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MANUAL OF MESOSTIGMATID
MITES

R. W. STRANDTMANN
&
G. W. WHARTON

THE INSTITUTE OF ACAROLOGY

1958

MAY 9 1960
^
 A MANUAL OF MESOSTIGMATID MITES
PARASITIC ON VERTEBRATES

by

R. W. Strandtroann
Texas Technological College, Lubbock, Texas

and

G. W. Wharton
University of Maryland, College Park, .Maryland

KEFEBSB PEST M6MI IMPO ANAIYSIS CIK

AFPMB, WEST GtSM SECTION, BRAMC
WASHIKGTOH, DC 20307

Edited by Conrad E. Yunker

Contribution No. 4
of
THE INSTITUTE OF ACAROLOGY
Department of Zoology
University of Maryland
College Park

20 December 1958
This investigation was supported by the Medical Research and Develop-
ment Board, Office of the Surgeon General, Department of the Army,
under Contract No. DA-49-007-MD-501 with the University of Maryland.
It was edited and published with the aid of National Science Foundation
Grant NSP-G4561.

ii
 PREFACE

This manual is an attempt to list all species of mites in the sub-

order Mesostigmata that parasitize vertebrates and to bring under one
cover a condensation of all information on their biology, distribution,
life history, and medical and economic significance. All papers pub-
lished on or before 31 December 1956 were considered in the prepara-
tion of the entire manuscript. Papers published during 1957 and 1958
that were found too late for inclusion in all sections of the book were in-
cluded, where feasible, in the later sections. It was found that to revise
the entire manuscript (especially those portions that had already been
mimeographed) whenever a new publication came to hand would have
been a perpetual task and publication would have been impossible.
The work was made possible by support received by the University
of Maryland from the Army from 1953 to 1957. The publication of the
Manual was facilitated by a grant to the University of Maryland by the
National Science Foundation. In the conduct of the work, the biblio-
graphic system, the original search of the literature, and the organiza-
tion were the responsibility of G. W. Wharton. The study of specimens
and the preparation of the manuscript were done by R. W. Strandtmann.
Both authors, of course, assume full responsibility for whatever recep-
tion this book merits.
Scientific names of the animals in the host list were checked by Drs.
Doris M. Cochran’(Reptilia), Herbert Friedmann (Aves) and David H.
Johnson (Mammals). These specialists arranged the hosts in their
proper classes and orders. We are grateful for their help. It should
be emphasized that the names for hosts are those used by the original
author sand are not necessarily the current valid names. The names used,
therefore, do not represent the opinions of these specialists concerning
the valid names of the species of hosts.
Extraordinary thanks are due Dr. Conrad E. Yunker who edited
the entire work, and Allie M. Brown who typed the final drafts and cut
the stencils. Especial thanks are dueAlena Elbl for translating the Rus-
sian literature and Richard 0. Albert for getting several Japanese papers
translated. We also wish to thank Martha Schultz Webster who typed the
first draft, and Leta Jane Holman for reading proof. Duvall A. Jones
andEmeron M. Montgomery assisted in the preparation of the final copy.
To all these and many more, we are deeply grateful.

R. W. Strandtmann

G. W. Wharton
 TABLE OF CONTENTS

PREFACE Page iii ...

’

INTRODUCTION Page 1

,
Medical and Economic Importance 3
Control............................ 5 .

. ." ’’,’. BEHAVIOR Page 1. . ,.

\’ LIFE CYCLES Page, 10

............. 10 Mating and Insemination ...,.
.

Dermanyssidae. 13
Laelaptidae ................ 13 Longevity .................. 14

ANATOMY Page 15
Segmentation ............... .15 _ Internal Anatomy ........... 18
External Anatomy ........... 15 The Life Stages ............ 23
’
CLASSIFICATION Page 26
The Parasitic Mespstigmatid Mites ..... 27
^ Key to the Families of Parasitoidea ..... 28
,

Family LAELAPTIOAE Berlese, 1892.. . 29

Key to the Genera of Laelaptinae 30
Genus Haemolaelaps ....... ,32 Genus Heterolaelaps ........ 55
Genus Cavilaelaps .......... 47 Genus Trichosurolaelaps. .... 56
Genus Bolivilaelaps ...,...,. 48 Genus Steptolaelaps ......... 56
Genus Eubrachylaelaps ...... 48 Genus Laelaps ............. 57
Genus Gigantolaelaps ........ 50 Genus Longolaelaps ......... 71
Genus Neoparalaelaps ....... 53 Genus Oryctolaelaps ........ -71
Genus BadfordUaelaps ........ 53 Genus Echinolaelaps ........ 72
Genus Mesolaejaps .......... 54 Genus Mysolaelaps .......... 76
Genus T.richolaelaps ........ 55 Genus Aetholaelaps ......... 76
.Family DERMANYSSIDAE Kolenati. 1859.. 77

the Dermanyssidae ...................

Key to the Subfamilies and Genera of

Subfamily Macronyssinae Oudemans, 1936.. 80

77

Genus Macronyssus ......... 81 Genus Neospinolaelaps....... 98

Genus Ornithonyssus ........ 81 Genus Austrolaelaps. ........ 98
Genus Ichoronyssus ......... 89 Genas Hirstionyssus ........ 99
Genus Spinolaelaps .......... 95 Genus Neoichoronyssus ...... 108
Genus Sauronyssus .......... 96 Genus Patrinyssus .......... 109
Genus Ophionyssus .......... 97 Genus Echinonyssus ......... 109
 Subfamily Macronyssinae Oudemans, 1936 (contd.)
Genus Lepronyssoides ...... 110 Genus Steatonyssus ....... 113
Genus Hu-stesia ........... 110 Genus Pellonyssus 116
Genus Neolaelaps .......... ’Ill Genus Radfordiella
_ _
117
Genus Liponysella ......... 112 Genus .7,............ 118
Tur^.
Subfamily Dermanyssinae Kolenati. 1859.. 118
Genus Allodermanyssus .... 11.8... Genus Dermanyssus........ 120
Genus Liponyssoides ....... 120
Subfamily My onyssihae. New subfamily... 124
Genus Myonyssus .......... 124
Family HAEMOGAMASIDAE Oudemans,
1926 ................................ 126
Key to the Genera of-Haemogamasidae .... 126
Genus Eulaelaps ........... 126 Genus Brevisterna ......... 136
Genus Haemogamasus ......: 129 Genus Ischyropoda ......... 137
Family IXODORHYNCHIDAE Ewing, 1923.. 138
Key to the Genera of Ixodorhynchidae ..... 138
Genus Ixodorhyhchus ....... 138 Genus- Hemilaelaps......... 139
Family RAILLIETIDAE Vitzthum. 1941 140 ..-.
Genus Raillietia ........... 141
Family SPELEORHYNCHIDAE Oudemans,
,. 1902 .’......................,........
..
141
Genus Spelaeorhynchus ..... 142 .......
Family DASYPONYSSIDAE Fonseca, 1940. 142

Genus Dasyponyssus- .. 143 i. ..

Key to the Genera of Dasyponyssidae ..... 142
Genus Manitherionyssus .... 143
Family .ENTONYSSIDAE Ewing, 1922 ..... 143
Key to the- Genera of Entonyssidae. .;’..... 144
Genus Entonyssus .......... 144 Genus Hamertonia ......... 146
Genus Ophiopneumic ola ..... 145 Genus Pneumophionyssus ..’. 147
. .Family HALARACHNIDAE Oudemans, 1906. 147
. .
,
.
... Key to the Genera of the Halaraehnidae .., 147 ;.
.
.
Genus .Halarachne .......... 1.48 Genus Rhinophaga ;.;........ 153
Genus .Qrthohalarachne ..... 149 Genus PneumoBysspides... .... ’’ 154
Genus .Pneumonyssus ....... ISl’ ".’ ;’.
’

. .... .Family KHINONYSSIDAE Trouessart, 1895.-’isa

.Key to the Genera of Rhinonyssidae....... ’155
.
.
Genus Rhinonyssus ......... 157- Genus Cas .......... ...... 169
Genus Larinyssus .......... 160 Genus Ptilonyssus ,.’........ 169
Genus Rallinyssus ......... 160 Genus Flavionyssus ........ 171
Genus Tinaminyssus. New .
_
Genus RochanyssuS ........ 171
genus ... .,»... .._,..........
161.. Genus Tra-vanyssus ........ 171
Genus Neonyssus........... 161- Genus Rhino.nyssoides ...... 172
Genus Rhmoecius 164 ..........
Genus .Vit^nyssus ............
173’
G.enus Ruandanyssus . ....... 165. Genu^ Paraneonyssus 176 ......
Genus Sternostoma .........
165. Genus Ptilonyssus. .;......... 178-
,
_, Family SPINTURNICIDAE Oudem.an3..1.901. 182
, ,. Key to the Genera of- Spinturnicidae........... 183
 Family SPINTUBNICIDAE (contd.)
Genus Spinturnix ..........
133 Genus Periglischrus ....... 189
Family HETEROZERCONIDAE Berlese,
1892................................ 191
Genus Heterozercon ....... 192
Trigynaspida
Family PARAMEGISTIDAE Tragardh, 1946.. 192
Genus Ophiomegistus ...... 192
Genera of Uncertain Familian Relation-
ships .......................
Acanthochela ....... 193
193
Genus Mungosicola ........ 195
Genus
Genus Alphalaelaps ........ 193 Genus Tympanospinctus . .. . 195
Genus Manisilaelaps ....... 194 Genus Ugandolaelaps ....... 195
Genus Myonyssoides ....... 194
Genera and Species Not Rightly Belonging
In this Manual ....................... 196
Conclusion ,..................... 197

DISTRIBUTION Page 198

Faunal List ........................... 201
North America...... ...... 201 Australia ................. 211
Central America .......... 202 Islands of the Arctic ....... 211
South America ............ 202 Islands of the Atlantic ...... 211
Europe ................... 204 Islands of the Indian Ocean. . 212
Asia ..................... 206 Indonesia ................. 213
Asia Minor ............... 207 Islands of the Pacific ...... 213
Africa ................... 207
Hosts ................................. 215
Mammalia ............... 216 Reptilia................... 259
Aves ..................... 247
Host Specificity ........................ 261

APPENDIX Page 263

Collecting Methods ........ 263 Culture Methods ........... 267
Mounting Methods ......... 264

LITERATURE CITED Page 271

INDEX Page 317

 LIST OF PLATES AND FIGURES

Plate -No’.

Ventral view of gnathosoma of a. laelaptid mite . ;

Illustration of three types of tecta . . . . . . .. . ’

Tritosterna. . . . . . . . . . .
Eemale chelae . . . . . . . . . . .
. 1. ...
.,... . . .’
. . .
. . . .: .
Digestive system . . . . . . . . . . . . . . ..;...’...
Side .view of gnathosoma of Haemolaelaps glas-

’
Sectional view. of a male gamasid mite (Neonyssus
melloi) . . . . . . . . . . . . . . . . . . . . . . .
Ambulacrum of tarsus I of two snake mites . .’.. .’.
Diagrammatic sketch showing the respiratory sys-
tem . . . . . . . . . . . . . . . . . . . . . . .’;. ...".’
Sectional view.showing internal organs of a female
gamasid.mite (Neonyssus melloi) . . . . . . ’.’...;.. 4
Laelaps nuttalli (.larva) . . . . . . ...
. . . . . . .
Haemolaelaps .glasgowi (protonymph) . . . . . . .
5
6
Haemolaelaps .glasgowi (deutonymph) . . . .’. ,-’.? 6
Echinolaelaps .echidninus (female) , . . . . . . .. 7
Echinolaelaps .echidninus (male) ...’.’.’. . . . . . . .’
Types of male chelae . . . . . s ’s. ._.. . . .
Haemolaelaps glasgowi . . . . . ."’ . . . .
. .
:,: .-. .
...
7
8
9
’

Cavilaelaps .bresslaui . , . . . , ..., . ^ .’. / ,

Bolivilaelaps tricholabiatus . . .^.’.. i ^ ’’i’i..^ ’,
l&
11
..
Eubrachylaelaps jamesoni . » . .
’. .
;.’. . ’t^. .’ i 13
’
Gigantolaelapscricetidarum ,’ ., V’..:.’.’;.::..;’. . . . . 13.; 14
Neoparalaelaps bispinosus’. ; v’..’ .’.;; . . . .
..’,
Radfordilaelaps meridionalis . .’.. . . . ’.’:.’. . ^ .
15
15 1

Mesolaelaps anomalus ..
: . , ; /.’.. .
,; .:..:,; .:...... ’ 16
Heterolaeiaps antipodianus . , . . , . . . :. ’. . . ’»’ 1&
Tricholaelaps .comatus , . .
, ’.^:.;;; , . . . . ... 17
Tri.chosu.rolaelaps.crassipes . . . . . . . . . . . .’ : 17
Steptolaelags heteroniys < . . . ..;;.
j :. ...
. . 18-.
i .
Laelaps nuttalli , . , . .
. . , . . . .’J. ’,’-1 . . 19’’
i

Longolaeilaps longulus . . . . . . . .;’. ., ’. . . . . 20-

Oryctolaelaps -bibikovae . . . . . . . ;,..;. . . . . 20.
Mysolael-apsparvispinosus . . . . . . ,. . . . ’.. ... .’ 21
Aetholaelapssylstra-i. . . < . . . . ;:.’,.’. . . .; ’ 22-
Macr.onyss.us lon-gi-manus . . . . .... ...
. . . . . . 23-
Qmithonysaus-sylviarum. . . . . ,’. . . .’. . . . 24
Qrnithonyssus burs a .. ... . . . . . . ..’. . . . . , 25
Ichor.onyssus longisetosus , . , .
. . ;.. . . . . . 2 &
Ichoronyssus quadridentatus . . . . . . . . . . . . 27
 Plate No.

39. Spinolaelaps sp. . . . . . . . . . . . . . . . . . . 27

40. Sauronyssus gordonensis . . . . . . . . . . . . . . 28
41. Ophionyssus natricis . . . . . . . . . . . . . . . . 29
42. Neospinolaelaps miniopteri , . . . ... ... . . . . 30
43. Austral olaelaps mitchelli . . . . . . . . . . . . . 30
44. Hirstionyssus br’eviseta. . . ... . . . . . . . . . 31

’
45. Neoichoronyssus werneoki........... . . . . . . ..32-
46. Patrinyssus hubbardi. . . .........
47. Echinonyssus nasutus . . .. . . . .
. .: . . . . . .
. . . . . . .
33-
34’
’

48. Lepronyssoides pereirai. . . . . . . . . . . . . . 34 ’

49. Liponysella madagascafiensis . . . . . . . . . . . 35

50. Hirstesia sterhalis . . . . . . . . . . . . . . . . .
’

35

’
51. Neolaelaps rnagnistigmatus . . . . . . . . . . . . 36
52. Steal onyssus joaquimi . . . . . . . . . . . . . . . 37
’

53. Pellonyssus’ passer! . . . . . . . . . . . . . . . . 38 "

54. Radfordi.el.la oudemansi . . .. . . . . . . . . . . . 39

55. Tur uriis’cutatus’. . . . . . . ... . . . . . . . . . . 40
56.. Allodermanygsus sanguineus . . . . . . . . . . . . 41
57. Dermanyssus galliriae . . .
58. Myonyssu’s ’jamesorii . . . . .
....
. . .
. . .
. . . . . . .
. . . . . . .
41
42
’

’
60. Haem’ogamas’us keegani . ...
59. Eulaelaps stabolaris . . . . . . . . .
. . .
61. Brevisterna morlani . . . . . . . . .
. . . . . .’.
. . . . . . .
. . . ... .
43
44
45
’

62. Ischyropod’a armatus. . . . . . . . . . . . . . . . 46 ’

63. Spelaeorhynchus praecursor . . . . . . . . . . ...46.

’

64. Hemilaelaps’ tanneri . . . . . . . . . . . . . . . .

’

65. Raillietia auris . . . . . . . . . . . . . . ..... . .

47
48
66. Ixodorhynchus liponys spides . . . . .

..
67. Manitherionyssus- heterotarsus . . .
68. Dasyporiyssus’ neivai . . . . . . .
. . , ...
...
. . . ..
. . . .,. . .
....
.
48
49
50
’
’
69. Entonyssus heterodontus.. . . . . . . . . . . .., . 50
’
70. Halarachne aftiericana . . . . . . . . . . . . . , .’ 51
71. Orthohalarachne attenuata . . . . . . . . . . . . . 51
72. Pneumonyssus simieola . . . . . . . . . . . . . . 52
73. RhinopRaga p’apionis . , . . . . . . . . . . ... . .’ 52
74. Pneurnonyssoides caninum.. . ..
. . . . . . . . . 53
75. Rhinonyssus coniventpis . . . . . . . . . . . . . . 54
76. Larmyssus orblcularis . . . . . . . . . . . . . . . 54
77. Ratlinyssus caudistignaus . . . . . . . . . . . . . 55
’
78. Neon’yssus zenaidurae . . . . , , . . , . . . . . . 56
-
79. RhinoeciuS co-oremani . . . . . . . . . . . . . . . 57
80. Sternostbrha boydi . .. . . . . . . . . . . . . . . . 58
81. Cas angrensis . . . . . . . . . . . . . . . . . . . 59
82. Ptilonyssoides triscutatus . . . . . . . . . . . . . 60
83. Tinamiriyfesus trappi . . . . . . . . . . . . . . . . 61
84. Flavionyssus rabelloi . . . . . . . . . . . . . . . 61
Fig. 85. Rochanyssus werneri . . . . . . . . . . . . . . . . 61
Pig. 86. Travanyssus paranensia . . . . . . . . . . . . . . 61
Fig. 87. Bhinonyssoides donatoi. . . . . . . . . . . . . . . 62
Fig. 88. Vitznyssus sp. . . . . . . . . . . . . . . . . . . . 63
Fig. 89. Paraneonyssus hirsti . . . . . . . . . . . . . . . . 64
Fig. 90. Ptilonyssus nudus . . . . . . . . . . . . . . . . . 65
Fig. 91. Spinturnix iiiyoti . . . . . . . . . . . . . . . . . . 66
Fig. 92. Periglischrus strandtmanni . . . . . . . . . . . . 67
Fig. 93. Acaathochela chilensia . . . . . . . . . . . . . . . 68
Fig. 94. Alphalaelaps aplodontiae. . . . . . . . . . . . . . 68
Pig. 95. Myonyssoides spiaosus . . . . . . . . . . . . . . . 69
Pig. 96. Ugandolaelaps protoxera. . . . . . . . . . . . . . 69
 INTRODUCTION

The order Acarina is made up of two distinct groups of arachnids

that share a number of characteristics that distinguish them from other
orders of the class Arachnida. Of these characteristics, the most im-
portant is the complete obliteration of the separation of the cephalo-
thorax and abdomen from each other. Another important characteristic
is thegnathosomaor the section of the body that bears the mouthparts,
chelicerae and pedipalps. In the Acarina the gnathosoma is usually set
off from the remainder of the body or idiosoma. Because in many aca-
rines, such as the ticks, it appears to be headlike, the gnathosoma has
been frequently called the capitulum. It is, however, more comparable
to the mouth than the head since it contains the pharynx, mouth and
moutbparts but never the brain or eyes. Acarina are smaller than other
arachnids, but some species and groups of acarines such as the ticks
are characteristically larger than the pseudoscorpions, and some spi-
ders are smaller than some of the acarines. The section of the Acarina
that includes the mesostigmatid mites is characterized by having lateral
openings or stigmata for the respiratory system. The respiratory sys-
tem of the other section has anterior openings or lacks primary res-
piratory organs altogether.
The suborder Mesostigmata is one of three suborders whose repre-
sentatives have lateral stigmata. The Onychopalpida have more than a
single pair of stigmata and are larger than most species of the Meso-
stigmata. The Ixodides or ticks are also large. They have a single
pair of lateral stigmata but these are posterior to coxa IV and are sur-
rounded by the peritreme, whereas the mesostigmatid peritreme is elon-
gate and sinuous. Ixodides also have a hypostome bearing recurved
barbs, at least in the immature stages. Mesostigmatid mites are unique
in that the gnathosoma is composed of a basal ring that encloses the
chelicerae dorsally. A revision of the suborder was made by Camin
and Gorirossi in 1955. Their classification follows:

Suborder Mesostigmata
Supercohort Monogynaspida Camin and Gorirossi. 1955
Cohort Liroaspina Tragardh. 1946
Superfamily Liroaspoidea Tragardh, 1946
Superfamily Microgynioidea Tragardh, 1942
Cohort Gamasina Leach, 1815
Superfamily Zerconoldea Tragardh. 1944
Superfamily Parasitoidea Banks, 1915
Cohort Uropodina Kramer, 1881
Superfamily Uropodoidea Kramer, 1881
Superfamily Trachytoidea Tragardh, 1937
Superfamily Diarthrophalloidea Tragardh, 1946
 Supercohort Trigynaspida Camin and Gorirossi, 1955
Cohort Antennophorina Camin and Gorirossi, 1955
Superfainily Celaenopsoidea Tragardh, 1938
Superfamily Megisthanoidea Tragardh, 1943
Superfainily Antennophoroidea Camin and Gorirossi, 1955
Superfamily Fedrizzioidea Tragardh, 1937
Cohort Cercomegistina Camin and Gorirossi, 1955
Supertamily Cercomegistoidea Camin and Gorirossi, 1955

Many of the species of the suborder Mesostigmata are parasitic in

the sense that they live on or in larger animals and use the body fluids
or tissues of their hosts as food. Those that parasitize vertebrates are
of two main sorts: they are either internal parasites of the respiratory
passages or they are ectoparasitic nidlcoles that reproduce in the nests
of their hosts. Those that are internal parasites reproduce in the host.
Those that live in the nests of their hosts breed in the nest and go to
the host only tor food. With one exception the mesostigmatids that para-
sitize vertebrates belong to one or another of twelve families that are
grouped by Camin and Gorirossi (1955) in the superfamily Parasitoidea,
cohort Gamasina. The exception belongs to the cohort Antennophorina.
External parasites belong to the families Spelaeorhynchidae, Spinturn-
icidae. Haemogamasidae, Heterozerconidae, Paramegistidae, Der-
manyssidae, Laelaptidae. Ixodorhynchidae and Dasyponyssidae. The
Haemogamasidae, Dermanyssidae and Laelaptidae are by far the most
important families in numbers of species and economic and medical
importance. Spelaeorhynchids and spinturnicids are parasites of bats,
ixodorhynchids and heterozerconids of snakes, and dasyponyssids of
edentates. Internal parasites belong to the families Raillietidae, Hal-
arachnidae, Rhinonyssidae. and Entonyssidae. Only two species of
Raillietidae are known and these are found in the external auditory me-
atus of cattle and antelope. Representatives of the Halarachnidae, Rhi-
nonyssidae and Entonyssidae are parasites of the respiratory passages
of mammals, birds and snakes respectively.
Three volumes with excellent but limited coverage of parasitic meso-
stigmatids have recently appeared and are well worth consulting. The
first, Acarina ot the Rodent Fauna of .the USSR (1955), deals with the
mites associated with rodents in Russia. The portion dealing with the
parasitic Parasitoidea, by Bregetova and others, covers fully the spe-
cies occurring in Russia of the three most important families: Laelap-
tidae, Dermanyssidae and Haemogamasidae. Written in Russian, it in-
cludes keys to genera and species and has excellent illustrations. The
second recent volume, also in Russian, is Gamasjd Mites (Gamasoidea)
by N. G. Bregetova (1956), The third volume. A; Manual of the Para-
sitic Mites a! Medical w^ Economic Importance by E. W. Baker and
others (1956), gives a good discussion and excellent illustrations of the
more common parasitic mites.
A brief summary of the medically important Parasitotdea is given
by Fuller (1956).
 Medical and Economic Importance

Ectoparasitic mesostigmatid mites may harm their hosts in three

ways. They may transmit a disease; their bites may produce a derma-
titis and do result in the loss of blood. In the case of some dermanys-
sid species, heavy infestations may result in the loss of sufficient blood
to cause the death of the host. The Raillietidae invade the ear of cattle
and damage the internal structures. They have been reported inthe
middle and inner ear (Olsen and Bracken, 1950). The pathogenicUy of
the other internal species is less well known. Death of animais has
been attributed to respiratory acariasis (Grzimek, 1951; Gay and Branch,
1927). Nasal mites have been reported as associated v/ith rhinitis and
sinusitis, but no cause and effect relationships have been demonstrated
(Monlux, 1951). Lung mites, on the other hand, do produce character-
istic lesions, and the mites themselves are frequently encapsulated by
host tissue. Canaries sometimes suffer from asthmalike conditions
attributed to internal mites.
Parasitic mesostignaatid mites have been demonstrated to be capa-
ble of transmitting viral, rickettsial, bacterial, protozoan, and helmin-
thic infectious agents to their hosts. The method by which transmission
is accomplished has been demonstrated in only a few instances." It is
probable, however, that viral, rickettsial and bacterial agents are in-
troduced by the bite. Protozoan and helminthic infections are probably
accomplished when the mites are eaten or crushed by their hosts. Few
mites have been investigated for their disease carrying potential. In-
vestigations of mesostigmatid mites as vectors of disease have fre-
quently produced contradictory results. It is therefore difficult to assess
the true significance of their role as vectors. The experience of the
investigations of mites as vectors of encephalitides is somewhat typical.
Dermanyssus gallinae was found to be capable of harboring the vi-
ral agent of St. Louis encephalitis and of transmitting it to chickens
(Smith etal.. 1944, 1945, 1946, 1947, 1948). Following the publica-
tion of these papers it was accepted that D. gallinae played an impor-
tant role in the epidemiology of St. Louis encephalitis. J5. gallinae
was assumed to be the reservoir in which the virus overwintered and a
vector of the virus from bird to bird when mosquitoes were not avail-
able for this purpose. D. gallinae and other similar parasites of chick-
ens and birds were studied in an attempt to determine the role of these
mites in the transmission and maintenance of Western equine enceph-
alitis (WEE) and eastern equine encephalitis (EEE ) as well as St.
Louis encephalitis (SLE). Four recent papers (Reeves et al., 1955;
Chamberlain and Sikes, 1955; Sulkinetal., 1955; Chamberlain et al.,
1957) review previous work and report new findings. Each of the papers
concludes that dermanyssid mites do not play a significant or essential
role in the epidemiology of SLE, EEE. or WEE.
Brody (1936) and Hofstad (1949) respectively have shown that Or-
nithonyssus sylviarum is capable of becoming infected with .Newcastle
disease and fowl poac but infections do not last and the mite is probably
not involved in transmission of the diseases.
Extensive experimentation is lacking but recent work by Nelzina
and Barkov (1951) and Hopla (1951) shows that some mites may become
infected with and transmit the tularemia microbe. Pasteur ella tularensis.
The mites tested by Nelzina and Barkov were Laelaps muris and Hae-
molaelaps rnohrae. Hopla worked with a laboratory culture of Ornith-
onyssus bacoti.
Seddon (1951), reporting on diseases of domestic animals in Aus-
tralia, states that Permanyssus gallinae is an important vector of avian
spirochaetosis.
Several species of external mites are involved in the transmission
of rodent haemogregarmes. Miller (1908) first demonstrated that Echi-
nolaelaps echidninus transmits Hepatozoon from rat to rat. dark (1958)
has shown that Hepatozoon griseisciuri of squirrels may reproduce in
Haemogamasus ambulans and Echinolaelaps echidninus. Transmission
probably occurs when the mites are eaten by the squirrel,
There is evidence that a similar epidemiology occurs in some mites
and trypanosomal diseases of small mammals (Macfie and Thomson,
1929; Walton, Ph. D» dissertation. University of Maryland, 1956).
Camin (1948) proved the transmission of haemorrhagic septicemia
(Pseudomonas hydrophilus) by Ophionyssus natricis from snake to snake
via the bite of the mite. Recently Camin (verbal communication) has
found evidence that the same mite harbors and transmits a haemogreg-
arine of snakes.
The surprising discovery that a mite was the intermediate host of
a parasitic nematode was made by Williams and Brown (1945) who first
demonstrated this in Ornithonyssus bacoti. Their findings have been
verified by several investigators, including Bertram etal. (1946) and
Scott (1948). To date, no other helminths are known to be transmitted
by mesostigmatid mites, but the possibilities have net been explored.
Many attempts have been made tosfaow transmission of murine ty-
phus rickettsiae from animal to animal, or from animal to man via 0.
bacoti and Laelaps spp. A few workers have reported positive results,
but the work of many others is overwhelmingly negative,
Rickettsialpox is beyond a doubt transmitted from animal to animal
and to man through the house mouse mite, Allodermanyssus sanguineus.
Huebner etal. (1946) and Philip and Hughes (1948) demonstrated the
ability of Ornithonyssus bacoti to transmit this rickettsia under experi-
mental conditions. Zdrodovski and Golinevitch (1953) showed that the
rickettsia is transovarially transmitted in the house mouse mite,
Mites crawling over the body ox a person may produce various de-
grees of discomfort. When very abundant, any species may cause an
itch or an allergic condition, but except for the dermanyssid mites,
none of the Parasitoidea are known to bite man through the undamaged
skin. Of the true biters, the most notorious is Orntthonyssus bacoti.
It bites man readily and causes a very noticeable dermatitis. Several
species of laelaptid mites have been found capable of feeding on abraded
human skin (Wharton and Cross, 1956). Since few people have oom-
pletely unbroken skins, exposure to large numbers of these mites may
result in biting.
With the exception of the ability of dermanyssid mites to produce
dermatitis, and the transmission of rickettsialpox by A. sanguineus.
the parasitic mesostigmatid mites can be considered as only of poten-
tial medical importance. In this connection reports of the recovery of
the etiological agent of haemorrhagic fever from laelaptid mites by Ki-
tano and Kasahara is extremely important (quoted by Traub, et al.
1954).
Relatively few species of parasitic mesostigmatids are now recog-
nized as economically important. Each of the species will be discussed
separately. The relatively few species that are known to produce finan-
cial loss because of their habit of parasitizing vertebrates is probably
in part due to a lack of knowledge. The few species that are important
cause considerable damage to poultry and are frequently annoying pests
of man in his home or in commercial establishments,

Control

A few general control measures against parasitic mesostigmatid

mites are given below. For more detailed information, the reader is
referred to the Manual of Parasitic Mites by Baker et al. (1956).
Mites m Human Habitations. As far as direct attack on man is
concerned, none of the mites in this manual bite human beings by choice.
Control, then, is essentially a matter of killing most of the mites, erad-
icatingthe normal hosts from the premises and then destroying the mites
that remain. It is important to kill most of the mites before elimina-
ting their hosts because, in the absence of a preferred host, man will
be attacked more readily. Thus, effective control can be obtained with
any good miticide without regard to its residual effect. We have per -
sonally obtained good results against Ornithonyssus bacoti in homes
with the use of a .pyrethrum-DDT bomb. Presumably rotenone or py-
rethrum should both be effective. Other recommendations are: spray-
ing the floors, furniture, etc. with kerosene (Herms, 1939); pyreth-
rum or DDT with kerosene (Lamb, 1952).
Presumably the same treatment would be effective against any mite
invading human habitation.
Poultry Mites. The mites attacking poultry and other domestic
birds are Dermanyssus gallinae, Ornithonyssus bursa and Ornithonys-
sus sylviarum. Habits of the first two are somewhat similar and con-
trol that is effective against one will have an effect on the other. 0.
sylviarum spends much more time on the birds so that special control
measures must be instituted for it as indicated in the next. paragraph.
Since these mites inhabit the nesting material or the cracks of the walls
and roosts, it is not possible to overemphasize that attention should be
given to these places. Infested houses can be sprayed with kerosene or
a mixture of kerosene and crankcase oil, also kerosene and creosote
(five gallons to one pint). Apparently the chlorinated hydrocarbons are
not so effective.
The birds themselves, especially those infested with Ornithonyssus
sylviarum, may be successfully treated by spraying with nicotine sul-
phate or dusting with powdered sulfur. Nesting material of setting hens
can be treated with nicotine sulphate. Purman (1953) recommends. for
the control of the northern fowl mite, 0. sylviarum, during mild weath-
er, a powder-spray application directly on the birds, using a one per
cent aqueous suspension of R-242 + Triton X-100, one gallon per 35 - 50
birds. In very cold weather, he recommends dusts alone, either 10 per
cent R-242 dust or 10 per cent bis(p-chlorophenoxy) methane dust ap-
plied by hand. He also suggests painting the roosts with a mixture of
equal parts of 40 per cent nicotine sulphate anddiesel oil. Aramite (Beta-
chloroethyl-beta-(paratertiary-butyl-phenoxy)-alpha-methyl-ethyl sul-
fite] was found effective but blistered the skins of the birds. Ten per
cent Toxaphene. 0.5 - 1 per cent lindane, and 2.5 - 5 per ceat Chlor-
dane appeared to be toxic to chicks.
Wild birds, especially the English sparrow, starling and the com-
mon pigeon, also harbor these mites and their nests should not be tol-
erated around hen houses.
Snake Mites. Control of the snake mite, Ophionyssus natricis is
of concern primarily to zoos, reptile houses and professional and ama-
teur herpetologists. Pyrethrum and sulphenone dusts are recommend-
ed, but best control is obtained by frequent and thorough cleaning of
quarters, steam-bathing cages and grounds, and providing contaiflera
of water large enough to allow the reptiles to submerge completely.
Internal Mites. Canaries are sometimes so heavily infested with
the respiratory mite, Sternostoma tracheacolum, that they suffer ill
effects. Stephan and Kaschula (1950) recommend the use of Double
Benhex Powder (DDT and benzene hexachloride) as follows. Place the
birds in a cage, cover with linen, pump in a mist; after five minutes,
air the cage. Kepeat at fourteen day intervals. Some care should be
exercised as Double Benhex Powder is toxic to birds after prolonged
exposure.
Other cage birds and also ducks and geese (but apparently not the
galliforme birds such as pheasants and chickens) are frequently infested
with nasal mites but rarely ever seemto be seriously affected by them.
Dogs are also known to be infested with a respiratory mite, Pneu-
monyssus caninum. No treatment is known tor this infestation,
The Effect of DDT Under Natural Conditions. During the middle
and late 1940’s, the U. S. Public Health Service conducted an intensive
campaign against murine typhus throughout the gulf coastal states.. Part
of this program was aimed at reducing the number of rat ectoparasites
by dusting rat runs and rat harborages with DDT powder. In southeast
Georgia, where this program was conducted under controlled conditions,
the number of Ornithonyssus bacoti was greatly reduced in dusted areas
but remained about the same in undusted areas ( Morlan and Utterback,
1952). No results were reported for other species of mites. In Gal-
veston, Texas, however, there was, no apparent difference before and
after dusting in the numbers of either Ornithonyssus bacpti or Echino-
laelaps echidninus. It should be noted that there were no undusted
control areas in the Galveston campaign; however, fleas were drasti-
cally reduced, just as they were in the dusted areas of southeast Geor-
gia. Although the Galveston results remain unexplained. DDT dust ap-
plied liberally to areas of high mite concentration has given good con-
trol.

BEHAVIOR

The behavior of mesostigmatid mites is known in a general way by

all who have dealt with living specimens, but controlled experiments
are rare. In fact, there is only one published record of extensive work
on behavior and that is Camin’s (1953) fine paper on the snake mite.
Ophionyssus natricis. All other observations are casual and desul-
tory.
Behavior patterns of mites are Interpreted mainly on the basis of
movement and rate of feeding. Whether the behavior of mites can be
conditioned is not known and as far as we know, no one has tried to find
out. The discussion on sensory behavior which follows is drawn almost
entirely from Camin, supplemented by a few observations made by others .
Heat. Camin measured the temperature preferences of the snake
mite by use of a glass tube 100 cm long and one cm in diameter. This
tube was laid in a metal trough filled with water and sand. Ice was
packed in one end and heat applied to the other, providing a temperature
gradient of 9.0to49C. Mites were introduced into the tube through
openings in the side. Within 20 minutes the vast majority of the mites
were congregated in the 20 - 23 range. Camin also found that within
this range the mite would travel in a straight line, but both above and be-
low these temperatures it immediately began to turn and weave.
Adult unfed females became akinetic at six degrees Centigrade while
fed females ceased movement at two degrees Centigrade. All other
stages stopped movement at 10C, Rate of movement increased directly
with increase in temperature till it reached its peak at 30- 40C. Above
40C speed greatly decreased and between 45 - 50C all mites were
akinetic. Death in five seconds occurred between 50 - 55C.
One might assume that mites of mammals would prefer warmer
temperatures than snake mites. To test this hypothesis, Virginia Riggs
of Texas Technological College (unpublished data) used the copperplate
from a slide drying oven, measuring 24 inches by six inches. A sheet
of white paper was placed on the plate and ringed with Baume’s Ben Gay
to confine the mites. Heat was applied to one end with a heating pad, the
other end was cooled with an ice pack. This provided atemperature grad-
ient from about 15 to 40C. The spiny rat mite, E^hmclaelaps eghid-
ninus, was tested. Individuals wandered freely within thia range bat with-
in half an hour all of them had come to rest between 23 - 35C. Within
the latter range there seemed to be no particular preference. These
mites (about 40 per test) were picked at random from the colony. No
attempt has yet been made to see if starved mites prefer the higher
temperatures.
Bertram et al, (1946) used heat to drive protonymphs of Ornitho-
nyssus bacoti through No. 8 silk cloth to separate them from the females.
The temperature was not stated but it was enough to kill the females.
Cross (1955) found that dermanyssids (0. burs a, 0^. bacoti, D. gal-
linae) and laelaptids (E, echidninus. Laelaps nuttali) fed better and more
consistently at higher temperatures than they did at lower temperatures.
Humidity. As a rule it is probably safe to say that the parasitic
Mesostigmata do best at relative humidities of 50 per cent or higher.
Camin soaked pieces of filter paper in various concentrations of
KOH so as to supply relative humidities of 20, 35, 50, 65, 85. 90, and
95 per cent. These were placed two at a time in a sandwich box, cov-
ered with organdy cloth, and 50 mites were introduced per test. By
this method it was shown that 100 per cent of the females of the snake
mite prefer 95 per cent relative humidity. The other stages were not
quite so responsive to 95 per cent although a majority were always in
that region. This is perhaps a natural corollary to the fact that the. eggs
hatch best in a 95 per cent relative humidity.
Camin found that the mite would come to rest in a short time at 95
per cent relative humidity. At higher or lower humidities it wandered
about and took longer to rest. The time required to come to rest in-
creased with a decrease in relative humidity to 20 per cent. At that
relative humidity the mites did not become akinetic.
Skaliy and Hayes (1949), working with Ornithonyssus bacoti, ob-
served that the highest percentage of eggs developed into adults at 25C
and 47 per cent relative humidity.
Bertram etal. (1946) noted that mites trapped in moisture of con-
densation would die quickly but also noted that if they floated free on the
surface of water they would survive as long as five days.
Wood (1917) found that the chicken mite, Dermanyssus gallinae,
would do very well at a relative humidity of 20 per cent. Sikes and
-
Chamberlain (1954) reared their cultures of bird mites at 60 70 per
cent relative humidity.
Cross (1955) found that low relative humidity was important in the
feeding of mites. Both dermanyssid and laelaptid mites were found to
feed better and more consistently at low rather than high relative hu-
midities.
Odor. Camin tested olfactory perception by drawing two currents
of air through a T-tube; one current from the open room and one from a
box containing a live snake. Hungry mites placed in the bottom of the
T-tube invariably crawled up that branch of the tube through which air
from the snake box was coming. It had to be a living snake, however.
They would not respond to the odor of snake skins, dead snakes, or
snake blood. The sense of smell was lost if the tips of tarsi I were re-
moved.
 The odor of a snake is, however, not a necessary stimulus for feed-
ing, as Camin observed that the mites apparently fed equally well on
frog and serpent blood, and also on man, if the ambient temperature of
the skin was lowered to 15C. It is only fair to add that. in the latter
case, the mites invariably died.
Odor sensitivity has not been tested for any other mites. Although
they may respond to the odor of the host, it is nevertheless true that
host odor is not at all; necessary to elicit a feeding response. Cross
(1954) and Cross and Wharton (1954) have shown that Ornithonyssus ba-
coti, 0. bursa, 0. sylviarum, Dermanyssus gallinae, JSchinolaelaps
echidninus, and Haemolaelaps glasgowi will take blood of various birds
and mammals through a silk cloth, and thrive on it. Owen (unpublished
data) of Texas Technological College has shown that Brevisterna ft. sp.
will also take blood in this manner. Allred and Marchette (1957) have
fed Brevisterna utahensis directly from the tip of small glass tubes.
Gravity. Apparently most species of mites will travel upward when
given a choice. Unfed protonymphs of the snake mite, the rat mite, and
several species of bird mites will ascend an obstacle rather than go
around it. When confronted with a flat surface, tilted at various angles
up to 90. most, but not all mites will travel up. Those that do ascend
generally wander about at the top and finally come to rest and do not
move again unless disturbed by an air current, light, or warmth.
Contact. In the case of the snake mite, Camin observed that con-
tact with the anterior dorsal region was necessary to elicit a -feeding
response. The mite would not feed unless it could crawl under a scale.
If the scale under which it was feeding was pulled forward, or other-
wise moved so it no longer made contact with the mite. the mite soon
detached itself and moved about till it found another scale to get under.
Such pronounced thigmotropism may exist for other species but there
is no experimental evidence for it.
Many species, especially among the Dermanyssidae, tend to form
clusters, apparently in response to contact. Camin (1953) found that
the snake mite moved more slowly over rough surfaces than over smooth
surfaces. Skaliy and Hayes (1949) observed that Ornithonyssus bacoti
preferred the roughened surface of filter paper for oviposition, rather
than the smooth sides of a glass tube.
Light. Although eyes are lacking in all Mesostigmata, many spe-
cies are nevertheless sensitive to light. The snake mite, when ex-
posed to diffused overhead light, will remain kinetic but in the dark it
soon becomes quiescent. If placed in a box, one half of which is darkened,
the mite moves in a straight line in the darker half, but as soon as it
reaches the lighted area it turns abruptly and is soon again in the dark.
When exposed to a horizontal beam of light, the mite moves away from
the source. This is especially true of the engorged females. The other
stages show a less marked revulsion to light.
In the snake mite the light sensitive organs are on the ambulacrum
of tarsus I. This is discussed more fully under "Special Senses" (p.21).
The chicken mite, Dermanyssus gallinae, is obviously repelled by
10

light. Normally it feeds only at night or in the dark. Starved individuals

will feed in the light but reluctantly so. ^_. bacoti. and 0. bursa feed as
well in the light as in the dark, but E. echidninus feeds more readily in
the dark (Cross. 1955).
Combinations of Stimuli. Camin (1953) tested several combinations
of stimuli to see which were stronger. To test the relative strength of
heat and light stimuli, a strong beam of light (150 foot candles) was
directedtoward a hot plate. When placed in this beam. the mites always
moved directly away from the source of light and to the hot plate, until
they reached the temperature range of 40 - 45C when light no longer
drove them forward. Here they wandered about very actively until they
chanced to get, out of the light beam. A few mites, however, failed to
be stopped by heat and continued away from the light source and walked
to their death under the hot plate. In general, then, tfe light sttmnlus
is dominant to heat up to 40C.
Response to gravity was dominant to red light or diffused white
light, but when a concentrated beam was directed at them they would
not walk up a tilted board into it.
In testing reactions of the snake mite to combinations of light, con-
tact, and relative humidities, it was found that clump formation occurred
only in the dark; so that light is apparently dominant over contact. Light
also was dominant over response to humidity because the mites would
congregate in the darkened area regardless of whether the relative hu-
midity was 20 per cent or 95 per cent.

LIFE CYCLES

Apparently the basic cycle consists of egg, hexapod larva, proto-

nymph, deutonymph and female and male, but this occurs only in some
species. Except in the Dermanyssidae, most species are larviparous
and occasional species even nymphiparous. They are, however, ovo-
viviparous, not truly viviparous. In-one family, the Raillietid&e, only
larvae and adults have been reported. :
Detailed life history studies are available for five species of mites:
Ornithonyssus bacoti, Dermanyssus gallinae, Allodermanyssus san-
guineus, Ophionyssus natricis and Echinolaelaps echidninus. There are
partial studies of Ornithonyssus sylviarum, Ornithonyssus bursa. Lac-
laps nuttaU, Pellonyssus passeri, Haemolaelaps glasgowi, and Brevj.-
sterna utahensis.

Dermanyssidae

Ornithonyssus bacoti. Many investigators have studied Ornitho-

nyssus bacoti. including Bertram et al. (1946), Ohmori (1935-1936) (as
jL. nagayoi, a synonym of 0. bacoti), Ovazza (1950), Williams (1946),
and Skaliy and Hayes (1949). The papers of Bertram et al. and of Oh-
mori are quite complete bat the most ’thorough study is that.of Skaliy
 11

and Hayes. There is essential agreement in the findings of all these

workers.
-
Eggs are laid within 2 10 days’.. after the female has fed to reple-
tion. From 1 - 20 eggs are laid over a period of 24 - 36 hours. After
this, another blood meal is taken and another batch of eggs is produced.
According to Skaliy and Hayes (1949) the number of eggs produced per
female varies from 8 - 141, with an average of 98.8 per female. The
mean life span of the female is 61 days. Ohmori (1935) found that the
number of eggs per female averaged 115 - 128, that they were laid at
intervals of two hours, and that survival time for females was only 22 -
28 days.
The eggs hatch into six. legged larvae in 1 - 4 days, generally
within 36 hours. The larvae molt to’protonymphs, without feeding,
within 24 hours. The protonymph is an active feeding stage and will
seek a blood meal within 12 - 24 hours after having molted. If a meal
is immediately tprthcoming it may molt to the deutonymph within 24 -
36 hours. However, if food is denied, the protonymph is capable of
surviving from $ .- .23 days and, in one observed case. a protonymph
lived for 43 days. .liven when fully fed it may not molt for several days.
Observed intervals range from 3 - 28 days, with the majority molting
between five and 1.4. days.
The deutoriymphs ofO. bacofi, and presumably of all Orniithpnyasus
species, as well as.pf many other related genera, including Ophionyssus,
do not feed. Although more sluggish than the protonymph, the deuto-
nymph is not an inactive stage and is not to be compared with the qui-
escent nymphochrysalis of trombidiform mites. It molts to the adult
within 24 - 48. hours. There seems to be no morphological difference
in pre-male and pre-female nymphs except size; the females on the
average are larger.
Adults will mate within 24 hours after reaching maturity and perhaps
at intervals thereafter." Camin (1953) found that Ophionyssus natricis
will mate before feeding, and according to Nelzina (1951) Ornithonyssus
bacoti behaves similarly. Regardless of whether or not they are im-
pregnated, the female will’feed .and lay eggs. Parthenogenetic eggs
develop invariably into .males, whereas a fertilised female will produce
both males and females. However, it is not known whether male off-
spring from mated female?, re suit from fertilized or unfertilized eggs .
Skaliy and. Hayes found that the shortest egg to adult cycle is seven
days and the shortest egg to egg cycle is 13 days.
’Ornithonyssus bursa and j3. sylyjarum. According to Sikes and
Chamberlain (1954), eggs of Ornithonyssus bursa hatched in 1. 5 - 2
-
days at room conditions (approximately 80 - 83F and 60 70 per cent
relative humidity) and within another 24 hours all had become proto-
nymphs. Protonymphs required at least two feedings to become neplete
-
and this generally took at least 36 48 hours. After complete engorge -
-
ment. molting to the .deutonymphs occurred in 24 36 hours and from
this to the adult required about another 24 hours. Adult females required
about two days and at least two feedings to become fully engorged. They
 12

averaged 2 ,- 5 eggs after a complete feeding. The entire cycle from the
initial feeding of the female to another generation of females can be
accomplished in as little as 5 - 7 days. No differences were noted in
the life cycles of Ornithonyssus bursa and Ornithonyssus sylvia.rum.
0. sylviarum is smaller in ..ail} stages; this .is most marked in adults.
Dermanyssus gallinae. Wood(191"7) gives an excellent account of
the life cycle of this common fowl mite. At a mean temperature between
75F and 85P the egg to egg cycle may be accomplished in as little as
nine days. This has been verified by Wisseman and Sulkin (1947) and
Sike sand Chamberlain (1954). The average egg to egg interval is around
12 - 13 days. The stages, as in Ornithonyssus and Ophionyssus, con-
sist of egg, larva, protonympb. deutonymph and adult. Unlike these two
genera, the deutonymph of Permanyssus is an active stage and avidly
sucks blood. The adults will mate either before or after taking blood
..and probably several times. Wood reported ari average of about 40 eggs
per female laid in batches of 1 - 8 eggs through nine layings, each lay-
ing being preceded by a blood meal. Wisserriah’and Sulkin (1947) con-
, curred with Wood’s findings and agreed that’ in nature probably more
eggs than this ’.’ "
’’.

’
are produced.
AUodermanyssus sanguineus. Fuller ’(.19’54) obtained life history
data of this important vector of rickettsialp’ox. He found’the life’stages
to be: egg, larva, protonymph, deutonymph’.. and adult male anAfemale,
The larva does not feed, but the protonymph and deutonymph each teed
one’e’ between molts. At a temperature of 24" - 25C and an approxi-
mate relative humidity of 80 per cent the complete life cycle ranged
from 17 to 23 days, viz.: egg, 4" 5 days; larva, three days; protonymph,
i - 5 days; deutonymph, 6 - 10 days. Both sexes ingested xnuBaqpie blood
meals and survived’on an average from 51 to 63 days. The’post-T.arval
-
stages were fed by placing a single mite wi’th a 6 8 day old suckling
.white mouse in a cotton-plugged vial.
., .. Ophionyssus nat’ricis. The life cycle of ttie snake mite was beauti-
.
;,
’fully worked out by’Camin (1.953). The stages agree exactly ^with Ophio-
riyssus bacoti but feeding is much different. Whereas 0. bacoij .’attaches
and engorges within minutes, Ophionyssus natricis may take weeks to
become fully engorged. Unless disturbed, the mite remains attached
at one point until it is completely filled. The total number of eggs pro-
duced average about 80 per female. The eggs hatch in about 24 hours
at temperatures between 25 - 30C. The larvae molt to protonymphs
in 14 -30 hours. The highest per cent of moltings occur at relative
humidities between 75 per cent and 95 per cent. The protonymph en-
gorges completely in 3’- 7 days at 25C but may take as long as 6 - 21
days at 15C. After" repletion, ecdysis occurs in 12 to 48 hours, The
deutonymph, a non-feeding stage, lasts only 13 houys at 30C, but as
.’.long as two days at 15C.
’

The male is ready to mate within a few hours after molting. It

may feed, but only for one or two days. It will mate with unfed or’par-
tially fed females but not with fully engorged females.
-
The female becomes replete in 5 8 days at 25C, It will lay from
 13

-
15 22 eggs after a single feeding and will total as many as 80 eggs in
its lifetime, which is approximately 35 days. The egg to egg cycle under
optimum conditions <25C, 95 per cent relative humidity) is from 13 "
19 days. This mite, therefore, requires a lower temperature butbigber
relative humidity than the species of Ornithonyssus.

Laelaptidae

Echinolaelaps echidninus. Employing the method of Cross (1954),

of feeding mites on drawn blood through silk bolting cloth, Owen (1956)
has worked out the life history of the spiny rat naite.
The female is larviparous. Within 24 hours after birth the larva
molts, without feeding, into the protonymph stage. This stage will take
a blood meal within a few hours and then molt within 4 - 8 days (average a-
five days) into the deutonymph stage. The deutonymph also will feed
within a few hours and likewise takes 4 - 8 days to change to the adult
form. The adult female will feed shortly after molting and begin to
produce larvae parthenogenetically in about five days but will mate readily
if given the opportunity. The female probably produces a larva every
two or three days thereafter. More exact data is lacking; the female
will eat her young if she comes in contact with it. so it was never cer-
tain whether the lack of a larva over a period of time was because none
had been born or because one had been born and immediately eaten. An
effort was made to see whether these mites would eat other small arth-
ropods. Animals tried were protonymphs and deutonymphs of the same
species (Echinolaelaps echidninus), adults of other, smaller gamasid
mites, and larvae and eggs of Prosophila. The female _E_. echidninus
refused all of these. Why they will eat their own larvae is not yet known.
Feeding studies of a mite having similar habits. Haemogamasus ambu-
lans, have demonstrated that this nidicole feeds on many things other
than the blood of its hosts (Furman. 1957).
Laelaps nuttalli and Haemolaelaps glasgowi. Details of the life
histories of these two species are lacking. However, they have been,
and are being reared in laboratories and therefore are mentioned here .
Apparently they have a life history very similar to that of Echinolaelaps
echidninus.

Mating and Insemination

The mode of coition is apparently similar in all the Gamasina. The

male approaches the female either from the rear or the side and slips
underneath her, so that their ventral sides are in apposition and the
anterior ends of both mites are in the same direction. He then lifts the
epigynial plate and dilates the female genital aperature with his chelae.
A clear bubble (the spermatophore) meantime forms at his genital aper-
ture and when this is complete he grasps it with "his chelae and inserts
one end of it into the female atrium. The inserted end then apparently
bursts and the elasticity of the bag, aided perhaps by pressure from the
14

male chelae, forces -the sperm into the female genital tract. After this
is complete the male withdraws the empty sac and cleans his chelicerae.
The male may ride on the back of the female for a ti»e ’after coition,
or for a while before coition if the female is not ready to accept him.

Longevity

Knowing the survival time of mites is obviously of the utmost impor-

tance for ecological, epidemiological, academic, and control purposes.
Yet, almost nothing is known for the vast majority of the mites. Desul-
tory observations show that Laelaptinae species, such as Haemolaelaps
glasgowi, Laelaps nuttalli, and Echinolaelaps echidninus will die within
48 hours in shell vials kept at room temperature and cork stoppered,
regardless of the developmental stage and whether they have fed or not.
However, if they are placed in a relative humidity above 85 per cent it
is possible to keep them alive more than a week. How long they live
under optimum conditions is not known exactly but it is probably two to
three months.
We have kept Rhinonyssidae alive for over two weeks when completely
submerged in either physiological saline or plain tap water and at tem-
peratures ranging from about 10C to 28C. They lived nearly that long
in small glass tubes at a relative humidity of near 100 per cent. Being
soft, internal mites, they naturally are extremely sensitive to moisture
and will die in a matter of hours at relative humidities of 50 per cent or
below. Skaliy and Hayes (1949), working with Qrnithonyssus bacoti.
recorded the survival time of 26 female mites which were allowed to
feed at intervals. Life spans varied from 21 to 90 days for an average
of 62 days. This was at 24 - 26C and 47 per cent relative humidity.
There was evidence to Indicate that at higher humidities the average
life expectancy would have been higher.
Protonymphs of Ornithonyssus bacoti live as long as a month. Qphio-
nyssus natricis nymphs may live up to 60 days. Dermanyssus gallinae
has the longest recorded longevity. Unfed protonymphs live up to 81
days under a variety of conditions; deutonymphs live as long as 139 days;
unfed adults survive up to 96 days and adults which had fed once. up to
108 days. Wisseman and Sulkin (1949) kept Dermanyssus gallinae alive
as long as four months in glass tubes at 4 - 5C. Hence 4 - 5 months
are necessary to starve these mites to death. There seems to be no
record of how long D. gallinae may live when allowed free access to
food (Wood. 1917),
Fuller (1954) found that unfed protonymphs of Allodermanyssus san-
gulneus could live as long as 10 days, and that unfed adults survived
for more than 51 days.
Longevity studies on Echinolaelaps echidninus are in progress at
Texas Technological College. Results to date indicate that unfed larvae
will live more than a week at 30C and 90 per cent relative humidity.
Females kept in vials and allowed to feed at will on drawn blood live
more than 90 days under these conditions.
 15

ANATOMY

’
Segmentation

The mesostigmatid mite is an extremely compact organism. Al-

though evolved from serially segmented ancestors, it has lost all trace
of segmentation except the serially arranged appendages and structures
associated with them. Head, thorax and abdomen are all fused into one.
Internally this is also the case. The nervous system is concentrated
into one large, central mass, and there is no serial arrangement of
muscle, nephridia, gonads, or any other systems. Therefore, to avoid
confusion, arbitrary terms have been given to the obvious body regions.
The palps and mouth parts are borne on a clearly delineated part
known as the gnathosoma. The rest of the body is known as the idiosoma.
The idiosoma is further differentiated into the podosoma and opistho-
soma. The podosoma is the region that bears the legs. In the Meso-
stigmata the legs are all contiguous or at least all equidistant from each
other so that the podosoma is not generally divided into propodosoma
and metapodosoma as it is in the sarcoptiform and trombidiform mites
(in which the first two pairs of legs may be clearly separated from the
last two).
Occasionally species of mesostigmatid mites may exhibit a con-
striction between the podosoma and opisthosoma but this is not an indi-
cation of primary segmentation.

External Anatomy
Gnathosoma. The gnathosoma is a cylindrical body attached sub-
terminally to the anterior end of the idiosoma and lying in the same plane
as the idiosoma. A segmented and movable palp arises from each side.
The region basad of these palps is known as the gnathosomai. base ring
(gbr. Fig. 1). This base ring encloses the chelicerae and includes the
fused palpal coxae, so that the moveable segments of the palp are: tro-
chsmter, femur, genu, tibia, and tarsus.
Extending forward from the base ring on the dorsal side is a mem-
brane of characteristic size and shape, the tectum (Fig. 2).
Distad of the base ring on the ventral side is the hypostome and its
associated structures (h. Fig. 1). The nypostome normally bears three
pairs of setae, known as hypostomal setae. These three plus the gnath-
osomai setae on the base ring make up the normal quota of four pairs
of setae on the ventral side of the gnathosoma. This number is fre-
quently less in the internal mites.
A median longitudinal, ventral band, or shallow groove, from the
base of the gnathosoma to the hypostome,is called the deutosternum (d.
Fig. 1). The deutosternum contains small structures called deutostemal
teeth (dst)Fig. 1). These may be arranged in single file or in combs.
16

The forward continuation of this groove is the protosternum (p, Fig. 1).
A pair of toothlike or hornshaped structures, the corniculi, termi-
nate the hypostome (c. Fig. 1). They are well developed in the Laelap-
tidae and Ixodorhynchidae but very poorly developed or lacking in the )
Dermanyssldae and the internal mites.
Projecting forward from the protosternum is a divided, serrate
structure, the hypopharyngeal process (hpp. Fig, 1). Ttv,s structure is
variable in size and shape and is rather difficult to see. -Dorsal to this
structure is a ciliated, generally pointed body, the epipharynx (e,,Fig. 1).
A pair of long, slender, very poorly, sclerotized structures are placed
between the hypo.stome and the palps. These are the salivary stylets
.(ss. Fig. 1). They are generally impossible to see because .they fold in
and lie behind the. epipharynx. In Ophiomegistus they are lacking.
Idiosoma. The idipsoma of the parasitic Gamasides AS generally
oval or elliptic in outline but it may also be almost circular, (Eubrachy-
laelaps) or conical (Rhinonyssus coniventris). It is never brpaderthan
long. The dorsum.is always strongly,,convex,:the venter only slightly
so. There is no sharp .
demarcation late rally, between dorsal and ven-
. -
.
.

tral sides.
^ ^
The coxae are all contiguous, with the occasional exception of coxa
I which may be slightly separated from the others.. ,Coxa I is cylindrical
and as long as or longer than wide. Coxae II.. Ill,, and IV are cup shaped
and wider than deep. The trochanter is attached to the coxa at-two ful-
cral points; mid-ventral and mid-do.raal, so that the leg moves-only
forward and backw,ard.qn the coxa. Up and down movements occur from
the trochanter out.
With the exception of Ophiomegistus, coxa I always has two setae,
both in the mid-ventral region, one apical and the other basal, but neither
one is on the extreme end. Coxa I of Ophiomegistus has three ventral
setae. Coxae II and III always have ..two setae each, one on the anterior
ventral margin, the other in the mid-posterior-ventral region.: Coxa IV
i

always has only one seta.. .It,is in the mid-ventral region just below the
point of articulation between the coxa and trochanter. Coxa II usually
has a triangular tooth projecting from the anterior margin (Fig. 37).
This is more pronounced in the Dermanyssidaethan in any other group.
Some species have one or more of the coxal setae thickened and spine-
like, and some may have accessory spurs or toothlike structures on the
coxae.
.
A bipartite structure, the tritosternum, (Fig. 3) originates mid-
ventrally between the. gnathospma and the. sternal-plate. This consists
of three parts: a short, undivided basal portion and two ciliated or fim-
briated arms. the lacinae.., In the Dermanyssidae there is always a
hyaline, denticulate membrane on each side of the basal portion. The
tritosternum is reduced to a small platelet in the Spinturnicidae and is
entirely lacking in most of the ..internal mites. Its function is unknown.
Other characteristics of the idiosoma are given under the various life
stages. .. .; .’
.
.
Integument. According to Vitzthum (1941), who quotes from other
 17

investigators, chiefj-y Winkler (1888) and Steding (1926), the integument

of the Mesostigmata consists of a basal hypodermis which secretes the
non-cellular cuticle. Two layers may be recognized in the cuticle, an
outer ectostracum and an inner hypostracum. The hypostracum takes
the ordinary histological stains but the ectostracuro does not.
Mites that are capable of great distention. as Ornithonyssus and
Dermanyssus, have the integument closely striated in the non-fed state
buta^ the body’swells with food, the atriations begin to smooth out. This
indicates that the integument is not necessarily elastic but may be laid
down in tiny folds to allow for expansion.
Various areas of the integument are thickened and sclerotized. form-
ing the very characteristic shields or plates. These are generally pig-
mented, the color varying from pale straw to dark reddish brown. The
parasitic Mesostigmata are never brightly colored with integumental
pigment. They may appear bright red immediately after feeding because
of the blood which they have drawn and which can be seen through the
walls of the gut and body. Most, but not"all, of the endoparasites are
colorless.
The setae are hair-like outgrowths of the integument that.originate
from both the sclerotized and non-sclerotized areas. They are mostly
smooth but they may be slightly pilose. Except for this pilosity, they
are simple in structure and origin; that is, none of them are associated
with special glands or functions. A few setae on the dorsal apex of tar-
sus I are an exception. These seem to be striated and hollow and are
probably more sensitive than the other setae. No one has yet tested
the true sensory nature of these setae, but is has been suggested that
they may be chemoreceptors,
There are many shapes and sizes of setae: short and thick, long
and slender, sharp, blunt, stiff, flattened, flexible, serrate, pilose,
and inflated setae. What their function may be is not known, but they
are probably tactile.
Some parts of the body always have certain setae; thus there are
always three pairs of setae associated with the female sternal plate, two
setae on each of the first three coxae and one on coxa IV. and so on,
Dorsallythe number may vary but there is probably a basic setal pattern
also. Zachvatkin (1948) worked out a pattern of positions and names
for the dorsal setae of Laelaps but this has not been superimposed on
other species. The legs very definitely have a uniform pattern but no
one has worked it out as yet, except for tarsus I. Haarl^v (1943) has
done the chaetotaxy of tarsus I of some of the Mesostigmata.
The base of the palpal tarsus bears a two-tined structure that is
called a specialized seta but it is not of the same origin as the body
setae. Camin «tal. (1957) gave evidence that this is a modified palpal
claw,
The fixed digit of the female and nymphal chela of many species
bears a setiform projection, the pilus dentiUn of unknown origin and
function (Fig, 4).
Also forming part of the integument may be various spurs, spines,
18

and apophyses. These are modified immovable protuberances, chiefly

from the legs, and generally from leg II. They exist more frequently
in males than in females and may be of some secondary sexual value.
The integument is pierced by several small, slitlike pores in var-
ious parts of the body. These are in orderly arrangement on the ventral
side. Thus the sternal plate of the female usually has two pairs and the
male sternal plate three pairs. The female generally has a third pair
below the sternal plate. There are also pores on the dorsal side, gen-
erally along the margins of the shields and there are two on the femur
and tarsus of each leg, one on the line between the basifemur and telo-
femur and the other on the line between the basitarsus and telotarsus.
The function of the pores is not known. One assumption is that they
connect with dermal glands and allow for the escape of a secretion of
some sort. It has also been suggested that they may be sensory (see
Special Senses). The internal mites, with few exceptions, possess only
the pores on the legs.

Internal Anatomy
Endoskeleton. The Gamasina, along with many other mites, pos’
sessan "endostemum" which appears to be a tree plate inside the podo-
somal region and is the point of origin of the leg muscles, and of muscles
which go to the dorsal body wall. This plate seems to be entirely con-
nective tissue fibers. Stanley(1931)callsthisthe "endoskeletal fructule"
Occasionally one finds small, dense bodies internally near the bases
of the coxae. and also near the female genital atrium. These are prob-
ably structures from which muscles originate.
Musculature. Muscles ofthe parasitic Mesostigmatahave been only
slightly investigated. From what is known, it appears that most of the
muscles are flexors. Extensor and flexor muscles are found in the
chelicera so that the chela may be opened and closed, and both dilators
and constrictors are found in the pharynx, enabling the animal to suck
blood. There are also extensors in the tarsi to activate the claws and
ambulacral apparatus. All other parts of the body are extended by the
hydrostatic pressure of the body fluid. Apparently there are no muscles
directly associated with the digestive system (except the pharyngeal
muscles) and the movements of food must be accomplished by peristal-
tic action of the body. Although Stanley (1931) stated that the pharyn-
geal muscles of JEchinolaelags^edudninus are non-striated, more recent
studies show that all mite muscles, including the pharyngeal, are stri-
ated.
Digestive System. The digestive tract of the parasitic Gamasina is
a relatively simple thing. It consists of a pharynx, a thin walled esoph-
agus, a rather small midgut. tour large cecae, a small hindgut, the
rectal sac. and the anus (Fig. 5).
In common with all arachnids, the mouth is simple. It is merely an
anterior opening surrounded by a few modified parts, including the epi-
pharynx. hypopharynx and hypostomal processes (Fig. 6). Immediately
 19

back of this opening is the pharynx with its several rows of constrictor
and dilator muscles. The chelicerae are modified appendages which
serve to puncture the host and cause blood or other body fluids to flow
to the surface. The epipharynx and hypostomal processes are appressed
to the wound and form a tight seal with tha host. The muscular pharynx
then pumps in the fluids as they accumulate at the surface.
Nearly all Mesostigmata have salivary glands which open through
the salivary stylets (Fig. 1). The saliva probably acts both as an anti-
coagulant and as a stimulant to blood flow.
The pharynx pumps the fluids through the esophagus into the midgut.
Four large diverticulae, or cecae, emerge laterally from. the midgut,
two curving anteriorly and two posteriorly. The anterior branches pro-
ceed partially into coxae II and then bend out again and enter coxae I.
The posterior branches go partially into coxae III and then proceed toward
each other at the posterior tip of the body. Crossley (1951, unpublished
data) has illustrated this very nicely (Fig. 5). Some species may have
small diverticulae in addition to the four large ones, or the large ones
may be branched. Michael (1892), in an excellent paper on internal
anatomy, found that females of Hae mogamasus hirsutus and Haemogama-
sus horridus had two pairs of anterior cecae and two pairs of posterior
cecae. The male of Haenaogamasus horridus, however, had only one
pair of posterior cecae.
The hindgut is a short tube which emerges from the midgut and
empties into the rectal sac. which in turn empties to the outside through
the anus. The anus is nearly always ventral near the posterior tip but
is occasionally exactly posterior (Sternostoma) and, rarely, even dorsal
(Hhinonyssus himantopus).
Circulatory System. The mites possess a circulating body fluid
containing cells of various kinds but not all of them have a circulating
device or heart. The ticks and many free-living Gamasina have a heart
dorsally located in the opisthosoma. According to Winlder (1888) such
a heart has a pair of ostia. and one aorta proceeding anteriorly. The
heart itself is roughly hexagonal and very small. However, a heart has
never been demonstrated in any species of parasitic Gamasina and they
are generally assumed to lack one. As in all arthropods, the blood cir-
culates in an "open" system. In parasitic Gamasina the body movements
are apparently sufficient to keep the blood circulating as much as is
necessary.
Excretory System. The excretory system of the Gamasina, inclu-
ding the parasitic forms, consists of two tubules reaching from coxae I
to the posterior end where they enter the rectal sac (see Vitzthum, 1941).
In cross section the tubule is seen to be composed of a single layer of
very large rhomboidal cells surrounding the central lumen. The cells
are so large that only three, rarely four or five, suffice to encircle the
central cavity. Anteriorly, the tubule extends into coxa I, sometimes
even into the femur, and occasionally as far as the genu. The whole
tubule is covered by a thin muscular sheath and the tubules exhibit peri-
staltic movements. The tubules are endodermal in origin and therefore
20

should not be called Malpighian tubules as is sometimes done, the latter

being of ectodermal origin. Also, the Malpighian tubules of insects are
not capable of peristalsis.
The excretory products enter the rectal sac and are voided through
the anus, either separately or with the teces. The chief excretory pro-
duct is said to be guanin (amino-oxypurine), which is white in color and
opaque.
Michael (1892) found a pair of small sacs on each side of the pos-
terior end of the body. just under the dorsal wall in Haemogamaaus hor-
ridus. They opened to the outside through a pore in the cuticle and we re
tilled with a yellowish, oily fluid. He believed the function of these sacs
to be excretory.
Nervous System. The central nervous system of all the Mesostig-
mata consists of a large mass or ganglion located medially in the podo-
soma. This ganglion is pierced by the esophagus and it is evident there-
fore that it is actually a mass consisting of the fused supra", circum-,
and sub-esophageal ganglia, plus the somatic ganglia that are charac-
teristic of the Arthropoda (Fig. 7). Paired nerves, each containing
both sensory and motor fibers, radiate from the central ganglion to the
various parts of the body. The nerves that enervate the gnathosoma
and dorsal light sensitive areas originate from that portion of the gang-
lion above the esophagus; the nerves supplying the legs and opistbosoma
originate from that portion below the esophagus.
Special Senses. The tactile sense in the Semites is apparently assoc-
iated with the body setae. In the parasitic Mesostigmata the body setae
are not greatly modified but, nevertheless, they are probably respon-
sible for the thigmotropic responses of the mite. It is doubtful that
setae, except certain specialized ones on tarsus I and on the tips of the
palpal tarsus, serve any other function.
A sense of equilibrium is well established in the free swimming
water mites and the centers of this sense have been demonstrated for
them. In the land mites, equilibrium can be easily maintained by pedal
contact with the substratum and hence specialized statocysts need not
be so highly developed. An equilibration center (or centers) has not
been demonstrated for the parasitic Mesostigmata, but geotaxis has
been demonstrated, tor some species (Camin, 1953), indicating that such
a center is present.
There is no indication that the mites under consideration here are
able to perceive sound waves.
The olfactory sense is apparently well developed. It is generally
assumedthatthe specialized hollow, striated setae on the dorsal anterior
tip of tarsus I are the chemoreceptors. Camin (195 3) demonstrated con-
clusively that the tip of tarsus I is the location of olfactory reception,
and assumed that the specialized setae are the receptors as they are
found only in that area. The total range of odor perception is not known.
Many ticks and mites possess eyes of the simple, or ocellus, type
on the podosoma that are probably only light sensitive and do not produce
actual images. These simple type eyes are lacking in the Gamasina,
 21

However, experimentation has shown that at least some species are able
to differentiate among lights of different intensity. Vitzthum (1941:244)
suggests that the dorsal ganglion may be directly stimulated by light
waves that pass through the overlying integument. He mentions tost
species with a heavily sclerotized dorsal plate, as in the genus Laelaps,
have a clear, or at least translucent, area immediately above the dor-
sal ganglion.
Camin (1953) has ingeniously shown that the snake mite, Ophionys-
sus natricis, has a pair of light sensitive spots in the pulvillar membrane
of tarsus I (a. Fig. 8). Leg I is not used in arobulation but is held aloft,
antenna-like, as the mite walks. In this position, the pulvillus is longi-
tudinally folded so that one light sensitive spot is on each side of the
fold. This enables the mite to detect with either leg whether the source
of light is from the left or the right.
A pair of small chitinized rings, each enclosing a very delicate
membrane, is present in the pulvillar membrane of tarsus I of some of
the internal snake mites of the family Entonyssidae <b. Fig. 8), These
have been dubbed "eye spots" because of their appearance but their actual
sensory nature has not been investigated.
It is interesting to speculate on the sensory adjustment that was
necessary in Ophionyssus natricis to make a pedal ganglion light sensi-
tive. As far as we know, light sensitivity is always associated with the
cerebral ganglion, never with the somatic ganglion as it must be in this
species. However, the possibility that these spots are actually heat
sensitive rather than light sensitive has not been ruled out.
The slitlike pores have been mentioned as a part of the integument;
they are again mentioned here because it has been suggested that they
may be sensory structures. This is credible because there is a nerve
to each pore. but what the nature of the sense may be is not known.
Vitzthum (1941) calls these pores "spaltorgane" and points out that they
are present not only on the sternal and dorsal plates but also on the
femora and tarsi of all the legs. They are located on the line that denotes
the division between the basi- and telofemur and between the basi- and
telotarsus. They are always present on the ventral side of the leg but
may be on the dorsal side also.
Respiratory System. The respiratory system of the Mesostigmata
consists of a pair of stigmal openings located one on each side between
legs III and IV. They are normally located ventrally but in the endo-
parasitic mites they may be lateral or even dorsal. Also, they may be
as far forward as coxae II (Ptilonyssus) or back near the posterior mar-
gin (Raillinyssus caudistigmus).
Interiorly the stigma opens into a very short trachea which branches
dichotomously, one branch going forward, the other back. The forward
branch very shortly branches again, one branch going ventrally and
supplying the first three pairs of legs and the gnathosoma, the other
going dorsally and supplying all the dorsal region that lies anterior
to the stigma. The posterior branch also branches quickly into ventral
and dorsal tubes, the ventral tube supplying leg IV and apparently all of
22

the opisthosoma, the dorsal tube apparently supplying only the mid-doraal
region. At least one branch of this dorsal tube seems to anastomose
with a similar branch from the other side (Fig. 9).
Running forward from the stigma is a prominent tube or depression
of unknown function known as the peritreme. This structure seems to
connect directly with the stigma. It is hollow, frequently chambered,
apparently has annular thickenings, and is fairly well sclerotized. It
generally extends forward to coxa I but may be longer or shorter. In
the internal mites it is always less than the diameter of a coxa in length;
in some genera (Sternostoma, Rhinonyssus) it is lacking entirely. It
is always short in the protonymph and completely absent in the larva.
Reproductive System. The parasitic Mesostigmata are dioecious
with apparent, though not exaggerated, dimorphism. All species that
have been studied are capable of parthenogenetic reproduction and this
capability probably extends to all species. Parthenogenesis always pro-
duces males, at least in the species whose life history is known, and
perhaps all males, even of fecundated females, are parthenogenetically
produced.
On the average, the ratio of malesto females is probably about equal
but the acarologist nearly always finds a preponderance of females in
any given habitat. This is probably because the male has a much shorter
life expectancy than the female. Mites of the genus Ptilonyssus are so
preponderantly female that some biologists have assumed that these
species produce females by parthenogenesis. However, males have
been found and although the possibility of parthenogenetic production of
females is not ruled out, it is more generally assumed that females are
produced from fecundated females.
Camin (1953) has indicated that the snake mite, Ophionyssus natri-
cis, has a reciprocal ratio of males and females. If the number of
males is low, many females fail to become fertilized and produce nothing
but males. This creates an overabundance of males and thus assures
a virtual absence of virgins in the next generation. Fertilized females
produce much higher than 50 per cent females so the next generation
will again be plagued with virgins, etc.
The male system (Pig. 7) consists of a single testis (or two fused
testes ?), two vasa deferentia, an accessory gland, and a single ejacu-
latory duct. The testis lie sin the middle of the opisthosoma and extends
from dorsal to ventral walls. It consists of relatively few very large
cells with the germinative area apparently at the posterior end. A short
vas deferens arises from each side anteriorly. The accessory gland
lies below the testis. The ejaculatory duct is medial and opens at the
anterior margin of the sternal plate. There is no penis. Spermato-
phoresare formed that are transferred on the chelae. How, and in what
part of the male system, the spermatophores are formed is not known,
Sperm of the Gamasina have seldom been seen. Vitzthum (1941:404).
quoting several authors, describes them as oval, elliptic, apindle-ahapsd
or spatulate bodies from three or four to 30 microns long. They lack a
flagellum (tail). They are probably transferred as immature sperm and
 23

become imature only after being transferred to the female. ^,

: The female system (Fig. 1.0.) consists of one ovary, a short oviduct,
uterus,, and-the genital pore. There is no vagina. The ovary is a com-
pact mass .at the posterior end of the body. .Only one egg or larva is
formed at ,a time .but this is of tremendous size, .frequently 1/4 the vol -
ume of the mite. The genital pore is a transverse opening just anterior
to legs IV. There seems to be no seminal receptacle but sperm are un-
doubtedly retained somewhere in the female system for some weeks after
copulation. Michael (1892) mentioned a central, domed cavity, anterior
and .dorsal to. the ovary, in Haemogamasus hirsutus. This he called the
camera spermatis because it contained sperm but was different in struc-
ture from, a .true receptaculum seminis.

... The Life Stages ., ,.:.

. The life stages consist of egg, larva, protonymph (Nympha I), deu-
tonymph (Nympha 11). and adult female and male. .There/is no evidence
of a so-cap,edtritonymph. As far as is known, there is no resting stage,
orpseudocfarysalis, ;m any instar.. The larval form has six legs; the two
nymphal forms .are octopod but can be readily recognized as immature
stages by the absence of genital openings. According to. Evans (1957)
chaetotaxy of the palps and intercoxal.area may be used to distinguish
between the nymphal.stages,.of the Mesostigmata. .

Egg. The majority of the.parasitic Mesostigmata are ovoviviparous

and produce active, hexapod, larvae, bu.t.most of the Dermanyssidae and
Haemogamasidae lay eggs. The eggs of all species that lay,them are
indistinguishable from.each other. They are pearly, white, ..smooth, and
elliptical, and have a pliable shell. Most eggs. hatch within.. 24 hours.
Eggs that will become males average slightly smaller than those that
will become females. . .,.
.
.
.
.
.Larva (F.ig. 11). Whether hatching friom a preparturient or a post-
parturient egg, thelaryais always hexapod.. (The term larva is perhaps
ill-choaen. as it. generally denotes a stage completely different .in form
from the. adult... The terro.proteronymph suggested,.by Fonseca U948)
seems more desirable, but we will follow the more usual custom of call-
ing the first stage the larva. ] This stage lacks ..respiratory, openings
.and has no plates. It is probably a non-feeding stag.e>,in all species. In
those .species of which we have records, the larva .molts to the proto-
r nymph within 24 hours. .
.
.
.
.
Protonymph (Fig. 12). Inthe two major, families of external mites
.(Laelaptidae and Dermanyssidae), the protonymph may be easily recog-
nized by the fact that (1) the peritre.me is very. short, .not reaching be-
yond cpxa III; (2), the sternal plate is not posteriorly attenuated and does
not,extend beyond the middle of coxae IV; (3) the dorsum has a large
,
podos.omal shield, a smaller opisthosomal shield and two or three pairs
. of’ intermediate platelets. In the Laelaptidae. there .are three pairs of
these small pl-atelets; in the Dermanyssidae only two pairs. In .other
families not enough is yet known about the immature stages to-charac-
24

terizethem. The chelae resemble those of the female although they are
not as Well’developed. It is probable that all species fee.d’in this stage,
Deutonymph (Fig. 13). In the Laelaptidae and Dermanyssidae the
sternal plate attenuates posteriorly and extends as far’as the posterior
margin of coxae IV; the dorsal plate i’s similar to that of the’ adult female ;
and the peritreme is nearly as long as in the adult. There is no genital
opening and. therefore, no epigynial plate.’ (The chelae resemble those
of the’female but are slightly smaller. The claws also are smaller than
in the adult. In most species this’is apparently a feeding stage, but in
Ornithonyssus and Ophionyssus, .and possibly some other Macronyssiriae,
it is’ a non-feeding stage of less than 24 hours duration. That is, the
replete protonymph molts into a deutonymph which, in turn, molts into
the adult within 24 hours. This is not comparable to the pseudochrysalis
of the chiggers because in the Mesostigmata this instar is active right
up to the time of molting and the new appendages form within the old ones.
Protohymphs and deutonymphs of internal mites are practically iden-
tical. That both forms exist is known because it is possible to find
nymphal forms within old nymphal skins just prior to ecdysis.
’Female (Fig. 14). The females of the parasitic Mesostigmata are
always distinguished by the presence of a genital plate or genital pore.
The- genital orifice lies between or anterior to legs IV. .The orifice is
cove’redby a membranous and more or less pleated flap which becomes
more sclerotized and therefore more obvious posteriorly. The anterior
and membranous portion is generally referred to as the genital plate
while the heavily sclerotized portion is thought to be the ventral plate.
Because these two plates are fused they are commonly called the genito-
ventral plate. However, we are not convinced that this plate’is actually
the product of two primitive plates fused’into one and therefore, through-
out this manual, w’e will call it the epigynial plate.
In the external parasites the plates of the ventral surface are: a
sternal, lying between legs II and III; an epigynial, lying between coxae
III and IV and extending posteriorly a greater or lesser distance; an anal,
nearthe posterior tip; a pair of metapodal-plates behind coxae IV; para-
podal plates outside the coxae; and a peritremal plate surrounding the
peritreme.
The sternal plate typically bears three pairs of setae and two pairs
of pores. Occasionally there may be accessory setae and sometimes,
by reduction of the plate, there are less than three pairs of setae on it.
but there are never less than six sternal setae. The plate varies from
a thin transverse rectangle to a large rectangle much longer than wide.
The epigynial plate has a membranous and poorly sclerotized an-
terior margin to allow for the deliverance of the relatively enormous
egg or young. The membranous portion of the epigynial plate extends
from the genital opening forward to the sternal plate and frequently over-
laps its posterior margin. The posterior- portion is sclerotized and
bears at least ’one pair of setae. The posterior margin is pointed or
rounded. .It is never tr.uncate or concave unless proximity to the anal
plate forces it to be. There may be one, two, or three additional pairs
 ’
’
. 25

of setae on the plate. When there are more. .as in the Haemogamasidae,
they become less constant in distribution and number.
The anal plate is ovate .or triangular, with the vertex pointed cau-
dally. It encloses the anal pore, bears three setae, and has a pubescent
caudal area commonly called the cribrum. Occasionally the single pos-
terior seta is missing; sometimes afew accessory setae may be present.
Some mites, especially the Laelaptidae, have a small plate pushed
in between coxae III and:.IV. This is the endopodal plate. The meta-
sternal plate is lacking in all the-parasitic Parasitoidea, but the meta-
sternal pore and the metasternal seta may be present.
Metapodal plates are generally present but are never strongly de-
veloped except in Eulaelaps (Fig. 59). Parapodal plates are generally
present and in the Dermanyssidae." coalesce with. the peritremalia and
embrace coxae IV. The peritremal plate is probably present in the ma-
jority of species but is often difficult to see. Presternal plates are
never present in the parasitic Gamasina but the presternal area may be
sclerotized, .

