number of solenostoma of

ventral gland gv2 (19), an oligotrichous opisthonotal area of the

idiosoma, with a maximum of 22 pairs of setae (26), a reduced number

of 7 pairs of opisthogastric setae (31) and a maximum of two pairs of 69 opisthonotal setae placed on the soft cuticle of the female

opisthonotum (36).

The Rhodacaroidea, on the other hand, evolved five

synapomorphies: the peritrematal shields fused to the exopodal IV

shields (17); the opisthonotal area of the idiosoma achieved

oligotrichy (although to a lesser degree than that of the

Laelaptonyssoidea), with a maximum of 27 pairs of setae (26); the

number of opisthogastric setae was reduced to a maximum of 9 pairs

(31); the peritremes gradually shortened (33) and the submarginal

(UR) series of setae became ever reduced to the point of their

complete loss (38),

From the biological standpoint, the Lalaptonyssoidea also

exhibit a more derivative modus vivendi: the Digamasellidae (with 8

genera and hundreds of species) are found in various decomposing

substrates or associated with wood and with the insects living in it

(especially bark beetles); the Laelaptonyssidae (with 2 monotypic

genera - Laelaptonvssus Womersley, 1956a and Puchihlungia Samsinak,

1964) have only been found associated with insects as well (on

termites and in their nests or, in fly cultures); the Panteniphididae

(with 2 genera - Panteniphis and Linda uistoseius) are found in

permanently moist habitats, in forest litter, rotting grass, etc.

As for the Rhodacaroidea, even though they can be found in a

myriad of habitats, there is no evidence of direct association with

other organisms, except for two genera of Euryparasitidae. These are

the closely related Cvrtolaelaps and Eurvparasitus. found associated

with nests of rodents, moles and rarely birds, with their deutonymphs 70 being phoretic on mammals, especially insectivores. Kethley (1983) reports that deutonymphs of Epiphis and Iphidosoma fimetarium Muller,

1859 (Epiphidinae) are also phoretic; the former having been collected from millipedes, the nest of a small mammal, a sparrow and a carabid beetle (Diacelus sp.), while the latter has been collected from a number of carabid species from Europe. Kethley points out that the microhabitat records for Epiphis rarior deutonymphs "seem more suggestive of a microhabitat correlated phoriant, rather than a taxonomically correlated phoriant", since it is "the ecological group of arthropods and small vertebrates in association with the microhabitat of ^ rarior (deciduous leaf litter in moist areas)" which is "more likely to be involved as phoriants rather than any particular species". Rare instances of phoresy by other rhodacaroid taxa include a single female of Gamasiphis Berlese, (Gamasiphinae) reportedly collected from a rat, in Puerto Rico (Fox, 1949) and a female of Gamasellus spiricornis G. & R. Canestrini, 1882

(Gamasellidae) collected from a carabid beetle (Cooreman, 1943, cited in Lee, 1970).

Within the three families comprising the Laelaptonyssoidea, there is a trend towards loss of one or two setae on basitarsus IV -

Laelaptonvssus mitis. Dendrolaelaspis. Lindquistoseius, L.

(Longoseius), L^ (Longoseiulus) - and more rarely, loss of one or two setae on basitarsi II and III - L^ (Longoseius). L. (Longoseiulus) and Dendrolaelaps.

Within the Rhodacaroidea, oligotrichous leg segments are uncommon, although a few trends are observed; the majority of 71

Rhodacaridae lack one seta on basitarsus IV, that seta consistently being pl4, therefore, reducing the total number of setae on that segment to seventeen; In the subfamily Gamasiphinae, there Is tendency towards the loss of one ventral seta on genu IV; In the

Euryparasitidae, the genera Heterogamasus and Notogamasellus have achieved neotrlchy In the genu and tibia III, where one seta was added to each of these segments; an Isolated case of these same conditions Is observed In the subgenus Perlselus « a member of the

Gamasellidae.

The taxon Rhodacaroidea constitutes an assemblage of groups which, aside from the synapomorphies that justify Its monophyletlc definition, exhibit numerous parallelisms (homologous character states Independently derived In closely related taxa), as Indicated by the results In this study and presented below.

Interfamlllal relationships;

The Euryparasitidae n. fam. Is the sister group of the remaining

Rhodacaroidea. This early derivative taxon differentiated from the ancestral stock by evolving a non pedunculate pretarsus I (30) and a female chellcera with 6-12 teeth on Its fixed digit (57). Its sister lineage evolved five synapomorphies: an oligotrichous opisthonotum, with 20-22 pairs of setae (26); female chelicerae with small and large teeth combined (34); female opisthonotum with 3-6 pairs of setae on soft cuticle (36); female with 2 pairs of opisthogastric setae placed on soft cuticle (37) and submarginal (UR) series of setae absent from the opisthonotum (38). 72

The first taxon to emerge from this lineage was the Rhodacaridae

(sister group to the Gamasellidae and Ologamasidae) with six apomorphies: spermadactyl recurved at basal or median level of the movable digit before projecting freely (4); four scleronoduli on the

dorsal hexagonal area of the podonotal shield (5), which

independently derived in the Digamasellidae, some Gamasellidae and

one Ascidae taxon as well; ventral gland gv2 produced into 1-2 pores

at most (19), a character state which also derived independently in

the Gamasellidae, Gamasiphinae and some Ologamasinae; pretarsus I not

pedunculate (30), independently derived in the Euryparasitidae as

well; opisthogastric setae reduced to 7 pairs (31); peritremes

reduced in length, reaching only up to median level of acetabula II

(33) - and one reversal - postanal seta longer than the adanal setae

(1). The lineage which originated the remaining Rhodacaroidea

(Gamasellidae and Ologamasidae) evolved four autapomorphies -

posterior margin of female genital shield not extending much beyond

acetabulum IV (9); opisthonotal setae Z5 less than one and a half

times as long as setae J5 (23); opisthogastric setae Zvl placed at

level below setae Jvl (55); female exopodal shields III not coalesced

but at least encasing peraxial edges of acetabula III (59) - and two

apomorphies: a maximum of one pair of podonotal setae placed on soft

cuticle of female podonotal shield (35) and no opisthonotal setae

placed on soft cuticle in females (36).

The Gamasellidae evolved with five synapomorphies: gland gv2

with 1-2 pores at most (19); no podonotal setae placed on soft 73 cuticle in females (35); no opisthogastric setae on soft cuticle in

females (37); female chelicerae with 6-12 teeth on its fixed digit

(57), a parallelism with Euryparasitidae; seta all on palp genu bifid

(60).

The Ologamasidae (with its three subfamilies represented on the

cladogram) evolved with two autapomorphies: dorsum of the idiosoma

usually entire in both sexes, but occasionally divided in males (49)

and female genital shield with tendency towards being wider than long

(54). The Ologamasinae differentiated from its sister

Epiphidinae-Gamasiphinae group, by acquiring two autapomorphies -

female chelicerae with predominantly large teeth (34); site of the

spermathecal opening commonly placed on coxa IV (61), and one

apomorphy: seta all on palp genu elaborated, being at least spatulate

(60). The lineage from which the Epiphidinae- Gamasiphinae group

originated uniquely derived a female ventrianal shield tending to

coalesce with the notai shield (15), dorsum of the idiosoma entire in

both sexes (49), female exopodal shields III coalesced (59), and

apomorphically gained the conditions where no podonotal setae are

placed on the soft cuticle of females (35) and no opisthogastric

setae are placed on the soft cuticle of females (37). The

Gamasiphinae evolved gland gv2 with 1-2 solenostoma (19) and a

tendency for the female sterno-metasternal shield to fuse with

endopodal shields IV (58). The Epiphidinae on the other hand,

achieved a unique combination of character states: eleven or more

pairs of opisthogastric setae (31), apparently, a neotrichous

condition; the peritremes became reduced in length (33), a 74 parallelism with the Rhodacaridae; the opisthogastric setae Zvl migrated to a level well above that of setae Jvl (35), a condition independently derived in Afrodacarellus (Rhodacaridae); seta all on palp genu elaborated, finely pilose (60); site to the spermatheca placed on coxa III (61) and postanal seta longer than the adanals

(1). Regarding the relationships among the Laelaptonyssoidea, each family has such unique sets of characteristics that, if it were not for the overwhelming number of synapomorphies they share (thirteen), one might not be inclined to group them together solely on the basis of their overall appearance. Examples illustrating these characteristics will follow.

The Digamasellidae differentiated from its ancestral and sister lineages with five autapomorphies: four scleronoduli placed on the dorsal hexagonal area (dha) of the podonotal shield (5); male sternogenital setae (st5) placed on separate platelets (8); ventrianal shield fused to opisthonotal shield (15); first sternal setae (stl) placed on area of weak sclerotization of sternal shield

(24) and pretarsus I set on a short peduncle (30). In addition, they achieved four apomorphic conditions: two pairs of opisthogastric setae usually placed on soft cuticle in females (37), a condition independently derived in Panteniphis also; movable digit of female chelicera with 4-8 teeth (56); fixed digit of female chelicera with

6-12 teeth (57); and spermathecae commonly opening on femora III

(61). Character states 56, 57 and 61 have independently derived in

Lindquistoseius as well. 75

The lineage from which the Panteniphididae-Laelaptonyssidae group arose, evolved three autapomorphies - podonotum oligotrichous, with 16-19 pairs of setae (25); femur I with a maximum of 12 setae

(39); female genital shield tending towards being wider than long

(54) - and three apomorphies: peritremes reduced in length (33); tibia I with a maximum of 12 setae (45) and opisthogastric setae Zvl tending to migrate to a level below Jvl (55).

The Panteniphididae differentiated from its sister group with six autapomorphies: male sternogenital shield fused to the ventrianal shield (12); female peritrematal shields fused to the exopodal shields IV (17); genu IV with 6 setae (44); tibia III with 6 setae

(46); tibia IV with 6 setae (47) and female exopodal shields III coalesced (59).

The Laelaptonyssidae achieved seven autapomorphies - opisthonotum paedomorphic, with 13-15 pairs of setae (26); arthrodial process of female chelicerae absent (27); peritremes extremely reduced, so that only the stigmata are present (33); female chelicerae tending towards toothlessness (34); femur I with 11 setae

(39); femur III neotrichous, with 7-8 setae (40); movable digit of female chelicera with 1-3 teeth at most (56) - two apomorphies - opisthonotal setae Z5 slightly longer than setae J5 (23); no podonotal setae placed on soft cuticle in females (35) - and one reversal - three to six pairs of opisthonotals placed on soft cuticle in females (36), a secondarily derived condition probably due to the reduction in size of the opisthonotal shield. Even though a detailed analysis of the Laelaptonyssoidea is beyond the scope of this study. 76 a few conunents concerning the composition of the Panteniphididae and

Laelaptonyssidae are worth making and these are presented below.

Family Panteniphididae: it includes the genera Panteniphis and

Lindquistoseius. the latter previously regarded as a subgenus of

Panteniphis (Hirschmann, 1983). Through this study I’ve found that

Lindquistoseius is distinctive enough to retain generic status. It has differentiated from Panteniphis with nine autapomorphic and six apomorphic states. Its autapomorphies include: first sternal setae

(stl) placed between the preendopodals and the sternal shield (24); arthrodial process of the female chelicera produced nto a conspicuous

(27), a convergence with some Rhodacaridae taxa; peritremes reduced, extending only as far as the midle of acetabula II (33) - another convergence with the Rhodacaridae; no opisthonotal setae placed on soft cuticle (36), a convergence with several Ologamasidae taxa; genu

I with only 11 setae (42); tarsus IV with 16 setae (48), a convergence with Tangaroellus (Ascidae) and a parallelism with some

Digamasellidae taxa; dorsum of the idiosoma may be entire in both sexes or the male may it divided (49) - a convergence with two taxa of Ologamasidae and one taxon of Gamasellidae; opisthonotal setae J5 placed at level posterior to that of setae Z5 (52), a convergence with Rhodacarus. Rhodacarellus. Nodacaroides (Rhodacaridae);

Notogamsellus.(Euryparasitidae); Tangaroellus. Gamasellodes.

Protogamasellus (Ascidae); Gamasolaelaps (Veigaiidae); setae all on palp genu spatulate (60), which is also observed in some rhodacaroid taxa. Its apomorphies are: spermadactyl less than one and a half times as long as the movable digit of the movable digit of the 77 chelicera (2); opisthonotal setae Z5 slightly longer than setae J5

(23); no podonotal setae on soft cuticle (35), a parallelism with

Laelaptonyssidae and Digamasellus; movable digit of female chelicera with four teeth (56); fixed digit of female chelicera with six or seven teeth (57); site of spermathecal opening on femur III (61), a parallelism with some Digamasellidae.

Panteniphis on the other hand, has evolved two autapomorphies: opisthogastric setae reduced to 4 pairs, a paedomorphic condition

(31), and dorsum of the idiosoma entire in both sexes (49), along with three apomorphies: usually one pair of opisthogastric setae on soft cuticle in females (37); tibia I with 12 setae (45), a reversal; spermathecae opening on coxa IV (61).

Family Laelaptonyssidae: it includes Laelaptonvssus and

Puchihlungia. Lee (1970) synonymized these two monotypic genera and placed them in the Laelaptonyssinae, which he regarded as one of the subfamilies of Rhodacaridae. My findings indicate that this taxon does not belong with the Rhodacaroidea and that Laelaptonvssus and

Puchihlungia are distinct genera.

Laelaptonvssus is defined by six autapomorphies - spermadactyl at least one and a half times as long as the movable digit of the chelicera (2); female sterno-metasternal setae placed on soft cuticle

(7); male genital setae absent (8); tibia I with 9 setae (45); tibia

III with 6 setae (46); fixed digit of female chelicera with a maximum

of two teeth (57), and four apomorphies: preendopodal shields absent

(14), a convergence with some Ascidae; podonotal setae zl absent

(22), a convergence with Gamasellopsis (Ologamasinae); tarsus IV with 78

17 setae (48), a parallelism with some Digamsellidae and a convergence with some Rhodacaridae; and finally, femur IV neotrichous, with 7 setae (53), a convergence with Pj_ (Periseius)

(Gamasellidae) and Tangaroellus (Ascidae).

Puchihlungia has evolved four autapomorphies: podonotum paedomorphic, with 13 pairs of setae (25); opisthogastric setae as 8 pairs (31); tibia IV with 8 setae (47), a reversal; opisthogastric setae Zvl placed at level below that of setae Jvl (55), also a reversal. Along with these, Puchihlungia evolved one apomorphy: ventral glands gv2 absent (19), a convergence with Evanssellus

(Gamasellidae), Notogamasellus (Euryparasitidae), and Protogamasellus

(Ascidae). On the basis of the characteristics listed above, I prefer to retain the generic status for Puchihlungia (Laelaptonyssidae) and

Lindquistoseius (Panteniphididae).

Intrafamilial relationships;

Family Euryparasitidae:

From the ancestral group, two lineages emerged: one, from which the Eurvparasitus - Cvrtolaelaps group of genera originated, developed a spermadactyl at least one and a half times as long as the movable digit of the male chelicera (2) and the female chelicerae composed of small and large teeth (34). The other lineage, which gave rise to the Heterogamasus - Acugamasus - Allogamasellus -

Notogamasellus group of genera, autapomorphically evolved: complex podonotal setae jl, at least barbed or strongly pilose (13); opisthogastric setae reduced to 8 or 9 pairs (31); a maximum of 2 pairs of opisthogastric setae placed on soft cuticle in females (37) 79 and submarglnal (UR) series of setae reduced to a maximum of 4 pairs

(38). An exception to character state 37 is observed in females of

Notogamasellus. which possess 3-4 pairs of setae on soft cuticle; this reversal seems due to a reduction in size of the ventrianal shield. The Heterogamasus - Acugamasus group achieved three autapomorphies - podonotal setae jl pilose and spatulate (13); female genital shield at least one and a half times as long as wide (54); spermathecae opening on trochantera III (61), and one apomorphy: dorsal setae of the idiosoma predominantly complex (29), a parallelism with Notogamasellus. Its sister Allogamasellus -

Notogamasellus uniquely derived four character states - opisthonotal setae Z5 subequal in length to setae J5 (23); opisthogastric setae reduced to 7 pairs (31); females with 2 pairs of podonotal setae usually placed on soft cuticle (35), a secondarily derived condition; opisthonotal setae J5 placed at level subequal to or posterior to that of setae Z5 (52) - as well as three apomorphies - gland gv2 with

1-2 pores (19), a parallelism with Acugamasus; peritremes reduced in length, reaching to mid acetabula II (33), a parallelism with

Eurvparasitus and with the Rhodacaridae; females with 3-6 pairs of opisthonotal setae placed on soft cuticle (36), a parallelism with

Cvrtolaelaps.

Cvrtolaelaps further became differentiated with six autapomorphies (13, 14, 26, 29, 35, 50) and one apomorphy (36), while its sister Eurvparasitus achieved two autapomorphies (50, 56) and two apomorphies (33, 60). Heterogamasus evolved three unique character states (6, 12, 31) along with four apomorphic states (32, 43, 46, 80

60); its sister Acugamasus in turn, achieved three autapomorphic conditions (1, A, 8) and five apomorphic ones (19, 21, 34, 55, 60).

Finally, Allogamasellus evolved three autapomorphies (14, 37, 56) and five apomorphies (2, 21, 32, 55, 60), whereas its sister

Notogamasellus differentiated with six autapomorphies (19, 25, 26,

27, 30, 38) and four apomorphies (29, 43, 46, 52). For a complete list of character states which define each stem refer to fig.2 legend.

Family Rhodacaridae:

The ancestral group gave rise to two lineages: on one side the group of genera formed by Litogamasus, Rhodacaropsis. Rhodacarus,

Rhodacarellus. Paragamasellevans. Solugamasus. Gen. n. #1 and

Nodacaroides new status, which will be referred to as lineage "A”; on the other side, the Afrogamasellus (including "mongii").

Mediodacarellus n. gen., Afrodacarellus (including "citri") and

Latogamasellus (including "lubalensis") group of genera, referred to as lineage "B".

Lineage "A” autapomorphically developed a spermadactyl freer than fused to the movable digit (3) and recurved at its base or at its median level (4) and female chelicerae with predominantly large teeth (34), while apomorphically developing a female genital shield at least one and a half times as long as wide (54). Its sister lineage ”B" evolved one autapomorphy - the female chelicerae with an arthrodial process produced into a conspicuous brush (27) - and six apomorphies: postanal seta slightly longer than adanals (1); podonotal setae zl shorter than setae jl (22); no opisthonotal setae 81 placed on soft cuticle in females (36); one pair of opisthogastric setae usually placed on soft cuticle in females (37); setae pl3 absent from tarsus IV (41); tarsus IV with a total of 17 setae (48).

It is interesting to point out that character states 41 and 48 independently derived in the Rhodacaropsis - Rhodacarus -

Rhodacarellus group as well.

From lineage "A", two others originated: one in which autapomorphically there are 3-6 pairs of opisthonotal setae on the soft cuticle in females (36) and apomorphically, the preendopodal shields have developed into three or more pairs (14) and the opisthonotal setae J5 have moved to a level subequal to that of Z5

(52), giving rise to the Litogamasus - Rhodacaropsis - Rhodacarus -

Rhodacarellus group. In the other, the spermadactyl is uniquely recurved distad of the movable digit before projecting freely (4), and the postanal seta is only slightly longer than the adanals (1) - the latter, a parallelism with lineage "B"; from this lineage evolved the Paragamasellevans - Solugamasus - Gen. n. #1 - Nodacaroides group. minvaspis Lee, 1973 (comb, ii) exhibits reversals of character states 35, 36 and 37; these conditions are secondarily derived and probably correlated with the overall reduction in extent of sclerotization - to my knowledge, this is the only rhodacaroid taxon with an anal (rather than a ventrianal) shield.

From lineage "B" arose one lineage with the first pair of sternal setae (stl) placed on the preendopodal shields (24) - from which Afrogamasellus originated - and another, with the peritrematal shields of females fused to the exopodals IV (17), the opisthonotal 82 setae Z5 subequal in length to setae J5 (23) and opisthogastric setae

Zvl placed at a level well above that of setae Zvl, nearly vertical to it (55) - which gave rise to the remaining rhodacarid taxa

(Mediodacarellus. Afrodacarellus and Latogamasellus).

It is important to notice that within the Rhodacaridae, there are generalized trends towards reduction (number of scleronoduli, leg chaetotaxy, degree of sclerotization and extent of shields). For instance, there is tendency for the preendopodal shields (14) to gradually decrease in number and degree of sclerotization: from at least three pairs (Litogamasus). to two pairs (Rhodacaropsis.

Solugamasus and Gen. n. #1), to one well sclerotized

(Paragamasellevans. Nodacaroides. Afrogamasellus) or weakly sclerotized pair (A. mongii. Mediodacarellus). Ultimately, these shields are lost, with only a remnant area of punctate sclerotization immediately before the sternal shield - this condition is observed in

Rhodacarus. Rhodacarellus. Afrodacarellus and Latogamasellus.

Reductions in the extent of shields include the peritrematals, the ventrianal, the opisthonotal and even the sternal shields, which may be accompanied by reduction in degree of sclerotization as well. Some consequences of these reductions involve the increase in number of

setae placed on the soft cuticle - notice the pathway taken by

characters 35, 36 and 37 in Fig. 3.

I recognize twelve genera within the Rhodacaridae: five of them

already established members of the family: Rhodacarus. Rhodacarellus,

Rhodacaropsis. Afrogamasellus and Afrodacarellus; three are newly

included: Paragamasellevans. Litogamasus and Solugamasus; three 83 represent new combinations and/or new genera - Nodacaroides new

status, including: N. costai (Sheals, 1962), comb, n.. N. minvaspis

(Lee, 1973), comb, n.. N. calidus (Karg, 1977), comb, n.. N.

brevispiritus (Karg, 1977), comb, n.. N. coniunctus (Karg, 1977),

comb, n. and N. levis (Karg, 1977), comb, n.; Mediodacarellus n. gen.

including M. bakeri (Hurlbutt, 1973), comb, n.; Latogamasellus Karg

(originally defined as a subgenus of Afrogamasellus). including L.

squamosus (Karg, 1977), L. bipilosus (Karg, 1979), L^ furculatus

(Karg, 1979), L^ myersi (Loots, 1969), and lubalensis (Loots,

1969). The last taxon to be included (referred to as Gen. n. #1) is represented by female specimens collected from tree holes in Ohio and a soil core from maple-basswood forest in Michigan. A brief discussion of these taxa follows:

Nodacaroides Karg, 1977 new status:

Type species: Rhodacaroides (Nodacaroides) coniunctus

Karg, 1977.

(= Rhodacaroides (Tenacaroides) Karg, 1977, new synonymy).

Type species: Rhodacaroides (Tenacaroides) calidus Karg, 1977.

This taxon is used to accommodate eight species previously attributed to Rhodacaroides by Lee (1970, 1973) and Karg (1977,

1979); Rhodacarus costai Sheals, 1962 (from South America),

Rhodacaroides minvaspis Lee, 1973 (from Australia), Rhodacaroides

(Tenacaroides) calidus Karg, 1977, Rhodacaroides (Tenacaroides)

brevispiritus Karg, 1977, Rhodacaroides (Nodacaroides) coniunctus

Karg, 1977 Rhodacaroides (Nodacaroides) levis Karg, 1977,

Rhodacaroides (Tenacaroides) unguellus Karg, 1979 and Rhodacaroides 84

(Nodacaroides) crinitus Karg, 1979, ail six from South America.

Lee (1970) transferred costai from the Rhodacarinae to

Rhodacaroides in the Ologamasinae as a temporary measure, admiting his uncertainty of its relationships. Later, Lee (1973) described a species from Australia (minvaspis) tentatively grouping it in

Rhodacaroides. but pointing out that ^ minvaspis was more similar to

R. costai than to the type species of Rhodacaroides, R. aeevptiacus

Willmann, 1959 (from Egypt). My results confirm that, N. costai (n. comb.) and N. minvaspis (n. comb.) are closely related and belong to the Rhodacaridae, while R^ aegvptiacus belongs to the Geunasellidae.

The two species share three apomorphies - opisthonotals Z5 subequal in length to setae J5 (23); opisthogastric setae reduced to

6 pairs (31), a parallelism with Solugamasus; opisthonotum oligotrichous, with 16-19 pairs of setae (26); opisthonotals J5 placed at level at most subequal to those of setae Z5 (52), a parallelism with the Litogamasus - Rhodacaropsis - Rhodacarellus -

Rhodacarellus group. The trend towards reduction can be particularly observed in Nj_ minvaspis which has developed a series of uniquely derived character states such as: the number of scleronoduli is reduced to two (5); gland gv2 absent (19); anal valves enlarged (20) and only an anal shield is present; females with 3 pairs of podonotals (35), at least 8 pairs of opisthonotals (36) and at least

4 pairs of opisthogastric setae (37) placed on soft cuticle. In addition, it has a paedomorphic opisthonotum with 12 pairs of setae

(26) - a parallelism with Solugamasus. 85

Although I have not examined the species described by Karg

(1977; 1979), there seems to be some evidence that they are more closely related to costai and N^_ minvaspis than to ^ aegyptiacus.

For example, these species share the following characteristics with

Nodacaroides: female tectum with central tine denticulate laterally

(in Rhodacaroides it is completely smooth); one pair of preendopodals

(14), a plesiomorphic condition (Rhodacaroides has two pairs); peritremes reduced, extending only to middle level of acetabula II

(33), whereas in Rhodacaroides they have full length, the plesiomorphic state; opisthonotum with a maximum of 19 pairs of setae

(26) - Rhodacaroides has 21 pairs; opisthogastric setae reduced to 6 pairs (31) - Rhodacaroides has 7 pairs; between 3-7 pairs of opisthonotal setae are placed on soft cuticle (36), except for N. calidus which apparently has no setae on soft cuticle (Rhodacaroides has 1 pair of opisthonotal setae on soft cuticle); female genital shield usually one and a half times as long as wide (54) - in

Rhodacaroides it is only slightly longer than wide, the plesiomorphic state; opisthogastric setae Zvl placed at level parallel to that of setae Jvl (55) - in Rhodacaroides setae Zvl are placed at level below

that of setae Jvl, an apomorphic condition; movable digit of female chelicera (56) usually with five teeth (and up to seven), except for

N. unguellus and crinitus which have three teeth on their movable

digit - Rhodacaroides females have only three large teeth on the

movable digit, the plesiomorphic condition; opisthonotal setae J5 are

commonly placed at anterior (plesiomorphic) or subequal (apomorphic)

level to that of setae Z5 (52), whereas in Rhodacaroides setae J5 are 86 placed at level posterior to that of setae Z5, the most derived condition; seta all on palp genu has at least 4 prongs (60) - in

Rhodacaroides all is trifurcate (with 2 lateral prongs); sternal shield entirely clearly defined (18) — in Rhodacaroides it is weakly defined at its posterior margin, the derived condition; podonotal glands gd4 not enlarged (21), the plesiomorphic condition

(Rhodacaroides has gd4 moderately enlarged, an apomorphic condition).

On the other hand, the pretarsus I (30) is absent in Nodacaroides,

coniunctus. N. levis and Nj_ crinitus. whereas in N^. calidus, Nj.

brevispiritus and Rhodacaroides it is present; in Nj^ unguellus

pretarsus I has sessile claws. Also, scleronoduli (5) are present in

Nodacaroides. N. coniunctus and Njj. crinitus : they are absent in

Rhodacaroides and apparently, in levis. calidus, IL.

brevispiritus and unguellus. It is possible that these two

conflicting characters (30, 5) represent respectively, a less derived

condition of pretarsus I in calidus. brevispiritus and unguellus and

a secondary loss of scleronoduli in levis. calidus. brevispiritus and

unguellus. It should be noticed that ]L_ minvaspis exhibits reduction

in number of scleronoduli (it has only two) and that another

Rhodacaridae taxon, Latogamasellus. from South America, also exhibits

secondary loss of scleronoduli. Unfortunately, the male

characteristics of Rhodacaroides cannot be compared, since no males

of Nodacaroides are known.

Mediodacarellus new genus:

Type species: Afrogamasellus bakeri Hurlbutt, 1973.

This taxon is created to include only Afrogamasellus bakeri 87

Hurlbutt, 1973. When describing this species, Hurlbutt pointed out the characteristics it had in common with Afrogamasellus - ..."tectum triangular and broadest at base; third sternal pore slitlike; tarsus

I with basal rodlike seta short; dorsal setae on femur IV minute, not thickened; seta pv2 on tarsus IV forked.", and with Afrodacarellus -

..."granulations anterior to sternal shield and posterior to genital

shield; genital shield relatively narrow posteriorly.", the length of

its genital shield being "intermediate between the two genera". He

chose to include it in Afrogamasellus.

I have observed that M. bakeri (n. comb.) does possess a few

more features in common with Afrogamasellus and Afrodacarellus so

that it seems to truly represent an "intermediate taxon" between the

two genera and yet it keeps its identity. For instance, VL bakeri

possesses the following characteristics; male genital setae (st5) on

separate platelets (8) - a condition also found in Afrodacarellus and

Rhodacaropsis. except that these platelets are the endopodals shields

IV (this is unique within the Rhodacaroidea); peritrematal shields

are fused to the exopodal shields IV (17) and opisthonotals Z5 are

only slightly longer than J5 (23) - as Afrodacarellus and

T^fnpflmgspniis; first Sternal setae (stl) placed on preendopodals

(24) - as AfrnpamaHellus - but these shields are weakly sclerotized,

produced into an area of punctate sclerotization, closely resembling

the anterior sternal shield of Afrodacarellus; pretarsus I set on

short peduncle (30), an autapomorphy; female tectum with central tine

divided (32); seta pv2 on tarsus IV bifid (50) and female exopodals

III not coalesced (59), both characteristics of Afrogamasellus. Also, 88 setae Zvl are placed well above setae Jvl (55) - as in Afrodacarellus

(including ^ citri). A. lubalensis and Gen. n. #1. In view of its unique combination of characteristics, I've chosen to erect a new genus, Mediodacarellus. to include this species.

Gen. n. #1:

Even though no males are known, this taxon is distinctive enough to be placed in a category of its own. It reunites characteristics of a number of rhodacarid taxa such as: four scleronoduli on dha of podonotum (5); at times, the central pair is partly coalesced producing a three-scleronoduli effect; two pairs of preendopodal shields (14), as Rhodacaropsis and Solugamasus; peritrematal shields are fused to the exopodal shields IV (17), as Solugamasus.

Afrodacarellus. Latogamasellus and Nodacaroides: pretarsus I absent

(30), as in Rhodacarus. Solugamasus and Nodacaroides; female genital shield at least one and a half times as long as wide (54), as

Afrodacarellus. Rhodacarellus. Nodacaroides. Paragamasellevans.

Solugamasus and Litogamasus; opisthogastric setae Zvl placed at level well above that of setae Jvl (55), as Afrodacarellus; movable digit of female chelicera with 4-8 teeth (56), as in Latogamasellus and N. minvaspis: fixed digit of female chelicera with 6-12 teeth (57), as in N_j_ minvaspis. Solugamasus and Rhodacaropsis ; site of spermathecal opening on coxa IV (61), as Afrodacarellus. Its uniquely derived features include: podonotal glands gd4 slightly enlarged (21); females with 2 pairs of opisthonotals (36) and 3 pairs of opisthogastric setae (37) on soft cuticle. Character states 35 and 37 are reversals, derived secondarily probably due to reduction in size 89 of the idiosomal shields.

A few notes concerning the systematic position of Afrogamasellus mongii Hurlbutt, 1973, Afrogamasellus citri Loots, 1969,

Afrogamasellus lubalensis Loots, 1969 and Afrogamasellus mversi

Loots, 1969 are also presented here.

Afrogamasellus mongii;

This species was included in my studies as an incertae sedis taxon for it seemed to have some resemblance to ^ bakeri - the elongated shape of the female genital shield, the coarsely punctate sclerotization of the presternal area and the position of setae Zvl relative to setae Jvl, for instance. It turned out, however, that A. mongii does hold more characteristics in common with Afrogamasellus than with any other taxon: preendopodals shields are present, although not as well sclerotized (14); first sternal setae are placed on the preendopodal shields (24); peritrematals are not fused to exopodals IV (17) and the male genital setae (st5) are placed on the sternogenital shield (8). Except for character state 24, the other states are plesiomorphic in ^ mongii. However, ^ mongii further differentiates with two apomorphies: females with no opisthogastric setae on soft cuticle (37) - Afrogamasellus may have 0-1 pairs on soft cuticle; setae Zvl placed at level below that of setae Jvl (55)

- Afrogamasellus has the plesiomorphic condition. Therefore, A. mongii is retained in Afrogamasellus.

Afrogamasellus citri:

Loots (1969) included this species in Afrogamasellus "till a male or deutonymph are found which may indicate whether it belong 90

(sic) to Afrogamasellus or Rhodacarellus”. He indicated the resemblance of its endopodal shields with those of ^ camaxiloensis

Loots, 1969 and its peritremes, peritrematal, genital and metapodal shields as well as the femora II with those of A. reticulatus. Loots,

1969; the above mentioned species were later transferred to

Afrodacarellus by Hurlbutt (1973). In the same paper, Hurlbutt conceded that ^ citri had a "puzzling” systematic position for it combines a triangular tectum (as Afrogamasellus). a long rodlike seta on tarsus I and a granulated area behind the genital shield (as

Afrodacarellus). However, he pointed out that according to a numerical phenetic study ^ citri was "most similar to ^ bakeri which in turn is similar to ^ franzoides which is very close to the type species of Afrogamasellus. A. franzi”, and therefore decided to retain the species in Afrogamasellus.

I have examined the female specimen of ^ citri (sensu Hurlbutt) and found that this species is closer to Afrodacarellus (and to

Mediodacarellus) than to Afrogamasellus. It shares two synapomorphies with Afrodacarellus - first sternal setae placed on area of punctate sclerotization of the sternal shield (24), a condition independently derived in Rhodacarus and Rhodacarellus; female tarsus I with a solenidium half as long as the length of tarsus (51), an autapomorphy of Afrodacarellus. along with some apomorphic conditions found in other rhodacarids: postanal seta shorter than adanals (1), as

Latogamasellus and Afrogamasellus; usually 2 pairs of podonotals on soft cuticle (35), as Rhodacarus; one pair of opisthonotals on soft cuticle (36), as N. costai. A. citri does have a unique pair of 91

shields - the paragenitals, as Hurlbutt called them - which are

relatively large, triangular in shape (resembling boomerangs) each

bearing a genital pore. In addition, the opisthonotal setae R5 have migrated ventrally and are not placed on soft cuticle but rather, are

placed on narrow platelets originating from an extension of the

opisthonotal shield. Its tectum is more similar in appearance to that

of Afrogamasellus than to that of Afrodacarellus.

This species also seems to reunite distinctive enough characters

that could grant a "new genus" status to it. However, since no males

or immature stages are known as yet, I prefer to provisionally include ^ citri with Afrodacarellus , the taxon closest to it.

Afrogamasellus lubalensis;

Loots (1969) noted the resemblance of this species with A. camaxiloensis (the distribution of podonotal setae and chaetotaxy of the opisthogaster), with ^ reticulatus (ornamentation of shields) and with A. myersi (structure of the sterno-metasternal shield and the punctated presternal area). Hurlbutt (1973) considered A. lubalensis "puzzling", citing (as examples) its three-tined tectum

("unusual for Afrogamasellus or Afrodacarellus"). the resemblance of

its sternal shield with that of ^ myersi and the kind of hypostome

("found in many Afrodacarellus"). Karg (1977) created the subgenus A.

(Latogamasellus) with the species ^ (L.) squamosus Karg, 1977 and transferred ^ lubalensis and ^ myersi to this subgenus on the basis of the position of the first sternal setae (placed on well sclerotized portion of the sternal shield, but immediately behind an area of punctate sclerotization on this shield). 92

My study revealed that ^ lubalensis is closer to Latogamasellus and Afrodacarellus than to Afrogamasellus. even though ^ lubalensis presents a mosaic of features found in Afrodacarellus and

Latogamasellus. For example, the shape of the female genital shield is closer to that of Latogamasellus females; the shape of the female ventrianal shield is intermediate between the two genera, but the position of setae Zvl relative to setae Jvl (55) is the same as that seen in Afrodacarellus; the preendopodal shields are absent (14), as in Latogamasellus and Afrodacarellus; the sternal shield is weakly defined anteriorly (18), as in Latogamasellus and Afrodacarellus; the podonotals zl are one and a half times as short as setae jl (22), a plesiomorphic condition observed in Afrodacarellus; the opisthonotals

Z5 are at least one and a half times as long as setae J5 (23), a plesiomorphy of Afrogamasellus also; two pairs of opisthogastric setae are placed on soft cuticle (37), a reversed condition, also shared by Latogamasellus. Also, ^ lubalensis possesses 4 scleronoduli on the dha of the podonotal shield - as most rhodacarid genera, except for Latogamasellus which has lost them - and its pretarsus I is not pedunculate (30), whereas the pretarsi of

Afrodacarellus and Latogamasellus are pedunculate.

On the other hand, a unique combination of three of the features mentioned above places A^ lubalensis closer to Latogamasellus than to

Afrodacarellus. These are: preendopodal shields absent (14), female sternal shield with an area of punctate sclerotization at its anterior margin (18) and first sternal setae (stl) placed on well sclerotized area of the sternal shield (24). The states for 93 characters 14 and 18 are apomorphic and are shared by ^ lubalensis.

Afrodacarellus and Latogamasellus; however, the state for character

24 as expressed above is plesiomorphic and is shared by ^ lubalensis and Latogamasellus. whereas Afrodacarellus has the apomorphic condition where setae stl are placed on the area of punctate sclerotization of the sternal shield. It is precisely the derived condition of character 24 in Afrodacarellus which prevents A. lubalensis from being closer to it. Furthermore, no males or immatures of ^ lubalensis are known; this makes matters a bit more complicated - the structure of the male sternogenital shield and the position of its genital setae (st5) could provide evidence of relationships with any of these genera. On the basis of the available evidence at hand, I choose to retain this species in Latogamasellus until males are found.

Afrogamasellus myersi;

This species (not included in Fig. 3) was also provisionally placed in Afrogamasellus and its similarities with other taxa noted by Loots (1969): with A. reticulatus. the ornamentation of the dorsal shields; with A. citri. the peritremes, peritrematal and metapodal shields, the aspect of the gnathosoma, the anterior margin of the tectum, the general structure of tarsus I; with ^ camaxiloensis. the chaetotaxy of the opisthogaster. Like L. lubalensis (n. comb.), A. mversi possesses the same states observed in Latogamasellus for a number of characters. These are: preendopodals absent (14); sternal shield weakly sclerotized anteriorly (18); first sternal setae on the sternal shield (24); two pairs of opisthogastric setae on the soft 94 cuticle (37). In addition to these, ^ mversi shares with L. lubalensis; four scleronoduli on dha (5); pretarsus I not pedunculate

(30); with Latogamasellus; podonotals zl slightly shorter than jl

(22), and the shape of the female genital shield; with

Afrogamasellus; shape of the female ventrianal shield and the

position of setae Zvl in relation to setae Jvl (55). As it can be

noticed, ^ myersi (like L. lubalensis) also seems to exhibit

characteristics of three genera (Afrogamasellus. Afrodacarellus and

Latogamasellus). Because I find that this species shares more derived

characters with Latogamasellus (and because no males are known that

could indicate a higher degree of affinity with any of the remaining

two genera), I choose to provisionally retain ^ mversi in

Latogamasellus as L. mversi (n.comb.).

Family Gamasellidae:

From the ancestral lineage, two lines of descent originated. One

gave rise to the Pvriphis - Hevdeniella - Onchogamasus - Hiniphis

group of genera and is defined by seven apomorphies: female

ventrianal shield fused to exopodals shields IV (11); male

sternogenital shield fused to the ventrianal shield (12); female

ventrianal fused to the notai shield (15); peritremes reduced,

reaching up to anterior level of acetabula II (33); female genital

shield at least as wide as long (54); female sterno-metasternal

shield fused to endopodals IV (58); seta all on palp genu with at

least five prongs (60). The other originated the Rhodacaroides -

Periseius - Gamasellus - Pilellus - Evanssellus - Euepicrius -

Gamaselliphis group of genera and is defined by two apomorphies: 95 preendopodal shields produced into two pairs (14); seta all on palp

genu with two lateral prongs (60).

The group Pvriphis - Hevdeniella differentiated from its sister

Onchogamasus - Hiniphis with one autapomorphy: site of the

spermathecal opening on trochanter III (61), and one apomorphy:

dorsum of the idiosoma entire (49). Onchogamasus and Hiniphis in

turn, evolved two apomorphies: pretarsus I set on short peduncle (30) and female chelicerae with small and large teeth combined (34).

The group Rhodacaroides and Periseius evolved two apomorphies:

opisthogastric setae reduced to seven pairs (31), independently

derived in Hevdeniella and Evanssellus also; female chelicerae with

small and large teeth (34), which independently derived in the

Onchogamasus - Hiniphis group. Its sister Gamasellus - Pilellus -

Evanssellus - Euepicrius group, acquired one autapomorphy - podonotal

setae jl complex, barbed (13) and five apomorphies: adanal setae

placed at mid-level of the anal valves (10); dorsal setae of the

idiosoma predominantly simple except for a few complex setae

distributed on the podonotum and opisthonotum (29); submarginal (UR)

series represented by 1-3 pairs of setae (38); female genital shield

as wide as long or wider than long (54) and, seta all on palp genu

with at least five prongs (60).

The Gamasellus species complexes further differentiated with two

apomorphies - preendopodal shields produced into three or more pairs

(14); females with usually one pair of podonotal setae on soft

cuticle (35), this being a reversed condition and independently

derived in Onchogamasus as well. Furthermore, at least one species in 96 each of these complexes developed 4 scleronoduli on the podonotum, an autapomorphic condition within the family, and a parallelism with the

Rhodacaridae.

Its sister group (Evanssellus - Pilellus - Euepicrius -

Gamaselliphis) achieved one apomorphy; pretarsus I on a short peduncle, at most (30); this condition eventually evolved into the complete loss of pretarsus I in Evanssellus and Euepicrius. In addition, this group also achieved the reversed state of character 14

- the preendopodal shields were reduced to one well sclerotized pair

(the plesiomorphic state in its transformation series), ultimately producing the autapomorphic condition of Euepicrius where these shields became incorporated to the sternal shield. The Pilellus -

Evanssellus group developed a male sternogenital shield coalesced with the ventrianal shield (12), while its sister Euepicrius -

Gamaselliphis autapomorphically achieved a spermadactyl at least one and a half times as long as the movable digit of the chelicera (2) and apomorphically, the podonotal setae jl developed further complexity (13), the female ventrianal coalesced with the notai shield (15) and the female sterno-metasternal shield became partly fused to the endopodal shields IV (58).