Dorsally the female, may have one or. two plates. If there .are two
dorsal plates there may be small accessory .platelets also, as in Ophio-
nyssus (Fig. 41) andin some ofthe Rhinonyssidae.
In the internal parasites, there tends to be a reduction of the ven-
tral plates. The first to disappear is the metasternal plate with its
associated seta and pore. The sternal plate tends to become longer and
narrower and less sclerotized until it disappears completely. The pre-
sternal area is never .sclerotized. The anal plate also becomes less
obvious. Its margin be comes poorly defined and the cribrum may dis-
appear; it may lose a seta,,; or all three setae and finally nothing but the
anal pore may remain.. The epigynial plate persists in most species
(except in the Halarachnidae), even in mites that have no other ventral
plates. It always has the pair of .genital setae, and conversely, it never
bears more than these two setae. Oddly enough, the dorsal plates do
not disappear. They are reduced in size, however, and are mostly
restrictedtothe podosoma, and in some species, as Larinyssue orbicu-
laris (Pig., 76), are reduced to a group of platelets. In other species,
as Ne.onyssus (Fig. 78), there may be two dorsal plates that nearly
cover the entire dorsum.
’
.
Male (Fig. 15). The male may be recognized by the genital pore,
a small, round opening between coxae II on the anterior margin of the
sternal plate. Apparently the primitive condition in the external para-
sites is a completely fused ventral armature called the holoventral plate
(Fig. 15). In its fully developed condition, the plate expands abruptly
posterior to coxae IV (Laelaptidae, Haemogamasidae). This is subject
to reduction, however, and the more common condition is for the plate
to narrow below the legs. In some cases the anal plate may be pinched
off and sometimes the holoyentral plate may be divided at about the level
of coxae IV.
In the internal parasites, the ventral plates are always greatly re-
duced. In no case are they all united; generally there is only a small
26

sternal plate, a so-called genital plate between coxae IV, and an.anal
plate.
The outstanding characteristic.of the male is th,e specially modified
spermatodactyl or spermatophore. carrier. The male gamasid lacks an
a.edeagus; the spermatodactyl, on the mcweable arm of the chela, serves
as a copulatbry organ. It "varies in the parasitic Gamasina. but tends
to be recognizably uniform within.each .^enus.(Pig. 16). A complete un-
derstanding of this one structure would enable one’to place a species into
its.proper genus. Wheth&r-or not it serves as-a selective barrier against
.hybridization is liot known. ’ "

’
;.

-
.., ,. ,.. ""
CLASSI.FICATI.QN ’.,;.
-..
..In. this outline of the classification of the parasitic me.sostigmatid
mites, the families, subfamilies, and genera are defined and keyed out.
The species in each genus or subgenus are listed according to the follow-
ingplan. The type species of each genus or subgenus is’placed first and
the other species follow in alphabetical order; under each species the
earliest, reference to the valid specific name is given first, followed in
chronological order by the various generic and specific combinations of
that. name. that: have been used; synonyms follow the valid specific.name
in chronological order. A comma between a scientific’name and. an
author’s, name .indicates that the name of the .original author of the spe-
cific name has been omitted. In the ...specific citations, a ’paper
that
simply refers to the’ species, without .extended discussion’or the pre-
sentation of new information about it, is ;cited only by the year of the
publication. The numbers of the pages that begin- discussions, of species.
follow such references as are important from tlie stahdpoirit’ot nomen-
clature, taxonomy, or because they cantata new information about the.
species. Letter designations follow these page nitmbers to indicate the;
kind. of information the papers contain. The letters used are:?
aanat-
omy,^ b--biology, c--chorology or.. distribution, Bdescription. .m.--
medical, Itaxonomy, vveterinary, The general scope of’the indi-
vidual .papers can be determined by .consultings’trie complete citations
;’
"
given in the bibliography at the end of this manual. ,

The location of the type specimen or the most important series of

cotype specimens, if known, is .indicated under the heading for each’
species. Omission of this’ information .means that the location, of type
material is unknown-to trie .author?.,... .
.,.... ’"

Many of the.^ species ai’-V still so pporly known-that little can be’.gained
by trying to define each"species’listed .here.’ -Much’work must yet be
done before it will be possible to make.a complete’specific revision of
the members.-uf these families. Coverage is. intended to be .complete
through .1956, although- selected references/subsequent to this datejiave
been added.
 27

The Parasitic Mesostigmatid Mites

The mesostigmatid mites are divided into two supercohorts, the

Monogynaspida and Trigynaspida. With one exception (Ophiomegistus
spp.), all of the species parasitic on vertebrates belong to the cohort
Gamasina of the supercohort Monogynaspida. Furthermore, they are in-
cluded in the superfamily Parasitoidea. The parasitoid species will be
discussed first. For a key to the cohorts and superfamilies of the Meso-
stigmata see Camin and Gorirossi (1955) or Baker et al. (1958).
The families that contain parasitic species may be separated from
others by their shields. Typically, the females have a sternal, an epi-
gynial or genito-ventral, and an anal shield. These may be greatly modi-
fied in the internal mites. The epigynial shield always has one pair of
setae associated with it. In the male the genital pore is always on the
anterior margin of the sternal plate, and the chelicerae are modified,
having a spermatodactyl,
Further separation may be based on the specialized forked seta, or
claw. at the inner base of the palpal tarsus (Pig. 1). Without exception,
in the species that parasitize vertebrates, this structure is two-tined
or entirely lacking. In the other groups it is always present and is fre-
quently three-tined or four-tined.
There are, unfortunately, a few groups of non-parasitic mites with
two-tined palpal setae that are very similar to the parasitic forms. They
are the Ascaidae, Iphiopsidae, Phytoseiidae, Aceosejidae, Hypoaapidaiae
and Hyletastinae (= Eviphididae). In the Ascaidae the dorsum is ewwed
by two large shields of equal size and shape. In the parasitic species, if
the dorsum has two plates, the anterior is always distinctly the wider,
except in Neonyssus, a highly specialized endoparasite of birds. The
other groups mentioned above have well developed, dentate chelae and
relatively sparse setation. and therefore may be confused only with the
parasitic Laelaptidae. Prom this family they may be distinguished as
follows:
In the Iphiopsidae, the inner margin of the fixed digit of the chela
is serrate and the peritreme does not extend beyond coxa III. This com-
bination of serrate chela and short peritreme is not found in parasitic
forms. The Phytoseiidae and Aceosejidae are distinguished by the short.
truncate epigynial plate. In the Laelaptidae the epigynial plate is rounded
posteriorly, or if it is truncate, it extends well beyond the middle of the
hysterosoma. Also, both Phytoseiidae and Aceosejidae commonly have
a ventrianal plate, while Laelaptidae never has. The Hypoaspidinae and
Hyletastinae are subfamilies of Laelaptidae and ffElEfereta.iating features
are given under the discussion of that family.
28

Key to the Families of the Parasitoidea

Containing Species Parasitic on Vertebrates

1. Base of the gnathosoina forming a narrow sheath for the cheli-

cerae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Base of the gnathosoma forming a broad cavity or camerostome

that encloses the chelicerae but not the palps. Pound on Cen- (
tral and South American bats . . . . . . . SPELAEORHYNCHIDAE

2. With a pair of prominent holdfast organs on the ventral region

of the opisthosoma; parasites of snakes . . HETEROZERCONIDAE
With a pair of prominent holdfast organs on the ventral region
of the opisthosoma lacking . . . . . . . . . . . . . . . . . . . . 3

3. Tritosternum always present and well developed, with lacinae. 4

Tritosternum lacking or very rudimentary . . . . . . . . . . . 8

4. Dorsal surface thickly beset with setae; epigynial shield with

ten or more setae. In some Breyisterna spp. there may be

. . . . . . . . . . . . . . . . . . . .
less than 10 setae, but they will be irregularly arranged. . . .
. . ... . HAEMOGAMASIDAE
Dorsal surface not thickly beset with Setae. Epigynial shield
with eight or less setae; if with more (some Myonyssus spp.)
then the dorsal setae are sparse and mostly very small, and the
anal plate is extremely broad . . . . . . . . . . . . . . . . . . 5

5. Chelae poorly sclerotized, nearly always edentate and rarely

withapilus dentilis. In the rare instances where teeth are pres-
ent. they are transparent and appear to be quite weak. Corni-
culi poorly defined . . . . . . . . . . . . . . . . . . . . . . . . 6

Chelae generally prominently toothed, or if not, the arms are

well developed and heavily sclerotized. Pilus dentilis rarely
absent. A corona of setae, or seta-like projections at the base
of the moveable arm in most cases. Corniculi well defined . . 7

6. Sternal plate about twice as long as wide. Parasitic in the audi-

tory meatus of cattle . . . . . . . . . . . . . . . . RAILLIETIDAE

Sternal plate not longer than wide, generally much wider than
long. . . . . . . . . . . . . . . . . . . . . . . . DERMANYSSIDAE

7. Corniculi long, harpoon-shaped or hooked at the tip; chelice-

 29

rae normal or with only one arm. Ectoparasifes of snakes . .

. . . . . . . . . . . . . . . . . . . . . . . . . IXODORHYNCHIDAE

Corhiculi may be long, but never hooked at the tip. Female

chelicerae always with two arms. Never parasitic on snakes. . .
. . .’.. . . . . . . . . , ;’’. . . . . . . . . . . . LAELAPTIDAE
8. Large, hairy, well sclerotized mites. Ectoparasites of bats
or ederitates . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Poorly sclerotized mites living in the respiratory passages of

reptiles, ’birds and mammals . . . . . . . . . . . . . . . . . . 10

9. Legs I much thicker’than tHe others. Parasites’of South Amer-

icari armadillos and scaly anteaters. ’. . . . . : ’DASYPONYSSIDAE
’
Legs I about equal to the others.’ Parasites of bats . .. . . .
. . . . . . . ’.’. . . . . .V; . . . . . . . . . . SPINTUKNICIDAE
"

10. Female epigynial plate generally lacking. Parasitic in the res- ’ ’

piratory passages or lungs of mammals . . . . HALARACHNIDAE

Female epigynial plate present, although it may be greatly re- ’

’
duced. . ;’.’.’. . . . . . . . . . . . . . . . . . . . . . . .’ . It

11. Parasitic in the respiratory .passages of birds. Tarsi strong

and with heavy claws . . . . . . . . . , . , , . RHENONYSSIDAE

Parasitic in the lungs and air sacs of snakes. Tarsi slender,

claws not excessively heavy . . . . . . . . . . . . ENTONYSSIDAE

Family LAELAPTIDAE B-erlese; 1892

Vitzthum’(1941) grouped all but the most specialized laelaptoid mites

as subfamilies under the Laelaptidae. -At present-most of these are sep-
arated and given full familial rank, and we think justifiably so since they
can be rather decisively differentiated from each other and from the
Laelaptidae sensu strictu. Two groups that cannot be readily differen-
tiated from the Laela’ptinae are the Hypoaspidinae and the Hyletastinae.
These are therefore temporarily retained in the family.
Description of the family. Modified palpal claw present and two-
tined, ’A single dorsal plate present that covers most of the body in the
male, female, and deutonymph. Protonymph with -two dorsal plates and
four or six platelets between them. Peritremes always extending be-
yond coxa III (except in the protonymph)^ Peritremal plate may or may
not be present. Epigynial plate of the female never pointed posteriorly,
separate from the anal plate, except in some species of Tur.
30

.Diagnosis of the.Laelaptinae.. .W.el,l..sclerotizi2.d.K.ite.s, with coarse

setation, frequently with heavy spurs or spines, especially on the coxa,
ventral plate, femur I andgenul. If spurs and spines are lacking, the body
setae are still-rather long, stiff. and cpar@e,., The tectum is. always a
transparent flap withoirt...ornamentatipn.. The female ventral; plate (al-
ways fused to the .genital) may extend posteriorly as far as the anal
plate and even partially surround it, but it is generally shorter and
rounded posteriorly. The fixed/arm of the chela.always bears a rather
elaborate seta which is generally inflated and of characteristic shape for
the species. In the male the chelae are never shearlike: themoveable
arm’is elongated and" is :.fused to an .even, longer spermatodactyl, The
fixed.arm is. always, much reduced, npnscleroti-zed and apparently func-
tionless. Especially-in this one character of the male do the other-aub-
families differ most drastically, fr.pm the Laelaptinae. In; them, the male
always has shearlike chelae., although the spermatodactyl. i-nay be quite
long. The Hyletastinae and Hypoaspidinae differ, as follows: never with
spurs or. spines .on the coxae or ventral plate; setae fine, slender, gen-
erally short- ’In, the rar? cases .where this is not so, other Characteris-
tics, such as the tectum, male chelae, or female ventral plate will sep^
aratethem: in the Hyletastinae,.the tectum has a narrow lanceolate pro-
jection; in the Hypoaspidinae, unfortunately nothing very definite can
easily .’be seen, but in well mounted specimens the pilus dentilis can be
observed as a small, non-inflated seta. The tectum of the Hypoaspidi-
nae has. a serrated margin, and the male chelae are shearlike. If the
epigynial plate has more than one pair of setae, they are very small
and slender and not prominent.

Key to the Genera of Laelaptinae

. Based, on the Females

1. Epigynial plate bearing only one pair of setae . . . . . . . . . . 2

Epigynial plate with three to eight setae . . . . . . . . . . . . . 8

2. Coxae I, or coxae I and II, with bifid, setigerous spurs . . . .

. . . . . . . . . . . . . . . . . . . . . . . Neoparalaelapa Fonseca

None of the coxae with setigerous spurs . . . . . . . . . . . . . 3

3. All of the coxae with a ventral spur, in addition to.ane or two

spurlike setae. . . .. . . . . . . .....Trichosurolaelaps Womersley

None of the coxae with ventral spurs, although the ventral setae
may he spurlike . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. The posterior seta.of coxa II. longer than the greatest diameter
of the.coxa. Large, coarse, hairy mites, generally over 1 mm
 31.

long. . ,’.". . . . . ; . . . . ..’. . . . . Gigantolaelaps Fonseca

Posterior seta of coxa 1.1 shorter than the length of the coxa . . 5

5. Hypostomal processes forming a prominent brush of stiff, api-

cally expanded -se’tules . . . . . . . . . . . . Bolivilaelaps Fonseca

Hypostomal processes may be fimbriated but they do. not form

a prominent brush . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Body short, almost circular. Legs I hardly longer than legs

II and III. A peritremal plate may.be present, Posterior seta
of coxa III generally heavier than any of the other coxal setae.
Spinelike dorsal setae on.femora and genua I and II . . . . .... 7

Body oval’ or elliptical; legs I distinctly longer than. legs II and

.’.’.". . . . . . . . .
III; posterior seta of coxa III similar to other coxal setae . . .
. . . . . . . . . . . . Haemolaelaps Berlese

7. Anal shield elongated, pyriform . . . . . . . Cavilaelaps Fonseca

Anal shield much broader, not pyriform. . Eubrachylaelaps Ewing

8. Epigynial plate with less than four pairs of setae . . . . . . . . 9

Epigynial plate with four pairs of setae . . . . . . . . . . . . . 13

S. Three setae on the epigynial plate . . . . . Radfordilaelaps Zumpt

Three pairs of setae on the epigynial plate . . . . . . . . . . . 10

10. Medial dorsal setae very small; dorsal plate with concave mar -
gins; female sternal plate twice as wide as long. the third pair
of setae may be off the plate . . . . . . . . . Oryctolaelaps Lange

Medial dorsal setae prominent; margins of dorsal plate convex;

female sternal plate nearly square, with three pairs of setae . 11

11. Coxae I. II. and III with heavy, thornlike setae . . . . . . . . .

.
. . . . . . . . . . . . . . . . Steptolaelaps Furman
No thornlike setae on the coxae but coxa II may have a re-
curved apophysis . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Coxa II with a recurved apophysis . . . . . . Heterolaelaps Hirst

No coxal ornamentation; all coxal setae slender. . . . . . . . .

Mesolaelaps Hirst
32

13. Sternal plate as long as or longer than wide. Epipharynx en-

larged apically, with a median, longitudinal groove. Epigynial
plate large and generally concave on the median posterior mar-
gin. Large mites ... . . . . . . . . . . . . . . ..,. . . . 18 ...
Sternal plate always wider than long. Epipharynx pointed, not
grooved. Hypostomal processes with slender, finger like pro-
jections. Generally less than 1 mm long . . . . . . . . . . . . 14

14. Coxa II with a prominent, recurved apophysis on the anterior

ventral surface. Epigynial setae varying from three to five
pairs . . . . , . . . . . . , . ., . .:. . . . . . Heterolaelapg. Hirst

Coxa II not so adorned . . . . . . . . . . . . . . . . . . . . . . 1-5

15. All setae long and slender . . . . . . . . . Tricholaelaps Vitzthum

Some setae, especially those of the coxae and ventral plates,

, heavy and thornlike . . . . . . ..
. . . . . . . . . . . . . . . 16

16. Body: long and narrow . . . . . . . . . . . Longolaelaps Vitzthum

Body oval, elliptic, or rounded . ... . . . . . . ... .. . . . . . . . 17

I?. Peritremalia embracing coxa IV; corniculi weak and flaccid; all
posterior coxal setae slender; legs I and II with spurlike setae
ventrally on trochanter and femur, . . . . . . . . . . . . ... .
. . . . . . . . . . . . . . . . Aetholaelaps Strandtmann and Camin

Peritremalia short, not encircling coxa IV; corniculi discrete;

at least some coxae with spurlike or at least heavy posterior
setae; none of the legs with ventral spurlike setae on trochanter
and femur . . . . . . . . . . . .. . . . . . . . Laelaps C. L. Koch

18. All hysterosomal setae long . . . . . . . . . Echinolaelaps Ewing

Dorsal hysterosomal setae minute . . . . . Mysolaelaps Fonseca

Genus Haemolaelaps Berlese, 1910

(Fig. 17)
Type: Haemolaelaps marsupialis Berlese.
Laelaps of authors, not of C. L. Koch. 1839.
Hypoaspis of authors, not of G. Canestrini, 1885.
Atricholaelaps Ewing, 1929.
Type: Laelaps reithrodontis Ewing, 1925: 7.
Ischnolaelap.s Fonseca, 1935: 19,
Type: Ischnolaelaps reticulatus Fonseca, 1935: 19.
 33

Diagnosis: Generalized roitess with only one pair of setae on the fe-
male epigynial plate, no spur-like or thornlike setae on the coxae. male
chela not shearlike, body oval and legs long. The female sternal plate
but
is wider than long, the male ventral plates are generally entire,
is separate. The better known species are
sometimes the anal plate
very definitely blood suckers, but many species are known only from
known
single specimens or from very small collections and nothing is
of their habits.

Haemolaelaps marsupialis Berlese

Type material in the Agricultural Entomology Station. Florence,

Italy.
Laelaps (Haemolaelaps) marsupialis Berlese, 1910: 261 (d).
Haemolaelaps marsupialis, Strandtrnann. 1949: 329(1); Tragardh, 1952:
66 (t); Womersley. 1955:423 (d, cf. ? illus.); Domrow and Smith.
1956: 202.
Locality; Australia.
Hosts: Mammal - "bandicoot, " a marsupial of the family Perameli-
-
dae. Bird nesting material from burrow of Puffinus tenuirostris.

Haemolaelaps aegyptius Keegan

Type material in the U. S. National Museum.

Haemolaelaps aegyptms Keegan, 1956: 246 (d, $ illus.).
Locality: Africa - Egypt.
Hosts: Mammals - Gerbillus gerbillus, Gerbillus pyramidum, Ger-
billus quadrimaculatus, Jaculus jaculus, Meriones crassus (type host).
Meriones libycus, Psammomys obesus, Rattus rattus.

Haemolaelaps aadrogypua Bregetova

Haemolaelaps androgynus Bregetova, 1952: 867 (c, d); 1956: 89 (illus.).

Locality: U.S.S.R. - Armenia; Russian Soviet Federal Socialist
Republic, Astrakhan Province; Turkmenistan.
Hosts: Mammals
musculus.
-
Citellus pygmaeus. Meriones persicus. Mug

Haemolaelaps angustiscutis Bregetova

Haemolaelaps angustiscutis Bregetova, 1952: 867 (c, d); 1956: 92.

Locality: U.S.S.R. - Tadzhikistan and Turkmenistan.
Hosts: Mammals - "sand rat" and its nests; "Redtail sand rat" and
its nests; Meriones erythrourus. Rhombomys opimfas.
34

Haemolaelaps arvicanthis (Radford)

Ischnolaelaps arviicanthis Radford, 1939: 246 (d).

-
Locality: Africa Uganda, Kakumiro and Kitgum.
Host: Mammal - Arvicanthis abyssinicus rubescens.

Haemolaelaps bathyergus (Radtord)

Ischnolaelaps bathyergus Radford, 1939: 250 (d),

Locality: Africa - Union of South Africa, Strandfontein, Capetown.
Host: Mammal - Bathyergus suillus.

Haemolaelaps bibbyi Strandtmann and Hunt

Type in the U. S. National Museum.

Haemolaelaps bibbyi Strandtmann and Hunt, 1950: 85 (d).
Locality: Philippine Islands - Samar.
Host: Mammal ~ Rattus frugivorus.

Haemolaelaps capensis Hirst

Haemolaelaps ? capensis Hirst, 1916: 79 (d); Bedford. 1932: 274 (c);

Strandtmann, 1949: 331 (t).
Locality: Africa - Union of South Africa, Cape Proiancfi, Gra.ba3Ec»-
town.
-
Host: Mammal Georychus hottentotus.

Haemolaelaps casalis (Berlese)

Iphis casalis Berlese, 1887: Fasc. XXXVIII, No. 8.

Haemolaelaps casalis, Bregetova, 1956: 89.
Hypoaspis oculatus Oudemans. 1915: 134. New synonymy.
Haemolaelaps molestus Oudemans. 1930: ll;Willmann, 1939: 108 (c. d).
Atricholaelaps megaventralis Strandtmann, 1947: 112 (d).
Haemolaelaps megaventralis, Strandtmann, 1949: 337 (b. c. t); Jameson,
etaL. 1952; Bregetova. 1952 (d); Pratt and Lane, 1953: 360; Asa-
numa, 1953: 14; Smith, 1955; Womersley, 1956: 563; Eads, 1956;
Strandtmann, 1956: 138.
Hypoaspis freemani Hughes. 1948: 129 (d).
Haemolaelaps haemorrhagicus Asanuma. 1952: 87. New synonymy.
Localities: Cosmopolitan.
Hosts: Mammals - Apodemus agrarius. Fells catus, Glaucoroys
volans, Mus musculus, Mus musculus albinus, Peromyscus nasutus.
Oryzomys palustris, Rattus rattus, Rattus norvegicus, Rattus turkes-
 35

tanicus, Sciurus carolinensis, Sciurus niger, Sciurus Vulgaris, Sigmo-

don hispidus. Birds - Centurus carolinus, Ceqphloeae pileatus, Dryo-
bates pubescens, Euphagus sp., Iridoprocne bicolor, Melanerpes eryth-
rocephalus, Passer domesticus, Petrochelidon fulya, Petfpchelidon’pyr-
rhonota, Riparia riparia, Sturnus vulgaris, SyphrapicuS varius.
Remarks; This mite is more frequently associated with birds than
is any other Haemolaelaps. It is also more frequently found in straw,
hay, detritus, etc. than is any other species of the genus.

Haemolaelaps chinchillulae (Strandtmann)

Type material in the U. S. National Museum.

Atricholaelaps chinchillulae Strandtmann, 1948: 189 (d).
Locality: South America Peru, S. W. Have.
Host: Mammal - Chinchillula sahamae.

Haernolaelaps coelogenys (Fonseca)

Type material in the Institute Butaritan,’Sao Paulo, Brazili

Ischnolaelaps coelogenys Fonseca, 1’9’36: 31 (d).
Locality: South America - Brazil, Rio de Janeiro.
Host: Mammal - Coelogenys pacca.

Haemolaelaps concur rens Berlese

Hypoaspis (Haemolaelaps) concurrens Berlese, 1918: 129 (d).

-
Locality: Europe Sardinia.
Host: ’?

Haemolaelaps cryptomius (Radford)

Type material in the collection. of.C. D. Radford.

Ischnolaelaps cryptomius Radford, 1939: 248 (d).
Locality: Africa - Union of South Africa, Cape Province.
Host: Mammal - Cryptomys capensis.
.
Haemolaelaps dasymys (Radford)

Type material in the collection of C. D. Radford.

Iscnaolaelaps dasymys Radford, 1939: 244 (d). :’
:

Locality: Africa - Uganda.

’
’

Host: Mammal - Dasymys helukus helukus.

36

Haemolaelaps -dayisi Zumpt and Till

Type material in the South African Institute .for Medical Research.

Haemolaelaps davisi Zumpt and Till, 1956: 287 (cr.and $, illus.).
Locality: Africa - Basutoland. .
’ ,;;

.
-
.Host: Mammal Mystromys- albicaudatus.

Haemolaelaps disimilis Asanuma

Type material in the collection of K. Asanuma.

Haemolaelaps disimilis Asanuma, 1953: 12 (d).
Locality: Asia - Japan, Miyake Island.
Host: .Bird - Emberiza rustica latifascia.

Haemolaelaps ellobii Bregetova

Haemolaelaps ellobii Bregetova, 1952: 870 (d); 1956: 93.

Locality: U.S.S.R. - Western Kazakhstan; Tadzhikistan, Tch-
kalov Province; and Turkmenistan. : -
Hosts: Mammal$ -’ Ellobius fuscocapillus. Ellobiustalpinus, Lagu-
rus lagurus.

Haemolaelaps eloffi Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps eloffi Zumpt and Till, 1953: 226 (d).
Locality: Africa - Union of South Africa, Orange Pree State,
Bloemtontein.
Host: Mammal - Cryptomys bigalkei.

Haemolaelaps elongatus Berlese

Hypoaspis (Haemolaelaps) elongatus Berlese. 1-918: 129 (d).

.Locality: Europe- Sardinia, i
.
Host: "Found in detritus in a stable. "

Haemolaelaps ewingi Keegan

Type material in the U. S. National Museum. ’

Haemolaelaps ewingi Keegan, 1956: 243 (d, 9 illus,).’

Locality: Africa - Egypt.
-
Host: Mammal Gerbillus gerbillus.
’
 37

Haenaolaelaps fahrenhoizi Berlese

Haemolaelaps fahrenhoizi Berlese, 1911: 432; Willmann, 1952: 402 (c,

d, t).
Locality: Europe - Germany; Italy.
Hosts: Mammals - Apodemus agrarius, Apodemus flavicollis, Apo-
demus sylvaticus, Arvicola amphibius, Clethrionomys glareolus, Micro-
tus agrestis, Microtus arvalis, Microtus oeconomus, Mus sylvaticus,
Putorius putorius, Talpa europaea.
Remarks: According to Willmann (1952), mohrae Oudemans is a
synonym of this species. But according to Bregetova (1952), mohrae
is a synonym of glasgowi Ewing. 1925. Therefore, if both Willmann and
Bregetova are correct, then the name glasgowi must be submerged and
fahrenhoizi Berlese, 1911 must replace it. However, we have seen
neither the type nor the original description and therefore prefer to list
fahrenhoizi as a separate species for the present.

Haemolaelaps galagus Lavoipierre

Type material in the Liverpool School of Tropical Medicine.

Haemolaelaps galagus Lavoipierre, 1955: 304 <d, illus.).
Locality: Africa - British Cameroons, Barombikang Forest.
-
Host: Mammal Galago alleni, a primate.
Remarks: Described from a single female specimen.

Haemolaelaps geomys Strandtmann

Type material in the U. S. National Museum,

Haemolaelaps geomys Strandtmann. 1949: 339 (c, d),
Locality: North America - United States.
Hosts: Mammals -
Cratogeomys castanops, Geomys bursarius,
Geomys cumberlandicus, Geomys floridanus. Geomys lutescens, Ge-
omys personatus, Geomys texensis, Geomys tuza, Thomomys bottae,
Thomomys bulbivorus.

Haemolaelaps glasgowi (Swing)

(Fig. 17)
Type material in the U. S. National Museum.
Laelaps glasgowi Ewing, 1925: 6 <d); Grant, 1947: 11; Strandtmann, 1949:
343 <t).
Atricholaelaps glasgowi, Wharton, 1938: 139; Jameson, 1947: 142 (c);
Ewing. 1947: 84 (t); Strandtmann, 1948: 187 (c); Harkema and Kart-
man, 1948: 183 (b).
Haemolaelaps glasgowi. Strandtmann, 1949: 339 (b, c, d, t); Aameson,
1950 (b); Asanuma, 1951: 5; Jameaon et al., 1952: 10; Irons et al. ,
38

1952: 2; Bregetova, 1952: 867 (t); Pratt and Lane. 1953: 360; Kee-
gan, 1953: 37; Bregetova, 1953: 309; Dubinin. 1953; Pratt and Goode,
1954: 125 (c); Burgess. 1955; Smith, 1955; Womersley, 1955: 421;
Bregetova, 1956: 86; Baker etal.. 1956.
Laelaps californicus Ewing, 1925: 5 <d); Grant, 1947: 10; Strandtmann,
1949: 434 (t).
Geneiadolaelaps califomicus, Augustson, 1941: 157.
Laelaps virginianus Ewing, 1925: 6 <d); Grant, 1947: 11; Strandtmann,
1949: 343 (t).
Haemolaelaps microti Oudemans, 1926 (d); Bregetova, 1952; 867 (t).
Haemolaelaps mohrae Oudemans. 1928: 374 (d); Nelzina and Barkov.
1951: 179 <m); Nelzina and Romanwa, 1951: 829 (m); Bregetova,
1952: 867 (t).
Hypoaspis cricetophilus Vitzthum, 1930: 417 (d); Strandtmann, 1949:
343 (t).
Laelaps stegemani Hefley, 1935: 22 (d); Strandtmann, 1949: 343 (t).
Haemolaelaps scalopi Keegan. 1946: 71 (d); Strandtmann. 1949: 343 (t),
Atricholaelaps strandtmanni Fox. 1947: 598 (d); Strandtmann, 1949: 343
(t).
Atricholaelaps sjgmodoni Strandtmann, 1946; 164 (d); Randolph and Eads.
1946: 599; Jameson, 1947: 142; Harkema and Kartman. 1948: 186;
Strandtmann, 1949: 343 (t).
Hypoaspis (Haemolaelaps) eos ZumptandPatterson, 1951: 79 (d); Zumpt
and Till, 1953: 246 (t).
-
Localities: Asia Japan; Korea; U.S. S.R. Australia -Fisher Is-
-
land. Bass Strait. South Australia. Europe Germany; Holland; Russia.
-
North America Canada; Mexico; United States; Puerto S4.co. South
-
America Peru.
-
Hosts: Mammals Allactaga saltator, Ammospermophilus leucurus,
Apodemus agrarius. Apodemus flavicollis, Apodemus speciosus, Apo-
demus sylvaticus. Auliscomys boliviensis, Baiomys tayloM, Blarina
breyicauda. Chraeomyg pulcherrunus. CiteUus armatus, Citellus beech-
vi, Citellus columbianus. Citellus lateralis chryppdeirus, Citellus mex-
manus. Citellus mollis. CiteUus pygmaeus, CiteUus richagdaoaM,, Ci-
tellua tridecemlineatus, Citellus undulatus. Clethrionomys glareolus,
Clethrionomys rufocanus. Clethrionomys rutilus, Cratogeomys casta-
nops. Cricetulua barabensis, Cricetulus migratorius. Cricetulus trx-
ton. Cricetus cricetus. Crypt otis parva. Cynomys gunnisoni, Cynomyg
ludovicianus, Didelphis virginiana, Dipodomys mxcrops, Dipodomys ordi,
Eutamiaa minimus, Eutamias sibericus, Evotomys gapperi, Geomye
bursarius, Geomys cumberlandicus, Geomys floridanus, Geomys lu-
tescens, Geomys personatus. Geomys tuza. Lepus califomicus, Mar-
mota monax, Marmota sibirica. Mephitis elongatum. Mephitis nigra,
Meriones persicus, Meriones tamariscinus. Micromys minutus. Micro-
tus. agrestis, Microtus arvalis. Microtus brandti, Micrptus breweri,
Microtus californicus. Microtus fortis, Microtus gregalis, Microtus
montanus, Microtua mordax dutcheri. Microtus mordax sierra, Micro-
tus nanu^ Microtus^ ochrogaster, Microtus oeconomus, Microtus ore-
 39

goni, Microtus pennsylvanicus, Microtus subterraneus, Microtus angu-

rensis, Myospalax dybowskii, Myotig grisescens, Napeozapus insignts,
Neofiber alleni, Neotoma cinerea, Neotoma floridana, Neotoma micro-
pus, Neotoma pennsylvanica, Neotoma sp., Nesokia indica, Ochotona
alpina, Ochotona daurica, Ochotona pusilla, Onychomys leucogaster,
Oryzomys palustris. Otospermophilus grammurus, Perognathus cali-
fornicus, Perognathus hispidus, Peromyscus boylii, Peromyscus cali-
fQmicus, Peromyscus crinitus, Peromyscus gossypinus, Peroroyscus
.leucopus, Peromyscus maniculatus, Peromyscus nuttaUi, Peromyscus
polionatus, Peromyscus sp,, Peromyscus truei, Phenacomys longicau-
dus, Phodopus bedfordiae, Phyllotis arenarius, Pitymys pinetorum.
Procyon lot or, Punoinys leminus, Rattus norvegicus, Rattus rattus,
Rattus turkestanicus, Reithrodontotnys humilis, Reithrodontomys mega-
lotus , Rhabdomya pumilio, Scalopus aquaticus, Scapanus latimanus,
Scapanus to^vnsendi, Sciurus carolinensis, Sciurus douglasi, Sciurus
hudsonicus, Sciurus niger, Sciurus sp., Sigroodon hispidus, Sorex ara-
neus, Sorex fumeus, Sorex vagrans, Sylvilagus auduboni, Sylvilagus
floridanus, Sylvilagus sp.. Synaptpmys sp., Talpa europaea, Talpa al-
taica, Tamias striatus, Tamiasciurus dpugla.sii albalunbatua, Thomoroys
bottae, Thomomys bulbivorus, Urocyon cinereoargentatus, Zapus hud-
sonicus. Birds - Dryobates villosus, Egretta alba, Colqmba sp., fairy
penguin, Lanius ludovicianus, Melospiza melodia, mutton bird, Pipilo
erythrophthalmus., Riparia riparia, Speotyto cunicularia, Stelgidopteryx
ruficollis.
Remarks: This commonnnte represents a species complex. Mites
from hosts of one species are slightly different from mites of a different
host species. There is every reason to believe that the above synonymy
is correct, yet this vast complex probably contains some good and dis-
tinct species.
Nelzina and Romanova (1951) have found this mite to be naturaUy
infected with tularemia in Russia and were able to transmit the disease
to laboratory white rats by allowing them to feed on the animals.

Haeroolaelaps hirsti Keegan

Type material in the U. S. National Museum.
Haeroolaelaps hirsti Keegan. 1956: 242 (d, 9, rf, NUlus.).
-
Locality: Africa Sudan.
Host: Mammal - a rat.

Haemolaelaps hystrici Zunapt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps hystrici Zumpt and Till, 1953: 253 (d); Keegan, 1956: 239.
Locality: Africa-Uganda, Kararooja district; Nigeria; Kenya; Egypt.
-
Hosts: Mammals Arvicanthisniloticus, Crpcidura manni, Hystrix
40

galeata (type host), Rattus rattus.

Haemolaelaps insculptus Keegan

Type material in the U. S. National Museum.

Haemolaelaps insculptus Keegan. 1956: 244 (d. d" and $. Sillus,).
Locality: Africa - Egypt.
Hosts: Mammals - Arvicanthis niloticus, Gerbillus gerbillus, Ger-
billus nanus, Gerbillus pyramidum, Jaculus jaculus. Mus musculus.

Haemolaelaps inversus Berlese

Hypoaspis (Haemolaelaps) inversus Berlese, 1918: 129 (d).

Locality: Europe - Italy. Florence.
Host: Found in detritus.
Kemarks: Probably not a true Haemolaelaps.

Haemolaelaps labuschagnei Zumpt and Patter son

Type material in the South African Institute for Medical Research.

Hypoaspis (Haemotaelaps) labuschagnei Zumpt and Patterson, 1951: 85
(d).
-
Localities: Africa Union of South Africa, Transvaal, Natal, Zulu-
land; Cape Province.
-
Hosts: Mammals Mastomya coucha. Otomys irroratus, Otomys
tropicalis, Rattus rattus.

Haemolaelaps lawrencei (Zumpt and Patterson)

Type material in the Natal Museum, Africa.

Liponyssus lawrencei Zumpt and Patterson, 1951; 89 (d).
Haemolaelaps lawrencei, Zumpt and Till. 1953: 8 (footnote).
-
Locality: Africa Union of South Africa. Cape Province.
-
Host: Mammal Georychus capensis.

Haemolaelaps longipes Bregetova

Type material in the Zoologic Institute of the Academy of Science,

U.S.S.R.
Haemolaelaps longipes Bregetova. 1952: 867 (d); 1956: 92.
Localities: U.S.S.R. -
Armenia, Turkmenistan, Tadzhikistan.
Hosts: Mammals - Citellus pygmaeus, Mer tones erythrourus, Mer-
iones tamariscinus, Meriones tristrami, Rhombomys opimus.
 41

Haemolaelaps lophuromius (Kadford)

Type material in the collection of C. D. Radford.

Cyclolaelaps lophuromius Radford, 1939: 243 (d).
Haemolaelaps lophuromius, Furman, 1955:- 59 (t).
-
Locality: Africa Uganda, Kampala.
Host: Mammal - Lophuromys aquilus.

Haemolaelaps macroventralis Asanuma

Type material in the collection of K. Asanuma.

Haemolaelaps macroventralis Asanuma, 1953: 14 <d).
Locality: Asia - Japan, Onsen, Matsuyama, Shikoku.
Host: Mammal - Petaurista leucogenys.

Haemolaelaps mauritanicus Hirst

Haemolaelaps mauritanicus Hirst, 1925: 98 (d),

Locality: Africa - French West Africa.
Host: Mammal - Gerbillus hirtipes.
Remarks: Hirst said "male" in the text but he described and fig-
ured a female.

Haemolaelaps mesopicus Radford

Type material in the collection of C. D. Radford.

Haemolaelaps mesopicus Radford, 1942: 192 (d).
Locality: Africa - Uganda.
Host: Bird - Mesopicus ruwenzorii.

Haemolaelaps morlani Strandtmann

Type material in the U. S. National Museum.

Haemolaelaps morlani Strandtmann. 1949: 337 (d).
Locality: North America - United States. Georgia.
Host: Mammal - Rattus rattus,

Haemolaelaps rourinus (Berlese)

Laelaps (Haemolaelaps) murinus Berlese, 1911: 432 <d).

Hypoaspis (Haemolaelaps) murinus, Berlese, 1918: 131.
Haemolaelaps murinus, Keegan, 1956: 233 (c, t).
42

Locality: Africa - Egypt; Kenya; Uganda; Unitfi of Soatfa AErica,

Zululand.
-
Hosts: Mammals Arvicanthis abyssinicus, Arvicanthis niloticus,
Crocidura olivieri, Dasymys incomptus (type host), Lemniscomys stri-
atus, Lophuromys aquilus.
Remarks: Keegan makes Haemolaelaps arvicanthis (Radford) a syn-
onym of murinus, but we are retaining the two as separate species.

Haemolaelaps mystromys Radford

Type material in the collection of C. D. Radford.

Haemolaelaps mystromys Radford, 1942: 305 (d).
Locality: Africa - Union of South Africa, Glen Craig, Albany.
-
Host: Mammal Mystromys albicaudatus,

Haemolaelaps namrui Radford

Type material in the U. S. National Museum.

Haemolaelaps namrui Radford, 1954: 310 (d).
-
Locality: Asia Yemen (Arabia).
Host: Mammal - Meriones rex buryi. a jird.

Haemolaelaps natalensis Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps natalensis Zumpt and Till, 1953: 230 (d).
Locality: Africa - Union of South Africa. Natal, Pieterniaaitebarg.
Host. Mammal - Cryptomys natalensis.

Haemolaelaps oliffi Zumpt and Patterson

Type material in the South African Institute for Medical Research.

Hypoaspis (Haemolaelaps) oliffi Zumpt and Patterson, 1951: 81 (d).
Locality: Africa - Union of South Africa, Orange Free State and
Transvaal.
-
Host: Mammal Tatera brantsii.

Haemolaelaps omnitectus Vitzthum

Hypoaspis (Haemolaelaps) omnitectus Vitzthum, 1928: 181 (d).

-
Locality: Island of the Pacific Ocean Java.
Host: Bird - Chrysocolaptes validus.
 43

Haemolaelaps pachyptilae Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps pachyptilae Zumpt and Till, 1956: 285 (d. ?).
Locality: Antarctica - Heard Island.
-
Host: Bird Nest of Pachyptila desolata, a petrel.

Haemolaelaps patersoni Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps patersoni Zumpt and Till. 1956: 286.
Locality: Africa - Union of South Africa. Transvaal, Rustenburg.
Host; Bird - Gyps coprotheres, a vulture.

Haemolaelaps phoeniculi Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps phoeniculi Zumpt and Till, 1954: 209 (d).
Locality: Africa-Union of South Africa, Transvaal, Potchfifatroom.
Host: Bird - Phoeniculus purpureus Miller (a red-billed hoopoe).

Haemolaelaps radfordi Keegan

Type material in the U. S. National Museum.

Haemolaelaps radfordi Keegan, 1956: 241 (d, $ illus.).
Locality: Africa - Sudan.
Host: Mammal - a rat.

Haemolaelaps razumovae Bregetova

Type material in the Zoologic Institute of the Academy of Science,

U.S.S.R.
Haemolaelaps razumovae Bregetova, 1952: 870 (d); 1956: 91.
Locality: U.S.S.R. -Georgia.
Host: Mammal - Prometheomys schaposchnikovi and its nests.

Haemolaelaps reithrodontis (Ewing)

Type material in the U. S. National Museum.

Laelaps reithrodontis Ewing, 1925: 7 (d).
Atricholaelaps reithrodontis, Ewing, 1929: 186 (t); Generotype of Atri-
cholaelaps.
Haemolaelaps reithrodontis. Strandtmann. 1949: 329 (t).
44

Locality: South America Argentina.

Host: Mammal - Reithrodon cuniculoides,

Haemolaelaps reticulatus (Ponseca)

Type material in the Institute Butantan. Sao Paulo, Brazil.

Ischnolaelaps reticulatus Fonseca, 1935: 5(d)r generotype of Ischno-
laelaps.
-
Locality: South America Brazil. Sao Paulo.
-
Hosts: Mammals Euryzygomatomys spinosus catellus, Mus mus-
culus, "rat, " "rat’s nest, " Sciurus estuans.

Haemolaelaps rhabdomys (Radford)

Type material in the collection of C. D. Radford.

Ischnolaelaps rhabdomys Radford. 1939: 249 (d).
-
Locality: Africa Union of South Africa. Transvaal.
Host: Mammal - Rhabdomys pumilio.

Haemolaelaps rhodesiensis Zumpt and Patterson

Type material in the South African Institute for Medical Research.

Hypoaspis (Haemolaelaps) rhodesiensis Zumpt and Patterson, 1951: 82
(d).
Locality: Africa - Southern Rhodesia.
Host: Mammal - Saccostomus campestris.

Haemolaelaps sangsteri (Radford). New combination

Liponyssus sangsteri Radford, 1942: 190 (d); Ponseca, 1948: 321.

-
Locality: Africa Uganda.
-
Host: Mammal Protoxerus stangeri centricola.
Remarks: The tongue-shaped epigynial plate, broadly rectangular
sternal plate, triangular anal plate, and short peritremalia indicate that
this species properly belongs in Haemolaelaps.

Haemolaelaps scapularis Berlese

Hypoaspis (Haemolaelaps) scapularis Berlese, 1918: 129 (d).

Locality: Africa ? -
"Durban."
-
Host: Mammal Spalax typhus.
 45

Haemolaelaps sciureus (Fonseca)

Type material in the Institute Butantan, Sao Paulo, Brazil.

Ischnolaelaps sciureus Fonseca, 1935: 36(d); Strandtmann, 1949: 331 (t).
Locality: South America - Brazil, Rio de Janeiro.
Host: Mammal - Sciurus estuans.

Haemolaelaps semidesertus Bregetova

Type material in the Zoologic Institute of the Academy of Science,

U.S.S.R.
1 Haemolaelaps semidesertus Bregetova, 1952: 870 (d); 1956; 91.
Locality: U.S.S.R. -
Russian Soviet Federal Socialist Republic,
Astrakhan, Stalingrad, and Tadzhikistan provinces.
’ Hosts: Mammals - Alactagulus acontion, Allactaga elater, Allac-
taga jaculus, Arvicola terrestris, Citellus pygmaeus. Cricetus overs -
manni, Mus musculus.

Haemolaelaps spatuliformis Lavoipierre

Type material in the Liverpool School of Tropical Medicine,

Haemolaelaps spatuliformis Lavoipierre, 1956: 293 (d. Ulus.),
Locality: Africa - British Cameroons, Kumba.
Host: Mammal - Anomalurus fraseri, a flying squirrel.
Remarks: Described from three female specimens.

Haemolaelaps spegazzinii Berlese

Hypoaspis (Haemolaelaps) spegazzinii Berlese. 1923: 254 (d).

Locality: Africa - Abyssinia.
Host: None given.
Remarks: This may not be a true Haemolaelaps.

Haemolaelaps spinitarsus Berlese

Hypoaspis (Haemolaelaps) spinitarsus Berlese. 1918: 129 (d).

Lpcality: Africa - Union of South Africa, Zululand.
Host: Mammal - Georychus sp.

Haemolaelaps spinosulus Berlese

Hypoaspis (Haemolaelaps) spinosulus Berlese, 1921: 152 (d).

Locality: ? - "Messico. "
46

Host: None given.

Remarks: This may not be a Haemolaelaps.

Haemolaelaps spreo Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps spreo Zumpt and Till, 1956: 290 (d, 9).
Locality: Africa
-
-
The only locality quoted is "Springs."
Host: Bird Spreo bicolor, the pied starling.
t-

Haemolaelaps sternalis (Ewing)

Type material in the U. S. National Museum.

Ichoronyssus sternalis Ewing, 1922: 14 (d, ). ;’.
Haemolaelaps sternalis, Strandtmann, 1956: 138 (t).
-
Locality: Nort h America United States, Maryland. Plummer Is-
land.
-
Host: Mammal Woodchuck.

Haemolaelaps subterraneus Berlese

Hypoaspis (Haemolaelaps) subterraneus Berlese, 1921; 153 (d).

Locality: Europe - Italy, Florence.
Host: None given.
Remarks: This is probably not a Haemolaelaps.

Haemolaelaps sudanicus Zumpt and Till

Type material in the South African Institute for Medical Research.

Haemolaelaps sudanicus Zumpt and Till. 1954: 213 (d); Kaagan, 1956:
237.
Locality: Africa - Sudan, near Torit; Kenya.
Hosts: Mammals - Arvicanthis sp., Lophuromys aquilus, Otomys
tropicalis, Tachyoryctes sp.

Haemolaelaps tachyoryctes Radford

Type material in the collection of C. D. Radford.

Haemolaelaps tachyoryctes Radford, 1941: 309 (d); Keegan, 1956: 236.
Locality: Africa - Kenya.
Hosts: Mammals - Tachyoryctes ruddi. Lemniscomys striatus.
 47

Haeiaiolaelaps taterae Zumptiand .Patterson

Type material in the South African Institute for Medical Hesearch.

Hypoaspis (Haemolaelaps) tateraa Zumpt and .Patterson, 1951: 83 (d).
Locality: Africa - Union ,ot S.outh Africa, Transvaal.
-
Host: Mammal Tatera .brantsii. .... . .. .

.
Haemolaelaps tateronis (Kadford)

Type material in the collection of C. D. Radford.

Ischnolaelaps tateronis; Radford, .19.3.9: .247 (d), ..
Locality: Africa - Uganda. : ....
Host: Mamraal - Tateroaa benvenuta. ;
,..,:
;
.. .

’
.

.
Haemolaelaps traubi (Strandtniann)

Type material in the U. S. National’Museum.’

Atricholaelaps traubi Straadtmahn, 19-48: 187 (d).
Haeniolaelaps tr&ubi, ’Asanuhia, !953: 10.
’

Locality: Asia -Burma.’

"
’
’
’
Host: Mammal -’ Callosciurus quihque striatus.

Haemolaeiaps viliosi.SB’fnius Berlese

Hypoaspis iHae’molaelaps) villosissitnua Berise, 1918: 129 (d);. Zumpt

.’ ",;
’^
’

and Patterson,-1951: 83’(d’).

Locality: Africa-Southern Rhodesia; Unioi’1. of South Africa, Zulu-
." ’’ " "
’ ’ ’

land.
-
Hosts: Mammals Saccostomus campestris, Mus coucha.

Haemolaelaps zulu Berleae

Hypoaspis (Haemolaelaps) inops var. zuln Berlese, 1918: 129 (d); K&e-
.gan,.19R6:.234 ($ illus.). : .

Locality: Africa - Union of South Africa, Zululand; Egypt,

..Hosts:. Mammals -Arvicanthis aor sails, Arvicanthis niloticas,
Crocidura orivieri, Lemniacoroys griselda, -Hattus rattus.

Genus Cavilaelaps Fonaeca, 1935

(Fig. 18)
Type: Cavilaelaps bresslaui Fonseca, 1935.
Piagaosis: Medium mites with rather sparse setation. The female
epigynial plate bears one pair of setae and has only one other pair near
48

it. The anal plate is elongated. The.ven.tral plates of the male are con-
tiguous but the anal plate retains its identity.

Cavilaelaps bressfaui Fonseca .:;’.

.. (Fig. 18) ’ ’",

-
.
.

Cavilaelaps bresslaui Fonseca, 1935: 11 (d).

Locality: South America - Argentina, Jujuy.
Host: Mammal - Caviella australis.

Cavilaelaps braziliensis (Ewing)

Type material in the U.’S, National Museum-’

Laelaps braziliensis Ewing, 1925: 3 (d).

"
’
Cavilaelaps braziliensis, Fonseca,. 1.938; 104.

.
Locality: South America - Brazil, Bahia.
Host: Mammal - Kerodon spixi.

Genus Bolivilaelaps Ponseca, 1940.

,...’ (Fig. 19)
’

.
.
Type: Bolivilaelaps tricholabiatus Fonseca.
Diagnosis: Medium mites. The female epigyniai -plate bears only
one pair of setae: the hypostomalprocesses have membranous extensions
bearing a brush of stout setules" with slightly expanded extremities,

Bolivilaelaps tricholabiatus Ponseca

(Fig. 19)
Type material, in the Institute Butantan, Sao Paulo,: Brazil.
Bolivilaelaps tricholaBiatus Fonseca,"i940:.’64 (d). .

.
Locality: South America -.Bolivia.
Host: Mammal - Isothrix bistrlatus.

’Genus Eubrachylaelaps Ewing, 1929

(Fig. 20)
Type: Laelaps. haUisteri Ewing. "’.
Cyclolaelaps Ewing, 1933.
Type: Cyclolael’aps circularig Ewing. .-
Diagnosis: Medium mites with a nearly -circular-body. The female
epigynial plate has;;(in,e pai,r of. setae. A peritremal’ plate is present.
Legs I are no longer., ;or only barely longer, than legs II. ’The posterior
seta of coxa III .n)ay ’.’he. thorniike,-; but -the.’ other coxal s-fetae’are slender.
Males have recently been described by Furman (1955) but have not been
illustrated.

Eubrachylaelaps bollisteri (Ewing)

..
Type material in the U. S. National Museum.
Laelaps hollisteri Ewing. 1925: 2 (d); Grant, 1947: 6(d)’.
 49

Eubrachylaelaps hollisteri, Ev/ing. 1929: 186 (t); Jameson, 1947; Jame-

son, 1950; Furman, 1955.
-
Locality: North America United States, California.
Hosts: Mammals - Neotoma sp., Per ognathus californicus, Pero-

’
myscus californicus, Peromyscus crinitus. Peroriiyscus maniculatus
gambeli; Thornomys bottae.
Remarks: According to Furman (1955), this species occurs abun-
dantly on Peromyscus californicus and P. crinitus and only rarely on
any other mammal.

Eubrachylaelaps circular is (Ewihg)

.
Type material in the U. S. National Museum.
Cyclolaelaps cireularis .Ewing, 1933: 6 <d); Radford, 1939.
Eubrachylaelaps cireularis, Jameson, 1950: 62 (t); Jameson, 1951; Kee-
gan. 1953; Furman, 1955. .’. -.

Locality: NorthAmerica - Mexico, Chiapas; United States, Arizona,

California. Colorado. Utah.
-
Hosts: Mammals Neotoma sp.., Neotoma ’mexicana, Peromyscus
boylii, Peromyscus californicus, Peromyscus hylocetes, Peromyscus
maniculatus, Peromyscus oaxacensis, Peromyscus truei.

Eubrachylaelaps crowei Jameson

Type material, in the U. S. National Museum. ’"

Eubrachylaelapa crowei Jameson, 1947: 391 (d);1950, 19 51;’Irons’etaL,
1952: 2(m); Keegan, 1953: 37 (c); Furman, 1955.
Locality: ..North America - United States, Colorado, Kansas, Tex-
as, Utah.
.

.
..
,
-
Host: Mammal Onychomys leucogaster.

Eubrachylaelaps debilis Jameson

Type material in the U. S. National Museum.

Eubrachylaelaps debilis Jameson, 1950: 64 <d); l’951;’Keegan, 1953: 37
(c); Furman, 1955: 58. .
’
;’’’
.
-
Locality; North America Mexico, D. P., Michocoan, ’Nuevo Leon;
United States, California, Oregon. Utah.
Hosts: Mammals - Microtus longicaudus, Peromyscus crinitus,
-
Peromyscus maniculatus. Birds Nucifraga columbiana, Oberholseria
chlorura. : : ’

Eubrachylaelaps janae-soni Turaian

(Fig. 20) -;;.’

- ^- ’.
.
Type material in the U. S. National Museum. ’
:
’’
’
Eubrachylaelaps jamesoni Furman, 1955: 52 (d).
Locality: North America
Host:
- Mexico, Chiapas.
Mammal - Peromyscus nie’xicanus saxatilis.
’
50

Eubrachylaelaps martini Jameson

Type material in the U. S-." National Museum.

Eubrachylaelaps martini Jameson, 1951: 556 (d); Furman. 1955.
Locality: Central America - Mexico. Mexico State.
Host; Mammal - Neotomodon alstoni.

Eubrachylaelaps rgtundus Fonseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Eubrachylaelaps rotundas Fonseca, 1935: 20<d); Jameson, 1951: 556 (d);
Furman, 1955: 59.
Locality: South America - Brazil, Sao Paulo.
Hosts: Mammals - Akodon mollis orophilus. Zygodontomys lasiurus.

Eubrachylaelaps spinosus Purman

Type material in the U. S. National Museum.

Eubrachylaelaps spinosus Furman.. 1955: 54 (d).
.Locality: North America - Mexico. Michoacan.
Host: Mammal - Peromyscus sp.
’

’
"-

Genus Gigantolaelaps Fonseca, 1939

(Fig. 21)
Type: Gigantolaelaps vitzthumi Fonseca, 1939.
Diagnosis: Large mites (over one mm); dark in color and with dense
setation. The male holoventral plate is expanded behind coxae IV. The
female sternal plate has a narrow anterior projection that reaches the
tritosternum and bears the first pair of sternal setae. The female epi-
gynial plate has one pair of setae. In bath sexes the posterior seta of
coxa II is longer than the coxa.

Gigantolaelaps vitzthumi Ponseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Gigantolaelaps vitzthumi Fonseca, 1,939: 28, 77 (d); Morlan, 1951.
-
Locality: South America Brazil. MinasGeraes and Goyaz boun-
dary.
Host: Mammal - "Wild rats."

Gigantolaelaps brachyspinosus (Fonseca)

Type material in the Institute Butantan, Sao Paulo, Brazil,

Macrolaelaps brachyspinosus Ponseca, 1935: 8 (d); 1936: 22 (d).
Gigantolaelaps brachyspinosus, Fonseca, 1939; 51 and 100 (d); Morlan.
1951.
Locality: South America - Brazil, Matto Grosso, Porto Joffre.
Host: Mammal - Holochilus vulpinus.
 51

G igantolaelap s butantanensis (Fonseca)

Macrolaelaps butantanensis Fonseca. 1935: 7 (d); 1936: 21 (d).

Gigantolaelaps butantanensis, Fonseca, 1931: 44, 93 <d); Morlan, 1951.
Locality: South America - Brazil, Butantan.
Host: Mammal - "Wild rat. "

Gigantolaelaps comatus Fonseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Gigantolaelaps comatus Fonseca, 1939: 39. 87 (d); Morlan. 1951,
-
Locality: South America Brazil. Butantan.
Host: Mammal - "An unidentified field rat. "

Gigantolaelaps cricetidarum Morlan

(Fig. 21)
Type material in the U. S. National Museum.
Gigantolaelaps cricetidarum Morlan. 1951: 274 (d, t); Pratt and Lane.
1953.
Locality: North America - United States, Florida, Georgia, South
Carolina, Texas.
-
Hosts: Mammals Oryzomys palustris palustris, Oryzomys palus-
tris coloratus, Rattus norvegicus, Battus rattus, Sigmodon hispidus ko-
mareki.
Remarks: Oryzomys palustris, the rice rat, is by far the predom-
inant host. (See also Morlan and Strandtmann. 1949: 217.)

Gigantolaelaps gilmorei Fonseca

Type material in the Institute Butantan. Sao Paulo. Brazil.

Gigantolaelaps gilmorei Fonseca. 1939: 22. 71 (d); Morlan, 1951.
-
Locality: South America Brazil. Goyaz.
Hosts: Mammals - Echimys sp., "wild rodents. "

Gigantolaelaps goyanensis Fonseca

Gigantolaelaps goyanensis Fonseca. 1939: 32, 81 (d); Morlan, 1951.

-
Locality: South America Brazil. Goyaz. Rio de Janeiro. Federal
-
District, Minas Geraes Goyaz boundary.
-
Hosts: Mammals Estrimys ? sp., field rats, Metachirops opos-
sum, Nectomys squamipes, rato paco, rats.

Gigantolaelaps mattogrossensis (Fonseca)

Type material in the Institute Butantan. Sao Paulo. Brazil.

52

Macrolaelaps mattogrossensis Fonseca, 1935: 8 (d); 1936: 22 (d).

Gigantolaelaps mattogrossensis, Fonseca, 1939; 40, 90 (d); Morlan,
1951.
Locality: South America-Argentina, Tobocai, Salta; Brazil, Ceara,
MattoGrosso.
Hosts: Mammals - Field rats, Holochilus sciureus, Holochilus vul-
pinus.

Gigantolaelaps maximus (Berlese)

Laelaps (Laelaps) maximus Berlese, 1904: 259 <d).

Gigantolaelaps maximus, Fonseca, 1939: 10, 60.
Locality: South America - Uruguay, Montevideo.
Host: Mammal - Hesperomys vulpinus.

Gigantolaelaps oudemansi Fonseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Gigantolaelaps oudemansi Fonseca. 1939: 15, 62 (d); Morlan, 1951.
Locality: South America - Brazil, Goyaz, Sao Paulo.
Hosts: Mammals - Echimys sp., field rats, Nectomys squamipes.
Sylvagus ; sp.

Gigantolaelaps peruvlanus (Ewing)

Type material in the U. S. National Museum.

Macrolaelaps peruvianus Ewing. 1933: 7 (d).
Gigantolaelaps peruvtanus, Fonseca, 1939:-8, 59 (t); Morlan, 1951.
Locality: South America Peru. -
Hosts: Mammals - mice and rats, wild rats.

Gigantolaelaps versteegi (Oudemans)

Laelaps versteegi Oudemans. 1904: 160<d); Oudemans. 1927;Buitendi3k.

1945.
Gigantolaelaps versteegi, Fonseca, 1939: 10, 60 (t); Morlan, 1951.
-
Locality: South America Surinam.
Host: Mus sp.

Gigantolaelaps wolffsohni (Oudemans)

Laelaps wolffsohni Oudemans. 1910: 67(d); Oudemans. 1927; Buitendijk.

1945.
 53

Gigantolaelaps wolffsohni, Morlan, 1951: 273 (t).

-
Locality: South America Chile.
Host: Mammal - "Probably found on Mus. "

Genus Neoparalaelaps Fonseca, 1936

(Fig. 22)
Type: Paralaelaps bispinosus Fonseca, 1935.
Paralaelaps Fonseca. 1935. (Not Paralaelaps Tragardh, 1908).
Type: Same.
Diagnosis: Small, well sclerotized species. The epigynial plate
is not expanded posteriorly and bears only one pair of setae; all of the
coxae have ventral spurs.
Remarks: Only females are known of this genus. A critical re-
examination may prove this to be a macronyssid rather than a laelaptid
mite.

Neoparalaelaps bispinosus (Fonseca)

(Fig. 22)
Type material in the Institute Butantan, Sao Paulo, Brazil.
Paralaelaps bispinosus Fonseca, 1935: 15 (d).
Neoparalaelaps bispinosus, Fonseca, 1936: 58 (t).
Locality: South America - Brazil, Sao Paulo.
-
Host: Mammal Cavia rufescens.

Neoparalaelaps bandicoota (Womersley). New combination

Type material in the South Australian Museum.

Paramelaelaps bandicoota Womersley, 1956: 574 (d, $).
Locality: Australia; Queensland, Brisbane,
Host: Mammal - Bandicoot.
Remarks: This species has broad spurs on all four coxae, rather
than only on coxae I and II, but otherwise does not differ generically from
bispinosus, the type of Neoparalaelaps.

Genus Radfordilaelaps Zumpt. 1950

(Fig. 23)
Type: Radfordilaelaps meridionalis Zumpt, 1950.
Diagnosis: Large mites, over one mm long. The posterior seta of
coxa I in both sexes is a heavy, knifelike spine. The female epigynial
plate bears three setae; the holoventral plate of the male is only slightly
expanded behind coxae IV.
54

Hadfordilaelaps meridionalis Zumpt

(Pig. 23)
Type material in the South African Institute for Medical Besearch.
Radfordilaelaps meridionalis Zumpt. 1950: 83 (d).
-
Locality: Africa Union of South Africa. Northern Transvaal.
-
Host; Mammal Pedetes caffer, a springhare.

Genus Mesolaelaps Hirst, 1926

(Pig. 24)
Type: Laelaps (Mesolaelaps) anomalus Hirst, 1926.
Diagnosis: Medium to large mites; epigynial plate small, bearing
three pairs of setae; coxae unarmed.
Remarks: Males of this genus are as yet undescribed.’

Mesolaelaps anomalus Hirst

(Pig. 24)
Laelaps (Mesolaelaps) anomalus Hirst, 1926: 840 (d).
Mesolaelaps anomalus, Womersley, 1937: 538;Domrow and Smith, 1956:
203.
Locality: Australia-Queensland. Townsville, Cairns Dis.; Islands
of the Pacific Ocean - New Guinea, Hyon Gulf.
Hosts: Mammals - Perameles macrura, Perameles nasuta, Peram-
eles sp., bandicoot.

Mesolaelaps australiensis Hirst

Laelaps (Mesolaelaps) australiensis Hirst. 1926: 840 (d).

Mesolaelaps australiensis, Womersley, 1937: 538; Derrick et al., 1939:
154 (m); Womersley, 1955: 425; Domrow and Smith, 1956: 203.
-
Locality: Australia New South Wales. Queensland, South Austra-
lia, Victoria.
-
Hosts: Mammals Isoodon torosus, "mice, " Perameles gunni,
Perameles macrura, Rattus lutreola. Birds - Nesting material of the
mutton bird.
Remarks: Derrick et al., in a search tor the infective agent of Q
fever, ground up examples of this mite from a bandicoot and injected
them into a guinea pig with negative results.

Mesolaelaps lagotisinus Hirst

Laelaps (Mesolaelaps) lagotisinus Hirst, 1931: 563 (d).

Locality; Australia Perth. -
-
Host: Mammal Thylacomys lagotis.
 55

Mesolaelaps thalacomys Womersley

Type material in the South Australian Museum.

Mesolaelaps thalacomys Womersley, 1956: 560 (d, ?),
-
Locality: Australia South Australia, Ooldea.
Host: Mammal - Thalacomys lagotis i

Genus Tricholaelaps Vitzthum. 1926

(Fig. 26)
Type: Laelaps (Tricholaelaps) comatus Vitzthum., 1926.
Diagnosis: Similar to Laelaps, except that all of the setae are slen-
der. The epigynial plate has four pairs of setae.
Remarks: This is a monotypic genus based on two female specimens
only.

Tricholaelaps comatus Vitzthum

(Fig. 26)
Type material in the Vitzthum collection.
laps (Tricholaelaps) comatus Vitzthum, 1926: 24 (d).
-
Locality: Indonesia Sumatra, Wai Lima.
Host: Mammal - Rattus whiteheadi.

Genus Heterolaelaps Hirst. 1926

(Fig. 25)
Type; Laelaps (Heterolaelaps) antipodiana Hirst, 1926.
Diagnosis: Medium mites, nearly circular in form. The epigynial
plate is small but bears three to five pairs of setae. Coxa II has a strong
ventral spur. Only the female is known.

Heterolaelaps antipodianus Hirst

(Fig. 25)
Laelaps (Heterolaelaps) antipodiana Hirst, 1926: 838 (d).
Heterolaelaps antipodianum, Womersley, 1937: 538; Derrick etal.. 1939:
154 (m).
Heterolaelaps antipodianus, Domrow and Smith, 1956: Womersley, 1956:
559.
Locality: Australia - Adelaide. Sydney, Victoria, Tasmania, Bris-
bane, Queensland, Betsy Island.
-
Hosts: Mammals Isoodon pbesulus, Isoodon torosus, Perameles
gunni, Peranaele s nasuta, rabbits.
Remarks: In a search for the infective agent of Q fever. Derrick
et al, injected ground up mites of this species taken from a bandicoot
into guinea pigs, with negative results.
56

Genus Trichosurolaelaps Woroersley, 1956

(Fig. 27)
Type: Trichosurolaelaps crassipes Womersley.
Diagnosis: The female sternal plate is quadrangular and nearly
square. The epigynial plate is rounded posteriorly and has one pair of
setae. Legs I have several ventral spurs and all tour coxae bear spurs
and spurlike setae. The male chela is shear like.
Remarks: This monotypic species resembles Hirstionyssus very
closely but the drawings of the peritreme and tritosternum indicate that
it is a laelaptid. Unfortunately, gnathosomal details are not given.

Trichosurolaelaps crassipes Womersley

(Fig. 27)
Type material in the South Australian Museum.
Trichosurolaelaps crassipes Womersley, 1956: 564 (d, rf and $),
Locality: New Zealand, Wellington; Australia, New South Wales and
Queensland.
Host: Mammal - Trichosurus vulpecula, an opossum.

Genus Steptolaelaps Furman, 1955

(Fig. 28)
Type: Neolaelaps heteromys Fox.
Diagnosis: The female chelae are very unusual, the moveable arm
is sclerotized. toothed and subtended by two very long setae; the other
arm is nonsclerotized. slender, edentate and apparently also moveable.
The male chela has a strongly toothed moveable arm bearing a moder-
ately slender spermatodactyl that is about twice the length of the move-
able arm; the immoveable arm is a straight, apically rounded stump
about half as long as the moveable arm. The holoventral plate of the
male is slightly expanded behind coxae IV; the male anal plate may or
may not be separate. The gnathosomal setae of the female are very
heavy; in the male they are slender.

Steptolaelaps heteromys (Fox)

(Pig. 28)
Type material in the U. S. National Museum.
Neolaelaps heteromys Fox, 1947: 117 (d).
Steptolaelaps heteromys, Furman, 1956 (in litt.) New combination.
Steptolaelaps heteromydis Furman, 1955: 521 (d. c? and $).
-
Locality: South America Venezuela, Kancho Grande, Aragua; Co-
lombia, Caracolicito, Magdalena.
-
Hosts: Mammals Akodon urichi, Heteromys anomalus, Heteromys
sp., Proechimys cayanensis trinitatis, Kattus rattus frugivorus. Bhip-
idomys nitela, Sigmodon hispidus hirsutus.
 57

Remarks: Purman erected the genus Steptolaelaps for his new spe-
cies S. heteromydis that he later found to be identical with Neolaelaps
heteromys Fox and so notified us by letter. Consequently, the new com-
bination, Steptolaelaps heteromys (Fox) should be credited to Furman.
Of 137 specimens at Furman’s disposal, over 90 were from the type
host, Heteromys anomalus.

Steptolaelaps liomydis (Grant)

Type material in the U. S. National Museum.

Laelaps liomydis Grant, 1947: 8 (d).
Steptolaelaps liomydis, Furman, 1955: 523 (d, d" and 9).
Locality: North America - Mexico. Jalisco, Michoacan, Oaxaca,
Chiapas, Vera Cruz, United States, Texas.
-
Hosts: Mammals Liomys irroratus acutus, Liomys irroratus u;-
roratus, Liomys pictus isthmius, Liomys pictus veraecrucis, Liomys
pictus parviceps.

Genus Laelaps Koch, 1836

(Fig. 29)
Type: Laelaps hilaris Koch, 1836.
Laelaps, subg. Hyperlaelaps Zachvatkin, 1948.
Type: Laelaps (Hyperlaelaps) amphibius Zachvatkin.
1948.
Laelaps, subg. Laelaps, Zachvatkin, 1948.
Type: Laelaps (Laelaps) muris (Ljungh, 1799)
Diagnosis: Medium mites. The female sternal plate is wider than
long; the epigynial plate always has four setae. At least some of the body
setae are spiniform. The male ventral plates are fused into a single holo-
ventral plate (which is expanded behind coxa IV) or, rarely, the anal
plate may be separate. Sternal setae I of the male overlap the bases of
the second setae.
Hemarks: C. L. Koch described and illustrated Laelaps agilis and
Laelaps hilaris in 1836 but did not designate a generotype until 1842:
88 when he designated hilaris as the type. (See Oudemans, 1936; ’WU.l-
mann, 1952).
In 1830 Robineau-Desvoidy proposed the generic name Cryptostoma
for a mite from Microtus arvalis. This was undoubtedly a Laelaps but
it is impossible to say whether the species was the present agilis or hil-
aris. The name Cryptostoma R-D., 1830 was preoccupied by that of
Blainview, 1818.
Zachvatkin (1948), in a detailed paper on Russian Laelaps spp., pro-
posed that the species of the genus be grouped into two subgenera, Lae-
laps sensu strictu and Hyperlaelaps. The former subgenus included
those species which are highly pigmented. have a very convex dorsal
plate, have long and subequal dorsal setae, and in which the female
58

sternal plate has a straight posterior margin, and the male anal and
ventral plates are confluent. The subgenus Hyperlaelaps included those
species that are pale in color, have an almost flat dorsal shield, have
short and thick medial dorsal setae, and in which the female sternal
plate is lunate, and the male anal plate is separate from the ventral
plate. He also suggests that Laelaps hilaris and Laelaps festinus are
synonyms of each other; that Laelaps pachypus, Oudemans. 1927, is not
the pachypus that Koch described but is what he now calls Laelaps (Hy-
perlaelaps) amphibius; that Laelaps arvicolae Oudemans, 1927, a male,
is not the same as arvicolae Oudemans, 1916, but is the new species
Laelaps micromydis Zachvatkin; and that Laelaps agilis is not determ-
inable. Lange (1955) reviewed the genus Laelaps in the U.S.S.B... and
accepted agilis as the type. He created four new subgenera (Myolaelaps,
Rattilaelaps. Microtilaelapg, and Cricetilaelaps) based mostly on chae-
totaxy and host relationships among cricetine and microtine rodents.
In a number of these efforts, the Russian acaroiogists are no doubt cor-
rect.. but we are not now in a position to critically evaluate their work.
Therefore, we are conforming to the earlier nomenclature. If subgenera
are recognized, the type of the subgenus Laelaps would have to be hil-
aris Koch. 1836 rather than muris (Ljungh. 1799) as designated by Zach-
vatkin or agilis Koch, 1836 as designated by Lange.
It is quite probable that host preference in the species of this genus
is much more marked than the collection data of some of the species
indicate. Sometimes mites are accidentally on a strange host and often
mites may be incorrectly identified. All are obligatory parasites of
small mammals, chiefly rodents. On the other hand, it is also true that
these mites are nidicoles. so that different mammals with similar nests
can provide suitable sites tor the non-feeding activities of these mites
(Wharton, 1957).

Laelaps hilaris Koch

Laelaps hilaris Koch, 1836: #4 (d); Murray. 1876; Oudemans, 1913: 195
(d); Oudemans, 1914; Hirst, 1916: 70 (c); Oudemans, 1927; Oude-
mans. 1936: #99. 100, 104 (t); Zachvatkin, 1948; Willmann. 1948 :
68; 1952: 393 (t); Bregetova. 1956: 109 (illus.).
Localities: Europe-Western Europe and the British Isles; U.S.S.R.
Hosts: Mammals - Apodemus flavicollis, Apodemus sylvaticus,
Clethrionomys glareolus. Microtus agrestis, Microtus arvalis, Micro-
tus gregalis, Microtus nivalis, Microtus oeconqmus, Microtus orcaden-
sis, Microtus socialis. Mus musculus, Mustela vulgaris, Sorex araae-
us, Talpa europea.
Remarks: The preferred hosts seem to be Microtus spp., especi-
ally Microtus arvalis.
 59

Laelaps aethlopicus Hirst

Laelaps aethiopicus Hirst, 1925: 56 <d); Keegan. 1956,

Locality: Africa Kenya. -
Host: Mammal - rats.

Laelaps agilis Koch

Laelaps agilis Koch, 1836: 4, 19 <d); Hirst, 1916; Oudemans, 1927;

Oudemans, 1936: #102 (t); Lange. 1948; Willmann, 1952: 392 (t);
1955: 181; Bregetova. 1956: 110 (illus.).
Laelaps festinus Koch, 1839: 24, 7 (d); Hirst, 1916 (d); Oudemans. 1927;
Hora, 1934; Willmann, 1952.
Laelaps crassipus Shrank, 1897 (d); Oudemans, 1903: 2: #55 <t); Oude-
mans. 1927: .198 (t). (Oudemans credits crassipus to Shrank; we
were unable to find Shrank’s publication.)
Locality: Europe -Western Europe and the British Isles; U.S.S.R,
Hosts: Mammals -Apodemus agrarius, Apodemus flavicollis, Apo-
demus fulvipectus, Apodemus sylvaticus, Rotelmaus (Clethrionomys gla-
reolus?), Talpa europea, water rat.
Remarks: Oudemans (1927 : 175) considered agilis a synonym of
muris (Ljungh) but Willmann (1952: 393) showed rather conclusively that
this is hardly likely.

Laelaps agrarius Lange

Laelaps pavlovskyi agrarius Lange. 1948: 77 (d); Dubinin. 1953.

Locality: U.S.S.R. - Russian Soviet Federal Socialist Republic,
Moscow, Astrakhan.
Host: Mammal - Arvicanthis agrarius.

Laelaps alaskensis Grant

Laelaps alaskensis Grant. 1947: 8 (d).

Locality: North America - Alaska, Jack River.
-
Host: Mammal Microtus sp.
Remarks: This species is perhaps a synonym of muris (Ljungh),

Laelaps algericus Hirst

Laelaps algericus Hirst, 1925: 57 (d); Dubinin. 1953; Bregetova, 1956:

106 (illus.); Keegan, 1956.
-
Locality: Africa -Algeria, Touggourt; Egypt. Asia U. S.S.R.
Hosts: Mammals - Mus algericus, Mus musculus, Apodemus al-
60

gerxcus, Crocidura olivieri.

Remarks: Somewhat resembling nuttalli.. but having a heavily scler-
otized band along the periphery of the dorsal plate.

Laelaps arvalis Zachvatkin

Laelaps (Hyperlaelaps) arvalis Zachvatkin, 1948.

Hyperlaelaps arvalis, Dubinin, l953;Bregetova, 1953; 1956: 117 (illus.).
Locality: U.S.S.R, -
Russian Soviet Federal Socialist Republic.
Stalingrad Province.
Hosts: Mammals - Clethrionomys rufocanus, Microtus agrestis,
Microtus arvalis, Microtus gregalfs, Microtus ungurensis, Rattus nor-

Laelaps arvicolae Oudemans

Laelaps arvicolae Oudemans, 1916: 132 (d); Willmann, 1952: 398 (d).
-
Locality: Europe Germany; Netherlands.
Hosts: Mammals - Arvicola arvalis, Microtus amphibius, Mus
minutus.
Remarks; Arvicola arvalis is probably the host of choice.

Laelaps assimilis Womersley

Type material in the South Australian Museum.

Laelaps assimilis Womersley, 1956: 557 (d, <J and $).
Laelaps sp. Domrow and Smith, 1956: 202.
Locality: Australia - Mt. Glorious. Brisbane, Mt. Nebo.
Host: Mammal - Rattus assimilis.
Remarks: This mite may prove to be the same as Laelaps nuttalli.

Laelaps bakeri Hirst

Laelaps bakeri Hirst. 1923: 690 (d).
Laelaps giganteus var. bakeri. Hirst. 1925: 67 (d); Bedford, 1832.
Macrolaelaps baken, Radford, 1950: 369.
-
Locality: Africa Algeria; Union of South Africa, Cape Province;
Uganda.
-
Hosts: Mammals Arvicanthis barbarus, Rattus rattus, "various
rodents,"

Laelaps bhutanensis (Turk). New combination

Macrolaelaps bhutanensis Turk, 1946: 351 (d).

 61

Locality: Asia - Bhutan, Hasimara.

Host: Mammal - Vandeleuria nilagirica nilagirica.
Laelaps buxtoni Radford

Laelaps buxtoni Radford, 1941: 307 (d).

Locality: Asia - India, Madras.
Host: Mammal - "a gerbille. "

Laelaps caucasicus Lange

Laelaps agilis caucasicus Lange, 1948: 77 (d).

Locality: Europe - Russia.
Hosts: Mammals - Apodemus flavicollis, Apodemus fulvipectus,
Apodemus sylvaticus.

Laelaps clethrionomydis Lange

Laelaps clethrionomydis Lange, 1955: 330 (d); Bregetova, 1953; Brege-

tova, 1956: 108 (illus.).
-
Locality: Asia U.S. S.R., North Korea, Siberia.
Hosts: Mammals - Apodemus speciosus, Clethriqnomys frater,
Clethrionomys glareolus, Clethrionomys rufocanus, Clethrionomys ru-
tilis, Microtus gregalis, Microtus ungurensis, Mogera robusta.
Remarks: Bregetova used this name in 1953, giving Lange credit
for it. Since the former worker gave only distribution and host data
and no description, there can be no question about the authenticity of
the above name.

Laelaps differens Fonseca

Laelaps differens Fonseca, 1935: 21 (d); 1936: 35 (d), female only.

Locality: South America - Brazil, Sao Paulo.
Host: Mammal - "wild rat. "

Laelaps ekstremi Zachvatkin

Laelaps (Laelaps) ekstremi Zachvatkin, 1948 (d).

Locality: U.S. S.R. - Russian Soviet Federal Socialist Republic,
Stalingrad Province.
Host: Mammal - Microtus arvalis.
62

Laelaps exceptionalis Ponseca

LaeXaps exceptionalis Fonseca, 1935: 22 (d); 1936: 34 (d),

Locality: South America - Brazil. Sao Paulo,
Host: Mammal - "rat."

Laelaps finlaysoni Womersley

Laelaps finlaysoni Womersley, 1937: 535 (d).

Locality: Australia - South Australia.
Host: Pseudomys apodemoides, Pseudomys minnie.

Laelaps grenieri Tauftlieb

Type material in the Laboratory of Medical Entomology, Pasteur

Institute, Paris.
Laelaps grenieri Taufflieb, 1954: 439 (d, $ illus.).
-
Locality: Africa French Equatorial Africa, Brazzaville.
-
Hosts: Mammals Leggada musculoides, Lempiscomys striatus.

Laelaps hapaloti Hirst

Laelaps hapaloti Hirst. 1931: 563 (d); Womersley, 1937: 532 (d); Wom-
ersley, 1956: 557.
-
Locality: Australia South Australia, Central Australia.
-
Hosts: Mammals Hapalotis sp., Pseudomys sp., Notomys mitch-
elli.

Laelaps hilaroides Oudemans

Laelaps hilaroides Oudemans, 1927: 375.

Locality: Europe - Germany.
Host: Mammal - Eptesicus serotinus.
Bemarks: This is the only species of Laelaps so far recorded trosa.
a bat. A description is given, but the species is not illustrated.

Laelaps hirsti Ponseca

Type material in the Institute Butantan. Sao Paulo, Brazil.

Laelaps hirsti Fonseca. 1938: 117 (d).
Locality: South America - Brazil. Sao Paulo.
-
Host: Mammal Oryzomys eliurus.

Laelaps jettnaari Vitzthum

Laelaps jettmari Vitzthum, 1930: 405 (d); Asanuma, 1951; Mayer. 1951:
21 (m); Jameson et al., 1952: 10 (c); Suyemoto et al., 1954; Brege-
tova, 1956: 108; Baker et al.. 1956: 43.
 63

Laelaps agUis subsp. jettaiari, Wi’llmann, 1-952:396.

’

;’
-
.
.

Locality: Asia Japan;.’ Manchuria. : s ’ .. ’

Hosts: Mammals - Apodernu^ag’rariu.s,’ Cricetulus griseus, Mus

agrarius, Microtus arvalis.
Remarks: This spec’ies is’quite similar .-to agUis C. L. Koch, but
probably distinct from it. According to Japanese workers, it is a car-
rier’and’transmitterofhaemort-hagicfever. The preferred host is prob -
ably Apodemus agrarius. .. ,..: .
’’’’ "
’" La&Iaps’keegani Thurman .’

Type material in the U. S. National Museum.

Laelaps berlesei Keegan, 1956r’ 264 <d, 1, $ illus.). Not Laelaps ber-
lesei Fonseca, 1938.
Laelaps keegani Thurman, ’i9^}8.’!’!"t’^t’^. ’ ’
’ ’ :’’-’ ’

’
Locality: Africa - Egypt, Giza. -’:"" ’. ’. : ’
’
.

Host: Mammal - Aryicanthis riiloti6us. .’

’

Remarks: Thurman proposed the new name keegani for berlesei

Keegan. The latter combination is preoccupied by bex’lesei (Fonseca,
1938), aBraailian mite of the genus EcM-nolaelaps. ’
Laelaps kolpakovae Zachvatkin

Laelaps (Laelaps) hilaris ’kolpakovae Zachvatkin, 1948 (d).

Locality: U.S.S.R. ’’:’- ’ : ’. ’’
".. ;i..:..
Host: Mammal - Microtus arvalis.’ : ’
’’’"
. ’ "
’

Laelaps- lamborni Hirst^ ’ ;’

’
-:

’’
’

Laelaps lamborni Hirst. 1925: 61 (d); Kieegan; 1956: 260.’ ".’

-
Locality: Africa Nyasaland, Karonga, Kenya, Rift Valley Prov-
ince; Nigeria, Onitri; Egypt, Giza.
Hosts: Mammals - rats called ’"kapuka," rat called "majancha. "
on rats, Arvicanthis niloticus, Mus musculus. Praeomys jacks oni, Bat-
;
tus rattua, Rhabdomys pumili’o. ’
"’-.’ ::’"
’
;

Laelaps lativentralis Fonseca

’
’
Type material in the Institflto Butantan, Sao Paulo, Brazil.
Laelaps lativentralis Fonseca, 1935: 22 <d); 1936: 36 (d).
-
Locality; South America Brazil, Parahyba, Joazeiro. "’..’.
’
-
Host: Mammal "Ah unidentified rat known under the common name
: .. :
of ’punari.’" :.
’
Laelaps lafrieri Taufflieb

Type^material in the Laboratory of Medical Entomology. Pasteur

’ ’. ".
’
Institute, Paris.
64

Laelaps lavieri Taufflieb. 1954: 442 (d, 9Ulus.).

Locality: Africa French Equatorial Africa, "Brazzaville.

-
Host: Mammal - Leggada musculoides.

Laelaps lavoipierrei Taufflieb

-
Type material in the Laboratory of Medica’l Entomology, Pasteur
Institute, Paris.
Laelaps lavoipierrei Taufflieb, 1954: 440 (d, S illus..).

.
Locality: Africa - French Equatorial Africa, Brazzaville.
Host: Mammal - Lophuromys sikapus-i.

Laelaps lemmi Grube

.
Laelaps lemni Grube, 1851: 38; Oudemans, 1927.
Laelaps grubei Oudemans, 1938.’
Laelaps lemmi, Bregetova, 1956: 108 (0’and $ illus.).
Locality: U.S.S.R. - Siberia.
Host: Mammal - Lemmus obensis.

’
Remarks: The above synonymy is that Of Bregetova. We have not
seen Oudeman’s 1938 paper.

Laelaps liberi-ensis Hirst ^

Laelaps liberiensis Hirst, 1925: 59 (d).

Locality: Africa - Liberia, Gpnyon Country,’ ’
Host: Mammal - Epimys defua.- "’ ’
~

Remarks: This mite seems to be intermediate between Echinolae-

laps and Laelaps: the sternal shield is’square and epigynial shield is
very slightly concave posteriorly. It was described from a single spec-
imen.

Laelaps. longiventris De Meillon and Lavoipierre

.
Laelaps longiventris De Meillon and Lavoipierre, ’1944: 65 (d).
Locality: Africa - Cape Province.
Host: Mammal - Myosorex.

Laelaps manguinhosi Fonseca

Laelaps manguinhpsi ’Fonseca, 1935: 20 (d); 1936: 34 (d).

Locality:, South America - Brazil, Matto Grosso, Porto Joffre.
Host: Mammal Holochilus vulpinus.
-
Laelaps mazzai Fonseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Laelaps mazzai Fonseca, 1938: 112, 135 (d).
 65

Locality. South America - Prwincia de Salta, Argentina.

Host: Mammal - "wild rat."
Laelaps micromydis Zachvatkin

Laelaps arvicolae Oudemans, 1927<. 200 (d); not arvicolae Oudemans,

1916.
Laelaps (Laelaps) micromydis Zachvatkin, 1948 (d); Lange, 1948: 77
(d); Bregetova. 1956: 110 (<f and $ illus.).
Locality: U.S.S.R, -
Russian Soviet Federal Socialist Republic,
Moscow. Leningrad.
Host: Mammal - Micromys minutus minutus.

Laelaps mosquensis Lange

Laelaps agilis mosquensis Lange, 1948: 77 (d).

Locality: U.S.S.R.
Hosts: Mammals - Apodemus flavicollis, Apodemus sylvaticus.

Laelaps multisplnosus Banks

Type material in the U. S, National Museum.
Leelaps multispinosus Banks, 1909: 136 (d); Banks, 1915: 84; Grant,
1947; Meyer. 1950; Knight. 1951; Bregetova. 1953; 1956: 105 (d- and
5illus.); Strandtmann, 1956: 134(t).
Liponyssus spiniger Ewing and Stover, 1915: 109 (d).
Tetragonyssus spiniger, Ewing, 1922: 6<t).
Laelaps parvanalis Willmann, 1952: 398 <d).
Locality: Europe
States. Asia U.S.S.R.
- -
Germany. North America Canada, United
Host: Mammal -
Ondatra zibetnica.
Remarks: The muskrat ’is’ the only known host of this mite.

Laelaps murie <Ljungh)

Acarug muris Ljungh, 1799: 5<d).

Laelaps muris Johnston, 1849: 367 (d);
Laelaps muris (Ljungh) Oudemans, 1027: 163 (c, d, t); Womersley. 1937:
535; Radford. 1939: 44; Grant. 1947: 6; Zachvatkin, 1948; Willmann.
1952; 392 (b, t); Bregetova. 1956: 104 (<S and 9 illus.).
Macrolaelaps muris (Ljungh) Dubinin. 1953.
Laelaps microti Oudemans, 1917: 309 (d); Oudemans. 1927; 200 (t).
- -
Locality: Australia South Australia, Adelaide. Europe British
Isles; Germany; Netherlands; U.S.S.R.
Hosts; Mammals -Arvicolaarvalis.^Arvjcola terrestris, Hydromys
chrysogaster, "long-tailed field mouse. "’Microtus amphibius, Microtus
oeconomus, Mus sylvaticus.
Remarks: The synonymy and biological limits of this species are
66

still in doubt. It should be noted that Bregetova (1956) lists Laelaps ar-
vicolae Oudemans. 1916, as a synonym. The water rat, or water vole
(Microtus amphibius), is the preferred host.

Laelaps nayasi Fonseca

Type material in the Institute Butantan, Sao Paulo. Brazil.

Laelaps nayasi Fonseca. 1938: 142 (d).
Locality: South America - Brazil, Sao Paulo.
Host: Mammal - "A wild rat. ’rato de taquaral.’ "

Laelaps novikovae Zachvatkin

Laelaps (Laelaps) muris novikovae Zachvatkin, 1948 (d).

Locality: U.S.S.R.
-
Hosts: Mammal "water rat."

Laelaps nuttalli Hirst

(Fig. 29)
Laelaps (Haemolaelaps) nuttalli Hirst, 1916: 183 (d).
Laelaps nuttalli, Berlese, 1918; Hirst. 1922; Hirst. 1926; Fonseca.
1932;Womersley. 1937; Fox. 1946: 447 (t); Pratt e^al.. 1947; Turk,
1950; Jameson etal., 1952; Strandtmann and Eben, 1953: 148 (m);
Pratt and Good, 1954: 125 (c); Smith. 1955; Domrow, 1955; Dom-
row and Smith. 1956: 202; Strandtmann, 1956: 138 (t); Bregetova,
1956: 110 ($illus.); Baker etal., 1956: 45 (illus.); Keegan, 1956.
Liponyssus bermudaensis Ewing, 1920: 290 (d).
Tetragonyssus bermudaensis. Ewing, 1922: 6 (t).
Laelaps bermudaensis, Radford, 1950.
Laelaps hawaiiensis Ewing. 1924: 8 (d); Ferris, 1933; Cole etal., 1934
(e. m); Fox, 1946: 447 (t); Randolph and Eads. 1946; ColeandKoep-
ke, 1946,
Echinolaelaps caraco Lange, 1947: 58 (d); Bregetova, 1953: 312 (t).
-
Localities: Africa Egypt, West Africa; Union of South Africa, Zu-
-
luland. Asia-India; Japan; Korea; U.S.S.R. Australia South Aus-
tralia, Queensland. Indonesia - Ceylon; Java; Malay Peninsula. Islands

-
-
of the Pacific Ocean Hawaiian Islands; Marquesan Islands. Islands of
the Atlantic Ocean Bermuda; Puerto Rico. North America - United
States. South America - Brazil; Surinam.
Hosts. Mammals - Arvicanthis dor salts, Arvicanthis niloticus,
Crocidura olivieri, Mastomys coucha, Melomys littoralis, Mus muscu-
lus, Nesokia bengalensis, Rattus conatus, Rattus coucha, Rattus culmo-
rum, Rattus hawaiiensis, Rattus norvegicus. Rattus norvegicus caraco,
Rattus rattus, Rattus rattus kandiyanus, wood rat.
Remarks; The preferred hosts of this mite are species of the genus
Rattus. The mite is found throughout tropical and warm temperate zones
wherever its host is found, with the apparent exception of Europe. We
know of no published record of its occurrence there. Because of its abun-
 67

dance on domestic rats it has been suspect as a disease carrier but as

far as we know, no one has yet demonstrated a disease. Cole and Koepke
(1946) demonstrated a positive correlation in the prevalence of this mite
with the prevalence of murine typhus, but Strandtmann and Eben (1953)
were unable to demonstrate either a natural or a laboratory infection of
mites with murine typhus.

Laelaps oraniensis Hirst

Laelaps oraniensis Hirst, 1925: 60 (d).

-
Locality: Africa Algeria. Oran, Mount Marabut.
Host: Mammal - "Field mice."

Laelaps oryzomydis Pratt and Lane

Type material in the U. S. National Museum.

Laelaps oryzomydis Pratt and Lane, 1953: 358 (d).
-
Locality: North America Southeastern United
Host: Mammal - Oryzomys palustris.
States.

Laelaps pachypus Koch

Laelaps pachypus Koch, 1839: #24 (d); Hirst, 1916; Oudemans. 1927:
179 (d); Willmann. 1952.
Liponyssus setiger Ewing, 1920: 290 (d). New synonymy.
Tetragonyssus microti Ewing, 1933: 9 (d); Augustson, 1941.
Laelaps kochi Oudemans, 1936: #101 (t); Grant, 1947; Jameson, 1947;
Jameson, 1950; Jameson et al., 1952.
Laelaps (Hyperlaelaps) amphibius Zachvatkin, 1948.
Hyperlaelaps amphibius, Dubinin, 1953; Bregetova, 1956: ll7(rfand$
illus.).
-
Localities: Asia - Manchuria. Europe Germany; Great Britain;
-
Netherlands; Russia. North America Alaska; Canada; United States.
Hosts: Mammals - Apodemus sp., Arvicola amphibius, Arvicola
arvalis, Blarina brevicauda, Clethrionomys glareolus, Lemmus arvalis,
Microtus agrestis. Microtus amphibius, Microtus californicus, Micro-
tus montanus. Microtus mordax, Microtus ochrogaster, Microtus oecon-
omus. Microtus orcadensis, Mustela vulgaris, Peromyscus sp.. Sorex
obscurus, Sorex vagrans.
Remarks: Oudemans (1936) states that Koch thought he was redes-
cribing Acarus pachypus Herman when he illustrated and wrote the des-
cription of this mite, and that therefore Laelaps pachypus Koch is a jun-
ior homonym. Oudemans then proposed the name kochi. However, since
Koch used the name Laelaps pachypus without reference to Acarus, Oud-
eman’s decision is not consistent with the International Rules of Zoolog-
ical Nomenclature.
Zachvatkin (1948) does not agree that pachypus of Oudemans (1927)
is the same as that of Koch and therefore proposes his new name. He
68

may be quite right but according to the rule of first reviser, it is desir-
able to retain the species as Oudemans has described it. Koch’s type is
lost and his description and illustration are insignificant. The name
pachypus must therefore be applied to this species or be relegated to a
name for a species inquirendae. Such action could be justified only by
a suspension of the Rules of Zoological Nomenclature.

Laelaps parvulus Hirst <

Laelaps parvulus Hirst, 1923: 691 (d); Hirst, 1925; Bedford, 1932; Had-
ford. 1942.
Locality: Africa - Union of South Africa. Cape Province, Grahams-
town.
-
Hosts: Mammals Arvicanthis dorsalis. Myosorex varius, Otomys
irroratus.

Laelaps paulistanensis Fonseca

Laelaps paulistanensis Fonseca, 1935: 19 (d); Fonseca, 1936: 33 (d).

-
Locality: South America Brazil, Sao Paulo,
-
Host: Mammal "Wild rat."

Laelaps pavlovskyi Zachvatkin

Laelaps (Laelaps) pavlovskyi Zachvatkin, 1948 (d); Lange, 1948: 77.

Laelaps pavlovskyi pavlovskyi, Lange, 1948: 77 (d),
Laelapa pavlovskyi, Bregetova, 1953; 1956: 110 (Stilus.).
Locality: U.S.S.K.
-
Host: Mammals Apodemus agrarius, Apodemus speciosus, Mi-
crotus oecononius.

Laelaps pitymidis Lange

Laelaps pity midis Lange, 1955; Bregetova, 1956: 108 ($ illus.).

Locality: U.S.S.R, - Northern Caucasia.
Host: Mammal - Microtus taajori.

Laelaps robu stipes Ewing

Type material in the U. S. National Museum.

Laelaps robustipes Ewing, 1925: 4 (d),
Locality: South America - Argentina, Buenos Aires.
-
Host: Mammal "A rodent. "
Remarks: The type specimen is in very poor condition, but it does
indicate this is a true Laelaps.
 69

Laelaps roubaudi Taufflieb

Type material in the Laboratory of Medical Entomology. Pasteur

Institute, Paris.
Laelaps roubaudi Taufflieb, 1954: 437 (d. ?illus.).
Locality: Africa - French Equatorial Africa, Brazzaville.
Host: Mammal - Dasymys incomtus.

Laelaps semitectus (Koch). New combination.

Sejus semitectus Koch, 1879: 123 (d).

Liponyssus semitectus, TragSrdh. 1904: 31 (d).
Ichoronyssus semitectus, Ewing, 1922: 13; Fonseca, 1948.
Laelaps pachypus Koch, Kramer. 1883, misidentification.
Laelaps hilaris Koch, Kramer, 1883, misidentification.
-
Locality: Holarctic Greenland, Siberia.
Host: Mammal My odes torquatus.
"

Remarks: Tragardh is responsible for the above synonymy through

1904. Koch’s types are no longer extant, so we must go to Tragardh’s
figures and description, which show clearly that this mite is a Laelaps.

Laelaps sicula Willmann

Laelaps hilaris C. L. Koch subspec. sicula Willmann, 1955: 182 (d, $).
-
Locality: Europe Italy, Sicily.
-
Hosts: Mammals Mus musculus brevirostris, Rattus noryegicus.

Laelaps simillimus Zumpt

Type material in the South African Institute for Medical Research.

Laelaps simillimus Zumpt. 1950: 81 (d).
Locality: Africa - Union of South Africa, Transvaal.
Host: Mammal - Thallomys namaquensis.

Laelaps sminthopsis Womersley

Laelaps sminthopsis Womersley. 1954; 117<d);DomrowandSmith, 1956.

Locality: Australia - Victoria. Mt. Glorious.
Hosts: Mammals " Sminthopsis leucopus, Antechinus flavipes.
Remarks? This is a very large mite (1.3 mm), with long. dense
setation, much of it plumose. Further study may prove it not to belong
in the genus Laelaps.

Laelaps soricis Oudemans

Laelaps soricis Oudemans, 1925: 29 (d); Oudemana, 1927b: 185 (d).

Locality: Indonesia - Ambon.
Host: Mammal - On a "tikoes moende, " (Soricidae).
70

Remarks: The setae are long and none seem to be thornlike. The
species may be a Tricholaelaps.

Laelaps spinifer Taufflieb and Mouchet

Type material in the Laboratory of Medical Entomology of the Pas-

teur Institute, Paris.
Laelaps spinifer Taufflieb and Mouchet, 1956: 302 (d, $ illus.).
Locality: Africa - French Camerqun, Yaounde.
-
Host: Mammal Arvicanthis rufinus.

Laelaps taprobanius (Turk). New combination

Macrolaelaps taprobanius Turk, 1950: 65 (d).

Locality: Indonesia - Ceylon.
Host: Bandicota malabarica.

Laelaps thamnomys Taufflieb

Type material in the Laboratory of Medical Entomology, Pasteur

Institute, Paris.
Laelaps thamnomys Taufflieb, 1954: 444 (d. Stilus.).
-
Locality: Africa French Equatorial Africa. Brazzaville.
-
Host: Mammal Thamnomys rutilans.

Laelaps thompsoni (Turk). New combination

Haemolaelaps thompsoni Turk. 1950: 67 (d).

Locality: Indonesia - Ceylon.
Host: Mammal - Rattus rattus kandiyanus.

Laelaps thori Fonseca

Type material in the Institute Butantan, Sao Paulo, Brazil.

Laelaps thori Fonseca, 1938: 111, 133 (d).
-
Locality: South America Brazil.
Host: Unknown,

Laelaps transvaalensis Zumpt

Type material in the South African Institute for Medical Research.

Laelaps transvaalensis Zumpt. 1950: 82 (d).
Locality: Africa - Union of South Africa, Transvaal.
Host: Mammal - Otomys irroratua.

Laelaps turkestanicus Lange

Laelaps turkestanicua Lange, 1955; Bregetova. 1956: 106 (tf, Stilus.).

 71

Locality: Asia U.S. S.R., Tadzhikistan.

Host: Mammal - Rattus turkestanicus.
Laelaps volgensis Lange

Laelaps agilis volgensis Lange. 1948: 77 (d).

Locality: U.S. S.R.
Host: Mammal - Apodemus silvaticus.

Laelaps wetroorei Ewing

Type material in the U. S. National Museum.

Laelaps wetroorei Ewing. 1925: 4 (d).
-
Locality: South America Argentina, Buenos Aires.
Hosts: Mammals " "rats. "
Laelaps zumpti Keegan

Type material in the U. S. National Museum.

Laelaps zumpti Keegan, 1956: 263 (d. <?, 9, Nil, $Illus.).
-
Locality: Africa Kenya.
-
Hosts: Mammals Lemniscomys striatus, Mus triton (type host).

Genus Longolaelaps Vitzthum, 1926

(Fig. 30)
Type: Longolaelaps longulus Vitzthum.
-
Diagnosis: Body long and narrow; coxae I 1X1 with spurlike setae;
epigynial plate slightly expanded and bearing four pairs of setae; legs I
and II much heavier than legs III and IV. This is a monotypic genus of
which only females are known.

Longolaelaps longulus Vitzthum

(Fig. 30)
Longolaelaps longulus Vitzthum, 1926: 74 (d).
-
Locality: Indonesia Sumatra, Wai Lima.
-
Host: Mammal Rattus whiteheadi,

Genus Oryctolaelaps Lange, 1955

(Fig. 31)
Type: Oryctolaelaps Mbikovae Lange
Diagnosis: According to the illustrations, this monotypic genus has
the general facies of Laelaps. The female sternal plate is reduced and
the third sternal setae may be off the plate. The epigynial plate bears
only three pairs of setae. The anal plate is roundly triangular. The
dorsal plate covers the dorsum almost completely and has concave mar-
gins. The male holoventral plate is just barely expanded behind coxae
72

IV, somewhat as in Hirstionyssus. It differs from Steptolaelaps in hav-

ing a broader anal plate, a larger and differently shaped dorsal plate,
and in lacking heavy spurlike setation.

Oryctolaelaps bibikovae Lange

Oryctolaelaps bibikovae Lange, 1955; Bregetova, 1856: 119 (tf and $ il-
lus.).
Locality: Asia - U. S. S. R., Primorsky region.
-
Host: Mammal Mogera robusta.

Genus Echinolaelaps Ewing, 1929

(Figs. 14. 15)
Type: Laelaps echidninus Berlese.
Macrolaelaps Ewing, 1929: 185.
Type: Laelaps sanguineus Vitzthum (sic). Lapsus calami
for sanguisugus.
Diagnosis: Large mites, one mm or more long. In the female the
epigynial plate is expanded behind coxae IV and frequently reaches to
the anal plate where it may conform to the anterior margin of that plate.
The female sternal plate is as long as, or longer than, wide and both
the anterior and posterior margins are generally somewhat projected.
The epipharynx is expanded toward the apex and has a medial longitudi-
nal groove. The hypostomal processes are modified into what appear
to be clasping structures.
Bernarks: Captain Vernon Tipton has pointed out to us that Macro-
laelaps Ewing, 1929, has page priority over Echinolaelaps Ewing, 1929.
the type species of both representing the same genus. However, in view
of the relative frequency of use of the two names, we are selecting Echi-
nolaelaps as the name to be used for that genus.

E chinolaelaps^ echidninus (Berlese)

(Pigs. 14, 15)
Laelaps echidninus Berlese, (fasc. 39) 1887 (d); Miller, 1908: 25 (m);
Hirst, 1913; Hirst, 1914: 119 (d); Hirst, 1915: 224; Banks. 1915;
Hirst. 1916; Hirst. 1922; Ewing. 1924; Hirst, 1926; Vitzthum, 1926;
Oudemans. 1927; Oudemans. 1929; Stanley. 1931: 1 (a); Mosser et
al.. 1931: 567 (m); Perris, 1933; Womersley, 1837; Radford. 1939;
Buitendl.jk. 1945; Lavoipierre, 1946; Grant. 1947; Alicata, 1948;
Jameson et al., 1952; Fujisaki, 1953; Strandtmann and Eben, 1953;
Domrow, 1955; Willmann. 1955; Bregetova. 1956: 106; Keegan,
1956.
Echinolaelaps echidninus, Ewing, 1929: 11, 185 (d. t); FonsecaandPra-
do. 1932; Mehta, 1937; ColeandKoepke, 1946; Finnegan. 1945; Fox.
1946; Randolph and Eads. 1946; Pratt and Fritz, 1947; Schweitzer.
1949; Turk, 1950; Morlan and Utterback. 1952; Smith, 1955; Baker
etal., 1956; Owen, 1956: 702 (b).
 73

Echinolaelaps echidninus sub sp, vitzthtnai Turk, 1950: 71 (t).

New name for Laelaps echidninus Vitzthum. 1926, notBerlese, 1887.
Echinolaelaps hirst! Turk, 1950: 71 (t). New name for Laelaps echid-
ninus Hirst, 1913. not echidninus Berlese, 1887.
Echinolaelaps hirsti subsp. ceylpnicus Turk, 1950: 72.
Localities: Africa - Belgian Congo; Egypt. Asia - India; Japan;
Korea. Australia - New South Wales. Queensland, Victoria. Europe -
British Isles; Italy; Switzerland. Indonesia - Borneo; Ceylon. Islands
-
of the Pacific Dutch East Indies; Marquesan Islands; Ocena Island; Ha-
waii; Samoa; Rosa Island. NorthAmerica - Mexico; UnitedStates. South
-
America - Brazil. West Indies Puerto Rico.
Hosts: Mammals - "brown rat, " "Marquesas rat, " Mus musculus,
Mus musculus molossinus, Mus norvegicus, Mus rattus, Mus rattus
alexandrinus. Nesokia bengalensis. "opossum. " Paradoxus hermaphro-
ditus, Perameles gunni, "roof rat. "Rattus crassus, Kattus culmorum,
Rattus norvegicus, Rattus norvegicus otomi, Rattus rattus, Rattus rat-
tus erythronotus. Rattus sp., "Rongeurs sylvatique. " "Texas cotton-
tail, " "white rats."
Remarks: The preferred host of this mite is the common brown rat,
Rattus norvegicus, but it will live as well on other members of the genus
Rattus. It is not known to bite man and is never found on birds. The
records from mammals other than Rattus probably represent accidental
and temporary associations. It is possible that all forms found on Rat-
tus are referable to this species.
Echinolaelaps echidninus is the intermediate host of the protozoan
Hepatozoon muris, a fatal disease of white rats (Miller. 1908). It is
apparently Incapable of harboring or transferring rickettsia of the typhus
group (Mosser et al., 1931). The life cycle has been worked out recently
by Owen (1956).

Echinolaelaps aragonensis Fonseca. New combination

Type material in the Institute Butantan, Sao Paulo, Brazil.

Laelaps aragonensis Fonseca, 1938: 108, 138 (d).
Locality: South America - Brazil, Goyas, Anapolis.
Host: Mammal - a rat.

Echinolaelaps atypicus Turk, New combination

Laelaps atypicus Turk, 1950: 63 (d).

-
Locality: Indonesia Ceylon,
Host: Mammal ~ Coelomys bicolor.
Remarks; The posterior margin of the female epigynial plate is
concave, and conforms with the anal plate, but the sternal plate is wider
than long. Each coxa is said to have only one seta. No pilus dentilis
is indicated in the figures.
74

Echinolaelaps berlesei (Fonseca). New combination

Type material in the Institute Butantan. Sao Paulo, Brazil.

Laelaps berlesei Fonseca, 1938: 104, 126 (d).
Locality: South America - Brazil, Sao Paulo.
Host: Mammal - Galictis vitatta.

Echinolaelaps giganteus (Berlese). New combination

Laelaps giganteus Berlese, 1918: 129(d); Hirst, 1925: 67; Bedford, 1932;
Keegan. 1956: 253 (? illus.).
Macrolaelaps giganteus, Radford, 1950: 369.
-
Locality: Africa Kenya; Liberia; Nigeria; Uganda; Union of South
Africa, Zululand, Transvaal; Belgian Congo; Sudan; Rhodesia,
Hosts: Mammals - Arvicanthis abyssinicus. Arvicanthis dor salts,
Dasymys incomtus, Dasymys rufulus, Dephomys defua. Lemniscomys
barbarus, Lemniscomys griselda, Lemniscomys macculus, Lemnis-
comys striatus, Mastomys coucha, Praeomys tullbergi, Rattus tulbergi,
Rhabdomys pumilis, Tatera schinzi.

Echinolaelaps grandis (Hirst). New combination

Laelaps grandis Hirst, 1925: 62 (d); Keegan, 1956.

Macrolaelaps grandis, Radford, 1950: 369.
Locality: Africa - Kenya; Uganda.
-
Hosts: Mammals rat, rodents.

Echinolaelaps muricola (Tragardh). New combination

Laelaps muricola Tragardh. 1910: 55 (d); Berlese, 1918: Hirst, 1925:

63 (d); Bedford, 1932; Keegan, 1956.
Macrolaelaps muricola. Radford, 1943; Lavoipierre, 1946; Radford,
1950: 369.
Localities: Africa - Abyssinia; Belgian Congo; British Sonaaliland;
Union of South Africa, Cape Province, Zululand; Gold Coast; Kenya,
Tanganyika, Nigeria; Nyasaland; Sudan; Uganda.
Hosts: Mammals - Aethomys chrysophilus, Arvicanthis dorsalis,
Choeromys gregorlanus. Cricetoinys sp., Lemniscomys striatus, Lo-
phiomys ibeanus, Mastomys coucha. Mug coucha, Mus crysophyllus,
Mus hildebrandtl, Praeomys jacksoni. Rattus rattus, Saccostomus cam-

Echinolaelaps pallidus (Tragardh)

Laelaps pallidus Tragardh, 1931: 616 (d).

Echinolaelaps hirsti subsp. pallidus Turk, 1950: 71,
Locality: Islands of the Pacific Ocean - Juan Fernandez Islands.
-
Host: Mammal Rattus rattus.
 75

Echinolaelaps sanguisugus (Vitzthum). New combination

Laelaps (Laelaps) sanguisugus Vitzthum, 1926: 58 (d).

Type of Macrolaelaps Ewing.
Macrolaelaps sanguineus. Ewing. 1929: 185 (Lapsus calami for sangui-
sugus Vitzthum); Ewing, 1933.
Locality: Indonesia - Java.
Host: Mammal - Mus lepturus.

Echinolaelaps sculpturatus (Vitzthum). New combination

Laelaps (Laelaps) sculpturatus Vitzthum. 1926: 64 (d).

Macrolaelaps sculpturatus, Radford, 1943; Turk, 1946; Radford. 1950:
369.
-
Locality: Indonesia Sumatra, Wai Lima.
-
Host: Mammal Rattus whiteheadi.

Echinolaelaps ugandanus (Hirst). New combination

Laelaps ugandanus Hirst, 1923: 971 (d); Hirst, 1925: 63 (illus.).

Macrolaelaps ugandanus, Fonseca, 1938: 11 (t); Radford, 1943; Radford,
1950.
Locality: Africa - Uganda.
Host: Mammal - An unidentified rodent.

Echinolaelaps vansomereni Hirst, New combination

Laelaps vansomereni Hirst. 1923: 690 (d); Hirst, 1925: 56; Bedford.
1932; Zumpt, 1950: 78; Keegan. 1956.
Locality: Africa - Kenya; Uganda; Southern Rhodesia; Sudan. Torit;
Union of South Africa, Zululand, Northern Transvaal.
-
Hosts: Mammals Aethomys chrysophilus, Aethomys namaquensis,
Mastomys coucha, Mus chrysophilus, Saccostomus campestris, Suncus
varilla.

Echinolaelaps yaoundensis (Taufflieb and Mouchet). New combination

Type material in the Laboratory of Medical Entomology, Pasteur

Institute, Paris.
Laelaps yaoundensis Taufflieb and Mouchet. 1956: 304 (d. Stilus,).
Locality: Africa - French Cameroun. Yaounde.
Host: Mammal - an unidentified rodent.
Remarks: Unfortunately, the illustration for this species in the
original publication is, by mistake, that of Andreacarus zumpti. A cor-
76

rection sheet published separately gives the correct illustration.

Genus Mysolaelaps Ponseca, 1935

(Pig. 32)
Type: Mysolaelaps parvispinosus Fonseca.
Diagnosis: Large species, well chitinized; epigynial plate broadly
expanded behind, with a straight posterior edge and with four pairs of
minute bristles.

Mysolaelaps parvispinosus Fonseca

(Fig. 32)
Type material at the Butantan Institute. Sao Paulo, Brazil.
Mysolaelaps parvispinosus Fonseca, 1935: 17 (d, ).
Locality: Brazil.
Host: Mammal - wild rat.

Mysolaelaps microspinosus Fonseca

Type material at the Butantan Institute, Sao Paulo, Brazil.

Mysolaelaps microspinosus Fonseca, 1935: 18 (d, ).
Locality: Brazil.
Host: Mammal - rat.
Mysolaelaps rothschildi (Hirst). New combination

Laelaps rothschildi Hirst. 1914: 325 (d); Domrow and Smith, 1956: 202.
Echinolaelaps rothschildi, Turk. 1950: 71.
Laelaps melomys Womersley, 1937: 534 (d); Domrow and Smith, 1956:
202.
Locality: Islands of the Pacific Ocean - Dutch New Guinea. Aus-
tralia - Queensland.
Host: Mammals
Melomys littoralis, Melomys
- Mus cervinipes.
sp,, Uromys lorentzi, Uroa’g’s stajkeri,
Rattus assimilis.

Genus Aetholaelaps Strandtmann and Camin. 1956

(Fig. 33)
Type; Aetholaelapa sylstrai Strandtmann and Camin
Diagnosis: Mites of medium size. The female epigynial plate has
four pairs of setae. The sternal plate is large and about two-thirds as
broad as long. The male ventral plates are united and slightly expanded
posterior to legs IV. The spermatodactyl is long and scimitar shaped
and typically laelaptid. Legs I and II are heavier than III and IV andhave
several heavy, spurlike setae.

Aetholaelaps sylstrai Strandtmann and Camin

(Fig. 33)
Type material in the U. S. National Museum.
 77

Aetholaelaps sylstrai Strandtmanajand Camia, 1956: 152 (d, d, $ and

nymphs, d’ and 9 illus.).
-
Locality: Africa Madagascar, by way of the Zoological Society of
San Diego, California, via New York harbor.
Host: Mammal - Lemur mongoz.

Family DERMANYSSIDAE Kolenati, 1859

Description of the family. A rather heterogeneous group of small

to medium mites. Many species are capable of great distention in the
nymphal and female stages. The tectum is an elongated, tonguelike,
transparent structure that may have an acuminate tip or a few small
apical denticles. The tritosternum is always present and typically has
a hyaline, serrated margin. Peritremes vary in length but they always
extend at least beyond coxa III. The dorsal plate is generally entire
but may be divided into two subequal plates (Steatonyssus and Pellonys-
sus) or into a large podosomal and tiny pygidial plate (Qphionyssus,
Allodermanyssus). The chelae vary from minute (Dermanyssus) to
nearly a third the length of the chelicerae, but are always hyaline and
poorly sclerotized. Denticles and a pilus dentilis are rarely present;
when they are, they are hardly obvious. The deutosternal denticles are
always in single file or in an alternating double file, never in combs.
The corniculi are poorly sclerotized and difficult to see. They may be
formed into half tubes and perhaps function as a preoral siphon. The
anterior dorsal spur of coxa II is generally quite long and narrow. Ac-
cessory spurs or protuberances on the coxae are common; accessory
spurs on the other leg segments are not infrequent. The typical life
cycle consists of egg, laa*va» two nymphal stages and adult. Some spe-
cies, however, are apparently ovoviviparous (Hirstionyssus Stafford!)
and in others the deutonymph is a nonfeeding stage of very short dura-
tion (Ornithonyssus spp.).
Members of this family parasitize birds, mammals, and reptiles.
They are primarily nest inhabitants and are the most harmful of the ga-
masine mite parasites.

Key to the Subfamilies and Genera of the Dermanyssidae

1. Anal plate of female more than half as wide as the opisthosoma;

concave on the anterior margin, Chelicerae attenuated in both
sexes, chelae long; male chela lacking spermatodactyl . . . .
. . . . . . . . . . . . . . . . . . . . . MYONYSSINAE Myonyssus

Anal plate not half as wide as the opisthosoma; not concave on

the anterior margin. If chelicerae are attenuated, then chelae
are minute; male chela with spermatodactyl . , , . . . , . . . 2
2. Chelicerae attenuated, the apical two-thirds or more niu<!h.nar-
rower than the basal portion; chelae minute . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . DERMANYSSINAE 3

Chelicerae of uniform diameter throughout their length, if not,

chelae are obvious . . . . . . . . . . . . . MACRONYSSINAE 5

3. Dor sum with two plates, the posterior small, pygidial. Anal
plate oval, anterior margin rounded. Parasitic only on mam-
mals . . . . . . .. . . . . . . . . . . . . . . . . Allodermanyasus

Dorsal plate entire . . . . . . . . . . . . . . . . . . . .. . . 4

4. Female sternal plate much wider than long; male holoventral

.
. . . . . . . . . . . . ..
plate slightly expanded behind coxae IV. Parasitic on birds. .
. . . . . . . . . . . . . . Dermanyssus

Female sternal plate as long or longer than wide; male holo-

ventral plate not expanded behind coxae IV, Parasitic on mam-
mals . . . . . . . . . . . . . . . . . . . . . . . . . Liponyssoides

5. Two dorsal plates . . . . . . . . . . . . . . . . . . . . . . . . 6

Only one dorsal plate . . . . . . . . . . . . . . . . . . . . . . 9

6. Dorsal plates subequal; posterior plate narrower. Parasitic

primarily on bats and on birds . . . . .. . . . . . . . . . . . 8

Posterior plate very small, pygidial. Parasitic on reptiles . . 7

7. Posterior dorsal plate with 2 - 3 pairs of setae. Usually found

on lizards . . . . . . . . . . . . . . . . . . . . .. . Sauronyssus

Posterior dorsal plate bare. Usually found on snakes . . ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . pphionyssus
8. Sternal plate of female about twice as wide as long, with a
strongly sclerotized posterior margin. Sternal setae I about
as large as sternal setae II. Chelicerae not especially long or
attenuate. Parasitic on bats . . . . . . . . . . . . . Steatonyssus
Sternal plate of female very narrow, arched, and without a
sclerotized posterior margin. Sternal setae I much smaller
than sternal setae II. Female chelicerae rather long and no-
ticeably attenuated. Parasitic on birds . . . . . . . Pellonyssus .
9. With anterior, dorsal, hooklike projection over the gnathosoma
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinonyssus
 79

Not as above . . . , . « . , . . .. ./.. « . . . . . . , . . . , . . 10

10. Some coxae with ventral or posterior spurs . . . . . . . . . . 11

No ventral or posterior coxal spurs, but coxal setae may be

heavy and thornlike . . . . . . . . . . . . . . . . . . . . . . 14

11. Some coxal spurs setigerous . . . . . . . . . . . . . . . . . . 13

Coxal spurs not setigerous . . . . . . . . . . . . . . . . . . . 12

12. Epigynial plate with four pairs of setae, Peritreme lacking or
very short . . . . . . . . . . . . . . . . . . . . Australolaelaps .
Epigynial plate with one or two pairs of setae, Peritreme ex-
tending at least beyond anterior margin of coxa HI. Hirstionyssus

13. Females with spur on palpal trochanter; epigynial plate pointed

posteriorly. Male ventral plate undivided, confluent with the
anal . . . . . . .. . . . . . . . . . . . . . . . . . Neoichoronyssus

Palpal trochanter of female lacking spur; epigynial plate rounded

posteriorly. Male ventral plate divided just behind coxa IV. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patrinyssus

14. Epigynial plate with five or more setae . , . . . » . . . . , . 15

Epigynial plate with only one pair of setae. (Occasionally an-

otherpairmaybe so close as to appear to be on the plate.). . . 17

15. With five setae on the epigynial plate . . . . . . . Neospinataelape

With more than five setae on the epigynial plate . . . . . . . . 16

16. Sternal plate with two large, stigmalike structures . . . Hirstesia

Not as above .. . . . . . . . . . .. . . . . . . . . . . . . . . 22

17. Havingtwo large, stigmalike structures on the sternal plate. . .

. . . . . . . . . . . . . . . , . . Lepronyssoides

Sternal plate without large, circular stigmalike structures . . 18

18. Epigynial plate narrow, pointed; dorsal plate broad anteriorly

and tapering posteriorly . . . . . . . . . . . . . . . . . . . . . 19

Epigynial plate rounded or pointed posteriorly; dorsal plate gen-

erally parallel-sided and covering most of dorsal surface in un-
80

engorged specimens 20

19. With spur on ventral surface of the palpal trochanter. Sternal

plate with three pairs of setae (except in 0. sylviarum). Para-
sitic only on birds and mammals . . . . . . . . . . Ornithonygsus

Without spur on palpal trochanter. Sternal plate bearing only

two pairs of setae and generally not nearly as wide as in Ornith-
onyssus. Parasitic only on reptiles . , . . . . . . » Sauronyssus

20. Legs I and II unusually thick and heavy, especially legs II » .

. . . . . . . . . . . . . . . . . . .. . . . . . . . . . . Radfordiella

Legs I and II not exceptionally thick . . . . . . . . . . . . . . 21

21. Coxae without tubercles or elevations. Ventral plates of male

fused into single holoventral plate . . . . . . . . . . Macronyssus

Coxae II and III each with a slight semicircular protuberance.

Ventral plates of male divided into sternogenital and a ventro-
anal plate. Parasitic on bats only . . . .
. . . . . . Ichoronyssus

22. Epigynial plate with six setae . . . . . . . . . . . . . . . . . . 23

Epigynial plate with more than six setae; male ventral plate di-
vided into sternogenital and ventroanal plates. Parasitic on
bats . . . . . . . . . . . . . . . . . . . . . . . . . . Spinolaelaps

23. Peritreme unusually wide; tritosternum very broad at base;

gnathosomal setae heavy, spurlike. Male anal plate separate
from remainder of ventral plate complex . . . . . . . . Neolaelaps
Not as above . . . . . . . . . . . . . . . . . . . . . . . Liponysella

Subfamily MACRONYSSINAE Oudemans, 1936

Diagnosis: Dermanyssid mites in which the female chelicerae are

of uniform diameter throughout their length and the chelae obvious. Both
armsofthe chela are present and essentially equal. The male holoven-
tral plate is entire or divided; the male spermatodactyl is about twice
the size of the moveable arm; the fixed arm of the male chela is always
present and is generally as long as the moveable arm.
An excellent review of this group of mites was written by Dr. Flavio
de Fonseca (1948) under the name Macronyssidae.
 81

Genus Macronyssus Kolenati, 1858

(Fig. 34)
Type: Caris longimana Kolenati, 1856.
Diagnosis: Female epigynial plate rounded posteriorly, holoventral
plate of male entire and slightly expanded behind coxae IV; tibia I and II
long; coxae devoid of ventral spurs.

Macronyssus longimanus (Kolenati)

Caris longiroana Kolenati. 1856; Kolenati, 1857: 16.

Liponyssus longimanus. Hirst. 1921: 796 (d>.
Macronyssus longimanus, Kolenati, 1858: 5; 1859: 178(d); Murray,
1876: 173; Fonseca, 1941; Fonseca. 1948: 269 (d).
Liponyssus ellipticus Kolenati, 1856; Hirst, 1921: 795 (d); Fonseca.
1948: 309.
Locality: Africa - Egypt.
Hosts: Mammals - Miniopterus schreibersii, Rhinolophus clivosus.
Rousettus aegyptiacus.
Remarks: Kolenati presumably first described longimanus in 1856,
but we were unable to locate this reference.

Macronyssus orcadensis (Turk)

Ichoronyssus orcadensis Turk, 1945: 786 (d).

Macronyssus orcadensis, Fonseca. 1948: 309.
Locality: Europe - England, Zoological Gardens.
-
Host: Mammal Microtus orcadensis,

Genus Ornithonyssus Sambon. 1928

(Figs. 35. 36)
Type: Dermanyssus sylviarum Canestrini and Fanzago.
1877
Liponyssus of authors, not of Kolenati, 1856,
Macronyssus of authors, in part, not of Kolenati, 1856.
Leiognathus Canestrini, 1884. Preoccupied by Leiognathus La-
cepede, 1802 (Pisces).
Type: Dermanysszis sylviaruoa Canestrtoi and Fanzago, 1877
Bdellonyssus Fonseca, 1941: 264.
Type: Leiognathus bacoti Hirst, 1913.
Fonsecaonyssus Radford, 1950: 223,
Type: Lsiognathus sylviarum, Canestrini and Fanzago. 1877
Diagnosis: Dorsal plate tapering posteriorly in the female, broadly
rounded in the male; female epigynial plate narrow and pointed posteri-
orly. male holoventral plate undivided and not expanded behind coxae IV;
female palpal trochanter with or without a palpal spur, male palpal fe-
mur generally with a prominent seta on a noticeable elevation; all coxae
devoid of ventral spurs or spines.
82

Remarks: Many of the mites of this genus are among the most im-
portant acarine parasites. Ornithonyssus bacoti readily attacks man
and has been implicated as a disease transmitter; it also acts as the vec-
tor of filarial worm in cotton rats. 0. burs a and 0. sylviarum may be-
come so abundant in bird nests as to kill the young birds by exsanguina-
tion.

Ornithonys sus sylviarum (Canestrini and Fanzago)

(Fig. 35)
Dermanyssus sylviarum Canestrini and Fanzago, 1877 (d).
Leiognathus sylviarum, Canestrini, 1884; Ewing, 1922; Sambon, 1928;
Fonseca. 1948: 259 (review).
Liponyssus sylviarum, Vitzthum, 1918: 17 (d); Hirst, 1922; Hirst, 1923;
Vitzthum, 1926; Payne, 1929: 819 (control); Cutright. 1929: 422
(control); Gibson, 1930; Taylor. 1930; Bishopp and Wood, 1931;
Maw, 1935: 79 (b); Brody. 1936 (v); Peters, 1936; Cameron, 1938:
230 (b, c, review, control); Radford, 1939; Spencer. 1941; Bunyear,
1941; Herms. 1939; Reeves etal., 1947 (m); Hammon, Beeves, et
al., 1948: 92 (m); Hofstad. 1949: 370 (v); Zumpt. 1950; Boyd, 1951.
Ornithonyssus sylviarum. Sambon, 1928: 105 (t); Bregetova, 1956: 165;
Baker et al., 1956: 26.
Macronyssus Bylyiarum. Buitendijk, 1945.
Bdellonyssus sylviarum, Strandtmann and Hunt. 1951: 467 (t); Furman,
1952: 926 (control); Furman, 1953: 822 (control); Jameson, 1952;
Bregetova. 1953; Chamberlain ertal., 1955: 106 (m); Reeves etal.,
1955: 93 (m).
Fonsecaonyssus sylviarum. Radford, 1950: 223 (t); Womersley, 1956:
595.
Lophotes patavinus Megnin, 1891. (Cited by Ponseca, 1948: 304).
Liponyssus americanus Banks, 1906: 136 (d); Ewing, 1922; Ewing. 1947:
87 (t).
Liponyssus canadensis Banks, 1909: 134 <d); Ewing, 1922; Gibson, 1930;
Ewing. 1947: 87 (t).
Liponyssus pacificus Ewing, 1922:1 (d);Augustson. 1941; Furman, 1948:
27 <t).
.
Localities: Asia-Japan; Korea; U.S.S.R. Australia -Adelaide.
- -
Europe Germany; Italy, Spain. North America Canada, BritishCo-
lumbia, Ontario; United States,
Hosts: Mammals - Eutamias alphinus. Homo sapiens, Mus muscu-
-
lus. Birds brewer’s blackbird, Carduelis carduelis weigoldi, chick-
ens, kingbird, meadowlark, pigeons, poultry, purple grackle. rod-eyed
vireo, robin, rook. Riparia yiparia, sparrow, starling, swallow, yelltw-
headed blackbird.
Remarks: Known as the Northern fowl mite, this species is most
common in the north temperate zones where it becomes a very serious
pest of domestic fowl and wild birds. It apparently does not attack mam-
mals except man, and then not readily. Brody (1936) found sylviarum
capable of transmitting fowl pox in the laboratory. Reeves et al. (1947)
 83

and Hammon and Reeves (1948) found it, taken from.various wild birds,
to be naturally infected with Western equine encephalitis, .and with mixed
infections of Western equine encephalitis and St. Louis encephalitis. Hof-
stad (1949) was able to infect the mite with Newcastle disease of chickens
but could not transfer it through the bite alone. Chamberlain arid Sikes
(1955), and Reeves et al. (1955), after .exhaustive tests, independently
reached the conclusion that sylviaruna is unimportant as a reservoir of
the equine encephalrfides.

Ornithonyssus bacoti (Hirst)

Leiognathus bacoti Hirst, 1913: 119 (d); Hirst. 1914 (c).

Liponyssus bacoti. Hirst, 1920; Ewing, 1922: 19 (c); Bishopp, 1923: 1
(b, ro); Cory, 1923 (c); Hirst, 1925; 1926; Holdaway, 1926; Sambon.
1928; Shelmire’.sin.d Dove. 1930: 115 (m); Dove and Shelmire, 1931:
1506 (rn); Shelmire. and Dove. 1931: 579 (b. m); Oudemans, 1931;
Bedford. 1932; Dove and Shelmire, 1932 (m); Ewing.. 1932; jFonseca,
1932;Fonseea, l’935-;Mdntosh, 1935;0hmori, 1936(b); Mehta, 1937:
353 (m); Smith and Mehta, 1937: 349 (m); Spencer. 1937: 44 (b); Ev-
ans. 1938 (control); Radford. 1939; Herms, 1939; Willmann, 1939;
Riley, 1940;’Weber. 1940; Pang. 1941 (m); Spencer, 1941; Newton,
1942; Baker, l943;Hungerford. 1943; Smart etal., 1943; Anderson,
1944; De Meillbn.and Lavoipierre. 1944; Pinnegan, 1945; Williams
and Brown, 1945: 482 (v); Bertram. 1-946: 209 (b); Lavoipierre, 1946;
Lowell, 1946: ’278 (m); Olson and Dahms, 1946: 56 (b); Randolph
and Eads, 194S; Taylor and Murray, 1946; Bertram, Unsworth, and
Gordon. 1946: 228 (b, v); Williams, 1946: 256 (b); Liu, 1947: 58 (m);
Pratt and Frits, 1947;. Scott et al.. 1947: 138 (b); Philip and Hughes.
1948: 6@7 (m); Scott.’1948:,4.81 <v);’Williams. 1948: 24 (v); Bertram,
1949:’ 313 (b, v)’;’ Haghes. 1949: 349 (a); Pirila and Kilpio. 1949’;
Skaliy and Hayes. ’1949: 759 (b); Baker. 1950; Bertram. 1950: 55 (b);
Hughes. 1950: 285 (v); Qvapzai-.;l950: 178 (b); Worth, 1950; Zumpt,
1950; Worth ’and Rick’ard.. 1951; 295 (m); Bishopp ’and Philip, 1952;
Schwab etal., 1952:, 982 (ro); Fujisaki, 19.53.
.

Bdellonyssus bacoti. Fohseca.,1941:. 262 (t>; Fox, 1946; Ponseca, 1948:

257 (review); Browning, 1.950: 2 (review); Gorirossi, 1950: 301 (a);
CanttinandRogoff.,1952;. H<?pls, .1951: 7.68 (m); Scott and Biynn, 1951:
.
519(b); Hughes, .1952: 54 (m); :(rons etal.. 1952: 2 (m); Lamb,. 1952;"
MacDonald and’ Scott, 1852: .283 (b); Morlan. 19-52: 89(c); Morlan
and’Utterback, 1952 (ecology); Scott, 1952; Willmann, 1952; Freer,
1953: 13 (b); Strandtmann and Eben. 1953: 148 (m); Woke. 1953:523
(m); Pratt and Good, 1954; Smith. 1955; Domrow, 1955; Bregetova,
19.53; Willmann,’1955. .

Macronyssus bacoti, Buitendijk, 1845.

Orm.thonyssus fcacoti, Bregetova, 1956: 165 (b, illus.); Baker’
^t’^1.,
’
1956: 22; Keegan, 1956; Strandtmann. 1956: 137 (t);
’
.

Haemogamas us sanguineus Ewing and Stover, 1915 (d); Ewirig, 1950;

Strandtmann. 1956.
84

Liponyssua tenuiscutatus Ewing. 1922: 22 (d); Strandtaiann. 1956."’

Liponyssus nagayoi Yamada, 1932 (d); Kodama and Kono, 1933:’ S9 (in);
Iviorishita and Ohmori, 1933; Ohmori, 1935: 3250 <b), in Japanese;
Asanuma, 1948 (t), in Japanese; Zumpt, 1950; Janieson e^ al.. 1952.
Bdelloriyssus nagayoi, Fonseca,"1948: 259.
localities: Asia - China; Formosa; India; Japan; Korea; U.S.S.R.
-
Africa Abyssinia; Belgian Congo; Egypt; Ethiopia; Liberia; Union of
South Africa; WestAfrica. Australia - New South Wales. Europe’-Eng-
-
land; Germany; Norway. Islands of the Atlantic Cuba; Puerto Rico;
West Indies, Islands of the Pacific - New Zealand. North America -
Canada; United States. South America - Argentina; Brazil.
-
Hosts; Mammals Acbmys cahirinus, Apodemufe. syXvaticus.-’A’FVi-
canthis niloticus. Cavia aperea, Citellus mexicanus, cotton mouse, cot-
ton rat, domestic animals, domestic cat, domestic rat, Florida skunk,
Gerbillus sp., golden mouse, gray squirrel, hamsters, house mouse.
human beings. Mus musculus, Mus musculus moloss’inus, Mus riorveg-
icus, Mus rattus, old-field mouse. Ony chbmy s ’leucogaster, op os sum,
Perognathus hispidus, raccoon, rats. Muridae, Rattus’<t6rve’gi.cus, Rat-
tus norvegicus otomi, Rattus rattus, ’Rattus rattus’ alexandrih’us. Rattus
rattug erythrondtus, rice rat. roof rat, sewer’rat, shrews (Soricidae),
Sigaiodon hispidus, spotted skunk, Swiss white mice, tropical rat, white
rats, wild rat, wood rat. Birds’-poultry, yellow-bellied sapsucker.
Remarks: Haemogamasus sanguineus Ewing and Stover was recog-
nized as bacoti by Ewing (1950). Tlie type slide has a line drawn through
the original name and "Liponyssus bacoti" written under it in Ewing’s
hand. Liponyssus tenuiscutatus Ewing is’’based on a’unique specimen
that is in rather poor condition but the chaetotaxy of the. dorsal plate in-
dicates it is bacoti. Liponyssus nagayoi Yamada was compared with
bacoti in great detail by Asanuma (1948), who concluded that they are
synonymous. Jam’eson et al.’came to the same conclusion independently
:"
’ ’
’
’
inl952.
The bite of Ornithbnyssus bacoti produces a’ dermatitis of man of
varying degrees of severity, depending, at least in part; ’oh the sensi-
tivity of the host. Pang (1941) and Liu (1947) claim to nave demonstra-
ted a natural infection of ’murine’ typhus in this species’.-’Dove and"Shel-’
mire(1931; 1932) used bacoti to produce laboratory infections of Murine
typhus in guinea pigs. Hopla (1951) was able to infect the mite’.;w.itti tu-
laremia and to transmit it to laboratory animals. Williams and Brown
(1945) discovered that Litomosoides carinii, the filarial worm of cotton
rats, is transmitted by bacoti and thus opened up a new line of’research.
Philip and Hughes (1948) showed it to be a successful experimental vec-
tor of rickettsialpox. Morishita and Ohmori (1933) were’abte to infect
the mite with Spirochaeta auttoni but could not demonstrate’transmission.
Yamada (1932) expressed the beli-ef that the mite can carry Pasteuryily:
pestis. Schwab ertal. (1952) were unable-to effect transmission
Dallas M. B. strain of Coxsakie virus from mouse to mouse arid con-
. o^tha’
cluded that bacoti can play but a small role, if any, in the transmission
of this strain of the virus.
 85

Ornithonyssus bacoti was first collected in 1913 and 1914 fromEgypt

as a parasite of domestic rodents and is therefore commonly known as
the tropical rat mite. The first record from the New World was a decade
or so later and was also from domestic rats. Swing (1932) therefore
assumed that Rattus was the original host, that Egypt was the country
of its origin, and that it was only recently introduced into North Ameri-
ca. Its association with the filarial parasite of the cotton rat, Sigmodon
hispidus, however provides evidence that bacoti is of New World origin.
Sigmodon is a New World rodent that has a high incidence of infection
with the filarial worm, Litomosoides carinii. 0. bacoti is an efficient
and apparently singular vector of this nematode, indicating a long and
close association with Sigmodon. It is conceivable then that bacoti be-
came secondarily adapted to Battus and that this host was a means of
emigration from rather than immigration to the New World.

Ornithonyssus aethiopicus (Hirst)

Liponyssus aethiopicus Hirst, 1921: 783 (d).

Bdellonyssus aethiopicus, Fonseca, 1948,
Chelanyssus aethiopicus, Zumpt and Till, 1953: 7. ,

Onnthonyssus aethiopicus, Keegan, 1956: 208 (9 illus.).

Locality: Africa - Nyasaland; Sudan.
Hosts: Mammals - bats, Chaerophon pumilis, elephant shrew.
Remarks: It is quite possible; that this mite, in the sense of Zumpt
and Till, and of Keegan, is actually an Ichoronyssus. Hirst took the
mite from an "elephant shrew"; all subsequent records have been from
bats. Keegan (ibid.) states that Chelanyssus forsythi Zumpt, also taken
from a bat, might be a synonym; we consider Chelanyssus to be a syno-
nym of Ichoronyssus, not of Ornithonyssus. We are retaining it in Qr-
nithonyssus because the original description indicates such, and be-
cause the host was not a chiropteran. If Zumpt and Till, and Keegan,
have correctly identified the mite, it is one of the very rare cases of an
Ornithonyssus parasitizing a bat.

Ornithonyssus banksi. New name

Haemogamasus americanus Banks, 1906: 137; 1907: 609.
Liponyssus americanus, Vitzthum.. 1930 (Not Liponyssus americanus
Banks,. 1906: 136); Keegan,’ 1951: 205.
Locality: North America - United States, Arizona.
-
Host:. Mammal Nest of Peromyscus eremicus.
Remarks: Liponyssus americanus Banks has page priority over
Haemogamasus americanus Banks, therefore Vitzthum’s combination
is’a junior homonym.

Ornithonyssus braziliensis (Fonseca). New combination

Type material in the Institute Butantan, Sao Paulo, Brazil.
Liponyssus braziliensis Fonseca, 1939: 147 (d).
86 .
, : ’

Bdellonyssus brazillensis, Fonseca, 1’948: 249.

Locality: South America-’Brazil,’Sao Paulo.
-
Hosts: Mammals .Cayia apera; Didelphis aunta, Epimys norveg-
icus, Euryzygomatomys sp.jnosus catellus, Galictis vj.ttata, Grison yit-
tatus, Homo sapiens, Marmosamarsupialis, Nectomys squamipes, Oxy-
mycterus judex, Sylvilagus minensis.
Bemarks: This mite is remarkably; similar to bacoti., both in habits
.and structure.

Ornithonyssus bursa (Berlese)

(Fig. 36)
Leiognathus bursa Berlese, 1888: 208 (d).
Liponyssus bursa. Hirst, 1916: 243 (c); Hirst, 1930: 32; Wood. 1920 (b);
Ewing, 1922; Hirst, 1922: 89 (c); Miller, 1925; Gibson, 1930; Rob-
erts, 1930: 595(m); Bedford, 1932; Arnold and Arnold. 1943: 41 (m);
Herms. 1939; De Meillon and Lavoipierre, 1944; Zimmerman, 1944;
Taylor and Murray, 1946; Fonseca, L948: 258 (v, m, t); Mathewson,
1950; Zumpt, 1950; Chamberlain and Sikes, 1950: 461 (b); Seddon,
1951.
Bdellonyssus bursa. Chamberlain and Sikes, 1955: 106 (m); tiomrow and
Smith, 1956.
’
Ornithonyssus bursa, Sambpn. 1928: 107 (t); Baker et al., 1956: 30;
Keegan, 1956. ’- .
.
Leiognathus morsitans Hirst, 1915: 55 (d); 1916: 244 (t).
Leiognathus constrictus Ewing, 192.2: 16 (d), new synonymy.
Ichoronyssus constrictus. Fonseca, .1948: 299.
Localities: Africa - Egypt; Nigeria; Nyasaiand; Union of South Af-
rica, Zululand. Asia - China; India. Indonesia - Mauritius. Australia -
New South Wales, Queensland.’ South Australia. Central America - Canal
Zone.’ Islands of the Indian Ocean-Comoro Is.; Zanzibar. Islands of
the Pacific - Hawaii; New Guinea. North America - Canada, Ontario,
Ottawa; United States. SouthAmerica-Argentina; Columbia. West In-
dies - Bahamas Islands. .
.
.

Hosts: Mammals. - Bandicoot, Homo sapiens. Birds canaries.

-
chickens, common sparrow, ducks, English sparrow, English starling,
kingbU-d, meadowlark. Passer, domesticus, pigeons, red-eyed vireo,
setting hen, turkeys, wild birds, wood thrush.
Remarks: This mite is commonly known as the tropical fowl mite.
It is outdone, only by Dermanyssus g_allmae as a serious pest of domes-
tic fowl and wild birds. It is almost never found on wild mammals al-
though there are many records of it .biting humans. It has never been
implicated as a disease transmitter, but neither have its. potentialities
been completely investigated. Chamberlain and Sikes (1955) concluded,
after exhaustive tests, that the mite is unimportant as a reservoir or
transmitter of the equine encephalitides. 0. bursa is almost entirely
restricted to warm and tropical regions, so there is some reason to
doubt that records from Canada are actually referable to this species.
 87

Ornithonyssus dendropicus (Radford). New combination

Liponyssus dendropicus Radford, 1942: 306 (d).

Bdellonyssus dendropicus, Ponseca. 1948: 289.
Locality:. Africa - Uganda.
Host: Bird - Dendropicus fuscesens lapidus.
Ornithonyssus dogieli (Bregetova)

Type material in the Zoological Institute of the Academy of Sciences,

U.S.S.R.
Bdellonyssus dogieli Bregetova, 1953: 311 (d, all stages illus.).
Ornithonyssus dogieli, Bregetova, 1956: 165 (illus.).
Locality: Asia - U.S. S.K., Tadzhikistan.
Host: Mammal - Dyromys nitedula.
Ornithonyssus eruditus (Ponseca). New combination

Liponissus eruditus Fonseca, 1935: 82 (d).

Bdellonyssus eruditus, Ponseca. 1948: 289.
-
Locality: South America Brazil. Sao Paulo.
Hosts: Probably bats, or swallows.

Ornithonyssus galagus (Zumpt and Till). New combination

Type material in the South African Institute for Medical Research.

Liponyssus galagus Zumpt and Till, 1953: 10 (d),
Locality; Africa - Uganda. Karamoia.
Host: Mammal - in nests of Galago senegalensis.
Ornithonyssus hirsti (Fonseca). New combination

Liponyssus hirsti Fonseca, 1935: 87 <d).

Bdellonyssus hirsti, Ponseca, 1948: 289.
-
Locality: South America Argentina.
Host: Mammal - Caviella australis.

Ornithonyssus iheringi (Fonseca). New combination

Liponissus iheringi Fonseca. 1935: 84 (d).

Bdellonyssus iheringi, Fonseca, 1948: 259. 282.
Locality: South America - Brazil, Sao Paulo.
-
Hosts: Mammals Bradypus tridactylus, man. Bird - Ponacobius
atricapillus.

Ornithonyssus lutzi (Fonseca). New combination

Type material in the Institute Butantan. Sao Paulo. Brazil.

88

Liponissus lutzi Fonseca, 1941: 104 (d).

Bdellonyssus lutzi, Fonseca, 1948: 290.
-
Locality: South America Brazil, Sao Paulo.
-
Host: Mammal "wild rat. "

Ornithonyssus meprai (Manso and Pletneff). New combination

Liponyssus meprai Manso and Pletneff, 1951: 13 (d).

Locality: South America - Argentina.
Host: Mammal - Rattus rattus.
Remarks: This mite could very well be Ornithonyssus bacoti..

Ornithonyssus monteiroi (Fonseca). New combination

Type material in the Institute Butantan. Sao Paulo, Brazil.

Liponissus monteiroi Fonseca, 1941: 107 (d).
Bdellonyssus monteiroi, Fonseca, 1948: 290.
Locality: South America - Brazil, Sao Paulo.
Host: Mammal - Zygodontomys lasiurus.

Ornithonyssus ondatrae (Willmann). New combination

Bdellonyssus ondatrae Willmann, 1952: 410 (d).

Locality: Europe - Germany.
Host: Mammal - Ondatra zibethica.

Ornithonyssus rose-innesi (Zumpt and Till). New combination

Liponyssus rose-innesi Zumpt and Till, 1953: 8 (d).

Locality: Africa - Union of South Africa, Cape Province.
Host: Mammal - Hhabdomys pumilio.

Ornithonyssus rhinolophi (Zumpt and P.atterson). New combination

Type material in the South African Institute for Medical Research.

Liponyssus rhinolophi Zumpt and Patterson, 1951: 87 (d).
Locality: Africa ~ Union of South Africa, Transvaal, Sterkfontein
caves.
Host: Mammal - Rhinolophus geoffroyi.
Remarks: It is doubtful that this species should be retained in Or-
nithonyssus but until such time as tlie type may be restudied, it seems
best to leave it here.

Ornithonys su s tinae (Lombardini). New combination

Liponyssus tinae Lombardini, 1953: 182 (d).

Locality: Europe - Italy (?), Cave of Pignone.
Host: Rhinolophus ferrumequinum.
 89

Remarks: I do not believe this is atrueOrnithonyssusbutarestudy

of the type will be necessary to place it correctly. It may be congeneric
with rhinolophi Zumpt and Patterson. Dr. Lombardini says it is very
similar to Liponyssus uncinatus, a species with which we are not famil-
iar.

Ornithonyssus vitzthumi (Ponseca). New combination

Type material in the Institute Butantan. Sao Paulo, Brazil.

Liponyssus vitzthumi Fonseca, 1941: 110 (d).
Bdellonyssus vitzthumi, Ponseca, 1948: 291.
Locality: Soutli America - Brazil, Butantan.
Host: Mammal - "wild rat."
Remarks: This species is very similar to bacoti.

Genus Ichoronyssus Kolenati, 1858

(Figs. 37, 38)
Type: Ichoronyssus scutatus Kolenati. (By designation of
Ewing, 1922: 13.)
Lepronyssus Kolenati, 1858: 4.
Type: Lepronyssus leprosus Kolenati. (By designation of
Fonseca, 1948: 269.)
Chiroptonyssus Augustson, 1945: 46.
Type: (Chiroptonyssus texensis Augustson) = Liponyssus
robustipea Ewing.
Chelanyssus Zumpt and Till. 1953: 5.
Type: Chiroptonyssus forsythi Zumpt,
Diagnosis: Dorsal plate entire and covering most of the dorsum; fe-
male epigynial plate pointed or rounded posteriorly, bearing one pair of
setae and having scale-like lines; male ventral plate divided just poster-
ior to coxa IV; at least coxae II and HI with ventral, semicircular ele-
vations; female sternal plate generally with a thickened medial or pos-
terior transverse band; opisthosoma fairly densely covered with stiff
setae. Parasites of Chiroptera.
Remarks: Ichoronyssus scutatus Kolenati is not very well described
or illustrated but it was chosen as the type for the genus by Ewing.
Fonseca (1948), although recognizing Ewing’s designation, made it clear
that the species he included as Ichoronyssus were similar to Ichoronys-
sus hasei Vitzthum, 1931.
Lepronyssus is considered a synonym of Ichoronyssus because there
are no apparent differences in the published descriptions and illustra-
tions. Fonseca (1948) found the only apparent difference to be the scaly
appearance of the epigynial plate of the female, but this plate is scaly in
all Ichoronyssus species.
Chiroptonyssus Augustson is placed in synonymy because its gener-
otype, texensis, is a synonym of Liponyssus robustipes, a species that
we consider to be Ichoronyssus. It is atypical only in having two rather
90

than three pairs of setae on the sternal plate. This character Is variable.
Zumpt and Till (1953) erected Chelanyssus because the chela of the
generotype, Chiroptonyssus forsythi, has a "grappling hook" structure,
unlike that of any other mite. To us, however, the illustrations they
give indicate that the moveable arm of the chela is bent so far back as
to give the appearance of such a hook. It is placed in synonymy because
in all other respects it seems to have the general characteristics of Xcho-
ronyssus. We cannot be sure of this, however, until the type can be
restudied and until males are found; forsythi is known only from the fe-
male and protonymph. As in robustipes. it has only two pairs of setae
on the sternal plate’.
Dr. Fonseca also lists Inchoronyssus Kolenati, 1858 and Chirop-
terolaelaps Vitzthum, 1927 as synonyms of Ichoronyssus. The former
is a lapsus calami and the latter is a nomen nudum.

Ichoronyssus scutatus Kolenati

Ichoronyssus scutatus Kolenati, 1858: 6 (d); Kolenati, 1859;Ewing, 1922;

Fonseca. 1948: 266 (t).
-
Locality: Europe Serbia. Steiermark,
Host: Mammal - Miniopterus schreibersii, Rhinolophus clivosus,
Rhinolophus ferrumequinum.

Ichoronyssus australicus (Womersley). New combination

Chiroptonyssus australicus Womersley, 1956: 597 (d. n, cf and $ illus.).

Locality: Australia-South Australia, WarblaCave, Nullabor Plains.
Host: Mammal - Bats.
Remarks; The male and nymph illustrated by Womersley have very
long, flexible setae, completely unlike that of the female. Also, the
male has an undivided holoventral plate unlike any other ichoronyssid.
We wonder if the male and nymph are actually conspecific with the female.

Ichoronyssus cornutus (Keegan). New combination

Type material in the U. S. National Museum.

Ornithonyssus cornutus Keegan, 1956: 211 (d, $ illus.).
Locality: Africa - Egypt.
Host: Mammal - Tadarida aegyptiaca.
Remarks: The broadened posterior setae so characteristic of icho-
ronyssid mites found on Tadarjda species suggest very strongly that this
is not an Qrnithonys sus.

Ichoronyssus crosbyi. (Ewing and Stover)

Type material in the U. S. National Museum.

Liponyssus crosbyi Ewing and Stover. 1915: 109 (d).
Leiognathus crosbyi, Ewing. 1922.
 91

Ichoronyssus crosbyi, Fonseca, 1948: 299; Strandtmann, 1956.

Locality: North America - United States. Missouri.
Host:’’’ Mammal’-,. Vesper subulatus i
Xchoronyssus diversipilus (Vitzthum)

Liponyssus diversipilus Vitzthum, 1918: 23 (d).. ;

Ichoronyssus-diversipilus, Fonseca, 1948: 299; Furmah, 1950.
Locality: EuropeGermany, ;;: ,

.
.
Host: Mammal - "a small bat, probably Vesp.erugo.pipistrellus."

Ichoronysgus flavus. (Kolenati’) .

:’

Dermanys.sus flavus Kolenati, 1857: 19... ,

Leprdhyssus flavus. Kolenati, 1859: 184; Fonseca, 18.48: 306 (t),
...
Liponyssus flavus. Hirst. 1921: 791 (d); Vitzthum, .1931: 10; Zumpt,
1950: 198.
Ichoronyssus flavus, Bregetoya, 1953; 330; Bregetova. 1956: 162 (c).
Lepronyssus lobatus Kolenati, ’18 57; Kolenati, 1859: 182.
Liponyssus lobatus, Oudemans. 1904: 18,(d);::Vit2thum, 1918: 17.
Allonyssus lobatus, Buitendijit, 1945... ,
Locality: Europe; Asia -U.S.. S.R.
.
.
Hosts: Mammals - Barbastella barbastellus, Myotis daubentonii,
Myotis myotis, Myotis murinus, Nyctalus noctula, Panugo noctula, Ves-
pertilio murinus, Vespertilio nathusii, Vespertilio nxissonii. Vespertilio
pipistrellus, Vesperugo noctula. . ,
.

Remarks: Zumpt (1950). remarks, that this mite may sometimes an-
noy man, and in Italy has been known t.o cause dermatitis.

Ichoronyssus forsythi (Zumpt). New combination

Type material in the South African Institute for Medical Research.

Chiroptonyssus forsythi Zumpt, 1950: 169 (d).
Liponyssus forsythi, Zumpt and Patterson, 1951.
Chelanyssus forsythi, Zumpt and Till, 1953: 5.
Locality: Africa - Madagascar.
Host; Mammal - Chaerephon leucostigma.

Ichoronyssus granulosus (Kolenati). New combination

Type material in the British Museum of Natural History.

Permanyssusgranulosus Kolenati. 1857: 20 (vide Hirst, 1921).
Lepronyssus granulosus, Kolenati, 1859: 181; Fonseca. 1948: 260.
Liponyssus granulosus. Hirst, 1921: 794 (d); Radford, 1939; Hoffman,
1944: 261. ,
,

Dermanyssus glutinosus Kolenati.. 1857: 20 (vide Hirst, 1921).

Localities: Europe - Croatia. "C. Stayal." North America Mex-
-
92

Hosts: Mammals - Miniopter.us schreibersii, Natalus mexicanus,

Vespertilio myofis, "whiskered hat. "

lehoronyssus haeiaatophagus (Fonseca). New combination

Liponissus haematophagua Ponseca, 1935: 25 (d).

Bdellonyssus haematophagus, Fonseca, 1948: 289.
-
Locality: South America Brazil. Rio de Janeiro.
-
Hosts: Mammals Molossus abrassus, Molossus .rufus, Nyctino-
^
mus macrotis. ... .
.

,
.
Remarks; Only the female of this species is known. It is very sim-
-
ilar to ^obu^st^es^ (Ewing). ..The drawing Fonseca gives indicates that

.
there are three pairs Of setae on the sternal plate; this seems to be the
only character differentiating it from robustipes, which has only two
pairs of setae on the plate. , . .

.
.
Morlan and Strandtmanh (.1949). Morlan .(1952) and Yunker (1958)
incorrectly referred to this species. What they actually had was Icho-
rpnyssus robustipes.. . .
.. - :; :. . .
.
Icboronyssus hasei Vitzthum ..;’ ’.

’
Ifchoronyssus hasei Vitzthum, 1932: 23 (Sj; Fonseca, 1948": .299.
Locality: South America - Venezuela,. Puerto la Cruzt
-
Hpst: " Mammal Myotis nigricans. ,
.

.
.
:-
’ -lehoronyssus hoogstraali (Keegan). New combination
’Type material in the U. S. National. Museum. .. ..

Qrnithonyssus hoogstraali Keegan, 1956: 210 (d, $,illus.).

Locality: Africa - Egypt. . . ; ,. ..
.

Host: Mammal - Tadarida teniotis, a bat; .

.
.
.
Remarks: The hysterosomal. body setae are broadened.as in robus-
tipes (Ewing). This. plus’the fact. that the mite is from ;a bat, would
indicate that it is an lehoronyssus. ~-;’.
.
lehoronyssus koc.bi Fonseca .

^
,

lehoronyssus kochi Fonseca, 1948: 278 (d).

Locality: South America - Brazil, Sao Paulo. ., ...
-
Host: Mammal Artibeus jamaicansis ’lituratus.

IchorQnys sus kplenati Fonseca .. ;

Ichoronyssus kolenati Fonseca, 1948: 280 (d). ..; .

.
.
Locality: South America - Brazil, Sao Paulo. .;

-
Host: Mammal Myotis ruber. ., ., , ,
.
.
.
Remarks: Fonseca mentions that this is not to be .confused with
Liponyssus kolenati Oudemans, 1902, a species known only from the
 93

male. which cannot be an Ichoronyssus because the ventral plate is un-

divided. Its proper placement is not known.

Ichoronyssus lepidopeltis (Kolenati)

Macronyssus lepidopeltis Kolenati, 1858; 1859: 179 (d).

Liponyssus lepidopeltis, Tragardh. 1912: 581 (d).
Allonyssus lepidopeltis, Buitendijk, 1945.
Ichoronyssus lepidopeltis, Fonseca, 1948: 302.
Locality; Africa - Egypt. Europe-France; Germany; Netherlands;
U.S.S.R.
Host: Mammal - Rhinopoma roicrophyllum.

Ichoronyssus leprosus (Kolenati). New combination

Lepronyssus leprosus Kolenati, 1858; 1859: 180 (d); Fonseca, 1948: 268.
308.
-
Locality: Africa Egypt.
Host: Mammal - Rhinolophus clivosus.

Ichoronyssus longisetosus Furman

(Fig. 37)
Type material in the U. S. National Museum.
Ichoronyssus longisetosus Furman, 1950: 479 (d).
-
Locality: North America United States, California.
Hosts: Mammals - Corynorhinus rafinesquii intermedium, Coryno-
rhinus rafinesquii pallescens.

Ichoronyssus mohrae Vitzthum

Ichoronyssus mohrae Vitzthum., 1932: 32 <d); Fonseca. 1948: 302.

Locality: Europe - Germany, Holstein.
Host: Mammal - Vespertilio nattereri.

Ichoronyssus nyctinomi (Zumpt and Patterson). New combination

Type material in the South African Institute for Medical Research.

Liponyssus nyctinomi Zumpt and Patterson, 1951: 89 (d). Not Liponys -
sus nyctinomius Radford, 1938.
-
Locality: Africa Cape Province.
-
Host: Mammal Nyctinomys boeagei.

Ichoronyasus quadridentatus Strandtmann and Hunt

(Fig. 38)
Type material in the U. S. National Museum.
Ichorpnyssus quadridentatus Strandtmann and Hunt. 1951: 462 (d); Mor-
lan, 1952: 89.
Locality: North America - United States. Georgia.
94

Host: Mammal - Eptesicus fuscus.

Ichoronyssus robustipes (Ewing)

Type material in the U, S. National Museum.

Liponyssus robustipes Ewing, 1925: 20 (d).
Chiroptonyssus robustipes, Ponseca. 1948: 291 (t).
Ichoronyssus robustipes, Strandtmann and Hunt, 1951: 465 (t); Morlan,
1952: 89; Strandtmann. 1956: 137 (t).
Liponyssus chilensis Ewing, 1925: 21 (d); Strandtmann and Morlan, 1953;
Strandtmann, 1956; 137.
Hirstionyssus chilensis, Fonseca, 1948: 298.
Liponyssus nyctinomius Radford, 1938: 431 (d).
Liponyssus venezolanus, Hoffmann, 1944: 43 (d).
(Misidentification, not venezolanus Vitzthum.)
Chiroptonyssus texensis Augusteon, 1945: 46 (d).
Bdellonyssus hematophagus. Morlan and Strandtmann, 1949; Morlan,
1952: 89. (Misidentification. not hematophagus Fonseca.)
Qmithonyssus haematophagus, Yunker, 1958.
(Misidentification, not hematophagus Fonseca.)
Localities: North America-Mexico; United States. SouthAmerica -
-
Chile, West Indies Jamaica.
-
Hosts: Mammals Myotis lucitugus, Natalus mexicanus. Nycti-
nomius brasiliensis, Tadarida brasiliensis, Tadarida cynocephala, Ta-
darida mexicana.
Remarks: The type specimen of Liponyssus chilensis (in the U. S,
National Museum) was seen by us in 1954. It is in very poor condition
but it is possible to see that the sternal plate has only four setae and
has a heavy transverse band. Also, the opisthosomal setae are flattened
apically. This, plus the fact it is from the same host as robustipes. in-
dicates its synonymy with the latter species. Fonseca (1948), confer-
ring with E. W. Baker, place nyctinomius Radford in synonymy, Chi-
roptonyssus texensis was compared with robustipes by both Ewing and
Baker and on their advice placed in synonymy by Fonseca. Bdellonys-
sus hematophagus Fonseca. as used above, was a misidentifxcation. as
was Liponyssus venezolanus Vitzthum as used by Miss Hoffman. It
should be noted that Ichoronyssus venezolanus (Vitzthum) and L hema-
tophagus (Fonseca) are species very similar to I. robustipes.
According to Hoffman (1944). robustipes attacks man and causes
some discomfort.

Ichoronyssus spinosus Oudemans

Ichoronyssus sptnosus Oudemans, 1902; Turk, 1945: 141 (t); Fonseca,

1948: 302 (t).
Liponyssus spinosus, Oudemans, 1902; Tragardh, 1912: 577.
Psilognathus spinosus, Oudemans, 1902,
Macronyssus spinosus, Oudemans, 1902; Buitendijk, 1945.
 95

Locality: Europe - France; Germany.

Host: Mammal - Vespertilio murinus.
Remarks: We have not seen the original descriptions nor any sub-
sequent description or illustration of this species. The synonymy is
taken from Ponseca (1948), and the host record is that given by Turk
(1945),

Ichoronyssus venezolanus (Vitzthum). New combination

Liponyssus venezolanus Vitzthum, 1932: 9 (d).

Bdellonyssus venezolanus. Fonseca, 1948: 259.
Ichoronyssus hermanni Fonseca, 1948: 277 <t).
-
Locality: South America Venezuela.
Host: Mammal - Molossus nagutus.
Remarks: Fonseca (1948) considered the protonymph and female of
Vitzthum’s species to represent a different form than the male and pro-
posed the name Ichoronyssus hermanni tor the male. The female and
nymph he considered to be a Bdellonyssus. We see no reason for this
as all stages of the mite as illustrated and described by Vitzthum fit the
generic characterization of Ichoronyssus.
There is a remarkable similarity between this mite and robustipes
(Ewing). In fact. Hoffman (1944), in her excellent paper on ectopara-
sites of Mexican bats, confused this species with Ichoronyssus robusti-
pes (Ewing).

Genus Spinolaelaps Radford, 1940

(Fig. 39)
Type: Spinolaelaps jacksoni Radford
Diagnosis: Female epigynial plate somewhat expanded posteriorly
and bearing nine accessory setae; chelae edentate but with a small seta
onthelmmoveable arm; male holoventral plate divided just posterior to
coxae IV. Parasitic on Chiroptera.
Remarks: Except for the enlarged epigynial plate, this genus is
identical with Ichoronyssus and was listed as a synonym by Strandtmann
and Hunt (1951). However, a restudy of Radford’s material and new
material received from Dr. E. W. Jameson, Jr.. has led us to believe
that the form of the epigynial plate is significant and that it deserves
generic rank.

Spinolaelaps jacks oni Radford

Spinolaelaps jacksoni Radford, 1940: 96 (d).

Ichoronyssus jacksoni, Strandtmann and Hunt. 1951: 465 (t).
Locality: Africa - Kenya.
-
Host: Mammal Rhinolophus lobatus.
96

Genus Sauronyssus Sambon, 1928

(Fig. 40)
Type: Liponys sus saurarum Oudemans.
Neoliponyssus Ewing, 1929.
Type: Liponyssus gordonensi.B Hirst
Oudemansiella Fonseca, 1948: 269.
Type: Liponyssus saurarum Oudemans,
Diagnosis: Coxae unarmed; female sternal plate trapezoidal, not
much broader than long and bearing only four setae; dorsal plate undi-
vided and extending beyond the middle of the dorsum, or occasionally
divided into a large podosomal and small pygidial shield; female epi-
gynial plate pointed posteriorly and bearing one pair of setae; male anal
plate discrete; ectoparasites of reptiles.
Remarks: Oudemansiella was proposed by Ponseca to be used in
the event that it should be shown that saurarum is not congeneric with
gordonensis. Fonseca was not aware of Sambon’s name, Sauroayssus.
We have seen very little of the original literature on this genus and no
specimens at all.

Sauronyssus saurarum (Oudemans)

Liponyssus saurarum Oudemans, 1901; 1902; 1903.

Neoliponyssus saurarum, Fonseca. 1948: 269 and 312; Radtord, 1950:
373; Zemska. 1951.
Sauronyssus saurarum, Sambon, 1928: 106; Bregetova, 1956: 159 (blol.,
illus.).
Locality: ? Europe.
Host: Reptile -Lacerta agilis agilis, Lacerta muralis, Lacerta
viridis, Lacerta vivxpara.
Remarks: Zemska (1951) mentions that this mite is the intermediate
host of Karyolysus lacertarum, a haemogregarine parasite of lizards.
The references to Oudemans are cited from Fonseca.

Sauronyssus arnhemlandensis (Womersley). New combination

Type material in the South Australian Museum.

Neoliponyssus arnhemlandensis Womersley, 1956: 594(d, cf and $ illus.).
-
Locality: Australia Arnhem Land.
-
Host; Reptile Leiolopisma fusca (a skink).

Sauronyssus gordonensis (Hirst). New combination

(Fig. 40)
Liponyssus gordonensis Hirst, 1923: 973 (d).
Neoliponyssus gordonensis, Ewing. 1929; Ponseca, 1948: 269.
Locality: Africa - Sudan, Khartoum.
Host: Reptile - Mabuia quinquestriata.
 97

Sauronyssus lacertinus (Ber-lese). New combination

Leiognathus lacertinus Berlese, 1892: 70 (d).

Neoliponyssus lacertinus, Fonseca. 1948: 312; Radford, 1950: 373.
Liponyssus natricis. Berlese, 1917: 55. (Not Gervais. 1844).
Locality; Europe.
-
Host: Reptile Lacerta viridis viridis.

Saur&nyssus myrmecophagus Fonseca. New combination

Neoliponyssus myrmecophagus Fonseca, 1954; 84.

Locality: South America - Brazil, Mato Grosso.
-
Host: Mammal Myrme c ophaga tridactyla tridactyla.
Remarks: The species myrmecophagus Fonseca does not tit the
generic diagnosis in all details and it is parasitic on a mammal but it
is retained here because the male anal plate is separate.

Genus Ophionyssus Megnin. 1884

(Fig, 41)
Type: Dermanyssus natricis Gervais, 1844: 223.
Serpenticola Ewing, 1922; 6.
Type: Ichoronyssus serpentium Hirst.
Diagnosis: Female with two dorsal shields, a podosomaland a small
pygidial shield; podosomal shield not extending beyond the fourth pair
of legs. with four tiny platelets posterior to it (much like the normal
protonymphs of other species of Macronyssinae); sternal plate of female
trapezoidal and bearing only four setae; male ventral plates consisting
of a short sternal-genital-ventral complex and a separate anal plate; all
coxae unarmed; external parasites of snakes and lizards.