Beginning with the Gamasellidae and proceeding through the

Ologamasinae, Gamasiphinae and Epiphidinae, there is a trend toward increased sclerotization, extension and consequently, fusion of idiosomal shields. This will become evident as I discuss them. First, however, some comments concerning the genera Onchogamasus. Hiniphis.

Rhodacaroides. Periseius. Gamasellus and Laelogamasus (the latter not 97 included in Fig. 4) are presented.

Onchogamasus Womersley, 1956b:

Lee (1970) recognized two species complexes in this genus: the communis complex, which included the type species 0^ communis

Womersley, 1956 and the pumilio complex including 0^ pumilio Lee,

1970 and Oj_ quasicurtipilus Lee, 1970. These groups could be distinguished by the fusion (or lack of) of the peritrematals to the exopodals, the splitting (or non-splitting) of exopodals II and III, and the angle formed by sternal setae st2-st4. He indicated that the genus had similarities with Gamasellopsis. Gamasitus and Sessiluncus and placed it in the Sessiluncinae Lee. In 1973, he described 0. virguncula. which combined characteristics of both complexes: the position of the female sternal setae was diagnostic of the communis complex, while the absence of fusion between the peritrematals and the exopodals was typical of the pumilio complex. Concurrently, 0. virguncula had some unique features: three pairs of preendopodals, dorsal shield divided and seta pd4 absent from tarsus IV (like the genus Gamasellopsis). The representatives of the two complexes have 1 pair of preendopodals which, although rarely, can be incorporated to the sternal shield; dorsal shield is entire and the leg chaetotaxy has the full complement. Lee then revoked his previous concept of two species complexes, adding that the attributes of 0. virguncula had lessened the gap between Onchogamasus. Gamasellopsis and Gamasitus and if males of the three genera (which are still not known) proved to be similar, "the possibly synonomy of these names should be considered". 98

I chose to Include 0^ communis and 0^_ virguncula as two separate entities in the computer runs to determine if there were any true affinities (apomorphically based) between them and to compare them with Sessiluncus and Gamasellopsis. Because Gamasitus is only known from a fragmented female specimen, I was not able to gather enough character states to include it in the computer runs. The results showed that Oj_ communis and Og_ virguncula are indeed closely related and should remain in the same genus. At this point, it is difficult to say if subgenera should be erected; because males are not known, I prefer to keep 0^ communis. 0. pumilio. 0. quasicurtipilus and 0. virguncula as species of Onchogamasus.

No affinities with Sessiluncus or Gamasellopsis were detected.

Sessiluncus was placed with the Gamasiphinae while Gamasellopsis was grouped with the Ologamasinae.

Hiniphis Lee, 1970:

Lee established this genus with a single species from Australia,

Hiniphis hinnus Lee, 1970. It combined the unususal features of a divided dorsal shield with an extensive fusion of the ventral shields in both males and females. His diagnosis included: divided dorsal shield (i. e.. a podonotal and an opisthonotal shield present); ventrianal shield broadly fused to the opisthonotal, peritrematal and exopodal IV shields on both sexes and to the sternogenital shield on the male; two pairs of preendopodals; dorsal setae simple; pretarsus

I pedunculate; opening to the spermathecae not known. Other characteristics of this species included: split in exopodals II and

III; female sterno-metasternal fused to endopodals II, III and IV; 8. 99 pairs of opisthogastric setae, none on soft cuticle; seta all on palp genu as a tapering spine and seta al2 slightly spatulate.

In 1973, he described Hiniphis bipala which, unlike IL. hinnus. had "...the dorsal and ventral shields widely separated by striated cuticle except for the narrow anterior fusion of the peritrematal and

podonotal shields..." in the female. Additionally, exopodals III not

split; 9 pairs of opisthogastric setae - one pair on soft cuticle;

seta all on palp genu spine-like with A pairs of short, lateral

prongs. No recognizable site of spermathecal opening was located. Lee

called attention to "...a pair of conspicuous pits near the anterior

margin of the opisthonotal shield..,", a unique feature within the

Rhodacaroidea. He remarked that although the males of both species

were similar, the females differed in the extent of sclerotization of

their shields; on the other hand, they shared attributes "...not

found together on other females of Ologamasinae, although they occur

together in Euepicrius (Gamasiphinae) and Onchogamasus virguncula

(Sessiluncinae)." These attributes were: fusion of the

sterno-metasternal with the endopodals IV and separate podonotal and

opisthonotal shields. Interestingly enough, all of the results

obtained through the computer runs placed Hiniphis. Euepicrius and

Onchogamasus (including Oj^ virguncula) in the same family group

(Gamasellidae). The fusion of the sterno-metasternal with the

endopodals IV (58), is one of the seven apomorphies uniting the

Pvriphis - Hevdeniella - Onchogamasus - Hiniphis group, which share

this character state with Euepicrius as well. 100

From my examination of both species, I found that H. hinnus has

4 scleronoduli on the dha of the podonotum, the site of the spermathecal opening Is on the postero-dorsal margin of coxa IV and the dorsal shield is entire, but with an apparent "groove of demarcation" between the podonotal and opisthonotal regions; a similar condition occurs in Rvkellus, an Ologamasinae genus. H. bipala in turn, lacks scleronoduli, the dorsal shield is divided, and the site to the spermathecal opening is (very probably) on the antiaxial margin of acetabulum IV, inserted between the exopodals IV

(which bear gland gv2) and a posterior extension of the peritrematal shield. The male of H. bipala does have a conspicuous pair of dorsal

"pits" (I am uncertain as to whether they are glands or pores) anteriorly on the opisthonotal shield, immediately posterior to podonotal setae z6). The female however, does not have these "pits" enlarged; it is possible that Lee interpreted the male dorsal pits and the female spermathecal ringed tubes as the same structures, since they look very similar.

Furthermore, in hinnus the podonotal setae zl are slightly shorter than setae jl, whereas in H. bipala. setae zl are almost two and a half times as long as jl - a parallelism with Periseius

(Psammonsella). The shape of the female genital shield is pointed anteriorly and rounded postero-laterally in Hj_ hinnus: in bipala it is rounded anteriorly and angled postero-laterally, exhibiting a subpentagonal appearance.

The differences between these two taxa seem to be more than simply interspecific. The different shapes of seta all on palp genu 101

and sites of spermathecal opening, for instance, should be given

important consideration, for these characteristics are usually very

stable within genera of Rhodacaroidea. The presence (or lack of) of

scleronoduli and enlarged dorsal glands/pores, as well as the

opposite development of setae zl, should also be taken into account.

On the basis of the observations made, it seems that in the future it

will be necessary to subdivide the genus Hiniphis into two subgenera.

Rhodacaroides Willmann, 1959;

In this study, Rhodacaroides is recognized as a monotypic genus.

Its species R^. aeevptiacus Willmann, 1959, is defined by four

autapomorphies - spermadactyl less than one and a half times as long as the movable digit of the chelicera (2); male genital setae (st5)

placed on soft cuticle (8); female sternal shield weakly defined posteriorly (18); opisthonotal setae J5 placed at level posterior to

that of setae Z5 (52) - and three apomorphies: dorsal gland gd4

slightly enlarged (21); pretarsus I set on short peduncle (30); females with one pair of opisthonotal setae usually set on soft cuticle (36), a reversal which also derived independently in species of the Gamasellus tragardhi complex; character state 52 is also independently derived in costai and Nj_ unguellus and character state 30 is independently derived in calidus and J N L _ brevispiritus; these four species have been previously allocated to this genus by

Lee (1970) and Karg (1977, 1979), respectively.

Lee and Hunter (1974) examined a species from the Crozet Islands and although they attributed it to Rhodacaroides. they noted that it was most similar to minvaspis from Australia and both were 102 probably congeneric with ^ costai from South America. Adding that

"...unfortunately no males of these three species are known...", they concluded by suggesting that these species should probably be grouped in a different genus than ^ aeevptiacus. On the basis of the results obtained, I am in full agreement with them. R. aeevptiacus is not related to any of the species previously included in the genus. As already discussed, the genus Nodacaroides has been elevated and expanded to include those species.

Periseius Womersley, 1961:

This genus is here recognized to include two subgenera: P.

(Periseius) - with the species (P. ) hammeni Womersley, 1961; P.

(P.) braziliensis Hirschmann, 1966 - and _P^ (Psammonsella) - with the species P^ (P.) nobskae Haq, 1965; and P^ (P. ) schusteri Hirschmann,

1966.

Even though maintaining the subgeneric categories for Periseius.

Lee (1970) suggested that it might be preferable not to recognize subgenera within this genus since Hirschmann*s key (1966) did not distinguish between P^_ (P.) nobskae and Pj. (P.) schusteri. two species of the genus Psammonsella Haq transferred by Hirschmann

(1966) to Periseius. with subgeneric status.

I have examined Pj. (P.) hammeni. P. (P.) littoralis (known only from deutonymphs and synonymized with Pj_ (P.) hammeni by Lee (1970)),

P. (P.) braziliensis and P^_ (P.) nobskae and found that P.

(Periseius) is defined by three autapomorphies and four apomorphies, whereas P^ (Psammonsella) is defined by three apomorphies. P.

(Periseius) autapomorphically expresses neotrichy through three 103 character states: genu III with 9 or 10 setae (43); tibia III with 9 setae (46) and femur IV with 7 setae (53) - this state is also independently derived in three non-related taxa (therefore, a convergence): mitis (Laelaptonyssidae), porosus (Ascidae) and

Gamasolaelaps bondwanensis Hurlbutt, 1983 (Veigaiidae). Its four apomorphies include: female ventrianal shield fused to the peritrematal shield (16); dorsal setae of the idiosoma predominantly simple except for a few complex podonotal and opisthonotal setae

(29); female genital shield as wide as long (54); female sterno-metasternal fused to endopodals IV (58). (Psammonsella) apomorphically possesses podonotal setae zl twice as long as setae jl

(22) - a rarely seen character state, independently derived in H. bipala: pretarsus I set on short peduncle (30) and female genital shield at least one and a half times as long as wide (54) - which

independently derived in Evanssellus also. In addition, the female metasternal setae (st4) are placed on the metasternal plates (7);

together with Epiphis (Epiphidinae), these are the only rhodacaroid

taxa possessing the plesiomorphic condition of this character.

Because (Periseius) and P^ (Psammonsella) have evolved such

distinctive character states, I prefer to retain them as separate

subgenera.

Gamasellus Berlese, 1892:

Lee (1970) recognized three species complexes within this genus

(falciger. discutatus and pyriformis), distinguished by the

chaetotaxy of the opisthonotum, number of opisthogastric setae placed

on soft cuticle, degree of fusion of the male ventrianal shield to 104 the opisthonotal and peritrematal shields, degree of fusion of the female ventrianal shield to the opisthonotal and exopodals IV shields, shape of seta all on palp genu and structure of pretarsus I.

I provisionally recognized a fourth species complex (tragardhi) formed by tragardhi. G. cophinus and G^ grossi (previously included in the falciger complex), on the basis of the following characteristics: all three species have 4 scleronoduli on the

podonotum (5) - G^ concinnus is the only species of the falciger

complex with scleronoduli; posterior margin of the female genital

shield extending well beyond level of acetabula IV (9) - this is not

the condition found in the falciger complex; podonotals jl at least

one and a half times as long as setae zl (22) - in the falciger

species, setae jl are only slightly longer than setae zl;

opisthonotals setae in 21 pairs (26) - falciger species have 22

pairs; usually 1 pair (G. cophinus with 2 pairs) of opisthonotals on

soft cuticle in females (36) - in falciger species there usually are

4 or 5 pairs of opisthonotals on soft cuticle; G. cooperi is the

exception, with 2 pairs on soft cuticle; submarginal (UR) series of

setae present in 1 pair (38) - falciger species have 2 pairs of

submarginals; podonotals J5 placed anteriorly to setae Z5 (52) - in

falciger species setae J5 are placed at level subequal to that of

setae Z5; finally, female genital shield is at least longer than wide

(54), in Gj_ cophinus is one and a half times as long as wide -

whereas in the falciger species this shield is as wide as long.

The four complexes were treated as distinct taxa in the computer

runs; the results of the analysis, however, indicate that in fact. 105 these groups of species are closely related while keeping their distinctive features. The pyriformis complex is the earliest derivative, defined by one autapomorphy - female with 2 pairs of opisthonotals on soft cuticle (36) - and four apomorphies; postanal seta slightly longer than adanals (1); female ventrianal fused to notai shield (15); opisthonotum with 23-27 pairs of setae (26); pretarsus I not pedunculate (30). Its sister discutatus - falciger - tragardhi differentiated with one autapomorphy - female with usually

2 pairs of opisthogastric setae on soft cuticle (37) - and one apomorphy - female with 3-6 pairs of opisthonotals on soft cuticle

(36), a secondarily derived condition.

The discutatus complex evolved a unique combination of character states, two of which are autapomorphic - opisthogaster hypertrichous, with at least 11 pairs of setae (31); dorsum of the idiosoma usually entire in both sexes, but male may have it divided (49). Its apomorphic states include: female ventrianal shield fused to exopodals IV (11); opisthonothum hypertrichous, with at least 29 pairs of setae (26) - a parallelism with Pilellus and Euepicrius; peritremes reduced, reaching to anterior level of acetabula II (33) - a parallelism with Hevdeniella and Hiniphis hinnus; movable digit of female chelicera with four teeth (56) - a parallelism with Euepicrius and Hiniphis bipala: female sterno-metasternal shield partly fused with endopodals IV (58) - a parallelism with Gamaselliphis; and seta all on palp genu bifid (60),a parallelism with Pvriphis. Pilellus and

H. hinnus. 106

The falciger - tragardhi sister complexes are defined by one apomorphy - podonotal gland gd4 moderately enlarged (21); this state

is independently derived in Rhodacaroides and Pilellus also. The

falciger complex autapomorphically achieved the condition where there

is at most, 1 pair of podonotal setae on soft cuticle (35). The

tragardhi complex in turn, achieved four apomorphic states; four

scleronoduli on dha of podonotum (5); adanal setae placed at level

anterior to the anal valves (10); females with 1 pair of opisthonotal

setae on soft cuticle (36), and seta all on palp genu at least

trifurcate (60). Of the characteristics I used to provisionally form

the tragardhi complex and compare it to the falciger complex,

character states 9 and 22 are plesiomorphic in tragardhi species;

character state 54 is apomorphic in cophinus but is plesiomorphic

in the other two species of the complex; additionally, character

state 3 (spermadactyl fused to movable digit of the chelicera through

most of its length) is also plesiomorphic in this complex. Therefore,

the recognition of the tragardhi complex seems justifiable at the

present. As a final note concerning the genus Gamasellus, there is a

trend towards reversal of certain character states (such as character

states 35-38), just as seen in the Rhodacaridae. It is intersting

that, outside of the Rhodacaridae, Gamasellus is almost the only

taxon (the other being hinnus) including species with fully

developed scleronoduli. 107

Laelogamasus Berlese, 1905;

I have not seen any specimens of this genus which Lee (1970) regarded as belonging to the tribe Gamasellini within the

Ologamasinae. There is one nominal species - Laelogamasus simplex

Berlese, 1905 - but Lee reports having seen at least two other

(unnamed) species. On the basis of the information available in the literature alone, it was not possible to obtain all the data necessary to include this genus in the computer analysis. However, there is evidence indicating definite relationships between this taxon and the Gamasellidae.

Laelogamasus shares the following apomorphic states with the

Gamasellidae: spermadactyl freer than fused to the movable digit of the chelicera (3); two or three pairs of preendopodal shields (14) - two pairs occur in Hiniphis and Rhodacaroides; three pairs are present in Gamasellus and Periseius: opisthonotum oligotrichous, with

25 pairs of setae (26) - a parallelism with the G^ pyriformis complex; dorsal setae of the idiosoma predominantly complex (29) - a parallelism with Evanssellus and Gamaselliphis; opisthogastric setae in 11-12 pairs (31) - a parallelism with the G. discutatus complex;

seta all on palp genu with at least five prongs (60) - a parallelism

with Hevdeniella. H. bipala. G. pyriformis complex. G. discutatus

complex and Evanssellus. Furthermore, Laelogamasus has the

submarginal (UR) series of setae present in five pairs (38) - a

plesiomorphic condition, also observed in the G^ pyriformis and G.

discutatus complexes as well as in Pilellus. Laelogamasus possesses

one autapomorphic condition: the site of opening to the spermathecae 108 is on the dorsal surface of coxa III (61).

Therefore, on the basis of the characteristics listed above, I place Laelogamasus in the Gamasellidae, as the probable sister group to the genus Gamasellus.

Family Ologamasidae

Subfamily Ologamasinae:

The Ologamasinae is here recognized to include ten genera seven of which were previously included in the Ologamasini of the

Ologamasinae sensu Lee and three are newly included: Antennolaelaps

Womersley, 1956; Queenslandolaelaps Womersley, 1956 (including Q.

berlesei Womersley, 1956) and Gamaellopsis Loots and Ryke, 1966.

These three genera have been transferred from the Sessiluncinae Lee,

1970.

The genus Rvkellus Lee, 1970 is the earliest derivative (and

sister to the remaining Ologamasinae) with two autapomorphies - male

sternogenital fused to the ventrianal shield (12); opisthogastric

setae Zvl placed at level well above that of setae Jvl, nearly

vertical to them (55) - and four apomorphies: ventral gland gv2

produced into a maximum of two pores (19); dorsal shield entire in

both sexes, but with an apparent line of demarcation between the

podonotal and opisthonotal regions of the idiosoma (49); opisthonotal

setae J5 placed at level subequal to that of setae Z5 (52); female

exopodal shields III entire (59). Its sister group Athiasella -

Hydrogamasellus - Ologamasus - Cvmiphis - Geogamasus -

Neogamasellevans - Antennolaelaps - Queenslandolaelaps -

Gamasellopsis is defined by four apomorphies: seven pairs of 109

opisthogastric setae (31); usually 1 pair of opisthogastric setae on

soft cuticle in females (37); opisthogastric setae Zvl placed at

level below that of setae Jvl (55); seta all on palp genu with at

least five prongs (60). The first taxon to differentiate in this

lineage is Athiasella. with 1 pair of opisthonotal setae usually on

soft cuticle in females (36) and the site of the spermathecal opening on trochanter III (61). The lineage which originated the remaining

Ologamasinae shares two apomorphies: females with no podonotal (35) or opisthogastric (37) setae placed on soft cuticle.

Hydrogamasellus has evolved one autapomorphy - spermathecal opening placed on coxa III (61) - and one apomorphy - female chelicerae composed of small and large teeth (34), a secondarily derived state which is also present in Ologamasus. Its sister lineage evolved two autapomorphies: dorsum of the idiosoma entire (49); fixed digit of the female chelicera with 6-12 teeth (57). From this lineage, two others arose: one in which the female tectum has a divided central tine (32), gave origin to the Ologamasus - Cvmiphis -

Geogamasus - Neogamasellevans group of genera. The other, is defined by a secondarily derived autapomorphy - females with 1 pair of podonotals on soft cuticle (35) - and three apomorphies: gland gv2 produced into 1-2 pores (19); dorsal setae of the idiosoma predominantly simple, except for a few complex podonotal and opisthonotal setae (29), a parallelism with the Ologamasus - Cvmiphis group; female exopodals III entire (59), a parallelism with Rvkellus; this lineage gave origin to the Antennolaelaps - Queenslandolaelaps -

Gamasellopsis group of genera. 110

The Ologamasus - Cvmiphis group has evolved two autapomorphies - female ventrianal shield fused to the notai shield (15); podonotal gland gd4 moderately enlarged (21), and two apomorphies: podonotal setae jl usually simple, but they may be complex at times (13) and dorsal setae of the idiosoma predominantly simple, with a few complex podonotal and opisthonotal setae (29). Ologamasus has differentiated from Cvmiphis by further achieving the autapomorphic development of gland gd4 into a conspicuous structure (21), while apomorphically combining small and large teeth in the female chelicerae (34). Its sister genus in turn, has three autapomorphies - female ventrianal shield fused to exopodal shields IV (11); dorsal setae of the idiosoma predominantly complex (29); seta all on palp genu bifurcate

(60), and one apomorphy: female sterno-metasternal shield fused to endopodals IV, a parallelism with Antennolaelaps and

Queenslandolaelaps. Its sister group, Geogamasus - Neogamasellevans. autapomorphically possesses peritremes reduced in length, reaching only up to mid-level of acetabula II (33) and apomorphically have a ventral gland gv2 with 1-2 solenostoma (19). The state 1 of gland gv2 has independently evolved three times in the Ologamasinae, with the general trend being towards reduction in the number of solenostoma.

Geogamasus has separated from its sister genus by autapomorphically achieving a spermadactyl which is recurved at distal level of the movable before projecting freely (4), a pretarsus

I set on short peduncle (30), dorsum of the idiosoma usually divided in both sexes but female may have it entire (49), and apomorphically possessing a spermadactyl which is at least one and a half times as Ill

long as the movable digit of the chelicera (2), a parallelism with

the Queenslandolaelaps - Gamasellopsis group of genera, and the site

of opening to the spermathecae is placed on trochanter III (61), a

parallelism with Athiasella. I was unable however, to find any

apomorphies which could define Neogamasellevans; instead, this taxon

is differentiated by two plesiomorphic character states: female

peritrematal shield not fused to the exopodals IV (17), a

symplesiomorphy with Antennolaelaps; opisthonotal setae Z5 at least

one and a half times as long as setae J5 (23) - a symplesiomorphy with Queenslandolaelaps.

It seems that Neogamasellevans is not a monophyletic taxon, for

although its species share a wide variety of derived character states, they sharply differ with respect to a few, but important

characters. For example, in the seven species attributed to the

genus, the length of the spermadactyl (2) may vary from shorter than

the movable digit (two species), to as long as the movable digit (two

species) to nearly three times as long as the movable digit (two

species) - the male of one species is not known; the first two

conditions are plesiomorphic whereas the latter is apomorphic.

Regarding the structure of the preendopodal shields (14), they may be

produced into one large, well sclerotized pair (four species) or two

pairs, one of them very small and placed anteriorly to the large pair

(three species); the second condition is apomorphic. The shape of

seta pv2 on tarsus IV (50) is simple in four species and complex

(bifurcate or trifurcate) in the other three; the first condition is

plesiomorphic and commonly found throughout the Rhodacaroidea, while 112 the complex shape is uniquely derived within the Ologamasidae and it has independently evolved in Afrogamasellus. Mediodacarellus

(Rhodacaridae) and Cyrtolaelaps (Euryparasitidae) as well. Finally, the site of opening to the spermathecae (61) may be on coxa IV (four species), coxa III (one species) or endopodals IV (one species); in

N. preendopodalis Loots & Ryke, 1967a (the type species), these openings could not be determined. All three conditions are apomorphic but the last one is unique within the Rhodacaroidea. Therefore, I am not ruling out the possibility that Neogamasellevans may indeed be a valid taxon. Even though at present Neogamasellevans cannot be apomorphically defined, I prefer to maintain it as a distinctive group for as more species are found, the complexity of seta pv2 on tarsus IV (50) may prove to be the rule rather than the exception.

Regarding the remaining Ologamasinae, Antennolaelaps is the first taxon to differentiate with two autapomorphic conditions - opisthogastric setae reduced to six pairs (31); females usually without opisthogastric setae placed on soft cuticle, but occasionally

1 pair may be removed from the ventrianal shield and be placed on the

integument (37) - and two apomorphic ones: opisthonotum

oligotrichous, usually with a maximum of 19 pairs of setae, rarely 20

pairs (26), a parallelism with Gamasellopsis; female

sterno-metasternal shield fused to endopodals IV (58), a parallelism

with Queenslandolaelaps. Its sister lineage in turn, is defined by

three apomorphies: opisthogastric setae in 8 pairs (31); female

tectum with central tine at least trifurcate (32), and opisthonotal

setae J5 placed at level subequal to that of setae Z5 (52), a 113

parallelism with Rvkellus. The taxon "berlesei" has achieved two

autapomorphies and four apomorphies - its relationships shall be

discussed below. The group Queenslandolaelaps - Gamasellopsis has

evolved two apomorphies: spermadactyl at least one and a half times as long as the movable digit of the chelicera (2), a parallelism with

Geogamasus: females with 1 pair of opisthogastric setae usually

placed on soft cuticle (37), a parallelism with Rvkellus and

"berlesei". Queenslandolaelaps further developed two apomorphic states: females with 1 pair of opisthonotal setae on soft cuticle

(36), a parallelism with Athiasella: female sterno-metasternal shield fused to endopodals IV (58).

Gamasellopsis has achieved a number of uniquely derived states

(eight) but it still shares some apomorphict conditions with several other taxa. Its relationships will be discussed below, followed by some notes on the relationships of Ologamasus. "berlesei". and

Gamasitus (which is not included in Fig. 5).

Gamasellopsis Loots & Ryke, 1966:

Lee (1970) regarded this genus as a member of his Sessiluncinae and noted that there were similarities between it and the genera

Onchogamasus. Gamasitus and Sessiluncus.

My results have shown that Gamasellopsis is a member of the

Ologamasinae, defined by eight autapomorphies and three apomorphies.

Its uniquely derived characteristics (within the Ologamasinae) are:

preendopodal shields incorporated to the sternal shield (14), a

parallelism with Sessiluncus (Gamasiphinae), Euepicrius

(Gamasellidae) and Gamasitus (incertae sedis); podonotal setae zl 114 absent (22), a convergence with P. chinensis (Laelaptonyssidae); females with 2 pairs of opisthonotal setae on soft cuticle (36), a parallelism with Sessiluncus; genu I with 12 setae (42), a convergence with Puchihlungia. Panteniphis and Digamasellidae, all members of the Laelaptonyssoidea; genu III with 8 setae (43), a parallelism with Gamasitus and a convergence with Puchihlungia

(Laelaptonyssidae), Digamasellus (Digamasellidae), Panteniphididae,

Protogamasellus and Gamasellodes (Ascidae); genu IV with 8-9 setae

(44), a parallelism with Gamasitus and with Sessiluncus. Gamasiphis,

Gamasiphoides and Hvdrogamasus (Gamasiphinae), as well as a convergence with several genera of Ascidae; tibia IV with 9 setae

(47), a parallelism with Hvdrogamasus and a convergence with

Protogamasellus (Ascidae); finally, tarsus IV with a total of 17 setae (48), a parallelism with seven taxa of Rhodacaridae and

Onchogamasus virguncula (Gamasellidae) and a convergence with

Laelaptonvssus (Laelaptonyssidae). Its apomorphic conditions include: podonotal setae jl usually simple, although they may be complex at times (13) - a parallelism with the Ologamasus - Cvmiphis group, and a convergence with Lasioseius (Ascidae); opisthonotum oligotrichous, with 16-19 pairs of setae (26). The state "3" for this character is also observed in Rhodacaropsis. Rhodacarellus. Nodacaroides

(Rhodacaridae); Antennolaelaps (Ologamasinae); Gamasiphis,

Hvdrogamasus and Parasitiphis (Gamasiphinae); and opisthogastric setae in 7 pairs (31), an apomorphic condition observed throughout the Rhodacaroidea. As it is evident, Gamasellopsis shares many derived characters with a wide variety of taxa, including non-related 115 ones. However, its closest relationships are with members of the

Ologamasidae as indicated by the number of genera sharing apomorphies with it. Therefore, I regard Gamasellopsis as a taxon of the

Ologamasinae and Queenslandolaelaps as its sister group with which it shares seventeen apomorphic states.

Ologamasus Berlese, 1888:

The morphological diversity encountered in this genus has incited taxonomic reviews by Lee (1966; 1970), Karg (1976) and Antony

(1980), as well as phylogenetic studies to determine its internal and external relationships (Antony, 1980; Antony and Johnston, 1984).

Because Ologamasus exhibits such an elasticity of characteristics, it has been confused with Gamasiphis (Ryke, 1961b). Also, Lee (1966) described seven species as belonging to this genus and later (1970) found that six of those species had to be accommodated in a newly established genus, Cvmiphis Lee, 1970, and the seventh species in the genus Pyriphis Lee, 1970. Lee (1970) also pointed out the similarity of appearance between females of Ologamasus and Hydrogamasellus and between females of Hydrogamasellus and Heydeniella, adding that where only females were available, alloting a species to one of these genera could be very difficult.

Antony (1980) preliminarily found that Athiasella Lee, 1973 appeared to the sister group of Ologamasus. This conclusion is revoked here and Cvmiphis is regarded as the sister group of

Ologamasus on the basis of the results obtained in this considerably more extensive (and inclusive) analysis. 116

Queenslandolaelaps berlesei Womerslev. 1956;

Lee (1970) removed this species from the genus

Queenslandolaelaps which he attributed to the Sessiluncinae, stating

that "berlesei" belonged to the Ologamasinae and although uncertain of its relationships, he provisionally transferred it to the genus

Neogamasellevans until information on the male anatomy was known. I

concur with the placement of this species in the Ologamasinae, but

not with its transfer to Neogamasellevans «

The results of this analysis indicate that "berlesei" is more

closely related to Queenslandolaelaps than to Neogamasellevans. In

fact, "berlesei" seems to represent an intermediate taxon between

Antennolaelaps and Queenslandolaelaps. It is defined by two

autapomorphies - preendopodal shields produced into two pairs (14);

female tarsus I with a solenidium half as long as the length of

tarsus (51), a parallelism with Afrodacarellus (Rhodacaridae) and

Sessiluncus (Gamasiphinae), and four apomorphies: no podonotal setae

placed on soft cuticle (35); there are 2 pairs of opisthogastric

setae on soft cuticle (37), a parallelism with Rvkellus; dorsum of

the idiosoma entire, but with an apparent line of demarcation between

the podonotal and opisthonotal regions (49), a parallelism with

Rvkellus and with Gamasellevans « some species of Gamasiphis and

Gamasiphoides (Gamasiphinae) and Hiniphis (H.) hinnus (Gamasellidae);

finally, female exopodal shields split (59). It is likely that

"berlesei" may represent a taxon distinctive from Queenslandolaelaps,

but until the male is known, I prefer to retain it within the genus

in which it was originally included. 117

Gamasitus Womersley, 1956:

This monotypic genus (which Lee, 1970 placed in his

Sessiluncinae) is known from one fragmented and incomplete female specimen (Gamasitus obscurus Womersley, 1956), which I was able to examine but could hardly extract any additional information to complement that already provided by Lee. Therefore, my efforts to disclose its relationships are tentative.

Gamasitus seems to be more closely related to the Ologamasinae, with which it shares sixteen derived characters, than to the

Gamasellidae, with which it shares fourteen derived characters.

Within the Ologamasinae, Gamasitus shares three derived states with

Gamasellopsis: preendopodals incorporated to the sternal shield (14);

genu III with 8 setae (43); genu IV with 9 setae (44). If Gamasitus

is excluded, these three character states are autapomorphic for

Gamasellopsis. Gamasitus also shares one character state with

Antennolaelaps (which otherwise is autapomorphic for the genus) -

opisthogastric setae reduced to 6 pairs (31). Other apomorphic states

of Gamasitus include: opisthogastric setae Zvl placed at level below

that of setae Jvl (55) — observed in most Ologamasidae taxa, except

for Rvkellus. Cvmiphis and Epiphis); fixed digit of the female

chelicera with six teeth (57) - observed in Ologamasus, Cymiphi^,

Neogamasellevans Antennolaelaps and Queenslandolaelaps; female

sterno—metasternal shield fused to endopodals IV (58) — observed in

Cvmiphis. Antennolaelaps and Queenslandolaelaps. Finally, Gamasitus

has the movable digit of the female chelicera with four teeth (56) -

a uniquely derived condition within the Ologamasinae. On the basis of 118 the evidence presented above, I therefore place Gamasitus in the subfamily Ologamasinae within the family Ologamasidae.

Subfamilies Epiphidinae and Gamasiphinae:

These two subfamilies share fourteen apomorphies. The

Epiphidinae (here represented by the genus Epiphis) is the first taxon to differentiate from its sister lineage with three autapomorphies - number of opisthogastric setae in at least 11 pairs

(31), a parallelism with the G. discutatus complex (Gamasellidae) and a convergence with Veigaia (Veigaiidae); peritreraes reduced, reaching up to mid-level of acetabula II (33), a parallelism with

Rhodacaridae; opisthogastric setae Zvl placed at level well above that of setae Jvl (55), a parallelism with Afrodacarellus.

Mediodacarellus (Rhodacaridae), Rvkellus (Ologamasinae), and a convergence with Protogamasellus (Ascidae) - and three apomorphies: spermadactyl at least one and a half times as long as the movable digit of the chelicera (2), a parallelism with the Gamasiphis -

Sessiluncus - Gamasellevans group of genera (Gamasiphinae),

Geogamasus. Queenslandolaelaps. Gamasellopsis (Ologamasinae),

Euepicrius. Gamaselliphis (Gamasellidae), Cyrtolaelaps.

Eurvparasitus. Allogamasellus (Euryparasitidae), Rhodacarellus.

Paragamasellevans. Solugamasus (Rhodacaridae), and a convergence with

Laelaptonvssus (Laelaptonyssidae); seta all on palp genu finely pilose (60), also independently derived in Sessiluncus

(Gamasiphinae), Heterogamasus. Eurvparasitus (Euryparasitidae) and

Litogamasus (Rhodacaridae); site of opening to the spermathecae on coxa III (61), independently derived in Latogamasellus 119

(Rhodacaridae), Hydrogamasellus (Ologamasinae), Sessiluncus

(Gamasiphinae) and in several taxa of Ascidae (therefore, a convergence) as well.

Its sister lineage (which gave origin to the Gamasiphinae) shares five autapomorphies: female peritrematal shields fused to the exopodals IV (17); ventral gland gv2 produced into a maximum of two pores (19); opisthogastric setae in a maximum of 8 pairs (31); female chelicera with small and large teeth combined (34); opisthogastric setae Zvl placed at level below that of setae Jvl (55), and two apomorphies: postanal seta shorter than the adanal setae (1) and peritremes reduced, reaching the anterior margin of acetabula II

(33).

Within this lineage, Caliphis is the sister group to the remaining Gamasiphinae and the first taxon to differentiate, achieving two autapomorphies: adanal setae placed at mid-level to the anal valves (10); opisthonotum hypertrichous, with at least 29 pairs of setae (26), and two apomorphies: female sterno-metasternal shield fused to the endopodals IV (58), a parallelism with the Gamasiphis -

Sessiluncus - Gamasellevans group; seta all on palp genu bifurcate

(60), a parallelism with Hvdrogamasus. Cvmiphis (Ologamasinae),

Pyriphis. H. hinnus. Pilellus and Gj_ discutatus complex

(Gamasellidae). Its sister lineage has autapomorphically attained seven pairs of opisthogastric setae (31), submarginal (UR) series absent from the opisthonotum (38) and apomorphically possesses an oligotrichous opisthonotum, with a maximum of 22 pairs of setae (26).

This lineage further split into two others, giving origin on one side 120 to the Gamasiphis - Sessiluncus - Gamasellevans group of genera which is defined by two autapomorphies: adanal setae placed at level anterior to the anal valves (10); female ventrianal shield fused to the exopodals IV (11); and four apomorphies: spermadactyl at least one and a half times as long as the movable digit of the chelicera

(2); pretarsus I set (at most) on short peduncle (30); genu IV with 8 or 9 setae (44), a condition independently derived in Gamasiphoides.

Hvdrogamasus and also in Gamasellopsis (Ologamasinae) and Gamasitus; female sterno-metasternal shield fused to endopodals IV (58), a parallelism with Caliphis; Cvmiphis. Oueenslandolaelaps.

Antennolaelaps (Ologamasinae); Heydeniella. Hiniphis. Onchogamasus.

Periseius (Periseius) and Euepicrius (Gamasellidae).

On the other side, the Gamasiphoides - Laelaptiella -

Hvdrogamasus - Parasitiphis group of genera originated, with one autapomorphy, preendopodals produced into two pairs (14), and one apomorphy: fixed digit of the female chelicera with 6-12 teeth (57), a condition also independently derived in Gamasiphis and several other rhodacaroid taxa including five genera of Ologamasinae.

Gamasiphis has separated from its sister Sessiluncus -

Gamasellevans with one autapomorphy, female tectum with central tine divided (32), and three apomorphies: opisthonotum oligotrichous, usually with a maximum of 19 pairs of setae (26); movable digit of the female chelicera with four teeth (56) and fixed digit with seven teeth (57). Its sister group in turn, has lost the peduncle on tarsus

I (30). Sessiluncus is further defined by five autapomorphies (14,

31, 34, 36, 51) and two apomorphies (4, 60), while its sister 121

Gamasellevans is defined by one autapomorphy (37) and three apomorphies (49, 54, 59).

The Gamasiphoides - Laelaptiella group autapomorphically possesses the podonotal glands gd4 enlarged (21), a parallelism with

Ologamasus (Ologamasinae) and Allogamasellus and Acugamasus

(Euryparasitidae). Its sister Hvdrogamasus - Parasitiphis group autapomorphically has podonotal setae J5 placed at level subequal to that of setae Z5 (52), while apomorphically exhibiting an oligotrichous opisthonotum with a maximum of 19 pairs of setae (26), a parallelism with Gamasiphis. and the female movable digit of the chelicera has a maximum of eight teeth (56). Gamasiphoides has achieved three uniquely derived character states (35, 58, 59), along with four other derived states (36, 37, 44, 49). Its sister

Laelaptiella has one uniquely derived character state (2).

Hvdrogamasus possesses two autapomorphies (8, 47) and three apomorphies (4, 44, 60), while its sister Parasitiphis possesses two autapomorphies (37,38) and five apomorphies (30, 31, 36, 54, 60).

Within the Gamasiphinae, Hvdrogamasus and Parasitiphis have the

least extensive sclerotization of shields. They also exhibit a trend

towards reversal to the plesiomorphic state of certain characters

which otherwise would represent autapomorphies of the Gamasiphinae.

For instance, the postanal seta (1) is shorter than the adanal setae

in all Gamasiphinae, except in Hvdrogamasus and Parasitiphis; the

peritremes (33) are reduced in all Gamasiphinae (and Epiphidinae)

except for these two genera. In addition, the opisthonotal setae Z5

are, at most, slightly longer than setae J5 (23) in both Gamasiphinae 122

and Epiphidinae whereas in Hvdrogamasus and Parasitiphis. setae Z5

are at least one and a half times as long as setae J5 (the

plesiomorphic state for the character).

I have tried to understand (and find a viable explanation for)

the reduction in extent and/or degree of sclerotization of the

shields of these two taxa, members of a subfamily characterized by an

extensive fusion of idiosomal shields. The only factor common to both

genera (aside from their synapomorphies) is that they have been

collected in or near littoral zones. With that in mind, I then looked

for other taxa which have been collected in similar environments and

found that Litogamasus. Rhodacaropsis (Rhodacaridae), Periseius and

Rhodacaroides (Gamasellidae) - found in similar habitats - all

exhibit reduction in extent of idiosomal shields and/or reduction in

the degree of sclerotization of shields (the latter condition is

observed in Periseius). Regarding Litogamasus. there is no reduction

in degree of sclerotization, only in extent of sclerotization. As for

Rhodacaropsis. I would not have expected extensive shields for this

is not common to Rhodacaridae; however, its shields are less

extensive (as well as weakly sclerotized) than expected for that

family, Periseius and Rhodacaroides belong to a family (Gamasellidae)

of which most of its representatives are heavily and extensively

sclerotized and again, the same unexpected reduction in degree and

extent of idiosomal sclerotization is noticed. I am still unable to

find an explanation for such a trend but it appears that there is an

association of littoral habitats with a reduction in degree and

extent of sclerotization of shields. 123

As the final part of this section, some notes on the definition of the Epiphidinae are presented below.

Subfamily Epiphidinae Kethley, 1983:

Kethley (1983) established the Epiphidinae based on the genus

Epiphis Berlese, 1916 and included the genera Iphidosoma Berlese,

1892 and Stvlochirus G. & R. Canestrini, 1882, sensu Lee 1970, (=

Phvsallolaelaps Berlese, 1908, Periphis Berlese, 1914 and Megaliphis

Willmann, 1938, but not Epiphis Berlese). He placed it in the

Rhodacaridae sensu Lee (= Rhodacaroidea) and listed a series of characters considered "uniquely derived with respect to other

Rhodacaridae": "dorsal sclerite of the deutonymph holonotal, three or more pairs of dorsal setae elongate, tarsus I with an acrotarsal pseudosegmentation; exopodal sclerites II-IV of the adults are continuous with each other, but separate from the peritrematal sclerite, holonotal sclerite continuous with the ventrianal region posteriorly and the peritrematal sclerite at least anterolaterally, Sad “ss sensillum."

I agree that this group of genera is distinctive enough to merit

being placed in a category of its own; however, not all the

characters listed above are unique to this subfamily. For example,

the acrotarsal pseudosegmentation on tarsus I of the deutonymph is a

condition also observed in deutonymphs of the genera Cyrtolaelaps and

Eurvparasitus (Gamasellidae). The presence of three or more pairs of

elongate dorsal setae on the deutonymph is also observed in the

deutonymphs of Gamasellus discutatus and Gj. tragardhi; it should be

noted that immature instars of rhodacaroid mites are poorly known.

The important point to be noticed here is that in Epiphis. these 124 setae are elongate and slightly barbed in the deutonymph, whereas in the adult they are simple and not elongate. In the Gamasellus deutonymphs, these setae are also elongate and complex (barbed, pilose spatulate) and although their adult instars possess all dorsal setae of approximate lengths, the complexity of some of those setae

(previously elongate in the deutonymphal instar) is maintained. The condition in which the adult exopodal shields II-IV are coalesced with one another but are separate from the peritrematal shields is also observed in Antennolaelaps (Ologamasinae) and Sessiluncus

(Gamasiphinae). The fusion of the holonotal shield with the ventrianal shield posteriorly and with the peritrematal anterolaterally is also observed in Pyriphis. Hiniphis hinnus. some

Heydeniella species (Gamasellidae), Cvmiphis. some Ologamasus and

Hydrogamasellus species (Ologamasinae), Gamasiphis. Laelaptiella media and some Gamasiphoides species (Gamasiphinae). The presence of a large, bulbous sensillum on the tarsus I of the male is also observed in Afrodacarellus camaxiloensis (Rhodacaridae) and two non-related taxa, Panteniphis and Lindquistoseius (Panteniphididae) exhibit the same condition.