Ophionyssus natricis (Gervais)

(Pig. 41)
Dermanyssus natricis Gervais, 1844: 223 (d).
Ophionyssua natricis. Megnin, 1884: 107; Andre, 1937: 63(1); Piekarski.
1936: 615 (b); Fonseca, 1948: 260 (review); Camin. 1949: 583 (t);
Camin. 1953: 5<b); Zemska, 1951; Bregetova, 1956: 160<biol.,
illus.);Womersley. 1956: 599; Baker etal.. 1956: 33; Keegan. 1956;
Yunker. 1956 <c).
Liponyssus natricis. Hirst. 1921.
Ichoronyssus serpentium Hirst, 1915: 383 (d).
Serpenticola serpentium, Ewing. 1922: 6.
Ophionyasus serpentium, Fonseca, 1932; Schroeder, 1934: 1004 (con-
trol); Radford, 1939; Ferris, 1942; Radford, 1942; Camin, 1948:
345 (v).
Liponyssus arabicus Hirst, 1921: 365 (d); Hirst. 1921: 775 (t).
Liponyssus monodi Hirst, 1925: 95 (d),
Serpenticola easti Ewing, 1925: 18 (d).
98

Ophionyssus easti, Ponseca, 1948; 313.

- -
Localities: Africa Egypt. Asia South Arabia; U.S.S.R., Mos-
-
cow zoo. Australia, Europe England; Netherlands. South America -
Brazil. North America - United States.
-
Hosts: Mammals - Rat. Homo sapiens. Reptiles Acanthodactylus
scutellatus, Agama adramitana. Coluber florulentus. Coluber karelini,
Drymarchon corals coopen, Epicrates cenchris, Najahaje, Psammophis
sibilans. -Psaminophis scholtari. Python reticulatus, Sceloporus graci-
ous, Spalerosophis Clifford!, Telescopus dhara obtusus, Vipera lebetina.
Remarks: Known as the snake mite, this is probably the most ser-
ious parasite of captive snakes and lizards the world over. Yunker (1956)
verified its existence on reptilian hosts in natural situations. Camin
(1948) showed that the bacterium Pseudomonas hydrophilus, a cause of
severe and highly fatal hemorrhagic septicemia in captive reptiles, is
transmitted by this mite (see also, Zemska, 1951). An intensive report
on the biology and sensory behavior of this mite was given by Camin in
1953. -Recently. Camin (private correspondence) has found that this
mite harbors, and very likely transmits, a haemogregarine of snakes.
Baker et al. (1956) give control recommendations.

Ophionyssus variabilis Zemska

Ophionyssus variabilis Zemska, 1951: 53 (d);.Bregetova, 1956: 161 (il-

lus.).
Locality: Asia - U.S.S.R., Moscow zoo.
-
Hosts: Reptiles Echis carinatus.

Genus Neospinolaelaps Zumpt and Patterson, 1952

(Fig. 42)
Type: Neospinolaelaps miniopteri Zumpt and Patterson.
Diagnosis: Dorsal plate entire; epigynial plate with five setae; che-
lae edentate and without setae.
Remarks: Described from only two female specimens.

Neospinolaelaps miniopteri Zumpt and Patterson

Type material in the Natal Museum, Pietermaritzburg, South Africa.

Neospinolaelaps miniopteri Zumpt and Patterson, 1952: 159 <d).
-
Locality: Africa Union of South Africa, Natal.
Host: Mammal - Miniopterus natalensis.
Remarks: The original authors placed this genus in the family Lae-
laptidae but it is here placed in the Macronyssinae because the chelae
are said to lack teeth and setae and because it is parasitic on bats. A
restudy of the types will be necessary to establish its true affinities.

Genus Australolaelaps Womersley, 1956

(Fig. 43)
Type: Australolaelaps mitchelli Womersley.
 99

Diagnosis: Coxae toothed; female epigyniai plate broadly rounded

and bearing four pairs of setae; no metapodal plates; no peritreme; che-
lae edentate.
Remarks: Womersley placed this. genus in the family Laelaptidae
but the type of chela and the general fades of the mite indicate it is a
dermanyssid, rather closely related to Hirstionyssus.

AustralolaelapS mitchel’li Womersley

Type
- (Fig. 43)
material in the South Australian Museum. .
.

AustralolaelapS mitchelli Womersley, 1956: ;562.(d, illus.).

Locality: Australia - South Australia, Greenly Island.
Host: Mammal - Thylo’gale eugenii, a wallaby. .. .

.’ Genus Hirstionyssus Fonseca, 1948,

’
. ..
(Fig. 44) ",. .

Type: Dermanyssus arcuatus Koch. .

.
.
.
Diagnosis: Dorsal plate entire and covering most of the dorsum;
some of .’trie. coxae with ventral or posterior spurs or both; female epi-
gyniai plate rounded posteriorly; male with a holoventral plate that is
slightly-expanded behind coxae IV; female without palpal spurs; female
chelae generally quite long, frequently as muchas a.thirdofthe^total
length of the chelicerae. ’

Remarks: It. should be clearly ’understood that the generolype is.

arcuatus Koch as it is. described and.figured by Oudemans (1913).. C, L.
Koch’s drawings and descriptions are not in the least diagnostic and it
is quite possible that Koch’s species assigned to this genus do not be-
long here at all. The genus, then, actually stems from Oudemans’ fine
paper, "Acarologisches aus Maulwurfnestern".(1913), and the synonymy
prior to that date is his. .
’ .

According to Koch, arcuatus was found on a bat but as far as we

know, this species, as it is now understood, never occurs on the Chiro-
ptera. This applies to other members of the genus as well. Hence, it
should be borne in mind when reading the host list of these mites that
any references to Chiroptera are from early records and the mites in-
volved may not have been Hirstionyssus.

Hirstionyssus arcuatus (Koch) ;

Dermanyssus arcuatus Koch, 1839: #2. .

, ..

Leiognathus arcuatus,- Gonder, 1910: 298 (m).

Liponyssus arcuatus, Oudemans. 1913: 68 (d); Radford, 1939.
Hirstionyssus arcuatus, Fonseca, 1948: 259 (review); Strandtmann and
Morlan, 1953; Willmann, 1952; Womersley, 1956; 593 (illus.); Kee-
gan, 1956.
Dermanyssus albatus Koch, 1839: #5.
Liponyssus albatus, Oudemans, 1902: 22 (d). .. .
100

Permanyssus coriaceus Koch, 1839.

Dermanyssus noctulae Koch, 1839: #5.
Dermanyssus pipistrellae Gervais, 1841.
Parasitus musculi Oudemaris, 1897: 113.
Ichoronyssus decussatus Oudemans, 1897: 136.
Hirstionyssus talpae Zemska, 1954; Bregetova, 1956: 184 (d. illus.).
New synonymy;
Localities: Africa - Egypt, Giza Province. Australia-’Queensland.
Europe - Germany; Italy;. Netherlands. North America - United States.
Hosts: Mammals - Apodemus agrarius, Apodemus flavicollis, Apo-
demus sylvaticus, Arvicola .amphlbius, Blarina brevicauda, Clethrio-
nomys glareolus, Cryptotis parva, Eptesicus serotinus, "field vole, "
Microtus arvalis, Microtus pennsylvanicus, "mole, " Mus tnusculus,
Myotis dasycneme, Myotis myotis, Neurotrichus gibbsi, Parascalops
breweri, Pterygistes noctula, Putorius erininea, Putorius putorius,
"rats, " Rattus norvegicus, Sorex cinereus, Sorex fumeus, Talpa euro -
pea, Vespertilio dasycneme, Vespertilio murinus, Vespertilio noctula,
Vespertilio serotinus, Vesperugo noctula.
Remarks: See under discussion of genus. Keegan (ibid., p. 216)
lists a mite taken in Egypt from Mus musculus. as being either arcuatus
or carnifex, stating that he could find no characters separating the two
species. He also suggests that otomys (Radford) and latiscutatus (De
Meillon and Lavoipierre) are synonyms of arcuatus.
Gonder (1910). working ’on trypanosomes of bats, lists as a likely
intermediate host and vector, the mite, Leiognathus arcuatus. No date
or authority is mentioned. We have assumed the name to refer to Hirst-
ionyssus arcuatus (C. L. Koch, 1839) and-hence listed it in the above
synonymy. However," it is’doubtful that Gender’s mite was really this
species. As mentioned in the discussion under the genus, true Hirst-
ioriyssus spp. are not found on bats. What Gonder worked with was prob-
ably some species of Ichorohyssus.
’Zemska (1954) consideredthe Hirstionyssus arcuatus (C. L. Koch),
as described by Oude’mans (1913). to be a species unlike the original
arcuatus of Koch. The name talpae was therefore proposed for the form
described by Oudemans.’We have already pointed out. in our discussion
of the genus, the probability that Oudemaris’ arcuatus. is unlike Koch’s
arcuatus.’ But because Koch’s description and illustrations are nondiag-
no’stic, and the types are lost, much less confusion will result by accept-
ing’Oudemans1 arcuatus as the lectotype ’and letting the genus Hirstio-
nyssus stem from it. That, after-all," is the basis’ on which Fonseca
(1948) erected the genus.

Hirstionyssus affinis Jameson

’

Type material in the U. S. National Museum.’

’
!
.
Neoichoronyssus (Hirstionyssus) affinis Jameson, 1950: 163’(d). .;.
Hirstionyssus affinis, Strandtmann and Morlan, 19.53: 632. .

Locality: North America United States,’ California.

 101

Hosts: Mammals - Eutamias minimus, Peromyscug boylii, Tam-

iasciurus douglasii.

Hirstionyssus blanchardi (Trouessart)

Leiognathus blanchardi Trouessart, 1904: 558 (d).

Liponyssus blanchardi, Hirst, 1921. .
. ..

.
Hirstionyssus blanchardi, Fonseca, 1948: 296; Willmann. 1952; Strandt-
mann and Morlan, 1953; Bregetova. 1956: 117(illus.).
:’ Locality: Europe - England; France; Germany; Switzerland.
Hosts: Mammals - Cynomys ludovicianus (specimens kept in cap-
.

tivity, England), Marmota marmota.

Remarks: The European marmot, Marmota marmota, is undoubtedly
the preferred host-

Hirstionyssus bregetovae Razumova

’
Hirstionyssus bregetovae Razumova, 1953; Bregetova. 1956: 176(d"andS,
Ulus.).
Locality: Asia - U. S. S. R,
-
Host: Mammal Prometheomys schaposchnikovi.

Hirstionysgus breviseta Strandtmann and Morlan

(Fig. 44)
Type material in the U. S. National Museum.
Hirstionyssus breviseta Strandtmann and Morlan, 1953; 632 (d).
-
Locality: North America United States, New Mexico.
Host: Mammal - Neotoma albigula albigula.

Hirstionyssus butantanensis (Fonseca)

Type material in the Institute Butantan. Sao Paulo. Brazil.

Ichoronyssus butantanensis Fonseca, 1932: 135 (d).
Hirstionyssus butantanensis, Ponseca. 1948; Strandtmann and Morlan,
1953.
-
Locality: South America Brazil. Sao Paulo.
Host; Mammal - Mus musculus albinus.
Remarks: "White laboratory mice were heavily infested. " (Fon-
seca. 1932).
Hirstionyssus carnifex (Koch)

Dermanyssus carnifex Koch, 1839: #1 (d).

Liponyssus carnifex, Oudemans, 1913: 75 (d).
Ichoronyssus carnifex, Ewing, 1922: 14; Smith, 1955.
Neoichoronyssus carnifex, Jameson_et_al., 1952: 11; Traub ^t al., 1954.
Hirstionyssus carnifex, Fonseca, 1948: 296; Willmann. 1952: 406;
Strandtmann and Morlan, 1953; Bregetova, 1956 ($, illus.).
102

Neoichoronyssus (Hirstionyssus) carnifex. Keegan. 1953: 40. ^.

Dermanyssus lanius Koch, 1839: #4,
Ichor onyssus decussatus Kolenati, 1858.
Liponyssus albato-affinis Oudemans, 1904: 24.
Localities: Asia - Japan; Korea. Europe - U.S. S.R.; Germany;
Netherlands, North America - United States. -
Hosts: Mammals - Apodemus flavicollis, ArvlcOla arvalis, "bats,"
Clethrionomya glareolus, "house mouse, " Mus musculus, Peromyscus
maaiculatus. Pterygistes noctula, Mus rattus, "rodents. " Sorex sp.,
Sylvia atricapilla, Talpa europaea, Vespertilio sp., Mycromys minutus’.

Hirstionyssus confuscianus (Hirst)

Liponyssus berlesei Hirst, 1921: 362 (d). Preoccupied by berlesei Can-

estrini, 1888.
Liponyssus confuscianus Hirst, 1921: 789. Proposed for berlesei Hirst,
HirstiQnyssus confuscianus. Eonseca, 1948: 297; Strandtmann and Hunt,
"’
1953; Bregetova, 1956: 174 (cT and 9, illus.).
Locality: Asia - North China; U.S. S.R.
Hosts: Mammals -Myospalax scansus, Myospalax psilurus, Myo-
spalax dybowskii.’ ..i..:.~

Hirstionyssus craticulatus Keegan

Type material in the U. S. Natl,onai,]y(useum.

Hirstionyssus craticulatus Keegan. 1956:^216.(d, $, illus.)..
Locality: ’Africa - Egypt.
’
... ..
. ..,
Hosts: Mammals’-’Gerbiilus’-’gerbmus." Jaculus jaculus.

Hirsttonyssus cretghtbhi (Hirst)

Leiognathus crej.ghtoni Hirst....19’12: .370 (d). ... .... ... .

.
Hirstionyssus ci-eightoni.’ Foiiseca, 1.948: 29 7;’...Strandtmann and Hunt,
:- 1’953. .
’

Locality: Africa -Kenya, Nairobi... ,, ...: ’

....
.
.
Host: Mammal - "Young porcupine." . -
’"’ Hirstiohyssus cncgtj. (Sulzer)
"
.
Acarug criceti Sulzer, .1.774: .33; Qudemans,, 1929: 96; Ponseca, 1948:
319.
Liponyssus criceti, Oudemans. 1932: 202 (illus.). ;. . .
... .,
Hirstionyssus ci-icgti. Zemska. 19.55; Bregetova, 1955:. 185 (rfand?. il-

. . ’-’...
’
;.. ;;- ^..’
’
lus.).
"
(
.
Locality: Europe - Germany; U.S. S. 11.
Hosts: ..Mammals - Citellus citellus. Citellus major. Gitellus pyg-
maeus. Citellus suslicus. Cricetulusbarabensis. Cricetulus.tritQnj Cri-
cetus auratus, Cricetus cricetus. Ci-icetug eyersmannt."’
""~
 103

Remarks: The reference to Sulzer is quoted from Oudemans (1929:

96) and is given only as "1774, Vers. Naturg."
Hirstionyssus cynomys (Radford)

Liponyssus cynomys Radford, 1941: 313.

Hirstionyssus cynomys, Fonseca, 1948: 29-7; Strandtmann, 1956; 138 (t).
Locality: North America - United States, Colorado.
Host: Mammal - "a prairie dog, " Cynomys sp.
Remarks: Strandtmann and Morlan (1953) placed this mite in the
synonymy of occidentalis Ewing but a study of the type of occidentalis
and paratype of cynomys showed that this was an error.

Hirsti-onyssus ellobil Bregetova

Hirstionyssus blanchardi Zemska, 1951; Dubinin and Bregetova, 1952.

N ot blanchardi Trou’es’sart, 1904.
Hirstionyssus ellobil Bregetova, 1956: 177 (cf and 9, illus.).
Locality: Asia - V. S. S.R., Mongolia.
Hosts: Mammals - Ellobius fuscocapillus. Ellobius talpinus.

Hirstionyssus eusoricis Bregetova

Hirstionyssus soricis Zemska, 1955. Not H. soricis Turk.

Hirstionyssus eusoricis Bregetova, 1956: 184 (d" and $, illus.).
Locality: Europe - Estonia; Latvia; U.S. S.R. .

Hosts: Mammals - Neomys fodiens, Sorex ararieus.

Hirstionyssus eversmanni Zemska

Hirstionyssus eversmanni Zemska, 1955; Bregetova, 1956: 181 ($, illus.),

Locality: Europe -U.S. S.R., Stalingrad.
- ’

Host: Mammal - Cricetus ever.smanni.

Hirstionyssus geomydis (Keegan)

Liponyssus geomydis K.eegan, 1946: 69.
Hirstionyssus geomydis, Fonseca. 1948: 297; Strandtmann and Morlan,
I 95 3; Strandtmann, 1956: 138.
Locality: North America -United States.
"
Host: Mammal - Geomys bursarius. -’
Remarks: This species has two clawlike setae at the apex of tarsus
I, contrary to the statement of Strandtmann and Morlarii 1953: 629.
HirstionyssuS georgicus Bregetova

Hirstionyssus georgicus Bregetova, 1956: 172(d"and $, illus.).

Localities: Europe and Asia - U.S. S.R.. Georgia, Armenia.
104

Host: Mammal - Spalax leucodon.

Hirstionyssus hillJ.Ja.iriesbn

Neoichpronyssus (Hirstionyssus) hilli Jameson, 1950: 1.65 (d); Keegan,

19.53: 40 (c). ,’. ... . .

HirstiQnyssus hilli., Strandtmann.and Morlan, 1953. . .

-
Locality;’ North America United States, California, Kansas, Ne-
vada, , .

, Hoists; Mammals - Dipodomys ordii,

Neotoma lepida, Perognathus
hispidus, Perognathus parvus, Peromyseus maniculatus.

Hirstionysaus hirsfl’Fonseca

Liponyssus aethiopicus Hirst, 1921: 783. In part (male only).

Hirstionyssus hirsti Fonseca, 1948: 297 (t); Strandtmann’ and Morlan,
1953. ; .

Locality: Africa - Sudan. ’. ,-. .

.
Host: Mammal - "bats,"

Hirstionyssus incomptu.s. Eads and Hightower

Type material in the U. S. National Museum.

Neoichoronyssus (Hirstionyssus) incomptus Eads and Highlower, 1952:
32. .

Hirstionyssus incomptus, Strandtmann and Morlan, 1953.

-
Locality: North America United States, Texas.
Host: Mammal - Djpodomys ordil.

Hirstionyssus isabellinus (Oudemans)

Liponyssus isabellinus Oudemans. 191.3: 384 (d); 1913: 80 (d. illus.).

Ichoronyssus isabellinus, Ewing, 1922: 14; Radford. 1939.
Hirstionyssus isabellinus, Fonseca, 1948: .297; Willmann, 1952; Strandt-
mann and Morlan, 1953; Bregetova, 1956: 181 (t, d. cr and o’illua.h
’
Baker etal., 1956. ’
.
.
.
Neoichoronyssus isabellinus, James on ~
et al., 1952: 11; Suyemoto et al.,
’~
1954.
.
Hirstionyssus arvicolae Zemska, 1951; Dubinin, 1953; Bregetova, 1953...
(Synonymy according to Bregetova, 1956.)
Localities: Asia - Japan; Korea; U.S.S.R. Europe - England; Ger-
many; Netherlands. North America - Canada; United States.
-
Hosts: Mammals Apodemus agrarius. Apodemus flavicollis^Apo-
demus sylvaticus. Arvicola amphibius, .Arvicola terrestris, Clethrio-
nomys rutilus, Eutamias sibiricus, Lagurua curtatus, Lemmus lemmus,
Microtus arvalis, Microtus montanus,’ Micrbtus oeconomus, Microtus
pennsylvanicus, "mole, " Mus musculus, Mustela erminea, Mustela nx-
valis, Mycromys minutus. Ondatra zibetheca, Paludicola amphibius,..
 .. . 105

Putorms sp., Rafrtus norvegicus caraco, Talpa europaea.

Hirstionyssus latiscutatus (DeMeillon and Lavoipierre)

Liponyssus latiscutatus DeMeillon and Lavoipierre, 1944; 62 (d).

Hirstionyssus latiscutatus, Fonseca, 1948: 298; Strandtmann and Moi
lan, 1953. ,.
Locality: Africa - Union of South Africa, Transvaal.
Host: Mammal - Rattus rattus.
.
.
Hirstionyssus liberiensis (Hirst)

Type material in the BritishMua’eum (Natural History).

.
Leiognathus liberiensis Hirst.. 1912: 364,
Liponyssus liberiensis. Hirst, 1921; Radford,’ 1944.
Hirstionyssus liberiensis, Fonseca, 1948; Strandtmann and Morlan, 1953.
.Locality: Africa - Liberia; Uganda.

-
-
Host: Mammal Heliosciurus rufobrachium.

Hirstionyssus macedonicus (Hirst)

Liponyssus macedonicus Hirst, 1921: 789’. ’

1953; Bregetova, 1956: 172 (d-and ,
Hirstionyssus macedonicus, Fonseca, 1948; Strandtmann and Morlan,
illus.).
Locality: -Europe - "Macedonia"; U.S.S.R.
-
Hosts: Mammals Spalax monticola thermaicus, Spalax microph-
thalmus.

Hirstionyssus meridiahus Zemska

Hirstionyssus meridianus Zemska, 1955; Bregetova. 1956: 179 (d’and$,

i-llus.).
"
.
.

Localities: Europe and Asia - U.S. S.R.. Turkmenia.

Hosts: Mammals - Meriones erythrourus, Merione.s meridianus,
Rhombomys opimus.

Hirstionyssus musculi (Johnston)

Permanyssus musculi Johnston, 1849. Not Dermanyssus musculi’C. L,

Koch, 1836. ’ . . .
.
Hirstionyssus musculi, Bregetova, 1953: 329; 195.6: i84 (d’ and 9, illus.).
Macronyssus? johnstoni Oudemans, 1936: 284. (Oudemans incorrectly
proposed this name for Johnston’s species. By Oudemans’ own
statement, musculi Johnston was not a junior homonym of musculi
Koch, but a distinct species belonging to a different genus.)
Localities: Europe and Asia - U.S. S.R.
Hosts: Mammals - Apodemus agrarius, Apodemus speciosus, Apo-
demus sylvaticus, Clethrionomys rufocanus, Microtus gregalis, My-
106 , ’;-

cromys minutus, Rattus norvegicus caraco, Rattus turke.stanicus.

-
.

Hirstionyssus ngami (Lavtoipierre). New combioati.on’ .

Type material in the Liverpool School of Trop.ifc’al’ Medicine.

Liponyasus ngami Lavoipierrei 1953: 306’Cd,’ jtlu’s.).

’
"

Locality: Africa - "British-Cameroons, Barombikang’Forest. "

Host: Mammal - Galago allerii, a primate. ..’.
Remarks: Described from two female, specimens. ,,..
’

Hirstionyssus neotomae Eads and Hightower’.-. ^

.

Type material in the U. S. National Museum*’ ’ ..’’ ’

Neoichoronyssus (Hirstionyssus) neotomae Eads. and Hightosrgr, 1951:

’ %
-’ .

’
295 (d). ",-. ,

Hirstionyssus neotomae, Strandtmann and Morlan. .1953:-630.,

-
Locality: North America United States,...-
’

..;-’’ .. . .,-’
Hosts: Mammals - Neotoma albtgula, ’Neotoma. fuscipes, N’eotoina
’.’

’
micropus. .

Hirstionyssus obsoletus Jameson

Type material in the U. S, National Museum.

Hirstionyssus obsoletus Jameson, 1950: 32 (d); .Strandtmann and.Mor-
lan, 1953. , .:;’ ,.,
Locality: North America - United. .-States, California.-
Hosts: Mammals - Clethrionomys sp., Neure’fr’ichus sp., Pero-
myscus sp., Sorex trowbridgei. .’ ’"
’
.
.
Hirstionyssus occjdentalis (Ewing)

Type material in the U. S. National Museum. ,. ".

Liponyssus occidentalis Ewing, 1922: 20 (d); Augustson, 1941; Jameson,
:.
-
1947. .
.’. ..’
.
.
Hirstionyssus occidentalis, Strandtmann and Morlan, 1953: 632; Strandt-
mann, 1956: 138,
Locality: North America - United States.
Hosts: Mammals - Citellus (Callospermophilus) lateralis chryso-
deiaus, Cryptotis parva, Microtus montanus dutcheri, Ochotona schisti-
ceps muiai, Sciurus hudsonicus richardsoni. .’’.’.--
Remarks: Strandtmann and Morlan (1953) gave drawings of this mite
but they erred in showing acute spurs on coKae II and HI;.’actually they
are blunt, either truncated or rounded. Contrary to their^statement.
cynomys Radford is not a synonym of occidentalis. ....
Hirstionyssus otomys (Radford) ,. :

Liponyssus otomys Radford, 1942: 190 (d). ,’;.’..

 107

Hirstionyssus otomys, Fonseca, 1948; Strandtmann and Morlan, 1953.

Locality: Africa - Uganda. :. :.’
.

Host: Mammal -’ Otomys sp.

Hirstionyssus pachypus Willmann

Hirstionyssus carnifex subspecies pachypus Willmann, 1953: 467 (d)i

Hirstionyssus. pachypus, Bregetova, 1956: 185.
-
Locality: Europe Germany; Austria. ’,. .

Host: Mammal - Nest of Talpa’europaea.

Hirstionyssus pauU Willmann

Hirstionyssus pauli Willmann. 1952: 407; Strandtmann and Morlan, 1953;

Bregetova. 1956: 177.
Locality: Europe - Germany, Silesia.
Host: Mammal - "Squirrels. "

Hirstionyssua santos-diasi Zumpt

Hirstionyssus santos-diasi Zumpt, 1951; 7(d); Strandtmarm and Morlan,

1953.
Locality: Africa - Union of South Africa, Cape Province; Mozam-
bique. .
.
-
Host: Mammal Pedetes cafer,

Hirstionyssus sciurinus (Hirst)

Type material in the Paris Medical Faculty Museum.

Liponyssus sciurinus Hirst, 1921: 785.
Hirstionyssus sciurinus, Fonseca, 1948; Strandtmann and Morlan, 1953;
Bregetova, 1953; 1956: 161 (c? and 9, illus.).
Locality: Europe - France; U.S.S. R., "Crimea."
Hosts: Mammals - "Squirrels, " Sciurus vulgaris.

Hirstionyssus soricis (Turk)

Type material in the private collection of F. A. Turk, England.

Ichoronyssus soricis Turk, 1945: 133 (d).
Hirstionyssus soricis, Fonseca, 1948; Strandtmann and Morlan, 1953;
Bregetova. 1956: 194.
Locality: Europe - England.
Host: Mammal - Sorex minutus.
Remarks: Only the male and nympli are known.

Hirstionyssus staff ordi Strandtmann and Hunt

Type material in the U, S. National Museum.

108

Hirstionyssus Stafford! Strandtmann and Hunt, 1951: 460 (d); Morlan.

1952; Strandtmann and Morlan, 1953.
Locality: North America - United States.
-
Hosts: Mammals Mephitis elongata. Mephitis mesomelas, Spil-
ogale interrupta, Spilogale leucoparia. Spilogale putorius. .

Hirstionyssus triacanthus (Jameson) ." ..:.’’.

.
Type material in the U. S. National Museum.
Neoichoronyssus (Hirstionyssus) triacanthus Jameson, 1950: 166 (d).
Hirstionyssus triacanthus, Strandtmann and Morlan,. 1953.’
Locality: North America’-United States. .1 ,

Host: Mammal - Dipodomys merriami.. . ;

.
Hirstionyssus transiliensis Bregetova

Hirstionyssus transiliensis Bregetova, 1956: 179 (d, d’and$, illus.).

Localities: -Europe and’ Asia - U.S. S.R.’-
Hosts: Mammals - Clethrionomys frater, Microtus’ gregalis. .’

Genus Neoichoronyssus Fonseca. -1941

(Fig.-45)

Type: Liponissus wernecki Fonseca. .-’.’,

.
Diagnosis: Dorsal plate undivided and covering most of the dor-’
sum. Female sternal plate with’only four setae; epigynial plate pointed
posteriorly; coxa I with a setigerous spur; palp trochanter bearing a ven-
tral spur. Male holoventral plate undivided and not expanded behind coxa
’
’
IV.
.
Neoichoronyssus wernecki (Fonseca)
(Pig. 45)’.’ " .

Liponissus wernecki Fonseca, 1935:. 74 (d). ... .

.
Neoichoronyssus wernecki, Fonseca, 1941: 263 (t); Fonseca, 1948; Mor-
lan and Strandtmann. 194&(c); Morlan,. 1952; Strandtmann and Mor-
lan, 1953. ,: ... .
Localities: North America - United States. South America-Bra-
zil. Sao Paulo. . ’.
.
Hosts: Mammals - Didelphys aurita, Didelphys virginianus. Sig-
modon hispidus. ....; ’’i ;.; "./..; ,

Remarks: The opossum (Didelp&ys’): is proBably the true host of this .

mite. ’
’
Neoichoronyssus dentipes (Strandtmann and Eads)

Type material in the U. S. National Museum. :. .

 109

Ichoronyssus dentipes Strandtmann and Eads, 1947: 51 (d).

Hirstionyssus dentipes, Fonseca. 1948: 297; Jameson, 1950.
Neoichoronyssus dentipes,: Jaroeson, 1.950: 162 (t); Strandtmann and Mor-
lan, 1953.
Localitie.s; North America -.United States. South America Chile. -
Hosts: Mammals - Sigmodon hispidus, Proechimys sp.

Genus Patrinyssus Jameson,.: 19 50

,
, (Pig. 46) .

.
.
,
Types Ichoronyssus hubbardi Jameson.
Diagnoses: Dorsal plate entire and covering most of the dorsum.
Female palp without spurs; epigynial plate rounded posteriorly. Male
ventral plate divided just posterior of coxae IV; some of the coxae with
setigerous spurs.

Patrinyssus hubbardi (Jameson)

(Pig. 46) "". . .,

Type material in the U. S. National Museum.

Ichoronyssus hubbardi Jameson, 1949: 109 (d); Jameson, 1950.
Neoichoronyssus (Patrmyssus) hubbardi. Jameson, 1950: 162 (t).
.

.
’ ,

Neoichoronyssus hubbardi. Strandtmann and Morlan, 1953: 628. . . .

Locality: North America - United States, Washington state.

Host: Mammal - Aplodontia rufa.

Genus Echinonyssus Hirst, 1925

(Fig. 47)
.

Type: Echmonyssus nasutus Hirst.

Diagnosis: Similar to Hirstionyssus, but with dorsum, projecting
anteriorly over gnathosoma. Anterior spur of coxa II developed into an
enormous hook; all coxae with heavy spurs. In the generotype the dor-
sum projects forward prominently.
Remarks: Known only from the female. It is quite possible that
species of Echinonyssus should be placed with Hirstionyssus. Certainly
Echi-n.ony-ssus.< Neoichpronys sus, Patrinyssus, and Hirstionyssus form
a closely related group.

Echinonyssus nasutus Hirst

(Fig. 47)
Echinonyssus nasutus Hirst, 1925; 51 (d. 9 only); Fonseca, 1948.
Locality: Africa.
Host: Mammal - Tupaia picta.
110

Echinonyssu’s validipes Domrow

Type material in the Queensland’ Institute of Medical .Research,

Brisbane.
Echinonyssus validipes Domrow, 1955: 133 (d. cf and.S illus.); Domrow
.’ H
and Smith, 1956; 20.2, .

Locality: Australia - Queensland, Mt. Nebo. .:;. ^

Host: Mammal - Potorous tridactylus.
Remarks: Differs from the generotype in not possessing a dorsal,
anterior projection.’ In the male, the anal plate is separated from the
ventral plate.

Genus Lepronyssoides Fonseca’,

’
1941
; ’^igr48) ’,’;,.

.
Type: Liponissus pereirai Fonseca.
Diagnosis: Dorsal shield undivided; sternal plate with the usual
threepairsof setae and two pairs of slitlike pores, bearing, in addition,
two special structures in the form of large infundibuliform orifices a
little behind the anterior pair of setae; genital plate scalelike, tapering
posteriorly to a point; tibia long, as also are the other segments of the
legs; a ventral spine is present on coxa III. (Copied.from Fonseca, 1948:
268.)
’
’"’
Lepronyssoides pereirai (Fonseca)

-
(Pig. 48)
Liponissus pereirai Fonseca, 1935: 80 (d).
Lepronyssoides pereirai, Ponseca, 1941: 263 (t); Fonseca, 1948: 268,
-
Locality: South America Brazil, State of Parahuba, State of Rio
Grande.
Host: Mammal - Kerodon spixi, also a wild rat commonly known
as "punare."
Lepronyssoides matogrosso (Fonseca). New combination

Neoichoronyssus (Hirstionyssus) matogrosso Fonseca, 1954: 80 (d).

Locality: South America - Brazil, Mato Grosso.
,
Host: Mammal - Cercomys cunicularis forsteri.
Remarks: The description and illustrations of this mite fit the- diag-
nosis-of LeprMiys^oi^les perfectly, except for the absence of the infun-
dibuliform orifices on the sternal plate,

Genus Hirstesia Fonseca, 1948

(Fig. 50)
Type: Liponyssus sternalis Hirst,
Diagnosis: Epigynial shield rounded-posteriorly and bearing from
four to six setae; sternal plate with two large, stigmalike structures in
 .-.’’. ul

the anterior, corners’; male holoventral plate undivided or entire,

Remarks; The species at present included in this genus do not:-seem
’ 1;
.;
’

to be. congeneric. ’
. : .

.
.
.

.
,.

.
.
. ,
";
Hirstesia sternalis (Hirst)

.
.
(Fig. 50)
Liponyssus sternalis Hirst, 1921: 781 (d). :... .-"
Lepronyssoides sternalis, Ewing, 1947: 87 (t).

.
Hirstesia sternalis, Fonseca, .1948: 266 (t),’Turk, 1952: 479 (d).
Locality: Europe - Greece, Salonika; England, South Devon.
Hosts: Mammals - "bat m.cave,..";Rhinolophu’s hjpposiderds minu -
tus. ’.’ ... :’.:.’:’." -

.
: Hirstesia britteni Radford . . -..;

Hirstesia britteni Radford, 1953: 239 (d). ’.-’

Locality: Europe - England. .. ; ’.’..,. .

Host: Mammal - Pipistrellus oipistrellus.

Remarks: The female epigynial plate has five setae; the holoven-
tral plate of the male is undivided and’greatly expanded behind coxae IV.

Hirstesia kenyaensis Radford

Hirstesia kenyaensis Radford, .195,1; 99 .(d);.Keegah, 1956.

Locality: Africa - Kenya.
Host; Mammal - "a bat. "
Remarks: This mite does not appear to be congeneric with sternalis.
The legs are much shorter, the.re is a spur on c.oxa IV, and the epigyn-
ial shield has only two pairs of setae. The ventral armature of the male
consists of two widely separated plates. .,,’

Hirstesia transvaalensis Zumpt

Type material in the South African Institute for Medical Research.

Hirstesia transvaalensis Zumpt, .1950:. 89 (d); Keegan, 1956: 213.
Locality: Africa - Union of South Africa, Transvaal, Sterkfontein
’-;:
Caves.
Host: Mammal Miniop.terus natalensis, Rousettus lanosus.
-

Remarks: Differs from the generotype in having only two pairs of

setae on the epigynial plate. . , : ., ,
.
.
Genus Neolaelaps Hirst, 1926
(Fig. 51)
Type: Liponyssus magnistigmatus Vitzthum.
Diagnosis: Peritremal tube unusually wide; female epigynial plate
bearing three pairs of setae. .Holoventral plate of male undivided and
slightly expanded behind coxae IV. All body setae unusually strong.
112 ,, .

Remarks: T.his mite has a very strong superficial resemblance to

Laelaps but it has a pointed, typical macronyssid tectum, a single file
of 9 - 10 deutosternal teeth, peritremalia that extend behind coxae IV,
nonsclerotized chelae, and it lacks discrete corniculi. All of these in-
dicate its affinities are with the Dermanyssidae rather than the Laelap-

"
tidae. :’.. .
^, ,

Neolaelaps magnistigmatus (Vitzthum)

(Fig. 51)
Liponyssus .magnistigmatus Vitzthum, 19l8: 21 (d); Vitzthum, 1926: 93
(c, d).
’
Laelaps (Neolaelaps) magnistigmatus. Hirst, 1926: 836.
Neolaelaps magnistigmatus, ’Radford, 1950.
Liponyssus echinus Oudemans, 1925; 31; Oudemans. 1927b; 198 (d). New
synonymy.
Neolaelaps echinus, Radford, 1950: 370.
Locality: Indonesia - Dutch East Indies.
Hosts: Mammals - Pteropus edule’., Pteropus giganteus, Pteropus
vampyrus,
Remarks: It is quite probable, as Hirst (1926) suggested, that Leio-
gnathus spinosus- Berlese,. 1910 is this species. The trivial name sug-
gests this, and it conies from the same host and locality. However, the
origin.al description is not diagnostic and there were no illustrations .
Until the types can be studied it is perhaps best to leave it as is. Oud-
emans’ fine drawings (1927b) leave no doubt about the conspecificity of
magnistigmatus and echinus.
"
.

Genus Liponysella Hirst, 1925

(Fig. 49)
Type: Liponyssus madagas cariensis Hirst.
Gneidolaelaps Ewing, 1929. .... ’

Type: Laelaps barbatus Ewing.

Diagnosis: Epigynial plate of-female truncate posteriorly and bear-
ingthree pairs of setae; chelae edentate but with a basal brush of setae;
anterior seta of coxa II and III spiniform, all other setae slender. Holo-
ventral plate of male entire and slightly expanded behind coxa IV.

Liponysella madagascariensis (Hirst)

(Fig. 49)
Liponyssus madagascariensis Hirst, 1921: 780 (d).
Liponysella madaga-scariensis. Hirst, 1925: 52 (t); Fonseca, 1948; Zumpt.
1950: 166 (d, rf and deutonymph),
Laelaps barbatus Ewingy 1925: 2 (d). ;

Gneidolaelaps barbatus, Ewing. 1929 (1); Baker, and Wharton. 1952: 85

’
(synonymy). ’

...
Locality: Africa - Madagascar.
Hosts: Mammals - Lemur albifrons, Propithecus verreauxi.
 113

Genus Steatonyssus Kolenati. 1858

(Fig. 52)
Type: Dermanyssus murinus Lucas, 1840 ( = Steatonyssus
periblepharus Kolenati, 1858)
Ceratonyssus Ewing, 1922: 6.
Type: Dermanyssus musculi Koch.
Diagnosis: Dorsal plate of female divided into two large shields of
about equal length, but anterior plate wider than posterior; female stern-
al plate about twice as wide as long, posterior margin thickened. Dor-
sal plate of male entire; holoventral plate entire, not expanded behind
coxae IV. Primarily ectoparasites of bats but occasionally also of other
mammals.
Remarks: We are familiar with a few species in this genus and with
most of the recent literature dealing with it but we have seen no type ma-
terial, except that of Ewing, and only some of the original literature of
the generotype and the other older species. Hence, we are indebted to
Dr. Fonseca’s monograph of the Macronyssidae (1948) for the majority
of the synonymy and references,
Although Steatonyssus musculi (Schrank, 1803) is the oldest des-
cribed species in the genus, it cannot be the generotype (Fonseca, 1948).
When Kolenati created the genus Steatonyssus he included only two spe-
cies , Steatonyssus pe riblepharus and Steatonyssus brachypeltis, and
therefore one of these must be the generotype. Since both of these names
are included as synonyms of Steatonyssus murinus (Lucas, 1840), it
must necessarily be the generotype.
As the genus is here understood, the species contained therein should
c onform to Steatonyssus joaquimi (Fonseca, 1935: 115 or 1948: 317). Will-
mann(1936) synonymizedEwing’s Ceratonyssus with Steatonyssus in his
description of S^. spinosus.

Steatonyssus murinus (Lucas)

Dermanyssus murinus Lucas, 1840: 698. (Quoted from Fonseca.)

Steatonyssus murinus, Fonseca, 1948: 318; Turk, 1952: 484 (c); Will-
mann, 1955; Keegan, 1956.
Dermanyssus avium Wagner, 1841: 11. (Quoted from Fonseca.)
Gamasus yespertilionis Gervais, 1844: 222.
Steatonyssus periblepharus Kolenati, 1858; Kolenati, 1859: 186.
Steatonyssuj; brachypeltis Kolenati, 1858; Kolenati, 1859: 187.
Liponyssus chiropteralis Hirst, 1921: 776 (d).
Localities: Africa - Algeria. Asia - "Tiberias." Europe - Eng-
land; Germany; Hungary; Sardinia.
Hosts: Mammals - Brachyotus capaccinii ( = Myotis capaccinii),
Ctenodactylus gundi, Isotus ciliatus, Myotis myotis, Nannugo pipistrel-
lus (=Pipistrellus pipistrellus), Pipistrellus kuhli, Plecotus auritus.
114

Steatonyssus brucei Lavoipierre

Type material in the Liverpool School of Tropical Medicine.

Steatonyssus brucei Lavoipierre, 1956 (d, illus.).
Locality: Africa - Northern Nigeria, Bernin Kebbi.
Host: Mammal - Nycteris sp. .’".

Remarks: Described from six undamaged female raises.

Steatonyssus ceratognathus (Ewing)

Type material in the U. .&. National Museum.

Ceratonyssus ceratognathus Ewing, 1922:. 11 (d); Miller, 1925.
Steatonyssus ceratognathus, Fonseca, 1948:; 3,16. . ,

Locality. North America - United: States,. North Carolina.

Host: Mammal- "Taken.from a small bat. N., hume rails. " .

Steatonyssus cyclaspis (Oudemans) ...

.
Liponyssus cyclaspis Oudemans, 1906: 61 (new...name tor Liponyssus
pipistrelli Oudemans, 1906, not Oudemans. 3-.904); Oudemans, 1915.
Steatonyssus cyclaspia; .Fonseca, 1948: 316. ’,.

.
’

Liponyssus pipistr-elli Oudamans, 1906: 37 (prptony^mph). ,,’ ’

...

.
.
-
LocaIi+y:-’-.Europe: Netherlands. :. ;.i, ... . ; .

Host: Mammal- -. Pi.pj.strellus pipistrellus. ..

.
Remarks: All of the above references are taken from Fonseca (1948). /
We have not seen Oudemans’.papers. ; , ,
. ..;

’
: ’.Steatonyssus eos Zumpt and Till

Type material in’the South African Institute for Medical Research.

Steatonyssus eos Zumpt and Till, 1954: 55 (d); Keegan, ,1956,.
Locality: Africa -Kenya, Muhoroni. ..

Host: Mammal - Pipistrellus nanus. ; ,.

.
Steatonyssus javensis (Oudemans) t

Liponyssus javensis Oudemans, 1914: 69 (d); Oudemans. 1915: 167.

Ceratonyssus javensis, Ewing, 1922: 6.’ ,..., .

Steatonyssus javensis, Buitendi.jk, 1945; Eonseca, 1948: 316.--

Localities: Asia - India, Bombay. Indonesia -’Java,;.Dutch East
Indies. ’ ’ :
.. ,
.
.. ;

Host: Mammal - "Chiropteran-species. "

Steatonyssus joaquimj. (Fonseca)

.
(Pig. 52)
Ceratonyssus joaquimi Fonseca, 1935: 89 and 98 (d).
Steatonyssus joaquimi, Fonseca. 1948: 317. ..,.
Locality: South America - Brazil, Sao Paulo.
 115

Host: Mammal - Glossophaga soricina.

Steatonyssus musculi (Schraak)

Acarus musculi Schrank, 1803: 208 (d). (Quoted from Fonseca.)

Steatonyssus musculi (Schrank) Fonseca, 1948,: 270 and 318 (t); Breg-
etova, 1953; 1956. ’ : . . .

Dermanyssus musculi Koch, 1-836: 13 (d),.. Not musculi Schrank.

Liponysaus musculi (Koch) Oudemans, 1902: 17 (nympha II); Oudemans,
1912.’ ..

Ceratonyssus musculi (Koch) Vitzthum. 1926: 30.

Liponyssus pipistrelli Oudemans, 1904: 18.
Localities: Africa - "The Congo. " Asia - India, Bombay. Europe -
Germany; Netherlands; Spain, U. S. S. R.
Hosts: Mammals - Eptesjcus serotinus, Myotis .mystacina ’., Mus
musculus, Nyctalus noctula, Vesperugo pipistrelli, Vespertilio pip is -
-
trellus, Vespertilio serotinus. Bird Acanthis cannabina.

Stga^onyssus natalensis Zumpt and Patterson

Type material in the Natal Museum, Pietermaritzburg.

Steatonyssus natalensis Zumpt and Till, 1951: 86 (d); 1954: 54.
Locality: Africa - Union of South Africa, Natal, Pietermaritzburg.
-
Host: Mammal Mihiopterus natalensis.

Steatonyssus -nyassae (Hirst)

Liponyssus nyassae Hirst.- 1921: 777 (d). . . .

.
. Steatonyssus nyassae, Fonseca, 1948: .319; Zumpt and Till. 1954: 49.
Locality: Africa - Nyasaland. CHiromo.-
Hosts: Mammals - Elephant shrew. Elephantulus sp., Scotophilus
murinoflavus. .

Steatonyssus occidentalis (Swing)

’
Type material in the U. S. National Museum. ,
.
Ceratonyssus occidentalis Ewing, 1933: 10 (d).
Steatonyssus occidentalts,. Fonseca, 1948: 319; Radford, 1950; Yunker.
1958.
Locality: North America-
Virginia.
- United States, Maryland,
.
Oregon, West
Hosts: Mammals - "Bat (?) Myotis sp.," Eptesicusfuscus, Lasiurus
borealis. Myotis lucifugus.
.
.
Steatonyssus spinosus Willmann

Steatonyssus spinosus Willmann, 1936: 152 (d); Fonseca, 1948.

Locality: Europe - Germany, Hamburg, Zoological garden.
 .:
’

116 .

.
.
.
.
Host: Mammal - Solenodon paradoyu.s.. . ;. . ...

Remarks: The host had been imported from Haitu The mites were
found only on the fresh feces, not on the .host.

Steatonyssus sudanensis (Hirst)

Liponyssus Budanensis Hirst, 1926; 835 (a).

_
..’ ’- ..

.
Steatonyssus sudanensis, Fonseca,"1948; 319; Zumpt and Till, 1954.
Locality: Africa - Sudan; Khartoum..
-
Host: Mammal Liponycteris nudiventris,

Steatonyasus’superans Zemska

Steatonyssus superans Zemstea,. 1949; Bregetwra," 1953; 325; ’1956.-

Locality: Europe and Asia - U. S. S. R.-,"’P"rimor. :.
Host: Mammal - "Vespertilio .’super-ans.
.
Genus Pellonyssus dark and Yunker, 1956
(Fig. 53),
Type: Pellonyssus passeri dark and Yunker.
Diagnosis: Female with two dorsal plates as in Steatonyssus. Fe-
male sternal plate more than twice as wide as long and without a thick-
ened posterior margin; sternal setae I smallerthan II and III; chelicerae
gradually attenuated apically. Male with entire dorsal plate and undi-
vided holoventral plate. Parasitic on birds..
Remarks: Mites of this genus very closely resemble those of the
chiropterophagic genus Steatonyssus. However, the-.shape of the female
sternal plate and the male chela are constant and peal. differences. In
Steatonyssus the male chela is tri-prohged;’in Pellonyssus it is heavier"
and more "fistlike."

Pellonyssus passeri Clark- and Yunker

(Fig. 53)
Type material in the U. S. National Museum. ..’
.
Pellonyssus passeri Clark and Yunker. 1956: 94 (d, ,b,. all’stages lllus.).
Locality: North America - United States, Maryland.
Host: Bird - Passer domesticus.
Remarks: Clark and Yunker cultured this mite for 15 days. The
life cycle includes the egg (laid off the host), non-feeding,hexapod larva,
active and feeding protonymp.h, non-feeding deutonymph, and adult male
and female.

Pellonyssus biscutatus (Hirst)

Liponyssus biscutatus Hirst, 1921: 779 (d).

’
’

Steatonyssus biscutatus, Fonseca, .1948: 316; Zumpt and Patterson, 1952:

163 (c); Zumpt and Till, 1954.
 -117

Pe’lloriyssus biscutafus.-’-Clark and Yunker; 1956: 94 (t).

Locality: Africa - Union of South Africa ; Northern Rhodesia.
"’ ’Hosts: Birds - Campethera.abingoni, Dendropicus cardinalis, Den-
dropieus fuscescens. ... . .’,..

Pellonyssus malurus (Womersley). New combination

Steatonyssus malurus Womersley, 1956: 214 (d, S.illus.).

Locality: Australia - Northern Australia, Beseick Creek, Cather-
ine.
Host: Bird - Afalurus melanocephala.

Pellonyssus reedi (Zumpt and Patterson)

Steatonyssus reedi Zumpt and Patterson, 1952: 163 (d); Zumpt and Till,
1954: 53.
Pellonyssus reedi, dark and Yunker, 1956: 94 (t).
Locality: Africa - Union of South Africa, Johannesburg.
Hosts: Birds - Creatophora cinereus, Hyphantornis velatus, Ony-
. chognathus morio. Passer melanura, Ploceus velatus.

Pellonyssus sinnlis (Zumpt and Till)

Type material in the South African Institute for Medical Besearch.

Steatonyssus similis Zumpt and Till, 1954: 51 <d).
Pellonyssus simil’is, dark and Yunker. 1956: 94 (t).
Locality: Africa-Union of South Africa, Transvaal, Potchefstroom.
Host: Bird - Plocepas s er mahali.

Pellonyssus viator (Hirst)

Liponyssus viator Hirst. 1921: 775 <d, S.illus.).

Steatonyssus viator, Fonseca, 1948: 319; Bregetova, 1956: 161; Zumpt
and Till, 1954; Keegan, 1956.
Pellonyssus viator, dark and Yunker. 1956: 94 (t).
Locality: Africa - Egypt; Morocco, Tangier. Asia - India, Calcut-
ta; U. S, S. R., Tadzhikistan..
Hosts: Birds - Cypselus affinis, Gecmus vaillanti. Passer domesti-
cus. Passer hispaniolenais. Passer montanua..

Genus Radfordiella Fonseca, 1948.

(Fig. 54)
Type: Radfordiella oudemansi Fonseca.
Kolenationyssus Fonseca, 1948: 267.
Type: Kolenationyssus athleticus Ponseca.
Diagnosis: Dorsal plate entire and covering most of the dorsum.
Epigynial plate 01 female rounded posteriorly, with one pair of setae.
118

Coxae unarmed. Male with undivided holoventral p’late that is not ex-
panded behind coxae IV.
Remarks: Kolenatiohyssus athleticus is known only from the male
and differs from the male of Kadfordiella ’oudemansi. apparently only in
having unpedunculated claws on tarsus II. We believe this denotes male
dimorphism within the species and not a generic difference. Both types
of males were taken from the same host,

Radfordiella oudemansi Fonseca

(Fig. 54) .
Type material in the Institute Butantan, Sao Paulo, Brazil.
Radfordiella oudemansi Eonseca, 1948: 274 (d).
Kolenationyssus athleticus Fonseca. 1948: 276 (d). New synonymy.
-
Locality: South America Brazil, Butantan.
-
Host: Mammal Desmodus rotundus rotundus.

Genus Tur Baker and Wharton, 1952

(Pig. 55)
.
ProtonyssusTurk, 1946:347. Not Protonyssus Trouessart, 1915.
Type: Protonyssus uniscutatus Turk.
Diagnosis: Female epigynial and anal plates fused and bearing four
pairs of setae in addition to the endopodal and anal setae.
Bemarks: This genus is based on a single female specimen mounted
in Canada balsam. It does not appear to have the characteristics of the
Macronyssinae, but is retained in this group because it will not fit any-
where else. .

Tur uniscutatus (Turk)

(Fig. 55)
Protonyssus uniscutatus Turk, 1946: 348 (d).
Tur uniscutatus. Baker and Wharton, 1952: 85 (t).
Locality: South America - Ecuador, Bulun.
Host: Mammal - Proechimys calidius calidius.

Subfamily DERMANYSSINAE Kolenati, 1859

Diagnosis: Medium sized mites capable of great enlargement when

engorged. Female chellcerae greatly attenuated, with minute chelae;
coxae unarmed; epigynial plate with only one pair of setae.
Remarks: This group is in great need of revision.

Genus Allodermanyssus Ewing, 1923

(Pig. 56)
Type: Dermanyssus sanguineus Hirst. :
.
Diagnosis: Female with two dorsal shields, the anterior large and
 119

tapering posteriorly, the posterior small and pygidial; sternal plate about
as broad as long; epigynial plate narrow and .pointed. Male with a single
dorsal plate tapering, posteriorly.’ Primarily parasites of rodents.

AIlodermanyssus sanguinftitS’; (Hirst) ..

., (Fig. 56) .. /..,,

’., ’ -

’
" ’

Dermanyssus sanguineus Hirst, 1914:’210..(4),- .

.

AIlodermanyssus sanguineus; Ewing, 1922’;.8 (t); Ewing, -1942: 74 <c,); .

Greenburget.al.., 1.947: 901 (m); Huebner etaL," 1946: ’1605, 1677

(m); Huebner et al., 1947: 777 (m); Pratt and Frit z, 1947 (atlas);
Pratt etal., :;1949; Reiss, 1949: 390 (m); Zumpt. 1950; Bishopp and
Philip, 1952; Jameson (rt al., 1982; Rm.dge, 1952; Eustis and Ful-
ler,"1952: 546.(m);. Nichols et al., 1953: 92 (b); Fuller, 1954: .236
(b); Bregetova,. 1-956: 203 (illus.-)j Baker et al., 1956: .18 <c); Kee-
gan, 1956. ’
Dermanyssus (Allodermanyssusy’sanguineus, Hirs^, ’1923; Hirst, 1:926;.
Matheson, 1950. ’’/; ’.: :., ’’’ , .’,.

.
Localities: Africa.- Egypt; Sudan. Asia - U.S.S.R., Ukraine.
Europe. North America - United. States. ," ’

.
.
Hosts; Mammals,-.Acomy s cahirinus, Acomys dimidiatus, Acoroys
russatus, Alticola argentatus, Arvicanthis niloticus, Cricetulus migra-
torius. Homo sapiens, "man," .Meriones persicus, Kus musculus, ’.Mus
musculus’ gentil.is, ’^tattus alexandrinus, Rattus norvegicus,’ Rattus rat -
tus, Rattus turkestanicus, "rodents."
Remarks: The preferred host of this mite is apparently the common
house mouse,’ Mus musculus. Ewing (1942) wrote that’a specimen had
been found in the United States in 1909; but was,not reported again until
1938. (It has been frequently reported since that date.) In the/same
paper Ewing also commented that the mite bites man and may cause a
rash. In 1946, Huebner et aL. reported that Rickettsia akari, the causa-
tive agent of rickettsialpox, could be’recovered from this mite and sug-
gested it as a transmitter of the disease. According to Bishopp and
Philip (1952). there is a. report from Russia that the mite can transmit
tularemia. Fuller (1954) found the life cycle to include egg, larva, two
nymphal stages and adult male and female. The larva does not feed, but
both nymphal instars do. -
AIlodermanyssus aegyptius (Hirst)

Dermanyssus (Liponyssoides’) aegyptius Hirst, 1913: 119 (d).

Dermanyssus aegyptius. Hirst, 1-915: 222 (d); Radford, 1950... ..
Allbdermanyssus aegyptius, Keegan, 1956: 205 (t).
Locality: Africa -’Egypt: : ,
.
.
Hosts: ’Mammals - Acomys cahirinus, Arvicanthus niloticus. Hat- .

tus norvegicus, Rattus rattus. ; ;

Remarks: Wehave not seen the types of this species. It was placed
in Allodermanyasus by Keegan (ibid.). There is a strong possibility that
this maybe a synonym of sanguineus Hirst, in which case aegyptius would
 120

have to be the accepted name as it has priority by nearly a year.

. .:’.. "
Genus Liponyssoides Hirst, 1913
"

Type: Permanyssus (Liponyssoides)

’

’
muris Hirst.
Diagnosis: Like Allodermanyssus except the pygidial plate is lack-
ing; the undivided dorsal plate covers about half of the dorsum and in un-
fed specimens reaches nearly to the posterior tip. Parasitic on rodents.
Remarks: It is quite possible that a thorough study of the Derma-
nyssinae will show Liponyssoides and Allodermanyssus to be the same.

Liponyssoides muris (Hirst). New combination

Dermanyssus (Liponyssoides) muris Hirst, 1913: 119 (d); Baker and

Wharton, 1952.
Dermanyssus muris. Hirst, 1915: 2i’6(d); Hirst. 1916: 185 (c); Hirst,

. 1916: 63 (d); Hirst, 1920; Hirst. 1925; Hirst,,1926; Finnegan, 1945;

Lavoipierre, 1946; Radford. 1950; Keegan, ’1956,
.
Localities: Africa - Belgian Congo; Egypt; Nyasaland. Asia Arab-

-
ia; Ceylon; Formosa; India. ;
.
’"’
Hosts: Mammals - Acomys sp.. "Arvicanthis niloticus, Rattus nor-
vegicus, Rattus rattus. "’ .’.
Remarks: Hirst illustrates this" species as having a single dorsal
:plate but in 1916 (page 63) he states. "This mite is easily recognized by
the shape of the minute posterior scutum on.the dorsal surface." Hence,
this species is either variable in this respect, or Liponyssoides and Allo-
dermanyssus are truly synonymous.

,, ; Liponyssoides brasiliensis (Fonseca). New combination

.
Dermanyssus brasiliensis Fonseca. 1935: 52 (d); Radford, 1950.
Locality: South America - Brazil, Ceara.
Host: Mammal - Holochilus sciureus.

Genus Der’manyssus Duges, 1-934-

’.
-
" -:"’ ’.’.".
.
’ (Fig. 57) .\ .
... ,
’.
:; ’"
_
Type: Acarus gallinae DeGeer. .’i
Diagnosis: Medium sized mites. Female ohelicerae greatly atten-
uated and terminating in very minute chelae; the sternal plate much
broader than long and strongly arched; epigynial plate broad and rounded
posteriorly; dorsalplate entire and covering half or more of the dorsum.
Male holoventral plate slightly expanded behind.coxae-IV, not divided in-
to two ..separate parts but a transverse line at level of’ posterior margin
of coxae IV is present. Parasitic on birds; especially domestic and
cage birds.
Remarks: The genus is in need of revision. It is doubtful that all
the species listed below really are specific entities.
 121

Dermanyssus gallinae (DeGeer)

(Pig. 57)
Acarus gallinae DeGeer; 1778: 106 (d) (=Pulex gallinae Redi; 1674)’.’
Dermanyssus gallinae, Duges, 1834: 18 (d. t); Berlese, ’1882; Osborn, ’

1896; Hassal, 1896; Huber, 1899; Herrick, 1902; Rainbow.’ 1906;

Banks, 1907;’Ewing, 1909; Ewing. 1911; Ewing,-1913;’Banks,’ 1915;
Herrick, 1915: 233 <b); Hirst, 1915; Barber, 1916; Hirst 1916;
Ewing, 1922; Hirst, 1922; Davidsori, 1924: I (control); Miller, 1925;
Hirst, 1925; Marcovitch, 1926; Sambon,’ 1928; Robertson, 1929;’
Ainslie. 1929; Macfie and Thompson;’1929: 185; ’Gibs’on, 1930;
’

Bishopp and Wood, 1931: 1 (b); Bedford, 1932; Ewing, 1933;’-Pie-’

karski, 1935: 615 (b); Ewing, 1936; Lawrence, 1938; Hoyle, 1938;
Riley and Johannsen. 1938; Fonseca, 1938: 1 (c); Radford, 1939;
Warren, 1949: 409 (b); Bunyear arid. Wehr. 1941; Spencer, 1941;
Smart etal., 1943; Herms, 1944; DeMeillon and Lavoipierre, 1944;
Smith et al., 1944: 362 (m); Vogelsang, 1944; Sulkin, 1945: 381 (m);
Taylor and Murray, 1946; Randolph and Eads,-1946; Wisseman and
Sulkin. 1947: 463 (b); Howitf’ et _al., 1948 (m); Ritcher and Insko,
’’
1948; Roman ^tal., 1950; Zumpt, 1950; Matheson, 1950; McCardle.
1950; Shaw and Pommerening, 1950 (m); Ba&’er, 1950; Sparazarii,
1950; WiUmann,. 19:50; Boyd, 1951; Seddon. 1951;’ Woodroffe and
Southgate, 1951; Jameson et al., 1952; Lamb, 1952; Morlan, 1952;
Woodroffe, 1953; Bregetova, 1953; 1956:’ 203 (b, illus.); Reeves et
al., 1955: 90 (m); Brown. 1953; Baker ert al., 1956: 12; Chamber-
lain and Sikes, 1955: 106 (m); Ke’egan, 1956.
’

’
Dermanyssus avium Puges, 1834; Barnesby, 1871; Lang, 1871; Murray,’.
’

’
’

1876; Rainbow. 1906; Buiteridijk; 1945. "

Localities: Africa - Morocco; Union 61 South Africa.’’ ’Australia -

New South Wales, Queensland. Europe "Croatia"; England; France;
-
Germany; Italy; Netherlands; Scotland; U; S.S.R. Islands of the Pacific -
Guam; New Zealand. North America - Canada; United States. South
America - Brazil; Venezuela. Asia - U.S. S. R. :" .’ :.
;. . .;
Hosts: Mammals - "Domestic animals, " "domestic rat," Homo
sapiens, "horses," "man." Birds - "cage "birds," "caged’chaffinches, "
"canaries, " "chickens;" "ducks/" "eagle owl, " "English owl," "English
sparrow," Gecirius vaillanti. "native song birds, "’"pigeons," "starling,"
"fdwhee, " "turkeys, " "wren house, " Riparia riparia. ’- ’ ’
"
Remarks: The common fowl .mite probably occurs everywhere in’
the temperate and tropical zones where domestic fowl or cage birds are
kept. Although they are recorded as having bitten man and a few other
mammals, they do not do so readily and must be considered obligate
parasites of birds. Wisseman’and’Sulkin (1947) in reporting on the biol-
ogy and feeding habits stated that they could not’induce the’mite to feed’
on white mice, white rats, or man. They may become so abundant in
hen houses and in bird nests ag’to kill the occupants by exsanguiriation.
Smith et al.( 1944), Sulkin (1945), and Howitt et _al. ( 1948’);Tecovered
St.’Louis encephalitis, ’western ’equine encephalitis; and eastern’equine
encephalitis respectively from this mite. ’However, ’after exhaustive’
122 ,,. .. ;; ,
. ..
’
’ .’’ ’

.
laboratory studies/Reeves et al’. (1955), and Chamberlain and Sikes
(1955), concluded that the chicken mite plays a minor role, if any, in
the perpetuation of the encephalitis viruses in nature^. Macfie. and Thomp-
son (1929) found presumptive evidence indicating that the mite is the
intermediate host and vector of a .trypanosome of canaries. -

.
.
Warren (1940) presented a-detailed study of the genital, system of
E>. gallinae. . ,

It is interesting to note that Redi in 1674, used the name gallinae

for this mite. However, DeGeer was apparently the first author to use
the name after 1758. and so he should properly be given, the credit. De-
Geer himself credits Redi with the name. ;

Dermanyssus americanus Ewing

Type .material in the U. S. National Museum.

Dermanyssus americanus Ewing, 1922: 24 (d); Ewing, 1936: 53 (c); Rad-
ford, 1950; Reeves (rtal.. 1955: 90 (m).

.
.
Locality: North America - Canada, Ontario Province; United
States. Maryland, Minnesota, District of Columbia.
Hosts: Birds - "Nuthatch," Passer domesticus.

Dermanyssus brevis Ewing

Type material in the U. S. National Museum.

Dermanyssus brevis Ewing, 1936: 54(d); Radford, 1950.
Locality: North America - United States, Oregon.
Host: Bird - "Horned lark. "

Dermanyssus chelidonis Oudemans

Dermanyssus chelidonis Oudemans, 1939: 303 (t); Radtord, 1950.

Locality: ? Europe.
Host: Bird - "Prom swallow nests. "

Remarks: Oudemans states: "In 1889. Berlese figured a Derma-

nyssus from swallow nests which had a peritreme entirely different from
either of the above two (hirundinus and gallinae). This is now given the
name chelidonis. "

Dermanyssus evotomydis Ewing

Type material in the U. S. National Museum.

Dermanyssus evotomydis Ewing, 1933: 11 (d); Ewing. 1936; Radford,
1950.
Locality: North America United States, New York.
-
Host: Mammal - Evotomys sp.
Remarks: The species is known only from the holotype. It is very
similar to gallinae and the rodent is probably not the normal host.
 123

Dermanyssus hirundinis (Hermann)

Acarus hirundinis Hermann, 1804.

Dermanyssus hirundinis, Murray, 1876; Berlese. 1882; Joyeux, 1915:
656 (c); Hirst, 1920; Sambon,. 1928; Oudemans,.-1-92.9: 111; Oude-
mans. 1936: 329 (#145); Buitendijk, 1945; Radford, 1950; Bregetova,
1956: 203. , .. . .,: .-. .’

.
.
Localities: Europe - England; Netherlands; U.S.S.R. Africa - Up-
per French Guinea. ’. ..

.
. .

,
.Hosts: Birds - Ant bus arboreus, Hi.rando rufula, Hirundo urbica,
"mart.en, " Micropus affinis, Passer montanus..
Remarks: Hirst (1920) suggests this ia,’ at most, .a variety of gal-
linae. It has been, accused of attacking -man,. : The original reference
.

(Hermann, 1804) was not foundby.us; the ..reference was .taken from- Oud-
emans (1929). ..: - ... .. : ,

.
. .

.
,
.
Dermanyssus oti Ewing

Type material in the U. S. National Museum.

Dermanyssus oti Ewing, 1925.:. 2,1 .(d); Ewiag. .1936;.Radford, 1950.
Locality: North America - United States, Indiana.
Host: Bird - Otus asio asio. :. , .... : s . ..
.
Dermany.ssus prognephilus Ewing

Type material in the U. S. National Museum.

Dermanyssus prognephilus Ewing, 1933; 12 (d); Ewing, 1936; Radford,
1950.
Locality;, North Alinerica -.United States, Maryland.
i

Host; Bird - Progne-s,ubis subis.

’
. .

’ , j^P-ermanyssus quintus Vitzthum

.
Dermanyssus quintus Vitzthum,, 1920: 7 (d).; Radford. 19,50; Turk. 1952;
Bregetova,’1956: 202 (Sillus.).
Locality: Europe - England; Germany; U.S.S.R.. "Crimea."
Hosts: Birds - Pendrocopus major, Dryobates leucotos, Dryobates

Dermanyssus scutatus Ewing

Type material in the U. S. National Museum.

Dermanyssus scutatus Ewing, 1936:, 4.9 .(d),;. Radford. 1950.
Locality:’ North America - United States, Oregon.
.
124

Host; Bird - Colaptes cafer cafer.

Subfamily MY ONYSSINAE. New’ subfamily :. .

.
,
Diagnosis: Medium sized mites; dorsal plate entire and covering
most of the dorsum. Holoventral plate of male slightly expanded behind
coxae IV. Epigynial plate of female somewhat expanded and frequently
bearing more than eight setae. Chelae in both sexes long, slender, and
poorly sclerotized, both arms equally developed, edentate, and without
setation. Male chela without spermatbdactyl.
Remarks: Furman and Tipton (1955).’placed Myonyssus lathe subfam-
ily Macronyssinae, thus emphasizing its dermanyssid characteristics .
However, we believe that the unmodified structure of the male ’chela is
basic in nature and therefore propose a’separate subfamily for it,"

Genus Myonyssus Tiraboschi, 1904

(Fig, 58)
Type: Myonyssus decumani Tiraboschi,
Tetragonyssus Ewing, 1922: 6. ’

Type: Liponyssus gigas Oudemans. ’ ’

"
Diagnosis: Chelicerae attenuate iriboth sexes; chelae long and eden-
tate. Male chelae shearlike and without a spermatod’actyl. Pe’male
anal plate very broad, with deeply concave anterior margin.

Myonyssus decumani Tiraboschi

Myonyssus decumani Tiraboschi, 1904: 337 (d); Hirst, 1916; Hirst,

1926; Hora, 1934; Ewing and Baker, 1947: 376 (t); Bregetova, 1949;
Radford, 1950; Asanuma, 1951; Furman and Tipton, 1955; Brege-
tova, 1956: 122 (illus.).
Localities: Europe-England; Italy; Netherlands; Scotland; Shetland
Islands.
Hosts: Mammals - Mus musculus; Rattus norvegicus, "rodent
nests," Apodemus flavicollis, Apodemus sylvaticus,

Myonyssus dubinini Bregetova

Type material in the Zoologic Institute of the Academy of Science,

U.S.S.R.
Myonyssus dubinini Bregetova. 1949: 751 (d); 1953; 1956: 122 (illus.).
Locality: U.S.S.R. - Russia, Maritime Province.
Hosts: Mammals - Apodemus speciosus, Mustela siberica, Cleth-
rionomys rutilus, Microtus fortis. ’’
 125

Myonyssus gigas (Oudemans)

Liponyssus gigas Oudeinans, 1912; Oudemans, 1914: 84 (d).

Myonyssus gigas. Hirst, 1916; Hora, 1934; Ewing and Baker, 1947: 376
.
.(.d); Bregetova,, 1949; Radford, 1950; Asanuma, 1951; WiUmann,
. 1952; Purman and Tipton. 1935; Bregetova;’’i956-
Tetragonyssus gigas, Ewing, 1822: 6. ’.’ " ’

Locality:. Europe - England; ’Netherlands;’ Scotland.

Hosts: Mammals - Apodemus flavicoHis, Apodemus sylvaticus, Tal-
pa europaea.

"
, .

.
.
. .’. Myonyssus ingricus’ Breget’ova

Myonyssus ingrjcus Bregetova, 1958: 121 (d. d’and ?, illus.).

’

Locality: Europe - U.S.S.R., Leningrad. ...,,

.
..

Host: Mammal - Clethrionomys glareolus.

Myonyssus jamesoni Ewing arid Baker

.
.
(Fig. 58)
Type material in the U. S. National Museum.
Myonyssus jamesoni Ewing and Baker; 194’7:"37’?’(d); Bregetova, 1949;
Jameson. 1950: 140 (c); Radford, 1950; Asanuma, 1951; Furman
..and Tipton. .1:955’.. ". . :

Locality: Worth America Canada; United States.

-

Hosts: Mamma.ls. - Blarina brevicauda,’ Sore’x ’fumeus.

MyOHyssus montan\is Furman and Tipton ..

Type material in the U. S. National Museum. , .,,:
Myonyssus montanus Furman and Tipton, 1955: 180 (d).
Locality: North America - Utah, San Juan Co.
’
Host: Mammal - Ochotona prineeps.
’
,. ,
.
Myonyssus rossicus Bregetova . ...
Myonyssus rossicus Bregetova,. 1956: 123 (d, o’and 9, illus.).
Localities; Europe and Asia - U.S.S.R., Voronov region. Northern
’
Ocetia. . .
,
.
.
Host: Mammal - Apodemus sylva’ticu’s.

. Myonyssus shibatai Asanuma

.
.
Myonyssus shibatai Asanuma, 1951: 79 (d); Jameson, et al.,1952; Pur-
man and Tipton, 1955.
Locality: Asia Southeastern Manchuria.
-
Host; Mammal - Microtus pelliceus.
Remarks: The types and some of the original drawings were des-
troyed during an air raid in 1945. . . .
.
126

Family HAEMOGAMASIDAE Oudemans, 1926

Diagnosis of the family. Medium to large mites; dorsum densely

covered with setae. The: epigynial plate of the female may be small and
drop-shaped or as large as to nearly cover the venter, but it is always
separate from the anal plate. :It’may: bear’from’ten to’bver’tifty’.setae.
The sternal and anal plates frequently bear’ "accessory setae. The tec -
turn may be tonguelike with serrated margins.’ ’or truncate and serrate,
or truncate and smooth. ’’ ’ . ; :

’
It is not yet definitely proven that these mites are. obligate parasites,
Perhaps some species, as Eulaelaps stabularis; are not, but others.
such as Haeroogamaaus liponyssoideB certainly seem to be. All species
are commonly found in nests of rodents and other small ve’rteB’rates."
A fine review of this family was written by Keegan (1951), a paper
from which we have borrowed treely;:’We have followed Asanuma (1952)
and placed Euhaemogamasus in synonymy with Haemogamasus. Also,
Eulaelaps and Brevisterna have been added to the family.^’ ’.’

.
Key to the Genera of Haemogamasidae

1. Metapodal plates large and -prominent, triangular in shape. An-

al plate triangular, broader than long, bearing only three setae
. . . . . . . .. . . . .. . . . ;., .
. -. .’-.:.i.^.^:.V: . Eulaelaps
Metapodal plates not triangular, not unusually large.^’Anal plate
oval. frequently with more than three setae’ .". ."’. . . . . . . . 2

2. Sternal plate poorly developed, bearing only sternal setae II.

(Occasionally one or more accessory setae may be’present, but
these can generally be recognized by the lack of sternal pores
associated with them.) . . . . . . . . . . . . . . . . . Brevisterna

Sternal plate normally developed, having at least three pairs of

setae and two pairs of pores . . . . . . . . . . . . . . . . . . . 3

3. Tectum with a truncate, fimbriated margin. Leg II with some

heavy spines and spurs . . . . . . . . . . . . . . . . . Ischyropoda

Tectum elongated, tonguelike, with a fimbriated margin. None

of the setae of leg II noticeably different from those of legs III
and IV . . . . . . . . . . . . . . . . . . . . . . . . Haemogamasus

Genus Eulaelaps Berlese, 1903

(Pig. 59)
Type: Gamasus stabularis Koch.
(Hemilaelaps Hull, 1918 was written in error and is therefore
 127

not considered a valid nomenclatorial entity. Hull ap-

parently had intended to write Haemolaelaps for Eulae-
laps stabularis.), ,S.ee. Turk, 18-15: 141.
Diagnosis: Medium to large mites. Female with large triangular
metapodal plates, a broad, triangular anal plate, and a greatly expanded
epigynial plate. Holowentral plate of male greatly expanded behind cox-
ae IV. Peritremalia in both sexes broad and short, bearing a large
stigmalike pore near the apex; chelae’dentate, well sclerotized.
Remarks: Although these mites are frequently found in association
with mammals and in nests of mammals, it is not yet definitely proven
that they are parasitic.
Most authors have placed Eulaelaps in the family Laelaptidae but
practically all its’m’orphological characters are exceptions to the family
characteristics. Jameson et al. (1952) were apparently the first to place
this genus in ’the. iHaemogamasidae, where it seems to fit very nicely.

;? :
.
Eulaelaps stabularis (Koch)
.
: ;;;.:.(Fig. 59>
.
Gamasus .stabularis Koch, 1836: i3 (d).
HyppaspiS stabularis, G. andR. Canestrini, 1832; Oudemans, 1913: 189.
Laelaps (Eulaelaps) stabularis. Berlese, 1903: 13; Halbert, 1915.
Eulaelaps stabularis. Hirst, 1914; Ewing, 1932; Hora, 1934; Willmann,
1939; Radford. ’1939;. Warren. 1940 (b); Turk, 1945: 135; Randolph
andEads, 1946; Ewing. 1947: 83(d); Jameson, 1947; Pratt and Pritz,
1947 (atlas); Hughes, 1948; 0,’Farrell and Butler, 1948; Schweizer,
1949; Asanuma, 1949;.Jameson, 1950; Zumpt. 1950; Asanuma, 1951;
Jameson et.al.. 1952; Mo.rlan. 1952; Willmann. 1952; 1955: 181;
^Dubinin, 1953;-B.regetova, 1953; 1956: 100 (illus. ); Baker et al.,
1956; Keegan, 1956.
Eulaelaps arcualis Tragardh, 1912: 577.
Localities: Africa-Egypt. Asia - China; Japan; Korea; Manchuria.
Europe - England; Germany; Isle of Lewis; Ireland; Norway; Outer Heb-
rides; Scotland,, Switzerland; North America - Canada, Ontario; United
States.
Hosts: Mammals-AlactagHlus acontion, Apodemus agrarius, Apo-
demus flavicollis, Apodemus geisha. Apodemus speciosus, Apodemus
sylvat-icus, ’ Aryicanthis niloticus, Arvicola amphibius, Arvicola ter-
.
restris, BIarina bre.vi,cauda, ; Gite’llus dauricus, Clethrionomys glareo-
lus,- Clethrionomys rufocanus, Clethrionomys rutilus, "cotton mouse, "
"cotton rat, " Cricetulns barabensis, Cricetus cricetus, Pidelphis sp..
Meriones erythrourus, Meriones meridianus, Meriones persicus, Mer-
iones tristrami, Microtus arvalis, Microtus oeconomus, Microtus greg-
alis, Microtus montebelli, Microtus pellicus, Mus musculus, Mustela
erminea, Myospa-lax dybowskii, Nesokia indica, Pitymys pinetorum,
Rattus norvegicus, Rattus turkestanicus, Sorex araneus, Talpa euro-
paea, Talpa aItaica,-Tscherskia triton. Birds - No specific birds are men-
tioned in the literature but we have, on occasion, found it in pigeon nests.
Remarks: It seems certain, that this species is ovoviviparous; we
128 :; .
.

have seen a few females in which the protonymph is clearly visible.

Eulaelaps ’citellus Radford , .

.
.

Eulaelaps citellus. Radford, 1942: 2’96’(-d).- , ;

Locality: North America - United States, California.
Host: Mammal - Citellus ’beecheyr. -
’

Eulaelaps cricetuli Vitzthum

Eulaelaps cricetuli Vitzthum, 1930: 413 (d); Asanuma, i.951; Jameson

etal., 1952; Bregetova, 1956: 104 (d- and 9, illus.).
Locality: Asia - North China; Manchuria; U,S.S.R.
Hosts: Mammals - Allactaga saltator, Ap.odemus speciosus, Cri-
cetulus barabensis, Dipus sowerbyi, Meriones tamariscinus, Microtus
ungarensis "Mongolian pika," .Ochotona daurica, Phodopus bedfordiae.

Eulaelaps kolpakovae Bregetova ’,. ;.

Eulaelaps kolpakovae .Bregetov’a, 1950; 1956: 102 (d’and.$, illus.).
Localities: .Europe and Asia - U.S. S.B,. ;

.
.

.
Hosts: Mammals - Alactagulus acontion, Allactaga saltator, Ar-
vicola terres.tris, C.itellus pygmaeus, Microtus- h.randti, Mus musculus,
Ochotona daurica, Scirtoppda telum. : ; . .

.
.
Eulaelaps novus Vitzthum. . ... ’.’
Eulaelaps novus Vitzthum," 1924; 1925; 165-’<d); Wiilmann.
1952; Turk,
1952. ’ /: .. .’ .

’Locality; Europe - England; Germany.

. . Host: Bird.- Nest’of Riparia’riparia.’. ;
’.: Remarks: Vitzthum discusses at sortie-length the possible synonymy
of this .species and suggests tha^ it might be any one. of several forms-
described by various authors as a small’form of stabularis. The date
of-the-original description, 1924, is quoted from Vitzthum, 1925: 165.-
N.o reference is given. .’ . ’,
’... .Eulaelaps pribatoides. (Michael),-

Laelaps oribatoides Michael, 1892.’ .

Eulaelaps. oribatoides, Oudemans, 1927; WiUmann. 1952.

-Locality: Europe - Germany, ; ,. .

Host: Mammal - Talpa europaea. .

.
Remarks: Oudemans (1927) suggests .this .might; be a synonym of
Eulaelaps stabularis.
 129

Eulaelaps pedalis (Banks)

Laelaps pedalis Banks, 1909: 136 (d); Banks, 1915; Grant, 1947.
.Eulaelaps pedalis. Radford, 1942.
Locality: North America - Canada, Ontario.
.,.Host: Mammal - "Chipmunk. " .,

Eulaelaps propheticus (Banks)

Laelaps propheticus Banks. 1909: 137 (d); Grant. 1’947.

Eulaelaps propheticus. Radford, 1942; 1950.
Locality: .North America - Canada. Ontario.
Host: "Groundhog. " .
’

Eulaelaps yitzthumi Ponseca

Eulaelaps vitzthumi Fonseca, 1935: 33, 39 (d).

Locality: South America - Brazil, Sao Paulo.
Host: Mammal -"Unidentified wild rat. "

Genus Haemogamasus Berlese. 1889

’
(Fig. 60)
Type: Haemogaroaaus hirsutus Berlese.
Euhaemogamasus Ewing, 1933: 3.
Type: Euhaemogamasus onychomydis Ewing [ = Haemoga-
masus ambulans (Thorell)].
Diagnosis: Female epigynial plate only slightly expanded; anal plate
ovate and mostly with accessory setae; chelae with or without teeth.
Holoventral plate of male fully expanded behind’coxae IV; spermatodactyl
equal to or only slightly longer than the moveable arm of the chela.

Haemogamasus hirsutus Berlese

Haemogamasus hirsutus Berlese. 1889: 117; Tragardh. 1912; Oudemans.
1913; 1914; Hirst, 1914; 1916; Ewing, 1925; Vitzthum. 1931; Ewing,
1933; Radford, 1939; Warren, 1940 (b); Buitendijk, 1945; Keegan.
1951 (c. d); Willmann, 1952; Bregetova, 1956(illus.).
Locality: Europe - England; Germany; Netherlands; Spain; Yugo-
slavia; "Slovenia," U.S.S.R,
Hosts: Mammals - Apodemus flavicollis, Arvicola amphibius, Cleth-
rionomys glareolus, "dead mole." Mus musculus, "mouse nest/’ Rat-
tus norvegicus, "rodents," Talpa alpina. Talpa europaea, "weasel. "
Microtus arvalis, Microtus gud.

Haemogamasus alaskensis Ewing

Haemogamasus alaskensis Ewing, 1925: 13.9 (d); Vitzthum. 1930; Jame-

130

son. 1950; Keegan, 1951: 213 (c, d).

Haemogamasus sternalis Ewing. 1933: 3 <d); Moore, 1949; Furman and
’’’’
’

Tipton, 1955.
Locality: North America - Alaska; Canada; northern United States.
Hosts: Mammals - Blarina brevicauda, Blaring brevicauda tal-
poides, Clethrionomys gapperi, Clethrionomys gapperi ochraceus, "field
mouse," "meadow mouse," Microtus montanus, Microtus oregoni, Mi-
crotus pennsylvanicus pennsylvanicus. Micr.otus sp., Myotis lucifugus
lucifugus, Napaeozapus insignis msignis, Peromyscus maniculatus gra-
cilis, Pitymys pinetorum.’ Sorex ’cinereus, Tamiasciurus hudsonicus
(nest).

Haempgamasuj ambulans (Thorell) :

Dermanyssus arnbulans Thorell, 1872: 164 (d). (Quoted from Keegan.

1951.)
Hypoaspis ambulans, Tragardh, 1904; 1.
Eulaelaps ambulans, Tragardh, 1910: 435 (Quoted fromKeegan. 1951.);
HaarlfSv, 1942.
Euhaemogamasus ambulans, Keegan, 1951: 228 (c, d); Willmann, 1952.
Haemogamasus ambulans, Jameson et al., 1952: 10; Bregetova, 1953;
1956: 147 (cf, .$. illus.); Furman~1957 (b),
Gamasus ovalis Koch, 1878: 121. (Quoted from Keegan, 1951.)
Laelaps ovalis, Tragardh, 1902: 61. (Quoted from Keegan, 1951.)
Haemogamasus nidi Michael. 1892: 314; Hirst, 1914; Hirst, 1916; Vitz-
thum, 1930; Radford, 1939; Buitendijk, 1945; Schweizer, 1949; Du-
binin, 1953; Lange, 1955; Bregetova, 1956: 134 (illus.).
Haemogamasus michaeli Oudemans, 1.904: 87; Oudemans, 1913; Hora,
1934: 363. . ;

Haemogamasus reidi Ewing. 1925; 140; Vitzthum, 1930.

Haemogamasus twitchelli Ewing, 1925: 142; Vitzthum, 1930.
Euhaemogamasus oregonensis Ewing, 1933: 5 (d); Augustson, 1941.
Euhaeinogamasus onychomydis Ewing. 1933: 4 (generotype of Euhaemo-
gamasus Ewing).
Euhaemogamasus sciuropteri Keegan, 1946: 72.
Localities: Asia-Japan; Korea; Siberia; U.S. S.R, Europe - Mid-
dle Europe; Western Europe; England; Germany; Isle of Lewis; Nether-
lands; Outer Hebrides; Scotland; Shetland Islands; Switzerland. Islands
of the Atlantic - Iceland. North America - Alaska; Greenland; United’
States.
Hosts: Mammals - Apodemus agrarius, Apodemus flavicollis, Apo-
demus hebridensis. Apodemus speciosus, Apodemus -sylvaticus (nest),
Aryicola terrestris, Blarina brevicauda, Citellus undulatus, Clethrio-
nomys amurensis mikado, Clethrionomys gapperi ochraceus, Clethrio-
nomys glareolus, Clethrionomys rufocanus, Clethrionomys rutilus. "cot-
tontail rabbit," Cricetulus barabensis, Cricetus cricetus. Evotomys
dawsoni, Eutamias sibiricus, Glaucomys sabrinus macrotis, Glaucomys
sp., Glaucomys volans, Lagurus lagurus, Lemnus lenmus, Lemnus
 131

obensis, Martes zibellina, Microtus agrestis, Microtus arvalis, Mi-

crotus californicus, Microtus gregalis,. Microtus majori, Microtus mon-
tanus, Microtus oeconomus, Microtus operarius operarius, Microtus
socialis, Mus musculus, Must’ela nivalis,. Mycromys minutus, Myotis
daubentonii. Myotis schistocolor, Neomys fodiens, Neotoma cinerea,
Neotoma .fuscipes, Neotoma fuscipes maerotis, Neotoma sp,, Ocbotona
alpina, 0’nychomys sp., Phen’acomys alfeipes, "raccoon," Rattus nor-
vegicus, Sciurus carolinensis, .Sciurus griseus nigripes, Sciurus niger’
rufiventer, Sorex araneus, Sorex palustris, Talpa altaica., Talpa euro-
paea, Tamiasciurus douglaaii, Tamiasciurus hudsonicus, Thomomys
fuscus, Thomoiriys monticola, Thomomys sp., Thomoinya talpoides.
Birds - "Blue jay," "bobwhite," "chicken nest," Circus sp., "red-
shouldered hawk," _R^pana r^paria.

’
Remarks: Bregetova (1956) and other Russian workers consider _H.
nidi and H, ambulang as two’ distinct species. .We have not. examined
type material and are therefore not qualified to judge. Keegan’s synon-
ymy is followed arbitrarily.
’

’
.

Furman (1957) has reported that H. ambulans will feed on a variety

of substances other than host Blood. ..;

’
’
..

.
Hafemogamasus avisugus Vitzthum

Haemogamasus avisugus Vitzthum, 1930: 396 (d); Keegan, 1951; Will-

mann, 1952. .
.
Locality: Europe - Germany. ; .
.
Host: Bird - Riparia riparia.

Haemogamasus ’barberi Bwing .

Type material in the U.’ S. National Mus’eum;

Haemogamasus barberi Swing, 1925: 140 (d); Vitzthum, 1.930.
Euhaemogamasus barberi, Keegan. 1951: -249 (c, d, t); Burgess, 1955.
Haemogamasus microti ’Ewing, 1925: 14’l’(d); Vitztfaum..l930.
’

Euhaemogamasus rnicroti; Jamesdn,. 1950: 139. .

.
Locality: North America - Canada (southeastern area); United
States, Maryland and New York. : ’
Hosts: Mammals - "Albermarle meadow mouse," Blarina brevi-
cauda, Citellus richardsorii, Microius pennsylvanicus, "mouse nest. "
"pine mouse," Pitymys pinetorum, "short-tailed s’hr’ew. "

Haentogamasus citelli Bregetova and.N’elzina .

Haemogamasus citelli Bregetova and Nelzina, 1952; Bregetova, 1956:

148 (rf and ?,illus.).
Europe and Asia U.S. S. R., Armenia.
Localities;
-
Hosts: Mammals - Alactagulus accintion, Citellus pygmaeus, Ci-
tellus suslicus, Citellua xanthoprymnus.
 132

Hae mogamasus daurjcus Bregetova

Haemogamasus dauric-us .Bregetova, :.:i950: 318; 1956: 133 ($, illus.).

Locality: Asia -^Mongolia; U.S.’S.R.
Hosts: Mammals.- Apodemus agrarius, Apodemus speciosus. Cleth-
rionomys rufocanus.-Crrcetus tritbn, Marmota sibirica, Myospalax dy-
bowskii. Och-otona alpina, ]j,a.ttus norvegicus. ’. ,. .

Haemogamasus ellipsoi.deus Asanurna

Haertiogamasus) ellipsoideus Asanuma-; 1952: 89. (4).’.

Locality: Asia - Manchuria. -/.
Hosts: Mammals - Apodemus agrarius, Cletbrionomys rutilus,
Crioetulus barabensis,. Microtus pellicus. . ,

Remarks: Asanuma states that he described’this species as new

with’some’hesitation; it may. .fee the same as alaskensis- Ewing.

Haemogamasus harperi-l^eegan

Type material in the U. S. National Museum. i

Haemogamasus harperi Keegan. 19.51: 223 (d).

Locality: North America-United-States, Florida, Georgia, Missis-
sippi, and South’Carolina.
Hosts: Mammals Cryptotis parva, Scalopus aquaticus australis,
-
Scalopus aquaticus howelli.

Haemogamasus hirsutosimilis Willmann

Haemogamasus hirsutosimilis Willmann, ’1952: 392 (d); Bregetova, 1956

(d’ and $, illus.);
Locality: -Europe -Germany; U.S.S.-R. ’
’
Hosts: -’Mammals -Apodemus agrarius, Apodemus .flavicollis, Apo-
demus sylvaticus, Clethrionomys glareolus, Talpa europaea. ’;

.Haemogamasus horridus. Michael

Ha-embgamasus horridus Michael, 1892: 312 (d); Oudemans, 1913:. 146

(d); Hirst, 1916; Vitzthum, 1930. . ...
Euhaemogamasus horridus,: Keegan, 1951: 235 (c, d, t);...Willmann, 1952.
Haemogamasus horridus var. arvicolarum Bertese, 1921: ,166 (d); Vitz-
thum, 1930:. ...
.
Haemogamasus arvicolarum, Turk, 1945 (c, d); Turfc, 1952; Willmann,
1955: 181. :,. .. ;,
.
Locality: Europe -England; Gertaany; ..Ireland; Island of Lewis;
Netherlands;. Italy; Outer Hebrides;. Sardinia;. Shetland Islands; Western
’

Europe. .;; .,.; ’

Hosts; Mammals - Apodemus flavicollis, Apodemus sylvaticus, Ar-
yicola arvalis, Clethrionomys glareolus, "field mouse," "mouse nest, "
 133

Mus hebridensis, Mus sylvaticus, "nest of" Arvicola arvalis, "nest of"
Talpa europaea, Sorex araneus, Rattus rattus frugivorus, Talpa alpina,
Talpa europaea.

Haemogamasus ivanovi Bregetova

Haemogamasus ivanovi’ Bregetova, 1955; 1956: 143 (cf and $, illus.).

Localities: Europe and Asia - U. S.S.R.,. Tadzhikistan. Grissar-
ski. ,
Host: Mammal - Microtus carruthersi.

Haeinogamasus japonicus Asanunaa

Haemogamasus japonicus Asanuma, 1952: 4.3 (d); Suye.moto et al., 1954.

Locality: Asia - Japan.
Hosts; Mammals - Apodemus speciosus, Microtus montebelli, Rat-

tus norvegicus.

Haemogamasus keegani (Jameson)

.,’ (Fig. 60)
Type material in the U. S. National Museum. ,

Euhaemogamasus keegani Jameson, 1952: 600 (d)..

Locality: North America - United States, California. .:... ,

Host: Mammal - Sorex trowbridgii.

Haemogan’iasus kitanoi Asanuma

Haemogamasus kitanoi Asanuma, 1948: .171 (d); Asanuma. 195.1; Kee-

.

gan, 1951; Jameson et al., 1952: 10; Bregetova, 1956: 148 (?, illus.).
Haemogamasus polychaeta Bregetova, 1949. ; .

Locality: Asia - Manchuria, U. S.S.R.

Hosts: Mammals - Marmota bobac sibirica, Ochotona alpina, Ocho-
tbna daurica. .
.

Remarks: Jameson et al. suggest this may be a synonym of alas-

kensis Ewlng.

Haemogamasus kusumotoi Asanuma

Type material in the collection of K. Asanuma.

Haemogamasus kusumotoi Asanuma, 1951: 18 (d); Jarneson et al., 1952;
Bregetova, 1956: 142,
’’ Locality: Asia - Manchuria, V.S.S.R.
-
. .
^
.
,
Hosts: Mammals - Apodemus agrarius, Citellus dauncus ramosus,
Cricetulus barabensis, Mus molossimus, Rattus norvegicus, Tsherskia
triton.
 134

Hae mogamasus’ liberiens.is-Hirst

’Haemogamasus liberiensis Hirst, 1916: TO ’(d); Vitzthum, 1930;..Keegan,

1951; 1956.
Locality; Africa Egypt; Liberia; Kenya.

-
Hosts: Mammals Arvicanthis sp., Chaerophon pumilis, Mus tri-

-
; . Haemogamasus liponyssoides Ewing

Type material in the U. S. National Museum..

Haemogamasus liponyssoldes Ewing, 1925: 139 (d); Vitzthum, 1930;
Augustson, 1941: 157; Bregetova, 1956: 140 (rf and $ Ulus.).
Euhaemogamasus liponyssoides. Spencer,. 1941: 14; Jameson, 1950: 139;
Keegan, 1951: 240 (c, d); Morlan, 1952; Bregetova, 1953.
.
.
Localities: Europe and Asia -.U.S.S.R.; Korea. North America -
’

Canada (eastern part); United States (eastern and southern area). Iowa.
Hosts: Mammals - Apodem,us agrarius, Apodemus speciosus, Ar-
vicola terrestris, Blarina brevicauda, Blarina brevicauda aloga, Bla-
rina brevicauda compacta, Blarina brevicauda talpoides, Clethrionomys
gapperi, Clethrionomys rufocanus, Condylura cristata, Didelphys vir-
giniana, Didelphis virginiana pigra, Microtus montanus, Microtus mor-
dax, Microtus ochrogaster (nest), Microtus pennsylvanicus, Parasca-
lops breweri, Peromyscus gossypinus, Peromyscus leucopus novebora-
censis, Peromyscus nest. Pitymys pinetorum,. Rattus norvegicus, Sca-
lopus aquaticus howelli, Scalopus arge.ntatus. (type host), Sigmodon his-
pidus, Sorex cinereus, Sorex fume.us, Sorex palustris, Sorex palustris
albibarbis, Talpa altaica. Talpa europaea.

Haemogamasus mandschuricus Vitzthum

,
Haemogamasus mandschuricus Vitzthum, 1930: 397 (d); Asanuma. 1948:
29; Asanuma, 1948: 171; Keegan, 1951: 218 (c, d); Asanuma. 1951;
Jameson et al., 1952; Purman and Tipton, 1955; Bregetova, 1956:
146(illus.).
Locality: Asia - Japan; North China; Manchuria; U.S.S.R.
, ,, .Hosts: Mammals - Apodemus agrarius, Apodemus geisha, Apo-
demus speciosus, Apodemus sylvaticus, Clethriononqys mikado, Cleth-
rionomys rufocanus, Crioetulus triton, Dipus sowerbyi, Eutamias si-
biricus, Marmota sibirica, Microtus brandti, Microtus fortis, Microtus
gregalis, Ochotona daurica. Phodopus bedfordiae, Scaptochirus gillie si,
Talpa altaica.

Haemogamasus nidiformes Bregetova

Haemogamasus mdiformes Bregetova, 1955; 1956: 136(d, cfand 9, illus.).

Localities: Europe and Asia - U.S.S.R.
Host: Mammal Microtus gregalis.
-
 135

Haemogamasus occidejrtalis (Keegan)

Type material in’the U. S. National Museum. ..

.
Euhaemogamasus liponyssoides dccidentalis Keegan,
"
1951: 255 (d);
Jameson et al., 1952. ’’

’
. .

Localities: Asia - Manchuria; North America - Canada (south-

western area); United States (northwestern area).
Hosts: Mammals - Blarina brevicauda talpoides, Microtus town-
sendii, Mustela saturata, Microtus townsendji.faest), Neurotrichus sp.,
Scapanus orarius, Scapanus orarius schefferi, Scapanus towns endii,
Sorex trowbridgii, Thomomys fuscus. L . .

.
Haemogamasus pontiger (Berlese)

Laelaps (Eulaelaps) pontiger Berlese, 1903 (1904); 260 (d).

Eulaelaps pontiger, Radford, 1950.
Haemogamasus pontiger, BaRer etal., 1956 (c); Bregetova, 1956(illus.).
Haemogamasus oudemansi Hirst, 1941: 122 (d); 1916; 1922; 1926; Sam-
bon. 1928; Vit^thvim,-’-19’30; Radford, 1939; Buitendijk, 1945; Hill
and Gordon, 1945: 46 (m);0’Farrell and Butler, 1948; Zumpt, 1950;
Jameson et al., ’1952.’ ’
’ ..’ .

Euhaemogamasus oudemansi, Hu’ghes; 1948; Keegan, 1951: 240 (c, d,

"

t); Willmann; 1952.’

Haemogamasus (Euhaemogamasus) oudemansi, Womersley, 1956: 540,
Eulaelaps mawsorii Womersley, 1937: 19 (d).
Localities; Africa - Union of South Africa. Asia - China; U.S.S.R.
Australia-Melbourne. Europe-England; Germany; Italy. North Amer-
ica - United States, Colorado, New York.
Hosts: Mammals - "laboratory mice and rats," Rattus norvegicus,
Talpa europaea, Tamlasciurus fremonti (nest). (Also frequently found
in store rooms, stables; warehouses, straw, packing material, etc.)
Remarks: Hill and Gordon (1945) report that American soldiers
stationed in North Wales suffered from dermatitis apparently caused by
this and other mites infesting the straw stuffed paliases on which they
were sleeping. Keegan (1951) is of the opinion that this species is only
a facultative parasite.

Haemogatnasus quadrisetatus -Vitzthum

Haemogamasus quadrisetatus Vitzthum, 1926: 52 (d); 1930; Jameson et

al., 1952; Asanuma et al., 1952.
Euhaemogamasus quadrisetatus, Keegan, 1951: 253 (c, d, t).
Locality: Asia - Japan. Indonesia - Java.
Hosts: Mammals Apodemus speciosus, Mus lepturus, Urotrichus
-
talpoides hondonis.
136

Haemogamasus serdjukovae (Bregetova)

Euhaemogamasus serdjukovae Bregetova, 1949; 1953: 323.

Haemoganaasus serdjukovae, Bregetova, 1955: 138 (c? and 9, lllus.).
Haemogamasus mandschuricus sylvaticus Bregetova, 1949; 1953: 318.
Localities: Europe and Asia - U>S.S.R.
Hosts: Mammals - Apodemus agrarius, Apodemus speciosus, Cleth-
rionomys rufbcanus, Eutamias sibiricus, Microtus fortis, Ochotona al-
pma, Rattus norvegicus, Rattus norvegicus caraco, Sicista caudata.

Haemogamasus zachvatkini Bregetova

Haemogamasus zachvatkini Bregetova, 1955; 1956: 136 (tf and 5, illus.).
Localities: Europe and Asia - U. S.S.R., Georgia.
Host: Mammal - Prometheomys schaposchnikovi.

Genus Brevisterna Keegan, 1,949

(Pig. 61) ,:

.
.

,
.
Type: Euhaemogamasus utahensis Ewmg.
Sternolaelaps Zumpt and Patterson, 1951: 63. ;.
Type: Euhaemogamasus utahensis Swing.
Diagnosis: Female sternal plate reduced, bearing only the middle
pair of sternal setae, although accessory setae may be present, Dor-
s’al plate undivided or a small pygidial plate may be present in the fe-,
male; fixed arm of chela in both sexes transversely striated and.bear-
ing a.piluj dentilis. .’

Brevisterna utahensis (Ewing)

Type material in the U. S. National Museum. :

Euhaemogamasus utahensis Ewing, 1933: 4 (d).
Brevisterna utahensis. Keegan, 1949: 222 (d, t); 1953; Strandtmann and
Allred, ’1956: 116 <.d, all stages illus.); Allred, 1957a: 51; 1957b:
72 (b).
.
Sternolaelaps utahensis. Zumpt and Patterson, 1951: 63.
Locality: North America - United States, Arizona. California, Utah.
Hosts: Mammals - Neotoma albigula, Neotoma fuscipes. Neotoma
streatori, Neotoma lepida (type host), Onychomys leucogaster, Peromys-
cus crinitus, Peromyscus eremicus, Peromyscus maniculatus.
Remarks: The life cycle includes the egg, non-feeding larva, two
feeding nymphal instars, and adult male and female. Species of N.eotoma
are by far the preferred hosts.

Brevisterna montanas (Ewing)

Type material in the U. S. National Museum.

 137

Liponyssus montanus Swing, 1922: 21 (d).

Brevisterna montanus, Strandtmann and.Allred, 1956: 131 (9, illus.).
Locality: North America - United States, Montana.
Host:.’ Mammal - Sciurus hudsonicus richardsoni. ..
.

Brevisterna morlani Strandtmann and .Allred

(Fig. 61)

,
,
Type material in the U. S.
National Museum.
Brevisterna morlani Strandtmann and Allred, 1956: 123 (d. rf and ?, il-
lus.>.
Locality: North America. - United States, Colorado, New Mexico.
Hosts: Mammals - Cynomys gunnesqni, Neotoma albigula (type
host), Neotoma micropus, Peromyscus leucopus, Peromyscus manicu-
latus, Peromyscus truei, Sylvilagus auduboni.
Remarks: Neotoma is the host of choice of.this mite. It is, so sel-
dom found on other hosts that its occurrence on them is probably acci-
dental. .

Genus Ischyropoda Keegan, 1951

(Fig. 62)
Type: Ischyropoda spiniger Keegan.
Diagnosis: Sternal shield with accessory setae; leg II, perhaps
others, conspicuously stout, with spurs and massive, often blunt, setae.
especially on tarsus. Anal shield of male separate. Accessory setae
present on ventral shield of nymph. (After Keegan, 1951.)

Ischyropoda spiniger Keegan

Type material in the U. S. National Museum..

Ischyropoda spiniger Keegan, 1951: 258 (d).
Locality: North America - United States, California.
Hosts: Mammals - Perognathus penicillatus angustirostris (type
host), Perognathus spinatus, Perognathus sp.

Ischyropoda armatus Keegan

(Fig. 62)
Type material in the U. S. National Museum.
Ischyropoda armatus Keegan. 1951: 261 (d); Burgess, 1955: 639.
Locality: North America - United States, Arizona, California, Col-
orado, New Mexico; Canada, Saskatchewan.
Hosts: Mammals - Citellus burrow, Citellus richardsoni, Pipo-
domys deserti deserti, Dipodomys merriami merriami, Dipodomys mer-
riami simiolus, Neotoma albigula, Onychomys leucogaster articeps,
Perognathus californicus, Perognathus inornatus, Perognathus sp . ,
Perognathus xanthonotus, Peromyscus maniculatus, Peromyscus mani
culatus rufinus, Thomomya bottae (type host). Thomomys sp,
 138

Ischyropoda furmani Keegan

Type material in the U. S. .National.Museum. ’ ; . .

Ischyropoda furmani Keegan, 1956: 311.(d...cf, ? and nymph, illus.).

Locality: -North America United, States, Utah.

-
Hosts: Mammals’- Dip odoiays ordii, Microdipodops megacephalus,
Microdipodops pallidus (type host), Onychomys leucogaster, Perognathus
longimembris, Peromyscus mani.culat.us.-: .

Family IXODORHYNCHIDAE Ewing, 1923

. .Description jdt the family. Robust mites having the ’general facies
of Laelaps. -Corniculi--elongate, barbed, recurved; or harpoon^shaped
at the tips. (’This is nof-easily seen unless-the gn’athosoma is mounted
separately on its side.) Chelicerae may be chelate (Hemilaelaps),.or
’

fixed arm may tie lacking (Ixodorhynchus). Coxae I and 11, or coxae I,
II, and Iir.ea.ch with one or two heavy, blunt spurs ventraUy. Epigyniai
plate rounded posteriorly and bearing one pair of setae. Dorsal plate
covering most of the dorsum, nearly always partially divided. External
parasites of snakes. : ’

.
.
.
,
Key to the Genera of Ixodorhynchidae’
Chelicera with only one arm, strongly developed and bearing
three or more heavy, recurved teeth , . . . . . . Ixodorhynchus >

Chela shearlike . . . . ^... . . .. . . :. . ’. . . ’), . . . . Hemilaelaps

Genus Ixodorhynchus .Ewing, l-@22

’
.
(Fig. 66) ’ .’
-
.
. .

Type: Ixodorhynchus liponyssoides Ewing.’’ .. .

.
.
Ixobl’oides Fonseca, 1934. ".,:’
’
.
.
.
Type: Ixobioides butantanensis Fonseca.
Diagnosis: Medium size mites. Female chelai. lacking fixed arm;
corniculi large and harpoon-shaped distally. Male chela shearlike.. _
Remarks: Vitzthum (1941: 774) apparently, was’the first to synony-
mize Ixobioides. "’
’
...

Ixodorhynchas liponyssoides Ewing

(Fig. .66) :
Type materlarin the U. S. National Museum.
Ixodorhynchua lipohyssotdes Ewing. 1922: 9 (d); Vitzthum. 1941: 774 (t).
Locality: North America - United States, Iowa.
Host: Reptile - "Snake." .
.
 139

Ixodorhynchus butantanensis (Fonseca)

Ixobioides butantanensis Fonseca. 1934: 508 (d).

Ixodorhynchus butantanensis. Baker and Wharton, 1952: 60.
Ixobioides butantanensis, Vitzthum. 1941: 774.
Locality: South America - Brazil.
Host: Reptile - Ophis merremi.

Genus Henailaelaps Ewing, 1933

(Fig. 64)
Type: Hemilaelaps americanus Ewing.
Ellsworthia Turk, 1945: 141 (Name proposed for Hemilaelaps
Ewing.)
Ophidilaelaps Radford, 1947: 239. New synonymy.
Type: Ophidilaelaps imphalensia Radford.
Diagnosis: Medium sized mites; dorsal plate entire to very deeply
indented laterally; chelae shear like.
Remarks: Turk (1945) proposed the generic name Ellsworthia be-
cause: "The generic name Hemilaelaps Ewing, 1933 had already been.
used (in error) by Hull in 1918 (Trans. Nat. Hist. Soc. North Durh. and
Newcastle, 5, new series, 1918), for the Eulaelaps group of species
which by mistake he referred to the genus Haemolaelaps Hirst and which
he wrote in error Hemilaelaps." This then is a subsequent erroneus
spelling, and therefore, has no nomenclatorial standing. Henailaelaps
Ewing is not invalidated and Ellsworthia Turk is a junior synonym. ,
That Ophidilaelaps is a synonym of Hemilaelaps Ewing is apparent
from a comparison of their type species.

Hemilaelaps americanus Ewing

Type material in the U. S. National Museum.

Hemilaelaps americanus Ewing, 1933: 8 (d).
Ellsworthia .americanus, Turk, 1945: 141 (t).
Locality:’ North America - United States, Texas.
Host: Reptile - Drymarphon corais melanurus.

’’’,’
’

’"’,’,, ’i Hemilaelaps distinctus Ewing

Type matefi/al in the .V. ’S.’National Museum.

Hemilaelaps disUnctus Ewjng. 1933: 8 (d),.
Locality: North America ’-l United States, Kentucky.
Host: Reptile - Elaphe obsoleta.

Hemilaelaps farrieri Tibbetts, New combination

Type matettal in the U. S. National Museum,

Ophidila’elapS farrieri Tibbetts, 1954: 70 (d, $. illus.).
140 .: ’.;,
Locality; A?ia Koreili
Host; Reptile - EJaphe di-orie.

Hemilaelaps Imphalensis (Hadford). New combination

Ophidilaelaps imphalensis Radford. 1947: 239 (d).

Locality: Asia - India, Manipur State, Imphal.
Host: Reptile - Coluber radiatus.

Hemilaelaps piger (Berlese). New combination

Laelaps piger Berlese, 1918: 70 (d).

Ophidilaelaps piger, Radford, 1947: 239 (t).
Locality: Europe - Italy.
Host: ReptUe - "Snakes. "
Hemilaelaps tanneri Tibbetts. New combination
(Fig. 64)
-Type material in the U. S. National Museum.
Ophidilaelaps tanneri Tibbetts, 1954: 67 (d. d-and ?, illus.).
Locality: Asia -Korea.
Host: Reptile - N.atrix tigrina lateralis.

Hemilaelaps’ triangulus (Ewing). New combination

Type material in the U. S. National Museum.

Liponyssus triangulus Ewing, 1922: 18 (d); Turk. 1947; Ponseca. 1948:
321.
Locality: North America - United States, Maryland.
Host: Reptile - Lampropeltis calligaster, a king snake.
Remarks: The original description was non-diagnostic and the true
generic affinity of the mite remained unknown until the types were ex-
amined recently by D. Johnston of the University of Maryland.

Family RAILLIETIDAE Vitzthum, 1941

Description of the family. With a single dorsal plate, restricted to

podosoma and about half as wide as the body. Peritremes extending
beyond coxa II. Tritosternum present with lacinae that are adorned with
short, sparse setae. Female sternal plate about twice as long as wide,
with three pairs of setae and two pairs otpores. Metasternal setae pres-
ent, but metasternal pores lacking; female epigynial plate reduced. Male
with a short sternigenital plate not extending beyond coxae IV; chela with
a greatly modified fused sperroatodactyl and moveable arm; immoveable
arm of chelicera degenerate. (The shape of the male chela is reminis-
cent of the horns of the American pronghorn antelope.) Femur II of male
 141

with ventral apophysis. Deutosternal groove present with six or seven

transverse rows pf denticles. Palpal claw with two tines.
This mite is found only in the auditory meatus of cattle and antelope
(and perhaps other ungulates as well). Although essentially an internal
parasite, about the only concession made to their special habitat is les-
sened sclerotization of the idiosoma. The leg sclerotization is not re-
duced, and is similar to that of Haemolaelaps. One genus, Raillietia.
Remarks: The presence of multiple rows ofdeutosternal teeth makes
it impossible to include Raillietia in the family Dermanyssldae, as has
recently been done by Evans (1955) and Baker et al. (1958).

Genus Raillietia Trouessart, 1902

(Fig. 65)
Type: Gamasus auris Leidy.
Diagnosis: With the characters of the family.

Raillietia auris (Leidy)

(Fig. 65)
Gamasus auris Leidy, 1872: 138; Osborn, 1896.
Raillietia auris, Trouessart, 1902: 231 (d, t); Trouessart. 1902: 806 and
1335; Preund, 1910; Banks, 1915; Radford, 1938; Olsen and Brack-
en. 1950: #8 (c. d, v); Bregetova. 1956: 128 (illus.).
Localities: Europe. North America - United States.
Host: Domestic cattle. Bos taurus.
Remarks: The most recent review of this species is that by Olsen
and Bracken (1950), who described symptoms apparently produced by
this mite. The mite is found only in the auditory apparatus of cattle
where it inhabits not only the outer and middle ear but may also pene-
trate into the inner ear. It is not known what percentage of cattle are
infected. It will probably be found everywhere that cattle are kept.

Raillietia hopkinsi Radford

Raillietia hopkinsi Radford, 1938: 428 (d); Olsen and Bracken. 1950.
Locality: Africa - Uganda.
Host: Mammal - Kobus defassa ugandae, a waterbuck.

Family SPELAEORHYNCHIDAE Oudemans. 1902

Description of the family. Large, ticklike mites. Gnathosoma pe-

culiar, much wider than long, with widely diverging palps. Chelae short
but very heavy; both arms well developed, each possessing two promi-
nent recurved teeth. Peritreme reduced to a short, rectangular plate
surrounding stigma. Body plates reduced to podosomal plate dorsally
and sternal plate ventrally.
142

Although this family is believed to parasitize only Central and South

American bats, the U.. S. National Museum has’a specimen taken from
a Venezuelan opossum. One genus, Spelaeorhynchus.

’
’Genus Spelaeorhynchus Neumann, 1902

’
(Fig. 63)
Type: Spelaeorhynchus praecursor Neumann.
’Diagnosis: Large, broad, .ticklike mites.

.
Spelaeorhynchus praecursor Neumann
(Pig. 63)
Spelaeorhynchus praecursor Neumann, 1902: 31 (d).; Banks, 1917.
Localities: Central America. South America. West Indies - Ja-
maica.
Hosts: Mammals - "Bats." ’

Spelaeorhynchus latug Banks

Spelaeorhynchus latus Banks. 1917: 197 (d); Radford, 1950..

Locality: Central America - Canal Zone. Obispci; ""
Host: Mammal - Bat.

Family DASYPONYSSIDAE Ponseca, 1940

’

Description of the family. Legs I very. stout, with strong; promi-

nent claws; legs II, III, and IV slender and with small claws. Forked
palpal claw apparently lacking. Tritosternum present. Dorsal plate
entire or divided, covering most of.the dorsum. Peritreme long. reach-
ing to-the middle of coxa I. Sternal plate of female reduced and weakly
sclerotized. Anal plate of male well separated from the short sterni-
ventral plate. ’’

These are large mites that parasitize pangolins and South American
armadillos. At least one species, Manitherionyssus heteroiarsus, is
known to run sideways when disturbed..
The genus Mariitherionyssus is placed in the same family as Dasy-
ponyssus on the basis of similar fore legs, similar reduction of ventral
plates and similarity of hosts.

Key to the Genera of Dasyppnyssidae

Dorsal plate divided. Female with highly modified, spatulate

setae. Male with short, slender setae. Parasitic on South
American armadillos . . , . . , . , . . < . ... .. . . Dasyponyssus
 ’

143
’

., .,

Dorsal plate entire. Posterior setae of both sexes, more than

half as long as the body.. Parasitic on the African pangolin
.
(scaly anteat’er) . . . . . .. . . . . . . . . . . . Manitherionyssus

Genus Dasyponyssus Fonseca, 1940

(Fig. 68)
Type: Dasyponyssus neivai Fonseca.
Diagnosis:- Medium sized mites in which the fore legs are slightly
longer and two or three times as broad as the other legs.

Dasyponyssus neivai Ponseca

(Pig. 68)
Dasyponyssus neivai Fonseca, 1940: 105 (d).
Locality: South America - Brazil, State of Parana.
Host: Mammal - Euphractus sexcinctus.

Genus Manithe rionys sus Vitzthum, 1925

(Fig. 67)
Type: Liponyssus heterotarsus Vitzthum.
Manisicola Lawrence, 1939; 451.
Type: Manisicola africana Lawrence.
Diagnosis: Large mites, nearly two millimeters long; fore legs
shorter but stouter than the others.

Manitherionyssus heterotarsus (Vitzthum)

.(Fig. -.67)
Liponyssus heterotarsus Vi’tzthum, 1925: 7,
Manitherionyssus heterotarsus, Vitzthum, 1925: 12; Kadford, 1950.
Manisicola africana Lawrence, .1939: 451 (d). New synonymy.
Manitherionyssus africana, Radford, 1950: 371. ,
Locality: Africa - Union of South Africa, Transvaal, Praetoria;
South West Africa. Grootfontein.
Host: Mammal - Manis temminckii.
Remarks: Vitzthum(1925) described this mite under the name Lip-
onyssus on page 7, but on page 12 he stated that should the. mite prove
to be specific for pangolins, it would.be. an indication of sufficient dif-
ference to warrant a new genus and in such event, suggested that the
name Manitherionyssus be used.

Family ENTONYSSIDAE Ewing. 1922

Description of the family. Long-legged, poorly sclerotized mites.

Dorsal plate undivided; tritosternum lacking; forked palpal claw present;
stigma with a very short peritreme (Pneumophionyssus). or peritreme
144

entirely lacking. Pulvillus of leg I frequently with two "eye spots." In-
’

ternal parasites of snakes. , ! "

.

.
Remarks:. Records of mites of this family taken from snakes in
zoos shouldbe considered in the light of the possibility that they. may be
transferred among snakes in captivity (as happens so frequently with
Ophionyssus natricis).

Key to the Genera of Entonyssidae

1. Chela withboth arms developed, the fixed arm harpoonlike and

the moveable arm falciform, or’both arms falciform . . . . . . 2

Fixed arm of chela aborted, moveable ^arm poorly sclerotized,

long and strongly recurved . . . . . . . . . . . Pneumophionyasus

2. Tarsus I without pulvillus; chela’of female bifid at apex; dor-

sal plate extending only slightly beyond coxa IV . . . . Hamertonia

Tarsus I with pulvillus; dorsal shield extending well beyond

coxa IV . . . . . . . . . . . . . . . . . . ’;’. . . . . . . . . .;. . 3

3. Fixed arm of chela harpoon-shaped, with a/recurved distal hook,

moveable arm falciform" and generally with a basal fringe of
setae, or setalike structures . ’,’’ , . . . . .; . . . .’. . Entonyssus
.
Fixed arm of chela not harpoon-shaped, moveable arm nearly
straight . . . . . . . . . . . . . . . . . . . . . . Ophiopneumicola

Genus Entonyssus Ewing, 1922

/. (Fig. 69)
Type: Entonyssus halli Ewing. ’;’...
.
. .
’
.

Diagnosis: Medium in size, legs slender. Parasitic in the air sacs

’
of snakes. ’
.
Remarks: Although many species have been described in-this genus,
very few have been illustrated.
.
.
Entonyssus halli Ewing ’.

Entonyssus halli Ewing, 1922: 9 (d); Schmidt, 1940; Keegan, 1943; Turk,
1947; Radford, 1953. ;. " ’,

Localities: North America-District of Columbia, National Zoolog-

ical Park. Europe - England, London Zoological Gardens.
Host: Reptile - Pituophis melanoleucus.
-
Entonyssus ewingi Hubbard

Type material in the U. S., National Museum.

.
.
 145

Entonyssus -ewingi Hubbard, 1939: 657 (d); Hubbard, 1940; Schmidt, 1’940;
’

Keegan, 1943; Turk, 1947. ’:’^’ ".’ "’ ’

Locality: North America - United States, Oklahoma.

’

-
Host: Reptile Crotal.us cinereous. ", .

Entonyssus fragilis Keegan

Type material in the U. S. National Museum.

’’

’
Entonyssus fragilis Keegan. 1946: 73 (d). ’; ’’
"

Locality: North America - United States, Florida. .

"
’
Host: Reptile - Lampropeltis getulus. . .

.
.
Entonyssus glasmacherj.. Vitzthum
(Fig. 69) ..^"’A..’""
Entonyasus glasmacheri Vitzthum, 1935:.710(d, cfand?, illus.); Schmidt,
’

’
1940; Keegan, 1943; Turk. 1947.

.
Locality: Europe - Germany, Berlin Zoo. ’.
’
’’’
’’’
Host: Reptile - Elaphe quadrivittata.. .

Entonyssus heterodontus Keegan ,

Type material in the U. S.. National Museum.

Entonyssus heterodontus Keegan, 1943: 128 (d); Turk, 1947. .’ .: ’
Locality: North America - United States, Iowa.
’ ’

Host: Reptile - Heterodon contortrix. ,

Entonyssus. rileyi Ewing

Type material in the U. S. National Museum.

Entonysaus r.ileyi Ewing, 19,24: 179 (d); Schmidt, 1’940; Keegan,’1943;
;’ . :
’
’
Turk, 1947. :
.
.
.
Locality: North America - United States, Texas’.
Host: Reptile - "Rattlesnake. "

Entonyasus vitzthumi Schmidt

Type material in the U.. S. National Museum.

Entonyssus vitzthumi Schmidt, 194Q: 319 (d); Keegan, 1943; Turk, 1947.
Locality: North America - United States, Oklahoma.
Host: Reptile - Pituophis sayi.

Genus Ophiopneumicola Hubbard, .1938

Type: Ophiopneumicola colubri Hubbard.

’
.’

Diagnosis: Medium sized mites similar to Entonyssus, except the

chelae are not recurved at the apex. All are parasites in the lungs and
air passages of snakes.
146

Ophiopneumicola colubri Hubbard

Type material in the U. S. National Museum. ;

Ophiopneumicola colubriHubbard, 1938: 400(d); Schmidt, 1940; Keegan,

’
’

1943; Turk, 19-47. ’:’ -

Locality: ? North America. ;:; ’’

’’
’

Host: Reptile - Coluber flagellum flavigularis.

Ophji.opneumicola americana Turk

Type material in the coUecti.on of E; A. Turk, England..
Ophiopneumicola americana Turk, 1947:’21.
Locality: ? North America.;.,;,,
Host: Reptile -Elaphe guttata. ’. :’. .’
Ophiopneuinicola- elaphes.. Ke e.gan

Type material in.the U,.’.S. ^National .Museum.

Ophiopneumicola elaphes Keegan, .1,943: 129 (d); Turk, 1947.
Locality: North America -; United States, Iowa. New York.
Host: Reptile - Elaphe obsoleta obsoleta.
"i
pphiopneumicpla hamertdni (Radford)

Entonyssus hamertoni Radford, 19.39: 251 (d). ’’’’

Ophiopneumicola hamertoni, Schmidt, 1940; Keegan, 1943; Turk, 1947.
Locality; Europe - England, London, Zoological Gardens.
Host: Reptile - ’Thamnophis si-rtalis.
’
’
:

Ophiopneumicola natricis Kee’gan

Type material in the 1.J,.:S.’ National ’Museum’.’

Ophiopneumicola natricis K.eegan,:.’1943’: 129 ’(d); Turk, 1947.
Locality:’ -North America - United States, Florida.
Host; Reptile - Natrix sipedon’.p-ictiventris: ’

Genus Hamer-tonia -Turk,;’ 1947

Type: Entonyssus bedfordi Radf-ord.

’

Diagnosis: Mites of medium size in which the "pulvillus on tarsus I

is lacking. Parasitic in the air, pas.s.ages of snakes.

Hamertonia bedfordi (Radford)

Entonyssus bedfordi. Radford, 1.937’: 39 (d),; :’’ ’.

Ophiopneumicola bedfordi, Schmidt, 1940; Keegan, 1943.
Hammertonia bedfordi, Turk, .1947: 21. ’.
 147

Hamertonia bedfbrdi, Radford, 1953.

:
Locality: Africa - Rhodesia, Mazabuka.
’

’
"
Host: Reptile - Dendraspis angusticeps.

Hamertonia schoutedeni Radford1

Hamertonia schoutedeni Radford, 1953: 106 (d).

Locality: Asia - India. Manipur State.
Host; Reptile - Naia tripudians fasciatus. ’

Genus Pneumophionyssus Fonseca. 1940

Type: Pneumophionyssus aristoterisi Fonseca.

Diagnosis:’ Mites of medium size in which the ’fixed arm of the che-
la is aborted. Internal parasites of snakes.
’
Pneumophionyssus aristoterisi Fonseca’

Pneumophionyssus aristoterisi’ Fonseca, 1940: 58 (d).

Locality: South America - Brazil, Sao Paulo.
’
Host: Reptile - Erythrolamprus aesculapii.
’
Family HALARACHNIDAE- Oudemans; 1906

Description of the family. Dorsal plate undivided in both sexes;

tritosternum absent; forked claw of palp present or absent, palpal tibia
and tarsus sometimes fused, ’so’that the palp has’, only four moveable
segments; peritreme very short, not reaching beyond’.’coxa III. Female
genital plate lacking, except in Pneumonyssus bakeri. Found only in
:
the respiratory system of mammals.
Remarks: The status of the five genera recognized here is rather
tenuous and is certainly not the final word. Very likely a separate genus
will need to be erected for the mites ’from each different group of hosts.
The following key is based entirely upon the literature and therefore
probably applies Only to the generotypes. We are quite aware that it
does not key out Pneumonyssus bakeri, which is the only halarachnid
from a rodent and no’doubt represents a separate genus.’

Key to the Genera of the Halarachnidae

’
’
1. Parasitic in the respiratory system of Pinnipedia (marine ver-
tebrates) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Parasitic in the respiratory system of terrestrial vertebrates. . 3

.148
2. Male and female with more than four pairs of setae’dorsally
between the posterior margin of the dors’al plate’and anal pt’ate,
and also ventrally between the level of’the.:foUrth pair of coxae
and the anal plate. Parasites of the hair seals . . . . Halarachne

Male and female with only two pairs of setae in the above areas..,
Abdomen of the gravid female greatly elongated in most spe-:
cies. Parasites of fur seals and walruses . . . . Orthohala.rachne

3. Palp with only four moveable segments, the palpal tibia and
palpal tarsus being fused. Combined length of the moveable
segments of the palp much less than. the fused coxae . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Pneumonyssus

Palp with four or five moveable segments, in either case, their

combined length is as great, or greater, than that of the fused
coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Palp with five moveable segments; modified forked palpal claw

lacking; peritremes slender and nearly as long as the diameter
of coxa IV. No constriction between podosoma and opisthosoma.
Parasitic in dogs and wart hogs . . . . . . . . . Pneumonyssoides

Palp with four or five moveable segments, forked palpal seta

present, peritremes much shorter than diameter of coxa IV.
Body constricted behind legs IV. Parasites of the lower Pri-
mates . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinophaga

Genus Halarachne Allman, 1847

""
" (Fig. 70)
.
.
Type: Halarachne halichoeri Allman.
Rhinixodes Nehring, 1884.
’
Rhinacarus Nehring. 1884.
Diagnosis: Abdomen of female not clavate, even when gravid. Sper-
mafodactyl of male not hooked at apex. Parasitic only in the nasal pas-
sages of the Phocidae (the hair seals).
Remarks; The genus has been ably reviewed by Newell (1947), from
whom we have borrowed freely. We did not make an exhaustive listing
of the literature for each species. For more complete references see
Newell (1947), and also Oudemans (1925), who gives a very thorough
and well illustrated account of the species known up to that time. Ac-
cording to Newell, Rhinixodes and Rhinacarus are synonyms because
"Both were proposed as possible substitutes for Halarachne on the grounds
that they were more appropriate."

Halarachne .halichoeri Allman

Halarachne. halichoeri Allman, 1847: 47 (d); Murray, 1876; Oudemans,

 1.49

1925; Vitzthum, 1936: 266; Ferris, 1942; Haarl^v. 1943: 273 (a);
Newell, 1947: 2.42 (c. d, t); Margolis, ’1954: 276.
Locality: Europe - Ireland to the Baltic Sea. North America-Uni-
ted States, West Coast.
Hosts: Mammals - Halichoerus grypus, Phoca vitulina richardi.
Remarks: Ferris (1942) reported this m-ite from Phoca richardi,
Pacific Grove, California. Newell doubted that it was actually this spe-
cies but Margolis. (1954) includes .it .in his.list of parasites from sea
mammal’s of the North American West Coast. .

Vitzthum (1936) reported it..from an elephant walrus in’the Berlin

.zoo which had been kept with hair seals. He expressed the opinion that
under natural conditions the elephant walrus would not harbor the mite.
Vitzthum also theorized that new hosts become infected on shore, when
larval mites are exhaled onto the ground, and ’crawl to new hosts.

Halarachne americaha Banks

(Fig. 70)- .;...: ’ ,
"
.
,
Type material’in the Museum of Comparative Zoology, Cambridge,
Massachusetts. .

Halarachne americana Banks, 1899: 213 (d); Ferris. 1925; Oudemans,

1925; Finnegan, 1934;. Ferris, 1942;. Doetschman, 1’944; Newell.
: :. l’-947: 244 (c. d, t).
Locality: Central America- "West Indies, CoastofYucatanto Flor-
ida and Bahamas. "
Host: Mammal - Monachus tropic alls.
’

’
Halarachne miroungae Ferris ’
Halarachne miroungae Ferris, 1925: 164(d); Doetschman, 1941; Ferris,
1942; Newell, 1947: 243 (c, d,.t); Margolis, 1954: 276.
Locality: North America - The Pacific Coast, from Baja California
to Washington State.
Hosts: Mammals - Phoca richardi, Mirounga angustirostris.
Remarks: Newell(1947) states that it is possible that the mites from
the two above hosts may not be the same species. Mirounga angustiros-
tris is the type’ host, ; . . .
’
,
Genus Orthohalarachne Newell, 1947
(Fig. 71)
Type: Halarachne attenuata Banks.
Diagnosis: Fairly large mites (one mm or more in length) in which
the opisthosoma of the gravid female is greatly elongated, giving the
mite a wormlike appearance. The male spermatodactyl forms a heavy
hook and bears a foliaceous appendage. Parasitic in the nasal passages
of the Otariidae and Odobenidae (fur seals and walruses respectively) ,
Remarks: Newell (194 7) .has fully reviewed the species included in
this genus. ...
. .
150

’Orthohalarachne attenuate (Banks)

.’ : ’..’ -(Fig. 71) -
/ .. ’; ;’

.
.

Halarachne attenuata .Banks, 1910:. 5 ( dh-’Ouaemahs,’ 1925; Finnegan,

’.’;
’

’
"

1934; Perris, 1942.

^ "’, ’
-
Orthohalarachne .attenuata, Newell. 1947: ’250 (c. d, t); Margblis. 1954:
’.:’
’’

’
’,... > -. ’-

’
.277. :- .

,
Locality: North America - Pribiloff Islands ’til’ the Bering Sea to
’

Depoe Bay, Oregon. ’ "

’
. .

.
.
Hosts: Mammals - Callorhirius ursinas cynoceprialus, Eumetopias

Qrthohalarachne diminuata (fioetschman) ,,

.
Type material in the U. S. National Museum.
Halarachne diminuata Poetschman, 1944: 72 (d).
Orthohalarachne diminuata, Newell,. 1947: 260 (c, d. t); Margolis, 1954.
Locality: North America - Goast of California to’St. Paul Island,-
Alaska.
Hosts: Mammals - .Callorhinus ursinus, Zaiophus californianus.

.. ., .Orthohalarachne magellanica (Finnegan)

Type material in the British Museum (Natural History).

Halarachne magellanica Pinnegan, 1934: 319 (d).
Orthohalarachne magellanica. Newell. 1947: 260 (c, t).
Locality: Islands of the South Atlantic - Falkland Islands.
:
Host: ’.Mammal - Otaria byronia.

Orthohalarachne reflexa (Tubb) ’

Halarachne reflexa Tubb, 1937: 417 (d).
Orthohalarachne reflexa, Newell, 1947: 258; Kadford, 1950.
Locality: Islands of the South Pacific - Lady Julia Percy Island,
Host: Mammal - "Tasmanian sea bear."
Remarks: Newell (1947) suspects that this is a synonym of attenuata.

Orthohalar achne rosmari (Oudemans)

Halarachne ro smart Oudemans, 1916: 132 (d); Finnegan, 1934; Ferris,

1942.
Orthohalarachne rosmari. Newell, 1947: 260 (c, t).
Locality: Islands of the Arctic - Franz-Joseph Land.
Host: Mammal - Odobenus rosmarus.
Remarks: This species was described from a walrus that had lived
in a zoological park in Hamburg for seven years after it was collected
in Franz-Joseph Land.
 151

Orthohalarachne zaiophi (Oudemans)

Halara.chne zaiophi Oudemans, 1916: 132 (d); Oudemans, 1’926; Finnegan,

1934; Doetschman, 1941: 23 (c); Ferris, 1942; D.petschman;; 1944.
Orthohalarachne zaiophi, Newell, 1947: 258 (c, d, t); Margolis, 1954.
Halarachne otariae Steding, 1926: 442 (d); Ass, 1935.
Locality: NorthAmerica - United States, coast of California.
Hosts: Mammals - Eumetopias jubata. Zaiophus californianus(=0ta-
ria californica).
Remarks: Newell (1947) believes that the types of otariae and zaio-
phi both came from the same sea lion’s head. Ferris (1942) first synony-
mized otariae. Ferris also suggests that zaiophi is a synonym of atten-
uata.

Genus Pneumonyssus Banks, 1901

(Fig. 72)
Type: Pneumonyssus simicola Banks
Pneumotuber Landois arid Hoepke, 1914: 393.
Type: Pneumotuber macaci Landois and Hoepke.
. Diagnosis: B/tedium to large mites, parasitic in the respiratory
tracts of the lower primates (except Pneumonyssus bakeri)..
Remarks: The most recent review of the genus is that by Furman,
1954.

Pneuroonyssus simicola Banks

(Pig. 72)
Pneumonyssus simicola Banks, 1901: 334 (d); Weidmann, 1915; Ewing,
1929; Vitzthum, 1930: 597; 1931: 49 (c. d...t); Oudemans. 1935: 494
(d. t); Radford, 1939; Buitendijk. 1945; Kirsch. 1950; @26 (v); Pur-
man, 1954: 39 (c, t); Zumpt and Till, 1.9.54; Baker et al., 1956: 35;
Hull, 1956: 653 (b).
Pneumonyssus griffithi Newstead, 1906: 49 (d, v); Weidmann, 1915;
Ewing, 1929; Vitzthum. 1930: 600.
Pneumotub.er macaci Landois and Hoepke, 1914: 393 (d); Oudemans, 1915;
Weidmann, 1915; Ewing, .1929; Vitzthum, 1930: 602,
Pneumonyssus foxi Weidmann, 1915: 37 (d); Helwig, 1925: 392 (m); Gay
and Branch, 1927: 49 (v); Ewing, 1929.
Localities: Africa - Abyssinia. Asia - Dutch East Indies; India;
Java and Sumatra. Also in hosts in many of the major zoological gar-
dens and research laboratories of Europe, North America and South
America.
Hosts: Ms-mmals - Macaca mulatta, Macacus rhesus, Pithecus
fascicularis, Pithecus nemestrinus, Pithecus rhesus, Theropithecus
gelada.
Hemarks: Helwig (1925),, Qay and Branch (1927), and Kirsch (1950)
have described in some .detail the effects of this mite on the host. It
does not seem to have a serious effect on the health of the host although
152

histological changes in the lung may be rather extensive. It was formerly

believed that the Halarachnidae have no nymphal stages, but Hull (1956)
found, described, and illustrated them in P. simicola. Apparently both
protonymph and deutonyrriph stages are nonfeeding and of very short du-
ration. ;: . . . . . .;

.
.

Pneumonyssus bakeri Furman

.
;

Type -material iri the’ collection of the .U. S, National Museum.

Pneumonyssus bakeri Furman, 1954: 35 (d>.
Locality: North America.-United States, California. ;

.
Host: Tamiasciurus douglasii moltipilosus...
Remarks: This mite probably-does not .belong in the genus Pneumo-
nyssus. It differs in having the sternal plate wider than long and in hav-
ing a distinct genital plate. Also, the host is a rodent rather than a
primate,

Pneumonyssus congoensis Ewing

Type material in the U. S. National Museum,

Pneumonyssus congoensis Ewing, 1929: 129 (d)i;i.Oudemans, .1935; Fain,
1952; Purman, 1954: 37 (c, d, t); .Fain, 1954; Zumpt and Till, 1954.
Locality: Africa Belgian Congo.
-
Host: Mammal - Papiosp; ’
Pneumonyssus dinoiti Oudemans

Pneumonyssus dinoiti Oudemans, 1935: 503 (d); Fain, 1952; Furman,

1954; Fain, 1954; Zumpt and Till, .1954.
Locality: "Probably from the Orient. "
Host: Mammal - Macaca mul’atta ( s Pithecus rhesus).

Pneumonyssus duttoni Newstead and Todd

Pneumonyssus duttoni Newstead and Todd, 1906: 41 (d); Weidmann. 1915;

Ewing, 1929: 127; Oudemans, 1935; Cooreman, 1946:. 331; Fain,
1952: 363 (d); Furman, 1954: 41; Fain, 1954; Vitzthum, 1930; Zumpt
and Till, 1954.
Locality: Africa - Belgian Congo.
Host: Mammal - Cercopithecus schmidti.
Remarks: According to Cooreman (1946), this mite is found only in
the trachea and bronchi, never in the lungs.

.Pneumonyssus mossambicensis Zumpt and Till

Type material in the South African Institute for Medical Research.

Pneumonyssus mossambicensis Zumpt and Till, 1954: 205 (d, cf and $);
Locality: Africa - Mozambique, Chicualacuala.
 . ’ ’

’: ,
. 153
. ’

/
Host: Mammal - Pap.io ursinus,

.
.

Pneumonyssus proc.avians Radford

Type material in the collection of-the Albany Museum, Grahamstown,

’

Union of South Africa. " .. .

,
Pneumonyssus procavians ’Radford, ,1938: ..147^); Furman, 1954: 40 (c,
d); Zumpt and Till, 1954: 210.^. .

Locality: Africa - Uriion of South Africa, Grahamstown.

Host: "Mammal - Procavia capensis.

Pneumonyssus rodhaini Pain

Pneumonyssus rodhaini Fain, 1952: 3.73 (d); Fairi. 1954.

Locality: Africa - "From the forests of Djugu. "
Host: Mammal - Colobus badius powelli.

Pneumonyssus santos-diasi Zumpt and Till

Type material in. the South African Institute for Medical Research.
Pneumonyssus. santos-diasi Zumpt and T.ill, 1954: 208 (d, d, ? and larva).
Locality: Africa. - Mozambique, Chicualacuala.
.Hosts: Mammals - Cercopithecus aethiops, Papio ursinus.

Pneumonyssus schoutedehi Fain

Pneumonyssus schoutedeni Pain, 1952: 369 (d); Fain, 1954.

Locality: Africa - Belgian -Congo, Ituri region (on the border of
Lake Albert. .;
’
. ’. . . .

Host: Mammal - Dendrohyrax arborens adolfi-friederici.-

’
Pneumonyssus ’stammeri Vitzthum

Pneumonyssus staromer.i Vitzthun-i, 1930: 606; Oudemans, 1935; Fur-

man, 1954.
Locality: South America. :
. Host: Mammal - Lagothrix poeppigii ( = Lagothrix infumatus).

;: Genus Rhinophaga Fain, ;1955

(Fig. 73)
Type: Rhinophaga cercopitheci Fain.
Diagnosis: Female dorsal plate nearly as long as idiosoma. Male
spermatodactyl sinuate and much longer than the gnathosomal palps.

Rhinophaga cercopitheci Fain

Type material in Musee Royal du Congo Beige.

154.’

Rhinophaga cercopitheci Fain, 1955: 318 (d, d" and 5, illus.).

Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Mammals - Cercopithecus ascanius montanus, Cercopithe-
cus mitis doggetti, Cercopithecus mitis stuhlmanni.
’

Bhinophaga atheruri Fain

.
Type material in the Musee Royal du Congo’ Beige.
’’ """ ’

’
Rhinophaga atheruri Fain, 1957: 77 (d, illus.).
Locality: Africa - Belgian Congo, Kivu.
Host; Mammal - Atherurus africanus centralis.

Rhinophaga leopoldi Fain

Type material in the Musee Royal du Congo Beige.

-
Rhinophaga leopoldi Fain, 1957: 71 (d, illus.).
Locality; Africa Belgian Congo, Kivu.
-
Host: Mammal - Atherurus africanus centralis.

Rhinophaga papionis- Fain

"
(Fig. 73) ,
,
:
^.
Type material in the Musee Royal du Congo Beige, ’.

-
.
Rhinophaga papionis Fain,. 1955: 309 (d, <f and’?, illus.). .

Locality: Africa - Belgian. Congo, .Lake Albert, Ruanda-Urundi,

Lake Tanganyika.
-
Host: Mammal Papio doguera tesseliatus.

Genus Pneumonyssoides Fain, 1955

(Fig. 74.) ,.
Type: Pneumonyssus caninum Chandler and Rune.
"

Diagnosis: Without constriction between pqdosoma and hystero-

soma. Dorsolateral expansions on podosoma absent. Parasitic in the
nasal fossae of dogs and wart hogs.

Pneumonyssoides caninum (Chandler and Ruhe) ’

:
’
(Fig. 74). .

Pneumonyssus caninum Chandler and Ruhe, 1940: 59 (d); Monlux, 1940;

Martin and Duebler, 1943; Camin and Rogoff, 1952; Douglas, 1951;
Gordon and Keep, 1951; Monlux and Turk, 1951; Monlux, 1951; Me
Clelland and Me Clelland, 1952; Schuiz and Thomas. 1953; Koutz
etal., 1953: 106 (m); Furman, 1954: 33 (c, d, t); Zumpt and Till,
. 1954: 204; Baker et al., 1956: 39.
Pneumonyssoides .caninum, Fa.in, 1955: 294.
’
:
.
Localities: Africa. Australia. North America - United States.
Cosmopolitan.
Hosts: Mammals - the common dog, Canis familiaris. Apparently
any breed of domestic dog may be affected.
 155

Remarks: These mites are found only in th-e upper respiratory tract
(the nasal passages and frontal sinuses), not in the bronchi and lungs.
There ’seems to be no definite clinical evidence that they have a serious
pathogenic effect on the dog.

Pneumonyssoides phacochoeri ’Fain

Type matei;?.al in Musee Royal du Congo Beige.

Pneumonyssoides phacochoeri Fain, 1955: 295 (d, o" and 9, illus.).
Locality: Africa - Belgian Congo, Ruanda’Urundi.
Host: Mammal - Phacochoerus aethiopicus,.

’Family RHINONYSSIDAE Trouessart. 1895

Description of the family. Legs rather thickset and stubby, with

well developed tarsal claws. Dorsal .plate entire or divided; ventral
plates reduced and poorly sclerotized. Peritre’m-e short or entirely’lack-
ing; tritosternum lacking or weakly developed.
These mites are found only in the respiratory tract of birds, yet it
is doubtful that’they represent a homogenous group. As in most endo-
parasites, host specificity is. marked. -
’
Key to the Genera of Rhinonyssidae
., Based, on the Females ’
.
.
1. Chelate portion of the chelicer.ae forming’only one-tenth or less
of their total length . . . . . . . . . . . . . . . . . . . . . . . . 2

Chelate portion .of the chelicerae forming one-seventh or more

of their total, length . . . . .. . . . . . . . . . . . . . . . . ... . 10

2. Stigma without p.eritreme., . . . . . . . . . . . . . . . . . . . . 3

Stigma with short peritreme that is elongate or circular . . . . 4

3. Mouthpartsv.entrally directed} not entirely visible from above.

(Whsn both fore coxae- are’.diree’ted forward, they i-neet above
the gnathosoma. Palps in some species curve upward rather
than downward.) Many species, found in a great variety of birds.
. . . . . . . . . . . . . . . . .. . . . . . . . . . . . . Sternostoma

Mouth parts apical. Opisthosonaal setae very thick, heavy,

blunt.Pound only in swallows . . . . . . . . . . . . . . . . . Cas

4. Chelicerae bulbous basally .. . . . . . . . . . . . . . . . . . . . 5

156

Chelicerae generally of uniform diameter throughout or grad-

ually attenuated. Not bulbous at base. :. . . . . . . . . :. . . . . 8

5. Dor sum with one or two plates . . . . . . . . . . . . ... . . . . 6

Dorsum with three or four plates . . . . . . . . . . . . . . . . . 7

6. Dorsum with two plates, one podosomal and one pygidial. (The
latter always bears two setae on the posterior margin.),Body
elongate; generally slightly constricted medially. Restricted
..to passeriform birds . . . . ;..’. . . , , ;’. . . . . Ptilonyssus ...
Dorsum with one plate. Body elongate; a well formed tritostern-
um present . . . . . . . . . . . . . . . . . . . . . . . Vitznyssus

7. Dorsum with four plates; one podosomal, one pygidial, and two
medial. (In the only species known, the two medial plates each
bear three prominent setae.,) From a Brazilian-brown creeper.
. ,, . . . . . ’. . . . ,... ...
. . . .... .-. ; . .^. . . . . Flavionyssus

Dorsum’with three distinct plates ’.’. . . . . . . . . Ptilonyssoides

8. With only one dorsal plate, the podosomal . , . . . Rhinonyssoides

Two or more dorsal plates present . . . . . . . . . . . . . . . . 9

9. With two dorsal plates: one podosomal and one opisthosomal.

Chelicerae of uniform diameter throughout or only slightly at-
tenuated. Restricted to passeriform birds . ... . Paraneonyssus

With six dorsal plates: one podosomal, one pygidial, and four

. . . . . . . . . . ... . . . ...
small medial plates. Cine species, from a Brazilian flycatcher
. . . .’.". . . . Trayanyssus ...
With six, dorsal plates: one podosomal, one opisthosomal and

flycatcher.. . . .
two on either side of the opisthosomal plate. From a Brazilian
. . . . . . . . . . . . . . . . . . . Ro.c.hanyssus

10. Peritreme lacking. Gnathosomal palps shorter than the rest

of the gnathosoroa. Found in shore and water birds , . Rhinonyssus

Peritreme present. Palps as long as or longer than the remain-

. der of .the gnathosoma . . . . . . .
. . . . . . . . . . . . . . . .11
11. Cheli’cera lacking the tixsd digit. The moveable arm well de-
veloped, slightly falcate and edentate , ; . . . . . . . . . . . . 14

Both digits of the chelicera present . . . . . . . . . . , .> . . . .12

 r57

12. Stigma and peritreme near the posterior end,of the body. Anal
pore surrounded by a free membrane. Found in ralliform birds
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rallinyasus

Stigma and peritreme in the usual position, above or anterior

,.
of coxa IV . . . . . . . . . . . ; . . . ...:.’. . . . . ....’..;. 13

13. Dorsum with a group of small, indefinite shields. Mouth parts

subventral. Found only in gulls and terns . . . . . . . Larinyssus

Dorsum with two shields. Reported from pigeons, herons, and

European nutcracker . . ... ;.’... . . . . . . . . . . . . Neonyssus

Dorsum with three shields. Parasitic in tinamiform birds . .

. . . . . . . . . . . . . . . . . . . . . . . . Tinaminyssus n. gen.

14. Without a tritosternum; with only three pairs of sternal setae

(no metasternal setae).. In strigiforra .birds . . . . . . Rhinoecius

With a tritosternum; metasternal setae present . . . Ruandanyssus

Genus Rhinonyssus Trouessart, 1894

... ..;; : (Fig. 75)
Type: :Rhinonysaus coniventris Trouessart. .

Sternostomum Trouessart, 1895 ( not Sternostoma Berlese and

Trouessart).
Type: Sterqostom.um rhinolethrum Trouessart.
Sommatericola Tragardh, 1904.
Type: Sommatericola levinseni Tragardh.
Diagnosis: Medium to large mites, having only a podosomal plate
or platelets; the sternal plate, lacking or -drastically reduced. Parasitic
in water birds. :

.Rhinonyasus coniventris Trouessart

(Fig. 75)
Type material in the British Museum (Natural History).
Rhinonyssus coniventris Trouessart. 1894: 723 (d); Hirst, 1921; Bedford,
1932; Vitzthum, 1935; Castro, 1948; Strandtmann, 1951: 130 (c, d);
Fain, 1956; Strandtmann, 1956: 130; Fain, 1957.
Sternostoma coniventris, Zumpt and Till, 1954: 84.
Rhinonyssus echinipes Hirst, 1921: 359; Castro, 1948; Bregetova, 1950;
Bregetova, 1953; 1956: 198.
Localities: Africa, Asia, Europe, and North America. (Probably
wherever the hosts may be found.)
Hosts: Birds - Arenaria interpres, Arquatella ptilocenemis, Ca-
toptrophorus semipalamatus, Charadrius alexandrinus, Charadrius hi-
aticula, Charadrius marginatus, Crocethia alba, Erolia alpina, Erolia
 158

maritina, Rhyacophilus glareola, Tringa ochrophus,. Tri-nga -strtata..

Rhinonyssus afribyx Pain

Rhmonyss-us afribyx gain, 1956: 1.50; .1957: .48 (illus.)- .

Locality: Africa - Belgian Congo, Ruanda-Urundi.

Hosts; Birds- - Afribyx senegallus,’ Stephanibyx lugubris.

Rhinonyssus alberti Strandtmann

Type naaterial in the U.. S.-National. Museum. ’

.
’.". .

’- .’ .
Rhinonyssus alberti Strandtmann, 1956: 140 (d, <S and 9, illus.); Fain,
:

’
1957: 48. ’ . .

.
Locality: North.Amer-ica - United States., California.. ,

.
Host: Bird - Colymbus caspicus.

Rhinonyssus apus P ain

Type material in Musee Royal du Congo Beige..

’
. . .

Bhinonyssus apus Fain, 1957: 151 (d, illus.); 1957: 48.

Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Apus caffer. ....
Remarks: This species has two. large dorsal plates, unlike other
. .

members of the genus. ’’This, -plus the fact that it is from a micropo-
diform bird, may necessitate a’separate generic status. ;.. .

Rhirioriyssus baledonicus Hirst

Rhinonyssus caledonicus ’Hirst,^l9?!: 357 (d); Strandtmann, 1956: 135;

.
’
Pam, 1957: 45. .’ .

Rhinonyssus (Rhinonyssus) ’caledonicus, Castro and Pereira;. 1947.

Sternostomum caledonicum. Vitzthum, 1935: 569; Bregetova, 1950; 1953:
332; 1956; 188.
U.’S.S.R. Europe - Shetland Islands; U.S.S.R.
Localities: Asia
-
Hosts: Birds - Aethia pus ilia, Cepphus columba, Cepphus grylle,
Uria aalge. Uria grylle. ’
’
.

Rhinonyssus himantopus Strandtmann .

Type mate’rial in the U. S.’ National Mu’seum;’ , !

Rhinonyssus himantopus Strandtmann, 1951; 1’36 (d); 1956: 432; Fain,

1956: 149; 1957: 44. ’ .’.
.
Localities: Africa Belgian Congo. Ruanda-Urundi. North Ameri-
-
ca - United States, Texas;/Florida.
’
Hosts: Birds - Charadrius tricollar.ia, Hemiparra crassirostris,
Himantopus mexieanus. . :::. ^
 159

Type
U.S.S.R.
Rhinonyssus minutus

.
(Bregetova)

material in the Zoological Institute of the

Sternostomum minutus Bregetova, 1950:..1007 (d); 1956: 198.

Sternostoma minutus, Furman, 1957: 483.
.
Academy of Sciences,
,

Rhinonyssus minutus. Fain, 1956: 150; 1957: 48.

Localities; Europe and Asia - U.S.S.R.
Host: Bird - Charadrius: hiaticula.

Rhinonyssus poliocephali Fain ..

Rhinonyssus poliocephali Fain, 1956: 149; 1957: 45 (illus.).

Locality: Africa - Belgian Congo, Ruanda-Urandi. :
Host: Bird - Poliocephalus ruficollis capensis.

Rhmonyssus rhinolethrum (Trouessart)

Sternostomum rhinolethrum Trouessart. 1895: 392 (d); Banks, 1915;

Vitzthum, 1935; Bregetova. 1950.
Rhinonyssus rhinolethrum, Castro, 1948; Strandtmann, 1951: 132 (c, d,
t); 1956: 137; Fain, 1956: 149; 1957: 43..
Sternostoma rhinolethrum, Zumpt and Till, 1955: 84 (t).
Sommatericola.levinseni Tragardh, 1904: 28 (d).
Rhinonyssus levinseni, Hirst. 1921: 357; Castro and Pereira. 1347.
Sternostomum levinseni,., Vitzthum, 1935; Bregetova, 1950; 1956: 198.
Rhinonyssus dartevellei Fain and Vercammen-Grandjean, 1953: 35 (d) .
Zumpt and Till, 1955: 84 (t); Fain. 1956: 149; 1957: 44.
Locality: Apparently wherever its hosts , the anseriforin birds
(ducks, geese, swans, and related birds) may be found, including North
America, Greenland, Europe, Asia, and Africa.
Hosts: Brids Alopochen aegyptiacus. Anas acuta. Anas caroli-
-
nenais. Anas erythrorhyncha, .Anas leucostigma. Anas platyrhynchos,
Anas strepera, Aythya affinis, Branta canadensis, Cygnus columbianus,
"domestic goose," Mareca americana, Mergus megganser, Plectgop-
terus gambensis, Sarkidiornis melanotus. Spatula clypeata, Somnaateria
mollissima.

Rhinonyssus water stoni Hir.st

Rhinonyssus waterstoni. Hirst, 1921: 359 (d); Castro and Pereira, 1947;
Strandtmann, 1956.
Rhinonyssus (?) waterstoni. Fain, 1957: 44.
Sternostomum waterstoni, Vitzthum, 1935; .Bregetova. 1950; 1956.
Locality: Europe - Shetland Islands; U.S.S.R.
Host: Bird - Aica torda. ...
 ’
’

160 -

-
Genus Larinyssus Strandtmann, 1948
.,. ,. (Fig, 76) ,. .
’;

.
,

.
.
Type: Larinyssus orbicularis Strandtmann.
Diagnosis: A medium to large mite with many small dorsal plate-
lets . There is a short peritreme but the mouthparts are mostly hidden
from above. Restricted to the nasal passages of .gulls and tern? (Lari-
’:
’’"
"
"
dae).

Larinyssus orbicularis Strandtmann

(Elg. 7G)
Type U. S. National Museum.
material in the
Larinyssus orbicularis Strandtmann,:.1948: 507 (d); Pereira and .Castro,
1949; Bregetova, 1950; Zumpt arid Patterson. 1951: 78; Zumpt and
Till, 1955: 68; Pain. 1956: 157; Bregetova. 1956: 198; Fain, 1957:
60. ..
.
.

.
Localities: Africa - Central and South Africa. Europe - U. S. S. R.
North America United States, California, Texas.
-
Hosts: B’irds - Chlidonias leucoptera, Gelochelidon nilotica, Larus
argentatus, Larus atricilla, Larus .delawarensis, Larus dominjcanus,
Larus occidentalis. Sterna hirundo, .Sterna maxima.

Genus Rallinyssus .Strandtmann, 1948

’

(Fig. 77)

.
Type: Rallinyssus caudistigmus Strandtmann.
Diagnosis: A large species (about one mm, long) with large chelae,
a single dorsal plate, .a free membrane surrounding the anal pore, and
posterior stigmata, ,
,

Rallinyssus caudistigmus Strandtmann

.’.
(Fig. 77)
’
.
’

’Type material in the U. S. National Museum.

Railinyssus caudistigmus Strandtmann, 1948; 512 (d); Pereira and Cas-
tro, 1949; Bregetova, 1950; 19,56: 198; Fa.in.. 1957; 58,. .

Locality: Europe - U.S.S.R. North America United States, New

-
York, Texas. Africa - Egypt.
Hosts: Birds - Fulica americana, Fulica atra, Rallua elegans.

Rallinyssus congolensis" Fain

Type material in Musee Royal du Congo Beige. ;

Rallinyssus congolensis Fain, 1956; 396; 1957: 58 (illus.).
Locality: Africa - Belgian.Congo, Akany.aru River.
Host: Bird - Limnocorax flavirostra.

Rallinyssus limnocoracis Fain

Type material in Musee Royal du Congo Beige.

 161

Rallinyssus limnocoracis Fain, 1956: 397; 1957: 60 (.illus.).

Locality: Africa - Belgian Congo, Akanyaru River.
Host: Bird - Limnocorax flavirostra.

Genus Tinaminyssus. New genus

(Fig. 83)
Type: Neonysaus (Ptilonyss.oides) trappi Pereira and Castro.
Diagnosis: Female with three dorsal plates, the middle plate rather
irregular. Chela from one-seventh to one-fifth the total length of the
chelicera. Stigma with a short peritreme. Parasitic in tinamiform
birds.
Remarks: See discussion under Ptilonyssoides.

Tinaminyssus trappi (Pereira and Castro). New combination

(Fig. 83)
Neonyssus (Ptilonyssoides) trappi Pereira and Castro, 1949; 231 (d).
Neonyssus trappi. Fain, 1957: 57.
Locality: South America - Brazil, Sao Paulo.
Host: Bird - Tinamus solitarius.

Tinaminyssus navajasi (Pere.ira and Castro). New combination

Neonyssus (Ptilonyssoides) navajasj. Pereira and Castro, 1949: 231 (d).

Neonyssus navajasi. Fain, 1957: 57.
Locality: South America - Brazil, Mato Grosso, Sao Paulo.
Host: Bird Nothura maculosa.
-
Genus Neonyssus Hirst, 1921
(Fig. 78)
Type: Neonyssus intermedius Hirst.
Neonyssoides Hirst, 1923: 975 (subgenus).
Type: Rhinonyssus (Neonyssoides) nuc ifragae Hirst.
diagnosis: Mites of medium size, with two large dorsal plates,
large chelae, and short peritremes.

Neonyssus intermedius Hirst

Neonyssus intermedius Hirst, 1921: 771 (d); Vitzthum, 1935; Castro,

1948; Pereira and Castro, 1949; Fain, 1957: 52.
Locality; Islands of the Indian Ocean Madagascar.
-
Host: Bird-"Sur un Oiseaude Madagascar. ’’

Neonyssus ardeae Zumpt and Till

Neonyssus ardeae Zumpt and Till, 1955: 63 ( d, sf and ?); Fain. 1956;
1957: 53 (illus.).
1.62 :

.
.
.
Locality: Africa Belgian Congo, Ruanda-Urundi; Union of South

-
Africa,Transvaal.
Hosts: Birds - Ardea melanocephala, Nycticorax nycticorax.
Remarks: It is quite likely that this is a synonym of Neonyssus be-
lopolskii Bregetova (Strandtmann,.,19’56: 137).

Neonyssus belopolskii Bregetova

Typ.e material in the Zoological Institute of the Academy of Sciences,

U.S.S.R,
Neonyssus belopolskii Bregetova, 19.50: 1005 (d); 195.3; 1956; Fain, 1957.
Localities: Asia-’U. S. S. R.,’Primer Territory, Sudzuknin Govern-
mental Reservation. North America - United States, Texas.
Hosts: Birds - Ardea cinerea, Hydranassa tricolor.

Meonyssus bubuici Zumpt and Till

Type material in the South African Institute for Medical Research.

Neonyssus bubuici Zumpt and Till, 1955: 66 (d, d", $, andnymph); Fain,
.1956: 134; 1957: 54. ,, ..
Locality: Africa - Belgian Congo, Astrida; Union of South Africa,
Johannesburg.
Hosts: Birds - Bubulcus ibis (type host), Egretta {Mesophoyx) in-

’
termedia brachyrhyncha!

"
. .

Neonyssus buteonis Fain

Type material in Musee Royal du Congo Beige.

Neonyssus buteonis Fain, 1956: 135 (d, 9); 1957: 51 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Birds Buteo rufofuscus augur, Lophaetus occipitalis.
-
Remarks: It is possible that this mite is not congeneric with others
of this group. It has only one dorsal shield and its host, falconiform
birds, differs greatly from hosts of other species of the genus.

Neonyssus columbae Crossley

, Type material in the U., S. National Museum.

Neonyssus (Neonyssus), columbae Crossley, 1950: 309 (d); Crossley,
1952. ,
,
Neonyssus columbae, Zumpt and Till, 1955; Fain, 1956: 134; 1957: 55.
Localities: Africa - Belgian Congo, Ruanda-Urundi. NorthAmerica-
United States, Texas.
Hosts: Birds - Columba livia (type host), Milvus (aegyptlus) tene-
brosus i, Streptopelia semitorquata.
Remarks: Milvus tenebrosus is a kite and we areinclinedtobelieve
that the presence of Neonyssus columbae in that host was purely acci-
dental.
 163

Neonyssus ^ijiobrychi gain

Type material in Musee Royal dtr Congo Beige. ’

Neonyssus ixobrychi Fain, 1956:’134 (d;’ S); ’1957:: 54’(illus.);.
Locality: Africa - Belgian Congo, Ruanda-Uruhdi.
Host: Bird Ixo’orychus minutus. "
’’’

’
-

Neonyssus melloi Castro

Neonyssus (Neonyssus) melloi Castro, 1948: 270 (d); Pereira and Cas-
tro. 1949; Crossley. 1952; Zu’mp.f and Till, 1955: 68’; Fain, 1957:
52. .-.. . , :.

.
Localities: Africa Union of SoutK Africa, Transvaal’. North Amer-
-
ica - United States, Texas. South America - Brazil, State of Sao Paulo.
Hosts: Birds - Columba livia (type host), Stigmatopelia senegalen-
sis, Turtur afer. ’. .’.;; "
’
.
;
Neonyssus nucifragae (Hirst)’
’ ’

Rhinonyssus (Neonyssoides)’ nucifragae- Hirst, 1923: 975 (d).

Neonyssus nucifragae, Vitzth’um;’1935; Castro, 1948; Pereira and Cas-
tro. 1949; Bregetova. 1953: 332; "1956: 199; Fain, 1957.
Localities: Asia U.S.S.R., Primor. Europe - Germany.
-
Host: Bird Nucifraga caryocatactes.
-
Neonys sus schoutedeni Fain

Type material in Musee’Royal du Congo Beige.

Neonyssus schoutedeni Fain, 1956: 134 (d, 9); 1957: 49 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird Halcyon leucocephala. ’
-
Remarks: This species has only a single dorsal plate and may not
be congeneric with other species of this group.

Neonyssus serraoi Castro

Neonyssus (Neonyssus) serraoi Castro, 1948: 271 (d); Pereira and Cas-
tro, 1949; Pain, 1957.
Locality: South America - Brazil. Mato Grosso.
Host: Bird - Rhynchotus rufescens.

Neonyssus treronis Fain

Type material in Musee Royal du Congo Beige.

Neonyssus treronis Fain, 1956: 394; 1957:’ 56 (illus.).
’

Locality: Africa - Belgian Congo, Akanyaru.

Host: Bird - Treron calva: ;’
’
 164

Neonyssus zenaidurae Crossley

’"(Fig. 78)
Typematerial in the U..S. National. Museum.
Neonyssus zenaidurae Crossley, 1952: .3.86 (d); Zumpt and Till;,..,1.955;
Fain, 1956; 1957. ..; , ;. ; _

.
.
Locality: North America United .States, ..Texas. :

-
. .

Hosts: Birds Columbigallina passerina, Zenaidura macroura,

-
Genus Rhinoecius Cooreman, 1946
.., (Fig. 79)-, ..
.

.
.
.
’:’ Type: Rhinoecius .oti Cooreman; ’.

"
’
Diagnosis: Medium to large mites, with one or more dorsal plates,
but .in which the.immoveable arm of the chela is lacking and. the move-
able arm is scimitar-shaped..

.
.. .

.
.
Rhinoecius oti1 Cooreman"

Type material in Musee d’Histoire Naturelle, Belgium.

Rhinoecius oti Cooreman, .1946: 1 (d); Strandtmann, 1948; Pereira and
Castro; -1949; Strandtmann, 1951; Zumpt and. Patt.erson, 1951;
.- Stpandtmann., .19.52: 20.5 (t); Zumpt and .Till, 1955: 82; F,ain, 1957; 130.
Locality: Europe - Belgium:. ..
.

"
.
Host: Bird: - Asio ptus otus. . .
.
i Rhinoecius africanus (Zumpt and Patterson)

’ Type material in the Natal Museum, Union of South Africa; para-

type sin the South African Institute for. Medical Research, .Johannesburg.
Rhinonyssus africanus. Zumpt and Patterson, 1951; 90 (d).
.Rhinoecius africanus, Zumpt and Till, 1955;..82; Fain, 19’57: 130.
Locality: Africa Union of. South Africa,, near Pietermaritzburg.
-
Host,:., .Bird - Asxo capensis.

Rhinoecius bisetosus Strandtmann

Type material in the U. S. National Museum.

Rhmoecius bisetosus Strandtmann, 1952: 212,(d>; Zumpt and Till. 1955;
Fain, 1957. . .

Locality: North America.- United States, .Texas.

Host: Bird - Speotyto cunicularia..,

Rhinoecius coor.emani Strandtmann

(Fig. 79)
Type materialin the U. S. National Museum.
Rhinoecius cooremani Strandtmann, 1952: 208 (d); Zumpt and Till, .1955;
’
Fain, 1956: 394; 1.957. .
.
.

Locality: North America - United States. Texas. .

 16:5

Host: Bird - Strix yaria.

. Rhinoecius grandis Strandtmann

Type material in the U. S. National Museum. ,:

Rhinoecius grandis Strandtmann, 1952: 205 (d); Zumpt and Till, 1955: 90;
Pain, 1957.
Locality: North America - United States, Texas.
Host: Bird - Bubo virginianus. .

.
.
, Hhinoecius tytonis Fain ..

.
Rhinoecius tytonis gain, t95S: 394; 1957: 131 (illus.).
Locality: .Africa - Belgian Congo, Akanyaru.
Host: Bird - Tyto alba affinis. .

.
.
Genus Ruandanyssus Fain, 1957

.
Type; Ruandanyssus terpsiphonei Fain.
Diagnosis: Chela with moveable digit only; metasternal setae and
tritosternum present. .; : ’. ,
.
Ruandanyssus terpsiphonei .Fain

Type material in Musee Royal du Congo Beige.

Ruandanyssus terpsiphonei Fain, 1957: 148 (d. illus.).
Locality: Africa - Belgian Congo, Akanyaru, Astrida.
Host; Bird - Terpsiphone viridis kivuensis,

Genus Sternostoma Berlese and Trouessart, 1889.

(Pig. 80)
Type: Sternostoma cryptorhynchum Berlese and Trouessart.
Diagnosis: Small to medium mites; chelae minute; mouthparts
mostly hidden from. above; peritreme absent; sternal plate present; one
or two. dorsal plates present.
Remarks: For an excellent discussion of the genus, see Furman
(1957). ..: . .
.
Sternostoma c ryptorhynchu m Berlese and Trouessart

Sternostoma cryptorhynchum Berlese and Trouessart, 1889: 126 (d);

Vitzthum, 1935; Castro and Pereira, 194’?; Bregetova, 1950; Strandt-
mann, 1951; Porter and Strandtmann,. 1952; Zumpt and Till, 1955;
Fain, 1956: 152; 1957: 70 <illus.); Fu.rman, 195.7: 475 (illus.).
Rhinonyssus (Sternostoma). cryptorhynchum, Castro, 1948:257,; Pereira
and Castro, 1949. .:.’
.
166
:
Locality: Europe - France.
Host: Bird - Passer domesticus.
Remarks: This species has recently been independently redescribed
and illustrated by both Furman (1957) and Fain (1957). In both cases,
the descriptions and illustrations were based on the type specimen that
had been loaned to both gentlemen’-by Professor Marc Andre.

Sternostoma boydi Strandtmann

(Fig. 80)
Type material in the’U. ’S. National. Museum.
Sternostoma boydi Strandtmann, 1951: 138 (d); Zumpt and Till, 1955;
Pain, 1956: 151; 1957: 66; Furman, 1857: 480. :
Localities: Africa - Belgian Congo, Ruanda-Urundi. North Amer-
ica - United States, Texas. . .;’

.
.
Hosts: Birds - Actitis hypoleucus, Arenaria interpres, Corcethia
alba (type host), Larus delawarensis, Larus atricilla, Phylomachus pug-
nax, Rhyacophilus glareola, Totanus nebularius, Tringa ocrophus.