Therefore, there are only a few unique characters left to

diagnose the Epiphidinae; nevertheless, they are of the utmost

importance. The presence of a holonotal shield in the deutonymph is

unknown in other Rhodacaroidea. A greatly elongate spermadactyl is

found in other Rhodacaroidea (Geogamasus. Euepicrius. Gamaselliphis

and Eurvparasitus. for instance), but these are never styletiforra, or

with a very reduced fixed digit and an absent corona. The two genera 125 of Laelaptonyssidae, a non-related taxon, lack a corona in their chelicerae and one of them, Laelaptonvssus. has a very elongate spermadactyl which is nearly styletiform but it differs from that of

Epiphis by being strongly recurved rather than straight. Another important characteristic of the deutonymph is the presence of a

distal, hyaline process on the fixed digit of the chelicera, which

Kethley described and illustrated but did not use as a diagnostic

character of the Epiphidinae.

Lastly, because the large, bulbous sensillum on tarsus I is so

rarely found in other Rhodacaroidea, this character should also be

considered one of the typical features of the Epiphidinae, when used

in conjunction with the other characteristics listed above and on

Fig. 6.

The newly proposed classification of the Rhodacaroidea and

Laelaptonyssoidea resulting from this analysis is given on Table 7. 126

Table 7. Proposed classification of the Rhodacaroidea and Laelaptonyssoidea. The arrangement of families is phyletically sequenced.

Superfamily Rhodacaroidea Oudemans, 1902

Family Euryparasitidae new family

Type Genus: Eurvparasitus Oudemans, 1902

Genus Cyrtolaelaps Berlese, 1887 (=* Protolaelaps Tragardh, 1912) Eurvparasitus Oudemans, 1902 Heterogamasus Tragardh, 1907 Acugamasus Lee, 1970 A. punctatus species complex Lee, 1970 A. natalensis species complex Lee, 1970 Allogamasellus Athias-Henriot, 1961 Notogamasellus Loots & Ryke, 1966 Subgenus (Notogamasellus) Loots & Ryke, 1966 N. (Podonotogamasellus) Loots & Ryke, 1966

Family Rhodacaridae Oudemans, 1902

Type Genus: Rhodacarus Oudemans, 1902

Genus Litogamasus Lee, 1970 Rhodacaropsis Willmann, 1935 Rhodacarus Oudemans, 1902 Rhodacarellus Willmann, 1935 Paragamasellevans Loots & Ryke, 1968 Solugamasus Lee, 1973 Gen. n. #1 Nodacaroides Karg, 1977 new status (= Rhodacaroides (Nodacaroides) Karg, 1977) (= Rhodacaroides (Tenacaroides) Karg, 1977) Afrogamasellus Loots & Ryke, 1968 (= ^ (Podalogamasellus) Karg, 1977) (= ^ (Jugulogamasellus) Karg, 1977) Mediodacarellus new genus Afrodacarellus Hurlbutt, 1973 (= Afrogamasellus (Foliogamasellus) Karg, 1977) Latogamasellus Karg, 1977 new status 127

Table 7. (Continued)

Family Gamasellidae Hirschmann, 1962 new status

Type Genus: Gamasellus Berlese, 1892

Genus Pyriphis Lee, 1970 Heydeniella Richters, 1907 Onchogamasus Womersley, 1956 Hiniphis Lee, 1970 Rhodacaroides Willmann, 1959 Periseius Womersley, 1961 Subgenus P.. (Periseius) Womersley, 1961 P. (Psammonsella) Haq, 1965 Genus Laelogamasus Berlese, 1905 Gamasellus Berlese, 1892 G. pyriformis species complex Lee, 1970 G. discutatus species complex Lee, 1970 G. falciger species complex Lee, 1970 G. tragardhi species complex new status

Genus Pilellus Lee, 1970 Evanssellus Ryke, 1961 Euepicrius Womersley, 1942 Gamaselliphis Ryke, 1961

Family Ologamasidae Ryke, 1962

Subfamily Ologamasinae Ryke, 1962

Type Genus: Ologamasus Berlese, 1888

Genus Rvkellus Lee, 1970 Athiasella Lee, 1973 Hydrogamasellus Hirschmann, 1966 Ologamasus Berlese, 1888 (= Ologamasellus Berlese, 1914) Cvmiphis Lee, 1970 Geogamasus Lee, 1970 Neogamasellevans Loots & Ryke, 1967 Antennolaelaps Womersley, 1956 Queenslandolaelaps Womersley, 1956 Gamasellopsis Loots & Ryke, 1966 Gamasitus Womersley, 1956 128

Table 7. (Continued)

Subfamily Epiphidinae Kethley, 1983

Type Genus: Epiphis Berlese, 1916

Genus Epiphis Berlese, 1916 Iphidosoma Berlese, 1892 Stvlochirus G, & R, Canestrini, 1892

Incertae sedis: Phvsallolaelaps Berlese, 1908 Periphis Berlese, 1914 Megaliphis Willmann, 1938

Subfamily Gamasiphinae Lee, 1970

Type Genus: Gamasiphis Berlese, 1904 (= Micriphis Berlese, 1914) (= Heteroiphis Tragardh, 1952) (= Neogamasiphis Tragardh, 1952)

Genus Caliphis Lee, 1970 Gamasiphis Berlese, 1904 Sessiluncus Canestrini, 1898 Gamasellevans Loots & Ryke, 1967 Gamasiphoides Womersley, 1956 Laelaptiella Womersley, 1956 Hvdrogamasus Berlese, 1892 Parasitiphis Womersley, 1956 (= Austrohvdrogamasus Hirschmann, 1966)

Superfamily Laelaptonyssoidea Womersley, 1956 new status

Family Digamasellidae Evans, 1957

Type Genus: Digamasellus Berlese, 1905

Genus Dendrolaelaps Halbert, 1915 Dendrolaelaspis Lindquist, 1975 Dendroseius Karg, 1965 Digamasellus Berlese, 1905 Insectolaelaps Shcherbak, 1980 129

Table 7. (Continued)

Genus Longoseius Chant, 1961 Subgenus L. (Longoseius) Chant, 1961 L. (Longoseiulus) Lindquist, 1975 Genus Multidendrolaelaps Hirschmann, 1974 Oligodentatus Shcherbak, 1980 Grientolaelaps Bregetova & Shcherbak, 1977

Family Panteniphididae new family

Type Genus: Panteniphis Willmann, 1949

Genus Lindquistoseius Genis, Loots & Ryke, 1969 Panteniphis Willmann, 1949

Family Laelaptonyssidae Womersley, 1956

Type Genus: Laelaptonvssus Womersley, 1956

Genus Laelaptonvssus Womersley, 1956 Puchihlungia Samsinak, 1964 SYSTEMATIC ACCOUNTS

The superfamily Rhodacaroidea Oudemans comprises four families:

Euryparasitidae new family; Rhodacaridae Oudemans, 1902; Gamasellidae

Hirschmann, 1962, new status; and Ologamasidae Ryke, 1962. The

Ologamasidae includes three subfamilies: Ologamasinae Ryke, 1962,

Gamasiphinae Lee, 1970 and Epiphidinae, Kethley, 1983.

DIAGNOSIS

Palp claw three-pronged; dorsal sclerotization varying from

separate podonotal and opisthonotal shields, to partially separate

shields (incompletely fused), to an entire (holonotal) shield;

ventral sclerotization variable, with usually more extensive shields

in the male; femur I with 13 setae including four ventrals; tibia I

with 14 setae including four ventrals; podonotal region with 20-23

pairs of setae, opisthonotal region with 20-22 pairs of setae and

opisthogastric region with 6-8 pairs of setae, but variations on

these patterns (reduction or increase in number) also occur;

submarginal (UR) series of setae commonly absent - if present, there

usually are 1-3 pairs of them and very rarely, up to 9 pairs may be

present; leg chaetotaxy usually follows that of the Rhodacaridae

sensu Evans (1963, excluding the genera Asca. Digamasellus and

Halolaelaps); peritrematal shield usually fused to exopodal IV

130 131 shields; peritremes often reduced in length, reaching up to anterior or midle level of acetabula II; almost always, a ventrianal shield is present and the anal valves are small in all postembryonic instars; adanal setae often placed at anterior or midle level to the anal valves; ventral gland gv2 usually present in 1-3 solenostoma; female with sternal and metasternal shields fused, so that the metasternal setae (st4) are almost always placed on a sterno-metasternal shield; female genital shield usually rounded anteriorly and almost always truncate posteriorly; accessory opening to the spermathecae usually associated with leg IV (coxa or acetabulum); female chelicerae usually with three teeth on the movable digit and five teeth on the fixed digit; male with a sternogenital shield always, which usually carries the genital setae (stS); male spermadactyl distally free and of various lengths; male leg II with antero-ventral (av) seta almost always modified into a conspicuous, thumb-like spur and seta av of genu and tibia often modified into a small spine; finally, almost always, deutonymph with separate dorsal shields.

DESCRIPTION

Size variable, ranging from small (less than 400u) to large

(more than 1200u) mites.

Gnathosoma: Palp claw always three-pronged, although the basal prong may be reduced (as in Heterogamasus and Evanssellus); seta all on palp genu variously shaped (simple, finely pilose, spatulate, bifurcate or with 2-15 lateral prongs); tectum usually triramous, with central tine varying from smooth on sides and undivided at its tip, to smooth and with a tip which may be bifurcate, trifurcate or 132 surrounded by spinules, to laterally denticulate with an undivided or divided tip; salivary stylets always well developed; corniculi almost always well developed (except for the very reduced corniculi of

Onchogamasus communis Womersley, 1956) and not divided (except for the bifid corniculi of Geogamasus furcatius Karg, 1976 and

Paragamasellevans vandenbergi Loots & Ryke, 1968); female chelicerae usually with small and large teeth, but chelicerae with predominantly large teeth are often observed and less commonly, chelicerae with predominantly small teeth; movable digit of the female chelicera usually with 3 teeth, although up to 8 teeth is not uncommon to find in representatives of all four families, and very rarely, there may be 3 or 4 large teeth plus up to 15 small teeth present (as in

Eurvparasitus and Cvmiphis): the fixed digit of the female chelicera usually has 5-7 teeth but is also subject to variability and up to 12 teeth (Rhodacaroides. Periseius. Hiniphis. Parasitiphis and some species of Rhodacarus) or even 19 teeth (Pyriphis) may be present; male spermadactyl always distally free, of various shapes (straight, basally or distally recurved, or rarely, curving around the movable digit as in Litogamasus and Acugamasus) and lengths which vary from shorter than the movable digit to at least one and a half times as long as the movable digit; arthrodial process of the chelicera is usually produced into a conspicuous corona but it may be modified into an arthrodial brush (as in the genera Afrogamasellus.

Afrodacarellus. Mediodacarellus. Latogamasellus and Notogamasellus); in the genus Gamasellevans. the membranous process of the female chelicera is a corona while in the male the process is modified into 133 a long, leaf-like flap; in the male of Queenslandolaelaps vitzthumi

Womersley, 1956 there is an extra, fimbriated flap between the spermadactyl and the corona.

Dorsum: Dorsal shield of deutonymphs is almost always divided except for the deutonymphs of the Epiphidinae which have a holonotal shield; dorsal shield of adults usually divided (Euryparasitidae,

Rhodacaridae, most Gamasellidae taxa), but a holonotal shield is often present (most Ologamasidae taxa); less frequently observed is the presence of extensively sclerotized shields which are separated by a groove or line of demarcation between the podonotal and opisthonotal regions (Rvkellus and some species of Geogamasus.

Gamasiphoides. Gamasiphis and Hiniphis hinnus); podonotal region usually with 20-23 pairs of setae (6 j, 6 z, 4, 5 or 6 s, 3, A or 5 r) or less frequently, more than 25 pairs may be present

(Notogamasellus. Euepicrius. Pilellus and Caliphis calvus Lee, 1970); podonotal setae rl always absent; opisthonotum usually with 20-22 pairs of setae (5 J, 5 Z, 5 S, 5-6 R, 0-3 UR), but a reduction in the number of setae is not uncommon; more rarely, hypertrichy (more than

28 pairs of setae) also occur (Caliphis. Euepicrius. Cyrtolaelaps ,

Pilellus and Gamasellus discutatus complex); submarginal (UR) series is commonly absent; when present, there are 1-3 pairs

(Euryparasitidae, Gamasellidae) or more rarely, at least 4 and up to

9 pairs (Eurvparasitus. Pyriphis. Epiphis. and the hypertrichous taxa); some podonotal and opisthonotal setae may be placed on soft cuticle - if only one pair of setae is on soft cuticle on each of these regions then, they usually are setae r4 and R5; dorsal setae of 134 the idiosoma usually simple, although they may be more elaborated as in most Euryparasitidae, Gamasellidae and a few Ologamasidae; podonotum may have two pairs of scleronoduli on the dorsal hexagonal area (most Rhodacaridae and a few Gamasellidae); podonotal glands gd4 usually small, but they may be enlarged in a few genera (for example,

Acugamasus. Allogamasellus. Ologamasus. some species of the

Gamasellus falciger and G^_ tragardhi complexes).

Venter: Preendopodal shields usually present as one pair, but

often there are two or more pairs present (some Rhodacaridae, most

Gamasellidae); less frequently, they may be weakly sclerotized or absent (five genera of Rhodacaridae) and very rarely, they may be

incorporated to the sternal shield (Gamasellopsis. Gamasitus.

Sessiluncus and Euepicrius): female with the sternal and metasternal

shields fused, the metasternal setae (st4) usually placed on a

sterno-metasternal shield, along with three pairs of sternal setae

and three pairs of lyrifissures; in the Epiphidinae, the metasternal

shields are narrowly fused to the sternal shield or, are discrete and

carry the metasternal setae ; in the subgenus Psammonsella and

Litogamasus gressitti Hunter, 1970, the metasternal setae are also

placed on discrete metasternal platelets; in Evanssellus. the

metasternal setae may be placed on the sterno-metasternal shield, on

the soft cuticle or may be absent; the sterno-metasternal shield may

or may not be coalesced with the endopodal shields; the first pair of

sternal setae (stl) are usually placed on the well or weakly defined

anterior margin of the sternal shield (weakly defined in

Afrodacarellus. Rhodacarus and Rhodacarellus). or very rarely, they. 135 may be placed on the preendopodal shields (Afrogamasellus and

Mediodacarellus) ; female genital shield with 1 pair of setae and usually rounded anteriorly or, less frequently, with a pointed anterior flap (Pyriphis. Gamasellevans. Afrodacarellus. Gen. n. #1 and some Rhodacarus and Afrogamasellus species), and almost always truncate posteriorly (the exception being Notogamasellus. with a rounded posterior margin); associated genital pores always on soft cuticle; posterior margin of female genital shield may extend well beyond acetabulum IV (Euryparasitidae, Rhodacaridae and some

Gamasellidae genera) or may not extend much beyond it (most

Ologamasidae and most Gamasellidae genera); accessory opening to the spermathecae usually associated with leg IV (placed on coxa, acetabulum, metapodal or even between the exopodal IV and the posterior end of the peritrematal shield), but association with leg

III also occur (on acetabulum, coxa, trochanter or femur); male always with a sternogenital shield carrying four pairs of sternal setae, three pairs of lyrifissures and usually carrying genital setae

(st5), although a few exceptions to this pattern are observed; the genital setae are placed on separate platelets in Rhodacaropsis and

Afrodacarellus; on shields fused to the endopodals IV in

Mediodacarellus: on the soft cuticle posteriorly to the sternogenital shield in Paragamasellevans and Rhodacaroides; on the ventrianal shield, anteriorly to setae Jvl in Hvdrogamasus: the genus Acugamasus may have setae st5 on separate platelets, on endopodals IV or on the soft cuticle; ventral glands gv2 usually present or rarely, absent

(Evanssellus. N. minvaspis): when present, they always open 136 posteriorly to acetabulum IV - on separate platelets, on exopodals IV or on soft cuticle - in 1-3 solenostoma, although up to 5 solenostoma can be found (Parasitiphis and Heterogamasus); peritrematal shields present and variously formed (wide or narrow throughout its length, narrow anteriorly and widened posteriorly), with variable degree of fusion to other shields (dorsal shield, exopodals II-IV, ventrianal and metapodal shields); except for Nodacaroides minvaspis (which has only an anal shield with moderately enlarged anal valves), there always is a ventrianal shield with small anal valves; the ventrianal shield may or may not be coalesced with the opisthonotal, peritrematals, exopodals IV and metapodal shields; the adanal setae are commonly placed at anterior or midle level to the anal valves, but may be placed at level of posterior margin of the anal valves in some taxa; the postanal seta is usually shorter or sligthly longer than the adanal setae (most Euryparasitidae, Gamasellidae and

Ologamasidae) but in some genera they are at least one and a half times as long as the adanal setae (Rhodacaridae: Litogamasus,

Rhodacaropsis. Rhodacarus. Rhodacarellus; Ologamasidae: Hvdrogamasus and Parasitiphis: Gamasellidae: Rhodacaroides); there usually are between 6-8 pairs of opisthogastric setae (4-5 Jv, 2-3 Zv) although up to 10 pairs (5 Jv, 3 Zv, 2 Lv) or even 14 pairs can be present; if

there are only 6 pairs, setae Jv4 and Zv3 are absent; if 7 pairs are present, Jv4 is absent; when setae are placed on soft cuticle, these

usually are Jv5 and Zv3 (if 7 or 8 pairs) or more rarely, setae Jvl,

Lvl and Lv2; opisthogastric setae Zvl are usually placed at level

below that of setae Jvl, but in a few genera Zvl are removed to a 137 level well above that of setae Jvl, almost vertically to them

(Afrodacarellus, Mediodacarellus « Gen. n. #1, Rvkellus and Epiphis); exopodal shields III usually not completely formed, so that they do not encase peraxial edges of acetabula III (in the Rhodacaridae and

Euryparasitidae) or, they are well formed and entire or narrowly split (in the Gamasellidae and Ologamasidae).

Legs: Legs II-IV always with pretarsi bearing two claws, bilobed pulvilli and often with a pair of pointed membranous processes between the bases of the claws ; leg I usually with a pretarsus which is smaller than the other pretarsi, set on a long peduncle and bearing claws and pulvilli, although often the peduncle is absent; more rarely, the pretarsus is lost (Notogamasellus. Rhodacarus,

Solugamasus. Nodacaroides. Gen. n. #1, Evanssellus and Euepicrius); in Afrodacarellus and Sessiluncus. tarsus I bears a long solenidion

(at least half as long as the length of tarsus) and in males of

Epiphidinae and Afrodacarellus camaxiloensis a large, bulbous sensillum is present; a rare, acrotarsal pseudosegmentation is observed in the tarsus I of the deutonymphs of the Epiphidinae and the genera Cyrtolaelaps and Eurvparasitus (Gamasellidae); leg II of males has the antero-ventral (av) seta almost always modified into a conspicuous thumb-like spur and seta av on genu, tibia and tarsus often modifed into a small spine (Evanssellus is the exception, having seta av on femur produced into a rather inconspicuous spur); the standard chaetotaxy of legs I-IV is generally that of the

Rhodacaridae-type of Evans (1963) and is listed as follows: 138

coxae I-IV: (2-2-2-1); trochantera I-IV: (6-5-5-S); femora I-IV;

(2-S/4-2) (2-5/3-1) (1-3/1-1) (1-3/1-1); genua I-IV; (2-3/2,3/1-2)

(2-3/1,2/1-2) (2-2/1,2/1-1) (2-2/1,3/1-1); tibiae I-IV: (2-3/2,3/2-2)

(2-2/1,2/1-2) (2-1/1,2/1-1) (2-1/1,3/1-2); tarsi II-IV: (3-4/3,3/2-3)

(3-4/3,3/2-3) (3-4/3,3/2-3).

Exceptions to this general pattern occur and they will be listed with the familial diagnoses and descriptions.

Family Euryparasitidae n.fam.

(Gamasellini Hirschmann, 1962 sensu Lee. 1970 in part).

Type Genus: Euryparasitus Gudemans, 1902.

DIAGNOSIS

Tectum three-tined, with central tine always denticulate

laterally; female chelicerae usually with 3-4 teeth on the movable

digit and 5-7 teeth on the fixed digit; spermadactyl freer than fused

to the movable digit, generally straight through most of its variable

length; dorsal shield divided in the deutonymph and in the adult;

dorsal setae usually complex; podonotal setae jl almost always

complex; podonotal region with 22-24 pairs of setae and opisthonotal

region with 20-23 pairs , but an increase in the number of setae on

each of these regions also occurs; submarginal (UR) series of setae

almost always present; ventral sclerotization usually more extensive

in the male; preendopodal shields almost always present; female

sterno-metasternal shield always carrying the metasternal setae (st4)

but never fused to the endopodal shields IV; first sternal setae

(stl) on sternal shield; female genital shield usually subrectangular

in shape, its posterior margin always projecting well beyond 139 acetabula IV; male with a sternogenital shield usually discrete, rarely fused to the ventrianal shield; male genital setae (st5) almost always on the sternogenital shield; ventral glands gv2 almost always present in 2 or 3 solenostoma; ventrianal shield present and always discrete (not fused to other shields) in the female; opisthogastric region usually with 6-8 pairs of setae, although up to

10 pairs may be present; peritrematal shields always fused to the exopodal shields IV and often widening posteriorly; exopodal shields

III not completely encasing peraxial edges of respective acetabula; pretarsus I almost always present; leg chaetotaxy usually as for the superfamily, with few exceptions.

DESCRIPTION

Size variable, ranging from small mites (less than AOOu and up to 600u e.g., Allogamasellus. Notoeamasellus) to medium-sized mites

(601-800u e.g., Acugamasus. Heterogamasus) to large mites (801u and above 1200u e.g., Euryparasitus. Cvrtolaelaps).

Gnathosoma: Palp claw always three-pronged, with basal prong reduced in Heterogamasus ; seta all on palp genu may be simple

(Cvrtolaelaps. Notogamasellus). finely pilose (Euryparasitus.

Heterogamasus) or spatulate (Allogamasellus. Acugamasus); tectum with central tine laterally denticulate, often divided at its tip

(Heterogamasus. Allogamasellus. Acugamasus natalensis species complex

Lee, 1970, Cvrtolaelaps mucronatus G. & R. Canestrini, 1881 and

Notogamasellus (Notogamasellus) vandenbergi Loots & Ryke, 1966); salivary stylets and corniculi well developed; female chelicera with a combination of small and large teeth (Cvrtolaelaps. Euryparasitus. 140

Acugamasus) or with predominantly small teeth (Allogamasellus.

Heterogamasus. Notogamasellus); movable digit of the female chelicera usually with 3-4 teeth, but in Euryparasitus there are 3 large teeth

plus up to 15 very small teeth distributed between the large ones;

fixed digit of the female chelicera with 5-7 teeth; male spermadactyl usually straight throughout its length and freer than fused to the movable digit; exceptionally, Acugamasus has a spermadactyl which curves around the movable digit and is fused to the digit for most of its length; the spermadactyl is shorter than the movable digit in

Acugamasus. Heterogamasus and Notogamasellus or, at least one and a half times as long as the movable digit in Cvrtolaelaps.

Euryparasitus and Allogamasellus; the arthrodial process of the chelicera is produced into a corona except for the genus

Notogamasellus which has an arthrodial brush.

Dorsum: Dorsal shield of deutonymphs and adults diyided; podonotal region usually with 22-23 pairs of setae (6 j, 6 z, 6 s,

4-5 r), but Notogamasellus exhibits hypertrichy on this region, with

28-32 pairs of setae; Cyrtolaelaps possesses a very unusual podonotal chaetotaxy with 23-24 pairs of setae distributed as 7 j, 6 z, 5-6 s,

5 r; podonotal setae rl always absent; opisthonotal region usually with 20-24 pairs of setae (5 J, 5 Z, 5 S, 5-6 R, 0-3 UR) -

Heterogamasus being the only taxon with 24 pairs of setae - but 27

pairs are present in Euryparasitus (5 J, 5 Z, 5 S, 5 R, 7 UR), while

Cyrtolaelaps exhibits opisthonotal hypertrichy (more than 28 pairs); female usually with some podonotal (1-2 pairs) and opisthonotal setae

(at least 3 pairs) placed on soft cuticle; on the podonotum these 141 usually are setae r4 and/or s6; on the opisthonotum they usually

include setae of the R and UR series and only rarely, setae of the S series (Heterogamasus spinosissimus Balogh, 1963 and Hj_ claviger

Tragardh, 1907 have setae SI and 82 on soft cuticle, along with the entire R and UR series); in Cvrtolaelaps there are no opisthonotal setae on soft cuticle; dorsal setae of the idiosoma predominantly complex in Acugamasus. Heterogamasus and Notogamasellus. whereas

Cyrtolaelaps has only a few complex setae distributed throughout the podonotal and opisthonotal regions; Allogamasellus and Euryparasitus have predominantly simple setae (some of them are at most, pilose); podonotal setae jl usually complex (strongly pilose, pilose-spatulate, modified into a strong spine), in Euryparasitus being only slightly pilose; podonotal glands usually small, but in

Acugamasus and Allogamasellus they are conspicuously enlarged.

Venter: Preendopodal shields usually well developed and produced into one pair, although in Allogamasellus there are three or more pairs; in Notogamasellus they are present as a weakly sclerotized pair, whereas in Cyrtolaelaps they are absent; female with a sterno-metasternal shield which always includes three pairs of

sternal setae, three pairs of lyrifissures and the metasternal setae

(st4) and is never fused with the endopodal shields IV; female

genital shield bearing one pair of setae, subrectangular, with

anterior margin usually rounded (except for Notogamasellus and

Heterogamasus spinosissimus which have it truncate) and posterior margin truncate (except for Notogamasellus which has it rounded),

always extending well beyond acetabula IV; genital pores always on 142 soft cuticle; accessory opening to the spermathecae placed on acetabulum IV (Allogamasellus), on posteroventral surface of trochanter III (Acugamasus) or on anteroventral surface of trochanter

III (Heterogamasus) ; in Cvrtolaelaps and Euryparasitus the accessory opening has not been determined with certainty although according to

Lee (1970), it may open on the sterno-metasternal shield, between st3 and acetabulum III; according to Michael (1892), the spermatophore is directly deposited in the vagina of Euryparasitus emarginatus; in

Notogamasellus the opening to the spermathecae is not known; male sternogenital usually discrete, except for Heterogamasus which has a holoventral shield (the sternogenital and ventrianal shields coalesced) which is also fused to the exopodals IV and peritrematal shields; sternogenital almost always bearing five pairs of setae

(including the genital setae) and three pairs of lyrifissures; exceptionally, males of Acugamasus may have st5 placed on separate platelets, on the endopodal shields IV or on the soft cuticle; ventral glands gv2 usually present in 2-3 solenostoma, although up to

5 solenostoma may be present (Heterogamasus); gv2 is apparently absent in Notogamasellus; peritremes often reduced in length (except for Heterogamasus and Acugamasus). reaching up to anterior or middle level of acetabula II; peritrematal shields always present and fused to the exopodal shields, usually well developed (reduced in

Notogamasellus). often fused anteriorly to the podonotal shield and widening posteriorly or, of same width along entire length

(Cyrtolaelaps. Euryparasitus and some species of Acugamasus); in males of Cyrtolaelaps. Euryparasitus and Heterogamasus. the 143 peritrematal shields may be fused to the ventrianal and to the metapodal shields as well; ventrianal shield always dicrete in the female, whereas in the male it is often discrete but it may also be fused with the metapodals, exopodals IV and peritrematal shields

(Euryparasitus. H. claviger), and with the opisthonotal shield posteriorly and laterally (Cvrtolaelaps. Acugamasus natalensis complex, H. euarmatus Karg. 1977) or, the ventrianal may be fused to the metapodals, exopodals IV, peritrematals and sternogenital shields, but not to the opisthonotal shield (H. claviger); anal valves small; postanal seta shorter than the adanal setae, except in

Acugamasus where it is longer than the adanals; adanal setae placed at level of anterior or posterior margin of the anal valves; there usually are 6-8 pairs of opisthogastric setae present (4-5 Jv, 2-3

Zv), but in Euryparasitus and Cyrtolaelaps there are 10 pairs (5 Jv,

3 Zv, 2 Lv); if 6 pairs are present, setae Jv4 and Zv3 are absent; if

7 pairs, setae Jv4 are absent; in the female, some opisthogastric setae may be set on soft cuticle (from 1 pair and up to 6 pairs of setae); setae Jv5, then Jvl and Zvl are the most commonly seen on soft cuticle, followed by setae Zv3, Lvl and Lv2; opisthogastric setae Zvl often placed at level parallel to or slightly above that of setae Jvl; in Allogamasellus and Acugamasus setae Zvl are placed at level below that of setae Jvl; exopodals 11 and 111 not completely formed, so that they do not encase peraxial edges of acetabula 11 and

111.

Legs: Legs 11-lV with pretarsi bearing two claws, bilobed

pulvilli and a pair of pointed membranous processes between the bases. 144 of the claws; leg I usually with a non-pedunculate pretarsus, which bears claws and pulvilli but it is smaller than the other pretarsi; very rarely, the pretarsus I is absent (Notogamasellus); tarsus I of deutonymphs of Cvrtolaelaps and Euryparasitus exhibit a rare, acrotarsal pseudosegmentation; leg II of male with the anteroventral seta (av) of femur always modified into a large thumb-like process, and seta av on genu, tibia and sometimes on tarsus, modified into conspicuous spurs or spines; the chaetotaxy of legs I-IV is usually that of the Rhodacaroidea pattern except that the genera

Cyrtolaelaps. Heterogamasus and Notogamasellus may have an extra posterolateral seta on genu and tibia III.

Family Rhodacaridae Oudemans, 1902

(Rhodacarinae sensu Lee, 1970).

Type Genus: Rhodacarus Oudemans, 1902.

DIAGNOSIS

Tectum three-pronged, with central tine variously produced; deutosternum usually with 7 rows of denticles; female chelicerae usually with 3 teeth on the movable digit and 5-7 teeth on the fixed digit; spermadactyl distally free and basally recurved before projecting anteriad or posteriad, of variable shape and length; dorsal shield divided in the deutonymph and almost always divided in the adult; dorsal setae usually simple; podonotal setae jl almost always simple; dorsal hexagonal area of the podonotum almost always with scleronoduli; podonotal region with 20-23 pairs of setae, opisthonotal region with 18-21 pairs, but further reduction in the chaetotaxy of both regions also occurs; submarginal (UR) series 145 absent; preendopodal shields usually present; female metasternal shield almost always bearing the metasternal setae, and almost never fused to the endopodal shields IV; first sternal setae usually placed on the sternal shield; female genital shield usually longer thah wide, posterior margin truncate and almost always projecting well beyond acetabula IV; accessory opening to the spermathecae almost always associated with proximal segments of leg IV; male sternogenital shield discrete, almost always bearing the genital setae; ventral glands gv2 usually present in 1-2 solenostoma; peritrematal shields almost always fused to the exopodal shields IV; peritremes reduced in length, reaching up to middle level of acetabula II; ventrianal shield almost always present, subtriangular, discrete in the female; anal valves almost always small; opisthogastric region with 6-7 pairs of setae; opisthogastric setae

Jv4 absent; pretarsus I usually present; leg chaetotaxy usually as for the superfamily, with few exceptions including the common absence of seta pl4 on tarsus IV.

DESCRIPTION

Size variable, ranging from small (less than 400u and up to

6Q0u) to large mites (801 - above 1200u).

Gnathosoma: Palp claw three-pronged; seta all on palp genu often simple, although it may also be spatulate (Afroeamasellus.

Latogamasellus). finely pilose (Litogamasus) or, with 4-6 lateral prongs (Solugamasus and some species of Nodacaroides; tectum usually three-tined, but the shape of the central tine varies from smooth on sides and undivided at its tip, to bifurcate, trifurcate, or bearing 146 spinules on its anterior half; the lateral tines may be smooth or denticulate on sides and very rarely, bifurcate at their tips; salivary stylets and corniculi well developed, corniculi usually entire (except for Paragamasellevans vandenbergi which has bifid corniculi); female chelicerae usually with small and large teeth, although the females of some genera possess chelicerae with predominantly large teeth (Rhodacarus. Rhodacaropsis. Litogamasus.

Paragamasellevans and Solugamasus); movable digit of female chelicera usually with 3 teeth, but 4-7 teeth may be present (Gen. n. #1,

Latogamasellus and Nodacaroides): fixed digit of female chelicera usually with 5-7 teeth, although up to 12 teeth are present in some species of Rhodacarus: spermadactyl always distally free and basally recurved (”S"-like base), its shape varying as follows: generally straight through most of its length (Afrogamasellus Afrodacarellus and Litogamasus): curving around the movable digit (Litogamasus setosus Kramer, 1898); recurved at basal, median or distal level of the movable digit before projecting freely anteriad (Mediodacarellus.

Latogamasellus and Paragamasellevans)) or posteriad (Rhodacarus.

Rhodacarellus Rhodacaropsis and Solugamasus); regarding length, the spermadactyl may be subequal in length to the movable digit of the chelicerae (Afrogamasellus. Afrodacarellus. Mediodacarellus.

Latogamasellus. Litogamasus and Rhodacarus). or less than one and a half times as long as the movable digit (Rhodacaropsis). or at least one and a half times as long as the movable digit (Rhodacarellus,

Paragamasellevans. Solugamasus and Litogamasus gressitti Hunter,

1970); the spermadactyl is fused to the movable digit through most of 147 its length in Afrogamasellus. Afrodacarellus. Mediodacarellus and

Latogamasellus. whereas in the remaining genera (for which the male is known) it is freer than fused to the movable digit; the arthrodial process of the chelicera is usually produced into a corona, but it may also be produced into a brush (Afrogamasellus. Afrodacarellus.

Mediodacarellus and Latogamasellus).

Dorsum: Dorsal shield always divided in the deutonymph; dorsal shield of the adult is almost always divided, except for

Afrogamasellus luberoensis luberoensis Loots, 1968 and ^ 1. kalibuensis Loots, 1968, which have extensively coalesced dorsal shields with a distinct line of demarcation between the podonotal and opisthonotal regions; podonotal region usually with 20-23 pairs of setae (6 j, 6 z, 4-6 s, 3-5 r) and although rarely, a reduction in the number of setae may also occur (some species of Nodacaroides); podonotal setae rl always absent; opisthonotal region often with

18-21 pairs of setae (5 J, 5 Z, 4-5 S, 4-6 R), but a reduced chaetotaxy is observed in several species of Nodacaroides (12-17 pairs) and in Solugamasus (which has only 11 pairs of setae); submarginal (UR) series absent; in the female, some podonotal and opisthonotal setae are commonly placed on soft cuticle and the setae involved usually belong to the r series (r2, r4 and r5) and to the R

series (R1-R5); exceptionally, Nodacaroides minvaspis also has 1 pair

of the s series (s6), 1 pair of the J series (J5), 1 pair of the Z

series (Z5) and all of its S series (4 pairs) and R series (Rl)

placed on the soft cuticle; opisthonotal setae J5 usually placed at

level anterior to that of setae Z5, but in Litogamasus. Rhodacarus. 148

Rhodacarellus and Nodacaroides. setae J5 are placed at a parallel or posterior level to that of setae Z5; dorsal setae of the idiosoma usually simple, except for Litogamasus Lee, 1970 (which has some of

its setae pilose) and Latogamasellus bipilosus (Karg, 1979), with most of its setae leaf-like and inconspicuously pilose; almost always, there are 2 pairs of scleronoduli set on the dorsal hexagonal area (dha) of the podonotal shield; in Rhodacarus and Rhodacaropsis.

the median pair is coalesced so that only three scleronoduli are present; in N. minvaspis. the outer pair is lost, leaving only two

scleronoduli on the dha; Latogamasellus has secondarily lost its

scleronoduli; the anterior half of the podonotal shield may be incised in "V" shape (Rhodacarus and Nodacaroides coniunctus) or in a semi-concave manner (Rhodacaropsis. Rhodacarellus minimus Karg,

1961); podonotal glands gd4 never enlarged.

Venter: Preendopodal shields often present in one well

sclerotized (Afrogamasellus. Paragamasellevans and Nodacaroides) or weakly sclerotized pair (Mediodacarellus and Afrogamasellus mongii

Hurlbutt, 1973), in two pairs (Rhodacaropsis. Solugamasus and Gen. n.

#1) or, in more than two pairs (Litogamasus); less frequently, preendopodals are absent (probably, secondarily lost) in Rhodacarus,

Rhodacarellus. Afrodacarellus and Latogamasellus; female always with

a sterno-metasternal shield (it may be weakly defined at its anterior and/or posterior margins) which usually bears the metasternal setae

(st4); in Litogamasus. setae st4 may be placed on the soft cuticle on

one side; sterno-metasternal shield usually not coalesced with the

endopodal shields IV, except for some species of Afrogamasellus; the 149 first pair of sternal setae (stl) are usually placed on the anterior margin of the sternal shield regardless of whether the shield is weakly sclerotized (Rhodacarus. Rhodacarellus and Afrodacarellus) or well sclerotized (Litogamasus. Rhodacaropsis. Paragamasellevans.

Solugamasus. Gen, n. #1 and Nodacaroides); more rarely, setae stl are placed on the preendopodal shields (Afrogamasellus and

Mediodacarellus). or on an area of punctate sclerotization between the sternal and the preendopodal shields (Rhodacarus berrisfordi

Loots, 1969); female genital shield usually longer than wide, with the anterior margin pointed or rounded and the posterior margin usually truncate, extending well beyond acetabula IV; genital setae on shield, genital pores off it; accessory opening to the spermathecae almost always associated with leg IV (acetabulum, coxa) but association with leg III also occurs (in Latogamasellus. the spermathecae open on the ventral surface of the coxae, while in

Rhodacarus they open on the basifemur); in some species of

Afrogamasellus. the spermathecae open on the metapodal shields; male with a sternogenital shield which almost always bears the genital setae (st5); in Rhodacaropsis and Afrodacarellus the genital setae are placed on discrete platelets, posteriorly to the sternogenital shield, whereas in Mediodacarellus. they are placed on the endopodal shields IV; in Paragamasellevans. they are placed on the soft cuticle; ventral glands gv2 usually present in 1-2 solenostoma, but there are 3 solenostoma in Solugamasus and Nodacaroides costai. whereas in Nodacaroides minvaspis they are absent; peritrematal shields always present, but not always fused to the exopodal shields 150

IV; although they are often reduced, in some species of

Afrogamasellus they are enlarged and fused to the metapodal shields;

peritremes usually reduced in length, reaching up to middle level of acetabula II; in Rhodacaropsis attenuatus Loots. 1969 they are so

reduced that only the stigmata are present; a ventrianal shield is almost always present (Nj^ minvaspis is the exception, with only an

anal shield), usually subtriangular (less often, subrectangular),

discrete in the female but often fused to the opisthonotal shield

(and sometimes to the peritrematal shields) in the male; anal valves

almost always small (N. minvaspis has moderately enlarged anal

valves); postanal seta usually longer than the adanal setae;

opisthogastric region with 6-7 pairs of setae (4 Jv, 2-3 Zv); Jv4

always absent; in the female, setae Jv5, Zvl and Zv3 are often placed

on the soft cuticle; in minvaspis. all 6 pairs of opisthogastric

setae (setae Zv3 are absent) are placed on soft cuticle;

opisthogastric setae Zvl are often placed at a level parallel to, or

slightly above that of setae Jvl (Litogamasus. Rhodacarellus.

Solugamasus. Nodacaroides. Afrogamasellus and Latogamasellus). or

they may be placed well above setae Jvl, almost vertically to them

(Afrodacarellus. Mediodacarellus. Gen. n. #1 and Afrogamasellus

mongii); more rarely, setae Zvl are placed at level below that of

setae Jvl (Rhodacarus. Rhodacaropsis and Paragamasellevans); exopodal

shields III almost never encase peraxial edges of acetabula III,

except for Mediodacarellus and some species of Afrogamasellus which,

although having their exopodals II-IV split, they all encase the

peraxial edges of their respective acetabula. 151

Legs: Legs II-IV always provided with pretarsal claws and

pulvilli; leg I usually with a pretarsus which is smaller than the

other pretarsi and may be set on a peduncle C Rhodacarellus.

Afrodacarellus. Mediodacarellus and Latogamasellus), or the pretarsus

is present but not pedunculate (Litogamasus. Rhodacaropsis.

Paragamasellevans and Afrogamasellus) or, the pretarsus is lost

(Rhodacarus. Solugamasus. Nodacaroides and Gen. n. #1); the tarsus I

is provided with solenidia, but Afrodacarellus possesses an

additional, long solenidion (at least half as long as the tarsus)

arising at median length of the segment; the leg II of the male has

the anteroventral (av) seta modified into a conspicuous thumb-like

spur, and the av seta on the genu and tibia is usually modified into

a spine; the pattern of leg chaetotaxy closely follows that of the

superfamily, with the few exceptions thus listed: on tarsus IV, seta

pi4 is absent in Rhodacarus. Rhodacarellus. Rhodacaropsis.

Afrogamasellus. Mediodacarellus. Latogamasellus and most species of

Afrodacarellus (the exception being A. camaxiloensis). Also, both

species of Rhodacaropsis lack one dorsal seta on femur II (2-4/3-1)

and Rhodacaropsis attenuatus Loots, 1969 lacks two dorsal setae on

femur III (1-2/1-0), whereas Rhodacaropsis inexpectatus Willmann,

•1935 lacks only one dorsal seta on femur III (1-3/1-0).