Sternostoma cp.l.ii Pain’. ; ’ ’

.

Sternostoma colii Fain, 1956: 392; 1957: 79(illus.).

Locality: Africa - Belgian Congo, Astrida.
Host: Bird - Colius st’r.iatus kivuensis. .:.

Sternostoma cooremani Fain ’

Type material in Musee Royal dil Congo Beige.

Sternostoma cooremani Fain, 1956: 154 (d; $); 1957: 72 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Birds - Mellitophagus lafresnayi, Merops apiaster. Merops
.: :
’
nubricoides. ’ ’..’
Sternostoma cuculorum Pain

.-Type material in Musee Royal du Congo Beige.

Sternostoma cuculorum Fain, 1956: 156 (d, 9); 1957: 74 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
.

Hosts: Birds - Chrysococcyx caprius, Clamator levaillanti, Cu-

culus canorus, Cuculus solitarius (type host).

Sternostoma dureni Fain

. Type material in Musee Royal du Congo Beige.

Sternostoma dureni Fain, 19-56:-153 (d, $); 1957: 68 (illus1;),
Locality: Africa Belgian Congo, Ruanda-Urundi.
-
Hosts; Birds - Turdoides jardinei, Turdoides melanops, Turdus
’
olivaceus graueri (type host).
 167

Sternostoma hirundinis Fain

Type material in Musee Royal du Congo Beige.

Sternostoma hirundinis..Fain, 1956: 154 (d. d- and 9);. 1957: 72 (illus.).
Locality: Africa - Belgian Congo,, Ruanda-Urundi,
Hosts: Birds - Hirundo smithi, Psalidoprocne albiceps .(type host).
Hemarks: "It is noted that specimens from Psalidoprocne have no
peritreme but those from Hirundo have a small circular perltreme. "
’> (Translated from Fain, 1956.) .
.

Sternostom.a. hutsoni Purman . ;;:;:

Type in the U. -S. .National Museum.

Sternostoma hutsoni Furman, 1957: 477 (d, illus.).
Locality: North America - United States, California.
.. .-
Host: Bird - Hylocichia ustulata ustulata.

Sternostoma lagonostictae Fain

Type material in Musee Royal du Congo Beige. ... -

Sternostoma lagonostictae Fain, 1956:-1.55 (d, $); 1957: 72<illus.).
Locality: Africa - Belgian Congo, Ruanda Urundi.
Host: Bird - Lagonosticta rubricata congica.

Sternostoma laniorum Fain

Type material in Musee.Royal du Congo Beige.

Sternostoma laniorum Fain. 1956: 156 <d, $); 1957: 76 (illus.),
Sternostoma laniorum var. batis Fain, 1957: 77.
-
Locality: Africa Belgian Congo, Kuanda-Urundi.
Hosts: Birds - Batis molitor, Lanius- collaria, Lanius collurio (type
host), Lanius excubitorius.

Sternostoma meddai Lbmbardini

Sternostoma meddai Lombardini. .1953: 187 (d).

Locality: Europe - Italy.
Hosts: Birds -’Canaries," "cardinals."

Sternostoma nectarinia Fain

Type material.in Musee Koyal..du Congo Beige.

Sternostoma nectarinia Fain, 1956: 152 (d, S); 1957: 68 (illus.).
Locality: .Africa - Belgian. Congo, Ruanda-Urundi.
Hosts: Birds - Ch’alcomrtra.senegalensis, Cinnyris cupreus, Cin-
nyris regius (type host), Nectarinia purpureiverrtris.
168

Sternostoma spatulatum Furman

..Type material in the U.. S. National.Museum.

Sternostoma spatulatum Furman, 1957: 480"(d, illus.). . ’..":
Locality: North America - United States, California.
Host: Bird - Hylocichia ustulata ustu’lata.

Sternostoma stran’dtmanni Furman

Type material in the U. S. National Museum.

Sternostoma strandtmanni Furman, 1957: 476 (d, illus.).
Locality: North America - United States, California.
Hosts: Birds - Agelaius tricolor (type host), Molotfirus ater cali-
fornicus. ... ’

’
.

Sternostoma sturnicola Fain

Type material in Musee Royal du Congo Beige.

Sternostoma sturnicola Fain, 1956: 151 (d, $); 1957: 65 (illus.).
Locality: Africa-.Belgian Congo, Ruanda-Urundi. ’’"’
Hosts: Birds - Buphagus. afric anu s, Lamprotornis ’purfaropter’us
(type host). ’

Sternostoma technaui (Vitzthum)

Sternostomum technaui Vitzthum. 1935: 569 (d); 1943: 657.

Rhinonyssus (.Rhinonyssus) technauiy-.-Castro, 1948:’ 257; Pereira and
Castro, 1949. .-:’’-"
.
Sternostoma technaui, Strandtmahri, 1951; Zumpt and Till, 1955; Pain,
1957; Furman,’,1957: 482 (illus.).
Locality; Europe -Germany. .
’
,
Host: Bird - Cinclus cinclus aquaticus.
’
’
.Remarks: Furman (1957) redescribed and illustrated this species
from- specimens obtained on loan .from Vitzthum’s collection.

Sternostoma thienponti Fain

Type material in Musee Boyal du Congo Beige’’.

’
Sternostoma thienponti Fata, 1956: 152 (d. ?); 1957: 68 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Dicrurus adsimilis.

Sternostoma tracheacolum Lawrence

Sternostoma tracheacolum Lawrence, 1948: 364 (d); Stephan and’Kas-

, chula, 1950; Strandtmann, 1’951; Lombardini, 1953; Zumpt and Till,
1955; Bregetova, 1956: 198; Baker et al., 1956: 41 (c);-Pain. 1956;
1957: 65 (illus.); Furman, 1957: 478 (illus.).
 169

Sternostoma castroae Fain, 1956: 393.

Localities: Africa - Union of South Africa, ’Pietermaritzburg; Bel-
gian Congo, Astrida. Europe and Asia - U.S.S.R., Volga Delta. North
America - United States, California.
. Hosts: Birds -"Canary" (type host), Acrpcephalus arundinaceus,
Agelaius tricolor, Hirundo rustic’a. Icterus bullocki, Macronyx croceus.
Remarks: StephanandKaschula (1950) state that the mite may cause
serious respiratory difficulties and even death in canaries. They recom-
mend exposing the birds to a vapor of DDT and benzine bexachloride mix-
ture for five minute periods at’fourteen day intervals for satisfactory
control. Furman (1957). found this mite in wild birds of North America.

Sternostoma turdi Zumpt and Till

Type material in the South African Institute for Medical Research.

Sternostoma turdi Zumpt and Till, 1955: 85 (d, 9 illus.); Fain, 1956: 151;
1957; Furman, 195’?: 481 (illus.).
Locality: Africa - Union of South Africa, Transvaal, Cape Province;
Belgian Congo, Ruanda-Urundi.
Hosts: Birds - Turdus olivaceii^, Turdus abyssinicus baraha.

Genus Cas Baker and Wharton, 1952

’(Fig: 81)
Type: Rhinonyssus (Bhiriacarus) angrensis Castro.
Rhinacarus Castro, 1948: 257 (subgenus). Preoccupied.
Type: As above.
Diagnosis: Medium sized mites with no peritreme; chelae minute;
mouth parts entirely visible from ab.ove; legs I longer than the others;
with two dorsal plates. Parasitic in swallows (Hirundinidae).

Cas angrensis (Castro)

(Fig. 81)
Rhinonyssus (Rhinacarus) angrensis Castro, 1948: 257 (d).
Cas angrensis. Baker and Wharton, 1952: 81,
Sternostoma angrensis, Zumpt and Till, 1955: 91; Fain, 1957: 84.
Localities: North America - United States, Texas. South America -
Brazil, Rio de Janeiro. Angra dos Reis.
Host: Bird - Progne chalybea domestica.
Remarks: Previously unpublished records that we have of this mite
are from the United States (Texas) from the following swallows: Irido-
procne bicolor, Petroch’elidon fulva, Petrochelidon pyrrhonota, Progne
subis.

Genus Ptilonyssoides Vitzthum, 1935

(Fig, 82)
Type: Ptilonyssoides triscutatus Vitzthum.
 170

Diagnosis; Medium sized, elongated mites having three dorsal

plates, short or circular peritremes, apparent sternal plate, .and che-
. lae that are .less than one-tenth the total length of. the chelicera.
i : Remarks; Vitzthum (1935) did not describe or illustrate the cheli-
cerae of P. triscutatus. When Per’eira and Castro (1949) found two new
nasal mites with three dorsal plates (as in. triscutatus but with large
chelae).they assumed that triec.utatus also had/large chelae and placed
their new species with it even though the hosts of triscutatus and of
Pereira and Castro’s two new .species belonged to two entirely different
.bird .orders. Recently Fain (’1956) .has .shown that .Ptilonyssoides tri-
scutatus has minute chelae and .that therefore P. triscutatus Vitzthum
is not congeneric with P. navajasi’Pereira and Castro and P_. trappi
Pereira and Castro. We are therefore deleting Pereira and Castro’s
two species from the genus and propose the generic name Tinaminyssus
for the latter two (see p. 161).

: -Ptilohyssoides triscutatus Vitzthum

.
.
.
(Pig. 82). . :.’<

,
Ptilonyssoides triscutatus Vitzthum, 19S5:,581 (d).

.
Neonyssus (Ptilonyssoides) triscutatus, Castro, .1948: 276; Pereira and
’

Castro.. 1949.
Neonyssus triscutatus,
’
Zumpt and
Till. 1955: 68.
Ptilonyssus triscutatus. Fain, 1956: 1.36; 1957: 121 (illus.).
Localities: Africa - Belgian Congo, Ruanda-Urundi. Europe.
Hosts: Birds - Merops apiastef, Merops persicus.

Ptilonyssoides’dicruri (Fain). New combination

Type material in Musee Royal du Congo Beige.

Ptilonyssus dicruri Fain, 1956: 137 <d, 9); 1957: 122 (illus.).
Locality:’ Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Djcrurus adsimilis.
Remarks: This and the following species are from passeriform
birds. They do not .agree entirely with triscutatus and me’littophagi,
which are from coraciiform birds and perhaps separate generic status
may eventually be warranted. ’,

Ptilonyssoides dioptrornis (Pain). New combination

’
Type material in Musee Royal du Congo Beige.
Ptilonyssus dioptrornis Fain, 1956: 137 (d,.. 9); 1957: 124 (illus.).
Locality:. Africa - Belgian Congo. Ruanda-Urundi.
Hosts: Birds - Bessonornis archeri, Cossypha heuglini, Dioptror-
nis fischeri toruensis (type host), Myrmecocichia nigra.

Ptilonyssoides melittop.hagi (Fain). New combination

Type material in Musee Royal du Congo Beige..

 171

Ptilonyssus melittophagi Fain, 1956; 136 (d, 9);. 1957: 121 (illus.),
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Birds - Melittophagus lafresnayi, Melittophagus pusillus.

Genus Flavionyssus Castro, 1948

(Fig. 84)
Type: Ptilonyssus (Flavionyssus) rabelloi Castro.
Diagnosis: Mites with one podosomal, one pygidial and two inter-
mediate dorsal shields.

Flavionyssus rabelloi Castro

(Fig. 84)
Ptilonyssus (Flavionyssus) rabelloi Castro, 1948: 266 (d); Pereira and
Castro, 1949.
Ptilonyssus rabelloi, Zumpt and Till, 1955; Pain, 1957: 109.
Locality: South America - Brazil, State of Sao Paulo.
Host: Bird - Sittasomus griseicapillus sylviellus.

Genus Rochanyssus Castro, 1948

(Ftg, 85) ,.... , .
.,. .,

Type: Neonyssus (Rochanyssus) werneri Castro.

Diagnosis: Two large dorsal plates and four small plates laterad
of the large opisthosomal plate.

Rochanyssus werneri Castro

(Fig. 85)
Neonyssus (Rochanyssus) werneri Castro, 1948: 272 (d).
Ptilonyssus (Rochanyssus) werneri, Pereira and Castro, 1949: 226.
Ptilonyssus werneri. Fain. 1957: 86.
Locality; South America - Brazil, State of Sao Paulo,
Host: Bird - Elaenia obscura sordida.

Genus Travanyssus Castro, 1948

(Pig. 86)
Type: Neonyssus (Travanyssus) paranensis Castro.
Diagnosis: Body elongated, with one podosomal and five small opis-
thosomal platelets.

Travanyssus paranens is Castro

(Fig. 86)
Neonyssus (Travanyssus) paranensis Castro, 1948: 276 (d).
Ptilonyssus (Travanyssus) paranensis, Pereira and Castro, 1949: 227.
Ptilonyssus paranensis, Zumpt and Till, 1955: 90; Fain, 1957: 100.
Locality: South America - Brazil, State of Parana.
Host: Bird Elaenia flavogaster flavogaster.
-
172

Genus Rhinonyssoides Hirst, 1921

..

.
(Fig. 87)
.
. Type: Rhinonyssoides trouessarti Hirst.
Diagnosis: Medium to large mites with attenuated chelicerae, one
dorsal plate, and short peritremes.
Remarks: As this genus now stands, it includes a number of mites
from several unrelated birds. We think it is quite likely that a critical
restudy of the mites will reveal valid criteria for generic separation.
Until that time, it is best to maintain their present status.

Rhinonyssoides trouessarti Hirst

Rhinonyssoides trouessarti Hirst, 1921; 770; Vitzthum, 1935.

Ptilonyssus (Rhinonyssoides) trouessarti, Castro, 1948: 266; Pefeira
and Castro, 1949. .,, . .

Ptilonyssus trouessarti, Zumpt and Till, 1955; Pain, 1957’:’. 109.

Locality: Australia.. ... .

Host: Bird - Sphecotheres’ maxillaris.

Rhinonyssoides cerchneia (Fain). New combination

Ptilonyssus cerchneis Fain, 1957: 133 (d, illus.).

Locality: Africa - Belgian Congo.",’...
Host; Bird - Cerchneis tinnunculus rufescens. ’’. ’

’
Rhinonyssoides cinnyris (Zumpt and Till). New combination

Type material in the South African Institute for Medical Research.

Ptilonyssus cinnyrjs Zumpt and Till. 1955: ’78 (d, $; illus.); Eain, 1956:
"’’
146; 1957: 86.
Locality: Africa - Belgian Congo, Ruanda- Urundi; Union of South
Africa, Cape Province,
Hosts: Birds - Chalcomitra senegalensis/-Cinnyrls afra (type host),
Cinnyris regius. Nectarinia kilimensis.

Rhinonyssoides donatoi Pereira and Castro

(Fig. 87)
Ptilonyssus (Rhinonyssoides) donatoi’ Pereira and Castro, 1949: 224 (d);
Zumpt .and Till. 1-955.
Ptilonyssus donatoi Fain, 1957: 84.
-
Locality: South America Brazil, Sao Paulo.
Host: Bird - Coragyps’atratus foeteris.

.Rhinonyssoides indicatoris (Pain)’^’ New combination

Ptilonyesus indicatoris Fain, 1957: 134 (d,’ fllus.).
Locality: Africa - Belgian Congo, Akanyaru.
Host: Bird - Indicator indicator.
 173

Rhinonyssoides nova-guineae Hirst, New combination

Rhinonyssus nova-aujneae Hirst, 1.921: 769 (d>; Vitzthum, 1935; Castro

and Pereira, 1947; Pain, 1957.

’
.
-
Locality: Islands of the Pacific Ocean New Guinea;
"

Host: Bird - Craspedophora magnifica.

Remarks: Arestudy of the type will be necessary to assign this spe-
cies to its. proper genus. It is removed from Rhinonyss’us primarily be-
cause it was found .in a terrestiral bird (bird-of-paradise). All other
species of.Rhinonyssus .are frona water bir’ds. Placing it in Rhinonys-
soides is purely provisional. ;

.
Rhinonys soides spuzai Pereira and Castro ’

Ptilonyssus (Rhinonyssoides) souzai Pereira and Castro, ’1949: 223 (d>;

Zunapt and Till, 1955. . .
’/
Ptilonyssus SQUzai, Fain,.:1957: 84.
"
;... ’.
.

Locality: South America - Brazil, State of Sao Paulo.

Hosts: Birds - Milvago chimachj.m.a, Rupornis magnirostris ’taa.g-

Rhinonyssoides squamosus (Vjtzthum)

Rhinonyssus squamosua Vitzthum, 1935: 576 (d).

Neonyssus (Vitznyssus) squamosus, Castro, 1948: 277.
Ptilonyssus .(Rhinonyssoides) squamosus, Pereira and Castro. I94’9:’222.
Neonyssus squamosus. Pain, 1957: 48. ’

’,,’"
Locality: Islands of the Atlantic Ocean - West Indies.
" ’
.’ :"

Host: Bird,- Sericotes holosericeus. ." ’

.
. . Rhinonysaoides strandtmanni (Fain), New combination
.
.Type material ;in Musee Royal du Congo Beige.
Ptilonyssus strandtmanni Fain. 1956: 148 (d, ?); Fain, 1957: 86 (illus.).
Locality; Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Apus caffer streubeli.

Genus Vitznyssus Castro, 1948

. ....

Type:
. (Fig. 88)
Dermanyssus nitzschi Giebel. By designation of Cas-
tro, 1948.
Astridiella Fain, 1357: 148; 1957: 91.
., Type: .Ptilonyssus scotornis Fain, 1956.
Diagnosis: Large, elongated nasal mites of caprimulgiforna and
certain gruiiform birds. Female .chelicera bulbous basally, greatly at-
tenuated and with minute chelae. Only one dorsal plate present, the
podosomal. With a large, well formed (although very poorly sclerotized)
174

tritosternum. Stigma, with short perit-reme..

Remarks: According to Vitzthum (1938) Nitzschwas probably the
first person .to find a bird nasal mite although Nitzsch did not recognize
it to be anything unusual and did not describe it. Later Giebel (1871)
recognized this as a new discovery, and described the mite, naming it
Permanyssus nitzschi in honor.of the original discoverer. The host was
Caprimulgus europeaus,, one of the nightha’wks.
In l935Vitzthumnotedthat nitzschi was not of the genus Dermanys-
sus and moved it. to the endoparasitic genus Rhinqny-ssus. At the same
time he also assigned to-this.-species a nasal mite recovered from the
european bustard, Otis tarda. He explained at. some length why he be-
lieved the bustard mite and the nighthawk mite to be the same species
although the two hosts were .entirely unrelated.
In 1948, Castro recognized that nitzschi was not congeneric with
the generotype of Rhinonyssus ..and.- moved the mite to the -genus Neonys-
sus, erecting for it a new subgenus, Vitznygsus.

.
.
In 1949 Pereira and Castro realized that .Neonyssus was not the
proper assignmentandplacedit in Ptilonyssus, subgenus Rhinony s s oides
Hirst. They contended that Vitznyssus Castro was not really distinct
from Rhinonyssoides Hirst and sank Vitznyssus into the synonymy. ,
Recently, Pain (1956, 1957) recovered nasal mites from caprimul-
giform birds and fqund.them .to be quite distinct in possessing a well de-
veloped tritosternum. He proposed for them the generic name Astridi-
ella, apparently unaware of the. earlier name, Vitznyssus Castro, 1948.
The Rules of Zoological .Nomenclature s.tate that generic-group names
are/coordinate. Vitznyssus .Castro is the earliest available name and
Astridiella Fain, 1957 must be a junior synonym.
Fain also had proposed (19S7: 93) that, because the type,specimen
ofDermanyssus nitzschi is lost and that the description isinqn-diagn-ostic,
the name nitzschi Giebel should be deleted from the literature. This is
both impossible and undesirable; impossible under, the Rules and unde-
sirable because it would ’destroy the first name given to a bird nasal
mite. It seems that the logical solution would be,to use the available
names and to designate’a iectotype to replace the lost. type ofGiebel’s.
Ordinarily the Iectotype would be the species, on which Fain. based
his genus Astridiella, i. e^., Astrj.dielia scotornis (Fain) <=Ptilonyssu3
scotornis Fain), but this is deemed undesirable by us because Fain has
designated two types for this species, "Holes: Scotornis fossil welwit-
schi Boc. a Muhero, pres .d’Astrida, en decembre 1955 (ancien type).
a 1’Akanyaru (Ruanda Urundi) Ie 3-3-1956 et en fevrier 1956; Caprimul-
gus tristigma Rupp ’a l’Akanyaru;le. 6-3-1956(oouv.eautype cidessus). .. "
(Fain, 1957: 96). It would seem more desirable to designate as Iecto-
type a species that has only one type designation and in which the type
host is identical with the host of..the original species. This would be
Fain’s second species of Astrldiella, A_. .caprimulgi [Pain, 1957; 98,
"Hotes: Caprimulgus .europaeus L. a 1’Akanyaru Ie 24-2-1956 (type) ..."].
Hence: Dermanyssus nitzschi Giebel becomes Vitznyssus nitzschi
(Giebel); Aatridiella Pain, .1957 becomes a,synonym of Vitznyssus Cas-
 175

tro, 1948; Astridiella caprimulgi Fain, 1957becomes a synonym of Vitz-

nyssus nitzschi (Giebel); and the type specimen of_A. caprimulgi Fain
becomes the lectotype of Vitznyssus nitzschi, by present designation.
The type is in the Musee.Royal dii Congo Beige.
This still leaves the,problem of the proper.identity of the mite des-
cribed by Vitzthum (1935: 572) from the bustard. Everyone who has
studiedthts species, including Vitzthum, has remarked upon the improba-
bility of the species from the nighthawk and the species from the bustard
being identical. But without actual specimens as evidence, a decision
can not be made. .Recently Pain (1957; 98) reported the recovery of a
mite from an African bustard, Neotis cafra, which is strikingly similar
to that from the nighthawks. .This also seems unusual, but in the face
of such evidence, we must accept the mites as being; congeneric and ac-
cept Vitzthum’s original position. However, the new lectotype for nitz-
schi is distinct fromVitzthuna’s description of nitzschi based on the Otis
tarda mite so that we must accept Vitzthum’s mite as only congeneric
with nitzschi Giebel, not identical with it. Also the new mite from the
African bustard does not fit Vitzthum’s description in all respects; so,
as Fain pointed out (1957: 95), the nitzschi of Vitzthum from Otis tarda
seems to be a distinct species and without a name. Fain (ibid.) proposed
the name vitzthumi, a choice that we readily accept,

Vitznyssus nitzschi (Giebel). New combination

Dermanyssus nitzschi Giebel, 1871: 31 (d).

Rhinonyssus nttzschi, Vitzthum. 1935: 572. In part.
Neonyssus (Vitznyssus) nitzschi, Castro, 1948: 277.
Ptilonyssus (Rhinonyssoides) nitzschi, Pereira and Castro, 1949: 222.
In part.
Ptilonyssus nitzschi, Zumpt and Till, 1955. In part.
Astridiella caprimulgi Fain, 1957; 96 (d, illus.). New lectotype.
Localities: Africa - Belgian Congo, Ruanda-Urundi. Europe.
Hosts: Birds - Caprimulgus europaeus, Cosmetornis vexillaris,
Scotornis fossii welwitschi.
Remarks: See under discussion of the genus.

Vitznyssus neotis (Fain). New combination

Astridiella neotis Fain, 1957: 98 (d, illus.); 1957: 127. :

Locality: Africa - Belgian Congo, Astrida.

Host: Bird - Neotis cafra.

Vitznyssus scotornis (Fain). New combination

Type rraterial in Musee Royal du Congo Beige.

Ptilonys sus s-cot orhis Fain. .1956: 148 (d, $); 1957: 95 (illus.); 1957: 125.
Locality: Africa - Belgian Congo, Ruanda-Urundi,
Hosts: Birds Caprimulgus europaeus, Caprimulgus tristigma,
-
176

Scotornis fossil welwitschi. ; ,:. ’,’ . : "’.-’ .

Remarks:-’In the ori’ginai.description (1956) -Fain designated the type

host Scotornis fossil welwitechi, .but in a later publication (1957: 125)
stated that ’a’new type had" be en chosen, from Caprimulgustristigma.

Vifaznyssus vitzthumi.(Fain). N’ew combination

Rhinonyssus nitzschi, Vitzthum; 1.935-.-572 (d, illus.). (In part; not Der-
manyssus nitzschi Giebel.)
. ..; .
"
Astridiella Vttzthumi Fain, 1957:.’,95,’ 127; .
-Locality: Europe. ;.<.

.
.

Host: Bird - Otis tarda.

.RemarRs: See under discussion of the genus.
.
. :.. Genus Paraneonyssus Castro, 1948 ’

....... .
:’"
’
(Pig., 89).. ;"^
’!
.: ’ Type: Neonyssus (Paraneonys:sus)..enrietti Castro.
.
Diagnosi’s: Mites of medium size; two dorsal plate’s, the’posterior
longer but-narrower than. the anterior; chelicerae barely or not at all
attenuatedi -efielae very small... Parasitic only in passeriform birds.

Paraneonys sus enrietti (Castro).’- New. combination

Neonyssus (Paraneonyssus) enrietti CaStr.o, 1948: 274. :^"-’.

Ptilonyssus (Paraneoayssus) .enrietti, P.ereira and Castro, 1949: 227’.,’.
Ptilonyssus enrietti, Zumpt’and Till,. 1955.; Fain, 1957: 89. j.Ti
Locality: South America - Brazil, i State, of Parana;.
; Host:’ Bird - Phylloscartes veritralis. .;;
.
.
.
.
Paraneonyssus astridae. (Fain) \ .New. conibination

Type-material in Musee Royal du Congo Beige,

Ptilonyssus astridae Fain,’ 1956: 141 (d, ?); .1957: 98(illus.).
Locality: Africa - Belgian Congo, Ruanda’Urundi.
Hosts: Birds - Amandava subflava, Estrilda paludicola, Granatina
granatina, Lagonosticta rhoaopareia (type host),..-Lagonosticta rubricata
cong’j.ca, I-.agonosticta senegaia ruberrima, Lonchura bicolor poensis,
Pytilia m&lba belli, TJra.eginthus .bengalus. ’ ’’ .’"

Paraneonyssus capehsis (Zumpt and Till).;.. New combination

Type material in the South African Institute’.io.r. Medical Research.

Ptilony^aus capensis Zump’f and Till, 1955: 72 (d, 9, illus.); Fain, 1956:
./ ’’.
’
138; 1957: 89 (illus.). ..
Locality: Africa Belgian Congo, Ruanda-’tJrundi; Union of South
Africa, Transvaal. .’ .. .: ’’:i-’..
.
Hosts: Birds - Macr.onyx capensis, Macrehyx..CQrceus.
 177

; . ! Paraneonyssus capita.tug Strandtmann . . -

Type material ip the U. S. National Museum. .; . ,

.
.
.
Paraneonyssmg capitatus. Strandtmann, 1956:. 133 (d, ?..iUus,). ’

Locality: North America - United States, .Utah. . . .

Host: Bird - Otocoris alpestris. :: . .

.
.

Paraneonyssus dryoscopi (Zumpt and Till). New combination

Type material in the South African Institute for Medical Research.

Ptilonyssus dryoscopi. Zumpt and Till, 1955: 1’39 (d, .?,. illus..). Fain,
1956: 139; .1957; 89 (illus.). ..

.
Locality: Africa - Belgian Congo, Ruanda-Urundi; Union of South
Africa,. Cape..Province,

.
Hosts: Birds - Dryoscopus. cubia (type host). Tchagra .senegala,

Paraaepnyssus emberizae (Fain). New combination

Type material in Musee’Royal du Congo Beige.

,
Ptilonyssus emberizae Fain. 195’6:;-l40 (d, rf and ?); P.ain, 1957: 95 (il-
lus.).
Locality: Africa - Belgian Congp, Ruanda-Urundi.
Host: Bird - Emberiza.flaviventris. .
.

Paraneonyssus hirsti (Castro and Pereira), New combination

.(Fig. 89)
Neonyssus hirsti Castro and Pereira, 1947: 129 (d); Porter and Strandt-
mann, 1952.
Neonyssus (Neonyssus) hirsti, Castro, 1948: 268.

.
Ptilonyssus (Paraneonyssus) hirsti, Pereira and Castro, 1949.
Ptilonyssus hirsti. Zumpt and Till, 1955; Fjiin, 1956: 1.40; 1957:. 86.
Ptilonyssus nudus Trouessart, in part; the male as described by Ber-
lese, 1889; the female as described by Hirst. 1923.
Localities: Africa - Belgian Congo, Ruanda-Urundi. Europe-Eng-
land; Italy; Portugal. North America - United States, Texas. South
America - Brazil.
Hosts: Birds - Passer domesticus. Passer ugandae griseus.

Paraneonyssus icteridius Strandtmann and Furmaa

Type material in the U. S. National Museum.

Paraneonyssus icteridius Strandtmann and Furman. 1956: 167 (d, d", $,
and nymph illus.).
Ptilonyssus icteridius. Fain, 1957: 96.
Locality: North America - United States, California, Texas.
Hosts: Birds - Agelaius phoeniceus, Agelaius tricolor, Euphagus
cyanocephalus, Molothrus ater (type host), Piranga ludoviciana, Quis-
calus quiscula, Sturnella magna, Xanthocephalus xanthocephalus.
:178

Paraneonyssus ploge anus-(Fain)... New’ combination

Type material in Musee Royal du.Congo Beige.

Ptilonyssus ploceahus Fain, 1956: 139 (d, 9); 19.57; 89 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Birds - Anaplectes rubriceps (type host), Euplectes orix,
Hyphanturgus ocularius, Quelea quelea centralis, Textor atrogularis.

Paraneonyssus serini (Fain), New combination

Type material’in Musee Royal du Congo Beige.

Ptilonyssus serini Fain. 1956: 139 (d, S); 1957: 96(Uius.).
’

Locality: Africa -’ Belgian Congo, Ruanda-Urundi.

Hosts: Birds - Serinus mozambicus bar-batus (type host), Serinus
.canieollis sassii, Serinus citrinelloides fronfailis. i !

Paraneonyssus travassosfilftoi (CastrO), New combination

Neonyssus (Paraneonyssus) trayassosfilhol Castro, 1948: 275.

Ptilonysaus -<Paraneonyssus) travassosfilhoi, Perefra and Castro, 1949:
227. ’
Ptilonyssus travassosfilhoi, Zumpt and Till, 1955; Fain,’1957: 88.
Locality; South America - Brazil. State ot Sao Paulo.
Host: Bird - PhUydor lichtensteini.

Paraneonys sus zumpti (Fain). New combination

Type material in Musee Royal du Congo Beige.

Ptilonyssus zumpti Fain. 1956: 138 (d. ?); 1957: 91 (illusj.
Locality: Africa ’’Belgian Congo, Ruanda-Urundi. . ’;
Host: Bird - Amblyospiza albifrons montaria.

Genus Ptilonyssus Berlese and Trouessart, 1889 ’

- "
(Fig. 90)
Type: Ptilonyssus echinatus Berlese and Trouessart.
Diagnosis: Small.to medium sized mites. Female with a large pro-
podosomal plate and a small pygidial plate; chelicerae attenuated and with
very small chelae; body elongated. Male with’-two large dorsal plates
and rather short, non-attenuated chelicerae. Parasitic only in passer-
iform birds. .

Ptilonyssus echinatus Berlese and Trouessart

Ptilonyssus echinatus Berlese and Trouessart, 1889; Vitzthum, 1935;

Castro, 1948; Pereira and Castro, 1949; Zumpt and Till, 1955: 74
(cT.and?, illus..); Fain, 1956: 146; 1957; 114.; ’:’’ "
Localities: Africa - Belgian Congo, Ruanda-Urundi; Union of South
 ; . . 179

Africa, Cape Province, Johannesburg. Europe-Germany. North Amer-

ica- United States. .

Hosts: Birds - Cecropis senegalensis, Hirundo smithi, Hirundo

rustica, Petrochelidon spilodera.
Remarks: Hosts from which we have taken the mite. in the United
States include: Hirundo rustica, Iridoprocne bicolor, Petrochelidon pyr-
rhonota, and Riparia riparia; these are previously unpublished records.

Ptilonyssus andropadj Fain

Type material in Musee Royal du Congo Beige.

Ptilonyssus andropadi Fain, 1956: 14T’(d.. ); 1957: 1J.4 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Hosts: Birds - Andropadus latirostris (type host), Phyliastrephus
fischeri.

Ptilonyssus aureliani Fain

Type material in Musee Royal du Congo Beige.

Ptilonyssus aureliani Fain. 1956: 146 (d, 9); 1957: 109 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Prinia subflava.graueri. .

.
Ptilonyssus calamocichlae Fain

Type material in Musee Royal du Congo Beige.

Ptilonyssus calamocichlae Fain, 1956: 142 (d. $); 1957: 114 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi,
Hosts: Birds - Apalis flavida, Calamocichia rufescens (type host),
Cisticola erythrops, Hippolais icterina, Prinia leucopogon, Serinus mo-
zambicus.

Ptilonyssus chloroclchlae Fain

Type material in Musee Royal du Congo Beige.

Ptilonyssus chloroeichlae Fain, 1956: 142 (d, 9); 1957: 114 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Chlorocichia flavicollis.

Ptilonyssus desfohtainei Zumpt and Till

Type material in the South African Institute for Medical Research.

Ptilonyssus desfontainei Zumpt and Till, 1955: 76 (d, <f and ?. illus.);
Fain. 1957: 84,
Locality: Africa - Union of South Africa, Transvaal.
Host: Bird - Thamnolaea cinnamomeiventris.
180

Ptilonysaus japuibensis Castro

Ptilonyssus (Ptilonyssus) japuibensis Castro, 1948: 264 (d); Pereira and

Castro, 1949; Zumpt and Till, 1955: 89; Fain. 1957.
Locality: South America - Brazil, State of Rio de Janeiro.
Hosts: Birds - Ramphocelus carbo centralis, Ramphocelus bresil-.
ius dorsalis.

Ptilonyssus lanii Zumpt and Till

Type material in the South African Institute tor Medical Research.

Ptilonyssus lanii Zumpt and Till, 1955: 79(d, d’and$, illus.); Fain, 1856:
143; 1957: 103 (illus.). ; .
..
,,.
.
Locality: Africa - Belgian Congo, Ruanda-Urundi; Union of South
Africa, Cape Province, Transvaal.
Hosts: Birds -Alseooax adustus, Calandrella cinerea, Campephaga
phoenicea, Cossypha heuglini, Geokichia piaggiae, Lanius collaris (type
host), Lanius collurio, .Lanius excubitorius, Muscicapa aquatica, Mus-
cicapa striata, Platystetra cynea, Poliospiza striolata.

Ptilonyssus motacillae Fain

Type material in Mus^e Royal du Congo Beige.

Ptilonyssus motacillae Fain, 1956: 143 (d, $); 1957: 104 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi. .

Hosts: Birds -Cercomela fanailiaris, Motacilla, aguimp (type host),

Motacilla capensis, Motacilla flava, Oenanthe oenanthe, Saxicola ru-
betra., Saxicola torquata.

Ptilonyssus nudus Berlese and Trouessart

(Fig. 90)
Ptilonyssus nudus Berlese and Trouessart, 1889: 128; Hirst, 1916; Vitz-
thum, 1935; Radford, 1939; Buitendrjk, 1945; Castro and Pereira,
1947; Castro, 1948; Pereira and Castro, 1949; Porter and Strandt-
mann, 1952 (c, d); Zumpt and Till, 1955: 79; Bregetova, 1956: 198;
Fain, 1956: 144; 1957.
Localities: Asia -..U.S..S.R. Europe - France; Italy; Portugal;
U.S.S.R. North America - United States, Texas.
Hosts: Birds - Emberiza cirlus, Pringilla coelebs, Hirundo rustica,
Hypochera funerea. Merganser sp., Muscicapa striata, Parus atrica-
pillus, Parus major. Passer domesticus (type host). Passer grisea,
Riparia riparia, Serinus canaria, Spinus spinus, Sturnus vulgaris.
Remarks: All the nonpasserine hosts listed above are from Brege-
tova (1956). Wethink it is possible that the mites from those hosts may
have been misidentified.
 181

Ptilonyssus olaioi Pereira and Castro

Ptilonyssus (Ptilonyssus) olaioi Pereira and Castro. 1949: 219(d); Zumpt

and Till, 1955; Fain, 1957. .:’.
Locality: South America - Brazil, State of Sao Paulo.
Host: Bird - Ampelion cucullatus. :’

’
Ptilonyssus orientalis (Swing)

Type material in the U. S. National Museum.

Sommatericola orientalis Ewing, 1933: 13 (d).
Sternostomum orientalis, Vitzthurn, 1943: 657.
Rhinonyssus (Rhinonyssus) orientalis, Castro, 1948; 257.
Sternostoma orientalis, Zumpt’and Till, ’1955: 91.
Ptilonysaua orientalis, Strandtroann, ’1956: 136 (t).
Neonyssus orientalis. Pain, 1957; 57.
Locality: Asia - Siam.
Host: Bird - Lanius nigriceps longicaudatus.
.
Remarks: We examined the type specimen in the summer of 1954.
:
It is, without question, a species of Ptilonyssus.

Ptilonyssus orioli Fain

Type material in Musee Royal du Congo Beige.

Ptilonyssus orioli Fain, 1956: 144 (d, 9); 1957: ll4(illus.).
Locality: Africa - Belgian Congo, Ruanda=Ururidi.
Hosts: Birds - Oriolus larvatus, Oriolus oriolu’s.

Ptilonyssus phyllastrephi Zumpt and Till

Type material in the South African Institute for Medical Research.

Ptilonyssus phyllastrephi Zumpt and Till, :i955: 78 (d, $ illus.); Pain,
1957: 104 (illus.).
Locality: Africa - Union of South Africa, Cape Province.
Host: Bird - Phyllastrephus terrestris.

Ptilonyssus psalidoprocnei Pain

Type material in Musee Royal du Congo Beige.

Ptilonyssus psalidoprocnei Fain, 1956: 146 (d,. 9); 1957; 117<illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi;
Hosts: Birds- Psalidoprocne albiceps, Psalidoprocneholomelaena.

Ptilonyssus pycnonofi Pain

Type material in ’Musee Royal du Congo Beige. :

Ptilonyssus pycnonoti Fain, 1958: 146 (d, $); 1957: 109 (illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
182

Host: Bird’’- Pycnbhotus barbatus.

Ptilonyssus ruaridae Fain . ’; ,’

Ptilonyssus-ruandae Fain, 1956: "395 (d); 1957: 109 (illus.).

.
Locality: Africa - Belgian Congo, Akanyaru.
Host: Bird - Zpsterops senegalensis.

Ptilonyssus sairae Castr-o

Ptilonyssus (Ptilonyssus) sairae Castro, 1948: 260 (d); Pereira and Cas-
tro, 1949; Zumpt and Till, 1955: 89; Fain, 1957. " :"

.
Locality:’ South America - Brazil, State of Sao Paulo. ...-.;
Host: Bird - Tangara seledon. .-."

Ptilonyssus stresemanni Vitzthum

Ptilonyssus stresemanni Vit’zthum, 1935: 579 (d); Castro.. 1948; Pereira

and Castro, 1949; Zumpt and Till, 1955: 89; Fain, 1957.

.
Locality: South America - Brazil.
Host: Bird - Procnias alba.

Ptilonyssus terpsiphonei Pain

Type material in Musee Royal du Congo Beige.

Ptilonyssus terpsiphonei Fain, 1956: 145 (d. 9); 1957: 104(illus.).
Locality: Africa - Belgian Congo, Ruanda-Urundi.
Host: Bird - Terpsiphone viridis (type host), Trochocerus cyanome-
las.

Ptilonyssus viduae Fain

Type material in Musee Royal du Congo Beige. .

Ptilonyssus yiduae Fain, 1956: 147 (d, 9); 1957: 120 (iilus..).. .

Locality: Africa - Belgian Congo. Ruanda-Urundi,

Host: Bird - Vidua macroura.

Family SPINTURNICIDAE Oudemans. 1.901

Description of the family. Dark brown mites with heavy, strong

legs, and very strong claws. Tritosternum reduced to a small plate or
entirely lacking; modified palpal claw lacking (or at least not forked).
Ventral plates reduced; one or two dorsal plates. Found only as ecto-
parasites of bats. Apparently all specie.s are ovoviviparous,.
.
.

Remarks: The mites of this family have an unmistakable tacies and

can be instantly recognized as spiriturnicid. This pronounced similarity
 183

of form has caused taxonomic difficulty at the generic and specific levels..
No less than-seven-generic names have been proposed for what is per-
haps really only one genus, although two genera are recognized here.
As far back as 1910, Oudemans gave a. page and a half of synonyms, for
Spinturnix vespertilioni-s alone, a. name that has been ruled invalid by
the International Commission.

.
We.list the species we have .found in the literature without any claim
at complete coverage or of correct or complete synonymy.

Key to the Genera of Spinturnicidae

(Modified after Zumpt, .1951)
.
Peritreme bent from the dorsal side to ventral side. Opistho-
soma not strikingly expanded in the engorged female. Pre-
sternal plate usually distinct,: rarely totally reduced . . Spinturnix

Peritremes confined to the dorsum. Opisthosoma strongly ex-

panded in the engorged female. Presternal plate, ’.always want-
ing . . . . . . . . . . . . . . . . . . . ... . . . .;. Periglischrus

Genus Spinturnix von Hey den, 1826

(Pig. 91)
Type:. Spinturnix myoti Kolenati, 1856. .

Pteroptus Dufour, 1832; 98.

Type: (Pteroptus vespertilionis Pufour; not Acarus vesper-
tilionis Scopoli) = Pteroptus murinus Walkanaer, ~

, ,..., ,. .., 1847: 545. .

.
Ancystropus .Kolenati, 1856.
Type: Ancystropus zeieborii Kolenati, 1856.
Diplostaspis Kolenati, 1857. (Type unknown to us.)
Leiostaspis Kolenati, 1657. (Type unknown to us.)
Meristaspis Kolenati, 1857.
. Type: Meristaspis lateralis Kolenati, 1857.
,

Diagnosis: Spinturnicids in which the Opisthosoma is shorter than

the podosoma, and has no lateral expansions.
Hemarks: Originally (according to Oudemans (1936: #153)] the type
of this genus was given as Acarus vespertilionis Scopoli, 1.763, but Aid-
rich (1936) states that. underOpinion 128 of the International Commis-
sion on Zoological Nomenclature, the specific name vesperttlionis is
invalidated for a long list of mite genera, including Spinturnix and its
synonyms. Consequently, myoti Kolenati, being the earliest available
name, was chosen.

. .’ . Spinturnix myoti Kolenati

(Fig. 91a);
Acarus vespertilionis Scopoli, 1763. (See discussion under the genus.)
184

Spinturnix vespertilionis, authors;;. Oudemans, 1910: 67; Hirst, 1927;

Oudemans, 1936: #153;’Radford, .1939; Bregetova, 1953; 1956: 204;
. WUlmann, 1955:. 183. ,
. , .

.
.
Spinturnix myoti Kolenati,. 1856; Aldrich, 1936: 29.
.Localities: Africa. Asia - U.S. S.R. Europe..

.
Hosts: Mammals - Various species of Chi’roptera, the most .com-
monly parasitized forms, being Myotis spp., but according to Bregetova
(1956) including also Plecotus auritus, Rhinolophus ferrum-equinum,
Vespertilio seratinua.
Remarks: The above references are only the highlights. Oudemans
(1910; 1936) gives the. most .complete summary of references and syno-
nyms. Aldrich (1936) gives the reasons for,.invalidating the name ves-
pertilionis.

Spinturnix abyssinicus Hirst

Spinturnix abyssinicus Hirst, 1927: .333 (d)-.

Locality: Africa - "Abyssinia. " ;

Host: Mammal - "Bats."

Spinturnix acuminatus (l^oon) . .

Pteroptus acuminatus Koch, 1836: 21; Oudemans, 1936: #153.

Spinturnix acuminatus. Hirst, 1927; Radford, 1950.
Locality: Europe.. ..
Host: Mammal - "The-N-octule bat."
Remarks; Oudemans (1936) gives this as a synonym of vespertili-
onis (=myotis) but Radford lists it as a good species.

Spinturnix aethiopicus (Hir.st) .

.
Ancystropus aethiopicus Hirst, 1923: 979 (d); Radford, 1950.
Spinturnix aethiopicus, Zumpt, 195.1: 80.
.
Locality: Africa - "Gambia"; Zanzibar; ;

Hosts: Mammals - Epomophprus minor, Micropteropus .pusillus.

Remarks: Hirst made this.a new species questionably, and stated
that it was possible that it was the same as Kolenati’s A. mulleri from
Pteropus vulgaris. Note: mulleri is a synonym of vespertilioriis (= my-
otj. ).

Spinturnix amboinensis Oudemans

Spinturnix ambolnensis Oudemans, 1925: 26 (d)’;’Radford, 1950.

Locality: Indonesia - Dutch East Indies, Ambon.
Host: Mammal - "Bat."
 185

Spinturn.ix ajriericanus (Banks)

Pteroptus americanus Banks, .1902: 173. :

Spinturnix americanus, Rysgaard, 1942:. 256; Yunker, 1958.

Locality: North America - United States, Indiana, Minnesota,
: .Hosts:.. Mammals,- E-ptesicusfuscu.s, Myotis .lucifugus/’bats."

.. .
. .
,. Spinturnix antipodjanus Hirst
.
.
Spinturnix antipodianua Hirst, 1931: 563 (d); Radford, 1950.
Locality: Australia - North C’ue’ensland. :
Host: Mammal - "Bat.". , .

.
Spinturnix araguensis Vitzthum

Spinturnix araguensis Vitzthuna. 1931: 34 (d); Radford, 1950.

^
Locality: South America - Venezuela.
Host: Mammal Myotis nigricans.
-
Spinturnix artibi-ensis Radford

Spinturnix artibiensis Radford. 1951: 97.

Locality: Islands of the,Atlantic Ocean - British West Indies. Trin-
idad.
Host: ..Mammal - Artibeus jamaicensis- palmarum.
Spinturnix calcaratus (Hirst)

AHcystropusdMeristaspis) calcaratus Hirst. 1923: 983 (d); Radford, 1950;

Zumpt. 1950. .. ..

Spinturnix calcaratus,’ Zumpt. 1951: 80. .

.
Localities: Indonesia - Dutch East Indies. Islands of the Indian
Ocean - Madagascar. ; .

Host: Mammal - Pteropus rufus, Pteropus sp.

Spinturnix,car loshoffmannj: Hoffmann

Spinturnix carloshoffmanni Hoffman. 1944; 185. M); 1944: 261; Radford,

’, .1.950. ...
Locality: North America’- Mexico, Federal District.
Host: Mammal - Natalua mexicanua.

Spinturnix echinipes (Banks)

Pteroptus echinipes Banks, 1910: 5.

Spinturnix echinipes. Stiles and Nolan, 1931: 708; Yunker, 1958.
Locality: North America - United States. New York.
Host: Mammal - Myotis lucifugus.
186

Spinturnix euryalis (Cane strini)

Pteroptus euryalis G. Canestrini, 1885. (Reference not seen.)
Spinturnix euryalis. Hirst, 1927: 323 (d); v.an Eyndhoven. i94l: 44 (1)-,
Radford, 1950; Turk, 1952. :
Locality: ’Europe - Corsica; England; Italy.
Hosts: Mammals - R-hioolophus euryale, Rhinolophus ferruro-equin-
um.
Remarks: VanEyndhoven(1949) discusses fully the taxonomic status
of this species..

Spinturnix ewingia Wharton

Type material in the U. S. National Museum. :
Spinturnix ewingia Wharton. .1938: 146 (d); Hoffmann, 1944; Radford,
’
"’: "
1950. .:
Locality: North America - Mexico, Central and Southern Mexico.
Hosts: Mammals - Artibeus jamaicensis, Leptonycteris niva.lis.
Spinturnix grossus (Banks)

Pteroptus grossus Banks, 1910: 6..

Locality: North America - United States, New Mexico.
Host: "Bat." ..
.
Spinturnix ianaai Lombardint . ,.; .... :

Spinturnix yespertilionis ianzai. Lombardini, 1944: 5

"
(d),JR.adford, 1950.
Locality:’ ?
Host: Mammal - Khinolophus sp,

Spinturnix iowae Keegan

.
.
Type material in the U. S. National Museum.
Spinturnix iowae Keegan, 1943: 55 (d); Radford. 195.0; .Morlan, 1952;
Yunker. 1958.- :..’ ’
Locality: North America - United States, Georgia, Iowa. Maryland,
New Jersey, ’
.
.:
Host: Mammal Myotis lucifugus.
-

Remarks: The possibly synonymy 6f this species with Banks’ amer-

.icanus and echinipes should be investigated (Yunker, 1954). Banks’ des-
criptions and drawings are non-diagnostic, and all. three parasitize the
same host species within the same geographical area. Keegan (1943)
compared iowae with ’araguensis Vitzthum, a’Venezuelan species, but
did not attempt to delineate it from the North American forms.

Spinturnix javensis Oudemans

Spinturnix javensis Oudemans, 1914: 85; Oudemans, 1951: 175 (d); Rad-
.....
’
ford, 1950. . .

Locality: Indonesia - Dutch East Indies; Malang. ^ ;;

.
Host: Mammal - "Bat. "
 .187

Spinturnix Jordan! (Radford) .;

Type material in the British Museum (Natural History).

Ancystropus jordani Radford, 1947: 309 (d); Radford, 1950.
Locality: Indonesia - Celebes.
Host: Mammal - "Bat. "
Spinturnix kenyaensis (Radford)

Type material in the British Museum (Natural History).

Atlcystropus kenyaensis Radford, 1947: 305; Radford, ,1950.

.
Locality; Africa - Kenya.
; Host: Mammal - Eidolon helvum.
Remarks: Zumpt (1951: 80) regarded kenyaensis as a synonym of
S. lateralis but we are retaining it here.

Spinturnix lateralis (Kolenati)

Type material in the British Museum’(Natural. History). .

Pteropus lateralis Kolenati, 1856: 29. (Reference not seen.)

Meristaspis lateralis, Kolenati, 1857: 29.
Ancystropus (Meristaspis) lateralis. Hirst, 1923: 985 (d); Radford, 1950.
Spinturnix lateralis, Zumpt, 1951: 80.
Locality: Asia - Palestine.
Host: Mammal Pteropus aegyptiaca.
- .
.
Remarks: See under S. kenyaensis.

Spinturnix lavellanua (Radford)

Type material in the British Museum (Natural History).

Ancystropus lavellanus Radford, 1951: 97 (d). . .
Locality: Islands of the Pacific - Vella Lavella.
Host: Mammal - Pteropus lavellanus.

Spinturnix lawrencel Zumpt

Type material in the Natal Museum. South Africa; also in the South
African Institute for Medical Research.
Spinturnix lawrencei Zumpt, 1951:’ 81 (d).
Locality: Africa - Union of South Africa, Natal.
Host: Mammal - Myotis tricolor.

Spinturnix macroglossi (Hirst) ., ..

.
.
.
Ancystropus’ (Meristaspis)’ macroglosai Hirst, l:923i 981 (d); Radford,
1950. .

Locality. ? GUolo.
Host: Mammal - Macroglossus minimus.
188

Remarks: Hirst gave.only the .name "Gilolo’-’.as a locality. However,

the host is restricted to the Malay region and, unless it was a zoological
garden resident, :ths mite.will probably be .found’, there ..’ .’.: :

Spinturnix murinus-CWalckenaer.)

Pteroptus murinus Walckenaer, 1847: 545 (d). (Reference not seen.)

Spinturni.x murinus, Oudemans, 1910:;67(t); Trag&rdh, 1912; Oudemans,
1936: <fT51 (t); Radford, 1950.
Locality:. Europe - Prance; (3ermaay;.N;etherland-s; Spain; U. S. S. R.
Host: Mammal- - ’Myotis my 6tis. .’;. .-. ;. ;..;, ’,. ..: :
. . .

.
Remarks: Oudemans (1910) gives a page of’synonymy for this mite,
covering the period from 1832-19.02.- ;: .: :.. ; ; ."

Spinturnix aovae-holtandae Hirst .. .’

.
Type material in the ; collection of-the South’ Australian Museum.
Spinturnix novae-hollandae Hirst, 193i: 564 (d); Radtord, 1950.
Locality: Australia - South Australia. .: .; ,. ; .’

.
Host: Mammal "Bat.." :; ’. :..’.
’
. -’ : ’::

.
.
,
.
, . :’ Spinturnix :omahonyi Turk.

Type material in the National Museum of Ireland, Dublin.

Spinturnix omahonyi Turk, 1,945:: .80,9, (d); Radford.- 1950.
Locality: Burope - England. ;: ’ .. . . .
.
.
Host: Mammal - Rhinolophus hipposideros minutus.

Spinturnix oudemansi van Eyndhoven

.Spinturnix oudemansi van .Eyndhoven,; 1941: 44 (d); Radford. 1950.

Localityr Europe. , ;.;’’
Host: Mammal - Rhinolophus ferr.um-equinum.

Spinturnix pipistrellus Radford

Spinturnix-pipistrellus Radford, ,1.951: 99. (d)..

Locality: Africa Sierra JLie’one.- .
’

. .
.
.
Host: Mammal - Pipistrellus stampflii.. ’.
.
Spinturnix plecotihus (Koch)

Pteroptus plecotinus Koch, L.838. .; ...

Spinturnm: plecotinus, Oudeinans, 1910: 67 (t); Hirst, 1927: 331 (d);
Turk, 1945:. 7@.6 (d, .<?);.Radford,. 1.950; ..
Pteroptus transversus Kolenati (vide Hirst, 1927).
Locality: Europe - "Croatia"; England; Germany; Netherlands.
Host; Mammal Ple.cotus auritua. ’.
-
.
 189

Spinturnix psi Kolenati

Type material in the British Museum (Natural History).

Spinturnix psi ’Kolenati, 1857; Hirst, 1927: 328 (d); Radford, 1950-.
Locality: Europe. :

’
’

’ ’Hosts: Mammals - Mirnopterns- schreibersi: ’(type host), Myotis sp.;

RKiholophus ferrum-equxnum. ’ ’’’’

’
’
Spinturnix scotdphili Zumpt and Till

Type material in the South African Institute’ for Medical Research.

Spinturnix scotophili Zumpt and Till, 1954: 214 (d).
Locality: Africa - Sudan.
Host:. Mammal - Scotophilus murinoflavus.

Spinturnix semilunaris DeMeillon and Lavoipierre

Type material in the South African Institute for Medical Research.

Spinturnix semilunaris DeMeillon and Lavoipierre, 1946: 79 (d); Zumpt,
1950; Radford. 1950;’’Zumpt, 1-951: 80.
Locality: Africa - Union of South Africa. Transvaal.
Host: -Mammal - Minibpteru& natalen’sis.

Spinturnix viduus Zumpt

Type material in the South African Institute for Medical Research.

Spinturnix viduus Zumpt, 1950: 92 (d); Zumpt, 1&51: 80.
Locality: Africa - Union of South Africa, Transvaal.
-
Host: Mammal Rhi-nolophus geoffroyi. ’

’. . ’"’
.

Spinturnix walkerae Zumpt and Till

Type material in the South African Institute for Medical Research.

Spinturnix walkerae Zumpt and Till, 1954: 216 (d).
:
Locality; Africa - Kenya, Muhoroni.
Host: Mammal - Pipistrellus nanus nanus.

Spinturnix zeiebori (Kolenati)

Ancystropus zeiebbri Kolenati, 1856; 1857: 23; Radford. 1950.’

Spinturnix zeiebori, Zumpt. 1951: 80.
Locality: Africa"- Egypt. :
Host: Mammal - Rhinopoma microphyllum.

Genus Periglischrus Kolenati, 1857

. (fig. 92)
’
.

Type: P-eriglischrus caligus Kolenati.

190

Diagnosis: Spinturnicids in which the opisthosorna is large and has

lateral expansions. .:’.’.. ’...
Remarks: The present authors know practically nothing about this
genus, Vitzthum (194l) and Baker and Wharton (’1952) give .the date as
1857 and record P_. caligus’ Kolenati; 1857, as the’generotype.. Radford
(l950)givesthedateas 1858 and records P. interruptus; (Kolenati, 1856)
as the senerotype. We have .been.Unable to find’the original literature
to check this point and have arbitrarily accepted Baker’and Wharton’s
statements. We think it is quite likely that Periglischrus will prove to
be a synonym of Spinturnix. ;;.’’...":..

Periglischrus caligus Kolenati . .’ :. ,

Periglischrus caligus Kolenati, 1857; Radford, 1950; Bak’er,,an,d Whar-

ton, 1952. .; : ; ; ’ .. ’. ’

Locality: ?
Host; ....;?. . ;.
.
...’ . : ’-’:. "’.’:- ’
"’

’"’
-Periglischrus africanus Zumpt ...
.
,..
.
.
.
^

Type material in the Souths African Institute for Medical Research.

’
Periglischrus africanus Zumpt,-: 1950: .98 (d^Tibbetts. 1957. ,

Locality: Africa - Union of South Africa, Transvaal.

Host: Mammal - Rhinolophus geoffroyi.

Periglischrus asema Kolenati

Periglischrus asema Kolenati. 18-57; Radford, 1950. .

.
,

’
’

Locality: .?.. ;. .,
..
.
.
? ": ":" ’""
Host: . .

, P er iglis chrus glutinimargo Kolenati

Periglischrus glutinimargo: Kolenati, ’’1857; ’aadtord,’ 195.0.

Locality: ? (pee remarks’.). ’’’ :
Host: Mammal Rhinolophus
- clivosus. .:.; . .

Remarks: The host is restricted.to’.the Red Sea Coast (African and

Arabian) and that of the Gulf of Aden. Barring contamination from cap-
tive hosts, the mite is probably, also from the same locality.

Periglischrus hippoaiderus’Kolenati : .;. .

Periglischrus hipposiderus Kolenati, 1857; Oudemans,’ 1910: 67.(c, t);

RadfordT’1950. ’.
’
’. :,:
Locality: ? (See remarks.)
Host: Mammal - Rhinolophus hippocrepis.
Remarks: This species is .probably.European. The host is a syn-
onym of the European form of R. ferrum-equinum. .
 191

Periglischrus iheringi Oudemans

Periglischrus iheringi Oudemans, 1902: 38; Buitendijk, 1945; Radford,

1950; Tibbetts. 1957.
Locality: South America - Brazil.
Host: Mammal - Vampyrops lineatus.

Periglischrus interruptus <Kolenati)

Pteroptus interruptus Kolenati, 1856; 1857: 27.

Periglischrus interruptus, Oudemans. 1910: 67 (t); van Eyndhoven, 1941:
44 (t); Radford, 1950; Tibbetts, 1957.
Locality: ? (See remarks.)
Host: Mammal Rhinolophus blasii.
Remarks: The host is a Mediterranean species, being found around
this sea from Italy and Greece to Palestine and North’Africa. Eastwardly
it extends into S. W. Russian Turkestan.

Periglischrus meridensis Hirst

1957. . ,
Periglischrus meridensis Hirst. 1927: 323 (d); Radford. 1950; Tibbetts,
’

Locality: Islands of the Atlantic - West Indies.

’

Host: Mammal - Artibeus jamaicensis.

Periglischrus strandtmanni Tibbetts

(Fig. 92)
Type material in the U. S. National Museum.
Periglischrus strandtmanni Tibbetts, 1957: 14 (d, illus.).
Locality: North America - United States, Texas.
Host: Mammal - Mormoops megalophylla.

Periglischrus vargasi Hoffmann

Type material in the Institute de Salubridad y Enfermeaades Trop-

icales de Mexico.
.Periglischrus vargasi Hoffman, 1944: 91 (d); 1944: 261 (c); Tibbetts,
1957. .

Locality: North America - Mexico, States of Baja California, Chi-

apas, Guerrero, San Luis Potosi, Tamaulipas.’ ’
Hosts: Mammals Anoura geoffroyi, Leptonycteris nivalis, Ma-
-
crotus californicus, Molossus nigricans, Sturnia lilium.

Family HETEROZERCONIDAE Berlese, 1892

Description of the family. With a pair of large, bell-shaped hold-

192

fast organs flanking .the anus. The sternal plate is divided into a pair
of anterior lateral plates and a pair of posterior lateral plates that are
fused with the endopodal plates. Body setae are short and stout; claws
are weak. .’’. ’::
Most members of the family are phoretic on insects. .One .species
has been found on snakes. It is discussed below.

Genus Heterozercon Berlese

Type: Heteroaercon degeneratus Berlese, 1888.

,

Diagnosis: With the characters of the family. .., ." ""... ^

Heterozercon oudemansi Finnegan

Heterozercon oudemansi Finnegan, 1931: 1349. .

.
. . .:

.
Locality: Europe - England.

,
.
Host: Reptile - Epicrates cenchris.
Remarks: This mite was found under the ventral scales of the host.
The host had been collected from the region of the Upper Amazon in South
America. The mites were.discovered in the Zoological. Society Gardens,
"
’
London.
, There are anumber of species in the genus Heterozercon, but oude-
mansi is the only one recorded as parasitizing a vertebrate/and therefore
is the only one included in’ this manual, ,., : ,,’.’’:. :.. ,..,.,,,.,. ,.,.., .,^ ..

.
.
r,,’,’.,’
’

. , ... Family ^ARAMEGISTIDAE Trsigardh, .1,946 . ,; , ,.,,:,,;,

’iiescription of the family.’ Trigynaspida.. The male genital opening

lice posterior to the front margin of-the sternal plate. The female has
three genital covers,. The. chela,is composed, of. two long and tapering
digits, of which the moveable digit bears^two elaborate excrescences.
Remarks:, The maj ority. of ..the Paramegistidae, are ..associated with
ar’throRpds bttt’ specie s 6f-:the genus ^OphiQmegiatus seem to’be obligate
ectoparasites" of .snakes arid therefore are included in this manual.. Dr.
Joseph Camin, of Kansas University,, is describmg,sbme new species of
Qphiomegigtus from’sriake’s’-in fhe Sovth Pacific’ area.
,
Since this is. atrigyna.spid.fami,]y,.,it do.esnot.appearinthekeyto
the families;: The key includes .pnly the parasitic Monogynaspida.

.Genus; Qphipmegistus Banks;

’Type: Ophiomegistus.luzoherisis.Banks, ..1914: 58.
;’Celaenopsoides Gunther, .1942. .... ,, ,.....,. ,,.,
, .,, ,. ,.

..
...’Type:7. Celaenppspj.degbulolgehs.ig.Giinther,’;. 19
. ,8.7.,;
 193

Dia.gnos.is: Strongly orbicular, mites. The two latigynial plates and

the mesogynial plate of the .female about equally developed. Venter with
numerous small setae, the posterior 2 - 4 rows of which are flattened
or otherwise distinct from the. other setae.

Ophiomegistus luzonensis Banks

Type material in the U. S. National Museum.

Ophiomegistus luzonensis Banks, 1.914: 58; Banks, 1915; Grant, 1947:
22 (d, illus.); Gunther, 1951.
Locality: Islands of the Pacific - East Indies; Philippine Islands,
Los Banos; Dutch New Guinea, Hollandia,
Host: Reptiles - Unidentified snakes.

Ophiomegistus buloloensis (Gunther)

Type material in the School of Public Health and Tropical Medicine,

University of Sydney.
Celaenopsis buloloensis Gunther, 1942: 87 (:d). .

Ophiomegistus buloloensis, Gunther, 1951 (t).

Locality: Islands of the Pacific -.East Indies, New Guinea, Bulolo.
Host: Mammal - Rattus browni.
Remarks: Although this species was taken from a rat; it is doubtful
that the rat is a normal host. (See remarks underfamily Paramegistidae.)

Genera of Uncertain Familial Relationships

Genus Acanthochela Ewing, 1933

(Fig. 93)
.
Type: Acanthochela chilensis Ewing, 1933.
Diagnosis: Sternal plate of female .square, bearing three pairs of
sternal setae and three or four pairs of accessory setae; epigynial plate
not expanded, bearing one or two pairs, of setae-;’ fixed arm of chelicera
with two or more recurved setae; moveable arm subtended by a crown-
let of setae.
Remarks: The true affinities, of this mite are not at all obvious.
Ewing originally placed it with the Haemogamasidae, as did also Vitz-
thum (1942) and Baker and Wharton (1952). However, Keegan (1951) in
his revision of the haemogamasids did not think it belonged there and
removed it to the Laelaptinae, where it most certainly does’not belong.

Genus Alphalaelaps Radford, 1951

’
, (Fig. 94) .

Type: Laelaps aplodontiae Jellison,’ 1945.

Diagnosis: Epigynial plate of female elongated, bearing six pairs
194

of setae; anal plate of male separate; spermatodactyl as long as the. body.

Remarks:’ This is certainly one of the most unusual mites that we
have seen. Its familial relationships are obscure but it .may belong with
the Dermanyssidae. A truly astounding feature of this mite is the ex-
treme length of the male spermatodactyl: it is. slender, whiplike, and
fully as long as the body of the mite.

Alphalaelaps aplodbtttiae (Jellison)

(Fig,-94)
Laelaps aplodontiae 3 ellis on, 1945: 3)3.
. / . ’’. ’.

.
.
.
Alphalaelaps aplodontiae, Radford, ’1951: 201. ".
Locality: North America - United States, ’Washington State.
Host: Mammal - Aplodontia rufa.
Remarks: Aplodontia is an unusual and apparently phylogenetically
isolated rodent. All ectoparasites reported from it are also unsual.

Genus Manisilaelaps Lavoipierre, 1956

,
’ "-Type: Manisilaelaps coronis Lavoipierre, .Type by mono-
’
’
’

,’ ’^\.
’

^py- .., ;. .:
Diagnosis: Female’sternal plate large.; rectangular; epigynial plate
very:saort. extending only to the posterior bor’der of c’oxae IV, but rounded
posteriorly and with two setae on the posterior margin.

Manisilaelaps coronis Lavoipierre

Type material in the Liverpool School of Tropical Medicine.

Manisilaelaps coronis Lavoipierre, 1956: 291 (d, illus.).
Locality: Africa - West Africa. :’’:.:
Host:- Mammal - Manis tricuspis. :. .. .’. ... .

Remarks: Described from eight female specimens that were col-

’
lected from the edge of the scales of the host.

Genus Myonyssoides Hirst, 1925

(Fig. 95)
’
’
.
.
.
’
Type: Myonyssoides capensis Hirst. ., ,

Diagnosis: Female epigynial plate very short, truncate; ventrianal

plate large, covering most of the opisthosoma; sternal plate with three
or four pairs of setae; coxa I with a ventral apophysis.
Remarks: We have seen no specimens otthis genus but judging from
the illustrations and descriptions, these mites do not rightly belong in
any of the families of mites included in this paper. The very short and
truncated epigynial plate, and the very large ventrianal plate closely ad-
joining it are reminiscent of some of the Phytoseiidae,
 195

Myonysspides capensis Hirst

Myonyssoides capensis ’Hirst, 1925: 49; Bedford, 1932; DeMeillon and

Lavoipierre, 1944.
Locality: Africa - Union of South Africa, Grahamstown.
Host: Mammal - Cryptomys hottentottus.

Myonyssoides spiposus DeMeillon and Lavoipierre

.
: .: (Fig.-95)
Myonyssoides spinosus DeMeillon and Lavoipierre, 1944: @3, .

, Locality: Africa’-- Union of South Africa, Cape Province,

Host: Mammal - Amblysommus hottentottus. ’,’

Genus Mungosicola Radford, 1942

Type: Mungosicola ugandae Radford, 1942:- i’86.

Remarks: Said to have been described from two^ females and two
males; however, the two "females" are certainty deutonymphs. The
family placement is not-obvious from a study of the male and deutonymph
only. The specimens were’ taken in Africa on the banded mongoose,

.-’Genus-Tympanospinctus B’erlese, 1918

.
Type: Tympanospinctus paradoxus Berlese, 1918.
-.. Remarks:.’:’The-true affinities of this mite are riot known. "Berlese
.
placed it with the Spinturnicidae but it does not resemble other members
o:f. the family and .its host was not a bat.
According to Dr. C. E. Yunker, -who examined, .the type specimen
at the Agricultural Entomology Museum, Florence, Italy, the specimen .

bears the following data: "Tympanospmctus paradoxua Berl., Tipico,

rf, 193/16. Su Crossarchus fasciafus. Greenwood Park. D. R. Boyce -
1914." Crossarchus is a viverid (Carnivora), and the locality would in-
dicate a British zoo. Dr. Yunker noted that the specimen is certainly
not a spintumicid, but hesitated to assign it to any known family because
of its morphological uniqueness. Its dorsum bears a large (+ l6’0.^), el-
liptical spiracular plate immediately over.leg III; a well-developed, la-
ci’nate .tritosternum is present; -its leg chaetotaxy is more laelaptidthan

turMcidae. ..
spintumicid; and the .mouthparts are not modified as are those of Spin-
.
’
.
.
.
.. . Genus Ugandolaelaps Radford, 1942 . .

(Fig. 96)
Type: Ugandolaelaps protoxera Radford.
Diagnosis: Female epigynial shield greatly expanded behind coxae
.
IV and joined’to the anal pla’te. Only the genital setae are on the epi-
196

gynial plate. Monotypic; known only from the female.

Remarks: This genus seems to be intermediate between the Der-
manyssidae and the Laelaptidae. A study of the gnathosoma would be
necessary to place it correctly.

Genera and Species Not Rightly Belonging in This Manual

The laelaptine genera Androlaelaps and Hypoaspis are not included

because they are believed, by the present authors, to be non-parasrtic.
Davisiella reticulata Zumpt and Patterson, 1951 (p.’ 78).is rightly one
of the Hypoaspidinae, as Zumpt and Patters on stated. .It-is mentioned
here because these authors, in their key to the genera of the Hypoaspid-
inae, included all laelaptoid genera which have only one pair of setae on
the epigynial plate, some of which are truly Laelaptinae (such as Gigan-
tolaelaps and Eubrachylaelaps). The mite was taken in Southern Rho-
desia on Crocidura luna. ’
.:.
Laelaps (Eugynolaelaps) coriaceus Berlese, l9i8 (p. 128) has been
listed as a subgenus under Laelaps. However, the description and illus-
tration given by Berlese indicate it is a hypoaspid rather than a laelap-
tid mite. It was collected in Durban on Spalax typhus, a mole-rat.
G eneiadolaelaps spencei Gunther, 1942 (p. 89), according to the
sketch given in the original description, is not a Geneiadolaelaps in the
sense of E wing. The genital and ventral plates of the female are separate.
It is not possible to tell definitely where it belongs. It was discovered in
Papua crawling on the legs of soldiers. ..
.
T.he genus Pseudolaelaps Berlese, 1916, has been listed as a sub-
genus of Eulaelaps but it is actually a member of the. non-parasitic fam-
ily Pachylaelaptidae. The generotype. Laelaps (Hoplolaelaps ) doderoi
Berlese, 1910, wasexaminedbyDr. C.E.Yunkerat.the Agricultural Ento-
mology Museum. Florence,’ Italy. Although.be was unable to see the
tines on the modified claw of the palpal tarsus, in all other respects the
mite appeared to be-a member.of the Pachylaelaptidae: metapodal, peri-
tremal and parapodal plates coalesced; igenltal plate with three pairs of
setae; leg structure typically pachylaelaptid; chelicerae huge, typical of
the tree-living parasitoids. .

Pililaelaps longiseta (Banks) Radford (1947: 229) has been listed

among the parasitic mites’but it definitely does not belong there. It was
collected in Canada on a carrion beetle. Banksia Radford. 1942 (p. 229)
and Rad Baker and Wharton, 1952 (p. 97) are synonyms.
Scissuralaelaps nova-guinea Womersley, 1945(p. 225) was listed un-
der the subfamily Laelaptinae by Baker and Wharton (1952). It rightly be-
longs in the subfamily Hypoaspidinae. In addition to the generotype, one
other species was described: S. queenslandica Womersley, 1945. The
host of the generotype was a millipede; no host was given for the second
species,
Indogynium lindbergi Sellnick, 1954 (p. 285) (Antennophorina: Schizo-
gyniidae) is probably not a pa.rasite of vertebrates even though the collec-
 197

tion data relating to it does not preclude such a possibility. It was taken
on three occasions from snakes of three different genera (Platyplectrurug
madurensis. Uropeltis pulnejensia and Teretr.urus rhodogaster) in Kodi-
kanal, India. Members of the superfamily Celaenopsoidea, to which it
belongs, are free-living or associated with insects. However, species
ofOphiomegistus are parasitic on snakes and belong to a related super-
family (Antennophoroidea) whose other members are associated with
arthropods. In this case, however, the parasitic nature of Ophiomegis-
tus cannot be disputed. They have been observed with their chelicerae
inserted beneath the. scales of their hosts (Camin, personal communi-
cation). We prefer "to have more definite evidence of a parasite-host
relationship between I. lindbergi and the reptiles before including it as
a parasite.

Conclusion

The foregoing synopsis of the parasitic mesostigmatid mites includes

13 families, 3 subfamilies, 83 genera, and 590 species. It is intended
to be complete only through. 19 5.3, though occasional later references have
been included.
It is certain that this is not the total number of species that exist in
nature, but at the same time, it is evident that the present number of
names is greater than the’actual number of species described. It is also
true. that some of the families and genera are not clear entities. This
is due to the fact that so very many species have been described from
only one or a very few specimens, and frequently from only one sex.
The present concept of taxonomy, which embodies populations rather
than individuals, needs to be more universally applied in mite system-
atics before real progress can be made. Also, and this cannot be too
forcefully emphasized, descriptions and illustrations oS new species
must be drawn ]^p with painstaking and patient thoroughness.
Theacarologistciustmakeuseofthe gnathosoma. The deutosternal
teeth, the hypostomal processes, the epistome, the tectum, and the de-
tails of the chelae must be seen and accurately figured and described.
Also, the acarologist must figure and accurately describe the tritostern-
um. Here are subtle but real characters for differentiation, frequently
even on the family level. To omit these parts of the mite anatomy is
just as serious an omission as a description of a bird without any men-
tion of the beak, or of a mammal without describing the dentition. Also,
descriptions should be made to fit the speciss, not only a specimen. In
addition to mentioning the special features of the .type specimen, the var-
iability of all available material should be included.
Not included in the foregoing synopsis are the common genera An-
drolaelaps and Hypoaspis, nor a great number of genera ending in "-lae-
laps. " We consider Androlaelaps as belonging -to the hypoaspidine com-
plex, in spite of a strong resemblance to the laelaptine complex, and
198

that the hypoaspidine complex is non-parasitic. Nor can we agree with

those acarologists who consider all laelaptine mites having only one
pair of setae’on the genital plate as being Hypoaspis. Our reasons for
this are already stated under-the discu.ssion of the family Laelaptidae.
Many genera of mites having the suffix "-laelaps" and not included here- .

in, such as Pachylaelaps, Gamasolaelaps, etc., are obviously not para-

sitic mites. Others, such .as Cosmolaelaps, Gymnolaelaps, Myrmo-
laelaps, although’often found in association with vertebrates are never
found in such-numbers nor under such circumstances as to suggest that;
they are parasitic, and are therefore not included. :

DISTRIBUTION

Representatives of the parasitic Mesostigmata have been found on

every major land mass in the world and on a great many of the smaller
islands. No doubt they may be found in every habitat that supports ter-
restrial vertebrates.
In parasites that have no free-living stages, one might assume that
their distribution equals that of their hosts. To an extent this is true of
the parasitic Mesostigmata, but there are limiting factors. Probably
the greatest of these is that the majority of species are primarily nest
and roost inhabitants and are not permanently on the host. This would
certainly limit the distribution of-the ectoparasites of reptiles, which
have no nests, and of the ectoparasites of birds., which may nest in one
area and never spend two nights in the-same place during the non-nesting
period. .;., , ,
.’.

.
.
The distribution and presence of the external bird-, mites depends
largely upon a continuous source of food for the mite. If one assumes
the nest of some wild bird to become infested with a few mites just at
the time the first egg is laid, then during the period of .incubation, plus
the period before the young leave the. nest. a tremendous population of
mites can be built up. Yet, after the young have flown-and the parent
birds leave the area, all of these mites will diminish in abundance, and
perhaps die long before the birds return the following year. Presumably,
if the grown birds divest themselves completely of any.hpldover nest-
mites, there should be no mites like Ornithonyssus sylviarum, 0. bur-
sa, or Dermanyssus gallinae in nests of strictly sylvan areas. The dis-
tribution of these mites, then, is probably determined by a continuous
population of indigenous birds such as house sparrows, pigeons, and do-
mestic fowl, and all migratory birds very probably become infested with
these mites through contamination-from such permanent residents.
It would seem .that the distribution of mites of mammals should be
continuous and uniform over the range, of their hosts. Here is a situa-
tion in which nests, or at least burrows, are constantly occupied and the
host itself provides a microhabitat. of constant temperature and fairly
uniform relative humidity. If we take, for example, the mite Laelaps
 J.99

multispinosus and its host the muskrat (Onadatrae zibethica) we seem

to have proof of this hypothesis. This mite has been found on the musk-
rat in every region of North America and in regions into which it has
been introduced (e.g., Europe). However, if we consider Laelaps nut-
tali and its host, the common rat (Rattus norvegicus), we quickly see
that out hypothesis is not always true. Laelaps nuttalli has followed its
host around the world, but only in the tropical and subtropical regions;
and even in these regions its distribution is not continuous.
The environmental requirements of some few species of mesostig-’
matid mites have been investigated. For all the others, nothing is known.
We do know that in some areas a host will have one species of mite; in
another area that does not seem greatly different physically, the same
host will have an entirely different species. The packrat <Neotoma spp.)
is a good example. It may be parasitized by Brevisterna sp. in one area
and Haemolaelaps glasgowi in another; or Hirstionyssus neotomaeat one
place and Hirstionyssus breviseta in another, etc.
We know that a limiting factor is the presence of predatory arthro-
pods in the nest. We have seen predatory mites andpseudoscorpions avid-
ly devouring parasitic’mites. Certainly these would play an important role
in regulating the abundance of a mite species, but we do not believe that
it is the most important factor in determining whether a species will be
present or absent.
Probably the most critical factor in determining the distribution of
parasitic mites is the physical presence of the host. Drummond (1957)
studiedthe factors influencing acarine populations in the nests of Pero-
myscus leucqpus in Maryland. He showed the nesting activity of the
mice to be the most important factor governing these fluctuations. Yunker
(1958b) investigated the ecological factors that determine mite popula-
tions in desert and semi-desert burrows of Egyptian gerbilles (Ger-
billus gerbillus and Ci. pyramidum), He found the presence or absence
of mites inanesttobe dependent upon the presence or absence of a host,
regardless of area or climatic state of the burrow, while population size
was affected by the type of area. He was unable to demonstrate any
specificity of certain laelaptid species for. either species of host,’ while
individual species couldbe shown to prefer burrows situated in a particu-
lar area. This must be considered in the light of Wharton’s (-1957) state-
ment concerning intraspecific variation of parasitic mites’that live and
breed in the nests of their hosts. He considered host specificity in these
forms to be largely a matter of nest specificity. He cited the ability of
these forms’to utilize many hosts under- laboratory conditions, and stated:
"Givenanestwithcertainenvironmentalfeatures favorable to the repro-
duction of the mites, it is quite possible that the mites will’be found in
the nest. " Physical ecological factors rather than host specificity may
well explain the non-random distribution of nest-dwelling mites.
Geographically, it is apparent that many areas ofthe’world have not
yet been extensively collected. Africa has a very impressive list for the
Union of SouthAfrica but not much from the rest of the continent. Asia
has been grossly neglected until the last two decades. Russian workers
200

have recently reported many. species from the U.S.S.R. and it seems
assured that many more will be reported in the future’. Japanese and
Americans are discovering more and more species in-Korea, Manchuria
and Siam. as well as in Indonesia, Japan, and .the Islands of the Pacific.
Australia has not yet recorded.more .than a fraction of the species it is
bound to have. One reason for the scarcity of records is that prior to
World’War II practically all. acarological work in Asia. the Pacific Is--
lands, and Australia was devoted to the Trombiculidae.
South America has a fairly complete list only from Brazil, chiefly
through the efforts of Fonseca, and of Pereira and de Castro. Except
for these workers. South America does not seem to have any native aca-
rologists interested in the .Mesostigmata. Most of the re-cords, outside
of Brazil were contributed by foreign scientists. .:

. : Ther-e are fairly complete-lists from Europe but not:as complete as

.one might -expect. This is probably because the Europeans have inter-
estedthemselves more in the free-livingforms such as.the Oribatei; etc.
Because every continent has at least one large area in which collec-
tions have been fairly .thorough, it is possible to comment on the cosmo-
politanism of some genera and species. The genera .Haemolaelaps, Lae-
laps, Ornithonyssus, Ichoronyssus, Eulaelaps, Haemogamasus and Spin-
turnix have been found everywhere that more than a casual collection has
been made. Haemolaelaps glaagowi, Ornithonyssus bacoti; and Eulae-
laps stabularis seem to be truly cosmopolitan. La-elaps- nuttal’li, Echino-
laelaps echidninus and Dermanys.sus gallinae encircle the-globe but only
within the limits of the tropic and-temperate zones. Many of the genera
of the internal mites of birds and mammals have a -world-wide distribu-
tion; a fact which would be difficult to explain if it were otherwise".
As a rule, the mites of world-wide distribution are those that para-
sitize animals having a world-wide distribution. Laelaps nuttalli, for
example, is certainly circumglobal because its hosts, the domestic rats,
have carried it around the world. Ornithonyssus bacoti likewise has
been carried all over the world by commensal rats, but because it will
parasitize other animals as well, it has a much wider distribution than
nuttalli. Haemolaelaps glasgowi. shows no host preference and may have
become universally distributed by a variety of hosts including the- Nor-
way and roof rats.
With. one .exception, the information available is not yet sufficient
to allow us to deduce the points of origin of any of these mites. In fact,
many of them, all of the Laelaptinae for instance, are still so much like
the,ir free-living predatory cousins that a thorough knowledge of the dis-
tribution of .the free-living forms is necessary before one can theorize
on points of origin of the parasitic forms. The association of Ornitho-
.
nyssus bacoti, with Sigmodon tuspidus and its nematode parasite, Lito-
mosoidea cariniij ..certainly suggests a ’new world origin for the mite.
Perhaps similar studies on other vectors of .disease may give clues to
the origins of other mites.
The following geographical list follows political rather than physical
boundaries. This. is for two reasons: (1) It is the purpose of this man-
 201

ual to indicate what mites are known in the various political divisions.
and (2) we do not have enough data to provide meaningful ecological dis-
tributions.

’FAUNAL LIST

NORTH AMERICA

Alaska .’. .

.
Laelaps alaskensis, L. pachypus, Haemogamasus alaskensis, H. am-
bulans, Orthohalarachhe diminuata.

Canada

Haemolaelaps glasgowi, Laelaps multispinosus, L. pachypus, Qrnith-

onyssus bacoti, (3. bursa, jO. sylviarum, Hirstionyssus isabellinus, Der-
manys sus americanus, ^. gallinae, Myonyssus jamesoni, Eulaelaps
pedalis, 13. prppheticus, E^. stabularis, Haemogarnasus alaskensis, H.
barberi, H. liponyssoides, H. occidentalis.