Diagnoses for the newly established genus (Mediodacarellus) and

for Nodacaroides Karg, 1977 and Latogamasellus Karg, 1977 new status

are given below. 152

Genus Mediodacarellus new genus

Type species: Afrogamasellus bakeri Hurlbutt. 1973

DIAGNOSIS

Tectum uniramous, laterally denticulate, with a divided or undivided tip; seta all on palp genu simple; female chelicera with 2 teeth on the movable digit and 4 teeth on the fixed digit; spermadactyl fused to the movable digit for most of its length, recurved at median level of the digit before projecting anteriad, shorter than the digit; arthrodial process of the chelicera produced into a brush; dorsal shield divided; dorsal setae simple; dorsal hexagonal area with two pairs of scleronoduli; podonotal region with

23 pairs of setae, setae r4 placed on soft cuticle; opisthonotal region with 20 pairs, all setae on the shield; preendopodal shields present as one weakly sclerotized pair defined by punctate sclerotization, and bearing the first pair of sternal setae; female sterno-metasternal shield not coalesced with the endopodal shields

IV, bearing the metasternal setae; female genital shield longer than wide, roughly bell-shaped rounded anteriorly and truncate posteriorly, projecting well beyond acetabula IV; spermathecae

opening on the posterior margin of acetabula IV; male sternogenital

shield discrete, bearing 4 pairs of sternal setae and 3 pairs of

lyrifissures; male genital setae (st5) placed on endopodal shields

IV, closely abutting the sternogenital shield; peritrematal shields

fused to the exopodal shields IV; exopodal shields III split, but

encasing peraxial edges of respective acetabula; peritremes reduced,

reaching only up to the posterior level of acetabula II; ventrianal 153 and metapodal shields discrete in both sexes; anal valves small; opisthogastric region with 7 pairs of setae, setae Zv3 placed on soft cuticle in the female; setae Zvl placed well above setae Jvl, almost vertically to them; pretarsus I present, set on short peduncle; seta pv2 on tarsus IV bifurcate; leg chaetotaxy as for the superfamily, except that seta pl4 is absent from tarsus IV,

Genus Nodacaroides Karg, 1977 new status

(= Rhodacaroides (Tenacaroides) Karg, 1977 new synonymy)

Type species: Rhodacaroides (Nodacaroides) coniunctus

Karg, 1977

DIAGNOSIS

Seta all on palp genu with 3-6 lateral prongs; tectum three-tined, with central tine laterally denticulate and undivided at its tip; female chelicera with 3-7 teeth on the movable digit and 4-9 teeth on the fixed digit; arthrodial process of the chelicera produced into a corona; dorsal shield always divided; scleronoduli may be absent from the dorsal hexagonal area of the podonotal shield; podonotal region with 20-23 pairs of setae and opisthonotal region with 18-21 pairs, but a reduction in the chatotaxy of both regions often occurs; opisthogastric region always with 6 pairs of setae, with setae Zvl usually set at a level parallel to that of setae Jvl; preendopodal shields present in one pair; female sterno-metasternal shield bearing the metasternal setae; female genital shield subrectangular, with anterior margin rounded and posterior margin

truncate, projecting well beyond acetabula IV; ventrianal shield nearly always present, subtriangular, discrete; anal valves nearly 154 always small; peritrematal shields present, peritremes always reduced in length, reaching up to middle level of acetabula II; pretarsus I usually absent; pattern of leg chaetotaxy as for the superfamily, including the presence of seta pl4 on basitarsus IV.

Genus Latogamasellus Karg, 1977, new status

Type species: Afrogamasellus (Latogamasellus) squamosus

Karg, 1977

DIAGNOSIS

Tectum usually three-tined, central tine smooth on sides, divided at tip, lateral tines denticulate; seta all on palp genu usually slightly spatulate; female chelicerae with usually 5 teeth on the movable digit, but only 2-3 may be present (in the African species); fixed digit with 5 teeth; arthrodial process of the chelicerae produced into a brush; spermadactyl fused to the movable digit through most of its length, basally recurved before projecting anteriad, shorter than the movable digit; dorsal shield divided; dorsal setae almost always simple; scleronoduli usually absent from the dorsal hexagonal area of the podonotum; podonotal region with

22-23 pairs of setae, opisthonotal region with 20 pairs of setae; almost always, all podonotal setae on the shield; all opisthonotal setae on the shield; preendopodal shields absent; female sterno-metasternal shield weakly defined anteriorly (with area of punctate sclerotization), not coalesced with the endopodal shields

IV, always bearing the metasternal setae; first sternal setae placed on sternal shield, immediately posterior to the area of punctate sclerotization; female genital shield longer than wide, usually 155 subtriangular, rounded anteriorly, truncate posteriorly, projecting well beyond acetabula IV; accessory opening to the spermathecae placed on the ventral surface of coxa III; ventral glands gv2 present in 1 or 2 solenostoma; peritrematal shields almost always fused to the exopodal shields IV; peritremes reaching up to middle level of acetabula II; exopodal shields III not encasing peraxial edges of respective acetabula; ventrianal shield usually subtriangular, discrete; anal valves small; opisthogastric region with 7 pairs of setae (setae Jv4 absent); setae Jv5 and Zv3 consistently placed on soft cuticle; pretarsus I present, pedunculate; seta pl4 absent from tarsus IV.

Family Gamasellidae Hirschmann, 1962 new status

(Gamasellini Hirschmann, 1962 sensu Lee, 1970 in part;

Ologamasini Lee, 1970 in part).

Type Genus: Gamasellus Berlese, 1892

DIAGNOSIS

Seta all on palp genu usually with lateral prongs; tectum usually three-tined, with central tine undivided, smooth or denticulate on sides; female chelicerae usually with 3-4 teeth on the movable digit and 5-8 teeth on the fixed digit, but an increase in number of teeth also occurs; spermadactyl may be generally straight

through most of its length or curved, almost always projecting anteriad; arthrodial process of the chelicerae produced into a corona; idiosomal shields usually well sclerotized and of similar extension in both sexes; dorsal shield always divided in the deutonymph and usually divided in the adult; shape of the dorsal. 156 setae usually in a combination of simple and complex setae; podonotal setae jl often complex, sometimes arising from tubercles; podonotal region with 21-23 pairs of setae, opisthonotal region with 20-22 pairs, but hypertrichy of each of these regions also occurs; submarginal (UR) series often present, usually in 2 or 3 pairs but as many as 9 pairs may be present; except for Gamasellus. all podonotal setae are placed on the podonotal shield; preendopodal shields almost always present in at least one pair; metasternal setae (st4) almost always on the sterno-metasternal shield, which may be coalesced with the endopodal shields IV; first sternal setae on sternal shield; female genital shield usually as wide as long, anterior margin almost always rounded, posterior margin truncate and usually projecting only as far as the posterior margin of acetabula IV; the opening to the spermathecae is almost always associated with the proximal segment of leg IV; male sternogenital shield almost always bearing the genital

setae and often fused to the ventrianal shield; ventral glands gv2 almost always present, opening in 2-3 solenostoma; ventrianal shield may be discrete or fused posteriorly and laterally to the dorsal

shield; opisthogastric region usually with 6-8 pairs of setae,

although up to 14 pairs may be present in the hypertrichous taxa;

opisthogastric setae Zvl usually placed at level below that of setae

Jvl; peritrematal shields present, fused anteriorly to the podonotal

shield and posteriorly to the exopodal shields IV (more rarely, to

the ventrianal shield as well), commonly widening posteriorly;

peritremes often reaching full length; exopodal shields III usually

well developed, but split (not entirely coalesced); pretarsus I 157 almost always present; leg chaetotaxy usually as for the superfamily, with the few exceptions occurring in Periseius and Euepicrius.

DESCRIPTION

Size range includes small mites with less than 400u (Hiniphis.

Periseius). medium-sized mites with 601-800u (Pvrvphis) and large mites with 801-1200u (some species of Hevdeniella).

Gnathosoma: Palp claw three-pronged, the basal prong being reduced in Evanssellus ; seta all on palp genu is usually produced with lateral prongs, being at least bifurcate; it may also be trifurcate, or it may have four or more lateral prongs (Hevdeniella.

Evanssellus. H. bipala. G. pvriformis and falciger complexes); exceptionally, all is simple (at most lanceolate) in Euepicrius: tectum usually three-tined (uniramous in Pvriphis). with central tine often denticulate on its sides (or just at its base), undivided at tip; salivary stylets and corniculi usually well developed, except for Onchogamasus communis which has reduced corniculi; female chelicerae with predominantly small teeth, although in some taxa, there is a combination of small and large teeth, while other taxa have chelicerae with predominantly large teeth (Euepicrius. 0. communis and Gamasitus obscurus); movable digit of the female chelicera usually with 3-4 teeth, although 5 teeth (H. bipala). 7 teeth (Periseius (Periseius) braziliensis Hirschmann, 1966), 11 teeth

(Gamasellus concinnus Womersley, 1942) or even up to 19 teeth

(Pvriphis) may be present; on the other hand, the female of

Evanssellus foliatus Ryke, 1961 has only 2 teeth on this digit; the fixed digit of the female chelicera commonly has 5-8 teeth, but 11, 158

17 or as many as 19 teeth may be present as well (P. (P.) braziliensis. G. concinnus and Pvrvphis. respectively); spermadactyl is commonly straight throughout its length, but it may also be curved

(as in some species of Gamaselliphis) or sinuous (some species of the

G. discutatus complex), almost always projecting anteriad; Pvriphis

is the exception, with a strongly recurved spermadactyl which projects posteriorly; the length of the spermadactyl varies from shorter than the movable digit, to at least one and a half times as long as the movable digit (as in Euepicrius and Gamaselliphis); spermadactyl usually freer than fused to the movable digit, except for (Hevdeniella. Hiniphis. and the G^_ tragardhi complex); the arthrodial process of the chelicerae is produced into a corona.

Dorsum: The dorsal shield is divided in the deutonymph and usually divided in the adult, the few exceptions including

Hevdeniella. Pvriphis and males of the Gj_ discutatus complex (with an entirely coalesced shield), and Hiniphis hinnus (with extensively fused shields separated only by grooves of demarcation between the podonotal and opisthonotal regions); dorsal setae may be all simple, although most often seen is a combination of simple and complex setae; in Evanssellus. Euepicrius. Gamaselliphis. Laelogamasus and several species of Gamasellus the dorsal setae are predominantly complex; podonotal setae jl are commonly complex, sometimes arising from strong tubercles (as in Evanssellus and Euepicrius): in

Pvriphis. they are produced into wide, leaf-shaped processes; in

Hevdeniella. Onchogamasus. Hiniphis and Rhodacaroides they are simple; podonotal region usually with 21-23 pairs of setae (6 j, 6 z. 159

5-6 s, 4-5 r), with 22 pairs most often observed; Pilellus and

Euepicrius exhibit hypertrichy on this region (more than 25 pairs of setae); opisthonotal region is subject to more variation, but there commonly are 20-22 pairs of setae (5 J, 5 Z, 5S, 5-6 R, 1-2 UR);

Pvriphis has the holotrichous set (28 pairs), whereas Pilellus,

Euepicrius and most species of the Gj_ discutatus complex exhibit hypertrichy (with 29 or more pairs); submarginal (UR) series is often present, its numbers varying from 1 to 9 pairs, although it is common to find 2-3 pairs; this series is absent in Hevdeniella. Hiniphis,

Periseius. Rhodacaroides and Evanssellus; all podonotal setae are placed on the shield, except for the species of Gamasellus which usually have setae r4 set on soft cuticle; regarding the opisthonotal setae, Gamasellus species have between 1-6 pairs on soft cuticle

(these always involving setae of the R and UR series); Rhodacaroides has setae Rl, H. bipala has setae S5, and Pilellus rykei Hunter, 1967 has all of its extra setae (7 pairs) on soft cuticle; podonotal glands gd4 usually not enlarged, except for Rhodacaroides,

Evanssellus and the G_j_ tragardhi and ^ falciger complexes which have glands gd4 enlarged; in several species of Gamasellus and in Hiniphis hinnus the dorsal hexagonal area (dha) of the podonotum is provided with four scleronoduli.

Venter: Ventral sclerotization of similar extension in both sexes; preendopodal shields almost always present, produced into 1 pair (Hevdeniella. Pvriphis. Pilellus. Evanssellus and

Gamaselliphis). 2 pairs (Hiniphis. Rhodacaroides) or 3 or more pairs

(Periseius. Laelogamasus and Gamasellus); exceptionally, Euepicrius 160 has Its pair incorporated to the sternal shield; female sterno-metasternal shield usually discrete, although it may be fused or partly fused to the endopodal shields IV (Hevdeniella. Hiniphis.

P. (Periseius). G. discutatus complex, Euepicrius and Gamaselliphis). an usually bearing the metasternal setae (stA); in Periseius

(Psammonsella). stA are placed on discrete metasternal plates; in

Evanssellus. setae stA may be placed on the sterno-metasternal shield, on the soft cuticle, or rarely, they may be absent; first sternal setae always on the sternal shield; female genital shield usually as wide as long (or wider than long), although it may also be sligthly longer than wide (Pvriphis. Rhodacaroides. Pilellus. G. tragardhi complex, and H. hinnus): exceptionally, in Evanssellus and

P. (Psammonsella) the genital shield is at least one and a half times as long as wide; the anterior margin of the genital shield is usually rounded (except for the pointed anterior flap of Pvriphis). its posterior margin is truncate and often extends only as far as the posterior margin of acetabula IV; in a few taxa however, it may extend well beyond these acetabula (Rhodacaroides. Pilellus,

Evanssellus foliatus. G. pvriformis and Gç^ tragardhi complexes); the spermathecae almost always open on the edge of acetabula IV; in

Pvriphis and Hevdeniella they open on the dorsal surface of trochanter III, whereas in H. hinnus thev open on the posterior margin of coxa IV; male sternogenital shield is commonly discrete, or it may be fused to the ventrianal shield - forming a holoventral shield - in Pvriphis. Hiniphis. Periseius. Pilellus. Evanssellus foliatus. Hevdeniella sherrae Lee and Hunter, 197A, Gamasellus 161 falciger and Gamasellus nepotulus Berlese. 1908; male genital setae

(st5) usually on the sternogenital shield, except for Rhodacaroides which has st5 on soft cuticle; ventral glands gv2 usually present

(they are absent in Evanssellus) in 2-3 solenostoma; peritrematal shields always fused anteriorly (sometimes laterally as well) to the podonotal shield and posteriorly to the exopodal IV shields, often widening posterioly; peritremes usually reaching full length,

extendind to the anterior margin of the podonotal shield, next to setae zl; more rarely, they may be reduced (in two species of

Hevdeniella. H. bipala and Gamasellus cophinus Lee, 1973); ventrianal

shield may be discrete or fused posteriorly and laterally to the dorsal shield (in both sexes), or less frequently, also fused to the

peritrematal shields in the male and sometimes in the female as well; anal valves small; postanal seta usually shorter than the adanal

setae, except for Rhodacaroides. P. (Periseius) and the G. pvriformis

complex with a longer postanal seta; opisthogastric region usually

with 6-8 pairs of setae (4-5 Jv, 2-3 Zv), although 10 pairs (5 Jv, 3

Zv, 2 Lv) are present in Euepicrius and Pilellus rvkei. and up to 14

pairs in species of the G. discutatus complex; often, setae Jv5 and

Zv3 are placed on soft cuticle in species of Gamasellus;

opisthogastric setae Zvl are usually placed at level below that of

setae Jvl (in Evanssellus and Pilellus setae Zvl are placed at a

parallel or sligthly above level relative to setae Jvl; exopodal

shields III usually well developed but not entire (split); in

Evanssellus they do not encase acetabula, whereas in H. bipala and

Euepicrius they are entire. 162

Legs: Legs II-IV with pretarsal claws, pulvilli and the pointed

processes arising from the bases of the claws; leg I usually with a

pretarsus, set on peduncle; less frequently, the pretarsus is not

pedunculate and rarely, is lost (Evanssellus and Euepicrius); leg II

of male with anteroventral (av) seta usually modified into a

conspicuous thumb-like spur; less often, av seta on genu and tibia

also modified, produced into small spine; the spur of femur II is

rather inconspicuous in Evanssellus: from the basic Rhodacaroidea

pattern of leg chaetotaxy there are a few exceptions: in the subgenus

Periseius. there may be one extra posterolateral seta on genu III

(2-2/1,2/1-2) and tibia III (2-1/1,2/1-2); in Euepicrius there is one

less ventral seta on genu IV.

Family Ologamasidae Ryke, 1962

(Ologamasinae Ryke, 1962 sensu Lee, 1970 in part; Gamasiphinae

Lee, 1970 in part; Sessiluncinae Lee, 1970 in part).

Type Genus: Oloeamasus Berlese, 1888

DIAGNOSIS

Seta all on palp genu usually with lateral prongs; tectum usually

three-tined, central tine smooth or denticulate on sides, usually undivided at tip; female chelicerae usually with 3-4 teeth on the movable digit and 4-7 teeth on the fixed digit; spermadactyl freer than fused to the movable digit, commonly with "S"-shape base, almost

always projecting anteriad; arthrodial process of the chelicera almost always produced into a corona; dorsal shield almost always

divided in the deutonymph and usually entire in both sexes; podonotal region usually with 20-22 pairs of setae and opisthonotal region with 163

19-21 pairs, but variation in the chaetotaxy of both regions also

occurs; submarginal (UR) series is usually absent, occurring in only

three taxa; idiosomal setae usually simple; preendopodal shields almost always present, in at least one pair; metasternal setae usually placed on the sterno-metasternal shield which is usually

separate from the endopodal shields IV; first sternal setae on the

sternal shield; female genital shield usually as wide as long,

rounded anteriorly, truncate posteiorly and projecting only as far as

the posterior margin of acetabula IV, although a few exceptions to

this pattern occur; male sternogenital shield almost always discrete,

but always bearing the genital setae; ventral glands gv2 always

present, opening in 1-3 solenostoma; ventrianal shield usually discrete in the female and fused to other shields in the male; anal

valves small; peritrematal shields fused anteriorly to the dorsal

shield and often, to the exopodal shields IV posteriorly; peritremes

usually partly reduced in length, reaching anterior margin of

acetabula II; opisthogastric region usually with 6-8 pairs of setae,

although up to 13 pairs may be present; setae Zvl usually placed at

level below that of setae Jvl; exopodal shields III almost always

completely encasing peraxial edges of respective acetabula; pretarsus

I always present, smaller than other pretarsi; leg chaetotaxy usually

as for the superfamily, but several exceptions occur in a number of

taxa.

DESCRIPTION

Well sclerotized mites, mostly medium-sized (6Gl-800u), although

the size range varies from small (less than 400u and up to 600u) to 164 large (801-1100u).

Gnathosoma: Palp claw three-pronged; seta all on palp genu variously produced, although most often it has at least 4 (and up to

15) lateral prongs (nine genera); it may also be finely pilose

(Epiphis and Sessiluncus), bifurcate (Cymiphis. Caliphis and

Hydrogamasus). spatulate (Rykellus. Gamasiphoides and Parasitiphis) or rarely, simple (spine-like as in Gamasiphis and Laelaptiella); this character is not known for Gamasellevans: tectum usually three-tined (uniramous in Rykellus). central tine smooth or denticulate on sides and undivided at tip, although a divided tip is present in some taxa (Ologamasus. Geogamasus. Neogamasellevans.

Queenslandolaelaps. Gamasellopsis and some species of Gamasiphis); salivary stylets and corniculi well developed, corniculi usually entire, except for the bifurcate corniculi of Geogamasus furcatius; female chelicerae may have predominantly small teeth (Cymiphis.

Epiphis). or predominantly large teeth (most Ologamasinae and

Sessiluncus) or, a combination of small and large teeth (most

Gamasiphinae, Hydrogamasellus and Ologamasus: movable digit of the female chelicera with 3-4 teeth (except for Parasitiphis. which has 8 teeth) and fixed digit with 4-7 teeth (Parasitiphis may have up to 12 teeth on this digit); spermadactyl commonly curved at its base

("S"-shape base), but generally straight throughout most of its length and projecting anteriad; exceptions to this pattern include

Sessiluncus. Hydrogamasus and Geogamasus. where the spermadactyl

recurves at the median or distal level of the movable digit before

projecting posteriad or anteriad; the spermadactyl is usually freer 165 than fused to the digit (except for Gamasiphoides) and often, of subequal length relative to the digit; however, in some taxa it is at least one and a half times as long as the movable digit (Epiphidinae,

Geogamasus. Queenslandolaelaps. Gamasellopsis. Sessiluncus and

Gamasiphis); the arthrodial process of the chelicerae is usually produced into a corona, but in the male of Gamasellevans it is modified into a long, leaf-shaped flap and in the male of

Queenslandolaelaps there is an extra, fimbriated flap between the

spermadactyl and the corona.

Dorsum: Dorsal shield almost always divided in the deutonymph

(entire in the Epiphidinae) and usually entire in both sexes (in

Geogamasus it is usually divided in both sexes, but the female may

have it entire); in some males of Athiasella and Hydrogamasellus the

dorsal shield is divided; in Rykellus and Gamasiphoides. the

holonotal shield has a groove or line of demarcation between the

podonotal and opisthonotal regions; podonotal region usually with

20-22 pairs of setae (6 j, 5-6 z, 4-6 s, 4-5 r), but only 19 (some

species of Gamasellopsis) or up to 23 pairs (Gamasiphoides) may be

present; opisthonotal region usually with 19-21 pairs of setae (5 J,

5 Z, 5 S, Q-6 R, 0-1 UR), although hypertrichy (29 or more pairs)

ooccurs in Caliphis. hypotrichy (as few as 15 pairs) ooccurs in some

species of Antennolaelaps. and even holotrichy (28 pairs distributed

as 5 J, 5 Z, 5 S, 7 R, 6 UR) ooccurs in Epiphis; submarginal (UR)

series usually absent; it is only present in Parasitiphis (1 pair),

Epiphis (6 pairs) and the hypertrichous Caliphis (at least 6 pairs);

although not commonly observed in this family, some podonotal and 166 opisthonotal setae may be placed on the soft cuticle, and they usually include setae of the r series (r4 or r5 or r6), and of the R series (R5 and/or R4); exceptionally, Parasitiphis has its only pair of UR setae placed on soft cuticle; dorsal setae usually simple, although in Ologamasus. Queenslandolaelaps and Antennolaelaps a few setae are complex among the simple setae, and in Cymiphis the dorsal setae are predominantly complex; podonotal setae jl usually simple, but in Ologamasus, Cymiphis and Gamasellopsis they tend to be complex; podonotal glands gd4 usually small, but in Ologamasus.

Cymiphis. Gamasiphoides and Laelaptiella they are conspicuously enlarged.

Vente'r: Ventral sclerotization usually more extensive in the male; preendopodal shields almost always present, in at least 1 pair

(2 pairs in Gamasiphoides. Laelaptiella and Hydrogamasus): in

Gamasitus. Gamasellopsis and Sessiluncus. the preendopodal shields have become incorporated to the sternal shield; female sterno-metasternal shield may be separate from the endopodal shields

IV (eleven genera), or it may be partly (Gamasiphoides) or entirely coalesced with them (Cymiphis. Queenslandolaelaps. Antennolaelaps.

Gamasitus. Caliphis. Sessiluncus. Gamasiphis and Gamasellevans), and almost always bears the metasternal setae (st4); in the Epiphidinae setae st4 are placed in discrete metasternal platelets which may be narrowly fused to the sterno-metasternal shield; first sternal setae

(stl) always placed on the sternal shield; female genital shield usually as wide as long (or wider than long), rounded anteriorly, truncate posteriorly and projecting only as far as the posterior 167 margin of the acetabula IV; exceptions to this pattern include:

Athiasella and Caliphis with a shield longer than wide; Parasitiphis and Gamasellevans with a shield at least one and a half times as long as wide; Gamasellevans. with a sharply pointed anterior margin; two species of Hvdroeamasellus. with a rounded posterior margin;

Parasitiphis. Laelaptiella and some species of Hydrogamasellus with genital shield projecting well beyond acetabula IV; accessory opening to the spermathecae usually associated with leg IV, placed on the acetabulum (most Gamasiphinae) or on the coxa (most Ologamasinae); however, association with leg III also occur - on the acetabulum or on the coxa (Hydrogamasellus. Epiphis and Sessiluncus). or on the trochanters (Athiasella and Geogamasus); exceptionally,

Neogamasellevans macrochela Karg, 1975 has the opening to the spermathecae placed on the endopodal shields IV, this being unique in the superfamily; male sternogenital shield almost always bearing the genital setae (st5), except in Hydrogamasus where setae st5 are placed on the ventrianal shield, anteriorly to setae Jvl; sternogenital shield discrete, except in Rykellus where it is produced into a holoventral shield; ventral glands gv2 always present, usually in 1-3 solenostoma, although up to 5 solenostoma may be present in Parasitiphis: ventrianal shield may be discrete in the female and fused posteriorly to the dorsal shield in the male (most

Ologamasinae) or, it may be fused to the dorsal shield in both sexes, and often also fused to the peritrematal shields in the male (and sometimes in the female) - Epiphidinae, most Gamasiphinae and several

Ologamasinae; anal valves small; postanal seta usually shorter than 168 the adanal setae, except for Gamasellopsis. Epiphis. Hydrogamasus and

Parasitiphis; peritrematal shields usually well developed and fused anteriorly to the dorsal shield; posteriorly, they are often fused to the exopodal shields IV (not fused in Gamasitus. Antennolaelaps.

Epiphis. Sessiluncus. Gamasellevans and most species of

Neogamasellevans); peritremes usually reduced in length (except for 4 genera), reaching up to anterior margin of acetabula II; in the

Epiphidinae, Geogamasus and Neogamasellevans. they reach only up to middle level of acetabula II; opisthogastric region with 6-8 pairs of setae (most often 7 pairs) distributed as 3-5 Jv, 3 Zv; exceptionally, Epiphis has up to 13 pairs of setae; although not very commonly observed, setae Jv5 and/or Zv3 may be placed on soft cuticle; setae Zvl usually placed at level below that of setae Jvl; in Rykellus and Epiphis. setae Zvl are set well above setae Jvl, almost vertically to them; in Cymiphis. setae Zvl are placed at a parallel or slightly above level to that of setae Jvl; exopodal shields III may be split or entire, but almost always encase the peraxial edges of respective acetabula, the exception being

Parasitiphis with incompletely developed exopodal shields.

Legs: Legs II-IV provided with preclaws, pulvilli and a pair of pointed membranous processes arising from the bases of the claws; pretarsus I always present, smaller than the other pretarsi and usually set on a peduncle, except for the non-pedunculate pretarsus of Sessiluncus. Parasitiphis and Gamasellevans: leg II of male with anteroventral (av) seta on femur always modified into a conspicuous thumb-like spur and av seta on genu and tibia usually modified into 169 spine-like processes; from the Rhodacaroidea pattern of leg chaetotaxy there are several exceptions listed as follows; one ventral seta is absent from genu I in Gamasellopsis. and from genua

III and IV in Gamasellopsis. Gamasitus. Sessiluncus. Hydrogamasus. some species of Gamasiphis and some species of Gamasiphoides: one anterolateral seta is absent from genu IV in some species of

Gamasellopsis: one posterolateral seta is absent from genu IV in some species of Gamasiphis; one posterolateral seta is absent from tibia

IV in Gamasellopsis. some species of Gamasiphis and some species of

Hydrogamasus; on tarsus IV, seta pd4 is absent in Gamasellopsis and

Gamasiphoides propinqua Womersley, 1956; finally, some species of

Gamasiphoides have one extra posterolateral seta on genu III (

2-2/1,2/1-2) and tibia III (2/1/1,2/1-2).

Diagnoses for the three subfamilies of the Ologamasidae are

given below.

Subfamily Ologamasinae Ryke, 1962

(Ologamasinae Ryke, 1962 sensu Lee, 1970 in part; Ologamasini

Lee, 1970 in part; Sessiluncinae Lee, 1970 in part).

Type Genus: Ologamasus Berlese, 1888

DIAGNOSIS

Seta all on palp genu almost always with at least 4 lateral prongs;

tectum variable but usually three-tined, central tine with lateral

denticles or spinules, and a divided or undivided tip; female

chelicerae with 3 teeth on the movable digit and usually 5-7 teeth on

the fixed digit; spermadactyl usually with "S"-shape base, always

freer than fused to the movable digit and projecting anteriad. 170 subequal in length to the movable digit or, at least one and a half times as long as the movable digit; arthrodial process of the chelicera almost always produced into a corona; dorsal shield always divided in the deutonymph and almost always entire in both sexes; podonotal region with 19-22 pairs of setae; opisthonotal region usually with 20 pairs of setae, but reduction in chaetotaxy also occurs; submarginal (UR) series absent; preendopodal shields almost always present, produced into one well sclerotized pair; metasternal setae on sterno-metasternal shield which is usually separate from the endopodal shields IV; female genital shield usually as wide as long, rounded anteriorly, truncate posteriorly and projecting only as far as the posterior margin of acetabula IV; male sternogenital shield almost always discrete, always bearing the genital setae; ventral glands gv2 opening in 2 or 3 solenostoma; ventrianal shield usually discrete in the female, but it may fused posteriorly to the dorsal shield and to the peritrematal shield in the male; opisthogastric region with 6-8 pairs of setae; setae Zvl almost always placed at level below that of setae Jvl; adanal setae placed at level above the anterior margin of the anal valves; postanal seta shorter than the adanal setae; peritrematal shields usually fused posteriorly to the exopodal shields IV; peritremes partly reduced in length; exopodal shields III split or entire, but always completely encasing peraxial edges of respective acetabula; pretarsus I present and always pedunculate; leg chaetotaxy usually as for the Rhodacaroidea, with the few exceptions occurring in Gamasitus and Gamasellopsis. 171

Subfamily Epiphidinae Kethley, 1983

Type Genus: Epiphis Berlese, 1916

DIAGNOSIS

Seta all on palp genu finely pilose; tectum uniramous, smooth, undivided at tip; deutonymph with a distal hyaline process on the fixed digit of the chelicera; female chelicerae with 3 teeth on the movable digit and 5 teeth on the fixed digit, arthrodial process produced into a corona; spermadactyl greatly elongate, straight, styletiform, fixed digit very reduced, arthrodial process absent; dorsal shield always entire in the deutonymph and in the adult; idiosomal shields extensively sclerotized in both sexes, with all idiosomal setae placed on shields; several idiosomal setae complex in the deutonymph but always simple in the adult; podonotal region with

21 pairs of setae, opisthonotal region with 28 pairs; submarginal

(UR) series present in 6 pairs; one pair of preendopodal shields present; metasternal setae placed on discrete platelets which may be narrowly fused to the sterno-metasternal shield; female genital shield discrete, wider than long, rounded anteriorly, truncate posteriorly, projecting only as far as the posterior margin of acetabula IV; spermathecae opening on ventral surface of coxa III; male with a holoventral shield; ventral glands gv2 opening in 2-3 solenostoma; female ventrianal shield discrete anteriorly, fused posteriorly to the dorsal shield; male ventrianal shield fused anteriorly to the sternogenital and to the exopodal shields IV, fused laterally and posteriorly to the dorsal shield; peritrematal shields extensive, widening posteriad, fused anteriorly to the dorsal 172 shield in the female, fused anterolaterally and posteriorly to the dorsal shield in the male, but always discrete from the exopodal shields II-IV in both sexes; peritremes reaching the anterior margin of acetabula I in the deutonymph, but reduced in the adult, reaching only up to middle level of acetabula II; opisthogastric region with

11-13 pairs of setae; opisthogastric setae Zvl placed at level well above that of setae Jvl; exopodal shields II-IV forming a continuous strip and completely encasing the peraxial edges of respective acetabula; pretarsus I pedunculate; tarsus I of deutonymph with an acrotarsal pseudosegmentation; tarsus I of male with a distal, large bulbous sensillum; leg II of male with the anteroventral (av) seta on femur modified into a thumb-like spur and seta av on tibia may be produced into a short, bullet-shaped process; seta av on genu not modified; leg chaetotaxy follows the Rhodacaroidea pattern.

Subfamily Gamasiphinae Lee, 1970

Type Genus: Gamasiphis Berlese, 1904

DIAGNOSIS

Tectum one or three-tined, central (or single) tine almost always undivided; female chelicerae usually with 3-4 teeth on the movable digit and 5-7 teeth on the fixed digit; spermadactyl with

"S"-shape base, almost always freer than fused to the movable digit and projecting anteriad; arthrodial process of the chelicera produced

into a corona; dorsal shield always divided in the deutonymph and always entire in the adult; idiosomal setae simple; podonotal region with 20-22 pairs of setae; opisthonotal region with 18-21 pairs of

setae, but hypertrichy occurs in one genus; except for two genera. 173 the submarginal (UR) series is absent; preendopodal shields almost always present, as at least one pair; metasternal setae on the sterno-metasternal shield, which is usually fused to the endopodal shields IV; female genital shield discrete, almost always (except for one genus) rounded anteriorly, truncate posteriorly, usually as wide as long and projecting as far as the posterior margin of acetabula

IV; male sternogenital discrete, almost always (except in one genus) carrying the genital setae; ventrianal shield discrete or fused to the dorsal shield in both sexes; opisthogastric region with 6-8 pairs of setae; opisthogastric setae Zvl always placed at level below that of setae Jvl; except for one genus, exopodal shields III completely encase peraxial edges of respective acetabula; pretarsus I always present, usually pedunculate; leg II of male with anteroventral (av) seta on femur, genu and tibia modified; leg chaetotaxy with several exceptions from the Rhodacaroidea pattern (listed in the familial description).

Taxa removed from the Rhodacaroidea Oudemans:

Tangaroellus porosus Luxton, 1968:

Luxton (1968) pointed out certain similarities of this genus to the Rhodacaridae sensu lato, but suggested that its strong sexual dimorphism and distinct leg chaetotaxy appeared to be a good case for establishing a new family group for it. Lee (1970), in agreement with

Luxton's point of view, erected the taxon Tangaroellinae Lee, 1970, within his Rhodacaridae.

In fact, Tangaroellus does possess several unique

characteristics that could justify the formation of a distinct group. 174 to accommodate it. These are: the chaetotactic pattern of femur II

(2-3/1,2/3-0) and femur IV (0-2/1,2/2-0); podonotal region oligotrichous, with 16 pairs of setae (z3, z6, si, s2, s3, rl, r4, and r6 absent); opisthonotal region paedoraorphic, with only 12 pairs of setae (Jl, SI, S2, and R1-R5 absent); opisthogastric region with 5 pairs of setae (Jv3, Jv4 and Zvl absent); coxa IV of male with a robust spur placed posteroventrally; on tarsus IV, setae ad3 and pd4 are absent. Regarding its relationships with other groups however, the results of my analysis indicated a closer relationship with the

Ascidae Voigts & Oudemans, 1905 sensu Lindquist & Evans, 1965, than with the Rhodacaridae sensu Lee. Tangaroellus shares four apomorphies with the Ascidae: palp claw two-pronged (6); metasternal setae (st4) placed on the soft cuticle posteriorly to the female sterno-metasternal shield (7); femur I with 12 setae (39); opisthonotal setae J5 placed at level posterior to that of setae Z5.

Other characteristics common to Tangaroellus and several taxa of

Ascidae include: seta all on palp genu simple ; female chelicerae with small teeth; spermadactyl straight, base not "S"-shaped; dorsal shield divided into subequal shields; podonotal setae j2 displaced anteriad, nearly parallel to setae zl; podonotal setae Z5 complex and

typically much longer than simple setae J5; preendopodal shields small, inconspicuous; female sterno-metasternal shield wider than

long, short, projecting only as far as the middle of coxae III;

endopodal shields II-IV free; female genital shield longer than wide, closely abutting the wide, rounded ventrianal shield, the ventrianal

shield resembling that of Gamasellodes regarding both its shape and 175 the position of the anal valves (nearly in the center of the shield); metapodal shields free in the female; peritrematal shields fused anteriorly to the dorsal shield, but free posteriorly (not fused to the exopodal shields IV); male sternogenital shield partly coalesced with the endopodal shields IV (a similar condition is found in males of Arctoseius Thor. 1930 and Leioseius Berlese, 1916); opisthogastric region with 5 pairs of setae; male leg II with the anteroventral (av) seta on femur, genu and tibia modified into a spur or a spine, although not as conspicuous as the spur commonly observed in the

Rhodacaroidea.

On the basis of the evidence presented above I choose to transfer this taxon to the Ascidae, retaining its subfamilial status.

Protogamasellus Karg, 1962 and Gamasellodes Athias-Henriot,

1961:

Protogamasellus was first included in the Rhodacaridae sensu lato by Karg (1965) who was followed in that placement by some authors (Shcherbak, 1976; Bregetova and Shcherbak, 1977;) but not by others (Lindquist & Evans, 1965; Lee, 1970). When first described,

Gamasellodes was referred to the Rhodacaridae sensu lato. In the same paper, Athias-Henriot described five new species of "rhodacarids" attributing them to the genera Rhodacarellus (R. mica. R. arcanus) and Rhodacaropsis (R.angustiventris. R. cognatus and ^ massula).

Lindquist & Evans (1965) transferred Protogamasellus and Gamasellodes to the Ascidae, noting that Rhodacarellus mica. Rhodacaropsis. angustiventris and Rhodacaropsis cognatus also appeared to belong to

the genus Protogamasellus. on the basis of their studies of the leg 176 chaetotaxy of Rhodacaropsis massula. My studies of all of the above mentioned taxa showed that except for Rhodacarellus arcanus. they share greater affinities with the Ascidae than with the other familial groups with which they were compared (Rhodacaridae,

Digamasellidae, Veigaiidae). Therefore, I concur with the retention of Protogamasellus and gamasellodes in the Ascidae and with the inclusion of four of Athias-Henriot* species in Protogamasellus as suggested by Lindquist & Evans.

Laelaptonvssus Womersley, 1956 and Puchihlungia Samsinak, 1964;

Womersley (1956) described Laelaptonvssus as a "laelaptid" mite, but erected a new family to include it.

When first described, Puchihlungia was referred to the

Rhodacaridae sensu lato. Lee (1970) synonymized the genera and placed them in the Laelaptonyssinae Womersley within his Rhodacaridae. My studies indicate that these two taxa are distinctive enough to retain their generic identities, while sharing apomorphies which justify their placement in the same family group. Therefore, the original familial status is retained and expanded to include Puchihlungia

Samsinak.

Panteniphis Willmann, 1949 and Lindquistoseius Genis, Loots &

Ryke, 1969:

The genus Panteniphis was first included in the Rhodacaridae sensu lato by Athias-Henriot (1968). Lee (1970) transferred it to the

Ascidae on the basis of its leg chaetotaxy, a two-pronged palp claw and inconspicuous enlargement of setae on the male leg II.

When first described, the genus Lindquistoseius was placed in 177 the Ascidae mainly on the basis of its leg chaetotaxy and some gnathosomal characters. Hurlbutt (1975) regarded Lindquistoseius as congeneric with Panteniphis and reduced its status to the subgeneric level, while considering it to be more closely related to the

Rhodacaridae sensu lato than to the Ascidae.

On the basis of the results of this study (refer to "Results and

Discussion" Section) I have erected a new family group,

Panteniphididae within the Laelaptonyssoidea Womersley, 1956 new status to include both taxa.

A key to the families and subfamilies of Rhodacaroidea is presented, followed by systematic accounts on the Laelaptonyssoidea. 178

Key to the families and subfamilies of Rhodacaroidea

(Based on adults)

1 __ Dorsal shield usually entire or with a line of fusion demarcating podonotal and opisthonotal regions or rarely divided.

Seta all on palp genu usually with several lateral prongs; if bifurcate or spatulate, many dorsal setae complex and/or dorsal shield entire. Arthrodial process of female chelicera always produced into a corona. Dorsal hexagonal area of podonotum never with scleronoduli. Submarginal series usually absent. Dorsal setae usually simple. Preendopodal shields present or incorporated to sternal shield, but never absent; if incorporated to sternal, then dorsal shield entire. Metasternal setae on sterno-metasternal shield 0£ rarely on discrete platelets which may be narrowly fused to sterno-metasternal shield. Female genital shield usually as wide as long and projecting posteriorly only as far as the posterior margin of acetabula IV. Pretarsus I always present ......

Family OLOGAMASIDAE ...... 2

Dorsal shield usually divided or rarely entire. Seta all on palp genu simple or complex; if simple, then most dorsal setae simple and dorsal shield divided; if complex, then most dorsal setae almost always complex (except for Hevdeniella. Solueamasus. Nodacaroides). dorsal shield divided and/or ventral shields extensive in both sexes. 179

Arthrodial process of female chelicera produced into a corona or a

brush. Dorsal hexagonal area of the podonotum with or without

scleronoduli. Submarginal series usually present. Dorsal setae

usually complex or rarely simple. Preendopodal shields present or

incorporated to sternal shield or absent; if incorporated to sternal,

then dorsal shield divided. Metasternal setae on sterno-metasternal

shield or on discrete platelets or, on soft cuticle. Female genital

shield usually longer than wide and projecting posteriorly well

beyond posterior margin of acetabula IV. Pretarsus I present or

absent ...... 4

2 __ Metasternal setae on discrete platelets which may be narrowly

fused to the sterno-metasternal shield. Fixed digit of male chelicera

very reduced. Arthrodial process of male chelicera absent.

Spermadactyl greatly elongate, styletiform, projecting anteriad.

Tarsus I of male with a distal bulbous sensillum. Idiosomal shields

extensive in both sexes. All idiosomal setae on shields. Submarginal

series present. Peritrematal shields extensive, but never fused to

the exopodal shields IV. Setae Zvl placed at level well above that of

setae Jvl ...... OLOGAMASIDAE - EPIPHIDINAE

Metasternal setae always on sterno-metasternal shield. Fixed

digit of male chelicera never reduced. Arthrodial process of male

chelicera always present. Spermadactyl shorter or longer than movable

digit of chelicera, projecting anteriad or posteriad, but never

styletiform. Tarsus I of male never with a distal bulbous sensillum. 180

Some idiosomal setae on soft cuticle. Submarginal series usually absent. Peritrematal shields may or may not be extensive, but usually fused to exopodals IV. Setae Zvl usually placed at level below that of setae Jvl ...... 3

3 __ Seta all on palp genu usually with at least 5 lateral prongs;

if bifurcate, then exopodal shields III split; if spatulate, then dorsal shield with line of fusion between podonotal and opisthonotal

regions and with holoventral shield in the male. Spermadactyl always projecting anteriad. Peritrematal shields usually fused to the exopodal shields IV in both sexes, and to the ventrianal shield in the male or if not fused to any of these shields, seta all on palp genu with at least 5 lateral prongs ......

OLOGAMASIDAE - OLOGAMASINAE

Seta all on palp genu variously produced (spine-like, bifurcate, spatulate or finely pilose); if bifurcate, then exopodal shields III entire; if spatulate, dorsal shield entire and male sternogenital shield discrete. Arthrodial process of male chelicera produced into a corona or rarely into a leaf-shaped flap. Spermadactyl projecting anteriad or posteriad. Peritrematal shields usually fused to exopodal shields IV in both sexes or if not fused, then seta all on palp genu at most finely pilose, but never with several lateral prongs

...... OLOGAMASIDAE - GAMASIPHINAE 181

4 __ Degree of development of shields similar in both sexes.

Arthrodial process of chelicera always produced into a corona.

Preendopodal shields present or incorporated to sternal shield.

Female genital shield usually projecting posteriorly as far as

posterior margin of acetabula IV. Exopodal shields well developed

(except in Evanssellus) Family GAMASELLIDAE

Degree of development of shields more extensive in the male.

Arthrodial process of chelicera produced into a corona or a brush.

Preendopodal shields present or absent, but never incorporated to

sternal shield. Female genital shield usually projecting posteriorly well beyond posterior margin of acetabula IV. Exopodal shields

usually not well developed ...... 5

5 __ Submarginal series absent. Dorsal hexagonal area of podonotum with 2-4 scleronoduli or if scleronoduli absent, then arthrodial

process of chelicera produced into a brush. Metasternal setae on

sterno-metasternal shield or on soft cuticle. Peritrematal shields

not always fused to exopodal shields IV ...... Family RHODACARIDAE

Submarginal series usually present or_ if absent (Notogamasellus).

then all dorsal setae complex. Dorsal hexagonal area of the podonotum

never with scleronoduli. Metasternal setae always on

sterno-metasternal shield. Peritrematal shields always fused to

exopodal shields IV ...... Family EURYPARASITIDAE 182

Superfamily Laelaptonyssoidea Womersley, 1956a new status

The superfamily Laelaptonyssoidea Womersley is regarded here as including three families: Laelaptonyssidae Womersley, 1956a;

Digamasellidae Evans, 1957; Panteniphididae n. fam. Diagnoses and descriptions for the superfamily and the newly established family are presented below. Diagnoses and descriptions of the Digamasellidae and

Laelaptonyssidae are given by Lindquist (1975) and Lee (1970) respectively.

DIAGNOSIS

Palp claw almost always two-pronged; deutosternum usually with 5 rows of denticles, the fifth row (or the basal, if 6 rows are present) widened ; movable digit of female chelicera with 3-4 teeth; male spermadactyl distally free, recurved basally before projecting anteriad or posteriad; dorsal shield divided in the deutonymph and almost always divided in the adult; podonotal region with 19-22 pairs of setae, opisthonotal region with 16-20 pairs, but further reduction in chaetotaxy also occurs; podonotal setae rl and r6 absent; submarginal (UR) series absent; almost always, female with a

sterno-metasternal shield which usually bears the metasternal setae;

female genital shield bearing 1 pair of setae and projecting

posteriorly well beyond margin of acetabula IV; male with a

sternogenital shield; male genital setae almost always present, set

on the sternogenital shield or on separate triangular platelets;

ventrianal shield usually discrete in the fëmale and fused to other

shields in the male; anal valves almost always small; adanal setae

elongate, often longer than the postanal seta; opisthogastric region 183 usually with 6-8 pairs of setae, although further reduction in chaetotaxy also occurs; femur I usually with 13 setae (rarely, 11 or

12 setae) including 4 ventrals; genu I usually with 12 setae (rarely,

11 or 13 setae) including 3 (or 2) ventrals; tibia I usually with 12 setae (rarely, 9 or 11 setae) including 3 (or 2) ventrals; tibia IV usually with 7 setae (rarely, 6 or 8) including 2 ventrals.

DESCRIPTION

Size small, varying from less than 400u and up to 6Q0u.

Gnathosoma: Palp claw almost always two-pronged, the exception being the Laelaptonyssidae with a secondarily derived three-pronged palp claw; seta all on palp genu usually simple (spatulate in

Lindquistoseius): tectum usually three-tined, with central tine smooth and undivided; salivary stylets and corniculi well developed; deutosternum with 5 rows of denticles (rarely, 6 rows may be present in some Digamasellidae), the fifth (or basal row) widened; female chelicerae commonly with small teeth; there are 3-4 teeth on the movable digit and usually 5-7 teeth on the fixed digit, although a reduction to 3 teeth on the fixed digit occurs in the

Laelaptonyssidae and Panteniphis; male spermadactyl of various lengths, always distally free and recurved basally before projecting anteriad (Digamasellidae, Panteniphis) or posteriad

(Laelaptonyssidae, Lindquistoseius); arthrodial process of chelicerae produced into a corona (most Digamasellidae, Panteniphis) or into an arthrodial brush (Lindquistoseius and several species of

Dendrolaelaps); the corona may be reduced in some species of

Digamasellidae and it is absent in the Laelaptonyssidae. 184

Dorsum: Dorsal shield always divided in the deutonymph and usually divided in the adult (Panteniphididae has it entire); podonotal region usually with 19-22 pairs of setae (6 j, 6 z, 4-6 s,

1-4 r) and the opisthonotal region with 16-20 pairs (4-5 J, 5 Z, 5 S,

1-5 R), but a further reduction in the number of setae on both regions is observed in the Laelaptonyssidae (Laelaptonvssus with 14 pairs of opisthonotals and Puchihlungia with 13 pairs of podonotals and 14 pairs of opisthonotals) and Digamasellidae (Loneoseius may have only 12 pairs of podonotals); podonotal setae rl and r6 always absent; in the female (or rarely, in the male), some podonotal and opisthonotal setae may be set on soft cuticle, these usually including setae of the r series (r2-r5) and of the R series (Rl); exceptionally, Panteniphis exhibits setae jl and zl on the soft cuticle, whereas its r series is set on the shield; opisthonotal setae Z5 commonly placed at a level parallel or anterior to that of setae J5, often conspicuously longer than setae J5; submarginal series absent; podonotal glands gd4 small; podonotal shield bears four scleronoduli on the dorsal hexagonal area of most of the

Digamasellidae, except for Digamasellus which has secondarily lost them.

Venter: Preendopodal shields commonly absent (most

Digamasellidae, Laelaptonvssus) or they may be rudimentarily developed as a coarsely granulated pair (some species of the

Dendrolaelaps cornutus species group of Hirschmann, 1960) or they may be produced into one weakly (Puchihlungia) or well sclerotized pair

(Panteniphididae); female almost always with a sterno-metasternal 185 shield (except for Laelaptonvssus which has a sternal shield), often weakly defined anteriorly (sometimes posteriorly also); metasternal setae (st4) usually on sterno-metasternal shield, rarely on soft cuticle (Panteniphis. Laelaptonvssus); first sternal setae (stl) usually placed on sternal shield, but in Lindquistoseius they are

placed on the soft cuticle between the sternal and the preendopodal

shields; in several females of the Dendrolaelaps cornutus group,

setae stl are placed on the soft cuticle on an intermediate area

between the weakly sclerotized sternal shield and a pair of coarsely

granulated shields (these are the structures to which I refer as rudimentary preendopodal shields); peritrematal shields usually present and often fused to the podonotal shield anteriorly, free posteriorly and rarely fused to the exopodal shields IV

(Panteniphididae and the male of Digamasellus australis Lindquist,

1975); peritremes extremely reduced in length and peritrematal shields absent in the Laelaptonyssidae; exopodal shields III never completely encasing peraxial edges of acetabula III; female genital

shield commonly subrectangular, rounded or truncate anteriorly, always truncate posteriorly and extending well beyond acetabula IV,

bearing the genital setae; genital pores usually off shield, rarely

on it; accessory opening to the spermathecae commonly associated with

the proximal segments of legs III (acetabulum, coxa, trochanter and

femur) and IV (acetabulum and coxa); male with a sternogenital

shield; male genital setae (st5) placed on the sternogenital shield

(Panteniphididae, Puchihlungia). on separate platelets

(Digamasellidae) or rarely, absent (Laelaptonvssus); ventral glands 186 gv2 usually present in one solenostomum; ventrianal shield always

present, usually discrete in the female but in the male it may be

fused to the opisthonotal shield posteriorly and laterally (most

Digamasellidae), or to the opisthonotal and peritrematal shields

(some Digamasellidae) or, extensively fused to the dorsal shield and

to other ventral shields (Panteniphididae); female ventrianal shield

usually subrectangular, rarely subtriangular (as in Digamasellus

australis. for instance); anal valves almost always small (except for

Digamasellus. with secondarily enlarged anal valves); adanal setae

elongate, tending to be longer than the postanal seta, and often

placed at level anterior or above that of the anal valves;

opisthogastric region usually with 6-8 pairs of setae (3-5 Jv, 1-3

Zv), although a total of 4 pairs (3 Jv, 1 Zv) is observed in

Panteniphis; if only 4 pairs are present, Jv3, Jv4, Zvl and Zv3 are

absent; if 6 pairs, Jv4 and Zv3 (or rarely, Zvl) are absent; Jv4 is

almost always absent except for Puchihlungia. which has the full

complement of 8 pairs of setae; setae Jv5 and/or Zv2 (rarely Zv3)

often placed on the soft cuticle.

Legs: Legs I-IV almost always provided with two preclaws,

bilobed pulvilli and a pair of pointed membranous processes arising

from the bases of the claws; the Laelaptonyssidae are clawless, with

pretarsus I larger than the other pretarsi, and exhibiting acrotarsal

pseudosegmentation in protonymph, deutonymph and adult instars;

tarsus I of males of Panteniphididae with a large apical bulbous

sensillum; male leg II with anteroventral seta (av) always modified

into a spur of moderate size, seta av on genu and tibia not modified; 187 the chaetotactic pattern for legs I-IV exhibits a general trend toward reduction (when compared to the Rhodacaroidea) and it is separately listed for each of the three families.

Family Digamasellidae:

coxae I-IV: (2-2-2-1); trochantera I-IV: (6-5-5-S); femora I-IV:

(2-5/4-2) (2-5/3-1) (1-4/1-0) (1-3/1-1); genua I-IV: (2-3/2,2/1-2)

(2-3/1,2/1-2) (2-2/1,2/0 or 1-1) (1-2/1,2/0-1); tibiae I-IV:

(2-3/2,2/1-2) (2-2/1,2/1-2) (2-1/1,2/1-1) (1-1/1,2/1-1); tarsi II-IV:

(2 or 3-3/3,1,3/2-2 or 3); (2 or 3-3/3,1,3/2-2 or 3) (2 or 3-3/3,1,2 or 3/2-2 or 3).

Family Panteniphididae:

coxae I-IV: (2-2-2-1); trochantera I-IV: (5 or 6-5-5-5); femora

I-IV: (2-5/4-1) (2-5/3-1) (1-4/1-0) (1-3/1-1); genua I-IV: (2-3/1 or

2,2/1-2) (2-3/1,2/1-2) (2-2/1,2/0-1) (1-2/1,1/0-1); tibiae I-IV:

(2-3/1 or 2,2/1-2) (2-2/1,2/1-2) (2-1/1,2/1-0) (1-1/1,2/1-0); tarsi

II-IV: (3-3/3,1,3/2-3) (3-3/3,1,3/2-3) (3-3/3,1,2 or 3/2-2 or 3).

Family Laelaptonyssidae:

coxae I-IV: (2-2-2-1); trochantera I-IV: (6-6-5-5); femora I-IV:

(11 - 10 - 7 or 8 - 6 or 7); genua I-IV: (12 or 13 - 9 or 10 - 8 or 9

- 7 or 10); tibiae I-IV: (9 or 11 - 7 or 8-9 - 6 or 8 - 7 or 8);

tarsi II-IV: (17 or 18 - 17 or 18 17 or 18). Where 17 setae, av3

absent. 188

Family Panteniphididae new family

Type Genus: Panteniphis Willmann, 1949

DIAGNOSIS

Palp claw two-pronged; deutosternum with 5 rows of denticles, fifth row widened; movable digit of female chelicera with 3-4 teeth; male spermadactyl longer than the movable digit, distally free, recurved basally and projecting anteriorly or posteriorly; dorsal shield usually entire; podonotal region with 19-21 pairs of setae, opisthonotal region with 19-20 pairs; preendopodal shields present in one pair; female with a sterno-metasternal shield, usually bearing metasternal setae; male with a sternogenital shield bearing genital setae; ventrianal shield subrectangular and discrete in the female, whereas in the male it is always extensively fused to other ventral shields (sternogenital, peritrematals and exopodals IV) and posteriorly and laterally to the dorsal shield; peritrematal shields fused anteriorly to the dorsal shield and posteriorly to the exopodal shields IV in the female, additionally fused to the ventrianal shield in the male; anal valves small; adanal setae elongate and conspicuously longer than the postanal seta; opisthogastric region with 4-7 pairs of setae; tarsus I of male with large apical bulbous sensillum; femur I with 12 setae including 4 ventrals; genu IV with 6 setae; tibia III with 7 setae, tibia IV with 6 setae.

DESCRIPTION

Mostly small mites, size ranging from 300-350u in males and from

330-450u in females.

Gnathosoma: Palp claw two-pronged; seta all on palp genu simple 189

(Panteniphis) or spatulate (Lindquistoseius); tectum three-tined and smooth, with central tine arising more ventrally than the lateral tines (Lindquistoseius) or, with central tine truncate and the lateral tines denticulate (Panteniphis); deutosternum with 5 rows of denticles, fifth row widened; corniculi well developed, simple; female chelicerae with 3 (Panteniphis) or 4 teeth (Lindquistoseius) on the movable digit, and 3 (Panteniphis) or 6-7 teeth in the fixed digit (Lindquistoseius); arthrodial process of chelicera produced into a corona in Panteniphis or into an arthrodial brush in

Lindquistoseius; male spermadactyl distally free, longer than the movable digit and basally recurved before projecting anteriad

(Panteniphis) or posteriad (Lindquistoseius).

Dorsum; Dorsal shield entire, except for the male of L. africanus sensu Hurlbutt which has extensive, but divided shields; dorsal setae simple; podonotal glands gd4 not enlarged; podonotal region with 19-21 pairs of setae (6 j, 6 z, 4-5 s, 3-4 r), setae rl and r6 always absent; opisthonotal region with 19-20 pairs (5 J, 5 Z,

5 S, 4-5 R); submarginal (UR) series absent; podonotal setae jl and zl and opisthonotal setae Rl set on soft cuticle in the female of

Panteniphis.

Venter: Preendopodal shields present in one pair; female with a sterno-metasternal shield usually including the metasternal setae

(st4) - in females of Panteniphis they are often set on the soft cuticle on one side; first sternal setae (stl) placed on the sternal shield (Panteniphis) or on the soft cuticle between the sternal and the preendopodal shields (Lindquistoseius); peritrematal shields 190 present and well developed, fused anterioly to the dorsal shield and posteriorly to the exopodal shields IV in the female, additionally to the ventrianal shield and laterally to the dorsal shield in the male; exopodal shields II-IV coalesced, but not completely encasing peraxial edges of respective acetabula; female genital shield as wide as long, anterior margin rounded, posterior margin truncate, projecting well beyond acetabula IV, bearing 1 pair of setae; genital pores on soft cuticle; accessory opening to the spermathecae placed on the ventral surface of coxa IV (Panteniphis) or on the ventral surface of basifemur III, near the suture (Lindquistoseius); male with a holoventral shield (sternogenital shield coalesced with the ventrianal shield), genital setae (st5) on shield; ventral glands gv2 present in one solenostomum; ventrianal shield subrectangular and discrete in the female, fused posteriorly and laterally to the dorsal shield and to the sternogenital, exopodals IV and peritrematal shields in the male; anal valves small; adanal setae elongate, placed at level well above that of the anal valves and conspicuously longer than the postanal seta; opisthogastric region with 4-7 pairs of setae

(3-4 Jv, 1-3 Zv); in the female of Panteniphis setae Jv3, Jv4, Zvl and Zv3 are absent, and setae Jv5 are set on soft cuticle; in the female of Lindquistoseius setae Jv4 are absent and setae Jv5, Zvl and

Zv3 are set on soft cuticle.

Legs: Legs I-IV provided with preclaws, pulvilli and membranous processes from between the bases of the claws; tarsus I of male with a large apical bulbous sensillum; leg chaetotaxy generally reduced

(listed above) and including a unique arrangement for the following 191 segments: femur I with 12 setae including 4 ventrals; genu IV with 6 setae; tibia III with 7 setae and tibia IV with 6 setae. 192

Figure 1. Suprafamilial and interfamilial relationships of the

Rhodacaroidea. Numbers on figure refer to stems.

Characters are listed following sequence of stems.

Character states are listed between parentheses.

STEM 1; 1(2); 3(1); 7(1); 10(2); 22(1); 25(1); 35(3); 37(1). STEM 2:

4(1); 6(2); 18(1); 19(1); 26(2); 31(2); 36(3); 38(2); 42(1); 43(1);

44(2); 45(2); 47(3). STEM 3; 17(1); 26(1); 31(1); 33(1); 38(1). STEM

4: 30(2); 57(1). STEM 5: 26(2); 34(1); 36(1); 37(2) 38(2). STEM 6

1(1); 4(1); 5(1); 19(1); 30(2); 31(2); 33(2). STEM 7: 9(1); 23(1)

35(4); 36(5); 55(1); 59(1). STEM 8; 19(1); 35(5); 37(5); 57(1)

60(2). STEM 9; 49(2); 54(-l). STEM 10: 5(1); 8(1); 15(1); 24(-l)

30(1); 37(2); 56(1); 57(1); 61(4). STEM 11: 45(3); 54(-l); 55(2)

STEM 12: 34(2); 60(1); 61(1). STEM 13: 15(1); 35(5); 37(5); 49(4)

59(2). STEM 14: 12(1); 17(1); 44(3); 46(1); 47(4); 59(2). STEM 15

23(1); 26(4); 27(2); 33(3); 34(-l); 35(5); 36(1); 39(2); 40(-l)

56(-l). STEM 16: 1(1); 31(-1); 33(2); 55(-l); 60(-l); 61(2). STEM

17: 19(1); 58(2). tti < Ml o < a w Ul Ul w to < < < 3 > I o z mt I z !: o M Ul 3c o K 3 « < M a. < < -J < < B & K O Ul : & s < < M < « < J & o < o X o 5 Ul o s o & < 3 X S o < < & a ij Ul K o Ul o 14 18 10

10 12 J J Î 3 Î LAELAPTONYSSOIDEA Î RHODACAROIDEA s 194

Figure 2. Intrafamilial relationships of the Euryparasitidae n. fam.

Refer to Figure 1 legend for explanation.

STEM 1: 1(2); 3(1); 7(1); 10(2); 17(1); 22(1); 25(1); 26(1); 30(2);

33(1); 35(3); 37(1); 57(1). STEM: 2(2); 34(1). STEM 3: 13(2); 31(1);

37(2); 38(1). STEM 4: 13(3); 29(2); 30(0); 33(0); 54(1); 61(3). STEM

5: 19(1); 23(1); 31(2); 33(2); 35(2); 36(1); 52(1). STEM 6: 13(-1)

14(-2); 25(0); 26(-l); 2991); 35(5). STEM 7: 10(0); 33(2); 50(-l)

60(-l). STEM 8: 6(1); 12(-1); 22(0); 31(3); 32(1); 43(-l); 46(-l)

57(0); 60(-l). STEM 9: 1(1); 3(0); 4(-l); 8(1); 10(0); 19(1); 21(2)

34(1); 55(1); 60(1). STEM 10: 2(2); 14(2); 21(2); 32(1); 37(3)

55(1); 56(1);60(1). STEM 11: 14(0); 19(2); 25(-l); 26(-2); 27(-l)

29(2); 30(3); 37(0); 38(2); 43(-l); 46(-l); 52(2); 57(0). 3 3 a # • # a m # « E « « « m E a ë E « o m a k m 9 # 9 o o c 111 3 4 4 2

<> ( 1 <> 44 <1 4 > 6 7 8 9 10 11

i> 4> 4» <> 44 <4 T

EURYPARASITIDAE

a 196

Figure 3. Intrafamilial relationships of the Rhodacaridae. Refer to

Figure 1 legend for explanation.

STEM 1: 5(1); 7(1); 10(2); 19(1); 25(1); 26(2); 30(2); 31(2); 33(2);

34(1); 35(3); 37(2); 38(2). STEM 2; 3(1); 4(1); 34(2); 54(1). STEM 3:

1(1); 22(1); 27(-l); 36(5); 37(3); 41(1); 48(1). STEM 4: 14(2);

36(1); 52(1). STEM 5; 1(1); 4(2). STEM 6; 2(2); 8(2); 35(5); 36(4);

37(3); 55(1). STEM 7: 17(1); 30(3); 57(1); 60(4). STEM 8: 24(2). STEM

9: 17(1); 23(1); 55(-l). STEM 10: 4(-l); 7(-l); 10(0); 17(1); 33(1);

60(-l). STEM 11: 14(1); 18(1); 28(1); 41(1); 48(1); 55(1). STEM 12:

2(2); 14(1); 19(0); 26(4); 31(3); 32(1); 36(5). STEM 13: 1(2); 22(1);

34(1). STEM 14: 4(1); 8(1); 24(2); 30(1); 32(1); 50(1); 59(1). STEM

15: 10(1); 14(-2); 18(1); 54(1); 61(1). STEM 16: 2(1); 5(2); 8(1);

10(1); 23(1); 26(3); 33(3); 40(1); 52(0); 54(0); 57(1). STEM 17:

7(2); 14(-2); 24(-10; 52(2). STEM 18: 14(1); 21(1); 37(1); 55(1);

56(1); 60(0); 61(1). STEM 19: 23(1); 26(3); 31(3); 52(1). STEM 20:

24(-l); 51(1). STEM 21: 37(2); STEM 22: 5(2); 22(1); 30(3); 33(1);

35(2); 54(0); 61(4). STEM 23: 2(2); 26(3); 28(0); 30(0); 34(1);

35(0); 55(0). STEM 24: 1(0); 19(0); 26(3); 36(1); 37(5); 52(2);

54(0); 57(0). STEM 25: 5(3); 10(1); 17(0); 19(2); 20(1); 26(4);

35(1); 36(0); 37(0); 56(1). STEM 26: 1(2); 9(1); 35(4); 50(1);

54(-l); 59(1); 60(1). STEM 27: 37(5); 55(1). STEM 28: 8(1); 22(0);

30(0); 32(1). STEM 29: 1(2); 23(0); 35(2); 36(3). STEM 30: 10(2);

22(0); 23(0); 55(-l). STEM 31: 1(2); 5(0); 30(0); 32(1); 35(5);

54(-l); 55(0); 56(1); 60(1). o # - o - .Lltogam atus

» o- .Rhodacaropaia N N R h o d a c a ru a

M M M L(y_^Rhodacarallua

e» # — 0- in a

M # — O- .Solugamasua » O- .Gan. n. #1 6>

Nodacaroldea c o a ta i M 01 N. minyaspia

M fl» « _0 __Q,Afrogamaaallua M N . m o n g ii 7 3 Z u H)—G- O ♦ — O- .Madiodacarallua o b a k a ri M» > M o O .Afrodacarallua > N 7 3 » A . c it r i O » > O m M L. lubalanaia Ci Lo—^Latogamaaellua L 6 l 198

Figure 4. Intrafamilial relationships of the Gamasellidae* Refer to

Figure 1 legend for explanation.

STEM 1; 1(2); 3(1); 7(1); 9(1); 10(2); 17(1); 19(1); 22(1); 23(1);

25(1); 26(2); 31(1); 35(5); 36(5); 37(5); 38(2); 55(1); 57(1); 59(1);

60(2). STEM 2; 11(1); 12(1); 1591); 3391); 54(1); 58(2); 60(4). STEM

3; 14(1); 60(3). STEM 4; 31(2); 34(1). STEM 5: 10(1); 13(2) 29(1)

38(1); 54(-l); 60(4). STEM 6; 14(2); 35(3);. STEM 7; 14 (0); 30(1)

STEM 8; 19(0); 22(0); 49(4); 61(3). STEM 9; 30(1); 34(1). STEM 10

1(1); 9(0); 15(1); 26(1); 30(2); 36(2); 38(0). STEM 11; 23(0); 36(1)

37(2). STEM 12: 33(2); 35(3); 61(1). STEM 13: 3(0); 14(1); 23(0)

STEM 14: 1(0); 2(1); 8(2); 9(0); 18(1); 19(0); 21(1); 30(1); 36(3)

52(2). STEM 15: 11(1); 12(1); 14(2). STEM 16: 4(1); 11(1); 22(0)

26(-l); 31(-1); 33(1); 38(0); 49(2); 56(1); 58(2); 60(2). STEM 17

13(3); 21(1). STEM 18: 9(0); 10(0); 12(1); 55(0); 57(0). STEM 19

2(2); 13(3); 15(1); 58(1). STEM 20: 4(1); 13(-2); 26(0); 29(1)

33(0); 38(0); 54(0); 56(2); 57(2); 58(0); 60(2). STEM 21: 3(0)

12(0); 16(1); 31(2); 57(0); 60(4). STEM 22: 1(1); 30(0); 34(2)

37(2); 49(4). STEM 23: 11(0); 14(2); 17(0); 30(2); 48(1); 51(1)

56(1). STEM 24: 1(1); 5(1); 16(1); 49(3); 54(0); 57(0); 60(2); 61(1)

STEM 25: 10(0); 11(0); 22(2); 30(2); 33(2); 34(0); 36(4); 52(1)

56(1); 59(2); 60(4). STEM 26: 7(0); 22(2); 30(1); 54(1). 199

Figure 4. (Continued)

STEM 27; 1(0); 13(3); 16(1); 23(0); 29(1); 43(-l); 46(-l); 53(1)

54(-l); 58(2). STEM 28: 35(4). STEM 29: 3(0); 5(1); 10(2); 22(0)

36(3); 54(0) 60(3). STEM 30: 21(1); 25(-l); 26(-l); 31(0); 37(2)

54(0); 54(0) 60(2). STEM 31; 6(1); 7(-l); 19(2); 22(0); 23(0)

29(2); 30(2) 31(2); 38(2); 54(1); 59(0). STEM 32: 10(2); 11(1)

14(-1); 25(-l); 26(-l); 31(0); 30(3); 34(2); 44(1); 60(0). STEM 33

11(0); 29(2); 52(1); 60(3). » l-e& O - Pyrlphis -*-0- M H#yd#nl#lla

M O" Onehogamasus co m m u n ia K» w Onchogamaaua v irg u n c u la M Hiniphia u h in n u a C—» M 01 H ln ip h la b ip a ia "O-#- o - . Rhodacaroidaa aagyptiacua M .Q . Paammonsaila

O .Gamaaaliut w -•-o - pyrlform la gr.

# - o - jQ. diacutatus gr. M œ D .G. falclgar gr. M > tt S ,G. tragardhi gr. > W CO e m • rS— O-' P lia llu a I" C illu a

ta m M g p#—O- Euapicrlua * - 0 - H u ta 003 * 0 m Gamasalliphia 201

Figure 5. Intrafamilial relationships of the Ologamasinae. Refer to

Figure 1 for explanation.

STEM 1; 1(2); 3(1); 7(1); 9(1); 10(2); 17(1); 22(1); 23(1); 25(1)

26(2); 31(1); 33(1); 34(2); 35(4); 36(5); 37(2); 38(2); 49(2)

54(-l); 59(1); 60(1); 61(1). STEM 2; 12(1); 19(1); 49(3); 52(1)

55(-l); 59(2). STEM 3; 31(2); 37(3); 55(1); 60(4). STEM 4: 36(3)

54(0); 61(3). STEM 5; 35(5); 37(5). STEM 6: 34(1); 61(2). STEM 7

49(4); 57(1). STEMS; 32(1). STEM 9: 19(1); 29(1); 3593); 59(2). STEM

10: 10(0); 17(0); 26(3); 31(3); 37(4); 58(2). STEM 11: 31(1); 32(1);

52(1). STEM 12: 13(1); 15(1); 21(1); 29(1). STEM 13: 19(1); 33(2).

STEM 14: 1(0); 14(1); 19(0); 35(5); 37(2); 49(3); 51 (1); 59(1). STEM

15; 2(2); 37(3). STEM 16; 21(2); 34(1). STEM 17; 11(1); 29(2); 32

(0); 34(0); 55(0); 58(2); 60(2). STEM 18; 2(2); 2(2); 4(2); 30(1)

49(1); 61(3). STEM 19: 17(0); 23(0). STEM 20: 36(3); 58(2). STEM 21

3(0); 13(1); 14(-1); 22(-l); 26(3); 29(0); 31(2); 36(2); 42(1)

43(1); 44(1); 47(1); 48(1); 52(0); 57(0). ie 17 18 20 21

12 13 14 18

10 11

2 OLOGAMASINAE I 203

Figure 6. Intrafamilial relationships of the Epiphidinae and

Gamasiphinae. Refer to Figure 1 legend for explanation.

STEM 1; 1(1); 3(1); 7(1); 9(1); 15(1); 22(1); 23(1); 25(1); 35(5);

36(5); 37(5); 49(4); 54(-l); 59(2). STEM 2; 2(2); 7(0); 31(-1); 33

(2); 55(-l); 60(2) 60(-l); 61(2). STEM 3: 1(2); 17(1); 19(1); 31(1);

33(1); 34(1); 55(1). STEM 4: 10(1); 26(-l); 54(0); 58(2); 60(2). STEM

5; 26(2); 31(2); 38(2). STEM 6; 2(2); 10(2); 11(1); 30(1); 44(1)

58(2). STEM 7: 14(1); 57(1). STEM 8: 15(0); 17(0); 30(2);,. STEM 9

26(3); 32(1). STEM 10; 21(2); STEM 11: 1(0); 23(0); 26(3); 33(0)

52(1); 56(1). STEM 12: 4(1); 14(-1); 22(0); 31(3); 34(2); 36(2)

51(0); 60(-l); 61(2). STEM 13: 19(0); 37(2); 44(0); 49(3); 54(1)

59(1). STEM 14: 3(0); 35(3); 36(3); 37(4); 44 (1); 49(3); 58(1)

59(1). STEM 15: 2(10); 9(0); 22(0); 44(0). STEM 16: 4(2); 8(3)

44(1); 47(1); 60(2). STEM 17: 9(0); 14(0); 15(0); 19(0); 22(0)

30(2); 31(1); 36(3); 37(3); 38(1); 54(1); 60(1). m « « a > « a # E m a o a k E E •o a « ■ >» Ul o (0 o o 12 t " t 14 { "t "t IZ • Î "( 6 I L Î J J

GAMASIPHINAE ro 205

SUMMARY

Concepts of the Rhodacaroidea (= Rhodacaridae Gudemans, 1902 sensu lato) proposed by three recent authors - Lee, 1970, 1971;

Lindquist, 1975; Shcherbak, 1976, 1979, 1980 - were reviewed in an attempt to determine the internal and external relationships of the group. Using phylogenetic methods and Farris' algorithm "Wagner 78", a comparative study of sixty-one characters in seventy-six generic taxa resulted in the construction of six consensus cladograms illustrating suprafamilial and familial relationships. The

Laelaptonyssoidea Womersley, 1956 new status emerged as the sister group to the Rhodacaroidea, with the sharing of eigth apomorphic character states. Both taxa were found to constitute monophyletic groups, the Laelaptonyssoidea as the more derived taxon with thirteen apomorphies, and the Rhodacaroidea with five apomorphies.

The Rhodacaroidea was redefined to include four families:

Rhodacaridae Oudemans, 1902, Euryparasitidae new family, Gamasellidae

Hirschmann, 1962, new status and Ologamasidae Ryke, 1962, with the

Ologamasidae consisting of three subfamilies: Ologamasinae Ryke,

1962, Gamasiphinae Lee, 1970 and Epiphidinae Kethley, 1983. The new arrangement of the Rhodacaroidea includes fifty-two genera, one of which (Mediodacarellus) is newly established and one undescribed. The subfamily Tangaroellinae Lee, 1970 was transferred to the Ascidae

Voigts & Oudemans, 1905 and the Laelaptonyssidae Womersley, 1956 206 sensu Lee, 1970 was transferred to the Laelaptonyssoidea. The prior placement of Protogamasellus Karg, 1962 and Gamasellodes

Athias-Henriot, 1961 in the Ascidae as was done by Lindquist and

Evans (1965), was corroborated in this study. Panteniphis Willmann,

1949 and Lindquistoseius Genis, Loots & Ryke, 1969 were found to be distinct genera as well as confamilial taxa and therefore, a new family group was erected for them. The Laelaptonyssoidea is regarded as consisting of three families: Laelaptonyssidae Womersley, 1956a,

Digamasellidae Evans, 1957 and Panteniphididae new family. Diagnoses and descriptions of the superfamilies, families, subfamilies and newly created taxa as well as a key to the families and subfamilies of Rhodacaroidea are given. 207

LIST OF REFERENCES

Antony, L, M. K. 1980. Systematic studies on the genus Ologamasus Berlese (Acari: Mesostigmata) in Southern Brazil. Unpublished M.Sc. thesis. The Ohio State University, Columbus.

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Johnston, D. E. 1968. An Atlas of Acari. Part I. The Families of Parasitiformes and . Acarology Laboratory, The Ohio State University, Columbus, p: 1-140.

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APPENDIX A. MATERIAL EXAMINED (LISTED ALPHABETICALLY BY REPOSITORY).

ACAROLOGY LABORATORY, THE OHIO STATE UNIVERSITY (ALOSU):

Afrodacarellus concavus; Male. TANZANIA. Mt. Bondwa, 2120 m. Under Philippia. 30 May 1972. H. Hurlbutt.

Afrodacarellus femoratus; Four males, two females. TANZANIA. Morogoro, 3600 ft. Tree in field, edge of clearing. 30 August 1967. H. Hurlbutt.

Afrodacarellus leleupi: Three females, one deutonymph. UGANDA. Katabane Swamp. Soil under Papyrus. Hurlbutt. Katebora Swamp. Soil under Papyrus. Patch of high forest. Hurlbutt.

Afrodacarellus lupangaensis; Three males, five females. TANZANIA. Above Morogoro, 3600 ft. Forest litter, tree in field. 30 Oct. 1966, 30 Aug. 1967, 9 Dec. 1967, 27 May 1972. H. Hurlbutt. Det. L. Antony.

Afrodacarellus machadoi; Twenty-five specimens. TANZANIA. Eight females, one deutonymph. Mt. Kilimanjaro, 2100-2250 m, forest litter, 9-10 June 1972. Three females. Arusha National Park, 1650 m, forest litter, 11 July 1966, 11 June 1972. Seven females, two deutonymphs. Lyamungu forest litter, 9 June 1972. Female. Rim of Ngorongoro crater, 7500 ft. litter, Jan. 3, 1971. Coll. H. Hurlbutt.

Afrodacarellus ? machadoi. Male. TANZANIA, Lyamungu forest. 19 ?. H. Hurlbutt. Det. L. Antony.

Afrodacarellus machadoi. Two females. UGANDA. Makerere Univ. Spruce forest litter, patches of high forest, 2 July 1975. H. Hurlbutt. Det. L. Antony.

Afrodacarellus minutus; Female. TANZANIA. Morogoro, 2900 ft. April 25, 1967. Coll. H. Hurlbutt.

Afrodacarellus mossi; Seven specimens. TANZANIA. Two females, two males. Uluguru Mts. Moss from trees. 30 May 1972. Female. Uluguru Mts, 2120 m. Moss on tree, 10 June 1968, Female. Uluguru Mts., 4700 ft. Forest litter. Feb. 29 1968. Female. Mt. Bondwa, 6850 ft. Leaf litter. May 1, 1968, H, Hurlbutt, Det, L, Antony,

Afrodacarellus msituni; Fifteen specimens, TANZANIA, Six males, three females. Above Morogoro, 3600 ft,, edge of clearing. Dec. 9, 1967, 27 May 1972. Male. Morogoro, 3000 ft., tree in field, June 12, 1966. Two males, one female. Morogoro, 3600 ft., tree in field, 30 Aug. 1967. Female, Morogoro, Eucalyptus litter, 24 May 1972, Male, Uluguru Mts., 2125 m, 30 May 1972. Coll. H. Hurlbutt. 214

Afrodacarellus ngorongoroensis; Five females. TANZANIA, Mt. Kilimanjaro, 2100 m. Forest litter. 10 June 1972. H. Hurlbutt.

Afrodacarellus novembus: Female. TANZANIA. Uluguru Mts., roots & soil. Nov. 4, 1967, Coll. H. Hurlbutt. Det. H. Hurlbutt.

Afrodacarellus novembus: Female. Uluguru Mts. Rain forest, 4700 ft. April 12. Coll. H. Hurlbutt. Det. L. Antony.

Afrodacarellus ? novembus; Female, Mt. Kilimanjaro, 2100 m. 10 June 1972. H. Hurlbutt. Det. L. Antony.

Afrodacarellus ? novembus; Female. Mt. Kilimanjaro, 2400 m. Moss & litter. 19 June 1972. H. Hurlbutt. Det. L. Antony

Afrodacarellus novembus; Deutonymph. Morogoro mud. 16 Sept. 1966. Coll. H. Hurlbutt. Det. L. Antony.

Afrodacarellus pili; Three specimens. TANZANIA. Two females. Uluguru Mts., 2120 m, moss. 30 May 1972. H. Hurlbutt. Female. Above Morogoro, edge of clearing. 27 May 1972.

Afrodacarellus pocsi; HOLOTYPE Male. TANZANIA. Summit of Mt. Bondwa, 2125 m. Moss on branch of tree. 30 May 1972. H. Hurlbutt.

Afrodacarellus pocsi; Male. Same locality and date of Holotype. H. Hurlbutt.

Afrodacarellus pocsi; Female. TANZANIA. Morogoro. Tree in field. Aug. 30 1967. H. Hurlbutt. Det. L. Antony.

Afrodacarellu pocsi; Male. TANZANIA. Uluguru Mts., 1680 m. Humus on camphor tree. 30 May 1972.

Afrodacarellus ? pocsi: Male. TANZANIA. Morogoro. Eucalyptus litter. 24 May 1972.

Afrodacarellus nr. ruwenzoriensis: Female. TANZANIA. Uluguru Mts. Forest. 30 May 1972. Coll. H. Hurlbutt. Det. H. Hurlbutt.

Afrogamasellus sp.: Ten females, six males. TANZANIA. Above Morogoro, 3600 ft. 27 May 1972. Uluguru Mts., 4700 ft. Rain forest. June 10 1967, Feb. 29 1968, May 30 1972. KENYA. Aberdares Natl. Park., 9000-11.500 ft. Under Cupressus.

Afrogamasellus sp.: Two females, two males. KENYA. Aberdares Natl. Park, . Under St. John Wart, 11.500 ft. 1965. 215

Afrogamasellus bakeri (= A. decembus) Hurlbutt: One male, one female. TANZANIA. Uluguru Mts., 4700 ft. Litter, roots. July 20 1968, 30 May 1972. H. Hurlbutt.

Afrogamasellus near citri: Female. TANZANIA. Morogoro. March 26 1968. H. Hurlbutt.

Afrogamasellus isthmus: One male, one female. TANZANIA. Above Morogoro, 3600 ft. Litter. (Feb. 21 1968, May 27 1972). Male. Mt. Bondwa, Uluguru Mts., 2120 m. Moss on tree. (30 May 1972). Male. Morogoro. Edge of clearing.(Aug. 30 1967). One deutonymph, two females. Near Morogoro river. Dead leaves. (October 29 1967, Nay 20 1972). Two females. Morogoro, 3600 ft. Tree in field. (Aug. 30 1967). H. Hurlbutt.

Afrogamasellus near isthmus; TANZANIA. Female. Lyamungu forest litter (9 June 1972). Female. Morogoro, 2900 ft. (April 26 1967). Female. Above Morogoro, edge of clearing. (27 May 1972). female. Mt. Kilimanjaro, 2100 m. 10 June 1972. H. Hurlbutt.

Afrogamasellus franzi: Eighteen males, nine females. TANZANIA. Mt. Kilimanjaro, 2100 m. Forest floor. (10 June 1972). Above Morogoro, 3500 ft. Soil & stems. (Jan. 23 1968). Ngorongoro forest, rim of crater, 7500 ft. (Jan. 3 19 ?). Above Morogoro, 3600 ft. Litter, soil. (Feb. 21 1968). Above Morogoro, 4400 ft. Forest litter. (23 Feb. 1967). Uluguru Mts., 4700 ft. Rain forest. April 12. Col. H. Hurlbutt.

Afrogamasellus franzoides: Female. TANZANIA. Uluguru Mts.^ 4700 ft. Twigs. Mar 6 1968. H. Hurlbutt.

Afrogamasellus lootsi: Eleven males, thirteen females, one deutonmyph, one larva. TANZANIA. Pugu Forest Reserve. (25 Nov. 1979, 5 Feb. 1980). Uluguru Mts. 4700 ft. (30 May 1972). Morogoro, 2900 ft. (April 26 1967). Morogoro. Leaves, twigs. (28 May 1972). Above Morogoro. forest. 4400 ft. (23 Feb. 1967). Near Morogoro river. Litter under bushes. (May 20 1972). Duff near Morogoro river. (July 18, 1968). Duff under tree near Morogoro river (27 Sept. 1967). Morogoro. Eucalyptus litter. (24 May 1972). Above Morogoro, 3600 ft. Litter. (Feb. 21 1968). Above Morogoro. Dead grass near stream. (18 Jan. 1967). Morogoro. Edge of clearing. (Aug. 30 1967). Coll. H. Hurlbutt.

Afrogamasellus lyamunguensis: Two females. UGANDA. Kasozi. Lake Nabugabo. Soil under Caraniuon thicket. (2 July 1965).coll. W. Block. Jubiya Forest. Sandy soil. #7. coll. W. Block.

Afrogamaasellus lyamunguensis: Fiye females, one deutonymph. TANZANIA. Lyamungu forest litter. 9 June 1972. Coll. H. Hurlbutt. Det. L. Antony. 216

Afrogamasellus maskamensis; One maie, two females. TANZANIA. Mt. Kilimanjaro, 2100 m. Under weeds, edge of Marangu track. 10 June 1972. H. Hurlbutt.

Afrogamasellus mongii; HOLOTYPE Male, three males, two females. TANZANIA. Mt. Kilimanjaro, 2100 ra. 10. June 1972.

Afrogamasellus mongii; TANZANIA. Mt. kilimanjaro, 2100 m. Forest floor. 10 June 1972. Coll. H. Hurlbutt. Det. L. Antony.

Afrogamasellus near mongii; One male, one female. UGANDA. N.W. Jinja. Forest litter. 1965. Coll. H. Hurlbutt. Det. Hurlbutt.

Afrogamasellus near meyersi; Female. TANZANIA. Rim of Ngorongoro Crater, 7300 ft. Litter, twigs. Jan. 3, 1968. Coll. H. Hurlbutt. Det. H. Hurlbutt.

Afrogamasellus paratruncatus; Two females. TANZANIA. Uluguru Mts., 1750 m. Rain forest, near stream. 23 March 1967, 30 May 1972. H. Hurlbutt.

Afrogamasellus near rugegensis; Male. UGANDA. Anthill thicket. H12.

Afrogamasellus succintus; Male. Mt. Kenya. Forest Station. H. Hurlbutt.