Mexico

Haemolaelaps casalig, H. glasgowi, H. spinosulus, Eubrachylaelaps

circularis, E. debilis, f[.martini, E. jamesonj, E, ..spinosus.,. Stepto-
laelaps liomydis. Echmolaelaps echidninus, Ichoronyssus granulosus,
I. robu stipes, Halarachne americana, H. miroungae, Periglischrus
vargasi, Spinturnix carloshoffmanni, S. ewingia.

Uriited States
Haemolaelaps casalis, H. geomys, H. glasgowi, H. morlani, Eubrachy-
laelaps eircularis, E_. crowei. E. debilis, E. hollisteri, Gigantolaelaps
cricetidarum, Steptolaelaps liomydis,. Laelaps multispinosus, L. nut-
talli, L. oryzomydis, L; pachypus, Echinolaelaps echidninus, Ornitho-
nyssus americanus, 0. bacoti, 0. burs.a, 0. sylviarum, Ichoronyssus
crosbyi, I_. longisetosus, I_. quadridentatus. L robustipes, Ophionyssus
natricis. Hirstionyssus affinis, H. arcuatus, H. breviseta, H. carnifex,
H. cynomys, II. geomydis, H. hilli, H. mcomptus, H. isabellinus. H.
neotomae, H. obsoletus, H. occidentalis, H. staffordi, H. triacanthus,
Neoichoronyssus dentipes, N. wernecki, Patrinyssus hubbardi, Steato-
nyssus ceratognathus, S. occidentalis. Pellonyssus passeri, Alloderma-
nyssus sanguineus, Dermanyssus americanus, D. brevis, D. evotomy-
202

dis, D. gallinae, D. oti, D. prognephilus, D. scutatus, Myonyssus jame-

soni) Eulaelaps citellus,, 13. stabularis, Haemogamasus alaskensis, H.
ambulans, H, barberi, H. harperi, H. keegani, H. liponyssoides, H.
occidentalis, H. oudemansi, Brevisterna montanus, B. morlani, B.
utahensis, Ischyropoda furmani, L spiniger, Ixodorhynchus liponya-
soides, Hemilaelaps americanus, IL diatinctus, H. triangulus, Rail-
lietia auris, Entonyssus ewingi, :E_. fragilis, E. heterodontus, TE_. rileyi.
]E. vitzthumi. Ophi.opneumi.c ola elaphes, 0. natricis, Halarachne amer-
icana, H. miroungae, Orthohalarachne attenuata, 0. diminuata, 0^. zai-
ophi, Pneumonyssus bakeri, P_. caninum, Rhinonyssus alberti, R. coni-
ventris, R. himantopus, R. rhinolethrum, Larinyssus orbicularia, Ral-
linyssus caudistigmus, Neonysaus belopolskii, N. columbae, N. melloi,
N. zenaidurae, Rhinoecius bisetosus, _R_. cooremani, R. grandis, Stern-
ostoma boydi, S. hutsoni, S. spatula-turn, S. strandtmanni, Paraneonys-
sus capitatus, Paraneonyssus hirstt, Ptilonyssus nudus, Spinturnix
anaericanus, S. echinipes., S. lowae, Periglischrus strandtmanni.
Spinturnix grossus. ,’ ,

CENTRAL AMERICA

Spelaeorhynchus praecursor. .-.-.

Canal Zone

Ornithonysaus burs a, Spelaeorhynchus latus.

SOUTH AMERICA .
Argentina

Haemolaelaps reithrodontis, Cavilaelaps bresslaui, Gigantolaelaps mat-

togrossensis, Laelaps mazzai, L_. robustipes, L^. wetmorei, Ornitho-
nyssus bacoti, 0. bursa, 0. hirsti, 0. meprai.

Bolivia

Bolivilaelaps- tricholabiatus.

Brazil

Haemolaelaps coelogenys, H. reticulatus, H. sciureus, Cavilaelaps bra-

ziliensis, Eubrachylaelaps rotundus, Gigantolaelaps brachyspinosus, G.
butantanensis, _G. comatus, G. gilmorei, G. goyanensiSj G. mattogros-
 203

sensis, G. oudemansi, Neopara’laelaps bispinosus, Laelaps aragonensis,

L. differens, i^. exceptionalis, L^. hirsti, ;L. lativentfalis, L. ’mangui-
hosi, i^. nuttalli, X;. parvulus, L. tnori, Echinplaelaps berlesei, E.
echLdnmus, Mysolaelaps parvispmosus, M_. microspi.nosns, Onuthonys-
sus b’a&oti, 0. braziliensis, 0. erudi.tus, 0. ihenngi. 0. lutzi., 0. mon-
teiroi, 0, vitzthun’ii, Ichoronyssus haematophagus, I. kochi, I. kolenati,
Sauronyssus myrniecophagus, Ophionyssus natricis, Hirstionyssus bu-
tantanensis, Neoichoronyssus wernecki, Lepronyssoides pereirai, L.
matogrosso, Steatonyssus joaquimi, Radfordiella oudernansi, Li.ponys-
aoides brasiliensis, Derinanyssus gallinae, Eulaelaps vitzthumi, Xxo-
dorhynchus butantanensis, Dasyponyssus neivai, Pneumophionyssus ar-
istoterisi, Tinaminyssus navajasi, T. trappi, Neonyssus melloi, N. ser-
raoi, Cas angrensis, Flavionyssus’rabeUol, Rochanyssus werneri, Trav-
anyssus paranensis, Rhinonyssoides donatoi, R. souzai, Paraneonys-
sus enriettii, P. hirsti, P. travassosfilhoi, Ptilonyssus japuibensis, P.
olaioi, P. sairae, P, stresemarini, Periglischrus iheringi.

Chile

Haemolaelaps glasgowi, Gigantolaelaps wolffsohni, Ichoronyssus robus-

tipes, Neoichoronyssus dentipes, ’’

.. ; ,. ’. Columbia

Ornithonyssus bursa, Steptolaelaps heteromys.

Ecuador

Tur uniscutatus.

’ ’ ’Peru .

Haemolaelaps casalis, H. chinchillulae, H. glasgowi, Eubrachylaelaps

rotundus, Gigantolaelaps peruvianus.

Surinam .
.
Gigantolaelaps versteegi, Laelaps nuttalli.

, i’ Uruguay

Haemolaelaps casalis, Gigantolaelaps maximus.

Venezuela

Ichoronyssus hasei, I. venezolanus. Steptolaelaps heteromys, Derma-

nyssus gallinae, Spinturnix araguensis.
204

EUROPE

Ichoronyssus’ flavus,. Sauronyss’us lacertinus, S_. saurar.um, Permanys-

3us chelidonis, E>. quintus, Haemogamasus ambulang, Raillietia auris,
Pneumonyssus simieola, Rhinohyssus coniventris. Ptilonyssoides tri-
scutatus, Vitznyssus:.vitsi.thumi, V. nitzschi,. Spinturnix acjiminatus, S.
myoti, S. oydemansi, .S. psi.’.’ . . . ;
Belgium"
Rhinoecius pti.

’British Isles

Haemolaelaps casalis, Laelaps agilis, L. hilaris, L. muris, L. pachy-

pus, E chinolaelaps echidninus, Macronyssus orcadensis, Ornithonyssus
bacoti, 0. sylvlarum, Ophionyssus natricis, Hirstionyssus blanchardi,
H. isab’ellinus, H. ’soricis,- Hirstesia britteni, H. sternalis, Steatonys-
sus murinua, Dermanyssus gallinae, D. hirundinis, D. quintus, Myo-
nyssus decumani, M. gigas, Eulaelaps novus, E. stabularis, Haemo-
gamasus ambulans, H. hirsutus, H. homdus, H.oudemansi, Entonys-
sus halli., Ophiopneumicola hamertoni, Halarachne halichoeri, Rhino-
nyssus caledonicus, R.- waterstoni, Paraneonyssus hirsti, Spinturnix
euryalis, £^. omahonyi, £^. plecotinus, Heterozercon oudemansi.

Corsica

Spinturnix euryalis.

France

Ichoronyssus lepidopeltis, I_. spinosus, Hirstionyssus bianchardi, H.

sciurinus, Dermanyssus gallinae, Sternostoma cryptorhynchum, Ptilo-
nyssus nudus, Spinturnix niurinus.

Germany
Haemolaelaps casalis, H. fahrenhoizi, H. glasgowi, Laelaps arvico-
lae, L. hilaroides, \^. multispinosus, L. muris, L, pachypus, Ornith-
onyssus bacoti, 0. ondatrae, .0. sylviarum, Ichoronyssus diversipilus.
L lepidopeltis, I. mohrae, I. spinosus, Hirstionyssus arcuatus, H.
blanchardi, H. carnifex, H. isabellinus, H. pachypus, H. pauli, Steato-
nyssus musculi, S. murinus, £[. spinosus, Dermanyssus .gallinae, D.
quintus, Eulaelaps nbvus, K. oribatoides, E. stabularis, Haemogamasus
ambulans, H. avisugus, H. hirsutosimiiis,’ H. hirsutus, H. horridus,
H. oudemansi, Neonyssus nucifragae, Ptilonyssus echinatus, Spinturnix
murinus, S. plecotinus,
 205

Hungary

Steatonyssus murinus.

Italy

Haemolaelaps casalis, H. fahrenholzj, H. inversus, H. subterraneus,

Laelaps sicula, Ornithonyssus sylviarum, 0. tinae, Hirstionyssus ar-
cuatus. Steatonyssus murinus, Permanyssus gallinae, Myonyssus de-
cumani, Eulaelaps pontiger, E, stabularis, Haemogamasus horridus,
Hemilaelaps piger, Sternostonia rneddai, Paraneonyssus hirsti, Ptilo-
nyssus nudus, Spinturnix euryalis.

Netherlands ’’’
Haemolaelaps casalis, H. glasgowi, Laelaps arvicolae, L. muns, JL.
pachypus, laMoronyssus lep.idopelti.8, Ophi,onyssus natricis, Hirstionys-
sus arcuatu’s/ H. ca.rnifex, H. isabellinus, Steatonyssus cyclaspts, S.
musculi. Dermanyssus gallinae, D’. hirundinis, Myonyssus decurnani,
M. gigas, Haemogamasus ambulans, H. hirsutus, H. horridus, Spin-
turnix murinus, S. plecotinus.
’

Norway ’
’

’
Ornithonyssus bacoti, Eulaelaps stabularis. ; .
.
Portugal

Ptilonyssus.hudus, Paraneonyssus, hir^ti.

’; ’’i Sardinia
’
.

Haemolaelaps concurrens, H. elongatus, Steatonyssus murinus, Haemo-

gamasus horridus.

. ’. -.: ,,,Spain.,; .
.

Ornithonys sus sylviarum, Steatonyssus musculi, Haemogamasus hir-

sutus, Spinturnix murinus.

’.. Switzerland
’
.’’,’.
Echinolaelaps echidninus, Eulaelaps stabularis, Haemogamasus ambu-
lans.

Yugoslavia

Hirstionyssus macedonicus, Haemogamasus hirsutus, Ichoronyssus

206

granulosus, _I. s.;utatus, Dermanyssus gallinae, Spinturnix plecotinus.

ASIA

Bhutan’

Laelaps bhutanensis.

Burma

Haemolaelaps traubi.

China

Ornithonyssus bacoti, 0^. bursa, Hirstionyssus confuscianus, Eulaelaps

stabularis, E. cricetuli, Haemogamasus oudemansi, H. mandschuricus.

India

Laelaps buxtoni, L. nuttalli, Echinolaelaps echidninus, Ornithonyssus

bacoti,0. bursa, Steatonyssus javensis, S. musculi, Pellonyssus viator,
Liponyssoides muris, -Hemilaelaps imphalensis, Hamertonia schoute-
deni, Pneumonyssus simicola.

Korea

Haenxolaelaps glasgowi, Laelaps jettaiari, L. nuttalli, E chinolaelaps

echidninus, Ornithonyssus bacoti, 0. sylviarum, Hirstionyssus carni-
fex, H. isabellinus, Eulaelaps stabularis, Haemogamasus ambulans,
Hemilaelaps farrieri, Hemilaelaps tanneri.

Malaya

Laelaps nuttalli, Longolaelaps longulus.

Manchuria

Hae molae lap s haemorrhagicus, Laelaps jettmari, L. p achypus, Myo-

nyssus shibatai, Eulaelaps cricetuli, E. stabularis, Haemogamasus el-
lipsoideus, H. kitanoi, H. kusumotoi. H. mandschuricus, H. occidental-

Siam

Ptilonyssus orientalis.
 207

..U.S.S..R.
Haemolaelaps androgynus, H; angustiscu.tis, H. casalis, H. .ellobii, H.
longipes, H. glasgowi, H. razunji’pvae, H. semidesertus, Laelaps agilis,
L. agrarius, L. algericus, L. arvalis, L. caucasicus, L. clethriono-
mydis, L. ekstremi, L. hilaris, L. jettmari, L. kolpakovae, L. lemni,
L. inicromydis, L. mosquensis, L. multispmosus, L. muris, L. novi-
kovae, L. nuttalli. L. pachypus, L. pavlovskyi, L. pitymydis, L. semi-
tectus, L. volgensLs, Oryctolaelaps bibikovae, Echinolaelaps echidninus,
Ornithonyssus bacoti, 0. sylviarum, Q. dogieli, Ichoronyssus Havus,
I. lepidopeltis, Sauronyssus saurarum,;. Ophionyssus natricis, 0. vari-
abllis, Hirstionyssus arcuatus, H. .blanchardi, H. bregetovae, H. car-
nifex. H. confucianus, H. cric.eti, H.. ellobii, H. eusoricis, H. evers-
manni~H-- georgicus, H7 isabelli.n.ui, H.. macedonicus, H. meridianus,
H. musculi, H. pachypus, H. pauli, H. sciurinus, H. soricis, H. trans-
iliensis, Steatonyssus musTuli, S. superans, Pellonyssus viator, AUo-
dermanyssus sanguineus, Dernaanyssus gallinae, D. hirundinis, D. qum-
tus, IVlyonyssus decumani, M. dubinini, M. gigas, M. ingricus, M.
rossicus, Eulaelaps stabularis, E. cricetuli, E. kolpakovae, Haemo-
gamasus ambulans, H. hirsutus, H. citelli, H. dauricus, H. hirsuto-
sinnlis, H. horridus~ H. ivanovi,~H. kitano.i,.-.’H. :kusurnotoi, H, lipo-
nyssoides, H. mandschuricus, H. nidiforines. ~H.. pontiger, H. serd-
jukovae. H. zachvatkini, Haillietia auris. Rhinonyssus caledonicus, R.
coniventris, R. minutus, R. rhinoletbrum, R. waterstoni, Larinys-
sus orbicularis, Rallinyssus caudistigmus, Neonyssus belopolskii, N.
nucifragae, Sternostoma tracheacolum, .Ptilonyssus nudus, -Spinturnix
~
" r
murinus, S. myoti.

ASIA MINOR

Arabia

Ophionyssus natricis, Liponyssoides muris.

Israel

Steatonyssus murinus, Spinturnix lateralis.

AFRICA

Haemolaelaps davisi, H. mystromys, H. spegazzinii. Laelaps grenieri.

L. lamborni, L. lavieri, L. lavoipier’rei, L. robaudi, L. spinifer, L.
thamnomys, L-.’ vansomereni. Echinolaelaps muricola,’ 15. yaoundensis,
Allodermanys.sus sanguineus, OrnUhonys sus bacotii Steatonyssus mur-
inus, Pneumonyssus mossambicensis, P. procavians,_P. rodhaini, P.
schoutedeni, P. simicola, P. santos-diasi, Rhinophaga papionis,- R.
cercopitheci, Pneunaonyssoides phacochoeri, Spinturnix myoti.

. .
Ethiopia (Abyssinia) ’’; :

Haemolaelaps patersoni, H. ’spegazzmii, H. 8preo,^Echin61aelaps mur-

icola, Ornithonyssus bacoti, Steatonyssus~eos, S.’natalensis, Pellonys-
ius: biseutatus, P, reedi, P. similis,- Pneunionyssus simicola. Lari-
nysstts orbiculariT, Neonyssus ardeae, N. bubuici, N... mellot, Rhinoecius-
africaftus, Sternostoma rhinolethruna, §’. traCheacolu-Ki, S.’turdi, Para-
neonyssus capensis, P. dryoscopi, Ptilonyssus echinatus, F. cinnyris,
P. desfontainel, P. lahii, P. nudus, P. phyllastrephi, Spinturnix abys-

Algena ’ _’;
Laelaps algericus, L. oraniensis, Echinolaelaps bakeri, Steatonyssus
rourinus. . . .’ ’;

Belgian Congo ’.’

Echinolaelaps echidninus, E, giganteus, E. muncola, Ornithonyssus
bacoti, Steatonyssus musculi, Liponyssoides muris, Pneumonyssus con-
goensis, P. duttoni, "Rhinophaga atheruri, R. cercopitheci, R. leopoldi,
R. papioms, Pneumonyssoides phacochoeri, Rhinonyssus afribyx, R.
apus, R. coniventris, R. himantopus, R. poliocephali, R. rhinolethrum,
Larinyssus orbicularis, Rallinyssus congolensis, R. llmnocoracis, Neo-
nyssus ardeae, N. bubuici, N. buteonis, N, columbae, N. ixobrychi,
N. schoutedeni, N. treronis, Rhinoecius tytonis, Ruandanyssus terp-.
siphonei, Sternostoma boydi, S, colii, S. cooremani, S. cuculorum, S.
dureni. S. hirundinis, S. lagonostictae, S. laniorum, S. nectannia, S.
sturnTcola, S. thienponti, S. tracheacolum, S. turdi, Rhinonyssoides
cerchneis, R. indlcatoris, Ptilonyssoides melittophagi, P. triscutatus,
Vitznyssus ^Titzschi, V. neotis, V. scotornis, V. vitzthuini, Paraneo-
nyssus astridae. P. capensis7 PT dryoscopi, P~. emberizae, P. hirsti,
P. ploceanus, P. "serini, P. zumpti, Ptilonyss^us andropadi, P, cala-
mocichlae, P. ^hloroclchlae, P. cinnyris, P. dicruri, P. dioptrurnis,
P. echinatusT P. lanii, P.motacillae, P. nudusT P. orioli, P. psalido-
procnei, P. pycnonoti, IP, ruandae, P. terpsiphonei, P. viduae.

British East Africa

Hirsti onyssus creightoni.

 209

British Somaliland

Echinolaelaps muricpla.

Egypt

Haemolaelaps aegyptius, H, ewingi, H. hystrici, H. insculptus, H. mur-

inus, Laelaps algericus, L. keegani, L. lamborni, L. nuttalli, Ec’hino-
laelaps’echidmnus, Macronyssus longirnanus, Ornithonyssus aethiopicus,
0. bacoti, Ichoronyssus cornutus, I. hoogstraali, I. lepidopeltis, I. le-
prosus, Ophionyssus natricis, Hirstionyssus arcuatus, H. carnifex, H.
craticulatus, Allodermanyssus aegyptius, A. sanguineus, Steatonyssus
murinus, Pellonyssus viator, Liponyssoides muris, Derrnanyssus gal-
linae, Eulaelaps stabularis, Haemogamasus.liber.iensis.’

French Guinea

Derrnanyssus hirundinis.

French West Africa

Haemolaelaos mauritanicus.

Gambia

Spinturnix aethiopicus.

Gold Coast

Echinola^laps. niuricola.

Kenya

Haemolaelaps hystrici, H. murinus, H. sudanicus, H. tachyoryctes.

Laelaps aethiopicus, L. lamborni, vansomereni, L. zumpti, Echino-
^.

laelaps giganteu’s, E. grandis. E^. muncola, Ichoronyssus jacksoni,

Hirstesia kenyaensis, H. transvaalensis, Steatonyssus eos, Dermanys-
sus gallinae, Haemogamasus liberiensis, Spinturnix kenyaensis, S. walk-

’.
’

’; ’ ’

Liberia. ’
’
’. ’
-
,
.
Laelaps liberiensis, Echinolaelaps giganteus, Ornithonyssus bacoti,
Hristionyssus liberiensis, Haeroogamasus liberiensis.

Morocco
’
Steatonyssus viator, Derrnanyssus gallinae.
210

Natal

Haemolaelaps labuschagnei, H. natalensis, Omithonyssus bacoti, Steato-

nyssus natalensis, Neospinolaelaps .ininiopteri, Rhinoecius africanus,
Spirrturnix lawerencei.

Nigeria . .. .

Haemolaelaps galagus, H. hystrici, H. spatuliformis, Laelaps lamborni,

Echinolaelaps giganteus, E. muricola, Omithonyssus bursa, Hirstionys-
sus ngami.
’
: ’. ’

’1 Nyasatao.d ’. "

’:...’.. ..’
Laelaps laniborni, Echinolaelaps muricola, Ornithonysaus aethiopicus,
.0. bursa, Steatonyssus nyassae, LiporiyssQldes muris^
’ ’..;
1
Rhodesia

.
.
Haemolaelaps rhodesiensis, H. Yillosissimus, Hamertonia bedfordt.

Sierra Leone

Spinturnix pipistrellus.

Sudan

Haemolaelaps hirsti. H. radfordi, H. sudanicus, Laelaps vansomereni,

Echinolaelaps muricola, Omithonyssus aethiopicus, Hirstionyssus hirsti,
Sauronyssus gordonensis, Steatonyssus nyassae, S. sudanensis, Allo-
dermanyssus sanguineus, Permanyssus gallinae, ’Neoliponyssus gordon-
ensis, Spinturnix scotophili.

Tanganyika

Haemolaelaps spirostrepti, Echinolaelaps muricola.

Uganda

Haemolaelaps arvicanthus, H. dasymys, H. hystrici, H. mesopicus, H.

murinus, H. sangsteri, H. tateronis, Eubrachylaelaps lophuroniius.
Laelaps vansomereni, Echinolaelaps bakeri, E. giganteus, E. grandis,
E. muricola, Omithonyssus dendropicus, 0. galagus, Hirstionyssus
liberiensis, H. otomys, Raillietia hopkinsi.

Union of South Africa

Haemolaelaps bathyergus, H. capensis, H. cryptomius. H. eloffi, H.

 211

eos, H. labuschagnei, H. inops,. H. lawrencei, H. murinus, H. oliffi.

H. phoeniculi, H. rhabdomys, H. spinitarsus, H. taterae, H. villosis-
iimus, H. zulu^adfordilaelaps’meridionalis, Laelaps longiwntris, L.
nuttalli, L. parvulus, L. simillimus, L. transvaalensis, L. vansom-
ereni, Echinolaelaps bakeri, E. giganteus, E. muncola, Omithonyssus
bacoti, 0. bursa, 0. rhinolophi, 0. roseinnesi, Ichoronyssus nyctinorni,
Hirstionyssus latiscutatus, H. sahtos-diasi, Hirstesia .transvaalensis,
’Pellonyssus biscutatus, P. Teedi, P. similise Dermanyssus gallinae,
Haemogamasus oudemansi, Mamtherionyssus heterotarsus, Pneumo-
nyssoides caninum, Pneumonyssus procavians, Rhinonyssus coniven-
tris, Larinyssus orbicularis, Sternostoma tracheacolum, Periglischrus
africanus, Spinturnix semilunaris, S. viduus.

West Africa

Haemolaelaps galagus, H. spatuliformis, Laelaps nuttalli, Ornithonys-

sus bacoti, Hirstionyssus ngami, Steatonyssus brucei, Manisilaelaps
coronis.

AUSTRALIA

Haemolaelaps casalis, H. marsupialis, Mesolaelaps anomalus, M. aus-

traliensis, M. lagotisinus,,,.]VI. thalacomys, HeteroTaelaps antipoSianus,’
Laelaps assirnilis, L. finlaysoni, L. hapaloti, L. muris, L. nuttalli,
T7. sminthopsis", EcTunolaelaps echidninus, Mysblaelaps roths childi,
Omithonyssus bacoti, 0. bursa, 0. sylviarum, .Sauronyssus arnhern-
landensis, Australolaelaps mitchelli, Hirstionyssus arcuatus, Echinonys-
sus validipes, Pellonyssus. malurus,. Dermanyssus gallinae,~Haen-ioga-
masus oudemansi, Pneumonyssoides caninum, Rhinonyssoides troues-
sarti, Spinturnix antipodiaiiusi,,. S. novaehollandae.-

ISLANDS OF THE ARCTIC

Franz-Joseph Land

Orthohalarachne rosmari.

ISLANDS: OF THE ATLANTIC

, Bahamas

Omithonyssus bursa. Halarachne americana.

212

... Bermuda . .
.. . :

.
.
Laelaps nuttalli. ; . . .. : : . ; ’.
.."., ..’ "" ".
’

Cuba

Ornithonyssus bacoti. ,. ’,.. ., ... .

Falkland Islands . .

.
Orthohalarachne magellanica.; .
^

Greenland

Laelaps semitectus, Haemogamasus ambulans.

. Iceland ,.
^
.
.
.
Haenaogamasus ambulans.

Jamaica

Ichoronyssus robustipes, Spelaeorhynchus praecursor.

Puerto Rico . . . »

-’Haemolaelaps glasgowi, Laelaps nuttalli, Echinolaelaps e.ehi.dninus. Or-

riithonyssus, bacoti. .

West Indies . .:.

Ornithonyssusbacoti, Halarachne americana, Rtiinony s s oide s squaino-
sw, Periglischrus meridensis, Spinturnix artibiensis.

ISLANDS OF THE INDIAN OCEAN

Ceylon
’
. ,
.
.
Laelaps atypicus, L. nuttalli, L. taprobanius, L. thompsoni, Echino-
laelaps echidninus, Liponysspides muris.

Comoro Islands

Ornithonyssus bursa.
 213

Madagascar

Aetholaelaps sylstrai, Ichoronyssus forsythi, Liponysella madagascar-

iensis, Neonyssus i.ntermedi.us, Spinturnix calcaratus.

..
.
Zanzibar

Orni.thonyssus bursa, Spinturnix aethiopicus. .

.
INDONESIA . . .

.
Laelaps soricis, Ornithonyssus bursa.

Borneo

E chinolaelap s echidninus.

Celebes

Spinturnix jordani.

Dutch East Indies

Echinolaelaps echidninus, Neolaelaps magnistigmatus, Steatonyssus .ja-

yensis, Liponyssoides murinus, Pneurnonyssus simicola, Spinturnix ana-
boinensis, S_. calcaratus, S. javensis.

Java

Haemolaelaps casalis, H. ornnitectus, Laelaps nuttalli, Echinolaelaps

sanguisugus, Steatonyssus lavensis, Liponyssoides murinus, Haemoga-
masus quadrisetatus, Pneurnonyssus simicola.

Sumatra

Laelaps longulus, Echinolaelaps sculpturatus, Tricholaelaps comatus,

Pneumonyssus simicola.

ISLANDS OF THE PACIFIC

Formosa

Ornithonyssus bacoti, Liponyssoides muris.

-214

Guam

’
Permariyssus gallinae. "
" .’.". ’... ;;:

Hawaii

Laelaps nuttalli, Echinolaelaps echidninus, Ornithonyssus bursa.

Japan

Haenaolaelaps disimilis, H. glasgowi, H. macroventrali.s, Laelaps jett- ,

mari, L^. nuttallt, Echinolaelaps echldninus, Ornithonys sus bacoti, 0.
sylviarum, Hirstionyssus arcuatus, H.
carnifex, jl_. isabeU.inus, Eu-
laelaps stabularis, Haeinogamasus air.bulans, H. japonicus, _H. mand-
schuricus, H. quadrisetatus. :: ’

.’ 1

.
,
Juan Fernandez’Islands

Echmolaelaps pallidus. . ; .. : .;,.,

.
Lady Julia Percy Island

Orthohalarachne reflexa. ..

Marquesas ’Islands

Laelaps nuttalli., Echinolaelaps echidninus. ,.

^
New Guirie’a. .

Mesolaelaps anomalus, Mysolaelaps roths childi, Ornithonyssus bursa,

Rhinonyssus nova-guinea, OphioEnegistus buloloensis, 0. luzonensis.

New. Zealand .
.
.
Trichosurolaelaps crassipes, Ornithonyssus bacoti, Dermanyssus gal-
linae ; ’!’

Ocean Island

Echinolaelaps echidninus. ...

Philippines

Haemolaelaps bibbyi, Ophiomegistus luzonensis.

Pribilof islands

;
Orthohalarachne attenuata. ’
’
.
 215

Samoa (Ros.e Island)

Echiriolaelaps echidninus. ., ’

Solomon Islands (Ve.lla Lavella)

Spinturnix lavellanus,

.Hosts

The parasitic Mesostigmata parasitize only the three higher classes

of vertebrates: reptiles, birds and mammals.
In the Reptilia only the snakes and lizards are affected to any degree.
There is only one published record from a chelonian and none at all from
the crocidilians.
Warm blooded animals are most frequently parasitized. Probably
all species of birds are affected either by internal or external mites, or
both. It is probably safe to say that all birds are subject to infestation
by external mites during the nesting period. During the non-nesting
period they are probably free of mesostigmatid mites unless they habit-
ually return to the same roosting areas. Internal mites of birds are
very common and perhaps all species of birds will eventually be shown
to have them. :
’
Perhaps all mammals can serve as hosts, but primarily only those
that live in burrows or nests, or congregate at common meeting places
are affected. Mammals that are individualistic or that do not have nest-
ing places, or that are primarily aquatic may hot have external mites,
although they may be attacked by the mites of other animals.
The majority of hosts, therefore, are found among the Rodentia,
Insectivora, Chiroptera, small Marsupialia, and the small, burrowing
Carnivora such as the weasels, skunks, etc. Some of the scaly Eden-
tata have mites that hide under the scales and remain permanently with
the host. The lower Primates and the Pinnepedia have internal mites
in the respiratory system. Cattle may have mites in the internal meatus
of the ear.
It should be noted thai, in the following host list, all published rec-
ords have been taken at face value. It is not only possible but in fact
quite likely that sometimes mites were incorrectly identified. Also,
the host list makes no indication of which mite is the most frequently
found on a certain host; If only one specimen of a species has been found
only once on a host, it appears just as conspicuously in the list as a spe-
cies that has been found repeatedly and in large numbers on that same
host.
Wehave.listed the hosts according to the following plan. The mam-
mals are listed first, followed by the birds and then the reptiles. Under
216

each class, the orders are listed phylogenetically, starting .with .the .most
primitive, and under each order the genera’and species or common names
are listed alphabetically. The latter appear in quotations. Under each
host species, the mite parasites are’listed in the same order as they ap-
pear in the text. ...:,, ;.
All of the host records are given as they appeared-uithe reviewed pub-
lications. No attempt was made to combine synonyms or to give the
probable scientific name for hosts that were recorded only by the com-
mon name.

HOST L-IST

MAMMALIA

Marsupialia

Antechinus flavipes . Marmosa marsupialis , . .

Laelaps sminthopsis.. .
Ornithonyssus braziliensis

"Bandicoot"
Haemolaelaps marsupialis
Metachirops opossum ,
Gigantolaelaps goy.anensis
Heferplaelaps antipodianus
Mesolaelaps australiensis "Opossum" ;

.
M.’ anomalus .. Echinolaelaps echidninus
Ornithonyssus bnrsa Ornithonys sus bacoti
Neoparala’elaps bandicoota Eulaelaps’’ stabularis

Dtdelphis aurita Perameles sp.

Ornithonyssus braziliensis Mesolaelaps anomalus
Neoichoronyssus wernecki
Perameles gunni
Didelphis virginiana Mesolaelaps australiensis
Haemolaelaps glasgowi Heterolaelaps antipodianus
Neoichoronyssus wernecki ;
’’Echinolaelaps .echidninus
Haemogamasus liponyssoides
Perameles ma.crura
Isoodoh obesulus Mesolaelaps anomalus
Heterolaelaps antipodianus M. australiensis

Isoodon torosus : Perameles nasuta

Heterolaelaps antipodianus Mesolaelaps anomalus
Mesolaelaps australiensis Heterolaelaps antipodianus
Potorous tridactylus Thylogale eugenii
Echinonyssus validipes Austrolaelaps mitchfelli.

Sminthopsis 1-eucopus TrichpSurus vulpecula

Laelaps sminthopsis Trichosurolaelaps crasaipes

Thylacomys lagotis (=Thalacomys

lagotis!)
jVlesolaelaps lagotisinus
M. thalacomys
Insectivora

Blarina brevicauda "Mole" (contd.)

Haemolaelaps glasgowi Hirstionyssus isabellinug
Laelaps pachypus Haemogamagus harperi
Hirstionyssus arcuatus H. hirsutus
Myonyssus jamesoni H. liponyssoides
Eulaelaps stabularis
Haemogamasus alaskensia
Hi ambulans Myosorex sp>
Hi barberi Laelap^ longiventris
Hi occidentalis
Hi liponyssoides My os or ex varius
Laelaps parvulus
Condylura cristata
Haemogamasus liponyssoides Neomys fodiens
Hirstionyssus eusori.cis
Cryptotis parva Laelaps hilaris
Haemolaelaps glasgowi Haemogamasus ambulans
Hirstionyssus arcuatus
H. occidentalis
Haemogamasus harperi Neurotrichus sp. : ’’ ’

Hirstionyssus obsoletus
"Elephant shrew" Haemogamasus occidentalis
Orn ithonys sus aethiopicus
Steatonyssus nyassae Neurotrichus gibbsi
Hirstionyssus arcuatus
"Maryland shrew"
Haemogamasus liponyssoides Paras calops breweri
Hirstionys sus arcuatus
Mogera robusta Haemogamasus liponyssoides
Laelaps clethrionomydis
Oryctolaelaps bibikovae Scalopus aquaticus
Haemolaelaps glasgowi
"Mole" Haem ogamasu s harperi
Hirstionyssus arcuatus H. liponyssoides
218

Scalopus argentatus Sorex minutus

Haemogamasus’liponyssoides Hirstionyssus soricis ,’.’.

Haemogsmasus horridus

.
Scapanus latimanus
Haenaolaelap’6- glasgowi .’.t’.’J. Sorex obscurus ’-

.
Laelaps pachypus
Scapanus orarius
Haemogamasus occidentalis Sorex-’palustris ;
.. ;

-
Haenaogamasus ambulans
Scapanus townsendi H. liponyssoides -.’:. .,;
Haemolaelaps glasgowi
Haemogamasus occidentalis Sorex trowbridgei
Hirstionyssus obsoletus
Scaptochirus gilliesi Haemogamasus occidentalis
Haemogamasus mandschuricus H. keegani

"Short-tailed shrew" Sorex vagrans,,

Haempgamagjts liponyssoides Haemolaelaps glasgowi

.
H. barberi ’; .’. ? Laelaps, paehypus

"Shrew" Suncus varilla ; . :.

Ornlthonyssus bacoti Laelaps vansomereni

.
Haemogamasus liponyssoides
Talpa alpina
Solenodon paradoxus Haemogamasus hirsutus

.
Steatonyssus spinosus H. horridus .’

Sorex sp. Talpa altaica

Hirstionyssus carnifex Haemolaelaps glasgowi
Eulaelaps stabularis
Sorex araneus Haemogamasus mandschuricus
Haemolaelaps glasgowi H. nidi
Eulaelaps stabularis TH. liponyssoides
Haemogamasus ambulans
H. horridus Talpa europaea
H. nidi Haemolaelaps fahrenhoizi
H_.glasgowi
Sorex cinereus Laelaps agilis
Hirstionys sus arcuatus L. hilaris
Haemogamasus alaskensis Hirstionyssus arcuatus
H. liponyssoides H_. carnifex
H. isabellinus
H_. pachypus
Sorex fumeus Myonyssus gigas
Haemolaelaps glasgowi Eulaelaps oribatoides
Hirstionyssus arcuatus E_. stabularis
Myonyssus jamesoni Haemogamasus ambulans
Haeinogamasus liponyssoides H. hirsutus
Talpa europaea (contd.) Tupaia picta
Haemogamasus horridus Echinonyssns nas’ntus .

H. nidi
H_. liponyssoides. .’.’.., "Townsend mole"
Haemogamasus ambulans
"Tikoes moende"
Laelaps soricis Urotrichus talpoides
Haemogamasus quadrisetatus

Chiroptera

Anoura geoffroyi "Chiropteron’species "

Periglischrus vargasi Steatonyssus javensis
.
Artibeus jamaicensis Corynorhinus raflriesquei
Ichoronyssus kochi Ichoronyssus lon’gisetosus
Spinturnix artibiensis
^. ewingia Desmodus roturidus
Periglischrus meridensis Radfordiella oudemansi

Barbastella barbastellus Eidolon helvum

Ichoronyssus flavus. Spinturnix kenyaensis

"Bats" Epoinophorus minor

Ornithpnyssus aethiopicus’ Spinturnix aethiopicus
Ichoronyssus. australicus
Hirstionyssu.g cagnifex Eptesieus fuscus
H. hirati "" . Ichoronyssus quadridentatus
Hirstesia kenyaensis Spinturnix americanus
H. sternalis ’.’ ’

Spelaeorhynchus praecursor Eptesieus serotinus

S. latus. Laelaps hilaroides
Spinturnix abyssinicus Hirstionyssus arcuatus
^. amboinensis Steatonyssus musculi
S. antipodianus
S. javensis Glossophaga soricina
S_. jordani Steatonyssus joaquimi
S. myoti.
S. novae -hollandae Isotus ciliatus
?. grossus Steatonyssus murinus
Chaerephon leucostigma
Ichoronyssus forsythi Lasiurus borealis
Steatonyssus occidentalis
Chaerophon pumilis !
Ornithonyssus aethiopicus Leptonycteris nivalis
Haemogamasus liberiensis Spinturnix ewingia
220

Leptonycteris nivalis (contd.) Myotis dasycneme

Periglischrus vargasi Hirstionyssus arcuatus

Liponycteris nudivenWis^."; Myotis daubentonii ,^’"..1

-
Steatonyssus sudaqensisi^ Ichoronyssus flavus ,..>.

’
.
Haemogamasus ambuians .

Macroglos sus minimus’’ ".’’t’’ Spinturnix vespertillpni’s

Spinturnix macrogl&sisy"
Myotis grisescens
Macrotus californicuS.
Periglischrus yarga-si . ^"’ Haemolaelaps glasgowi

Myotis lucifugus
:.

Micropteropus pusillus Ichoronyssus robustipes

Spinturnix aethiopicus Steatonyssus occidentalis
Spinturnix americanus
Miniopterus natalensis S. echinipes
Neospinolaelaps miaiopteri S^. iowae
Hirstesia transvaalensi.8 Haernogalnasus alaskensis
Steatonyssus natalensis
Spinturnix semilunaris Myotis myotis (=M. murinus)’
Hirstionys sus arcuatus
Miniopterus schreibersi Ichoronyssus; flavuS’.:
Macronyssus longimanus Steatonyssus murinus ’-: :;
Ichoronysgus granulosus Spinturtfiji rnurinus ’:
’

I. scutatus S^. vespertilionis

Spinturnix psi
Myotis mystacirius
Ichoronyssus flavus
Molossus abrass’tis . Steatonyssus musculi
Ichoronyssus haematophagus Spinturnix vespertilionis

Molossus nasutus ’^::. Myotis nigricans .

’

^lehijronyssus venezolanus Ichoronyssus hasei

Spinturnix araguensis
Molossus nigricans
Periglischrus vargasi Myotis ruber
Ichoronyssus kolenati
Molossus rufus
Ichoronyssus haematophagus Myotis tricolor
Spinturnix lawrencei

Myotis sp. Natalus mexicanus :

Steatonyssus occidentalis Ichoronyssus granulosus
Spinturnix psi L robustipes
Spinturnix carloshoffmanni
Myotis capaccinti
Steatonyssus murinns "Noctule bat"
Spinturnix vespertilionis Spinturnix acuminatus
Nyctalus noctula Pteropus edulis
Ichoronyssus flavus Neolaelaps magnistigmatus
Steatonyssus musculi
Pteropus giganteus
Nycteris sp. Neolaelaps magnistigmatu s
Steatonyssus brucei
Pteropus lavellanus
Nycticeius hurqe rails Spinturnix lavellanus
Steatonyssus ceratognathus
Pteropus rufus
Nyctinomus bocagei_ Spinturnix calcaratus
Ichoronyssus nyctinomi
Pteropus vampyrus
Nyctinomus brasiliensis Neolaelaps magnistigmatus
Ichoronyssus robustipes
Pterygistes noctula
Nyctinomua macrotis Hirstionyssus arcuatus
Ichoronys sus hae mat ophagu s H. carnifex

Panugo noctula Rhinolophus sp.

Ichoronyssus flavus Spinturnix ianzai

Rhinolophus blasii
Pipistrellus sp. Periglischrus inter ruptus
Steatonyssus murinus
Rhinolophus clivosus
Pipistrellus kuhlii Macronyssus longimanus
Steatonyssus murinus Ichoronyssus leprosus
I. scutatus
Pipistrellus nanus Periglischrus glutinimargo
Steatonyssus eos
Spinturnix walkerae Rhinolophus euryale
Spinturnix euryalis
Pipistrellus pipistrellus
Hirstesia britteni Rhinolophus ferrum-equinum
Steatonyssus cyclaspis (= R. hippocrepis)
S. murinus Haemolaelaps glasgowi
Ornithonyssus tinae
Pipistrellus stampflii Ichoronyssus scutatus
Spinturnix pipistrellus Spinturnix euryalis
S. psi
Plecotus auritus S. oudemansi
Steatonyssus murinus S. myoti
Spinturnix plecotinus Periglischrus hipposiderus
S. myoti
Rhinolophus geoffroyi
Pteropus aegyptiacus Ornithonys sus rhinolophi
Spinturnix lateralis Spinturnix viduus
222

Rhinolophus geoffroyi (contd.) Vesper sublatus 1

Periglischrus africanus Ichoronyssus crosbyi

Rhinolophus hipposideros
Spinturnix omahoriyi Vespertilio auraus

"
.
Hirstesia steriialis’’ ’ ’" Ichoronyssus flavus
:

’
’
Rhinolophus lobatus

’
Vespertilio dasycneme
Ichoronyssus jacksont Hirstionyssus arcuatus

Vespertilio rnurinus
Rhinopoma microphyllum Hirstionyssus arcuatus
Spinturnix zeiebori Ichoronyssus spinosus
Ichoronyssus lepidopeltis I. flavus

Rousettus aegyptiacus Vespertilio myotis

Macronyssus longimanus Ichoronyssus granulosus

Rousettus lanosus Vespertilio nathusii

,
.
Hirstesia transvaalensis Ichoronyssus flavus

Scotophilus murinonavus Vespertilio nattereri

Spinturnix scotophili Ichoronyssus mohrae
Steatonyssus^ nyassae
Vespertilio nilssonii
Sturnira lilium Ichoronyssus flavus
Penglischrus yargasi
Vespertilio noctula
Synotus barbastella Hirstionyssus arcuatus
Ichoronyssus flavus’ Ichoronyssus flavus
Tadarida aegyptiaca Vespertilio pipistr.ellus
Ichoronyssus cornutus Ichoronyssus flavus
Steatonyssus musculi
Tadarida brasiliensis
Ichoronyssus robustipes Vespertilio seratinus
Hirstionyssus arcuatus
Tadarida cynocephala Steatonyssus musculi
Ichoronyssus robustipes Spinturnix myoti

Tadarida mexicana Vespertilio superans

Ichoronyssus robustipes Steatonyssus superans
S. musculi
Tadarida teniotis
Ichoronyssus hoogstraali
Vesperugo noctula
Vampyrops lineatus Hirstionyssug arcuatus
Periglischrus ihertngi Ichoronyssus flavus
 2,23.

Vesperugo pipistrellus "Whiskered bat"

Steatonyssus musculi Ichoronyssus granulosus
Ichoronyssus diversipilus

Primates

Cercopithecus aethiops Lemur mongoz

Pneumonyssus santos-diasi Aetholaelaps sylstrai

Cercopithecus schmidti
Pneumonyssus duttoni Macaca mulatta (=Pithecusthesus)
Pneumonyssus dinoiti
Cercopithecus; ascanius P. simicola .

Rhinophaga cercopitheci

Cercopithecus mitis Papio sp.

Rhinophaga cercopitheci Pneumonyssus congoensis
Colobus badius
Pneumonyssus rodhaini, Papio doguera ,; ,

.
Rhinophaga papionis
Galago alleni
Haemolaelap.s galagus
Hirstionyssus ngami Papio ursinus . .

.
Pneumonyssus. santos-diasi
Galago senegalensis P. mossambicensis
Ornrthonyssus galagus

Homo sapiens Pithecus fascicularis

Ornithonyssus bacoti Pneumonyssus simicola
braziliensis
0. bursa
0. iheringi Pithecus nemestrinas
0. sylviarum Pneumonyssus simicola
Ophionyssus natricis
Allodermanyssus sanguineus
Dermanyssus gallinae Pithecus rhesus (=Macacus rhesus)
Pneumonyssus simicola
Lagothnx poppigi
Pneumonyssus stammeri
Propithecus verreauxi
"Lemur" Liponysella madagascariensis
Liponysella madagascariensis

Lemur albifrons Theropithecus gelada

Liponysella madagascariensis Pneumonyssus simicola
2’2-t

Edentata

Bradypus tridactylus Myrmecophaga tridactyla

Ornithonyssus iheringi Sauronyssus myrmecophagus

Euphractus sexcinctus
Dasyponyssus neivai

Pholidota

Manis teniininckii Manis tricuspis

Manitherionyssus heterotarsus Manisilaelaps- coronis

Lagomorpha

"Cottontail" Ochotona rutila :

Ornithonyssus bacoti AIlodermanyssus sanguineas
Haeniogamasus ambulans

Ochotona schisti-ceps
Lepus californicus Hirstionyssus occidentalig
Haemolaelaps .glasgowi Myonyssus montanus

Ochotona alpina "Rabbit"

Haemolaelaps glasgowi Heterolaelaps antipodianus
Haernogamasus dauricus
H. naandschuricus
H. ambulans Sylvilagus sp.
H. kitanoi Haemolaelaps glasgowi
H. serdjukovae

Sylvilagus auduboni.
Ochotona daurica Haemolaelaps glasgowi
Haemolaelaps glasgowi Brevisterna morlani
Eulaelaps cricetuli
E. kolpakovae
Haemogamasus mandschuricus Sylvilagus minensis
H.. kitanoi .
Ornithonys sus braziliensis
.
Ochotona pusilla "Texas cottontail"’
Haemolaelaps;;glasgowi .Echinolaelaps echidninus
 Rodentia

Acomys sp. Allactaga saltator

Liponyssoides muris Haemolaelaps glasgowi
H. semidesertus .y
Acomys

"
"
caharinis , Eulaelaps .cricetuli’:;,
Ornithonyssus bacoti E. kolpakovae

.
Allodermanyssus sanguineus
A_. aegyptius Alticola argentatus
Allodermanyssus sanguineus
Acornys russatus
Allodermanyssus sanguineus Ammospermophilus leucurus
:
Haemolaelaps glasgowi
Acomys dimidiatus ".
Allodermanyssus sanguineus Anomalurus fraseri
Haemolaelaps spatuliformis
Aethomys chrysophilus
Laelaps vansomereni Aplodontia rufa
Echinolaelaps muricola Patrinyssus hubbardi
Alphalaelaps aplodontiae
Aethomys namaguensis
Laelaps yansomereni Apodemus sp.
Laelaps pachypus
"African rat" Haemogamasus arobulans
Laelaps yansomereni
Apodemus agrarius .’’

’
Akodon mollis Haemolaelaps fahren’hoizi
Eubraclr^laelaps rotundus H. glasgowi
Laelaps agilis
Alactagulus^ acontion L_. algericus
Haemolaelapa semidesertus L_. }ettmari~
Eulaelaps stabularis
E. kolpakovae L_. pavlovskyi
Hirstionyssus arcuatus.
Haemogamasus crtelli H. arvicolae
H. isabellinus
"Alaska short-tailed mouse" H. musculi
Haemogamasus ambulans Eulaelaps stabularis
"Albemarle Haemogamasus ambulans
meadow mouse"
H. dauricus
Haemogamasus barberi H. ellipsoideus
H. hirsutosimilis
H. horridus
Allactaga elater
Haemolaelaps semidesertus
" H. kusumotoi
H. liponyssoides
H. mandschuricus
Allactaga jaculus
Haemolaelaps semidesertus H_. nidT
H. serdjukovae
226

Apodemus flavicollis Apodemus sylvaticus (contd.)

Haemolaelaps fahrenhoizi Laelaps agilis ,;

H_. glasgowi’ L. caucasicus

’
.

,
Laelaps agilis I^_. mosquensis

’
.
L. caucasicus L^. volgensis .,.
L. mosquensis ’. . Ornit.honyssus bacoti
Hirstionyssus arcuatus Hirstionyssua arcuatus
H. carnifex H. isabellinus.^.

-
.

H. isabeUinus ’ ’. H. musculi
H^inusculi- Allodermanyssus .sanguineus

’
"
’
Myonyssua decumani Myonyssus decumani ... .

M. gigas ~~’^.::;;’. KI. gigas

Eulaelaps stabularis J. M. rosslcus . .

Haemogamasus ambulans
~
Eulaelaps stabularis
H. hirsutosimilis~ Haemogamasus ambulans
H. hjrsutus H. hirsutosimilis
H. nidi H. hirsutus

.
H. horridus
Apodemus fulvipectus H. mandschuricus
Laelaps agilis H. nidi
L. caucasicus
Arvicanthis abyssinicus
Apodemus geisha
"

Haemolaelaps arvicanthis
Eulaelaps stabularis "
H. murinns .:,,.

.
Haemogamasus mandschuricus Echinolaelaps giganteua

Apodemus hebridensis Arvicanthis agranus -

Haemogamasus ambulans Laelaps agrarius

Apodemus speciosus Arvicanthis barbarus

Haemolaelaps glasgowi Echinolaelaps .bakeri
Laelaps clethrionomy^is
L. pavlovskyi Arvicanthis dorsalis
Myonyssus dubtnini Haemolaelaps zulu
Eulaelaps cricetulii. Laelapg nuttalli
E. stabularis
~
L_. parvulus
Haemogamasus ambulans
~~
Echinolaelaps giganteus
H.
’
dauricus E_, muricola
H. japonicus
H. liponyssoides ;;." .
Arvicanthis niloticus
’""

H. mandschu’r’i.cus Haemolaelaps hystrici

.
,
’~~~
H_. nidr H-’ Jflsculptus
H. quadnsetatus H. niurinus. .
’
H. serdjukoyae ;

H. zulu
Laelaps keegani
Apodemus sylvaticus L_.^ lamborni- .

Haemolaelaps fahrenhoizi. Qrnithonyssus bacoti

H. glasgowi Allodermanyssus sanguineus
Arvicanthis niloticus (contd.) Baiomys taylori
Allodermanyssus aegyptius Haemolaelaps glasgowi
Liponyssoides muris
Eulaelaps stabularis Bandicota malabarica
Laelaps taprobanius
Arvicanthis rufinus
Laelaps melomys
’ Bathyergus suillus
Lr. spinifer Haemolaelaps bathyergus

Arvicola arvalis "Black rats"

Laelaps arvicolae Ornithonyssus bacoti
L. muris
L_. pachypus "Brown rat"
Hirstionyssus carnifex Echinolaelaps echidninus
Haemogamasus horridus Ornithonyssus bacoti
Eulaelaps stabularis
Arvicola amphi.bi.us
Haemolaelaps fahrenhoizi Callosciurus quinque stnatus
Laelaps pachypus Haemolaelaps traubi
Hirstionyssus arcuatus
H. i.sabellinus Cavia aperea
Eulaelaps stabularis Ornithonyssus bacoti
Hae mogama sus hirsutus 0. braziliensis

Arvicola terrestris Cavia rufescens

Haemolaelaps glasgowi Neoparalaelaps bispinosus
H. semidesertus
Laelaps amphibius Caviella australis
L. muris Cavilaelaps bresslaui
’
Hirstionyssus arvicolae . Ornithonyssus hirsti
~
H. isabellinus
Eulaelaps kolpakovae Cercomys cunicularis
E. stabularis Lepronyssoides matogrosso
Haemogamasus ambulans
H. citelli Chinchillula sahamae
H. horridus Haemolaelaps chinchillulae
H. liponyssoides ’.
fl. nidi "Chipmunk"
H. oudemansi Eulaelaps pedalis
Haemogamasus ambulans

Atherurus africanus
Rhmophaga atheruri Choeromys gregorianus
R. leopoldi Echinolaelaps muricola

Auliscomys boliviensis Chraeomys pulch.errimus !

Haemolaelaps glasgowi Haemolaelaps glasgowi
228

Citellus sp. Citellus richardsonii (contd.)

Ischyropoda armatus Ischyropoda armatus ; .. .,..,.,

Citellus armatus Citellus suslicus ; . .

:

.
Haemolaelaps glasgowi Hirstionyssus crieeti .
..,,.
Citellus beecheyi Citellus tridecemlineatus
Haemolaelaps glasgowi Haemolaelaps glasgowi :

.
Eulaelaps citellus
Citellus undulatus
Citellus citellus Haemolaelaps glasgowi .

Hirstionyssus crieeti Haemogamasus ambulans

Haemogamasus citelli
Clethrionomys sp.

-
Citellus columbianus Hirstionyssus obsoletus
Haemolaelaps glasgowi
Clethrionomys amurensis
Citellus dauricus Haemogamasu-s ambulans.
Eulaelaps stabularis H. mands fchur icu s
Citellus dauricus Clethrionomys frater
Haemogamasus kusumotoi Laelaps clethrioriomydis
Hirstionyssus transiliensis
Citellus lateralis
Haemolaelaps glasgowi Clethrionomys gapperi.
Hirstionyssus occidentalis Haemogamasus alaskensis
H. ambulans
Citellus major H. liponyssoides
Hirstionyssus crieeti
Clethrionomys glareolus
Citellus mexicanus Haemolaelaps fahrenhoizi
Haemolaelaps glasgowi H. glasgowi
Ornithonys sus bacoti
’

Laelaps agilis ’ :
.
L_. clethrionomydis
Citellus mollis L. hilaris ;
Haemolaelaps glasgowi L_. pachypus
Hirstionyssus arcuatus
Citellus pygmaeus H. carnifex
Haemolaelaps androgynus Myonyssus ingricus
H. glasgowi Eulaelaps stabularis
.
.
H. longipes Haemogamasus ambulans
H. semidesertus H_. hirsutosimilis
Hirstionyssus crieeti H. hirsutus
Haemogamasus citelli H. horridus
Eulaelaps kolpakovae H. nidi

Citellus richardsonii Clethrionomys rufocanus

Euhaemogamasus barberi Haemolaelaps glasgowi
Clethrionomys rufocanus (contd,) Cricetulus barabensis (contd.)
Laelaps arvalis Haemogamasus ellipsoideus
L. clethrionomydis H. kusumotoi

’
.

Eulaelaps stabularis
Haemogamasus ambulans Cricetulus griseus
H. dauricus Laelaps jettmari
H. liponyssoides

’
H. mandschuricus ’

Cricetulus migratorius

-
.
H. serdjukovae ’."’ Haemolaelaps glasgowi
Laelaps ekstremi
Clethrionomys rutilus L_. jettmari
Haemolaelaps glasgowi Allodermanyssus sanguineus
Laelaps clethrionomydis Eulaelaps stabularis
Hirstionyssus isabellinus Haemogamasus nidi
Myonyssus dubinini ’
’
Eulaelaps stabularis Cricetulus triton
Haemogamasus ambulans Haemolaelaps glasgowi
H. ellipsoideus Hirstionyssus criceti
H_. nidi Eulaelaps stabularis
Haemogamasus dauricus
Coelomys bi color H. mandschuricus
Laelaps atypicus
Cricetus auratus
Cpelogenys p-acca Hirstionyssus criceti
Haemolaelaps coelogenys
Cricetus cricetus
"Cotton mouse" Haemolaelaps glasgowi
Ornithoriyssus bacoti Hirstionyssus criceti
Eulaelaps stabularis Eulaelaps stabularis
’
Haemogamasus liponyssoides Haemogamasus nidi

"Cotton rat" Cricetus eversmanni

Ornithonys sus bacoti Haemolaelaps semidesertus
Eulaelaps stabularis Hirstionys sus criceti
H. eversmanni
Cratogeomys castanops
Haemolaelaps geomys Cricetus triton
H. glasgowi Haemogamasus dauricus

Cricetomys sp.
Echinolaelaps muncola Crocidura manni
Haemolaelaps hystrici
Cricetulus barabensis
’
Haemolaelaps glasgowi Crocidura olivieri
Hirstionyssus criceti Haemolaelaps murinus
Eulaelaps cricetuli H_. zulu
E. stabularis Laelaps algericus
Haemogamasus ambulans L. nuttalli
 230

Cryptomys bigalkei Dipodomys ordi ,

; ,:
Haemolaelapa eloffi ’
Haemolaelaps glasgowt,
Hirstionyssus hilli

.
Cryptomys capensis H. incomptus
Haemolaelaps cryptomiug Ischyropoda furmani, ’. ’.

’
Cryptomys natalensis Dipus sowerbyi ,
;;
Haemolaelaps natalensis Haemogamasus mandschuricus
Eulaelaps cricetuli
Ctenodactylus gundi

’
Steatonyssus murinus "Domestic mice"
Allodermanyssus sanguineus
C-ynomys sp. .
.’’;,: :.
Hirstionyssus cynpmys’ ’.’ "Domestic rat"
Ornithonyssus bacoti
Cynomys gunnisoni Dermanyssus gallinae
Haemolaelaps glasgowi .;.:;’ Eulaelaps stabularis
Brevisterna morlani
Dyromys nitedula
Cynomys ludoviclanus Ornithonyssus dogieli
Haemolaelaps glasgowi
Hirstionyssus blarichardi
Echiroys sp.
Gigantolaelaps gilmorei
. Dasymys helukus G_. oudemansi
Haemolaelaps dasymys
Ellobius fuscocapillus
Dasymys incomtus Haemolaelaps ellobii .
.

Haemolaelaps murinus Hirstionyssus eilobii

Laelaps roubaudi
Echinolaelaps giganteus Ellobius talpinus
Haenaolaelaps ellobii
Dasymys rufulus Hirstionyssus blanchardi
Echinolaelaps giganteus
Epimys defua
Laelaps liberiensis
Dephomys dephua Echinolaelaps giganteus
Echinolaelaps giganteus
Epimys norvegicus
Dipodomys deserti Echinolaelaps echidninus
Ischyropoda armatus Ornithonyssus braziliensis

Dipodomys merriami Euryzygomatomys spinosus

Hirstionyssus triacanthus Haemolaelaps reticulatus
Ischyropoda armatus Ornithonyssus braziliensis

Dipodomys microps Eutamias sp.

Haemolaelaps glasgowi Haemolaelaps glasgowi
Eutaroias alpinus Geomys bursarius
Ornithonyssus sylviarum Haemolaelaps geomys
H_.glasgowi
Eutamias minimus’ Hirstionyssus geomydis
Haemolaelaps glasgowi
Hirstionyssus affinis Geomys cumberlandicua
Haemolaelaps geomys
Eutamias stbericus H. glasgowi
Haemolaelaps glasgowi
Hirstionyssus isabeUinus Geomys floridanus
Eulaelaps stabu’laris Haemolaelaps geomys
Haemogamasus ambulans H_. glasgowi
H. maiids churicus
H. serdjukovae . Geomys lutescens
Haemolaelaps geomys
H. glasgowi
Evotomys sp.
Dermanyssus evotomydis Geomys personatus
Haemolaelaps geomys
Evotomys dawsoni H, glasgowi
Haemogamasus ambularis.
Geomys texensis
Evotomys gapperi Haemolaelaps geonays
Haemolaelaps glasgowi
Geomys tuza
Haemolaelaps geomys
"Field mice" H. glasgowi
Laelaps oraniensis
Haemogamasus alaskensis Georychus sp.
H. ambulans Haemolaelaps spinitarsus
H. horridus
Georychus capensis
"Field rat" Haemolaelaps lawrencei
Gigantolaelaps comatus
G. goyanensis Georychus hottentotus
G. mattogrossensis Haemolaelaps capensis
G. oudemansi
Haemogamasus ambulans "Gerbille"
Laelaps buxtoni
"Field vole"
Hirstionyssus arcuatus Gerbillus sp.
Haemogamasus ambulans Ornithonyssus bacoti

"Flying squirrel" Gerbillus gerbillus

Haemogamasus ambulans Hae molaelaps aegyptius
H. ewingi
"Fox squirrel" H. insculptus
Eulaelaps stabularis Hirstionyssus craticulatus
23:2-

Gerbillus hir-tipe-s-’,’ :"!.’.... Heteromys anomalus >.i ’

.
Haemolaelaps ro:auritani’cus Steptolaelaps’heteromys’
-’-* ’’:-
’ .
Gerbillus-faanus ’.. ’"’.:.’ Holochilus sciurei^s’,. .
Haemolaelaps insculptus Liponyssoidea. brasilietisis
’
Gigantolael’aps ’mattogrossensis
Gerbillus pyramidum
Haemolaelaps aegyptius Holochilus vulpinus
H. insculptus Gigantolaelaps mattogrossensis
G, bracRyspinosus
Gerbillus quadrimaculatus Laelaps manguinhosi : ;

.
.
Haemolaelaps aegyptiu’s ..

"House mouse"
Glaucomys sp. Laelaps hilaris
Haemogamasus ambulans Ornithonyssus bacoti
Hirstionyssus carnifex
Glauconays sabrinus Eulaelaps stabularis
Haeniogaitiasus ambulans Haemogamasus ambulans

Glaucomys volans Hydromys chrysogaster .

Haemolaelaps megaventralis Laelaps muris

Haemogamasus ambulans
Isothrix bistr.iatus
"Golden mouse" Bolivilaelaps tricholabiatus
Ornithonyssus bacoti
Jaculus iaculus
"Gray squirrel" Haemolaelaps a,egyptius
Ornithonyssus bacoti H. insculptus
Haemogamasus ambulaii’s...... Hirstionyssus craticulatus
"Groundhog" "Kapuka"
Eulaelaps propheticus Laelaps lamborni

Kerodon spixi
"Hamsters" Lepronyssoides pereirai
Ornithonyssus. bacoti Cavilaelaps braziliensis

Heliosciurus rufobrachiatus
Hirstionyssus libertensis’ Lagurus curtatus
Hirstionyssus isabellinus
Hapalotis sp.
Laelaps hapaloti Lagurus lagurus
: Haemolaelaps ellobii ;

Hesperomys vulpinus Haemogamasus nidi

.
Gigantolaelaps maximus
Leggada musculoides
Heteromys sp. Laelaps lavieri
Steptolaelaps heteromys’; L. grenieri
Lemmus sp. Lophuromys aquilus (contd.)
Eulaelaps stabularis Eubrachylaelaps lophuromius
Steptolaelaps liomydis
Lophuromys sikapusi
Lemmus arvalis Laelaps lavoipierrei
Laelaps pachypus

Lemmus lemmus "Majancha"

Hirstionyssus isabellinus Laelaps lamborni
Haemogamasus nidi
Marmota bobak
Lemmus obensis Haemogamasus kitanoi
Laelaps lemmi ’ H. mandschuricus
Haemogamasus nidi
Marmota marmota
Lemniscomys barbarus Hirstionys sus blanchardi
Echinolaelaps giganteus
Marmota monax
Lemniscomys griselda Haemolaelaps glasgcwi
Haemolaelaps zulu
-
E chinolaelaps giganteus Marmota sjbirica
Haemolaelaps glasgowi
Lemniscomys macculus Haemogamasus dauricus
Echinolaelaps giganteus H. kitanoi
.
.
Lemniscomys stnatus
H-- mandschuricus

Haemolaelaps murinus "Marquesas rat"

H. tachyoryctes Echinolaelaps echidninus
Laelaps grenieri
L. zumpti Mastomys coucha
Echinnlaelaps giganteus Haemolaelaps labuschagnei
E. muricola Laelaps nuttalli :
L_. vansomereni
Liomys irroratus Echinolaelaps giganteus
Steptolaelaps liomydis E_. muricola

Liomys pictus . "Meadow mouse"

Steptolaelaps liomydis Haemogamasus-alaskensis
H_. barberi
"Long-tailed field mouse" H_. liponyssoides
Laelaps muris

Lophiomys ibeanus Melomys gervinipes

Echinolaelaps. muricola My solaelaps roths childi

Lophuromys aquilus Melomys littoralis

Haemolaelaps mur inu s Laelaps nuttalli
H. sudanicus Mysolaelaps rothschildi
 234

Meriqnes crassus Micromys minutus (contd.)

Hatemolaelaps aegyptius Haemogamasus ni4i

.
.
Meriones erythrourus !
Haemolaelaps angustiscuUs Microtus sp.
H_. longipes ; . Laelaps alaskenais
’-Hirstionyssus meridianus Haeinogamajsus-aiaskensis
Eulaelaps stabularis H. ambulans

Meriones libycus .-’ Microtus agrestis

Haeinolaelaps aegyptius Haemolaelaps fahrenhoizi

.
H. glasgowi
Meriones meridianus ’..’.. . Laelapa arvalis
Hacaiolaelaps longipes L^. hilaris
Hirst! onyssus meridianus L_.pachy pus
Eulaelaps stabularis Haemogamasus ambulans
H. nidi
Meriones persicus
Haeinolaelaps androgynus Miorotus amphibius
H. gl’asgowi - Laelaps arvicolae
niuns*"
.i Allodernianyssus sanguineus L.
Eulaelaps stabularis L. pachypus

Meriones.rex Microtus arvalis

.Haeinolaelaps namrui Haeinolaelaps fahrenhoizi
H_. glasgowi
Meriones tamariscinus Laelaps arvalis
Haemolaelaps glasgowi L. ekstremi
B[,; longipes L. hilaris
Eulaelaps cricetuli L_. jettmari
.
L. kolpakovae
Meriones tristrami Htrstibnyssus arcuatus
Haemolaelaps longipes H. arvicolae
-
.
Eulaelaps stabularis . H. isabellinus
Eulaelaps stabularis
"Mice" ’Haemogamasus ambulans
Mesolaelaps australiensis H. hirsutus
Gigantolaelaps peruvianus. H. hon-idus
H. nidi
’
.

Microdipodops megacephalus
Ischyropoda furmani Microtus brandti
Haemolaelaps glasgowi
Microdipodops pallidus Eulaelaps kolpakovae
Ischyropoda furmani Hae mogamasu s mands churieus

Micromys minutus
Haemolaelaps glasgowi .
Microtus breweri
Laelaps micromydis ’

Haemolaelaps glasgowi
 235

Microtus californicus Microtus montebelli (contd.)

Haemolaelaps glasgowi Haemogamasus japonicus

.
Laelaps pachypus
Haemogamasus ambulans Microtus mordax
Haemolaelaps glasgowi
Microtus carruthersi Laelaps pachypus
Haemogamasus ivanovi Haemogamasus liponyssoides

Microtus fortis Microtus nanus

Haemolaelaps glasgowi Haemolaelaps glasgowi
Myonyssus dubinmi
Haemogamasus mandschuricus Microtus nivalis
H. serdjukovae Laelaps hilaris

Microtus gregalis Microtus ochrogaster

Haemolaelaps glasgowi Haemolaelaps glasgowi
Laelaps arvalis Laelaps pachypus
L. clethrionomydis ; ,. , Haemogamasus liponyssoides
L. bilaris
Hirstioriyssus musculi Microtus oeconomus
H. transiliensis Haemolaelaps fahrenholzi
Eulaelaps stabularis.’ H. glasgowi
Haemogamasus ambulans Laelaps hilaris
,
H. mands churicus L. muris
H^. nidi. : L_. pachypus
H. nidiformes L_.pavlovskyi
Hirstionyssus arvicolae
Microtus gud H. isabellinus
Haemogamasus hirsutus Eulaelaps stabularis
Haemoganaasus ambulans
Microtus longicaudus H. horridus
Eubrachylaelaps debilis H. nidi

Microtus rnajori Microtus operarius

Laelaps pitymydis Haemogamasus ambulans
Haemogamasus nidi
Microtus orcadensis
Microtus naontanus Laelaps hilaris
Haemolaelaps glasgowi L. pachypus
Laelaps pachypus Macronyssus orcadensis
Hirstionyssus isabellinus
H. occidentalis Microtus oregoni
Haemogamasus ambulans Haemolaelaps glasgowi
H, liponyssoides Haemogamasus alaskensis
H. alaskensis
Microtus pelliceus
Microtus montebelli Myonyssus shibatai
Eulaelaps stabularis Eulaelaps stabularis
236

Microtus’pelliceus (contd.) Mus coucha ."’’.’ ’.

’
’. . ..

Haemogamasus ellipaoideus liaemolgelaps villo^issim’us

Echinolaelaps, muricola ’..
Microtus pennsylvanicus
Haemolaelaps glasgowi Mus hebridensis
Hirstionyssus arcuatus Haemogamasu.g’horridus
H. iSabellinus
Haemogamasus alaskensis Mus hildebrandti
H. barberi Echinolaelaps muriccila
H. liponyssoides
Mus lepturus ’
Microtus socialis .
Echinolaelaps sanguisugus.
Laelaps hilaris Haemogamasu’s ’quadrisetatus
Haemogamasus nidi
Mus minutus
Microtus subterraneus Laelaps .arvicolae
Hae molaelap s glaagowj.
.
Mus molos sinus
Microtus townsendi Haemogamasus kusumotoi
Haemogamasus occidentalis
Mus musculus
Microtus ungurensis Haemolaelaps androgynus
Haemolaelaps glasgowi
Laelaps clethrionomydis
H^ glasgowi
H. insculptus
L. arvalis H. m egaventralis

’
. .
.
Eulaelaps cricetuli H, murinus
E. stabularis H. reticulatus
Haembgamasus ambulans iH. semidesertus
JH,zulu
"Mouse" Laelaps algericus
.
Echinolaelaps giganteus L. hilaris
E. muricola L, lambdrni . :
Haemogamasus barberi L^. nuttalli :

Mus sp.
t^. sicula
Echinolaelaps echidninus
.
Gigantolaelaps versteegi Ornithonyssus bacoU
G. wolffsohni Hirstionyssus arcuatus
Mysolaelaps rothschildi H. butantanensia
H. carriifex
Mus agrarius H. isabellinus.
Laelaps jettmarj^ H. musculi
Steatonyssus musculi
Mus algericus Allodermanyssus sanguineus
Laelaps algericus ’.;;’: Myonyssus decumani
Eulaelaps kolpakovae
Mus chrysophilus. . E, stabularis
Laelaps vansomereni Haemogamasus ambulans
Echinolaelaps muricola H. hirsutus
Mus musculus (contd.) Myospalax dybowskii (contd.)
Haemogamasus nidi Eulaelaps stabularis
H. oudemansi Haemogamasus dauricus

Mus norvegicus Myospalax psilurus

Echinolaelaps echidninus- Hirstionyssus confucianus
Ornithonyssus bacoti ’-.

’
Allodermanyssus aegyptius Myospalax scansus
Lipohyssoidea muris Hirstionyssus confuscianus

Mus rattus
Echinolaelaps echidninua
Ornithonyssus bacoti
Mystromys albicaudatus
Haemolaelaps mystromys
H. davisi
.
Hirstionyssus carnifex’
Allodermanyssus aegyptius Napeozapus insignis
Lippnyssoides muris Haemoiaelaps glasgowi
Haemogamasus alaskensis
Mus silaceus
Echinolaelaps muricola Nectomys squamipes
Gigantolaelaps goyanensis
Mus sylvaticus G_. oudemansi
Haemolaelaps fahrenholzi Ornithonys sus braziliensis
Laelaps muris ..
Haemogamasus horridus Neofiber alleni
Haemolaelaps glasgowi
Mus triton
Laelaps zumpti Neotoma sp.
Haemolaelaps glasgowi
Mus trivirgatus Haemogamasus ambulans
Haeniogamasus liberiensis
Neotoma albigula
Mycrpmys minutus 1 Hirstionyssus breviseta
Haemolaelaps glasgowi H. neotomae
Laelaps micromydis Ischyropoda armatus
Hirstionyssus carnifex Brevisterna utahensis
H_. isabellinus B_. morlani
H. rnusculi
Haemogamasus nidi Neotoma cinerea
Haemolaelaps glasgowi
Myodes torquatus Haemogamasus ambulans
Laelaps semitectus
.Neotoma floridana
Myopus schisticolor Haemolaelaps glasgowi
Haemogamasus nidi
Neotoma fuscipes
Myospalax dybowskii Hirstionyssus neotomae
Haemolaelaps glasgowi Haemogamasus ambulans
Hirstionyssus confucianus Brevisterna utahensis
238

Neotoma lepida Onychomys leucogaster

Hirstionyssus hilli Haemolaelaps glasgowi
Brevisterna utahensiB’ Eubrachylaelaps crowei
Ornithonys.sus bacoti
Brevisterria utahensis
Neotoma mexicana Ischyropoda armatus .’ .-
Eubrachylaelaps circularis
Oryzomys eliurus
Laelaps hirsti
Neotoma micropus .

.
Haemolaelaps glasgowi Oryzomys palustris

.
Hirstionyssus neotomae Haemolaelaps glasgowi
Brevisterna morlani
H_.megaventralis
Gigantolaelaps cricetidarum
Laelaps oryzomydis
Neotoma pennsylvariicus
Haemolaelaps glasgowi
Otomys sp.
Hirsti onyssus otomys
Neotomodon sp.
Eubrachylaelaps circularis Otomys irroratus
Haemolaelaps labuschagnei
Neotomodon alstoni Laelaps parvulus
Eubrachylaelaps martini L. transvaalensis
Nesokia bengalensis Otomys tropicalis
Laelaps nuttalli Haemolaelaps labuschagnei
Echinolaelaps echidmnus H. sudanicus

Nesokia indica Otospermophilus. grammurus

Haemolaelaps glasgowi
’
Haemolaelaps’ glasgowi
Eulaelaps stabularis
Oxymycterus judex
Notomys mitchelli Ornithonyssus braziliensis
Laelaps hapaloti
Pedetes cater
"Old field mouse" Radfordilaelaps meridxonalis
Ornithonyssus bacoti
-
Hirstionyssus santos-diasi
Eulaelaps stabularis

Ondatra zibethica ’Perognathus sp.

Laelaps multispinosus Ischyropoda armatus
Ornithonyssus ondat-rae I. spiniger
Hirstionyssus isabellinus
Perognathus californicus
Haemolaelaps glasgowi
Onychomys sp. Eubrachylaelaps hollisteri
Haemogamasus ambulans Ischyropoda armatus
 239

Perognathus hispidus .. ; Peromyscus hylocetes

Haemolaelaps glasgowi Eubrachylaelaps circularxs
Ornithonyssus ba-coti
Hirstionyssus hilli .’ .
Peronaygcus leucopus
Haemolaelaps glasgowi
Perognathus inornatus Haemogamasus liponyssoides
Ischyropoda armatus Brevisterna morlani

’
Perognathus parvus Peromyscus maniculatus
Hirstionyssus hilli Haemolaelaps glasgowi
Eubrachylaelaps circularis
Perognathus penicillatus E_. debilis
Ischyrop’oda spiniger E_. hollisteri
Hirstionyssus carnifex
Perognathus spinatus H. hilli
IschyroRoda spiniger Brevisterna morlani
B. utahensis
Perognathus xanthonotus Haemogamasus alas.kensis
Ischyropoda armatus : Ischyropoda armatus

Peromyscus sp. Peromyscus mexicanus

Haemolaelaps glasgowi Eubrachylaelaps jamesoni
Eubrachy.laelaps spinosus
Laelaps pachypus Peromyscus nasutus
Hirstionyssus obsoletus Haemolaelaps megaventralis

Peromyscus boylei Peromyscus nuttalli

.
Haemolaelaps glasgowi Haemolaelaps glasgowi
Eubrachylaelaps circularis
Hirstionyssus affiais Peromyscus oaxacensis
Eubrachylaelaps circularis
Peromyscus californicus
Haemolaelaps glasgowi Peromyscus polionotus
.
.
Eubrachyiaeiaps circularis Haemolaelaps glasgowi
E. hollisteri
Peromyscus truei
Peromyscus crinitus Haemolaelaps glasgowi
Haemolaelaps glasgowi. Eubrachylaelaps circularis
Eubrachylaelaps debilis’ Brevisterna morlani
Brevisterna utahensis
Petaurista leucogenys
Peromyscus eremicus Haemolaelaps macroventralis
Ornithonyssus banksi
Brevisterna utahensis Phenacomys albipes
Haemogamasus ambulans
Peromyscus gossypinus
Haemolaelaps glasgowi Phenacomys longicaudus. :.
Haemogamasus liponyssoides Haemolaelaps glasgowi
240

Phodopus bedfordiae Psammomys obesus

Haemolael-aps glasgowi Haemolaelaps aegyp’tius
:
.. .Bala.elaps cricetuli
Haemogamasus mandschuricus Pseudoi’nys.sp.’’
Laelaps hapaloti
Phodopus songarus
Eulaelaps cricetuli Pseudomys apodeinoides
Laelaps finlaysoni
Phyllotis arenariua
Haemola.elaps glasgowi Pseudomyg ’minnie
Laelaps finlaysoni
"Pine mouse"
Haemogamasus alaskensis Punomys leminus
H. barberi Haemolaelaps glasgowi
-
H. liponyssoides
"Punare"
Pityinys pinetoruni.":’ Laelaps lativentralis
Haemolaelaps glasgowi Lepronysaoides pereirai
Eulaelaps stabulai-ts ’:
Haemogamasus alagkensis "Prairie dog"
H. barberi Hirstionyssus cynomys
H. liponyssoides
"Prairie vole"
"Porcupine" Eulaelaps stabularis
Hirstionyssus creightoni
"Rat"
Praeomys jacksoni Haemolaelaps hirsti
Laelaps lambo.rni H. radfordi

.
Echinolaelaps muricola H. reticulatus
Gigantolaelaps goyanensis
Praeomys tullbergi Laelaps aethiopicus
E chinolaelaps giganteus L. aragonfensis
L. exceptionalis
Proechimys sp. ’’’’.’v L. lamborni
Neoichoronyss^s dentipes L. wetmorei
:
E chinolaelaps grandis
Proechimys caljdius E. muricola
Tur uniscutatus
’
Ornithonyssus bacoti
.
.
Ophionyssus natricis
Prometheomys schaposchnikovi Hirstionyssus arcuatus
Haemolaelaps razumovae
Hirstionyssus bregetovae
,H. prometheonaicus "Ratode taquaral"
Haemogamasus zachvatkini Laelaps navasi

Protoxerus stangeri "Rato paco"

Haemolaelaps sangsteri Gigantolaelaps goyanensis
Rattus sp. Rattus norvegicus (contd.)
Echinolaelaps echidninus- Allodermanyssus sanguineus
A. aegyptius
Rattus alexandrinus -’:’. Myonyssus decumani
Allodermanyssus sanguirieus Eulaelaps stabularis
Haemogamasus ambulans
Rattus assunilis H. dauricus
Mysolaelaps rothschildi H. hirsutus
Laelaps assimilis H. japonicus
H. kusumotoi
Battus browni H_. liponysspides
Ophiomegistus buloloensis H,. nidi
H. oudemansi
Rattus crassus H. serdjukovae
Echinolaelaps echidninus
Rattus rattus
Rattus conatus Haemolaelaps aegyptius
Laelaps nuttalli H_. glasgowi
H. hystrici
Rattus coucha H. labuschagnei
Laelaps nuttalli H_.megaventralis
H_.morlani
Rattus culmorum H. zulu
Laelaps nuttalli Gigantolaelaps cricetidarum
Echinolaelaps echidninus Laelaps lamborni
L_.nuttalli
Rattus frugivorus L_.thompsoni
Haemolaelaps bibbyi Echinolaelaps ecliMnlnus
E_. bakeri
Rattus hawaiiensis E. muricola
Laelaps nuttalli E_. pallidus
Ornithonyssus bacoti
Rattus lutreolus Q--itieprai
Mesolaelaps australiensis Hirstionyssus latiscutatus
Allodermanys sus sanguineus
Rattus norvegicus (= Epimys nor- A. aegyptius
vegicus) Liponyssoides muris
Haemolaelaps glasgowi Haemogamasus arvicolarum
H. megaventralis
Gigantolaelaps cricetidarum Rattus tulbergi
Laelaps arvalis E chinolaelaps giganteus
L. nuttalli
L. sicula Rattus turkestanicus
Echinolaelaps echidninus Haemolaelaps glasgowi
Ornithonyssus bacoti H.. megaventralis
Hirstionyssus arcuatus Laelaps turkestanicus
H. isabellinus Ornithonyssus bacoti
H. rnusculi Allodermanyssus sanguxneus
242

Rattus turkestanicus.(contd.) "Rodents" (contd.)

Eulaelaps stabularia .
. Echinolaelaps yaound^nsis
Hirstionyssus carnifeA
Rattus whiteheadi .:; Allodermanyssus san^uineus .

Laelaps longulus Myonyssus^decumani I

Tricholaelapa comatus Eulaelaps stabularis !
Echinolaelaps sculpturatus Haemogamasus ambuljans
"Red-backed mouse"
H. hirsutus .
Haemogamasus liponyssoides "Rongeurs sylvatique" |
Echinolaelaps echidnipus
"Red squirrel"
Haemogamasus ambulans "Roof rat"
Echinolaelaps echidnijnus
"Red-tail sand rat" Ornithonyssus bacoti I
Haemolaelaps angustiscutis Eulaelaps stabularis |

Reithrodon cuniculoides Saccostomus campestri^

Haemolaelaps reithrodontis Haemolaelaps rhodesiensis
H. villosissimus
Reithrodontomys humilis Laelaps vansomereni
Haemolaelaps glasgowi Echinolaelaps muric^la
.
Reithrodontomys megalotis "Sand rat"
Haemolaelaps glasgowi Haemolaelaps angustiscutis

Rhabdomys pumilio Scirtopoda telum

Haemolaelaps. eos Eulaelaps kolpakova^
H. murinus
H. rhabdomys
Laelaps lamborni Sciurus sp.
Echinolaelaps giganteus Haemolaelaps glasgowi
Ornithonyssus roseinnesi Haemogamasus ambulans

Rhombomys opiinus Sciurus estuans ,

Haemolaelaps angustiscutis Haemolaelaps reticujiatus
H. longipes H. sciureus
Hirstionyssus meridianus
Sciurus carolinensis
"Rice rat" Haemolaelaps glasgowi
Ornithonyssus bacoti H. megaventralis
Haemogamasus ambulans
"Rodents"
Laelaps vansomereni Sciurus douglasi
Echinolaelaps bakeri
~
Haemolaelaps glasgowi
E. giganteus
E. grandis’ Sciurus griseus
E". muricola Haemogamasus ambulans
Sciurus hudsonicus ... . ; "Squirrels" (contd.)
Haemolaelaps glasgowi Hirstionyssus sciurinus
Hirstionyssus oceidentalis Eulaelaps stabularis
Brevisterna montanus
"Swiss white mice"
Sciurus niger Ornithonyssus bacoti
Haemolaelaps glasgowl
H. megaventralis Synaptomys sp.
Haemogamasus ambulans Haemolaelaps glasgowi

Sciurus vulgaris Tachyoryctes sp,. .