Afrogamasellus tetrastigma; Two males, two females. TANZANIA. Mt. Kilimanjaro, 2800 m. Moss. (19 june 1972). Ngorongoro Crater. Litter, 7500 ft. (Jan. 3, 1968). Mt. Kilimanjaro. Under weeds, edge of Marangu track, 2000 m. (10 June 1972). Coll. H. Hurlbutt. West Usambara, Mazumbei Forest. Dec. 1978. Coll. K. Howell (?).

Afrogamasellus near tetrastigma; Six adults. TANZANIA. Mt. Kilimanjaro, forest floor, 2100 m. (10 June 1972). Ngorongoro Forest, 5800 ft. (Jan. 4, 1968). Coll. H. Hurlbutt.

Afrogamasellus truncatus; Four males, five females. TANZANIA, Above Morogoro, 3600 ft. Thicket. (27 May 1972). Above Morogoro, 3600 ft. Tree in field, edge of clearing. (30 Aug. 1967). Above Morogoro. Clump of trees. (18 January 1967). Coll. H. Hurlbutt.

Afrogamasellus uluguruensis; Female. TANZANIA. Above Morogoro, 4400 ft. Forest. 23 Feb. 1967. Coll. H. Hurlbutt.

Dendrolaelaps fallax; Two females, ex. straw & manure. Field south of OSU woods. Franklin Co., Ohio. D.L. Wrensch, col.. 23.VII,73.

Dendrolaelaps fallax; Male. Dorsal. 22.IX.74-1.

Digamasellus perpusillus (Berl.); Female dorsal. 23.VII.73. 217

Dendroseius reticulatus (Sheals): Two females. SWEDEN, ex. soil core, grassland. Ecological Station, Stensoffa. D.L.W, D.E.J, L.L. col. 5.IX.1974-3.

Gamaselliphis nr. potchefstroomensis; Four females. TANZANIA. Ngorongoro Forest, 5800 ft. Jan. 4, 1968. Coll. H. Hurlbutt. One male, one female. Rim of Ngorongoro Crater, 8000 ft. Jan. 2, 1968. Coll. H. hurlbutt.

Gamasellodes bicolor (Berl.); Three females, 4.X,74-5. Two females, 1.IX.74. One female, one deutonymph, 15.VII.73.

Gamasellodes cf« major A.H. (= bicolor Berl.): Female. Ex Quercus litter. Bois de Vincennes, Paris, FRANCE. Johnston coll. lA-249.

Gamasellodes cf. major A.H. (= bicolor Berlese): Female. W. Berlin. Johnston coll. 253.

Gamasellus morogoroensis: Four females, one male. TANZANIA. Morogoro, 1330 m. (24 May 1972). Morogoro. Morningside leaf litter. (March 10 1968). H. Hurlbutt,

Gamasellus muscosus: HOLOTYPE. TANZANIA. Uluguru Mts., 2125 m. Moss. 30 May 1972. H. Hurlbutt.

Gamasellus muscosus: Thirteen specimens. TANZANIA. Uluguru Mts., 2125 m. (30 May 1972). Uluguru Mts., humus next to log. (July 29 1968). Uluguru Mts., 1680 m., humus on camphor tree. O O May 1972). Morogoro, 2900 ft. (April 26 1968). Moss on tree, 2120 m. (30 May 1972). H. Hurlbutt.

Gamasellus uluguruensis: Adults, deutonymphs, protonymphs. TANZANIA. Uluguru Mts., 4700-5000 ft., Morogoro. Soil under fern; Morningside leaf litter; tree hole; humus next to log; moss and humus; in leaf mould; log; forest litter. 11 Sept. 1966, 31 January 1967; Dec. 1 1967; March 10 1968; July 29 1968; 24 May 1972; 30 May 1972; April 1980. Coll. H. Hurlbutt. Det. H. Hurlbutt.

Lasioseius allii Chant 1958: Two females. Chrysanthemum flowers. Germany: at Philadelphia, Pa. Sept. 12, 1959. I.P. Boeshore, coir. Lot 59-23399.

Lasioseius bispinosus Evans 1958: Female, ex. Tulip bulbs. Japan: at New York. C. Tuthill. Sept. 3. 1964. 64-29972. D.E.J. det.

Lasioseius bispinosus: Seven females. Chrysanthemum flowers. Germany: at Philadelphia, Pa. 12.IX.1959. I.P.Boeshore.

Lasioseius bispinosus: Three females. Ex Narcissus bulbs. Netherlands: at New Orleans. 29.IX.1950. K.E. Miller. 218

Lasioseius fimetorum Karg: Five females. D.L, Wrensch. 21.X.1973. D.E.J. det.

Lasioseius tuberculiger (Berl.): Six females, three males. Ex Polypore. Barnebey Cntr. Fairfield Co., Ohio. 29.VI.72.

Longoseius elongatus: Deutonymph. on Monochamus notatus. on Pine. Orono, Maine. July 20,1959. R.S. Soper, dr., coir.

Ologamasus megadenoides n. sp.: Twenty-six males, fourteen females, sixty-five deutonymphs, eleven protonymphs, one larva. BRAZIL. Pine plantation at Tupi (14 km from center of town), Piracicaba, Sao Paulo, ex. grass piled deep. 2-II-78. Col: D.E. Johnston. Det: L. Antony. (AL 1005).

Ologamasus megadenoides n. sp.: Fourteen males, nine females, fourteen deutonymphs, five protonymphs, one larva. BRAZIL. Horto Florestal "Navarro de Andrade", 35 km from Piracicaba, Sao Paulo. Eucalyptus forest at Rio Claro, ex. bamboo litter. 3-II-78. Col: D.E. Johnston. Det: L. Antony. (AL 1006).

Ologamasus megadenoides n. sp.: Three males, five females, three deutonymphs, one protonymph. BRAZIL. "ESALQ", Piracicaba, Sao Paulo, ex piled leaf litter (Pinus. etc.), park. 21 March 1978. Col: D.E. Johnston. Det: L. Antony. (AL 1046).

Ologamasus megadenoides n. sp.: One male, three females, one deutonymph, two protonymphs. BRAZIL. "ESALQ", Piracicaba, Sao Paulo. Piled grass clippings, park. 24 March 1978. Col: D.E. Johnston. Det: L. Antony. (AL 1048).

Ologamasus membranosus: Three males, two females, one deutonymph. BRAZIL. "ESALQ", Piracicaba, Sao Paulo. In litter, humus, second growth natural forest near cow pasture, above tributary of river. 2-III-1978. Col: D.E. Johnston. Det: L. Antony. (AL 1038).

Ologamasus simplicior: One female. BRAZIL. "ESALQ", Piracicaba, Sao Paulo. In litter, humus, second growth natural forest near cow pasture, above tributary of river. 2-III-1978. Col: D.E. Johnston. Det: L. Antony. (AL 1038).

Ologamasus simplicior: Two females. BRAZIL. "ESALQ", Piracicaba, Sao Paulo. Piled grass clippings, park. 24 March 1978. Col: D.E. Johnston. Det: L. Antony. (AL 1048).

Ologamasus striolatosimilis: Four males, seven females, one protonymph, one larva. BRAZIL. Hotel Pouso do Nhambu, Estrada Rio-Santos, km 93. ex. litter, humus, moist primary forest. 16-11-78. Col: D.E. Johnston. Det: L. Antony. (AL 1019). 219

Ologamasus striolatosimilis; One maie, one female, one deutonymph. BRAZIL. "ESALO”, Piracicaba, Sao Paulo. In litter, humus, second growth natural forest near cow pasture, above tributary of river. 2-III-1978. Col: D.E. Johnston. Det: L. Antony. (AL 1038).

Ologamasus tuberculatum n. sp.: Fifteen males, fourteen females, eleven deutonymphs, two protonymphs, on larva. BRAZIL. Hotel Pouso do Nhambu, Estrada Rio-Santos, km 93. ex. litter, humus, moist primary forest. 16-11-78. Col: D.E. Johnston. Det: L. Antony.

Panteniphis (Lindquistoseius) africanus: Six males, two females. TANZANIA. Lyamungu forest litter, 9 June 1972d. Morogoro, thicket, edge of clearing, August 30, 1967. Morogoro, dead grass near stream, 18 Jan. 1967.

Proctolaelaps bicklevi: Forty-two females. 1981.12.

Proctolaelaps bicklevi: Female. Ex European corn borer colony-lab, U. of D. Newark, Delaware. 27.X.75. G. Curl & P.P. Burbutis.

Protogamasellus mica; Two females. TANZANIA. Morogoro. Under Lantana. Dec 16, 1968. Coll: Hurlbutt. Det: L. Antony.

Protogamasellus primitivus: Female #5. TANZANIA. Morogoro. Under Lantana. Dec. 16, 1968. Coll. Hurlbutt. Det. Hurlbutt.

Protogamasellus primitivus similis: Four females. TANZANIA. Duff under tree near Morogoro river. 27 Sept. 1967. Above Morogoro, 3600 ft, soil. Dec. 9, 1967, Feb. 21, 1968. Col: Hurlbutt. Det: Hurlbutt.

Rhodacarellus silesiacus Willm.: Female dorsal. SWEDEN, ex. soil core, grassland. Ecology Station, Stensoffa. D.L.W., D.E.J., L.L. col. 5.IX.74-3.

Rhodacaridae. (Gen. n. #1): Seven females. Control - ELF. Soil Core 6. (30 August, 1983, 29 September 1983). Soil Core 8. (14 September 1983). Soil Core 10. (20 October 1983). Soil Core 12 (29 September 1983). Michigan, Dickinson Co. T43 N, R30 W, Sec.II. Maple-basswood forest. CMCPP 9. 14.XII.1984. Det. L. Antony.

Rhodacaridae. (Gen. n. #1): Three females. Ex Tree hole. Crane Hollow, Hocking Co. Ohio. 26.IV.1975. Chandler.

Rhodacarus cf. coronatus Berl.: Female. SWEDEN, ex. open meadow. Linkoping. A. Edler, col. 12.VI.74-1

Rhodacarus denticulatus: Four females. Somers, Conn. May 25, June 23 and August 4, 1956. H. Hurlbutt. 220

Rhodacarus denticulatus; Four females, one deutonymph. TANZANIA, Under trees in field. 16 Oct. 1966. Litter under trees in field. 18 Jan, 1967. Above Morogoro, A1 layer, forest soil. 30 Oct. 1966. Col: H. Hurlbutt. Det: H. Hurlbutt.

Rhodacarus near denticulatus: Female. TANZANIA. Morogoro. Duff under bushes. Feb. 26 1966. Col. H. Hurlbutt. Det. H. Hurlbutt.

Rhodacarus ? denticulatus; Female. TANZANIA. Uluguru Mts., 4700 ft. Forest litter. Feb. 29 1968. Col. H. Hurlbutt. Det. L. Antony.

near Sessiluncus: Female. TANZANIA, Lushoto, Mazurabi, natural forest. Det. Hurlbutt.

near Sessiluncus: Female. TANZANIA. Morogoro. Dead leaves near Morogoro river. Oct. 22, 1967. Col. H. Hurlbutt. Det. H. Hurlbutt.

BERNICE PAUAHI BISHOP MUSEUM (BPBM):

Acugamasus watsoni. Female. CAMPBELL IS. Courejolles (sic) Penin. 230m. 13-11-1963. Ex. Low plants. Mollymawk Colony. Berlese funnel. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Female. CAMPBELL IS. Perseverence (sic) Harbor, Look out Bay. 3 Feb. 1963. Leaf mould or under Tussock. Berlese funnel. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Female. AUCKLAND IS. (N) Ocean I. Dec. 29,1962. J.L. Gressitt. ex. Stilbocarpa leaf mould.

Acugamasus watsoni. Female. AUCKLAND IS. Ranui I. Jan. 3 1963. J.L. Gressitt. ex. Metros leaf mould.

Acugamasus watsoni. Female. CAMPBELL IS. Perseverence Harbor Look out Bay. 3 Feb. 1963. Leaf mould under Tussock. Berlese funnel. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Male. CAMPBELL IS. Beeman Hill. 2-II-1963. Moss. Berlese funnel. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Male. CAMPBELL I. Courejolles Pen. 200 m. 14-XII-1961. Gray-headed Mollymawk (Nest). J.L. Gressitt. Bishop Museum.

Acugamasus watsoni. Male. AUCKLAND IS. Ranui I. Jan. 3, 1963. J.L. Gressitt. ex. Metros leaf mould. 221

Acugamasus watsoni. Male. AUCKLAND IS. Ranui Cove. 2-1963. Rata leae mould. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Male ventral. CAMPBELL I. St. Col Ridge. 18ü-28üra. 24.XI.1961. J.L. Gressit - coll. Berlese funnel collection - moss on rocks. Bishop Museum.

Athiasella pecten. PARATYPE Female. AUCKLAND IS. Ranui Cove. Jan. 1, 1963. K.A.J. Wise, ex Blackbird or Bellbird nest. Berlese Funnel.

Athiasella scaphosternum. PARATYPE Female. CAMPBELL IS. Beeman. 19-XII-1961. Dracophyllum approx. 5 m. K. Rennell. Bishop Museum.

Athiasella scaphosternum. PARATYPE Female. CAMPBELL IS. Summit Mt. Dumas 503 m. 6-II-1963. Moss. Berlese. Tussock. K.A.J. Wise. Bishop Museum.

Athiasella scaphosternum. PARATYPE Maie. CAMPBELL IS. N. Slopes Mt. Dumas 150 m. 5-II-1963. Nest of Diomedea epomorphora. K.A.J. Wise. Bishop Museum.

Athiasella scaphosternum. DN. CAMPBELL IS. Rocky Bay S. Coast Below Mt. Dumas. 8-II-1963. Ex Dracophyllum longifolium leaf mould. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Athiasella viripileus. PARATYPE Male. CAMPBELL IS. Perseverence Harbor Look out Bay. 3-II-1963. Leaf mould under Tussock. Berlese Funnel. K.A.J. Wise.Bishop Museum.

Athiasella viripileus. Female PARATYPE. CAMPBELL IS. Perseverence Harbor Look Out Bay. 3-II-1963. Leaf mould under Tussock. Berlese funnel. K.A.J. Wise. Bishop Museum.

Caliphis schusteri. Male. CAMPBELL IS. Smooth water Bay. 16-11-1963. Leaf mould under Tussock. Berlese funnel. K.A.J, Wise. Bishop Museum.

Caliphis schusteri. Male. CAMPBELL IS. Rocky Bay S. Coast Below Mt. Dumas. 18-11-1963. Ex Dracophyllum longifolium leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum.

Caliphis schusteri. Female. AUCKLAND IS. (N) Rose I. Jan. 8, 1963. J.L. Gressitt. ex Ground, leaf mould.

Caliphis schusteri. Female. AUCKLAND IS. (N) Rose I. Jan. 8, 1963. J.L. Gressitt. ex Ground, leaf mould.

Cymiphis watsoni. Female. AUCKLAND IS. Ranui I. Jan. 3, 1963. J.L. Gressitt. ex Metros leaf mould. 222

Cymiphis watsoni. DN. CAMPBELL IS. Perseverence Harbor S. Side. 24-11-1963. Liverwort. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Cymiphis watsoni. Female. CAMPBELL IS. Perseverance Harbor S. Side. Liverwort. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Euepicrius caesariatus. Female PARATYPE. AUCKLAND IS. Ranui I. Jan. 3, 1963. J.L. Gressitt. ex Metros leaf mould.

Euepicrius caesariatus. Male PARATYPE. AUCKLAND IS. Ranui I. Jan. 3, 1963. J.L. Gressitt. ex Metros leaf mould.

Euepicrius caesariatus. Female. CAMPBELL IS. Beeman Point. 28-11-1963. Sedge leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum.

Euepicrius caesariatus. Male. CAMPBELL IS. Beeman Pt. 28-11-1963. Pittosporum leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum.

Evansellus foliatus. Female D. AUCKLAND IS. (N) Ranui Cove. Dec. 26, 1962. J.L. Gressitt. ex Ground moss.

Gamasellus gressitti Hunter. PARATYPE Proto. SOUTH GEORGIA. Bird Island. North Valley. 19-IV-1963. H.B. Clagg. ex South Georgian Diving Petrel nest 10.

Gamasellus gressitti Hunter. PARATYPE Deuto. SOUTH GEORGIA. Bird Island. Fresh Water Bay. 7-IV-63. H.B. Clagg. ex. Dove Prion Nest 12.

Gamasellus gressitti Hunter. PARATYPE Male. SOUTH GEORGIA. Bird Island. Cave Crag. 30-IV-63. H.B. Clagg. ex Black browed Albatross nest.

Gamasellus gressitti Hunter. PARATYPE Female. SOUTH GEORGIA. Bird Island. Cave Crag. 30-IV-63. H.B. Clagg. ex Black-browed Albatross nest 3-4 inch depth.

Gamasellus gressitti Hunter. PARATYPE Female. SOUTH GEORGIA. Bird Island. Fresh Water Bay. 7—IV-63. H.B. Clagg. ex.Dove Prion Nest 12.

Gamasiphoides costai. Male PARATYPE. AUCKLAND IS. French I. 2-1-1963. White-headed Petrel. J.L. Gressitt. Bishop Museum.

Gamasiphoides macquariensis. Female. CAMPBELL IS. St. Col. Peak. 250m. 23-11-1963. Low plants on Rocks. Berlese funnel. K.A.J. Wise. Bishop Museum.

Gamasiphoides macquariensis. Male. AUCKLAND IS. (N) Rose I. Jan. 8, 1963. J.L. Gressitt. ex Ground, leaf mould. 223

Gamasiphoides macquariensis. DN, CAMPBELL IS. Moubray Hill. 12-XII-1961. J.L. Gressitt. Bishop Museum.

Hevdeniella sherrae. PARATYPE Maie. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa.

Hevdeniella sherrae. PARATYPE Maie. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa.

Hevdeniella sherrae. PARATYPE Maie. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa.

Hevdeniella sherrae. PARATYPE Maie D. AUCKLAND IS. Crozier Pt. 28-XII-1962. Sedge. J.L. Gressitt. Bishop Museum.

Hevdeniella sherrae. PARATYPE Female. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa.

Hevdeniella sherrae. Maie 6. CAMPBELL IS. North West Bay, 5m. middle Cove. 5-II-1963. Dracophvllum leaf mould. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. Male 5. CAMPBELL IS. Shoal Point. 7-II-1963. Tussock. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. Male 4. CAMPBELL IS. North West Bay, 5m. middle Cove. 5-II-1963. Dracophvllum leaf mould. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. DN 5. CAMPBELL IS. Tucker Cove. 1-II-1963. Dracophvllum leaf mould. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. DN 6. CAMPBELL IS. North West Bay, 5m. middle Cove. 5-II-1963. Dracophvllum leaf mould. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. PN 2. CAMPBELL IS. Below Summit, Mt. Honey ca. 550m. 24-11-63. Moss & Lichen on Rock. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella sherrae. DN 4. CAMPBELL IS. Beeman Hill. 2-II-1963. Moss. Berlese Funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella womerslevi. PARATYPE Male. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa.

Hevdeniella womerslevi PARATYPE Male. AUCKLAND IS. Rose I. Jan. 8, 1963. J.L. Gressitt. ex Poa. 224

Hevdeniella womerslevi. PARATYPE Female. AUCKLAND IS. Ranui I. . Jan. 3, 1963. J.L. Gressitt. ex Metros leaf mould.

Hevdeniella womerslevi. Female 3. CAMPBELL IS. Monumental Harbour. 9-II-63. Tussock Near Beach. K.P. Kennel. Bishop Museum.

Hevdeniella womerslevi. PARATYPE Female. AUCKLAND IS. Rose I. Jan. 8, 1967. J.L. Gressitt. ex Poa.

Litoeamasus falcipes. PARATYPE Male 2, V. AUCKLAND ISLAND, rocky shore. 1967. Col. Colin Suramerhayes.

Parasitiphis aurora. MALE Ventral. CAMPBELL ISLAND, rocky shore. Col: Colin Summerhayes. 1967.

Parasitiphis aurora. Female ventral. CAMPBELL ISLAND, rocky shore. Col: Colin Summerhayes. 1967.

Parasitiphis aurora. Female dorsal. CAMPBELL ISLAND, rocky shore. Col: Colin Summerhayes. 1967.

Parasitiphis aurora. DN. CAMPBELL IS. Venus Cove. 2, Feb. 1963. on stone near Low Beach. K.A.J. Wise. Bishop Museum.

Pilellus rueipellis. PARATYPE Female 2, V. CAMPBELL ISLAND. CRL 17. moss, lÜ-lOüm, Tucker Cove. 21.XI.1961. Col. J.L. Gressitt.

BRITISH MUSEUM OF NATURAL HISTORY (BMNH):

Afrogamasellus evansi HOLOTYPE Female. BM(NH) 1972.33. Habitat- Soil, Nyinabitaba. Lokaliteit- Ruwenzori, Uganda. Versaraelaar- G.Ü. Evans. Datum- 1952. Familie- Rhodacaridae. Ident. deur- Loots. Nommer- R31.

Afrogamasellus evansi. ALLOTYPE Male. BM(NH) 1972.34. Habitat- Soil, Nyinabitaba. Lokaliteit- Ruwenzori, Uganda. Versaraelaar- G.O. Evans. Datum- 1952. Familie- Rhodacaridae. Ident. deur- G.C. Loots. Noraraer- R21-24.

Afrogamasellus nvinabitabaensis. HOLOTYPE Female. BM(NH) 1972.28. Habitat- Soil, Nyinabitaba. Lokaliteit- Ruwenzori, Uganda. Versamelaar- G.O. Evans. Datum- 1952. Familie- Rhodacaridae. Ident. deur- G.C. Loots. Nommer- R21-24. 225

Afrogamasellus nvinabitabaensis. ALLOTYPE Male. BM(NH) 1972.29. Habitat- Soil, Nyinabitaba. Lokaliteit- Ruwenzori, Uganda. Versaraalaar- G.O. Evans. Datum- 1952. Familie- Rhodacaridae. Ident. deur- G.C. Loots. Nommer- R21-24.

Cvrtolaelaps (Gamasellus) rvkei Hunter. Female HOLOTYPE. 1967.10.3.4. SOUTH SANDWICH IS. Candlemas Is. 1964 ?. P. Tilbrook. ex. Polvtrichum alpinum mat.

Cvrtolaelaps (Gamasellus) rvkei Hunter. Male PARATYPE. 1967.10.3.11. SOUTH SANDWICH IS. Candlemas Is. ? 1964. P. Tilbrook. ex. Polvtrichum alpinum mat.

Gamasellus (Hvdrogamasellus) antarcticus (Trag.). Female. 1970.99. South Georgia. Bird Island. Banderspateh. 17.III.1963. H.B. Clagg. ex. Giant Petrel nest. Det; P.E.H.

Gamasellus (Hvdrogamasellus) antarcticus. Maie. 1970.98. South Georgia. Bird Island. Fresh water Bay. 18.IV.'63. H.B. Clagg. ex. Dove Prion Nest.

Geogamasus coxalis (ex Gamasellus) Sheals. HOLOTYPE Female V. 1961.6.20.25. Det: J.G. Sheals, 19. 142/1.

Geogamasus coxalis (ex Gamasellus) Sheals. PARATYPE Female V. 1961.6.20.26. Det: J.G. Sheals, 19. 142/1.

Geogamasus howardi. PARATYPE Female. 1970.12 (2). AUSTRALIA. Plant litter & soil. Pinus forest, Mt. Burr, South Australia, 30.5.1966. Col. G.W. Howard.

Geogamasus howardi. PARATYPE Male. 1970.12 (1). AUSTRALIA. Plant litter & soil. Pinus forest, Mt. Burr, South Australia, 30.5.1966. Col. G.W. Howard.

Geogamasus skoshi. HOLOTYPE Female. 1970.199. ARGENTINA. University Grounds. Tucuman. 11.1957. Col; P. Wygodzynsky. Bol. 4. Det: D.C. Lee.

Geogamasus skoshi. ALLOTYPE Male. 1970.200. ARGENTINA. University Grounds. Tucuman, 11.1957. Col; P. Wygodzynsky. Bol. 4. Det; D.C. Lee.

Geogamasus skoshi. Male Lateral. 1970.201. ARGENTINA. University Grounds. Tucuman. 11.1957. Col; P. Wygodzgnsky. Bol. 4. Det: D.C. Lee.

Hevdeniella goei. HOLOTYPE Female. 1970.13. ENGLAND. Mould, pit props. Chalk mine, Chislehurst caves, Kent. 20.2.1955. Coll. P.N. Lawrence. 226

Hevdeniella goei. Male (minus chelicerae). ALLOTYPE. 1970.14. ENGLAND. Mould, pit props. Chalk mine, Chislehurst, caves, Kent. 20.2.1955. Col. P.N. Lawrence.

Hvdrogamasellus sp. Male. ARGENTINA. (136). Moss, Parc du Larain, Province du Neuquen. 20.3.1959. Col. C. Delaware Debouteville.

Hvdrogamasellus sp. Two deutonymphs. ARGENTINA. (164). Col. C. Delaware Debouteville.

Hvdrogamasellus cavei. PARATYPE Female (pt.). idiosoma & legs. 1961.6.20.22. ARGENTINA. Litter under Nothophagus dombevi. Nahuel Huapi Res., Puerto Blest. 7.3.1959. Col. C. Del Debouteville.

Hvdrogamasellus cavei. Male (pt.). gnathosoma & legs II. Locality data as for Paratype female.

Hvdrogamasellus cavei. Male (pt.). idiosoma. Locality data as for Paratype female.

Hvdrogamasus delamarei Sheals. Female HOLOTYPE. 39/2. 1961.6.20.16. Det: J.G. Sheals, 1961/1.10.

Hvdrogamasus delamarei Sheals. ALLOTYPE Male. 39/2. Male fig. 1961.6.20.17. Det: J.G. Sheals, 1961/1/10.

Hvdrogamasus delamarei Sheals. Male PARATYPE dissected. 39/2. 1961.6.20.18. Det: J.G. Sheals, 19.

Hvdrogamasus kensleri Luxton. PARATYPE. 1966.10.18.4. Rock fissures on upper shore. Takatu Peninsula near Matakana. North Auckland. NEW ZEALAND. Coll: Luxton. Dec. 1964.

Hvdrogamasus kensleri Luxton. PARATYPE. 1966.10.18.1. Rock fissures in upper shore. Takatu Peninsula near Matakana. North Auckland. NEW ZEALAND. Col: Luxton. Dec. '64.

Hvdrogamasus gamasiphioides Sheals. HOLOTYPE Female 68. 1961.6.20.3. Det: J.G. Sheals, 19.

Hvdrogamasus striatus Sheals. Nil. PARATYPE. 1961.6.20.6. Det: J.G. Sheals, 19. 33/1. Figd.

Laelaogamasus sp. Female. MALAYA. Forest soil. Ulu Ranching Templer Park, Selangor. 27.7.1962 c. Col. Nadchatzem. May 4. Female coxa & trochanter IV + spermathecal access, duct. Det: D.C. Lee.

Ologamasus cvmosus Lee. 1966. PARATYPE Female. 1966.9.81. Leaf mould. Botanical Gdns., Wellington, NEW ZEALAND. Coll. D.C. Manson. 12. 1960. 227

Ologamasus simplicior. Female ventral. ARGENTINA 5. Humus & rotting wood, 9 de Julio Park. Tucuman. 1953. Col. P. Wygodzynsky.

Parasitiphis aurora. Female ventral.Gough Is. assoc. Pertorgunia sp. Col. Dr. Hollgate. RM 39, by Lee (SAM), 1966. Det. D.C. Lee.

Rhodacarus costai. HOLOTYPE Female. 162. det. J.G. Sheals. 1961.6.20.2. ARGENTINA. Surface soil, Los Arrayanes, Nahuel Huapi Res., Andes Mts. 2-5.1959. Col. C. Delamare Debouteville.

Sessiluncus sp. Female. NEPAL. Litter in oak forest, 8000 ft. Sikha. 24-26.5.1954. Col. K.H. Hyatt.

CANADIAN NATIONAL COLLECTION (CNC):

Caliphis sp. Female. AUSTRALIA. , Toorongo River, 1 km. S. Toorongo Rd. 10.11.80. Det. L. Antony.

Ologamasus membranosus. Three females, two males, four deutonymphs. ARGENTINA. El Rosedal, Buenos Aires. V.1960.. Coll. 0. de Ferrariis. Hojarasca. Det. L. Antony.

Ologamasus striolatosimilis. Female. BRAZIL. Nova Teutonia, 27 degr. 11'S., 52 degr. 2pW^ Brazil, 300-5G0ra. XI. 1971. Coll. Fritz Plauraann. Det. L. Antony.

Panteniphis mirandus. Two females, two deutonymphs. 5mi. N. Writing on Stone Provincial Park, Alberta, 12.VIII.78. Coll. E.E. Lindquist, sedges and substrate by marsh.

CZECHOSLOVAKIAN ACADEMY OF SCIENCES (CAS);

Puchihlungia chinensis Samsinak. PARATYPUS Female. Det. Samsinak. CHINA. Coptotermes formosanus. 848. Col. Samsinak.

Puchihlungia chinensis Samsinak. PARATYPUS Male. Det. Samsinak. CHINA. Coptotermes formosanus. 825. Col. Samsinak.

Puchihlungia chinensis Samsinak. PARATYPUS PN. Det.Samsinak. CHINA. Coptotermes formosanus. 944. Col. Samsinak. 228

FIELD MUSEUM OF NATURAL HISTORY (FMNH);

Epiphis rarior. Male dissected. Det. J. Kethley. 81-134.

Epiphis rarior. Two pharate males. U.S.A. 111., Cook Co., Hills, 12-III-1981. F14NH # 81-128a, ex raked oak leaves on ridge top. Coll. L. Watrous. Det. J. Kethley .

Epiphis rarior. Female. Gnathosoma. 81-157.

Epiphis rarior. Female. Dissected. 81-157.

INSTITUT FUR PFUNZENSCHUTZFORSCHUNG, KLEINMACHNOW (KARG):

Afrogamasellus furculatus Karg, 1979. Female TYPE. ARGENTINIEN. Rio Negro, El Bolson, Mt. Piltriquitron. 1000m Gras + Erde unter Baumstamm. 16.5.61.

Gamasiphoides linealis Karg, 1976. Female TYPE. CHILE. Provinz Tarapaca bei Caquena, in feuchtem Boden mit Lamadung. 1965.

Gamasiphoides linealis Karg, 1976. Male, B. II, chel. CHILE. Provinz Tarapaca bei Caquena, in feuchtem Boden mit Lamadung. 1965.

Gamasiphoides longocuspis Karg, 1976. Female TYPE. CHILE. Provinz Tarapaca, Sopahuira, in 3100m Hohe in einer Bodenfalle. 1965.

Geogamasus filicuspidis Karg. 1976. Male TYPE. Chel., B. II. CHILE. Azapa (Prov. Tarapaca). Nematodenprobe v. Blattern unter Weidenbussch. 23.11.65.

Geogamasus foliaceus Karg. 1976. Male TYPE. Chel., B. II. gez. CHILE. Umgebung von La Union (Prov. Valdivia). Regenwurmer aus feucht. Waldi. tiefem Tal. 26.10.65.

Geogamasus furcatius Karg, 1976. Male TYPE. B. II, Chel. gez. CHILE. Concon (Prov. Valparaiso). Feuchtes Material aus d. Umgebung Quintero. 10.10.65.

Geogamasus pentaspinosus Karg, 1979. Female TYPE. ARGENTINIEN. Rio Negro, El Bolson. Mt. Piltriquitron, 720m. Erde von Vichweide. 24.4.61. 229

Geogamasus pentaspinosus Karg, 1979. Maie. B. II, Chel. ARGENTINIEN. Rio Negro, El Bolson. Mt. Piltriquitron, 720m. Erde von Vichweide. 24.4.61.

Geogamasus tuberosus Karg, 1979. Female Chel. CHILE. Umgebung von Azapa (Prov. Tarapaca). Bodenprobe am Bach. 2.12.65.

Geogamasus tuberosus Karg. 1979. Maie Hypost. gez., Chel., B. II gez. CHILE. Umgebung von Azapa (Prov. Tarapaca). Bodenprobe am Bach. 2.12.65.

Hevdeniella distorta Karg, 1976. Female TYPE. CHILE. Environs of La Union (Prov. Valdivia), Strasse n. Valdivia. 26.10.65.

Hevdeniella distorta Karg, 1976. Maie. Chel. gez., B. II. CHILE. Environs of La Union (Prov. Valdivia), Strasse n. Valdivia. 2b.10.65.

Hvdrogamasellus brevipilus Karg. 1976. Maie TYPE. B. II, Chel. CHILE. Umgebung von La Union (Prov.Valdivia). Gesammeltes Material. 26.10.65.

Hvdrogamasellus microcrinis Karg. 1979. Female TYPE. ARGENTINIEN. Rio Negro, El Bolson. Mt. Piltriquitron, Laubstreu nahe Fluss. 6.10.1961.

Ologamasus foliatus Karg, 1976. MALE. B. II, Chel. CHILE. Bofedal de Bacuyo. Berlese-Proben von Algen am See. 26.11.65.

Ologamasus foliatus Karg, 1976. Female TYPE. CHILE. Bofedal de Bacuyo. Berlese-Proben von Algen am See. 26.11.65.

Rhodacarellus kreuzi Karg, 1965. Female. NSG Kragenriss westl. Worlitz Verraoderungsschicht unter Laub (Hainbuche). 13.4.67.

Rhodacarus minimus (Karg, 1961). Female TYPE. STAHNSDORF b. Berlin. Grasansaat. 25.11.57.

ISTITUTO SPERIMENTALE PER LA ZOOLOGIA AGRARIA (ISZAF);

Ologamasellus aberrans. Female cotipo Berl. 215/28. Austro-America. (balzan).

Ologamasellus aberrans. Female, male tipico Berl. 9/16.

Ologamasellus simplicior. Maie, female tipico Berl. 165/45. La Plata, bruck.

Ologamasellus striolatus Berl. Maie 215/27. bello. La Plata. (Bruch!). 230

Ologamasellus striolatus. Male, female tipico Berl. 170/16, La Plata. (Bruch).

Ologamasellus striolatus Berl. Maie, female. 178/49. La Plata. Bruch.

Ologamasellus striolatus. Maie 170/17. rotto. Berl. La Plata. (Bruch).

MUSEE ROYAL DE L'AFRIQUE CENTRALE (MRAC):

Afrogamasellus celisi Loots. HOLOTYPE. Det: G.C. Loots, 1968. Loc: Congo. Lubero, Mulo 1880m. Rec: R.P. Celis, 3/7/1957. R. G. Mus. Afr. Centr. 99.559/99.563.

Afrogamasellus kahusiensis. Loots. HOLOTYPE. Det: G.C. Loots, 1968. Loc: Congo. Kalehe, N.O. de Kahisi. Rec: N. Leleup, 29/6/1951. R. G. Mus. Afr. Centr. 118.829.

Afrogamasellus luberoensis kalibuensis Loots. HOLOTYPE. Det; G.C. Loots, 1968. Loc: Congo. Ruwenzori, ruiss Kaliba, 23/1/1954. Rec: RR.PP. Celis, Collart et Massaux. R. G. Mus. Afr. Centr. 77.051/77.055.

Afrogamaellus luberoensis luberoensis Loots. PARATYPE. Det: G.C. Loots, 1968. Loc: Congo. Kivu, Lubero, riv. Kakolwe. Rec: R.P. Bergmans, 21/12/1953. R. G. Mus. Afr. Centr. 78.713/78.732.

Afrogamasellus muhiensis Loots. HOLOTYPE. Det: G.C. Loots, 1968. Loc: Congo. Kivu, Mt. Muhi, VII/1955. Rec: N. Leleup. R. G. Mus. Afr. Centr. 93.489.

Afrogamasellus rugegensis Loots. PARATYPE. Det: G.C. Loots, 1968. Loc: Ruanda. Rugege, V/l951. Rec: N. Leleup. R. G. Mus. Afr. Centr. 134.592.

Cvrtolaelaps (Gamasellus) kivuensis Ryke et Loots. Det: J. Ryke, 1965. Loc: Congo. Kivu. Kabore, Nyakasiba, 2350m. Rec: N. Leleup, 27/12/1950. R. G. Mus. Afr. Centr. 127.859.

Cvrtolaelaps (Gamasellus) leleupi. Ryke et Loots. Det; J. Ryke, 1965. Loc: Congo. Kivu. Kabore, Nyakasiba, 2350m. Rec: N. Leleup, 27/12/1950. R. G. Mus. Afr. centr. 118.863.

Cvrtolaelaps (Gamasellus) uviraensis congoensis Ryke et Loots. Det: J. Ryke, 1965. Loc: Congo. Kivu, Terv. Uvira, Mt. Kahololo, 2800m. Rec: N. Leleup, 1-1960. R. G. Mus. Afr. Centr. 127.860. 231

Cvrtolaelaps (Gamasellus) uviraensis uviraensis Ryke et Loots. Det: Ryke, 1965. Loc: Congo. Kivu, Terv. Uvira. Rec: N. Leleup, XI-1959. R. G. Mus. Afr. Centr. 127.861.

MUSEUM NATIONAL D'HISTOIRE NATURELLE (MNHN):

Allogamasellus aquafortensis Athias. SYNTYPE Female. lC-15. L/A 370. Registre 802.

Allogamasellus aquafortensis Athias. SYNTYPE Female, Male. lC-16. L/A 370. R. 802.

Allogamasellus squalidus Athias. HOLOTYPE Female. lC-14. L/B 86. Registre 802.

Rhodacarellus francescae Athias. HOLOTYPE Female. lE-3. L/B 636. Registre 802.

Rhodacarellus mica Athias. HOLOTYPE Female. lE-2. L/B 636. Registre 802. Museum National d'Histoire Naturelle. Paris. Laboratoire de Zoologie - Arthropodes Acariens.

Rhodacaropsis angustiventris Athias. HOLOTYPE Female. lD-7. L/B 636. N. registre 802. Museum National d'Histoire Naturelle. Paris. Laboratoire de Zoologie - Arthropodes Acariens.

Rhodacaropsis arcanus Athias. HOLOTYPE Female. lD-9. L/B 636. R. 802.

Rhodacaropsis cognatus Athias. HOLOTYPE Female. lD-10. L/ 249. N. registre 802. Museum National d'Histoire Naturelle Paris. Laboratoire de Zoologie - Arthropodes Acariens.

Rhodacaropsis massula Athias. SYNTYPES Females, lD-8. L/A 134. R. 802.

Rhodacarus clavulatus Athias. HOLOTYPE Female. lD-15. L/C 836. N. du registre 802.

Rhodacarus cuneatus Athias. SYNTYPES Females. lD-16. L/A 353. R. 802.

Rhodacarus laureti Athias. SYNTYPES Females. lD-14. L/B 636. Registre n.802.

Rhodacarus reconditus Athias. SYNTYPE Female. lD-12. sp 410 C. R: 802. 232

Rhodacarus reconditus Athias. SYNTYPE Maie. lD-13. sp 410 C. r: 802.

Rhodacarus tribaculatus Athias. HOLOTYPE Female. lD-11. L/ 276. N. registre 802.

NEW ZEALAND ARTHROPOD COLLECTION (DSIR):

Acugamasus watsoni. Female. CAMPBELL IS. Filhol Peak Summit. 9-II-63. Tussock & moss. Berlese. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Male. CAMPBELL IS. Northwest Bay, 5m Middle Cove. 5-II-1963. Moss on Dracophyllum. Berlese funnel. K.A.J. Wise. Bishop Museum.

Acugamasus watsoni. Male. CAMPBELL IS. Tucker Cove. O-lOOm. 26-11-63. Pittosporum leaf mould. K.A.J. Wise. Bishop Museum.

Athiasella longiseta (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - idiosoma etc. CAMPBELL ISLAND. CRL 18. Moss & lichen on rocks. Courrejolles Peninsula. Col: K.A.J. Wise. 13.2.1963.

Athiasella longiseta (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - legs III & IV. CAMPBELL ISLAND. CRL 18. Moss & lichen on rocks. Courrejolles Peninsula. Col: K.A.J. Wise. 13.2.1963.

Athiasella longiseta (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - gnathos. CAMPBELL ISLAND. CRL 18. Moss & lichen on rocks. Courrejolles Peninsula. Col: K.A.J. Wise. 13.2.1963.

Athiasella pecten (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - idios. etc. CAMPBELL ISLAND. CRL 19. Leaf mould of Dracophvllum. Tucker Cove. Col: K.A.J. Wise. 1.1.1963.

Athiasella pecten (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - gnathos. CAMPBELL ISLAND. CRL 19. Leaf mould of Dracophvllum. Tucker Cove. Col: K.A.J. Wise. 1.1.1963.

Athiasella pecten (ex Hevdeniella). HOLOTYPE (part,1/3). Female 1 - legs III & IV. CAMPBELL ISLAND. CRL 19. Leaf mould of Dracophvllum. Tucker Cove. Col: K.A.J. Wise. 1.1.1963.

Athiasella scaphosternum. PARATYPE Female. CAMPBELL IS. Moobray Hill. 16-11-1963. Low plants near Summit. Berlese funnel. K.A.J. Wise. Bishop Museum.

Athiasella scaphosternum. PARATYPE Male. CAMPBELL IS. Beeman Pt. 28-11-1963. Pittosporum leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum. 233

Athiasella viripileus. PARATYPE 1. Female. CAMPBELL IS. By Beeman Beach. 19-XII-1961. Azorella etc. K. Rennell. Bishop Museum.

Athiasella viripileus. PARATYPE 3. Maie V. CAMPBELL I. Courrejolles Penin. 200 ra. J.L. Gressitt. Azorella. Bishop Museum.

Caliphis schusteri. Female. AUCKLAND 13. (N) Rose I. Jan 8, 1963. J.L. Gressitt. ex. ground, leaf mould.

Caliphis schusteri. Male. AUCKLAND IS. (N) Rose I. Jan 8, 1963. J.L. Gressitt. ex. ground, leaf mould.

Cymiphis watsoni (ex Athiasella scaphosternum). Female. CAMPBELL IS. Perseverence (sic) Harbor S. side. 24.11.1963. Berlese funnel. Liverwort. K.A.J. Wise. Bishop Museum.

Cymiphis watsoni. Female. CAMPBELL IS. Perseverence Harbor S. side. 24.11.1963. Liverwort. Berlese funnel. K.A.J. Wise. Bishop Museum.

Euepicrius caesariatus. Female. CAMPBELL IS. North West Bay, 5m. middle cove. 5.II.1963. Dracophyllum leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum.

Euepicrius caesariatus. Male. CAMPBELL IS. Shoal Point. 7.II.1963. Tussock. Berlese funnel. K.A.J. Wise. Bishop Museum.

Evanssellus foliatus. 3 Female D. AUCKLAND IS. (N) Ranui Cove. Dec. 26, 1962. J.L. Gressitt. ex. ground moss.

Evanssellus foliatus. 2 Female D (one Leg I missing). AUCKLAND IS. (N) Ranui Cove. Dec. 26, 1962. J.L. Gressitt. ex. ground moss.

Gamasiphoides costai. HOLOTYPE (1/2 part), idiosoma. Female. AUCKLAND IS. RL 26. Ocean Is. Stilbocarpa leaf mould. Dec. 29, 1962. J.L. Gressitt.

Gamasiphoides costai. HOLOTYPE (1/2 part). Gnathosoma. Female. AUCKLAND IS. RL 26. Ocean Is. Stilbocarpa leaf mould. Dec. 29, 1962. J.L. Gressitt.

Gamasiphoides costai. ALLOTYPE Male. AUCKLAND IS. (N) Ocean I. Dec. 29, 1962. K.A.J. Wise. ex.under log.

Gamasiphoides macquariensis. Male. CAMPBELL IS. Above Tucker Cove. 28-11-1963. Lichens on Dracophyllum scoparium. Berlese funnel. K.A.J. Wise. Bishop Museum.

Gamasiphoides macquariensis. Female. CAMPBELL IS. Smooth water Bay. 16-11-1963. Dracphyllum leaf mould. Berlese funnel. K.A.J. Wise. Bishop Museum. 234

Hevdeniella sherrae. Male, venter of idi + est. CAMPBELL ISLAND. CRL/4. nest of Diomedea epomorphora. 150m. N. slopes of Mt.Dumas. Col: K.A.J. Wise. 5.2.1963.

Hevdeniella sherrae. Male. CAMPBELL IS. Below Summit Mt.Honey ca. 550m. 24-11-1963. Moss & lichens on rocks. Berlese funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella womerslevi. Female. CAMPBELL IS. Shoal Point. 7-II-1963. Tussock. Berlese funnel. K.A.J. Wise. Bishop Museum.

Hevdeniella womerslevi. PARATYPE Male. Auckland Is. Ewing Is. Jan 5, 1963. J.L. Gressitt. ex.shap nest & nest area.

Hvdrogamasus kensleri Luxton. PARATYPE Female. Crevices on upper shore. , Bay of Islands, N.Z. 3.11.65. Coll: C.B. Kender.

Hvdrogamasus kensleri Luxton. PARATYPE Male. Crevices on upper shore. Russel, Bay of Islands, N.Z. 3.11.65. Coll: C.B. Kender.

Litogamasus falcipes. HOLOTYPE Female 1. CAMPBELL ISLAND, rocky shore. 1967. Col: Colin Summerhayes.

Litogamasus falcipes. ALLOTYPE Male 1 - chelicerae. CAMPBELL ISLAND. CRL 14. Under rock. High Tide M., Lookout Bay Beach. 16-19.XII.1961. Col: J.L. Gressitt.

Litogamasus falcipes. ALLOTYPE. Male 1 - idiosoma. CAMPBELL ISLAND. CRL 14. Under rock. High Tide M., Lookout Bay Beach. 16-19.XII.1961. Col: J.L. Gressitt.

Litogamasus falcipes. ALLOTYPE Male 1 - some legs & part of gnath. CAMPBELL ISLAND. CRL 14. Under rock. High Tide M., Lookout Bay Beach. 16-19.XII.1961. Col: J.L. Gressitt.

Litogamasus setosus. Female 1 - gnathos. CAMPBELL ISLAND. CRL 13. On stone, beach near Low Water Mark, Venus Cove. 1.2.1963. Col; K.A.J. Wise.

Litogamasus setosus. Female 1 (part), idiosoma. CAMPBELL ISLAND. CRL 13. On stone, beach near Low Water Mark. Venus Cove. 1.2.1963. Col: K.A.J. Wise.

Parasitiphis ieanneli. 1 Female, 2 DNN. AUCKLAND ISLAND. Rocky shore. Col: Colin Summerhages. 1967.

Pilellus rugipellis. HOLOTYPE, 1/2 Female 1 (part), gnath. AUCKLAND ISLAND. CRL 15. ex-moss Dicromolona billardieri. Hooker Hills. 11th Feb. 1963. Col: J.A.K. Wise. 235

Pilellus rugipellis. HOLOTYPE, 1/2 Female 1 (part), idios. AUCKLAND ISLAND. CRL 15. ex-moss Dicromolona billardieri. Hooker Hills, llth Feb. 1963. Col; J.A.K. Wise.

Pilellus rugipellis. ALLOTYPE, 1/3 Male 1 (part), idiosoma. AUCKLAND ISLAND. CRL 16. ex-moss Dicromolona billardieri. Hooker Hills, llth Feb. 1963. Col: J.A.K. Wise.

Pilellus rugipellis. ALLOTYPE, 1/3 Male 1 (part), gnathos. AUCKLAND ISLAND. CRL 16. ex-moss Dicromolona billardieri. Hooker Hills, llth Feb. 1963. Col: J.A.K. Wise.

Pilellus rugipellis. ALLOTYPE, 1/3 Male 1 (part), legs II. AUCKLAND ISLAND. CRL 16. ex-moss Dicromolona billardieri. Hooker Hills, llth Feb. 1963. Col: J.A.K. Wise.

QUEENSLAND INSTITUTE OF MEDICAL RESEARCH (QIMR):

Gamasiphis (Laelaptiella) mackerrasae n.sp. PARATYPE MALE, leaf mould. Low Is. 19.8.54. E. N. Marks, coll. R. Domrow det.

Gamasiphis (Laelaptiella) mackerrasae n.sp. MALE chelicera. leaf mould .Low Is. 14.8.54. E. N. Marks, coll. R. Domrow det.

Gamasiphis (Laelaptiella) mackerrasae n.sp. FEMALE chelicera. leaf mould "freem Auck. Is.". Low Is., 24.8.54. E. N. Marks, coll. R. Domrow det.

QUEENSLAND MUSEUM (QM):

Gamasiphis (Heteroiphis) australicus Womersley. Female. Leaf mould. Low Is. 8/54. M.J. Mackerras. R. Domrow det. G 2335.

Gamasiphis (Laelaptiella) mackerrasae n. sp. HOLOTYPE Female. Leaf mould on cay. Low Is. 19.8.54. E.N. Marks coll. R. Domrow det. G 2339.

Gamasiphis (Laelaptiella) mackerrasae n. sp. PARATYPE Male. Leaf mould on cay. Low Is. 19.8.54. E.N. Marks coll. R. Domrow det. G 2343. 236

Rhodacarus marksae n. sp. HOLOTYPE Female. Leaf mould. Ant Is., Low Is. 24.8.54. E.N. Marks coll. det. R. Domrow. G 2332.

SOUTH AUSTRALIAN MUSEUM (SAM);

Acugamasus elachyaspis. Female v. NÎ970345. Paratype. ARA73. SOUTH AUSTRALIA. LF53 Moss under Callitris preissi. Grange Golflinks. 23.5.1965. Col. M. Fagg.

Acugamasus elachyaspis. Male. N1970344. ALLOTYPE. ARA73. SOUTH AUSTRALIA. LF53 Moss under Callitris preissi. Grange Golflinks. 23.5.1965. Col. M. Fagg.

Acugamasus punctatus. N1970341. ADELAIDE, S.A. 6/35. H. W. Det. H. Womersley. Rhodacaridae. HOLOTYPE Female. Digamasellus (=Cyrtolaelaps) punctatus sp.n. Det. H. Womersley.

Acugamasus punctatus. N1970342. National Park, S.A. 8/38 H. W. Det. H. Womersley. Rhodacaridae. ALLOTYPE Male. Digamasellus (=Cyrtolaelaps) punctatus n.sp. Det. H. Womersley.

Acugamasus semipunctatus. N1970349. ARA73. In moss. Muston, K. J. 23.8.43. H.M.C. Det. H. Womersley. ALLOTYPE Male. Digamasellus (=Cyrtolaelaps) semipunctatus Wom. Det. H. Womersley.

Acugamasus semipunctatus. N1970348. In moss. Bridgewater, S.A. 8/42. Det. H. Womersley. HOLOTYPE Female. Digamasellus (=Cyrtolaelaps) semipunctatus Wom. Det. H. Womersley.

Antennolaelaps affinis. Male N1968195. PARATYPE. S. AUS. MUS. I. PARATYPE Male. Antennolaelaps affinis g. et sp. n. In soil debris. Brookfield, Qu. 31.5/10.6.1949. E.H. Derrick. Det. H. Womersley.

Antennolaelaps aremenae Female V. N1970381. PARATYPE. ARA73. SOUTH AUSTRALIA. LF 233. Moss nr. summit, Mt. Lofty. Cleland Reserve. 7.6.1968. col. D.C. Lee.

Antennolaelaps aremenae Male. N1970380. ALLOTYPE. ARA73. SOUTH AUSTRALIA. LF 233. Moss nr. summit, Mt. Lofty. Cleland Reserve. 18.7.58. col. D.C. Lee.

Antennolaelaps celox Male. N1970387. ALLOTYPE. ARA73. SOUTH AUSTRALIA. LF 233/9M. Moss nr. summit, Mt. Lofty. Cleland Reserve. 4.7.1968. col. D.C. Lee.

Antennolaelaps celox. Female frag. N1970388. PARATYPE. ARA73. SOUTH AUSTRALIA. LF 233/6M. Moss nr.summit, Mt. Lofty. Clealand Reserve. 21.6.1968. col. D.C. Lee. 237

Antennolaelaps convexa. Female N1968199. HOLOTYPE. S.AUS,MUS.I. HOLOTYPE Female V. Antennolaelaps convexa g. et sp. n. In soil litter. Brookfield, Qu. 31.5/10.6.1949. E.H. Derrick. Det. H. Womersley.

Antennolaelaps convexa. Male N1968200 (part), idiosoma. ALLOTYPE. S. AUS. MUS. I. ALLOTYPE Male V. Stvloaamasus convexa g. et sp. n. In soil litter. Brookfield, Qu. 31.5/10.6.1949. E.H. Derrick. Det. H. Womersley.

Antennolaelaps convexa. Male N1968200 (part), chelicerae & legs II. S. AUS. I'lUS. I. ALLOTYPE Male, gnathosoma & legs II. Stvloaamasus convexa g. et. sp. n. In soil litter. Brookfield, Qu. 31.5/10.6.1949. E.H. Derrick. Det. H. Womersley.

Antennolaelaps testudo. Female N1968201. HOLOTYPE. Queensland. LF 171. Moss. Brookfield nr. Brisbane. 7.9.1966. col. D.C. Lee.

Antennolaelaps testudo. Male N1968202 (part). ALLOTYPE. Queensland. LF 171. Moss. Brookfield nr. Brisbane. 7.9.1966. col. D.C. Lee.

Antennolaelaps testudo. Male N1968202 (part), gnathos. & legs II. ALLOTYPE. Queensland. LF 171. Moss. Brookfield nr. Brisbane. 7.9.1966. col. D.C. Lee.

Athiasella relata. Female HOLOTYPE. N1970177. Hvdrogamasus relatus sp. n. Female HOLOTYPE. In moss. Glen Osmond, S. A. 7/34. Det. H. Womersley.

Athiasella relata. Male ALLOTYPE. N1970178. Hvrogamasus relatus sp. n. In moss. Mt Osmond, S. A. o/34. Det. H. Womersley.

Athiasella relicta. NEOPARASITIDAE. Hvdrogamasus relictus sp. n. HOLOTYPE Female. In moss. Brisbane, Qu. Oct./34. Det. H. Womersley.

Athiasella relicta. NEOPARASITIDAE. Hvdrogamasus relictus sp. n. ALLOTYPE Male. In moss. Brisbane, Q. Oct./34. Det. H. Womersley.

Athiasella relicta var major. ARA7314. NEOPARASITIDAE, Hvdrogamasus australicus. Female. Sassafras, Vic. Dec/31 H.C.A. Det. H. Womersley.

Caliphis calvus. Female HOLOTYPE. N1968231. gnathos. Female PARATYPE. N1968233. idios. TASMANIA. LF 191. Moss. Mt. Wellington, nr. Hobart. 14.12.1966. Col: L.W. Miller.

Caliphis calvus. Female N1968231. idios. HOLOTYPE. Female N1968233. idios. PARATYPE. TASMANIA. LF 191. Moss. Mt. Wellington, nr. Hobart. 14.12.1966. Col: L.W. Miller. 238

Caljphis calvus. Male N1968232. (part), idlosoma. ALLOTYPE. TASMANIA. LF 191. Moss. Mt. Wellington, nr. Hobart. 14.12.1966. Col; L.W. Miller.

Caliphis calvus. Male N1968232 (pt). gnathos. & legs II. ALLOTYPE. TASMANIA. LF 191. Moss. Mt. Wellington, nr. Hobart. 14.12.1966. Col: L.W. Miller.

Caliphis ?queenslandicus. Male D. Litter. Brookfield Rd., Brookfield, Queensland. 5.4.61. LF 19. Coll: E.H. Derrick.

Cymiphis cvmosus. Female HOLOTYPE V. N19661 RL21. det. D.C. Lee. NEW ZEALAND. Leaf mould. Botanical Gardens, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis cvmosus. Male ALLOTYPE. N19662 RL21. det. D.C. Lee. NEW ZEALAND. Leaf mould. Botanical Gardens, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis dumosus. HOLOTYPE (part). Female chelicerae. N196621. Det. D.C. Lee. NEW ZEALAND. RL31. Leaf mould. Botanical Garden, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis dumosus. Female V HOLOTYPE (part). N196621. det. D.C. Lee. NEW ZEALAND. RL31. Leaf mould. Botanical Gardens, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis leptosceles. Female V HOLOTYPE. N196620. det. D.C. Lee. NEW ZEALAND. RL30. Leaf mould. Khandallah, Wellington. 19.12.61. coll. D.C.M. Manson.

Cymiphis mansoni. Female HOLOTYPE. N196615 RL12. det. D.C. Lee. NEW ZEALAND. Moss. Botanical Gardens Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis mansoni. Male ALLOTYPE. N196616 RL21. det. D.C. Lee. NEW ZEALAND. Leaf mould. Botanical Gardens, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis nucilis. Female V. HOLOTYPE. N196622 RL21. det. D.C. Lee. NEW ZEALAND. Leaf mould. Botanical Gardens, Wellington. 12.1960. col. D.C.M. Manson.

Cymiphis validus. Male D. HOLOTYPE. N196624 RL14. det. D.C. Lee. NEW ZEALAND. Waimamuka. 21.10.1938. col. E.D. Pritchard.

Cyrtolaelaps cooperi n.sp. HOLOTYPE Female. In moss. Huston, K. I. 23.8.43. H.M.C. Rhodacaridae. Det. H. Womersley.

Cyrtolaelaps cooperi n.sp. ALLOTYPE Male. In moss. Huston, K. I. 23.8.43. H.M.C. Det: H. Womersley. Rhodacaridae. 239

Cyrtolaelaps mucronatus. Female. N196841. idlosoma & 2 legs a. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. Coll: R.M. Emberson.

Cyrtolaelaps mucronatus. Female. N196841. gnathosoraa c. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. col: R.M. Emberson.

Cyrtolaelaps mucronatus. Female. N196841. chelicerae & 2 legs b. SCOTLAND. Mole*s nest. Roxburgh, East Lothian. 1964. col: R.M. Emberson.

Cyrtolaelaps mucronatus. Male N196842. idlosoma & 7 legs a. SCOTLAND. Mole's nest. Roxburgh, East Lotnian. 1964. col: R.M. Emberson.

Cyrtolaelaps mucronatus. Male N196842. gnathosoma, leg I & chelicerae b. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. col: R.M. Emberson.

Euepicrius filamentosus. HOLOTYPE Female. Dorsal. N1970130. desig - D.C. Lee, 1971. S. AUS. MUS. I. Euepicrius filamentosus g. et sp. n. Female. In moss. Glen Osmond, S. A. 4.83. Det. H. Womersley.

Euepicrius filamentosus. ALLOTYPE Male dorsal. N1970131. desig - D.C. Lee. S. AUS. MUS. I. Euepicrius filamentosus g. et sp. n. Male V. In moss. Glen Osmond, S. A. 3.6.83. Det. H. Womersley.

Euepicrius lootsi. HOLOTYPE Female. N1968138 (part). Idiosoma. Victoria. LF 127. Moss & grass, Wannon River, Yarrara Gap. 14.5.1966. Col: D.C. Lee.

Euepicrius lootsi. HOLOTYPE Female N1968138 (part). Gnathosoraa. Victoria. LF 127. Moss & grass, Wannon River, Yarram Gap. 14.5.1966. Col: D.C. Lee.

Euepicrius lootsi. ALLOTYPE Male N1968139 (part). Idiosoma. Victoria. LF 127. Moss & grass, Wannon River, Yarram Gap. 14.5.19b6. Col: D.C. Lee.

Euepicrius lootsi. ALLOTYPE Male N1968139 (part). Legs & gnathosoma. Victoria. LF 127. Moss & grass, Wannon River, Yarram Gap. 14.5.19b6. Col: D.C. Lee.

Euepicrius lootsi. ALLOTYPE Male N1968139 (part). chelicerae. Victoria. LF 127. Moss & grass, Wannon River, Yarram Gap. 14.5.1966. col: D.C. Lee.

Euepicrius queenslandicus sp. n. HOLOTYPE Female V. 5. In soil litter, Brookfield, (ju. 21st May-2nd July, 1949. E.H. Derrick. Det: H. Womersley. 240

Euepicrius queenslandicus sp. n. PARATYPE Maie 4. In soil litter, Brookfield, l^u. 21st May-2nd July, 1949. E.H, Derrick. Det: H. Womersley.

Eurvparasitus emarginatus. Female N196876. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. col: R.M. Emberson.

Eurvparasitus emarginatus. Male N196877. gnathosoraa & legs b. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. col. R.M. Emberson.

Eurvparasitus emarginatus. Male N196877. idiosoma a. SCOTLAND. Mole's nest. Roxburgh, East Lothian. 1964. col: R.M. Emberson.

Evansellus medusa. Female HOLOTYPE (part), gnathosoma. N1968297. det: D.C. Lee. AUSTRALIA. Moss & leaf litter, nr. Hordemvale, Cape Otway, Victoria. 28.8.65. col. F.S. Mitchell.

Evansellus medusa. Female HOLOTYPE (part), idiosoma. N1968297. det: D.C. Lee. AUSTRALIA. Moss & leaf litter, nr. Hordemvale, Cape Otway, Victoria. 28-8-65. col. F.S. Mitchell.

Evansellus medusa. Male ALLOTYPE (part), gnathosoma. N1968298. det: D.C. Lee. AUSTRALIA. Moss & leaf litter, nr. Hordemvale, Cape Otway, Victoria. 28.8.65. col. F.J. Mitchell.

Gamasellus concinnus (Worn.). Female HOLOTYPE. N1970242. SOUTH AUSTRALIA. In moss, RL 103. Long Gully, Belair National Park. 8.1938. J.S. Womersley.

Gamasellus concinnus (Wom.). Male ALLOTYPE. N1970243. SOUTH AUSTRALIA. In moss, RL 104. Long Gully, Belair National Park. 8.1938. J.S. Womersley.

Gamasellus cophinus. Female ventral. HOLOTYPE. N1970281. SOUTH AUSTRALIA, ex-moss LF233/7, nr.summit, Mt.Lofty Cleland Reserve. 7.6.1968. col: D.C. Lee.

Gamasellus cophinus. Male ventral. ALLOTYPE. N1970282. SOUTH AUSTRALIA, ex-moss. LF233/7. nr.summit, Mt.Lofty Cleland Reserve. 7.6.1968. col: D.C. Lee.

Gamasellus cursor elongatus. Female N196851. D HOLOTYPE. Victoria. LF99. moss. Beauchamp Falls, Otway Ranges. 9.12.1965. Col: D.C. Lee.

Gamasellus cursor elongatus. Male ALLOTYPE (part). N196852. Idiosoma, V. Victoria. LF99. moss. Beauchamp Falls, Otway Ranges. 9.12.1965. col: D.C. Lee. 241

Gamasellus cursor elongatus. Male ALLOTYPE. N196852. Gnathosoraa & leg II. Victoria. LF99. moss. Beauchamp Falls, Otway Ranges. 9.12.1965. col; D.C. Lee.

Gamasellus (discutatus) discutatus. Female HOLOTYPE. N196655 V.F. Det: D.C. Lee. AUSTRALIA. Leaf litter nr. Sardine Creek. Victoria. 23.11.1959. Col: G.F. Gross. RL 8.

Gamasellus (discutatus) discutatus. Male ALLOTYPE. N196656 V.F. Det: D.C. Lee. AUSTRALIA. Leaf litter nr. Sardine Creek. Victoria. 23.11.1959. Col: G.F. Gross. RL 8.

Gamasellus grossi. Female V. HOLOTYPE. N1970307. Det: D.C. Lee, 1971. SOUTH AUSTRALIA. LF59. Moss under Callitris. Grange nr. Adelaide. 10 June 1965. Coll: D.C. Lee.

Gamasellus grossi. Male V. ALLOTYPE, squashed. N1970308. Det: D.C. Lee, 1971. SOUTH AUSTRALIA. LF59. Moss under Callitris. Grange nr. Adelaide. 10 June 1965. Coll: D.C. Lee.

Gamasellus (discutatus) litoprothrix. Female HOLOTYPE. N196685 D. Det: D.C. Lee. AUSTRALIA. Leaf litter & moss. Turtons Track, Otway Ranges, Victoria. 18.1.1962. Col: G.F. Gross & P.F. Aitken. RL 24.

Gamasellus (discutatus) southcotti. Female HOLOTYPE. v. N196690. Det: D.C. Lee. AUSTRALIA. LF 72. Plant litter. Gogerley Point, N.S.W. 7.8.1965. Col: R.V. Southcott.

Gamasellus (discutatus) tasmanicus. Female HOLOTYPE. Det: H. Womersley. AUSTRALIA. Moss. Mt. Wellington, Tasmania. 27.7.1943. Col: V.V. Hickman. RL 80.

Gamasellus (discutatus) tasmanicus. Male ALLOTYPE. Det: H. Womersley. AUSTRALIA. Moss. Mt. Wellington, Tasmania. 27.7.1943. Col: V.V. Hickman. RL 81.

Gamasellus (discutatus) tindalei. Female HOLOTYPE. N196688 V. Det: D.C. Lee. AUSTRALIA. Moss & plant litter. Mt. Ginini. A.C.T. LF 3. 17.10.1964. Col: N.B. Tindale.

Gamasellus (discutatus) tindalei. Male ALLOTYPE (part). V. idiosoma & legs. N196689. Det; D.C. Lee. AUSTRALIA. Moss & plant litter. Mt. Ginini. A.C.T. LF 3. 17.10.1964. Col: N.B. Tindale.

Gamasellus (discutatus) tindalei. Male ALLOTYPE (part), gnathosoma & legs. N196689. Det: D.C. Lee. AUSTRALIA. Moss & plant litter. Mt. Ginini. A.C.T. LF 3. 17.10.1964. Col: N.B. Tindale.

Gamasellus tragardhi. HOLOTYPE Female. N196849. SOUTH AUSTRALIA. RL63. Moss. Adelaide. 6.1935. col: H. Womersley. 242

Gamasellus tragardhi. ALLOTYPE Male. N196850. SOUTH AUSTRALIA. RL 64. Moss. Bridgewater. 8.1942. col: J.S. Womersley.

Gamasellus (discutatus) virgosus. Female HOLOTYPE V. N196692. Det: D.C. Lee. AUSTRALIA. LF 72. Plant litter. Gogerley Point, N.S.W. 7.8.1968. Col: R.V. Southcott.

Gamasiphis australicus. Female HOLOTYPE. N196891. S. AUS. MUS. I. HOLOTYPE Female. Gamasiphis (Heteroiphis) australicus sp. n. et subg. n. From moss. Mylov, S. A. 27th June, 1948. G.F. Gross. Det. H. Womersley.

Gamasiphis australicus. N196889. Male (in part), gnathos. (cont. chel.). N196890. Male (in part), gnathos + leg II. AUSTRALIA. LF 58. Grass & leaf litter. Heywood Park, Hyde Park, S. A. 6.6.1965. Col. R.V. Southcott.

Gamasiphis australicus. Male idiosoma. N196889 (in part). AUSTRALIA. LF 58. Grass & leaf litter. Heywood Park, Hyde Park, S. A. 6.6.1965. Col. R.V. Southcott.

Gamasiphis (Heteroiphis) australicus. QLD. INST.OF MEDICAL RESEARCH. Gamasiphis (Heteroiphis) australicus Wom. Male. ? setosus. Leaf mould on cay. Low Is. 16.5.54. E.N. Marks coll. R.Domrow det.

Gamasiphis fornicatus. N196892 (part). HOLOTYPE Female, idiosoma. SOUTH AUSTRALIA. LF 142. Moss. Mt. Remarkable, nr. Melrose. Col: H.M. Cooper. 9.8.1966.

Gamasiphis fornicatus. HOLOTYPE Female, (part). N196892. gnathos. SOUTH AUSTRALIA. LF 142. Moss. Mt. Remarkable, nr, Melrose. 9.8.1966. Col: H.M. Cooper.

Gamasiphis fornicatus. ALLOTYPE Male. N196893 (part). Male ventral. SOUTH AUSTRALIA. LF 142. Moss. Mt. Remarkable, nr. Melrose. 9.8.1966. Col: H.M. Cooper.

Gamasiphis fornicatus. ALLOTYPE Male. N196893 (part). gnathosoma+ legs II & stern. SOUTH AUSTRALIA. LF 142. Moss. Mt. Remarkable, nr. Melrose. 9,8.1966. Col: H.M. Cooper.

Gamasiphis (Neogamasiphis) hickmani sp. n. HOLOTYPE. Mt. Wellington, Tasmania. 3.000 ft. 27th July, 1943. V.V.H. Det. H. Womersley.

Gamasiphis (Neogamasiphis) hickmani sp. n. ALLOTYPE Male. Mt. Wellington Tasmania. 3.000 ft. 27th July, 1943. V.V.H.. Det. H. Womersley.

Gamasiphis (Neogamasiphis) hickmani sp. n. ALLOTYPE Male (mandibles). Mt. Wellington, Tas. 3.000 ft. 27th July, 1943. V.V.H. Det. H. Womersley. 243

Gamasiphis (Neogamasiphis) hickmani sp. n. var.tambourinensis nov. G 278. coll. on card. Mt. Tambourine, (ju. 14th April, 1952. E.H. Derrick. Det. H, Womersley.

Gamasiphis (Neogamasiphis.) hickmani sp. n. var.tambourinensis nov. G 278. coll. on card. Mt. Tambourine, Qu. 14th April, 1952. E.H. Derrick. Det. H. Womersley.

Gamasiphis lenifornicatus. HOLOTYPE (part). N197097. Female ventral-idiosoraa. SOUTH AUSTRALIA. LF 233. Plant litter nr. Summit, Mt. Lofty, Cleland Reserve. 24.4.1969. Col: D.C. Lee.

Gamasiphis lenifornicatus. HOLOTYPE (part). N197097. Female gnathosoma. SOUTH AUSTRALIA. LF 233. Plant litter nr. Summit, Mt. Lofty, Cleland Reserve. 24.4,1969. Col: D.C. Lee.

Gamasiphis lenifornicatus. ALLOTYPE (part). N197098. Maie ventral, legs, sternum and gnathos. SOUTH AUSTRALIA. LF 233. Plant litter nr. Summit, Mt. Lofty, Cleland Reserve. 24.4.1969. Col: D.C. Lee.

Gamasiphis lenifornicatus. ALLOTYPE (part). N197098. Maie ventral-idios. SOUTH AUSTRALIA. LF 233. Plant litter nr. Summit, Mt. Lofty, Cleland Reserve. 24.4.1969. Col: D.C. Lee.

Gamasiphis (Neogamasiphis) novae-zealandae (sic) sp. n. HOLOTYPE Haie. Mandibles. Beachlands, N. Z. (Nth. Isl.). March, 1949. Det. H. Womersley.

Gamasiphis (Neogamasiphis) novae-zealandae sp.n. HOLOTYPE Maie minus mandibles. Beachlands, N. Z. (Nth. Isl.). March, 1949. Det. H. Womersley.

Gamasiphis (Neogamasiphis) novae-zealandae sp. n. ALLOTYPE Female. Beachlands, N. Z. (Nth. Isl.). March, 1949. Det. H. Womersley.

Gamasiphis (Neogamasiphis) queenslandicus sp. n. HOLOTYPE Maie. Taringa, N. Qu. 29-31st Jan, 1949. E.H. Derrick. Det. H. Womersley.

Gamasiphis (Neogamasiphis) queenslandicus sp. n. ALLOTYPE Female. Taringa, N. Ou. 29-31st Jan, 1949. E.H. Derrick. Det. H. Womersley.

Gamasiphis saccus. HOLOTYPE Female v. N1970104. SOUTH AUSTRALIA. LF 233. Moss. Waterfall Gully, nr. Adelaide. 21.6.1968. Coll: U.C. Lee.

Gamasiphis ? setosus. Maie, Female. Leaf litter, corner Haven & Upper Brookfield Rd. Brookfield, OLd. 22.7.60. Det. H Womersley.

Gamasiphoides. Male. N1968283. in female metast-endop IV. Victoria. LF 106. Nothophagus litter, nr. Ferguson, Otway Ranges. 10.12.1965. D.C. Lee. 244

Gamasiphoides. Female. N1968282. metast-endop IV. Victoria. LF 106. Nothophagus litter, nr. Ferguson, Ütway Ranges. 10.12.1965. D.C. Lee.

Gamasiphoides aitkeni. Female N1968155. ALLOTYPE. Victoria. LF 36. Moss, etc. nr.Glenelg River, Hundreds of Caroline. 28.1.1965. Col: P. Aitken & H.B. Tindale.

Gamasiphoides propinquus. HOLOTYPE Female. N1970140. SOUTH AUSTRALIA. RL 101. ex moss. Long Gully, Belair National Pk. Sept, 1938. col: H. Womersley.

Gamasiphoides propinquus. Maie N1968152. SOUTH AUSTRALIA. LF 16. wood & lichen. Mt. Lofty nr. Adelaide. 20.11.1964. Col: G.F. Gross.

Gamasitus obscurus. Female N1968159. HOLOTYPE. TASMANIA. RL 52. moss, Mt. Wellington. 2.12.1934. col: V.V. Hickmamm.

Geogamasus minimus. N1970218. HOLOTYPE. Female Dorsal. SOUTH AUSTRALIA: LF233. Moss. Waterfall Gully nr. Adelaide. 1968 or 1969. nr. Mt.Lofty summit. Coll: D.C. Lee.

Geogamasus minimus. ALLOTYPE. N1970219. Male Ventral. SOUTH AUSTRALIA: LF233. Moss. Waterfall Gully, nr. Adelaide. 1968 or 1969. nr. Mt. Lofty summit. Coll: D.C. Lee.

Heterogamasus calcarellus. Female PARATYPE. ventral. N1968296. Det. D.C. Lee. Tierra del Fuego. Nothofagus litter. Cordillera . 22.2.1962. col. E. Shipton.

Heterogamasus (Evanssellus) medusa. Male ALLOTYPE (part), dorsal idiosoma. N19o8298. AUSTRALIA, moss & leaf litter, nr. Hordemvale, Cape Otway, Victoria. 28.8.1965. col. F.J. Mitchell.

Heydeniella (dentatus) dentatus. Female N1968117. HOLOTYPE. Hydrogamasus dentatus sp. n. Female HOLOTYPE. In moss. Long Gully, S. A. Aug/38. Det. H. Womersley.

Heydeniella (dentatus) dentatus. Male N1968118. ALLOTYPE. SOUTH AUSTRALIA. RL68. Moss. Long Gully, nr. Adelaide. 8.1938. col: H. Womersley.

Heydeniella (dentatus) markmitchelli. Female N1968123. HOLOTYPE. SOUTH AUSTRALIA. 5 miles south west of Wilmington. 2000*. Flinders Ranges. 25.9.1958. RL71. Col: H.M. Cooper.

Heydeniella (dentatus) markmitchelli. Male N1968127. ALLOTYPE. SOUTH AUSTRALIA. RL72. Moss, Hummock Ranges 1064'. 9.9.1958. Col. H.M. Cooper. 245

Heydeniella (Hydrogamasellus) racovitzai. Female N1963132, SOUTH ORKNEYS. Under shore stones, Michelsen Island, 3.1.1933. col: Discovery Investigations (British).

Heydeniella (Hydrogamasellus) racovitzai. Male N1968135. SOUTH ORKNEYS. Under shore stones. Michelson Island. 3.1.1933. col. Discovery Investigations (British).

Hiniphis bipala. HOLOTYPE. N1970358 (part). Female gnathosoma. SOUTH AUSTRALIA. LF 233. Waterfall Gully, nr. Adelaide. Core sample B262-Litter, Mt. Lofty Summit Reserve. Coll: D.C. Lee. 5.8.1968.

Hiniphis bipala. HOLOTYPE. N1970358 (part). Female idio. V. SOUTH AUSTRALIA. LF 233. Waterfall Gully, nr. Adelaide. Core sample B262-Litter, Mt. Lofty Summit Reserve. Coll: D.C. Lee. 5.8.1968.

Hiniphis bipala. ALLOTYPE. N1970359 (part). Male gnathosoma & leg I. SOUTH AUSTRALIA. LF 233. Waterfall Gully, nr. Adelaide. Core sample B262-Litter, Mt. Lofty Summit Reserve. Coll: D.C. Lee. 5.8.1968.

Hiniphis bipala. ALLOTYPE. N1970359 (part). Male idiosoma V. SOUTH AUSTRALIA. LF 233. Waterfall Gully, nr. Adelaide. Core sample B262-Litter, Mt. Lofty Summit Reserve. Coll: D.C. Lee. 5.8.1968.

Hiniphis hinnus. Female N1968222 (part), gnathosoma. HOLOTYPE. Victoria. LF1Ü4. Moss, Pinus plantation, Otways. 9.12.1965. col. D.C. Lee.

Hiniphis hinnus. Female N1968222. (part), idiosoma. HOLOTYPE. Victoria. LF104. Moss, Pinus plantation, Otways. 9.12.1965. col. D.C. Lee.

Hiniphis hinnus. Male N1968223 (part), gnathosoma. ALLOTYPE. Victoria. LF104. Moss, Pinus plantation, Otways. 9.12.1965. col. D.C. Lee.

Hiniphis hinnus. Male N1968223 (part), idiosoma. ALLOTYPE. Victoria. LF104. Moss, Pinus plantation, Otways. 9.12.1965. col. D.C. Lee.

Hydrogamasellus relicta. p.T.O. In leaf mould. Bourkes Bush. Nairaamaka. N221. 10.35. E.D.P. Det. H. Womersley. NEOPARASITIDAE.

Hydrogamasus australicus sp. n. HOLOTYPE Female. NEOPARASITIDAE. Det. H. Womersley. In moss, Brisbane, Q. Oct./34.

Hydrogamasus australicus sp. n. ALLOTYPE Male. Det.H.Womersley. NEOPARASITIDAE. In moss, Brisbane, Q. Chelicerae & mandibles. Oct./34. 246

Hydrogamasus australicus sp. n. ALLOTYPE Male, Det. H, Womersley. NEÜPAKASITIDAE. In moss, Brisbane, Q. Legs chelicerae & mandibles, Oct./34.

Hydrogamasus australicus. Female. In leaf mould. Bourkes Bush, Naimamaka, N. Z. 21/10/35. E.D.P. Det. H. Womersley. NEOPARASITIDAE. P.T.O.

Laelaptiella anomala. Female PARATYPE. N1968279. SOUTH AUSTRALIA. RL 83. Adelaide. 6.1935. Col. H. Womersley.

Laelaptiella anomala. Male idiosoma. N196850 (part). Victoria. LF 30. Small shrub litter nr. Apsley. 17.12.1968. Col: D.C. Lee.

Laelaptiella anomala. Male gnathos & leg II. N196850 (part). Victoria. RL 30. Small shrub litter nr. Apsley. 17.12.1968. Col: D.C. Lee.

Laelaptonyssus chinensis Samsinak,1964 (ex Puchihlungia). PARATYPE Male N1968269. CHINA. Off Coptotermes formosanus. Shiraki, near Koa-ho, Kanton Province. 1963. Col: K. Samsinak.

Laelaptonyssus mitis Womersley, 1956. HOLOTYPE Male N1968288. ALLOTYPE Female N1968284. Det: H. Womersley. AUSTRALIA. From fly culture. Zoology Department, University of Western Australia. 10.1950. Col; E.P. Hodgkin.

Neogamasellevans berlesei (ex Heydeniella). Female gnathosoraa. N11968107 (part). HOLOTYPE. Queensland. Soil litter, Brookfield. 31.5/10.6.1949. col. E.H. Derrick. RL67.

Neogamaselleyans berlesei (ex Heydeniella). Female idiosoma. N1968107 (part). HOLOTYPE. Queensland. Soil litter, Brookfield. 31.5/10.6.1949. col. E.H. Derrick. RL66.

Ologamasus simplicior. fragmented Male. N1968256. Drawn Lee (1970). ARGENTINA 5. Humus, etc. 9 de Julio Park, Tucuman City, 1953. P. Wygodzynsky.

Ologamasus simplicior. fragmented Female. N1968255. Drawn Lee (1970). ARGENTINA 5. Humus, etc. 9 de Julio Park, Tucuman City, 1953. P. Wygodzynsky.

Onchogamasus communis. Female. N1968185. HOLOTYPE. S. AUS. MUS. I. Onchogamasus communis g. et. sp. n. HOLOTYPE Female. In soil litter. Brookfield, S. A. 21.5/2.6.1949. E.H. Derrick. Det. H. Womersley.

Onchogamasus pumilio. Female N1968186 (part), idiosoma. HOLOTYPE. SOUTH AUSTRALIA. LF 97. Moss. Mallee, Formby Bay, Yorke Peninsula. 3.11.1965. col: N.B. Tindale & N.Mc. Farland. 247

Onchogamasus pumilio. Female N1968186 (part), palp. HOLOTYPE. SOUTH AUSTRALIA. LF 97. Moss. Mallee, Formby Bay, Yorke Peninsula. 3.11.1965. col: N.B. Tindale & N.Mc. Farland.

Onchogamasus quasicurtipilus. Female N1968187 (part), idiosoma. HOLOTYPE. Victoria. LF 79. Moss & litter. Ottway Ranges. 28.8.1968. col. F.J. Mitchell.

Onchogamasus quasicurtipilus. Female N1968187 (part), gnathosoma & leg I. HOLOTYPE. Victoria. RL 79. Moss H litter, Otway Ranges. 28.8.1968. col. F.J. Mitchell.

Onchogamasus virguncula. N1970396 (part). HOLOTYPE. Female gnathosoma. SOUTH AUSTRALIA. LF 233. B67. Litter nr. Mt. Lofty summit. Waterfall Gully nr. Adelaide, Summit Reserve. 5.8.1968. Coll: B.C. Lee.

Onchogamasus virguncula. HOLOTYPE. N1970396 (part). Female idiosoma. SOUTH AUSTRALIA. LF 233. B67. Litter nr. Mt. Lofty summit. Waterfall Gully nr. Adelaide, Summit Reserve. 5.8.1968. Coll: D.C. Lee.

Parasitiphis aurora. HOLOTYPE. Female v.f. N1968182. Between tide marks, Macquarie Is. 28.10.1913. Coll: T.H.J., A.A.Exp. RM 38, by Lee, 1966.

Parasitiphis ieanneli. Male N1968171. Heard Is. RL 34. Vegetation. Roger's Head. 31.1.1952. col. K. Brown. A.N.A.R.E.

Parasitiphis ieanneli. (Andre, 1947). det: D.C. Lee. N1968176. Hydrogamasus watsoni Female. Macquarie Island. Isthmus Pucinellia. Watson 1961.

Parasitiphis littoralis. Female N1968160. HOLOTYPE. SOUTH AUSTRALIA. RL32. Littoral zone. Mouth of American River, Kangaroo Island. 1.1946. col. H. Womersley.

Parasitiphis littoralis. Male N1968161. ALLOTYPE. SOUTH AUSTRALIA. RL33. Littoral zone. Mouth of American River, Kangaroo Island. 1.1946. col. H. Womersley.

Periseius hammeni. DN N1968214. PARATYPE. Amongst Cladophora socialis and red algae, on stones in intertidal zone, nr. base of R. Neth. N. Det: H. Womersley. Rhodacaridae.

Periseius hammeni. Female N1968212 (part), gnathosoma & sternum. PARATYPE. NEW GUINEA. RL 46. Intertidal. Biak Island. 10.12.1953. Coll: L. v.D. Hammen. 248

Periseius hammeni. Male N1968218 (part), gnathosoma. PARATYPE. NEW GUINEA. RL 47. Intertidal. 6iak Island. 10.12.1953. Col; L. v.D. Hammen.

Periseius hammeni. Male N1968213 (part), idiosoma. PARATYPE. NEW GUINEA. RL47. Intertidal. Biak Island. lU.12.1953. col. L. v.D. Hammen.

Periseius hammeni. Female N1968202 (part), idiosoma (excl. st.). PARATYPE. NEW GUINEA. RL4b. Intertidal. Biak Island. 10.12.1953. col. L. v.D. Hammen.

Periseius (Psammonsella) schusteri. Females, Males. N1968258-268. Nea Krini, Saloniki. (Flachstrand). Schotter sand. Schuster 1904. Griechenland. Rhodacaridae.

Pvriphis pvrenoides. Female HOLOTYPE V. N196625. det. D.C. Lee. AUSTRALIA. RL 28. Beach litter. Beauchamp Falls, Otway Ra., Victoria. 19.1.1962. col. G.F. Gross & P.F, Aitken.

Queenslandolaelaps vitzthumi. Female N1968I91 (complete). HOLOTYPE. S. AUS. MUS. I. Queenslandolaelaps vitzthumi g. et. sp. n. In soil litter. Brookfield, Qui 31.5/10.6.1949. E.H. Derrick. Det. H. Womersley.

Queenslandolaelaps vitzthumi. Male N1968190 (part), gnathosoma & legs. Queensland. LF 200. Leaf litter. Brookfield, nr. Brisbane. 10.5.1967. col. J. Herridge.

Queenslandolaelaps vitzthumi. Male N1968190 (part), chelicerae. Queensland. LF 2UQ. Leaf litter. Brookfield, nr. Brisbane. 10.5.19b7. col. J.A. Herridge.

Queenslandolaelaps vitzthumi. Male N1968190 (part). idiosoma. Queensland. LF 200. Leaf litter. Brookfield, nr. Brisbane. 10.5.19o7. col. J.A. Herridge.

Rhodacarellus. Female dorsal. N1968280. det. D.C. Lee. SOUTH AUSTRALIA. Tea^tree swamp, nr. Milliant Moss. LF 45. 11.3.1965. Col: N.B. Tindale.

Rhodacaroides minvaspis (ex Acugamasus). Female dorsal. N1970363. HOLOTYPE. SOUTH AUSTRALIA. LF 233/88/97. Litter, nr. Mt. Lofty summit, Cleland Reserve. 12.8.1968. Col. D.C. Lee.

Rhodacarus roseus. Male N19687984. Male N19688085. AUSTRALIA. LF 58. grass & leaf litter. Heywood Park & Hyde Park, S. A. 6.6.1965. col: R.V. Southcott. 249

Rhodacarus roseus. Female N19688478. Female N19688580. AUSTRALIA. LF 58. Grass & leaf litter. Heywood Park, Hyde Park, S. A. 6.6.1965. col; R.V. Southcott.

Sessiluncus hungaricus. Male N196874. ISRAEL. RL 65. Forest Litter (Wadi). Mishmar Haemek. 24.4.1967. col: M. Costa.

Solugamasus mustela (ex Rhodacaroides). HOLOTYPE. N1970368. Female V. SOUTH AUSTRALIA. LF 232. Moss nr. pond. Waterfall Gully, Cleland Res., nr. Adelaide. 9 May'68. Coll: D.C. Lee.

Solugamasus mustela (ex. Rhodacaroides). ALLOTYPE. N1970369. Male V. SOUTH AUSTRALIA. LF 232. Moss nr. pond. Waterfall Gully, Cleland Res., nr. Adelaide. 10 Oct'68. Coll: D.C. Lee.

Tangaroellus porosus Luxton. PARATYPE Female, ventral view. N1968271. Among barnacles. Magazine Point, Nelson, N. Z. 5.6.67.

Tangaroellus porosus Luxton. PARATYPE Male, ventral view. N1968273. Among barnacles. Magazine Point, Nelson, N. Z. 5.6.67

UNGARISCHEN NATURWISSENCHAFTLICHEN MUSEUMS BUDAPEST (UNMB):

Afrogamasellus bipilosus Karg. Female TYPE. ARGENTINIEN. Rio Negro, El Bolson, Mt. Piltriquitron NW, Tal 460 m. Laubstreu. 19.10.61.

Afrogamasellus squamosus Karg. TYPE. Einsendung. ARGENTINIEN. Rio Negro. Norquinco, 920 m. Laubstreu u. Grasbatzen. 23.2.61.

Gamasiphoides baloghi Karg, 1976. Male. Chel., B.II. gez. CHILE. Provinz Valdivia. Bodenstreu bei La Union an der Strasse nach Valdivia. 1965.

Gamasiphoides brevisetis Karg, 1976. Female TYPE. Chel. gez. CHILE. Misituni (Provinz Tarapaca). Bodenfalle ira Tal des Rio Lauca. 1965.

Gamasiphoides coniunctus Karg, 1976. Female TYPE. CHILE. Provinz Tarapaca. Uragebung von Cuesta el Melon unter Steinen. 1965.

Gamasiphoides longoventris Karg, 1976. Female TYPE. CHILE. Provinz Tarapaca, unter Steinen am. Fusse des Guallatiri-Vulkans. 1965.

Gamasiphoides octosetae Karg, 1976. Female TYPE. CHILE. Provinz Valparaiso, in feucht., verrott. Schilf am See. 5 km von Concon in Richtung Quintero. 1965. 250

Geogamasus apophyseus Karg. 1976. Maie TYPE, Chel., B.II. gez. CHILE. Curacavi (Prov. Santiago) Los Cerillos. Unter Baumrinde nahe Felsen, 3.10.65.

Geogamasus arcus Karg, 1976. Female. Chel. gez. CHILE. Azapa (Prov. Tarapaca). Nematodenprobe von Streu unter Weidensusch. 23.11.65.

Geogamasus ardoris Karg, 1976. Female. CHILE. Salto del Laja (Prov. Biobio). Berlese-Probe nahe Wasserfall, Moos am Felsen. 27.10.65.

Geogamasus bicirrus Karg, 1976. Male TYPE. B.II, Chel. gez. CHILE. Umgebung von (Prov. Tarapaca). Berlese-Probe von durchnasst. Moos. 20.11.65.

Geogamasus bisetosus Karg, 1976. Male TYPE, B.II, Chel. gez. CHILE. Azapa, (Prov. Tarapaca). Erdfalle m. Athylenglykol an einem Wassergraben. 2.12.65.

Geogamasus cochlearis Karg, 1976. Male TYPE. B.II. Chel. gez. CHILE. Sapahuira (Prov. Tarapaca). Bodenfallen mit Athylenglykol (3100 m). 1.12.65.

Geogamasus fibularis Karg. 1976. Male TYPE. B.II, Chel. CHILE. Maipu, Quebrada La Plata (Prov. Santiago).La Rinconada. Nematodenprobe aus Moos. 28.11.65.

Geogamasus flagellatus Karg, 1976. Female. CHILE. Umgebung von Azapa (Prov. Tarapaca). Bodenfalle aus cinein Flussbett. 2.12.65.

Geogamasus forcipis Karg, 1976. Female. CHILE. Misituni (Prov. Tarapaca), Aus Bodenfallen von feucht. Stellen. 1.12.65.

Geogamasus incisus Karg, 1976. Male TYPE. Chel., B.II. gez. CHILE. Pelchuquin (Prov. Valdivia), von Regenwurraern aus raodriger Erde. 25.10.65.

Geogamasus longisetosus Karg, 1976. Male TYPE. B.II. Chel. gez. CHILE. Misituni (Prov. Tarapaca), Grund wasserprobe vora Bach, sand. Ufer. 25.11.65.

Geogamasus monocuspidis Karg, 1976. Male TYPE. Chel., B.II. CHILE. Farellones (Prov. Santiago). Berlese-Probe m. nassem Gras (2100 m). 6.10.65.

Geogamasus pugionis Karg, 1976. Male TYPE. B.II, Chel. gez. CHILE. Farellones (Prov. Santiago) In 2000 m Hohe ira Gebusch, Erde m. Blattern. 6.10.65.

Geogamasus reticulatus Karg, 1976. Male TYPE. CHILE. Farellones (Prov. Santiago). Berlese-Probe unter Buschen. 6.10.65. 251

Geogamasus trispinosus Karg, 1976. Maie TYPE. Chel. gez., B.II. CHILE. Sapahuira (Prov. Tarapaca), an einem Salzsumpf (340Ü m). 24.11.65.

Heydeniella leei Karg. Maie. CHILE. Environs of La Union (Prov. Valdivia), an d. Strasse n. Valdivia. 26.1U.65.

Heydeniella mahunkai Karg. Female TYPE. ARGENTINIEN. Rio Negro, El Bolson, Mt. Piltriquitron 1170 m. Laubstreu, im Nothofagus Wald. 13.11.61.

Heydeniella tridentata Karg. Female. Chel. gez. CHILE. Collipulli (Prov. Malleco). Pan-Araerican-Highway, unter Steinen. 17.10.65.

Hydrogamasellus antennatus Karg, 1976. Female TYPE. CHILE. Farellones (Prov. Santiago). Berleso-Proben von Moos (2300 m). 6.10.65.

Hydrogamasellus armatissimus Karg, 1976. Female TYPE. CHILE. Collipulli (Proy. Malleco). Pan-Araerican-Highway. Regenwurmer unter Steinen. 27.10.65.

Hydrogamasellus avium Karg, 1976. Female TYPE. Chel. gez. CHILE. Umgebung yon Collipulli (Proy. Malleco), Pan-Amer-Highway. Unter Steinen. 27.10.65.

Hydrogamasellus cicatricosus Karg, 1976. Female TYPE. CHILE. Am Fusse des Guallatiri-Vulkans u. seiner Umgebung (Prov. Tarapaca), unter Steinen. 29.11.65.

Hydrogamasellus lanceolatus Karg, 1976. Female TYPE. CHILE. Umgebung von La Union (Prov. Valdivia), An d. Strasse n. Valdivia. 20.10.65.

Hydrogamasellus longopilus Karg, 1976. Female TYPE. CHILE. Umgebung von La Union (Prov. Valdivia), üesammeltes Material. 26.10.65.

Hydrogamasellus multospinosus Karg, 1976. Female TYPE. CHILE. Cerro El Roble, Cordillera de la Costa (Prov. Santiago). Blatter unter Strauchern. 29.9.65.

Hydrraamaselluc nasutus Karg, 1976. Female TYPE. CHILE. Farellones (Prov. Santiago). Berlese-Proben aus Gebusch. 0.10.65.

Hydrogamasellus testudinis Karg, 1976. Female TYPE. CHILE. Cuesta El Melon (Prov. Santiago), NW von Santiago de Chile. 3.11.65.

Laelaptiella media Karg. Female TYPE. CHILE. Provinz Santiago, Cuesta El Melon, unter Steinen, Laubstreu. 1965.

Neogamasellevans brevitremata Karg. Female TYPE. CHILE. Misituni (Prov. Tarapaca). Schleimige Algenproben aus klein Wasserpfuhl mit Nematoden. 29.11.65. 252

Neogamasellevans dentata Karg. Female TYPE, CHILE. Misituni (Prov. Tarapaca), in Bodenfallen unter Gestrauch. 1.12.65.

Neogamasellevans longocalcaris Karg. Female TYPE. Chel. CHILE. Misituni (Prov. Tarapaca), im nordl. Hochl., in Nematoden-proben aus Lamadung. 29.11.65.

Neogamasellevans macrochela Karg. Female TYPE. CHILE. Sapahuira (Prov. Tarapaca). In 3100m Hohe bodenfallen. 1.12.65.

Neogamasellevans ornata. Female TYPE. CHILE. Am Fuss des Vulkans Guallatiri (Prov. Tarapaca). 29.11.65.

Neogamasellevans serrata Karg. Female TYPE. CHILE. Misituni (Prov. Tarapaca). im Nordl. Hochland in Nematoden-proben aus Lamadung. 29.11.65.

ZOOLOGISCHE STAATSSAî-lMLUNG MÜNCHEN (ZSM);

Cyrtolaelaps minor n.sp. Jl/Fl. Knappenlicher bf Innsbruck.

Cyrtolaelaps minor n.sp. 7u,. WALDINACIS n GELBHALSSUAM. Wolfelagruech. 24/9 u 1/10.43. Det. C. Willmann.

Gamasellus % montanus. Maie. W6810-4cm.

Gamasellus montanus. Maie. W6810-4cm.

Gamasellus ? montanus. Female. Gamasolaelaptidae. W6810-4cm.

Gamasellus montanus. 69 a. Frenzel. Det. C. Willmann.

Gamasellus montanus. 64 a. Frenzel. Det. C. Willmann.

Gamasellus montanus Willmann 1936. 8.8.56. Nie 56. Female.

Gamasellus sertatus n.sp. Pubrava. Oesiua, Sudherre-geneiua. Ly. 215 la. Det. C. Willmann

Gamasellus ubatubaensis. TYPE. Female, Male. Felslitoral. Dr. Werner Hirschmann. Ubatuba. BRASILIEN. Schuster. 1960.

Hydrogamasus (Gamasellus) antarcticus. Female. Dr. Werner Hirschmann. Z 28 Nuggets Creek. Stilbocarpa litter. Macquarie Insel. Watson 1961.

Hydrogamasus (Gamasellus) antarcticus. Female. Wireless Hill Moss. Macquarie Insel. Watson 1961. 253

Hydrogamasus (Gamasellus) antarcticus. Female. Aerial Cove. Pucinellia. Macquarie Isl. Watson 1961.

Hydrogamasus (Gamasellus) antarcticus. Female. Handspike Point. Diomedea exulans. Macquarie Island. Watson 1961.

Hydrogamasellus (Gamasellus) antarcticus. Female chelicere. Nuggets Point. Stilbocarpa litter. Macquarie Isl. Watson 19bl.

Hydrogamasus (Gamasellus) antarcticus Tragardh 1907. 74. Dr. Werner Hirschmann. Langdon Point. Stilbocarpa soil. Macquarie Insel. Watson 1961.

Hydrogamasus (Gamasellus) antarcticus. Female TYPE. Langdon Point. Stilbocarpa litter. Macquarie Isl. Watson 1961.

Hydrogamasus (Gamasellus) antarcticus. Female. Brotherspoint. Cove scrapings. Macquarie Isl. Watson 1961. Z/146. Neoparasitid.

Hydrogamasus (Gamasellus) antarcticus. Male. Dr. Werner Hirschmann. Z 56 North head, Poa hamiltoni litter. Macquarie Insel. Watson 1961.

Periseius braziliensis. TYPE Female. Dr. Werner Hirschmann. 1966. Hartboden litoral. Recife. BRASILIEN.

Periseius braziliensis. (Eleven adult specimens). Dr. Werner Hirschmann. Br 152. Hartboden-litoral. Recife. BRASILIEN. Schuster. 1964.

Periseius braziliensis. Male. Dr. Werner Hirschmann. Hartboden litoral. Recife. BRASILIEN.

Periseius braziliensis neue Art. (Eleven adult specimens). Dr. Werner Hirschmann. Br 152. Hartboden-litoral. Recife/BRASILIEN. Schuster 1964.

Rhodacarellus corniculatus Willm. 1935. TYPE. Lpg. 20 d. Det. C. Willmann.

Rhodacarellus silesiacus n.sp. TYPUS. 33 b. Frenzel. Det. C. Willmann.

Rhodacarellus silesiacus Willm. Halle/S. Meschkat. XII-40. 80 cm. Det. C. Willmann.

Rhodacarellus silesiacus Willm. Male. N. 21.2. Det. C. Willman.

Rhodacarellus silesiacus Willmann. dorsal Female. B 10. cheliceren prep. Grasaussaat. Stahnsdorf. W. Karg. 7.1.58. 254

Rhodacarellus silesiacus Willm. 1936, 285/120. Halle/S. Meschkat. VII. üet. C. Willmann.

Rhodacarellus silesiacus Willm. Sechtem 57. Dikopshof III, B.6. 21.6.55. Det. C. Willmann.

Rhodacarellus silesiacus Willmann. Larve. 92. Kartoffelfela. Kleinmachnow b. Berlin. Vers. Br. 18.7.58, W. Karg.

Rhodacarellus silesiacus Willmann. Female ventral. Grasaussaat. Stahnsdorf. W. Karg. B 10 (5-lOcm). 27.11.58.

Rhodacarellus silesiacus Willmann. Female dorsal. Grasaussaat. Stahnsdorf. W. Karg. B 10 (5-lOcm). 27.11.58.

Rhodacarellus silesiacus Willmann. Pr. N. 106. Grasaussaat. Stahnsdorf. W. Karg. 25 B. 14.5.58.

Rhodacarellus silesiacus Willmann. D. N. 133. Grasaussaat. Stahnsdorf b. Berlin. B 1. 7.8.58.

Rhodacarellus silesiacus Willmann. Pr. N. 99. Grasaussaat. Stahnsdorf. W. Karg. C (10-15cra). 20.1.58.

Rhodacarellus silesiacus Willmann. Pr. N. Chel. Grasaussaat. Stahnsdorf b. Berlin. W. Karg. 25 C. 7.1.58.

Rhodacarellus silesiacus Willmann. Female. Chel. Grasaussaat. Stahnsdorf b. Berlin. W. Karg. C 10. 7.1.58.

Rhodacarellus subterraneus n.sp. 1935. TYPE. LPZG. 23 c. Det. C. Willmann.

Rhodacarellus subterraneus Willm. 1935. Female. LPZG. 20 d. Det. C. Willmann.

Rhodacarellus subterraneus Willm. 1935. Maie, Female. 22 b 4/5. Det. C. Willmann.

Rhodacaroides aeRVPtiacus n.g. n.sp.. Maie Atriph. Ghardaga. Mer Rouge. E. Schulz. 28.3.56. Det. C. Willmann.

Rhodacaroides aegyptiacus n.g. n.sp. Maie. Ghardaga. Mer Rouge, leg. Schulz. 28.3.56. Det. C. Willmann.

Rhodacaroides aegyptiacus n.g. n.sp. Female. Ghardaga. (Mer Rouge), leg. E. Schulz. 28.III.56. Det. C. Willmann.

Rhodacaroides aegyptiacus n.g. n.sp. Female. Ghardaga. Mer Rouge, leg. E. Schulz. 29.3.56. Det. C. Willmann. 255

Rhodacaropsis inexpectatus n.g.n.sp. Male, Female. Chelic. Kiel. Det. C. Willmann.

Rhodacaropsis inexpectatus. Male, Female. B. II. Kiel. Det. C. Willmann.

Rhodacaropsis inexpectatus Willm. Kurische Nebrung. Inti 1937. E. Schulz leg. Det. C. Willmann.

Rhodacarus c.f. strenzkei Willmann 1957. Female. Det. J.G. Sheals, 1963-iii.

Rhodacarus strenzkei n.sp.. TYPE. 1208. Strenzki. Gruue Insel. Eiderunsadung. Ig. 570.I—420. Det. C. Willmann.

Sessiluncus oculatus Vitzth. PARATYPOID Male. 28.8.1933. Zool. Staatssaramlg. München V893. TAHITI: Paparatal, 25Um, 7400m von der Kusten Zingiber sp. 21.12.1928. leg: Adamson, det: Vitzthura. Abt. Wirbellose Tiere. 256

APPENDIX B. DATA MATRIX

01 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 02 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 03 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 OUTGROUP

04 02 00 00 00 01 00 01 00 01 02 00 00 00 00 00 00 00 00 01 00 00 01 00 05 02 01 02 -1 00 00 02 02 00 02 01 04 05 03 02 00 00 01 00 00 00 00 00 06 00 01 00 01 00 00 00 -1 00 00 00 00 01 01 00 AFROGAMA

07 01 00 00 00 01 00 01 01 00 01 00 00 00 -2 00 00 01 01 01 00 00 00 01 08 -1 01 02 -1 00 00 00 02 01 02 01 03 05 03 02 00 00 00 00 00 00 00 00 09 00 01 00 00 01 00 00 01 01 00 00 00 00 00 01 AFRODACA

10 00 00 01 01 02 00 -1 00 00 02 00 00 00 -2 00 00 00 01 01 00 00 01 00 11 -1 01 02 00 01 00 03 02 00 01 02 02 01 02 02 00 00 01 00 00 00 00 00 12 00 01 00 00 00 02 00 00 01 00 00 00 00 00 04 RHODACAR

13 00 02 01 01 01 00 -1 00 00 02 00 00 00 -2 00 00 00 01 01 00 00 00 00 14 -1 01 03 00 00 00 00 02 00 02 01 00 01 02 02 00 00 01 00 00 00 00 00 15 00 01 00 00 00 02 00 01 00 00 00 00 00 00 00 CARELLUS

16 00 01 01 01 02 00 01 01 00 01 00 00 00 01 00 00 00 01 01 00 00 00 01 17 00 01 03 00 01 00 02 02 00 03 02 03 01 02 02 00 01 01 00 00 00 00 00 18 00 01 00 00 00 00 00 00 01 00 01 00 00 00 00 CAROPSIS

19 00 01 01 00 00 00 01 -1 00 02 00 00 00 01 00 00 01 01 00 00 01 01 01 20 00 01 02 00 00 00 01 02 00 00 01 05 03 05 02 00 00 00 00 00 00 00 00 21 00 00 00 00 00 01 00 00 01 00 01 00 01 03 00 CAROIDES

22 02 02 01 00 00 00 01 00 00 02 00 00 02 02 00 00 01 00 01 00 02 01 01 23 00 01 01 00 00 00 02 02 01 02 00 02 01 03 00 00 00 00 00 00 00 00 00 24 00 00 00 00 00 01 00 00 01 01 01 00 00 01 00 ALLOGAMA

25 00 01 01 00 00 02 -1 00 00 02 00 00 00 00 00 00 00 00 00 00 00 00 00 26 00 02 04 00 00 00 02 04 00 00 00 00 05 03 02 01 00 00 00 00 00 00 00 27 00 02 00 00 00 02 01 00 02 00 00 00 00 00 01 TANGAROE

28 02 00 01 01 00 02 01 00 00 02 00 01 00 00 00 00 01 00 01 00 00 01 00 29 00 02 02 00 00 00 00 05 00 00 00 03 03 03 02 01 00 00 01 01 03 02 01 30 04 00 04 00 00 00 00 -1 02 00 00 00 02 00 01 PANTENIP

31 02 01 01 01 00 02 01 00 00 02 00 01 00 00 00 00 01 00 01 00 00 01 01 32 01 01 02 -1 00 00 00 02 00 02 00 05 05 01 02 01 00 00 02 01 03 03 01 33 04 02 02 00 00 02 00 -1 00 01 01 00 02 01 04 LINDQUIS

34 02 02 01 01 00 00 01 00 01 02 01 00 00 -1 00 00 00 00 01 00 00 00 01 35 00 01 02 00 00 00 02 03 00 01 02 05 02 05 02 00 00 00 00 01 01 00 00 36 00 00 04 00 01 00 00 -1 01 00 00 02 02 -1 02 SESSILUN 257

37 02 00 01 00 00 00 01 00 01 00 00 00 00 00 00 00 00 00 01 00 00 01 01 38 00 01 03 00 00 01 00 03 00 01 02 03 05 04 02 00 00 00 00 00 00 00 00 39 00 00 04 00 00 00 00 -1 01 00 01 02 02 04 01 ANTENNOL

40 02 02 01 00 00 00 01 00 01 00 01 00 00 00 01 00 01 00 01 00 00 01 01 41 00 01 03 00 00 00 01 02 01 01 01 05 05 05 02 00 00 00 00 00 01 00 00 42 00 00 04 00 00 00 00 -1 01 01 01 02 02 00 00 MASIPHIS

43 02 00 01 00 00 00 01 00 01 01 00 00 00 00 01 00 01 00 01 00 00 01 01 44 00 01 -1 00 00 00 00 01 00 01 01 05 05 05 00 00 00 00 00 00 00 00 00 45 00 00 04 00 00 00 00 00 01 00 00 02 02 02 00 CALIPHIS

46 02 02 01 00 00 00 01 00 01 02 01 00 03 -1 01 00 01 00 01 00 00 01 01 47 00 -1 -1 00 00 01 03 00 00 00 02 05 05 05 00 00 00 00 00 00 01 00 00 48 00 00 00 00 00 00 00 -1 01 01 01 02 02 00 00 EUEPICRI

49 02 00 00 00 00 00 01 00 01 02 00 00 00 01 01 00 01 00 01 00 02 01 01 50 00 01 02 00 00 00 00 02 00 01 01 03 03 04 02 00 00 00 00 01 01 00 00 51 00 00 03 00 00 00 00 -1 01 00 01 01 01 00 00 MASIPHOI

52 00 00 01 02 00 00 01 03 01 00 00 00 00 01 01 00 01 00 01 00 00 01 00 53 00 01 03 00 00 00 00 02 00 00 01 05 05 05 02 00 00 00 00 00 01 00 00 54 01 00 04 00 00 01 00 -1 01 01 01 00 02 02 00 HYDROGAM

55 02 01 01 00 00 00 01 00 00 00 00 00 00 01 01 00 01 00 01 00 02 00 01 56 00 01 02 00 00 00 00 02 00 01 01 05 05 05 02 00 00 00 00 00 00 00 00 57 00 00 04 00 00 00 00 -1 01 00 01 00 02 00 00 LAELAPTI

58 00 00 01 00 00 00 01 00 00 00 00 00 00 00 00 00 01 00 00 00 00 00 00 59 00 01 03 00 00 00 02 01 00 00 01 05 03 03 01 00 00 00 00 00 00 00 00 60 00 00 04 00 00 01 00 01 01 01 01 00 00 01 00 PARASITI

61 02 00 01 01 00 00 01 00 01 02 01 01 -2 00 01 00 01 00 00 00 00 00 01 62 00 01 00 00 00 01 00 01 00 00 00 05 05 05 00 00 00 00 00 00 00 00 00 63 00 00 04 00 00 00 00 00 01 02 02 00 01 02 03 PYRIPHIS

64 02 02 01 00 00 00 01 00 00 00 00 00 00 00 00 00 01 00 00 00 00 01 00 65 00 01 01 00 00 00 02 00 00 02 01 03 00 01 00 00 00 00 00 00 00 00 00 66 00 00 00 -1 00 00 00 00 00 02 01 00 00 -1 00 EURYPARA

67 02 02 01 00 00 00 01 00 00 02 00 00 -1 -2 00 00 01 00 00 00 00 01 00 68 00 00 -1 00 00 01 02 00 00 01 01 05 01 01 00 00 00 00 00 00 00 00 00 69 00 00 00 01 00 00 00 00 00 00 01 00 00 00 00 CYRTOLAE

70 02 00 01 00 00 00 01 00 01 02 00 00 01 00 01 00 01 00 00 00 02 01 01 71 00 01 02 00 00 01 00 02 01 00 01 05 05 05 02 00 00 00 00 00 00 00 00 72 00 00 04 00 00 00 00 -1 01 00 01 00 01 04 01 OLOGAMAS 258

73 02 00 01 00 00 00 01 00 01 02 01 00 01 00 01 00 01 00 00 00 01 01 01 74 00 01 02 00 00 02 00 02 00 00 00 05 05 05 02 00 00 00 00 00 00 00 00 75 00 00 04 00 00 00 00 -1 00 00 01 02 01 02 01 CYMIPHIS

76 02 00 01 00 00 00 01 00 01 02 00 00 00 00 00 00 01 00 00 00 00 01 01 77 00 01 02 00 00 00 00 02 00 01 02 04 03 03 02 00 00 00 00 00 00 00 00 78 00 00 02 00 00 00 00 00 01 00 00 00 01 04 03 ATHIASEL

79 02 00 00 00 00 00 01 00 01 02 01 00 00 00 01 01 01 00 00 00 00 00 01 80 00 01 02 00 00 00 00 02 00 01 00 05 05 05 02 00 00 00 00 00 00 00 00 81 00 00 04 00 00 00 00 -1 01 00 00 02 01 04 03 HEYDENIE

82 02 02 01 02 00 00 01 00 01 02 00 00 00 00 00 00 01 00 01 00 00 01 01 83 00 01 02 00 00 00 01 02 01 02 02 05 05 05 02 00 00 00 00 00 00 00 00 84 00 00 01 00 00 00 00 -1 01 00 00 00 01 04 03 GEOGAMAS

85 02 00 01 00 00 00 01 00 00 02 00 00 00 00 00 00 01 00 00 00 00 01 01 86 00 01 02 00 00 00 00 02 00 01 01 05 05 05 02 00 00 00 00 00 00 00 00 87 00 00 02 00 00 00 00 -1 01 00 00 00 01 04 02 HYDROSEL

88 02 00 01 00 00 00 01 00 01 02 00 00 00 00 00 00 00 00 01 00 00 01 00 89 00 01 02 00 00 00 00 02 01 02 02 05 05 05 02 00 00 00 00 00 00 00 00 90 00 00 04 00 00 00 00 -1 01 00 01 00 01 04 01 NEOGAMAS

91 02 00 01 00 00 01 01 00 00 02 00 01 03 00 00 00 01 00 00 00 00 00 00 92 00 01 01 00 00 02 00 03 01 00 00 03 00 02 00 00 00 00 00 -1 00 00 -1 93 00 00 00 00 00 00 00 01 00 00 00 00 00 -1 03 HETEROGA

94 00 00 01 -1 00 00 -1 00 00 00 00 00 00 02 00 00 01 00 01 00 00 00 00 95 00 01 02 00 00 00 02 02 01 00 02 03 01 02 02 00 00 00 00 00 00 00 00 96 00 00 00 00 00 01 00 01 00 00 00 00 00 -1 00 LITOGAMA

97 00 00 01 00 00 00 01 00 01 02 01 01 03 02 00 01 01 00 01 00 00 01 00 98 00 01 02 00 00 01 00 02 00 00 01 05 05 05 02 00 00 00 00 -1 00 00 -1 99 00 00 00 00 00 00 01 -1 01 00 01 02 01 03 00 PERISEIU

CO 02 00 01 00 00 00 00 00 01 02 01 01 00 02 00 00 01 00 01 00 00 02 01 Cl 00 01 02 00 00 00 01 02 00 00 01 05 05 05 02 00 00 00 00 00 00 00 00 C2 00 00 00 00 00 00 00 01 01 00 01 00 01 03 00 PSAMMONS

C3 02 00 01 00 00 00 01 00 00 00 00 01 02 00 00 00 01 00 01 00 01 01 01 C4 00 -1 -1 00 00 01 01 01 00 00 00 05 05 00 00 00 00 00 00 00 00 00 00 C5 00 00 00 00 00 00 00 00 00 00 00 00 01 02 00 PILELLUS

C6 01 02 01 02 01 00 01 00 00 02 00 00 00 01 00 00 01 00 00 00 00 00 00 C7 00 01 04 00 00 00 03 03 01 02 02 03 05 02 02 00 00 00 00 00 00 00 00 C8 00 00 00 00 00 00 00 01 00 00 01 00 00 04 00 SOLUGAMA 259

C9 01 00 00 -1 00 00 01 -1 00 00 00 00 03 00 00 00 01 00 01 00 01 01 00 10 00 01 01 00 00 02 00 01 00 00 01 03 00 02 00 00 00 00 00 00 00 00 00 11 00 00 00 00 00 00 00 01 01 00 01 00 00 01 03 ACUGAMAS

12 02 00 01 00 00 00 01 00 01 01 00 00 03 02 00 00 01 00 01 00 01 01 00 13 00 01 02 00 00 01 00 01 00 00 00 04 01 02 01 00 00 00 00 00 00 00 00 14 00 00 00 00 00 00 00 -1 01 00 01 00 01 04 00 FALCIGER

15 02 00 01 00 00 00 01 00 01 01 01 00 02 02 00 00 01 00 01 00 00 00 00 16 00 01 -1 00 00 01 00 -1 00 01 00 03 01 02 00 00 00 00 00 00 00 00 00 17 00 00 02 00 00 00 00 -1 01 01 01 01 01 02 00 DISCUTAT

18 01 00 01 00 00 00 01 00 00 01 00 00 02 02 01 00 01 00 01 00 00 01 01 19 00 01 01 00 00 01 02 01 00 00 00 03 02 05 00 00 00 00 00 00 00 00 00 20 00 00 00 00 00 00 00 -1 01 00 01 00 01 04 00 PYRIFORM

21 02 00 00 00 01 00 01 00 00 02 00 00 03 02 00 00 01 00 01 00 01 00 00 22 00 01 02 00 00 01 00 01 00 00 00 03 03 02 01 00 00 00 00 00 00 00 00 23 00 00 00 00 00 00 00 00 01 00 01 00 01 03 00 TRAGARDH

24 02 02 01 00 00 00 01 01 01 02 00 00 00 00 00 00 01 00 01 00 00 00 00 25 00 01 02 00 00 01 00 01 01 00 02 03 03 03 02 00 00 00 00 00 00 00 00 26 00 00 04 00 00 01 00 -1 01 00 01 02 02 04 00 QUEENSLA

27 01 02 01 02 01 00 01 02 00 02 00 00 00 00 00 00 00 00 01 00 00 00 00 28 00 01 02 00 00 00 02 02 00 02 02 05 04 03 02 00 00 00 00 00 00 00 00 29 00 00 00 00 00 00 00 01 01 00 00 00 00 00 00 PARAGAMA

30 01 02 01 00 00 00 00 00 01 00 00 01 00 00 01 00 00 00 00 00 00 01 01 31 00 01 00 00 00 00 00 -1 00 02 00 05 05 05 00 00 00 00 00 00 00 00 00 32 00 00 04 00 00 00 00 -1 -1 00 00 00 02 -1 02 EPIPHIS

33 02 02 01 00 00 00 01 00 01 01 00 00 03 00 01 00 01 00 01 00 00 01 01 34 00 01 02 00 00 02 01 01 00 00 00 05 05 05 01 00 00 00 00 00 00 00 00 35 00 00 00 00 00 01 00 -1 01 00 01 01 01 03 00 GAMASELI

36 02 00 01 00 00 00 01 00 01 02 00 01 00 00 00 00 01 00 01 00 00 01 01 37 00 01 02 00 00 00 00 01 00 01 02 04 05 02 02 00 00 00 00 00 00 00 00 38 00 00 03 00 00 01 00 -1 -1 00 00 00 02 01 01 RYKELLUS

39 01 00 00 00 01 00 01 00 01 02 01 01 00 01 01 01 01 00 01 00 00 01 00 40 00 01 02 00 00 00 01 01 00 01 01 05 05 05 02 00 00 00 00 00 00 00 00 41 00 00 03 00 00 00 00 00 01 00 00 02 01 02 01 HIHINNUS

42 02 00 00 00 00 00 01 00 01 00 00 01 00 01 01 00 01 00 01 00 00 02 00 43 00 01 02 00 00 00 02 01 00 02 00 05 04 05 02 00 00 00 00 00 00 00 00 44 00 00 00 00 01 01 00 -1 01 01 01 02 02 04 00 HIBIPALA 260

45 02 00 01 00 00 01 -1 00 00 00 00 01 02 00 00 00 01 00 02 00 00 00 00 46 00 01 02 00 00 02 03 02 00 00 00 05 05 05 02 00 00 00 00 00 00 00 00 47 00 00 00 00 00 00 00 01 00 00 00 00 00 04 00 EVANSSEL

48 02 02 01 01 00 00 -1 -1 00 01 00 00 00 -2 00 00 00 01 00 00 00 01 01 49 00 02 04 02 00 00 00 03 00 03 -1 05 01 01 02 02 -1 00 00 00 00 04 02 50 03 01 00 00 00 00 01 00 02 -1 -1 00 00 00 00 LAEMITIS

51 02 00 01 01 00 00 01 00 00 01 00 00 00 00 00 00 00 01 02 00 00 -1 01 52 00 03 04 02 00 00 00 01 00 03 -1 05 01 01 02 02 -1 00 01 01 02 03 00 53 02 00 00 00 00 00 00 -1 01 -1 00 00 00 00 00 CHINENSI

54 01 00 00 00 01 00 01 00 00 02 00 00 00 00 00 00 00 00 01 00 00 01 00 55 02 01 02 -1 00 00 02 02 00 02 01 03 05 05 02 00 00 01 00 00 00 00 00 56 00 01 00 00 00 00 00 00 -1 00 00 00 00 00 00 MONGII

57 01 00 00 00 01 00 01 01 00 02 00 00 00 00 00 00 01 00 01 00 00 01 01 58 02 01 02 -1 00 00 01 02 01 02 01 03 05 03 02 00 00 01 00 00 00 00 00 59 00 01 00 01 00 00 00 00 -1 00 00 00 01 00 00 BAKERI

60 02 00 01 01 01 02 01 01 00 00 00 00 00 00 01 00 00 01 01 00 00 01 00 61 -1 01 02 00 00 00 01 02 00 01 00 00 03 02 02 00 00 00 01 00 02 02 00 62 03 00 00 00 00 01 00 00 00 00 00 00 00 00 00 DENDROSE

63 02 01 01 01 00 02 01 01 00 00 00 00 00 00 01 00 00 01 01 01 00 00 00 64 -1 01 03 01 00 00 02 02 00 00 00 05 05 03 02 00 00 00 01 01 02 02 00 65 03 00 00 00 00 00 00 00 01 00 00 00 00 00 01 DIGAMASE

66 01 01 01 00 00 02 00 00 00 00 00 00 00 00 00 00 01 00 01 01 00 00 00 67 00 01 00 01 00 00 00 00 00 00 00 02 01 00 00 01 00 00 00 00 01 01 00 68 00 00 04 00 00 01 00 01 01 -1 02 00 02 00 02 PROCTOLA

69 01 01 01 02 00 02 00 00 00 01 00 00 01 00 00 00 00 00 01 00 00 00 00 70 02 01 00 01 00 02 00 00 00 00 00 03 00 03 00 01 00 00 00 00 01 01 00 71 00 00 04 00 00 00 00 01 00 00 01 00 02 00 02 LASIOSEI

72 02 00 00 00 00 02 00 00 00 00 00 00 00 -2 00 00 01 00 02 01 00 01 00 73 00 01 01 01 01 00 01 03 00 02 00 00 00 03 01 01 00 00 00 01 01 01 00 74 01 00 00 00 00 02 00 01 -1 02 01 00 00 00 00 PROTOGAM

75 02 00 01 00 00 00 00 02 00 01 01 00 00 00 00 00 00 00 00 00 00 00 01 76 00 01 -1 00 00 00 00 -1 01 00 00 02 00 01 00 00 00 00 00 00 00 00 00 77 00 00 01 00 00 00 00 00 00 00 00 00 00 04 00 VEIGAIA

78 02 00 01 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 01 79 00 01 01 00 00 00 00 00 00 00 00 03 00 00 01 00 00 00 00 00 00 00 00 80 00 00 01 00 00 02 00 -1 00 01 01 00 00 01 00 GAMASOLA 261

81 02 02 01 02 00 00 01 00 01 02 00 00 00 00 00 00 00 00 00 00 00 01 01 82 00 01 02 00 00 00 02 02 00 01 01 05 05 02 02 00 00 00 00 00 00 00 00 83 00 00 03 00 00 00 00 01 01 00 00 02 01 00 ? MASELLEV

84 02 02 00 00 00 00 01 00 01 02 00 00 01 -1 00 00 01 00 00 00 00 00 01 85 00 01 03 00 00 00 00 02 01 01 02 03 02 03 02 00 00 00 01 01 01 00 00 86 01 01 04 00 00 00 00 -1 01 00 00 00 02 04 ? SELOPSIS

87 02 UK UK UK 00 00 00 UK 00 02 00 UK 03 00 00 00 00 00 02 01 00 01 01 88 00 -1 02 -1 00 02 03 02 00 02 00 00 01 01 02 00 00 00 00 00 00 00 00 89 00 00 00 00 00 01 00 -1 00 00 00 00 00 04 ? POOONOTO

90 02 00 01 00 00 00 01 00 00 02 00 00 02 00 00 00 01 00 02 01 00 01 01 91 00 -1 02 -1 00 02 03 02 00 02 00 02 01 00 02 00 00 00 00 -1 00 00 -1 92 00 00 00 00 00 02 00 00 00 00 00 00 00 00 ? NOTOGAMA

93 02 UK UK UK 01 00 01 UK 00 01 00 UK 00 02 00 00 01 01 01 00 00 01 00 94 -1 01 02 -1 00 00 02 02 00 02 01 02 03 03 02 00 00 01 00 00 00 00 00 95 00 01 00 00 01 00 00 01 -1 00 00 00 00 00 01 CITRI

96 01 UK UK UK 01 00 01 UK 00 02 00 UK 00 -2 00 00 01 00 01 00 00 00 00 97 00 01 02 -1 00 00 02 02 00 02 01 03 05 02 02 00 00 01 00 00 00 00 00 98 00 01 00 00 00 00 00 01 -1 00 00 00 00 00 ? LUBALENS

99 00 UK UK UK 00 00 01 UK 01 02 00 UK 00 01 00 00 01 00 00 00 00 01 01 DO 00 01 02 00 00 01 00 01 01 01 02 05 05 02 02 00 00 00 00 00 00 00 00 D1 00 00 03 00 01 01 00 -1 01 00 01 00 01 04 00 BERLESEI

D2 02 UK UK UK 00 00 01 UK 01 02 00 UK 00 02 00 00 00 00 01 00 00 01 01 03 00 01 02 00 00 00 02 01 00 02 01 03 05 05 02 00 00 00 00 00 00 00 00 04 00 01 00 00 01 00 00 -1 01 01 01 02 01 04 01 VIRGUNCU

05 01 UK UK UK 00 00 01 UK 01 02 01 UK 00 00 00 00 00 00 01 00 00 01 01 06 00 01 02 00 00 00 00 01 00 02 02 03 05 02 02 00 00 00 00 00 00 00 00 07 00 00 04 00 00 00 00 -1 01 00 01 02 01 04 01 ONCHOGAM

08 02 UK UK UK 03 00 01 UK 00 01 00 UK 00 00 00 00 00 00 02 01 00 01 01 09 00 01 04 00 00 00 03 03 00 02 01 01 00 00 02 00 00 00 00 00 00 00 00 10 00 00 00 00 00 01 00 01 00 01 01 00 00 04 ? MINYASPI

11 00 UK UK UK 01 00 01 UK 00 02 00 UK 00 00 00 00 01 00 00 00 00 01 01 12 00 01 03 00 00 00 03 03 00 02 01 03 01 05 02 00 00 00 00 00 00 00 00 13 00 00 00 00 00 02 00 00 00 00 00 00 00 04 00 COSTAI 262

14 02 UK UK UK 02 00 01 UK 00 01 00 UK 00 00 00 00 00 01 01 00 00 01 01 15 00 01 02 -1 00 00 00 02 01 02 01 05 05 02 02 00 00 01 00 00 00 00 00 16 00 01 00 00 00 00 00 -1 01 01 00 00 _-1 01 02 SQUAMOSU

17 02 00 00 00 00 00 01 00 00 01 00 00 -2 00 00 00 01 01 01 00 00 00 01 18 00 01 02 -1 00 02 00 02 01 02 01 05 05 02 02 00 00 01 00 00 00 00 00 19 00 01 00 00 00 00 00 -1 01 01 00 00 -1 01 02 BIPILOSU

20 02 UK UK UK 00 00 01 UK 00 01 00 UK 00 00 00 00 01 00 01 00 00 01 00 21 00 01 02 -1 00 00 00 02 01 02 01 05 05 02 02 00 00 01 00 00 00 00 00 22 00 01 00 00 00 00 00 -1 01 01 00 00 -1 01 ? FURCULAT

23 02 UK UK UK 01 00 01 UK 00 02 00 UK 00 -1 00 00 01 00 01 00 01 01 00 24 00 01 02 00 00 00 03 02 00 02 01 03 02 01 02 00 00 00 00 00 00 00 00 25 00 00 00 00 00 00 00 01 00 01 01 00 -1 00 01 MICHIGAN

26 00 01 01 00 00 02 -1 00 00 01 00 00 00 00 00 00 01 01 01 00 00 00 00 27 -1 01 02 00 00 00 01 01 00 00 00 00 01 00 01 01 00 00 00 01 01 01 00 28 00 00 00 00 00 02 00 01 01 -1 00 00 -1 00 02 SELLODES