,
.
Haemolaelaps megaventralis Haemolaelaps sudanicus
Hirstionyssus sciurinus
Haemogamasus oudemansi Tachyoryctes ruddi
Haemolaelaps tachyoryctes
"Sewer rat" ’:
Ornithonyssus bacoti Tamias striatus
Haemolaelaps glasgowi
"Short-tailed mouse"
Haemogamasus ambulans Tamiasciurus douglasi
Haemolaelaps glasgowi
Sicista caudata Hirstionys sus affinig
Haemogamasus serdjukovae Haemogamasus ambulans
Pneumonyssus bakeri
Sigmodon hispidus
Haemolaelaps glasgowl Tamiasciurus fremonti
H. megaventralis Haemogamasus oudemansi
Gigantolaelaps cricetidarum
Steptolaelaps heteromys Tamiasciurus hudsonicus "

Ornithonyssus bacoti Haemogamasus alaskensis

Neoichoronyssus dentipes H. ambulans
N. wernecki
Haemogamasus liponyssoides. Tatera brantsii
Haemolaelaps taterae
Spalax leucodon H. oliffi
Hirstionyssus georgicus
Tatera schinzi
Spalax microphthalmus Echinolaelaps giganteus
Hirstionyssus macedonicus E. muricola
Spalax i-nonticola Thallomys namaquensis
Hirstionyssus macedonicus Laelaps simillimus

Spalax typhus Thamnornys rutilans

Haemolaelaps scapularis Laelaps thamnomys

"Squirrels" . .
Thomomys sp.
Hirstionyssus pauli Haemogamasus ambulans
244

Thomomys sp. (corrtd.) "Water vole"

Ischyropoda armatus ’’ Laelaps muris

Thomomys bottae "White-footed mouse"

Haemolaelaps geomys Haemogamasus liponyssoides
S-’ glasg0’^!
Eubrachylaelaps holliateri "White rats"
Ischyropoda armatus Echinolaelaps echidninus
Ornithonyssus bacoti
Thomomys bulbivorus
Haemolaelaps geomys "Wild rats"
fL glasgowi Gigantolaelaps butantanensis
G. peravianus
Thomomys fuscus >, G. vitzthumi

-
Haemogamasus ambulans Laelaps differens
H. occidentalis L. mazzai
i:
,

i^ pauliatanensis
Thomomys monticola :;,i Ornithonyssus bacptj
.
Haemogamasus ambulans 0^. lutzi
0. vitzthumi

’
Thomomys talpoides Eulaelaps vitzthumi
Haemogamasus ambulans.
"Wild rodents"
"Tropical rat" Gigantolaelaps gilmorei
Ornithonyssus bacoti
"Wood mouse"
Tscherskia triton Laelaps hilaris
Eulaelaps stabularis
Haemogamasus kusumotoi "Wood rat"
Laelaps nuttalli
Uromys lorentzi .. .
Ornithonysaus bacoti
Mysolaelaps rothschildi
"Yellow-throated mouse"
Uromys stalkeri .,;, Laelaps hilaris
Mysolaelaps rotlischildi
Zapus hudsonicus
Vandeleuria nilagirjca Haemolaelaps glasgowi
Laelaps. bhutanensis
Zygo.dontomys las’turus
"Water rat" Eubrachylaelaps rotundus
Laelaps agilis Ornithonyssus monteiroi
L. novikovae

Carnivora

Canis_familiaris ;. . Crossarchus fasciatus

Pneumonys s oides canimuni, Tympanospinctus paradoxus
 Domestic cat" Mustela sibinca
Ornithonyssus bacoti Myonyssus dubinini,

Fells catus ^’-; ’..: . Paradoxurus’ hermaphroditus

Haemolaelaps megaventralis Echinolaelaps echidninus

"Florida skunk" "Pine marten"

’
Ornithonyssus bacoti . Haemogamasus amfaulans
Eulaelaps stabularis
Procyon lotor ..

Haemolaelaps glasgowi
Galictis vittata (= prison vittatus)
Echinolaelaps berlesei Putorms sp. ;. ’
.

.
Ornithonys sus braziliensis Hirstionyssus isabellinus

Putorius erminea
Martes zibellina Hirstionyssus a’rcuatus
Haemogamasus nidi
H_. ambulans . Putorius putorius
Haemolaelaps fahrenhoizi
Mephitis elongata Hirstionyssus arcuatus
Haemolaelaps glasgowi
Hirstionyssus staffordi
"Raccoon"
Mephitis mesomelas Haemogamasus ambulans
’
Hirstionyssus staffordi Ornithonyssus bacoti
:
Mephitis nigra Spilogale interrupta
’
Haemolaelaps glasgowi Hirstionyssus staffordi

Spilogale leucoparia
Mustela sp. Hirstionyssus staffordi
Haemolaelaps glasgowi
Spilogale putorius
Mustela erminea Hirstionyssus Stafford!
Hirstionyssus isabellinus
Eulaelaps stabularis "Spotted skunk"
Ornithonyssus bacoti
Mustela nivalis Eulaelaps stabularis
Laelaps hilaris
L. pachypus
Hirstionyssus isabellinus Urocyon cinereoargenteus
Haemogamasus ambulans Haemolaelaps glasgowi
H. nidi .

"Weasel"
Mustela saturata Laelaps hilaris
Haemogamasus pccidentalis Haemogamasus hirsutus
246

Pinnipedia

Callorhinus ursinus Odobenus rosmarus

Orthohalarachne attenuate Orthohalarachne rosmari
0. diminuata

Otaria byronia
Eumetopias j’ubata
’
Orthohalarachne att’ehuata
Orthohalarachne magellanica

0. zaiophi
Phoca richardii
’
Halarachne miroungae
Halichoerus grypus
Halarachne halichoeri
"Tasmanian sea bear"
Orthohalarachne reflexa
Mirounga angustirostris
Halarachne miroungae
Zaiophus californianus
Orthohalarachne zaiophi
Monachus tropicalis 0. diminuata
Halarachne americana

’.’’ Hyracoidea

’
;
Dendrohyrax arborens ’’ Procavia capensis
:
Pneumonysgua schoutedeni Pneumonyssus procavians

.’ Perissodactyla

Equus caballos
Ornithonyssus bursa
Dermanyssus gallinae

Artiodactyla

Bos taurus Phacochoerus aethiopi’cus

Raillietia auris Pneumonyssdides phacochoeri

Kobus defassa
Raillietia hopkinsi
 AVES

Tinamiformes

Nothura maculosa Tinamus solitarius

Tinaminyssus navajasi Tinaminyssus trapRi

Rhynchotus rufescens
Neonyssus serraoi

Colymbiformes

Colymbus caspicus Poliocephalus ruficollis

Rhinonyssus alberti Rhinonyssus pplioc.ephali

Procellariiformes

Pachyptila desolata Puffinus tenuirostris

Haemolaelaps pachyptilae Haemolaelaps marsupialis

Ciconiiformes

Ardea cinerea Egretta intermedia

Neonyssus belopolskii Neonyasus bubuici

Ardea melanocephala Hydranassa tricolor

Neonyssus ardeae Neonyssus belopolskii

Bubulcus ibis Ixobrychus minutus

Neonyssus bubuici Neonyssus ixobrychi

Egretta alba Nycticorax nycticorax

Haemolaelaps glasgowi Meonyssus ardeae

Anseriformes

Alopochen aegyptiacus Anas acuta

Rhinonyssus rhinolethruna Rhinonyssus rhinolethrum
248

Anas carolinensis .- /"Domestic goose"

Rhinonyssus rhinolethrum Rhinonyssus rhinolethrum

Anas erythrorhyncha "Ducks"

Rhinonyssus rhinolethrum .: .. Ornithonyssus burs a

.
Dermanyssus gallinae
Anas platyrhynchos
Rhinonyssus-rhinolethrum :;;;;’ Mareca americana .: . .

Rhinonyssus rhmolethrfUm
Anas sparsa
Rhinonyssus rhinolethrum Mergus merganser. ,,.. ...,;.
Rhinonyssus rhiriolethrum
Anas strepera
Rhinonyssus rhinolethrum Plectropterus gambensis
;.,. Rhinonyssus rhinolethrum
Aythya affinis
Rhinonyssus rhinolethrum
Sarkidiornis melanotos , : :

Rhinonyssus rhino].e1;hru’nii’
Branta canadensis
Rhinonyssus rhinolethrum Spatula clypeata
. Rhinonyssus rhinolethrum

Cygnus columbianus Soinmateria mollissima ’.

Rhinonyssus rhinolethr.UjBi;; Rhinonyssus rhinotethrum

Falconiformes
;’?i)r:c.i;;,’.-.;:’?

Buteo rufofuscus Milvago chimachima

Neonyssus buteoni-s, ,.; Rhinonyssoides souzai- : i:

Coragyps atratus Milvus tenebrosus

Rhinonyssoides .donatoi : Neonyssus-icolumbaei ,-.

Gyps coprotheres "Red-shouldered hawk"

Haemolaelaps pate.rs^ini- Haemogamasus arnbulans

Lophaetus occipitalis
Neonyssus buteonis .. Rupornis magnirostris
Rhinonyssoides souzai

Galliformes

"Bobwhite" . ; . "Chickens"
Haemogamasus aimbulans Ornithonys sus bursa
"Chickens", (contd.)
Orhithonyssus sylviarum "Turkeys"
Dermanyssus gallinae Ornithonys sus bursa
Haemogamasus ambulans Dermanyssus gallinae

"Poultry"
Ornithonyssus bacoti
Dermanyssus gallinae

.
Gruiformes

Fulica americana Limnocorax flavirostra

Rallinyssus caudistigmus Hallinyssus congolensis
R. limnocoracis
Fulica atra
Rallinyssus caudistigmus Otis tarda
Vitznyssus vitzthumi

Neotis cafra Rallus elegans

Vitznyssus neotis Rallinyssus caudistigmus

Charadriiformes

Actitis hypoleucos Cepphus columba

Sternoston’ia boydi Sternostoma. caledonicum

Aethia pusilla Cepphus grylle

Sternostoroa caledonicum Sternostoma caledonicum

Afribyx senegallus. Charadnus alexandnnus

Rhinonyssus afribyx Rhinonyssus coniventris

Aica torda Charadrius hiaticula

Rhinonyssus waterstoni Sternostoma minutus
Rhinonyssus coniventris
Arenana interpres
Rhinonyssus coniventris Charadrius tricollaris
Sternostoma boydi Rhinonyssus himantopus

Arquatella ptilacenemis ; Chlidonias leucoptera

Rhinonys sus coniventris Larinyssus orbicularis

Catoptrophorus semipalamatus Crocethia alba

Rhinonyssus coniventris Rhinonyssus coniventris
250

Crocethia alba (contd.) Rhyacophilus glareola

Sternostoma boydi Rhinonyssus coniventris
Sternostoma .boydi ’
Erolia alpina
’

Rhinonyssus coniventris Stephanibyx lugubris ,

Rhinonyssus afribyx
Erolia maritima
Rhinonyssus neglectus Sterna hirundo
Larinyssus orbicularis
Gelochelidon nilotica
Larinyssus orbicularis Sterna maxima
Larinyssus orbicularis
Hemiparra crassirostris
Rhinonyssus himantopus
Totanus flavipes
Himantopus mexicanus Rhinonyssus coniventris
Rhinonyssus himantopus
Totanus nebularis
Larus argeniatus Sternostoma boydi
Larinyssus orbicularis
Tringa ocrophus
Larus atricilla Sternostoma boydi
Larinyssus orbicularis
Sternostoma boydi Tringa striata
Rhinonyssus coniventris
Larus delawarensis
Larinyssus orbicularis Totanus nebularius
"
Sternostoma boydi Sternostoma boydi

Larus occidentalis Uria aalge

Larinyssus orbicularis Sternostoma caledonicum
Philomachus pugnax Uria grylle
Sternostoma bpydt Rhinonyssus caledonicus

ColumbUormes

Columba livia Stigmatopelia senegalensis.

Haemolaelaps glasgo’wi Neonyssus-melloi ; : .

Ornithonys sus bursa

0. sylviarum
Dermanyssus gallinae Streptopelia semiforquata
Neonyss.us columbae ; Neonyssus’ columbae.
.
N. melloi

Columbigallina passerina Treronicalva-

Neonyssus zenaidurae Neonyssus treronis
 Turtur afer
Zenaidura macroura
Neonyssus melloi
Neonyssus zenaidurae

Cuculiformes’

Chrysococcyx caprius Cuculus canorus

Sternostoma cuculorum Sternostoma cuculorum
Clamator levaillanti
Cuculus solitarius
Sternostoma cuculorum Sternostoma cuculorum

Coliiformes

Colius striatus
Sternostoma striatus

Strigiformes

Asio capensis
Otus asio
Rhinoecius africanus
Dermanyssus oti
Asio otus
Steotyto cunicularia
Rhinoecius oti
Haemolaelaps glasgowi
Rhinoecius bisetosus
Bubo yirginianus
Trix varia
Rhinoecius grandis
Rhinoecius cooremani
"Eagle owl"
Dermanyssus Tyto alba
gallinae
Rhinoecius tytonis

Caprimulgiformes

Caprimulgus europaeus
Vitznyssus nitzschi
Cosmetornis vexillaris
Vitznyssus nitzschi
V. scotornis

Scotornis fossil
Caprimulgus tnstigma
Vitznyssus nitzschi
, Vitznyssus~icotornis
V_. scotornis
252

_
;
i.Mieropodiformes

Apus caffer Mid-opus affinis

Rhinonyssus apus Dermanyssus hirundinis

’
’.

.
.
Rhinonyssoides strandtmanni

Cypselus affinis. . , .’.". Sericotes holosericeus

Pellohyssus viator Rhinonys soides squam osus

’;.;-;Coraciitormes .

Halcyon leucocephala ’.: Merops apiaster (contd,)

Neonyssus schoutedeni Sternostoma cooremani

Mellttophagus lafresnayt Merops nubicoides

Sternostoma cooremani Sternostonaa cooremani’
Ptilonyssoides mellttophagi
Merops persicus
Melittophagus pusillus Ptilonyssoides triscutatus
Ptilonyssoides meXittophagi

Merops apiaster . : : Phoeniculus purpureus :

.
Ptilonyssoides trisc.utatus Haemolaelaps phoenicuU

Piciformes

Campethera abingoni . : "

Dendropicus cardinalis
.
Pellonyssus biscutatus Pellonyssus biscutatus

Centurus carolinus Pendropicus fuscescens

Haemolaelaps megaventralis Ornithbnyssus dendropicus
Pellonyssus biscutatus
Ceophloeus pileatus
Haemolaelaps megaventralis Dryobates pubescens
Haemolaelaps megaventralis
Chrysocolaptes validus
Haemolaelaps omnitectus Dryobates villosus
Haemolaelaps glasgowi
Colaptes cafer
Dermanyssus scutatus
Gecinus vaillanti
Dendrocopus major Pellonyssus viator
Dermanyssu-s quintus Dermanyssus gallinae
Melanerpes erythrocephalus .: Sphyrapicus varius
Haem olaelap.s meg.aventralis Haemolaelaps megaveritralis

Mesopicus ruwenzorii "Yellow-bellied’sapsucker"

Haemolaelaps’ mesoptCus Ornithonyssus bacoti

Passeriformes

Acanthis cannabifla .
’
Bessonornis archeri
SteatonyssuS, musculi PtilonyssoCTes dioptrornis

Acrocephalus arundinaceus "Blue jay"

Sternostoma tracheae olum Haemogamasus ambulans

Agelaius phoeniceus "Brewer’s blackbird"

Paraneonyssus icteridius Ornithonyssus sylviaruro

Agelaius tricolor Buphagus africanus

Sternostoma strandtmanni Sternostoma sturnicola
S. tracheacolum
Paraneonyssus icteridius "Cage birds"
Dermanyssus gallinae
Alseonax adustus
Ptilonyssus lanii "Caged chaffinches"
Dermanyssus gallinae
Arnandava subflava
Paraneonyssus subflava Calamocichia rufescens
Ptilonyssus calamocichlae
Amblyospiza ’albifrons
Paraneonyssus zumpti Calandrella cinerea
Ptilonyssus lanii
Ampelion cucullatus
Ptilonyssus olaioi Campephaga Bboeflicea
Ptilonyssus lanii

Anaplectes rubric’eps "Canaries"

Paraneonyssus ploceanus Ornithonyssus bursa
Permanyssus gallinae.
Andropadus latirostrii Sternostoma tracheacolum
Ptilonyssus andropadi S. meddai
Anthus arboreus "Cardinals"
Permanyssus hirundinis Sternostoma meddai

Apalis flavida Carduelis cannabina

Ptilonyssus calamocichlae Steatonyssus musculi
254

Carduelis carduelis Dicrurus adsimilts

.Ornithonyssus sylyiarum..;. Ptilonys sus dicruri
Sternostoma thienponti
’""-. -,^_^-
Cecropis -senegalensis
Ptilonyssus echinatus : ’

Dioptrornis fis’cHeri
Ptilonyssus dioptrornis
Cerchneis tinnunculus
Rhtnonyssoldes cerchneis- Donacobius atricapillus
Qrnithonyssus iheringi
Cercomela familiaris
Ptilonyssus rnotacillae Dryoscopus cubia
Paraneonyssus^dryscopj.
Chalcomitra senegalensis
Pttlonyssus cinnyris
Sternoatoma nectarinia Elaenia obscura-’..’
Rochanyssus werneri
Chlorocichia flavicQllis
Ptilonyssus chlorocichlae Elaenia flavogaster
Travanyssus paranensis
Cinclus ci.ncl.us
Sternostoma technaui Emberiza cirlus
Ptilonyssus nudus

Cinnyris afra Emberiza flaviventris

Ptilonyssus cinnyris Paraneonyssus embenzae

Cinnyris cupreus Emberiza rustica

Sternostoma nectarinia Haemolaelaps disimilis

Cinnyris regius "English sparrow"

Sternostoma nectarinia Ornithonyssus’-jSursa
Ptilonyssus cinnyris Dermariys sus. gallinae’ ’-

Estrilda paludicola’.-.
Cisticola erythrpps .’ Paraneonyssus aatridae
Ptilonyssus calamocichlae
Euphagus sp.
"Common sparrow" Haemolaelaps megaventralis
Ornithonyssus .bu-rsa ....
Euphagus cyanocephalus
Cossypha heugl.ini .. . Paraneonyssus icteridius
.
Ptilonyssus lanii
Euplectes orix
Craspedophora magnifica Paraneonyssus ploceanus
Rhmonyssoides .nova-guineae

Creatophora cinereus . Fringilla coelebs ’

Pellonyssus reedi . : Ptilonyssus nudus.

Geokichia piaggiae Irodoprocne bicolor
Ptilonyssus lariii Haemolaelaps megaventralis
Ptilonyssus echinatus-
Granatina granatiria
Paraneonyssus astridae "Kingbird"
Ornithonyssus sylviarum
Hippolais ictenna- 0. bursa
Ptilonyssus calamocichlae
Lagonosticta rhodopareira
Hirundo rufula Paraneonyssus astridae
Dermanyssus hirundinis
Lagonosticta rubricata
Hirundo rustica Sternostoma lagonostictae
Ptilonyssus echinatus Paraneonyssus astridae
Sternostoma tracheae olum
Lagonosticta senegala
Hirundo smithi Paraneonyssus astridae
Ptilonyssus echinatus
Sternostoma hirundiriis Lamprotornis purpuropterus
Sternostoma sturnicola
Hirundo urbica
Dermanyssus hirundinis Lanius collaris
Ptilonyssus lanii
"Horned lark" Sternostoma laniorum
Dermanyssus brevis
Lanius collurio
"House sparrow" Ptilonyssus lanii
Dermanyssus gallinae Sternostoma laniorum

Hylocichlae ustulata Lanius excubitorius

Sternostoma hutsoni Ptilonyssus lanii
S. spatulatum Sternostoma laniorum

Hyphantornis velatus Lanius ludovicianus

Pellonyssus reedi Haemolaelaps glasgowi

Hyphanturgus ocularius Lanius nigriceps

Paraneonyssus ploceanus Ptilonyssus orientalis

Hypochera funerea Lonchura bicolor

Ptilonyssus nudus Paraneonyssus astridae

Macronyx croceus
Icterus bullocki Sternostoma tracheacolum
Sternostoma tracheacolum Paraneonyssus capensis.

Indicator indicator Macron.yx capensis .

Rhinonyssoides indicatoris Paraneonyssus capensis

256

Malurue melanocephala "Nuthatch"

Pellonyssus malurus ,". Ornithonyssus bankgi. :

.
"Martin" Oberholseria chlorura
Dermanyssus hirundinis :;. Eubrachylaelaps deb,ffl,is
"Meadow lark" Oenanthe oenanthe .

Ornithonyssus bursa Ptilonyssus motacillae

0. sylviarum ,

Onychognathus moria
Melosplza melodia Pellonyssus reedi
Haemolaelaps glasgowi
Oriolus larvatus ;. :
Molothrus ater Ptilonyssus orioli
Sternostoma strandtmanrii
Paraneonyssus icteridius Oriolus oriolus
Ptilonyssus orioli
Motacilla aguimp
Ptilonyssus motacillae Otocoris alpestris
Paraneonyssus capitatus
Motacilla capensis
Ptilonyssus motacillae Parus atricapillus :.;’:’
Ptilonyssus nudus
Motacilla flava. .;
^-
Ptilonyssus motaciilae Parus major .
.

.
Ptilonyssus nudus
Muscicapa aquatica
.
Ptilonyssus lanti. Passer domesticus . .

Haemolaelaps megaventralis
Muscicapa striata. ; Ornithonyssus’banksi
Ptilonyssus nudu s 0. bursa
P. lanii
~
’ 1

Pellonyssus passeiri
P. viator
Sternostoma cryptorhynchum
"Native song birds" Paraneonyssus hirsti-
Dermanyssus gallinae Ptilonyssus nudus

Nectarinia kilimensis Passer grisea . . 1;.

Ptilonyssus cinnyris Ptilonyssus nudus

Nectarinia purpureiventris Passer hispaniolensj.s

Sternostoma nectarinia Pellonyssus viator
Nucifraga caryocactes Passer naelanura
Neonyssus nucifragae Pellonyssus reedi
Nucifraga columbiana Passer montanus ’ -.’:’ .
Eubrachylaelaps "debilis Dermanyssus hirundinis
Passer montanus (contd.) Prinia subflava
Pellonyssus viator Ptilonyssus aureliani

Passer ugandae Procnias alba

Paraneonyssus hirsti Ptilonyssus stresenaani

Petrochelidon fulva Prggne subis

Haemolaelaps megaventralis Dermanyssus prognephilus

Petrochelidon pyrrhonota Progne chalybea

Haemolaelaps megaventralis Cas angrensis
Ptilonyssus echinatus
Psalidoprocne albiceps
Petrochelidon spilodera Sternostoma hirundinis
Ptilonyssus echinatus Ptilonyssus psalidoprocnei

Philydor lichtensteini Psalidoprocne holomelaena

Paraneonyssus travassosfilhoi Ptilonyssus psalidoprocnei

Phyllastrephus fischeri "Purple grackle"

Ptilonyssus andropadi Ornithonygsus sylviaruin

Phyllastrephus terrestris Pycnonotus barbatus

Ptilonyssus phyllastrephi Ptilonyssus pycnonotj.

Phylloscartes ventralis Pytilia melba

Paraneonyssus enriettii Paraneonyssus astridae

Piranga ludoviciana Quelea quelea

Paraneonyssus icteridius Paraneonyssus ploceanus

Platysteira cyanea Quiscalus quiscula

Ptilonyssus lanii Paraneonyssus icteridius

Pipilo erythrophthalmus Ramphocelus bresilius

Haemolaelaps glasgowi Ptilonyssus japuibensis

Plocepasser mahali Ramphocelus carbo

Pellonyssus similis Ptilonyssus japuibensis

Ploceus velatus "Red-eyed vireo"

Pellonyssus reedi Ornithonyssus bursa
0. sylviarum
Poliospiza striolata
Ptilonyssus lanii Riparia riparia
Haemolaelaps glasgowi
Prinia leucopogon Ornithonyssus sylviarum
Ptilonyssus calarnocichlae Eulaelaps novus
258

Riparia riparia (contd.) "Starlings" (contd.)

Haemogamasus ambulans Dermanyssus- gallinae
H. avisugus :
Ptilonyssus echinatus Stelgidopteryx ruficollis
P. nudus Haernolaelaps glasgowi

"Robins" Sturnella magna

Ornithonyssus sylviarum Paraneonyssus icteridius

"Rook" Sturnus vulgaris

Ornithonyssus sylviarum ’. Haemolaelaps aiggaventralis
Ptildnys sus-nudfls
Saxicola rubetra
Ptilonyssus mbtacillae "Swallows"
Ornithonys sus sylviarum
Saxicola torquata
Ptilonyssus motacillae "Swallow nest"
Dermanyssus chelidbnis
Serinus canaria
Ptilonyssus nudus
Tangara seledon- ’
Serinus canicollis ,-.’ . / .. . Ptilonyssus sairae
.
Paraneonyssus serini
Tchagra senegala’’ ;’ ’:

Serinus citrinelloides . . Paraneony’s’sus dryoscopi

Paraneonyssus serini
Terpsiphone virtdis ’" :
Serinus mozambicus : Ptilonyssus terpsiphonei
Ptilonyssus calamocichlae Ruandanyssus terpsiphonei
Paraneonyssus serini. : .’
Textor atrogularis
Sittasomus griseicapillus Paraneonyssus ploceanus
Flavionyssus rabelloi

"Sparrows" Thamnolaea cinnamomeiventris

Ornithonyssus sylviarum Ptilonyssus desfontainei

Spinus spmus "Towhee"

Ptilonyssus nudus Dermanyssus gallinae

Sphecotheres maxillaris Trochocercus cyanomelas

Rhinonyssoides trouessarti Ptilonyssus terpsiphonei

Spreo bicolor Turdoides jardinei

Haernolaelaps spreo Sternostoma dureni

"Starlings" . Turdoides melanops

.
.
Ornithonyssus bursa .
Sternostoma dureni
Turdus abyssinicus "Wood thrush"
Sternostoma turdi Ornithonys sus burs a

Turdus olivaceus "Wren"

Sternostoma turdi Dermanyssus gallinae
S. dureni
Xanthocephalus xanthocephalus
Paraneonyssus icteridius
Uraeginthus bengalus
Paraneonys-sus astridae "Yellow-headed blackbird"
Ornithonyssus sylviaruna
Vidua macroura
Ptilonyssus nudus Z osterops senegalensis
P. viduae Ptilonyssus ruandae

REPTILIA

Chelonia

Ophionyssus natricis’

Squamata

Sauria

Acanthodactylus scutellatus Lacerta vivipara

Ophionyssus natricis Sauronyssus saurarum

Agama adramitana
Ophionyssus natricis "Lizard"
Ophionyssus natricis
Lacerta agllis
Sauronyssus saurarum
Mabuya quinquestriata ;
Lacerta muralis Sauronyssus gordonensis
Sauryonyssus saurarum

Lacerta yiridis Sceloporus gracious

Sauronyssus lacertinus Ophionyssus natricis
260

Serpentes

"Captive reptiles" Erythrolamprus aesc.ulapi

.
.
Ophionyssus. natricis . Pneumophionyssus aristoterisi

Coluber flagellum ; Heterodon contortrix

Ophiopneumicola colubri Entonyssus heterodontus

Coluber florulentus Lampropeltis calligaster

Ophionyssus natricis Hemilaelaps triangulus

Coluber karelini Lampropeltis getulus

Ophionyssus natricis Entonyssus fragilis
Coluber radiatus i Leiolopisma fusca
Hemilaelaps imphalensis Sauronyssus arnhemlandensis
Crotalus cinereous ’. Naja haje
Entonyssus ewingi Ophionyssus natricis

Dendroaspis angusticeps ..Naja tripudians

Hamertonia bedfordi Hamertonia schoutedeni

Prymarchon corias Natrix sipedon

Ophionyssus natricis Ophiopneumicola .natricis:,..,.’:.
Hemilaelaps americanus
Natrix tigrina
. Hemilaelaps tanneri
Echis carinatus
Ophionyssus yariabilis Ophis merremi
Ixodorhynchus butantanensis
Elaphe dione
Hemilaelaps farrierj.
Pituophis melanoleucus
Elaphe guttata ; . . Entonyssus halli
Ophiopneumicola americanus
Pituophis sayi :
Elaphe obsoleta Entonyssus vitzthumi
Hemilaelaps distinctus
Ophiopneumicola elaphes Psammophis sibilans
Ophionyssus natricis
Elaphe quadrivittata .
.
.
Entonyssus glasmacheri Psammophis schokari
Ophionyssus natricis
Epicrates cenchria ;
Ophionyssus natricis Python reticulatus
Heterozercon oudenianst Ophionyssus natricis
,
 261

"Rattlesnake" Telescopus dhara

JShtonyssus rileyi Ophionyssus natricis

.
.
"Snakes": .

Opbionyssus natricis . Thamnophis sirtalis

Ixodorhynchus liponyssoides Ophiopneumicola hamertoni
Hemilaelaps piger :
Ophiomegistus luzonensis ,.
Vipera lebetina
Spalerosophis cliffordi . .. Ophionyssus natricis
Ophionyssus natricis

Host .Specificity

The parasitic Mesostigmata vary from very low specificity to com-

plete reliance on one species of host. But in no instance is specificity
so low that a mite will parasitize .both warm and cold blooded animals.
Generally even those mites .that indicate low specificity restrict them-
selves to one class, i.e. only reptiles, only mammals, or only birds.
Those species that seem to be more catholic may appear so only because
we do not thoroughly understand the species. Eulaelaps stabularis, for
example, has been found on a wide variety of mammals and birds but
it has not yet been proven that this species is a true parasite. Also,
Haemolaelaps glasgowi may be found on many birds and mammals, but
here we may be dealing with a species complex and more refined studies
may reveal that several species are involved.
; Numerous species will attack with equal gusto any host of the same
class. Ophionyssus natricis will do equally well on all kinds of snakes
as well as lizards. Ornithonyssus bacoti attacks any mammal, includ-
ing man, but will not readily attack birds. Ornithonyssus sylviarum,
0. bursa and Dermanyssus gallinae, on the other hand, will attack birds
of all kinds but do not often attack mammals. However, Sikes and Cham-
berlain (1954) have shown that if the mites are confined on the host they
will feed on mammals and produce viable eggs from such a feeding. Or-
nithonyssus bursa and 0. sylviarum would feed on mouse and rabbit but
not on man under such conditions. Dermanyssus gallinae would not feed
readily on either mouse or rabbit and not at all on man.
More commonly, a species of mite will restrict itself to a group
of related host species. For example, Haemolaelaps geomys will be
found only on the rodent family Geomyidae, but within this family ap-
parently all species are equally choice hosts; Laelaps nuttalli and Echino-
laelaps echidninus parasitize only rats and mice of the family Muridae;
many of the Spinturnicidae and several species of Ichoronyssus parasi-
tize only bats but do not restrict themselves to one species. This also
is the case in some species of internal mites. Rhinonyssus rhinoleth-
rum, for example, parasitizes only waterfowls of the order Anseriformes,
262

but any species will do.

.
It may also be noted that sometimes whole genera of mites are re-
stricted to certain types of hosts. For example, the genus Laelaps para-
sitizes only rodents; the genus Rhinoecius is fouad only in owls; ,Ptilo-
nyssus only in passeriform birds; Spinturnix, Steatonyssus,,. Ichoro
nyssus and Periglischrus only on bats, etc. ’,’". ""

’
’
True host specificity cannot be determined tor a species uriles.s it
has been accurately recorded from a number of localities. Because so
many species have been found only once, it is a bit hazardous to guess
how many are actually specific, but judging from better known species,
only a relatively few qualify. Laelaps multispinosus on the muskrat
Ondatra zibethicus appears to be an example of a truly specific host-
parasite relationship. This mite does not occur on any other species.
Recently, Cross (1954) has shown that some mites (including Ornith-
onyssus bacoti, 0. bursa, 0_. sylviarum, Dermanyssus gallinae, Echino-
laelaps echidninus and Haemolaelaps glasgowi) will feed through silk
bolting cloth on blood from an abnormal host. Apparently then. in such
instances, host specificity is more a matter of physiological, compatabil-
ity than a question of choice of diet. It has been previously noted that
conditions, rather than hosts, may determine specificity in certain forms
(see page 199).
Much more needs to be known about many more species before any-
thing very definite can be said about host specificity. La our present
state of knowledge, it appears as if the majority of species restrict them-
selves to certain host groups rather than to any one host species. How-
ever, Zachvatkin(1948) and Lange(l955) believe that the parasitic Meso-
stigmata, at least of the genus Laelaps, are very closely circumscribed,
both geographically and faunally. They have erected several new sub-
species on the basis of minute morphological differences, which they
claim are significantly associated with either geographical location or
host association or both.
 APPENDIX

Collecting Methods

There are really only two sources of parasitic mites: (1) the host,
and (2) the habitat of the host. Probably the best way to collect from
the nest is to place it in a paper sack or other mite-tight container, car-
ry it to the laboratory and place it in a Berlese funnel. Needless to say,
the maker of the nest must be positively identified. A glass jar should
be screwed into a lid that is soldered to the bottom of the funnel so the
mites can be collected alive. The jar can be periodically emptied into
awhite enamelpanand the mites aspirated into vials as they crawl away
from whatever debris may have fallen through with them. Not all mites
will crawl into the open, but the hidden ones can be quickly found by put-
ting the debris in a small dish and examining under a binocular micro-
scope. Collecting the mites alive is vastly superior to collecting them
in alcohol or water. For best results, nests should be placed in Ber-
lese funnels within twenty-four kaurs after collection.
To collect from the host ( and this is the only way to find internal
mites and the mites of free ranging reptiles ) it is first necessary to
catch the animal. It is probably preferable to live trap the host although
there is no real evidence that dead trapped animals lose many mites
within the first few hours after death. Whether taken dead or alive, the
animal should be carried to the laboratory in a tightly tied bag.
Iji the laboratory, the animal, if it is a bird or mammal, should be
anesthetized and then combed, brushed and vigorously shaken over an
enamel tray. We think this method gives better results than washing in
a detergent, at least as far as mesostigmatid mites are concerned. Spe-
cific areas of the animals may also be investigated under magnification,
such as in the ears of mammals and under the wings and around the eyes
of birds, but it is usually unrewarding to examine furred or feathered
animals in detail over the entire body. Mites on reptiles, however, are
best found by examining the body under a dissecting microscope. We
have never had much success in collecting mesostigmatid mites by plac-
ing the host in a Berlese funnel although this is sometimes the method
of .choice for chiggers.
Internal mites are found only by opening the animal and examining
the organs. As far as is now known, they inhabit only the body cavities,
primarily the respiratory system and the auditory apparatus.

263
264

Mounting Methods

The best method of mounting mites is probably that method in which

the user has most confidence. The most commonly used media are those
which do not require dehydration of the specimen. Purely temporary
mounts may be made by immersing the specimen in glycerin, phenol, or
lactic acid. These clear the specimens nicely and also have a suitable
refractive index. More permanent mounts may be made with any of sev-
eral chloral hydrate-gum acacia formulations (Berlese-type media) or
polyvinyl alcohol formulations. Formulae for these are given below.
Completely permanent mounts can be made by the standard histological
procedure of clearing in caustics, such as KOH, dehydrating with alco-
hol, and mounting in Damar, clarite, or Balsam. However, this has
the disadvantage of being time consuming and the even greater disadvan-
tage of shrinking or disfiguring delicate parts. Also, the mounting media
have almost the same refractive index as the mite, making staining neces-
sary.
Many acarologists use polyvinyl alcohol (PVA) exclusively. It is
certainly true that it gives excellent initial results but it has been our
experience that the specimen gradually becomes distorted in this medium.
Perhaps the proponents of PVA have found ways of obviating this. A
major difficulty with PVA formulations is that remounting is difficult
and discouraging. Not only must the mite be removed from the medium
(a sticky, time consuming task), but in order to restore the specimen to
its natural appearance, the medium must be removed from around and
within the mite so that reinfiltration with fresh medium can be made (a
nearly impossible task;).
In our opinion, the mesostigmatid mites are best mounted in Hoyer’s
medium or any other of the various modifications of Berlese’s chloral-
hydrate-gum acacia mixtures. Thus mounted, the specimen is never
permanently damaged. It may turn black, it may slip, it may partially
dry, but the specimen is not injured. All that is necessary is to soak
the mite in clear, warm water for a short time and remount. Mounted
in this medium, it is possible to find the most evasive structures because
the refractive index is excellent (especially for phase-contrast micros-
copy) and there has been no destruction of delicate characters by dehy-
dration.
If there are several specimens it is advantageous to mount them
alternately with dorsal and ventral sides up. If there is only one, there
is a slight advantage to mounting it ventral side up, as this is where
most of the critical characters are to be found. Never mount more than
one mite per slide. Always use a cover glass of #1 thickness or less,
which will allow use of the oil immersion objective, an essential for
finding the finer details.
Freshly caught specimens can be mounted directly into the medium.
If they are fully engorged with blood it is best to clear them in Nesbitt’s
solution or lactic acid. Also, specimens that have been kept for a long
 265

time in alcohol, formalin, or some other preservative, will not clear

well if mounted directly into the medium. We have found it desirable to
allow the specimens to soak in water for as long as a week before mount-
ing. Camin has routinely boiled all of his specimens in lactic acid be-
fore mounting. His method is to place the mites in a 1 mm vial, cover
with lactic acid, hold over a’ small flame until the lactic acid "begins to
bubble, wash the mites briefly in 90% alcohol and mount immediately.
Recently we (as well as Camin) have become strong advocates of clear-
ing in warm Nesbitt’s solution. The method is to put the uncleared speci-
mens in a small covered watch glass with the solution, and place in a
drying oven adjusted for a temperature of 50 - 55 C. The specimens
become clear in a few hours to a day. Further treatment may cause
destruction of the mites.
After the cover glass is put on, the slide may be heated over a lamp
or placed in an oven at around 50 C. This causes the legs to extend,
air bubbles to disappear, and also hastens the clearing process if the
specimen has not been previously cleared. Ordinarily it takes over 24
hours for a non-engorged specimen to clear in Hoyer’s alone. Cover
glasses may be sealed with any of several water insoluble substances
such as clarite, Damar, etc., only if the slide will not be used with im-
mersion oil. Xylolis often used to clean oil from the slide. In this case
the ringing substance must also be able to resist this organic solvent.
We have used cellulose cement and fingernail polish, but the most promts -
ing sealers appear tobe plastic lacquers that resist treatment with xylol.
A well mounted and properly cleared specimen is the first essential
for mite study. Beyond this it is necessary to have the best in micro-
scopes. An oil immersion objective is essential, and phase-contrast mi-
croscopy is also highly desirable.

Berlese’s Mounting Medium and Several Modifications

Berlese’s Mounting Medium: Ewing’s Modification:

20 cc water 35 cc water
15 g gum arable 20 g .gum arable
Up to 160 g chloral hydrate 30 g chloral hydrate
10 cc glucose syrup 12 cc glycerin
5 cc acetic acid 3 cc glucose syrup
,
Gater’s Modification:

10 ml distilled water 74 g chloral hydrate

8 g gum arable 5 ml glucose syrup

The glucose syrup is prepared with equal parts of glucose and water.
Honey may be substituted for the glucose syrup.
266

King. Bradley, and McNeel’s Doetschman’s Modification:

Modification:
’ 35 cc water
8 g gum arable 20 g gum arable
8 ml distilled water 20 g chloral hydrate

.
5 ’ml glycerin 20 cc glycerin
70’" g chloral hydrate 3 cc glucose syrup
3 ’ml glacial acetic acid 10 drops or more basic fucnsin.

Hoy er’s Modification:

50 g distilled water
30 g gum arable
200 g chloral hydrate
20 g glycerin

In all the foregoing formulae, the ingredients should be mixed in the

sequence given. It is especially’desirablethat all the gum arable be dis-
solved before the chloral hydrate is added. Warming in a water bath or
in an oven to hasten the process may be desirable.. Either crystalline or
powdered’ gum arable may be used. but we have found highly purified
flakes to be most efficacious. After mixing, the solution should be
strained through a lint-free filter to remove insoluble’ debris.

Polyvinyl Alcohol (PVA) Mounting Medium

’
1. Dissolve "Elvanol 71-24" (Du Pont polyvinyl alcohol) in tour volumes
of water by stirring at about 90 C.

2. Filter the solution until it is no longer murky.

3. Concentrate-the clear filtrate on a water bath Mhtxl’it has the vis-

cosity of a syrup. (A scum will form on the surface during the pro-
cess of evaporation but will redissolve when stirred into the solution.)

4. Add 22 parts of lactic acid to 56 parts of the PVA concentrate to

make the finished mounting medium. ’’"-

5. Use like any mountant. Material may be mounted directly from life,
any aqueous solution, or alcohol,

Methocellulose Formula

Workers at the University of California have developed a metho-

cellulose formula that appears to be promising for certain mites. Their
 267

formula is:

5 g metboceUulose
2 g carbowax 4, 000
1 ml diethylene glycol
,25 ml 95% ethyl alcohol
100 ml lactic acid
25 ml distilled water

They found the best procedure was to clear thoroughly in lactophenol

before mounting, although some of the more delicate mites needed no
special preparation as the lactic acid in the medium cleared the speci-
mens sufficiently. (See dark and Morishita, 1950.)

Nesbitt’s Clearing Solution:

40 g chloral hydrate
25 cc water . ,

2.5 cc hydrochloric acid

Keep preparation tightly stoppered to prevent evaporation and sub-

sequent recrystallization of the chloral hydrate.

. Culture Methods

Some parasitic Mesostigmata are easily cultured, but as is so often

the case, initial efforts to get a culture started can be most frustrating.
We should .hasten to add that only a few species have been cultured, and
all of these are external parasites belonging to the families Laelapti-
dae and Dermanyssidae, No internal parasites have yet been cultured;
neither have any of the snake mites (except Ophionyssus natricis) nor
any of the Spinturnicidae.
Mass cultures. Stock cultures are generally maintained by creating
an environment roughly similar to the natural environment of the host
but with the removal of such undesirable features as predators, fungi,
excessive moisture, and extreme dryness. The most widely used cul-
ture device for bird or mammal mites is a mite-tight container partially
filled with clean straw, hay. excelsior or wood shavings. The host and
a number of mites are introduced into the container and nature is allowed
to take its course. Although this method is generally successful it has
one great disadvantage. The box becomes excessively fouled with ex-
creta which encourages flies, cheese mites and beetles and makes fre-
quent cleaning of the cages necessary. This is a tedious task and may
result in a high loss of mites.
Various researchers have devised various means of circumventing
 268

this inconvenience. Wisseman and Sulkin (1947), in rearing the chicken

mite, Dermanys BUS gallinae, first plugged the anal orifice of a young
chicken before introducing it into’the culture. Chamberlain and Sikes
(1950) left young chicks in the cage only overnight. Cross and Wharton
(1954) fed the host chicks a bare minimum of food and water.
Perhaps the best method is that of Bertram et al. (1946). These
workers filled the bottom of containers with sand. A small tray was
set in the sand and over it was Set a small cage containing the host ani-
mal (white rats in this case). The cage was then surrounded with wood
shavings and mites introduced. In this mariner a nearly normal habitat
is obtained, a minimum of mites are lost because of the, restrained activ-
ity of the host, and th6 tray can be periodically removed and cleaned
without seriously disturbing the mite colony.
Wharton and his students at the University of Maryland have used a
modification of this for small mammals. A tin can,. No. 10 size or lar-
ger, is loosely filled atoout 3/4 full with slightly moistened wood shavings.
The host is placed in a cylinder of hardware cloth and this is immersed
in the shavings. The mites are introduced onto the host, and the can is
placed in a large pan of liquid detergent in water that acts as a moat.
To prevent excessive fouling of the Jar. minimum food and water may
be given to the host.
’:. Strandtmann and his students, in-attempting to create a suitable en-
vironment for rearing Brevisterna on pack rats (Neotoma spp.), devised
a two part container; one part consisting of a metal box 15" x 15" x 15"
and supplied with a lid, the other part being a box 24" x 15" x 8" and
covered with hardware cloth. The two parts are placed end to end and
communication between the two is by means of a 2 inch hole cut through
the apposed walls. Two inches of sand and activated charcoal are put
in the bottom of each box and this is covered with hardware cloth to pre-
vent the rats digging it up. A sheet of plaster of paris is tied into the
lid of the taller box and this is moistened to maintain a desirable rela-
tive humidity in the box. A natural pack rat nest is placed into the box
after first being autoclaved to remove undesirable organisms.
For stock cultures of snake mites, Camin (1953) kept snakes in
wooden boxes set over water on rubber stoppers. The water serves
both as a- barrier against the escape of mites and as a source of mois-
ture for the -high humidity required by this mite.
,
, Water moats are generally Sufficient to keep the mites where they
belong but they may occasionally cross such barriers. For greatest
safety detergent may be added to the water or moats of a nonvolatile
:oil maybe used. Suchmoats can be made an integral part of metal boxes
:or.- several boxes can be set in a large shallow tray filled with oil or
water..;’’ ’.. ..
.
.

Several methods of collecting mites from stock cultures are in.use,

One, method is to keep small squares of corrugated cardboard in.the
cages’; -Engorged mites will crawl into these and when mites are.needed
all that is necessary is to pick up these small squares, hold them.over
a;white,enamel-pan’and tap them with the finger.
 269

Another method is to remove the host a day or two before the mites
are needed. Hungry mites will crawl to the top of the cages and cluster
where they can be easily aspirated. [Chamberlain and Sikes (1950) des-
cribe .an excellent aspirator.]
A third method is to run some of the nesting material through a Ber-
lese funnel to the bottom of which has been screwed a glass jar.
Individual, or life; history cultures. The methods above described
are obviously unsuited for the close observations necessary for life his-
tory studies. For such studies the mite must be restricted for contin-
uous observation and at the same time must have a source of food cons-
tantly available. Several very ingenious methods have been devised for
this purpose.
Camin(1953) placed individual mites (Ophionyssus natricis) insmall
plastic cups and sewed these onto the skin of the snake. Kubber cream
glue was usedto seal any openings between the bottom of the cup and the
snake.
Bertram <rt al. (1946) kept Ornithonyssus bacoti under continuous
observation by confining them in glass vials and periodically inserting
the tail of a rat into the vial. The rat was immobilized during the feed-
ing period and a piece of modeling clay or plasticene was molded around
the base of the tail to keep the mites from escaping. (See also Skaliy
and Hayes. 1949.)
Sikes and Chamber lain (1954) used mass feeding techniques to work
out the life history of bird mites. Fully engorged females were isolated
in glass tubes plugged with black cotton. All the eggs collected within
24 hours were isolated, allowed to hatch and change to protonymphs.
These protonymphs were placed on chicks which were caged over a tray
of water and after a specified time engorged protonymphs were collected
and placed in vials. These were allowed to molt to deutonymphs (and to
adults in the case of Ornithonyssus) and the process repeated- Many
mites are lost by such techniques but the net results are quite satisfac-
tory.
Cross and Wharton (1954) found that mites could be successfully
cultured in small glass tubes by feeding them on heparinized blood through
#173 mesh silk bolting cloth. Owen (1956) has successfully studied the
life cycle of E chinolaelaps echidninus by this method.
There are several methods of collecting eggs or protonymphs of
known age. Bertram et aL (1946) placed several engorged females in
a section of glass tubing, one end of which was covered with #8 silk cloth,
and another tube placed end to end with it. As soon as protonymphs were
present and active, a heated metal ring was slowly passed over the first
tube and the protonymphs driven through the cloth into the second vial.
The females, which could not pass through the cloth, were killed by this
method.
Skaliy and Hayes (1949) placed a strip of roughened filter paper in
a vial with engorged females. The mites preferred to lay their eggs on
this strip and after any specified length of time the filter paper and at-
tached eggs could be withdrawn. Adhering females could be brushed off
270

and saved for future use.

Camin (1953) -divided a glass tube into two compartments with a
screen of 50 mesh Copper cloth. The tubing was then suspended verti-
cally over a pan of water and engorged females introduced into the up-
per half.. The eggs would fall through the screen and were easily col-
lected off the water.
We are not aware of any method for mass collecting of larvae of
known ages of larviparous mites. ; ’.
 LITERATURE CITED

(References marked with an asterisk were not seen]

AINSLIE, C. N. 1929 Note on the occurrence of the mite Dermanyssus

gallinae L. in the nest of a house wren. Canadian Entom. 61: 39-40.
ALDRICH, J. M. .1936 Nycteribia 1796, Pupipara. and Spinturrnx 1826.
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Acarina.
ALICATA. E. 1948 Ectoparasites of mammals and/poultry from Guam
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ALLMAN, G. J. 1847 Description of anewgenus and species of tracheary
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ALLRED, D. M. 1957a Notes on the life history and bionomics of the
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___________ 1957b Mites found on mice of the genus Peromyscus
in Utah. II. Family Haempgamasidae. Proc. Entom. Soc. Washing-
ton 59: 31-39. .
:.; .
.
1.957c. Idem III. Family Dermanyssidae. American
Midland Nat. 57: 450-460. .. ;.
__________ 1957d The male, deutonymph and protonymph of the
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__________
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1957e Anew species of ante, .Hirstionyssus bisetpsus,

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__________ 1.957f Setal variations on mites of the species Brevi-
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.
.
1958b Idem V. Trombiculidae and miscellaneous
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ALLRED, D. M. and BECK, .D. E. 1953 Mite fauna of woodrat nests

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. .

ALLRED,. D. M. and MARCHE.TTE,, .N. .J. 1957 Experimental feeding

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’

.. . .. .
.
ALLRED, D. M, and ROSCOE,. E. J. .1957 Parasitic mites in desert

271
272

wood rat nests with notes on free-living forms. Trans. American

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_______ 1951 Mites on man. Nat..Pest Control Ass., Inc.
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ARNOLD, H. L.. JR. and ARNOLD, H. L., SR. 1943 The diagnosis
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’
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_______’ l948b Notes on parasitic Acari collected from the Man-
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__________ 1948c Ontwo unrecorded mites parasitic on rodents in
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_________ 1949a An observation of the macroscopic fauna in the
nests of some mice and voles, a quantitative study of the samples ob-
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________ 1949b On the occurrence of a parasitic mite, Eulaelaps
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________ 1950 A study on some structures of each stage of para-
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’
’
24-32, .. :
1951a Redescription of Haemogamasus kitanoi Asanuma
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1951b Critical notes on the genera Euhaemogamasus
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from Manchuria. Ibidem No. 24: 15-21.
________ 1951c Myonyssus shibatai: a new parasitic mite of the
family Liponyssidae. Ibidem Nos. 19-21: 79-86.
________ 1952a Anewparasiticmite of the genus Haemogamasus
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________ 1952b Two new species of the blood-sucking mites para-
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________ 1953 Descriptive notes on two new species of parasitic
 273

mites belonging to the genus Haemolaelaps from Japan. Ibidem No.

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ASANUMA, K., JAMESON, E. W., JR.. and SEKIKAWA. K. 1952 On

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______________ 1945 A new genus, new species of dermanyssid
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___________ 1950b Chicken mite, Dermanyssus gallinae (Degeer)
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1950c Tropical rat mite, Liponyssus bacoti Hirst (Aca-
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BANKS. N. 1899 New species of the genus Halarachne. Proc. Entom.
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________ 1901 A new genus of endoparasitic acarians. Geneesk Tid-
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________ 1902 New genera and species of acarians. Canadian En-
tom. 34: 171-176.
_______ 1906 Descriptions of some new mites. Proc. Entom. Soc.
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_______ 1907a A catalogue of the Acarina or mites of the United
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_______ 1907b Mites and .lice on poultry. U. S. Dept. Agric. Bur.
Entom. Circ. 92: 1-3.
274

1909 New Canadian mites. Proc. Entom. Soc.. Washington

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_______1910 New American mites. Ibidem 12: 2-12.
_______1914 [Article describing Qphipmegistus luzonensis. ] Jour.
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.
1915 The Acarina or mites. A review of the group for the
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1917 New mites, mostly economic.; Entom. News 28: 193-
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BARABASH-NIKIFOROV, I. I. 1950 Fauna of muskrat-burrow as a
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.
BARBER. L. B. 1916 Report of the animal husbandman and veterin-
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.
BARNESBY, G. J. 1871 Parasites on canaries. Newman’s Entom. 5:
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BEDFORD, G. A. H. 1932 A synoptic check list and host list of ecto-

parasite’Si: found on South African Mammalia, Aves, and Reptilia. l8th
Rept..Dir.’Vet. Serv. and Animal Ind., U. S. Africa, pp. 233-533.
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of terrestrial vertebrates and man. Med. Parazitol. i Parazitar
Bolezni 17: 385-400.
BELIAEV, A. IA. 1950 Northsakhaline mite as the cause of a new mange
in man. Vestn. Venerol. i Dermatol. 3: 33-39.
BELOPOLSKA, M. M. 1952 Parasitofauna of marine water birds. Utah.
Zap. Leningrad Gos. Univ. #141 Ser. Biol. Nauk #28: 127-180.
BERLESE, A. 1882 Gamasidi nuovi e poco noti. Bull. Soc. Entom.
Italiana 14: 338-352.
__________ 1882-1892 Acari, Myriap.oda et Scorpiones hucusque in
Italia reperta: Mesostigmata. Fasc. 52, No. 2, 143 pp.
_* 190 3a Illustrazione iconografica degli Acari mirmecofili.
Redia 1, 299.
________ 1903b Diagnosi di alcune nuove specie di Acari italiani,
mirmecofili e liberi. ZooL.Anz., Leipzig 27: 12-28.
________ 1904 Acari nuovi,. Manipulus ii. Redia 1: 258-280.
1910 Lista di nuove species e nuovi generi de Acari.
Ibidem 6: 242-271.
1911 Acarorum species novae- quindecim. Ibidem 7:

________
429-435.
1917 Sul Liponyssus natricis (Gerv.) e su altri Derman-
issidi del Bettili.
Ibidem 13: 55-71.
1918 Centuria quarta di Acari nuovi (con 1 tigura nel
teste). Ibidem 13: 115-192.
1921 Centuria quinta di Acari nuovi. Ibidem 14: 143-
195.
__________ 1923. Centuria sesta di Acari nuovi. Ibidem 15: 237-262.
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 275

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1949 Studies on the transmission of cotton rat fila-
_
_
_
_
_
_
_
_
_
_
_
_
_
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_______: . .
1950 Idem, II, Factors influencing the efficiency of
.’BERTRAM,
.
.
,

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D. S.. UNSWORTH, K. and GORDON. R. M. 1946 .The
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BISHQRP,. F..C, 1923 The’ra^ mite.(Liponyssus bacoti) attacking man.

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.

BISHOPP,..F. C. and PHILIP, C. B. 1952 Carriers of human disease. .

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.BISHOPP, F. C. and.WOOD, H. P.. 1931 Mites and lice on poultry.
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.
.
BQYD, E. M. 1951a The external parasites of birds: A review. Wilson
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BREGETOVA, N.G. 1949a Contributioritothefaunaof the mites belong-
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;,
______________ 1949b Of parasitic mites of the genus Myonyssus
^

(Gamaspidea, Liponyssidae). Rept. Acad. Sci. U.S.S.R. 67; 751-

.
..
’753.’, .’, .
.
______________ 1950a New species of endoparasitic mites of the
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_____________ 1950b Mites parasitizing the nasal cavity of birds.
,Parazitol. Sb..Zool. .Inst. Acad. Sci. U.S.S.R. 13: 111-119.
_____________ 1.952a New species of mites of the genus Haemo-
laelaps (Gamasoidea,’Laelaptidae) parasitizing rodents. Zool. Zhur.
’
31: 866-874. _ .. ,. ,,, , ,
. .
..
_____________ 1952b Collection and study of gamasid mites .
_____________
Moskva-Leningrad. 33 pp.
1953a.To the. fauna of gamasid mites of the Far
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. .

_____________ 1953b Mites of the genus Bdellonyssus Fonseca

276

1941 (Gamasoidea, Liponyssidae). Tr. Zool. Inst. Acad. Sci. U.S.

S.R. 13: 310-319.
______________ 1954 Gamasoidea of the central basin of Riwer
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______________ 1955a Diagnosis of the genera Androlgelaps, ’Hae
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of Androlaelaps Berlese. Ibidem 21: 231-240.
______________ 1955b Mites of the rodent fauna of the U.S. S.R.
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_____________’ 1956a Gamasid Mites (Gamasoidea). ’Acad. Sci.
U.S.S.B., Moskva. 247pp.
’___’_____" 1956b New d’ata on the mites of family Haemoga-
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.;

’
’

Zhur. 3’5.
BREGETOVA, N’. G. and KOLPAKOVA, S. A. ’1952 Gamasid mites
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___________________ 1956 Garoasid mites (Parasitiformes, Ga-
masoidea) - parasites of small mouselike rodents and their nests in
the Volga Delta. Ibidem 16: 184-197.
BREGETOVA, N. G. and NELZINA.’E. N. 19’52 -The marmot mite
Haemogamasus citelli Breg. andNelz. sp. nov. (Gamasoidea, Haemo-
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BREGETOVA. N. G. and VYSOTSKAYA, S. 0. 1949 Gamasid Acarina
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BROWN, A. W. A. 1951 Insect Control by Chemicals. Chapt. 11, In-
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BROWN, J. H. 1953 A chicken mite infestation in a hospital. Jour.
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BROWNING, E. 1950 On the occurrence of the tropical rat raite, Bdel-
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BRUMPT, E. *1936 Bdellonyssus bacoti Hirst. Precis de Parasitol.
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.
BUITENDIJK, A. M. *1945 Voorloopige catalogus van de Acari in de
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 277.
BUSHNELL, L. D, andBRANDLY, C. A. 1929 Poultry diseases and
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CAMIN, J. H. 1948 Mite transmission of a haemorragic septicemia

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__________. 1949 .An attempt to clarify the status of the species in
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__________ .1950 An isolation chamber for; .the study of individual
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1953 Observations on the life history and sensory be-
_
_
_
_
_
_
_
_
havior of. the snake mite, Ophionyssus natricia (Gervais). Chicago
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CAMIN, J. H. and GORIROSSI, F. E. ’1955 A revision of the sub-order
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70 pp. .
..

.
CAMIN, J. H. and ROGOFF. W. M. 1952 Mites affecting domesticated
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.
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1888 Idem. Ibidem 6: 142-143.
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CASTRO, M. P. de 1948 Reestruturacao generica da familia: "Rhino-
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284. .. ..
CASTRO, M. P. de and PEREIRA, C. 1947 Acaros nasicolas (Para-
sitiformes: ^Rhinonyssidae) do Pardal -. "Passer Domesticus L."
’

Ibidem 18: 125-133.

CHAMBERLAIN, R. W. and SIKES,. .R. K. 1950 Laboratory rearing
methods for three common species of bird mites. Jour. Parasitol.
36: 461-465. .. : .

_______________ 1955 Laboratory investigations on the role of

bird mites in the transmission of eastern and western equine enceph-
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278

CHAMBERLAIN,.’R. W.," SIKES.. R. K. and: SUDIA,.W.. D. 1957 At-

, ’tempted laboratory infection of bird mites-with the’ virus of St. Louis
’’

’
.. encephalitis. Ibidem 6:..10’.47-105.3.: : : .’

CHANDLER, W. L. and .R’UHE^ L.-’S.. .1940 Pneum-onyssus caninum

.
26:. 59-70. ; .
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... ;. .

.
.
.
.
CLARK. iE. W. and MORISH.ITA, P..’: 1950 C-M medium: A .mounting
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CLARK, G. M, .-1958 Hepatozoon griseisciuri n. sp’;; a new species .of
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CLARK, G. M. and YUNKER, C. E. 1-956 A new-genus, and.-species of
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.
.

COLE, L. C. andKOEPKE, J.A.. 1946. Study of rodent-ectoparasites

.in-Mobile, Ala. .U..-S. Publ. Health Repts. 61: .1469-14.87.
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.

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’. . ’.’,.._____
Ibid.em: 25-41. . ... ....
1947b ’Study of rodent-ectoparasites-in.Honolutu.’

.
.
______________ 1947c Astudy of rodent-ectoparasites :in Savan-
.;.’ ’’
nah, Georgia. Ibidem: 42-60.
[_____________. l’947d A study of rodent-ectoparasites in Doth-
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COLLINS, B. J. 1931 Confused nomenclature- of’Nycteribia Latreil.le,.
179.6 and Spmturmx Heyden 1826. Nat. Inst. Health Bull. 155: 743-
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.
.
COOREMAN, J. 1943 Note sur la faune des Hautes-.Fa.nges en Bel-
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_____________
,
.
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’
.___. .: 1946a Observations sur Pneumonyssus duttoniNew-
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1946b Rhinoecius oti n, gen.,’.ii. sp. (Acarien,
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.
__________ 1955 A study of factors that may contribute to the host
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 279

Univ. Maryland. College Park. 16S pp.

CROSS, H. F. and WHARTON. G. W. 1954 Techniques for testing the
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CROSSLEY, D. A., JR. 1.950 A new species of nasal mite, Neonyssus
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________________ 1951 Nasal mites of some c’olumbiform birds.
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OUTRIGHT, C. R. 1929 A valuable aid in the control of the feather

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.
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’
. .

DEGEER, ’C.’ 1778 Des mittes qui vivant sur les oiseaux. Mem. Hist.
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.
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1944 A new species of endoparasitic mite of
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_
_
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_
_
_
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_
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:i

’ ":

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.
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_______________ 1951b A new mite from nests of the-wood rat,
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.
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.
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________ 1913 Some external parasites of poultry. OregonAgric.
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_
_
_
_
_
_
_
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1924a A new mite from the lung sac of a rattlesnake.
_
_
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_
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EWING, H. E. and BAKER, E. W. 1947 Myonyssus. jamesoni, a new

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282

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’ "

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FAIN, A. 1949 Contribution a 1’etude des Arthropodes piqueurs dans

Ie territoire de Bannmgville (Regions du Bas-Kwangoet Bas-Kwilu), ’

.".

’
’

Rev. Zool. Bot. Afr. 42: 175-182..

.
. .

______ 1952a Sur les acariens parasite.s du genre Pneumonyssus

auCongoBelge. Description de deux especes nouvelles chez Ie daman
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_______ 1952b L’acariase pulmonaire au Congo Beige. Ann, Soc.
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_______ 1954 Notes sur les acariens de genre Pneumonyssus an Colt-
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_______ 1955a Un nouvelacarien parasite des fosses nasiales.dupha-
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Rev. Zool. Bot. Afr. 51: 293-303.

,
_______

.
1955b Deux nouveaux acariens de la famille Halarachnidae
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_______ 1956 Les acariens de lafamille Rhinonyssidae Vitzthum 1935
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.
_______1957a Essai de classification des Rhinonyssidae (Acari: Meso-
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.
_______ 1957c. Les acariens du genre-Astridiella n. g. (Rhinonyssidae)
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_______ 1957d Les acariens des families Epidermoptidae et Rhino-

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_______ 1957f Notes on nasal mites of birds from South Africa .with
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.
,
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____________ 1932b Idena III - Parasitismo do homem e da Cavia
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___________ 1934 Per Schlangenparasit Ixobioides butantanensis
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___________ l935a Notas de Acareologia X - Occorrencia, em
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___________ 1935c Idem XIII New South American species of
-
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284

__________ 1935-1936b IdernXDC Generoseespeciesdeacari- -

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18.’
:
"’ ’:’’ :"-’" .’

’
’. ’ ’

.
__
1935-1936’e ’Idem XX -’ Especies novas deacarianos
__
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^S-l^Wd Idem XXII - Liponissus haem’atophagus
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1935-1936e Idem XXIV - A) Representante brasil-
_
_
_
_
_
_
_
_
_
_
eiro do genero Dermahyssus’Duges, 1834. Ibidem 10: 51-57.
__________ l935-1936f Idem XXIV
- B) Nota sobre o home gen-

___________
erico Parataelaps. Ibidem 10: 58-59.
1938-1939a Idem XXV - The giant Laelaptidafe,.para-
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___________ l938-1939b Idem XXVI - N-ew studies oh the genus

Laelaps Koch, 1836. Ibidem 12: 12 5-14-5.

.
’"’
’____"’ 1938-1939c Liponissus brasiliensis, sp. n. usual
parasite of rodents and accidental of man. Ibidem 12: 148-160.
’

’
’
1938-1939d Notas de Acareologia XXVIII - Occur-
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Dermanyssi’dae). Ibidem 12: 1. ’ .

___________ 1940a IdemXXIX-- Dasyponyssus neivai, gen. n.,

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_____ 1940b IdemXXX - Familias, genro e espeeie novos
’
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1940c Idem XXXI - Bolivilaelapstricholabi.atus, gen.
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___________ 1941a Idem XXXII - Novas especiesBrasileiras do
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194lb Idem XXXIV - Posicao do genero Liponissus
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___________ 1948 A monograph, of the genera and species ofMac-
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’

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___________
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 289

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 290

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_____
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.
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.
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.
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’"
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.
OLSUFEV, N. G. 1940a The role of ectoparasites in the dissernina.tion
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’

’
60: 42-55.

.
.
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.

1943 Parasitology of tularaemia. Sb. "Tularemic

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’
’:- ’

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’
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1898 Insects affecting domesticated animals. U. S. Dept.
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____________
*1902(i’Idem III. Ibidem (8). Deel 1. 1 Nov. 46-
49. .. _ .

____________
,

l9’02d New list of Dutch Ac.ar.i,; second, part. With

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 299

,". . 1903 Idem Fifth Series.’ Tijdschr. Entoia. Deel

.

.
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____________ 1904a Idem Seventh.Series,. Tijdscbr. Nederland.

’
Dierk. Vereen. 2. s., Deel 8(1), Feb. 17-34.
.

.
.
1904b Idem Eighth Series. Ibidem (2), Nov. 70-
~Q2~.
_
_
_
_
_
_
_
_
_
_
: ’....

.
1904c Acarologische aanteekeningen. XII .Entom.
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^^__
1910b Acarologische aanteekeningen. XXXII. En-
_
_
_
_
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____________ 1912b Idem XLIV. Ibidem (68), 1 Nov. 291-292.
____________ 1913a Idem XLVIII. Ibidem (72) Peel 3, 1 Juli.
384-387.

.
.
____________ 1913b Idem XLIX. Ibidem (73) Deel 4:. 2-9.
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____
1914 Acarologische aanteekeningen. LII. Entoin,
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__
l915a Pneumotuber macaci Landois and Hoepke.
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l915c Acarologische aanteekeningen. LVI. En-
_
_
_
_
_
_
_
_
_
_
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____________ 1916b Idem LX. Ibidem (91) Deel 4, ISept. 308-
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^^
____
1923 Studie overdesedert 1877 ontworpen system-
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___
1925 Acarologische aanteekeningen. LXXIX. En-
_
_
_
_
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_____________ 1926b Idem LXXXII. roidem (150) 7, 1 Juli. 119-
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_____________ 1926c Halarachne-Studien. Arch. Katurg., Berlin
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_____________ 1926d KritischHistorischOverzicht derAcarologie
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1927a Laelaps-Studien. Ibidem 70: 163-209.
300

______... 1927b Acari uit Ambon. Zopl. Mededeel. Rijks-

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1928 Acar.qlogische aanteekenihgen. XCIV. En-

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....
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.
_______’ 1929b Acarologische aanteekeningen. XCV. En-
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....

.
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-

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"

267. ’"

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302

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’
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. ’’
.. ’
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.
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____
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’
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’
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_______________ 1945 Further isolation of St.-Louis encephalitis
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,

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.
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.
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’
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.
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.
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._______

’
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.
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’: ’"’
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__________’
...’

’"’ ’
’
1949 ’The
’

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of the genus

_______
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.
_________’ :,:’ ’

1951 The’mesostigmatic nasal miteg of birds.

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.
.
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’

^
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 309

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 310

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"
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.
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’
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’ ’

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’
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’

’ ’
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____________

___
1943 Parasitische Milben von Kleinsaugern aus

:
’
.
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1948- Parasitologische Notizen. Zur Frage der

_____
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 313
"
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___
’

28: 530-538.’ "

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.
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’
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’

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.
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 315

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__ __
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.
.
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.
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_
_
_
_
_
_
_
_
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_
_
_
_
_
_
_
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316 ...

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: .
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’ :.
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’
’

__________________ 1954b -Four new bloodsucking mites from

; . . .
the Ethiopian region (Acarina; Laelaptidae and Spinturnicidae). Rev.
Ecuat. EntOTO.-Parasitol.-2; 209-218. .’,.

.
.
’
_____________/___^ 1954c The lung and nasal mites .of the genus
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new species from African primates. Jour. Entom-. S.oo. S.. Africa
17: 195-212.
’
’
- .’ . ... .. ....’.
’

____’ ’ 1955 Nasal mites of birds hitherto known

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_________________ 1956 -Notes on Haemolaelaps glaagowi (Ewing)
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:
17: 282-291.
 INDEX
[Underlined names’-’= synonymy; underlined numbers = major reference.]
abdomen, 1 americanus, Hemilaelaps, 139
abyssinicus, Spinturnix,. 184,, americanus, Ornithonyssus, 82,
Acanthochela, 193. ., i5
acariasis, respiratory, 3 ;
americanus. Spinturnix, 185, 186

.
Acarus, 65, 102., 115, ’1,20. 121. amphibius, Laelaps, 5,7, 67

’ "..
’

183 anal:plate, 24, 25, 26 ,

Aceosejidae, 27 .... anal pore, 25

.
.
acuminatus, Spinturnix, 184 anatomy, 15
aegyptius, Allodermanys.sus, 119 anatomy, external, 15
aegyptius, Haemolaelaps. 33 anatomy, internal, 18
.
aethippi.cus, Hirstionyssu.s, 104
’
Ancy.stropus, 183
aethiopicus, Laelaps, 59 androgynus, Haemolaelaps, 33
.
aethiopicus, Ornithonyssus, 85 Androlaelaps, 196. 197
aethiopicus, Spinfurnix, 18^ andropadi.. Ptilonyssus, 179
’
Aetholaelaps, 32, 7^ angrensis, Cas, J_69
affinis, Hirstionyssus, 100. angus’tiscntis, Haemolaelaps. 3.3
afribyx, Rhinonyssus, 158 anomalus, Mesolaelaps, 54
africa,na, Manitherionyssi,is. 143 Antennophorina, 2, 196 . .

africanus. Periglischrus. ,1.90 Antennophorotdea, 2, 197

africanus, Rhinoecius, 1,64, antipodUaa, Laelaps, 55
agilis, Laelaps, 57. 59,, 61, 63. antipodianttm, Heterolaelaps, 5.5
"*’
65, 71 antipodianus, Heterolaelaps, 55
agrarius, Laelaps, 59 antipodianus, Spinturnix, 185

.
ala?kensis. Haemogamasus,. 129, anus, 18
’i’32, 133 .’ aplodontiae, Alphalaelaps, 193,
.
alaskensis, Laelaps, 59 194
albatp-affinis, Hirs’ti.onyssus, 102 apus, Rhinonyssus, 158
albatus, Hirstionyssus, 99 arabicus, Ophionyssus, 97
alberti, Rhinonyssus, 158 Arachnida, 1
algericus, Laelaps, 59 aragonensis, Echinolaelaps, 73
allergy, 4 araguensis, Spinturnix, 185, 186
Allodermanyssus, 77, 78. 118, Aramite, 6
120 arcualis, Eulaelaps, 127
AHonyssus, 91_ arcuatus, Hirstionyssus, 99
Alphalaelaps, 193 ardeae, Neonyssus, 161
amboingnsis, Spinturnix, 184 aristoterisi, Pneumophionyssus,
ambularig, Haemogamasus, 4, 13, 147
139. 130, 131 armatus, Ischyropoda, 137
americana, Halarachne, 149 .? arnhemlandensis, Sauronyssus,
americana, Ophiopneumicola, 14.6
aniericsjius, Dermanyssus, 122
66.
artibiensis, Spinturnix, 185

[317]
318

arvalis, Laelaps, 60 berlesei, Echinolaelaps,. 74

arvicanthis, Haemolaelaps, 34. 42 berlesei, Hirstionyssus, 102
arvicolae, Hirstionyssus, 104 berlesei., Laelaps, 63
arvicolae, Laelaps, Oudemans, berirtu’daensfe. Laelaps, 66 ;: :
1916, 58, 60, 65, 66 bhutanensis, Laelaps, 60
arvicolae, Laelaps, Oudemans, bibbyi,,,,Haemolaelaps, 34

.
1927, 58, 65’ ’ ’’." bibikovae, Oryctolaelaps, 71. 72
arvicolarum, Haemogamasus, 132 biscutatus, Pellonyssus. 116.
;
Ascaidae, 27 ’’’" ’:i "

bisetpsus. Rhinoepius, 164

asema, Periglischrus, 190 -." ’

bi’spihosus. Neopars.laelaps, 53
assimilis, Laelaps,-60 ’ blanchardi, Hirstionyssns. Troues-

’
astridae, Paraneonyssus, 176 sart, 1904,. l0\_
’. :

’
Astridiella, 173, 174

’
’:
blanchardi.. Hirstionyssus, Zemska,
atheruri. Rhinophaga, 154 1951, 103 ., .’

_
athleticus, Radtordiella, 117, 118 Bolivilaelaps,. 31, 45

.
,
.
Atrlcholaelaps, 32^
’
boydi. Sternostoma, .166

.
attenuata, Orthohalarachne, 149,’ brachypeltis, Steatqnys.s.us, 113
150 : ’ brachy’spinosus.. .Gigantolaelaps, ^0
atypicus, Echinolaelaps, 73 brasiliensis, Liponysspides, 120
aureliani Ptilonyssus,’ 179
;1’’’-’’’
’ braziliensis,. Cavilaelaps, 48
auris, Ratllietia, 141 braziliensis, Ornithonyssus, 85
australicus, Ichoronyssus, 90 bregetovae, Hirstionyssus, 101

.
australiensis, Mesolaelaps, 54 ’ bresslaui, Cayilaelaps.. 47, 48

.
Australolaelaps, TS^ 98 ’ "’ brevis, Dermanyssus, 122 ..

.
avisugus, Haemogamasus,: 131 breviseta, Hirstionyssus, 199, 101
a»i.um, Dermanyssus, :.l2.1:’ Brevisterria, 9, 126, 136, 1.99, 268
’
avium, Steatonyssus, ’1’13
’
’
britteni, Hirstesia, 111
brucei. Steatonyssus,, 114
bacoti, Ornithonyssus, 4, 5, ’6,’7, b’ubuici, Neonyssus, .1.62
8, 9, 10, 11,. 12, 14,81, 82, buloloensis, Celaenopsoides, 192,
8^, 86, 200, 261, 262, 269 I93
’

bacteria, 3 ;’
bursa, 6.rrii.thony.s,sus,.. ,5. 8, 9. tO,
Bdellonyssus, 81 11, 12, QQ, 198, 261,. 262
bakeri, Laelaps, 60 ’ burrows, 198, 199. .
’
... .

bakeri, Pneumdnyssus, 147, 151, butantanensis, Gigantolaelaps, ,.51

"
152 butantanensis, Hirstionyssus.,. 101
bandicoota, Neoparalaelaps, 53 butantanensis, Ixodorhynchus,
banksi, Ornithonyssus, new nanie, 138, 139
..
’

t5 buteonis, Neonysaus, 16,2

’
.

Banksia, 196
’
buxtoni, Laelaps., 61_ .. .:
barbatus. Liponysella. ’112
barberi, Haemogainasus, 131 calamocichlae, I’Ptilony.ssus, 179
bathyergus, Haernolaelaps, 34 calcaratus, .Spinturnix, 1.8,5 .:.;,..
batis, Sternostoma, 167 caledonicutn., Rhinonyssus, 158.
bedtordi, Hamertonia, 146
behavior, T_ :
belopolskii, Neonyssus, 162
. ’
caledpnicus,. Rhinpn.yssu.s^ 158
califorriicus, Haemolaelaps, 38
.caligus, Periglischrus, 189, 190
benzene hexachloride, 6 camera spermatis, 23
 319

canadensis, Ornithonyssus, 82 circulatory system, 19

caninum. Pneumonyssoides, 6, citelli, Haemogamasus, 131
154 citellus, Eulaelaps. 128
capensis, Haemolaelaps, 34, . classification, 1,26
capensis., ..Mypnyssoides, 194,. 195 clearing solution, Nesbitt’s, 264,
capensis, Paraneonyssus, 176 265. 267
capitatus, Paraneonyssus, 177 clethrionomydis, Laelaps, 61
capitulum, 1 coelogenys, Haemolaelaps, 35
caprimulgi, Astridiella, 174, 175 coition, 13
caraco, Laelaps, .66 colii,’ Sternostoma, 166
carinii, Litomosoides, 84, 85, 200 collecting methods, 263
Cans, 81 colubri, Ophioneumicola, 145,’ 146
carlo.shoffmanni,. .Spinturnix. .185 columbae, Neonyssus, 162
carnifex, Hirstionyssus, 100,101,107 comatus, Gigantolaelaps, 51’
Cas, .155, 169 : .
comatus, Tricholaelaps, 55
.
.

casalis, Haemolaelaps, 34 . concur’rens, Haemolaelaps, 35

castroae, Sternostoma, 169 confuscianus, Hirstionyssus, 102
caucasicus, Laelaps, 61 congoensis, Pneum.onyssas, 152
caudistigmus, Rallinyssus, 21.,. congolensis, Rallinyssus, 160
160 . . .
coniventris, Rhinonyssus, 16, 1.57
CavUaelaps, 31, .47 constrictus, Ornithoriyssus, 86
cecae. 18 contact response, 9
Celaenopsoidea, 2 control, 5
Celaenopsoides, 19.2 cooremani, Rhinoectus, 164
.
cephalothorax, 1. cooremani, Sternostorna, 166
ceratognathus. Steatonyssus, 114 coriaceus, Eugynola’elaps, 196
Ceratonyssus, 113 coriaceus, Hirstionyssus, 100
cerchneis. Rhinonyssoide.s, 172 corniculi, 16
’
Cercornegistina, 2 : cornutus, Ichoronyssus, 90
Cercomeg’istoidea, 2. coronis, Manisilaelaps, 194
cercopitheci, Rhinophaga, 153 Cosmolaelaps, 198
ceylonicua, Echinolaelaps, 73. coxae, 16
chaetotaxy, 17 Coxsakie virus, 84
’
Ghelanyssus, 85, 89, 90 . .. ,
crarikcase oil. control with, 5
chelicerae, 1. 15, 1.9 crassipes, Trichosurplaelaps, 56
chelidonis, Dermanyssus, 122 . .
crassipus, Laelaps, 59
chemoreceptors, 17 craticulatus, Hirstionyssus, 102
chilensis, Acanthochela, 193 .
.
creightoni. Hirstionyssus, 102
chilensi.s, Ichoronyssus, 94 creosote, control with, 5
chinchillulae. Haemolaelaps, 3.5 cribrum, 25
chiropteralis, Steatonyssus, 1.13 criceti, Hirstionyssus, 102
Chiropterolaelaps, 90 cricetidarum, Gigantolaelaps, 51
Chiroptonyssus, 89^ ., .
Cricetilaelaps, 58
chlordane, 6 cricetophilus, Haemolaelaps, 38
chlorocichlae, Ptilonyssus, 179 cricetuli, Eulaelaps, 128
cinnyris, Ptilonyssus, 172 crosbyi, Ichoronyssus, 90’
circularis, Eubrachylaelaps, 48, crowei, Eubrachylaelaps, 49
49 cryptomius, Haemolaelaps, 35
320

cryptorhynchum, Sternostoma, disirnllis, Haemolaelaps, 36

165 ,
.
.. ..-. :. distinctus, Hemilaelaps, 139
Cryptostoma, _57 ., ’:’

’
.; .
distribution, 198 :;

.
.

cuculorum, Sternpstoma, 1.6.6 -.. div.ersipilus, Ichoronyssus, 91

cultures, individual, 269,. ^’.:
cultures,mass, 267 ..’..’ . .
dogiel-i, Ornithdnysisus, 87
donatoi, Rhinonyssoides, 172

.
culture methods. 267; - , dorsal plates, 25

-
.

cuticle. IT^ ;...:,. .

dubinini, Myonyssus, 124
cyclaspis, Steatpnyssus, -1 1:4 ...
dureni. Sternostoma,’ 166
Cyclolaelaps, 41. 48 : . .
duttoni, Prieurnonyssus’, 152

.
.
cynomys, .Hirstionyssus, 1&3, 106 duttoni, Spirochaeta, 84
dryoscopi. Paraneonyssus, 177
dartevelle.i, ..Rhinonyssus, 159
dasymys, Haemolaelaps. 35 easti, Ophionyssus, 97’
Dasyponyssidae, 2. 2.9. 1.42 echidninus, Echinolaelaps. 4, 7,
Dasyponyssus, 142, 143 8,.-9, 10; 13, 14, 18. ^2,

.
.
dauricus,, Haemogamasus, 132 > 200, 261, 262. 269
davisi, Haemolaelaps, 36 echiriatus, Ptilonyssus,’ 178
Davisiella, .196. . .
.
echinipes, Rhinonyssus, 15.7
DDT., 5. 6..7 .
echinipes, Spinturnix. 185, 1’86
debilis, Eubrachy.lae.lapa, -.49 .
Echinolaelaps, 32, n_
decumani, Myonyssus, 124 .
Echinonyssus, 78, 109 ’
.
decussatus, Hirstionyssus, 100,. echinus, Neolaelaps,’ 112
102 .. , ... ecological factors, 199
.
.
,
degeneratus, Heterozercon, 192 economic importance," ’3
dendropicus, Ornithonyssus, ..87: ectostracum, 17’.

.
dentipes, Neoichoronyssus,. 108’.. :
eggs,’ 11, 23^
dermal glands.,.. 18 :

’
,
ejaculatory duct, 22
Dermanyssidae, 2, 9, 10, 16, 23, ekstremi, Laelaps, 61 ’
24. 25, 28,. ^77, 1.41, 194.. elaph’es, Ophiopneunaicola, 146
.
196, 267 ellipsoideus, Haemogamasus, 132’
Dermanyssinae,. 78, 118 .
ellipticus, Macronyssus, JU
.
Dermanyssus, 17, 77.. 78,. 120, ellobii, Haemolaelaps; 36
1:74 ,,... ... .. .. ellobii, Hirstionyssus. 103
.
dermatitis, 4. .. ... . . .: Ellsworthia, 139
"

desfontainei,. Ptilonyssus, 179 :. eloffi, Haemolaelaps, 36

’
deutonyroph, .-11,. 2^4 ; ,; , ,
elongatus, Haemolaelaps, 36
:

deutosternal teeth,. 1& , :. .

emberizae, Paraneonyssus; 177’.
.
.
deutosternum, 15 ,.,.. .:. . .
encephalitides, 3, 86 ’ ’
’

Diarthrophalloidea,. 1 ;,: ; .!, encephalitis. Eastern equine’, 3,

.
.
dicruri, Ptil.onyssQides,:rl70’
.
;
.
121 "
"’’"’,
"

differens, Laelaps, 61 .:
encephalitis, St. Louis. 3; 83,"1.21
digestive system,, j^ ,. encephalitis. Western’equine, 3;
’
. , .

;:
diminuata. Orthohalarachne, ,150 83;- 121 : ’ ’:’

dinoiti, Pneumonyssus, 152 ’’

’
’
endopodal plate, 25
dioptrornis, Ptilonyssoides, 170 ~~
endoskeleton, ^8 ’ ’

Diplostaspis, 183 :, ,, .
enrietti, Paraneonyssus. 176
disease transmission, 3, 86 Entonyssidae, 2, 29. 143
 321

Entonyssus, 144; ,:, . .. ., ,... _ gallinae, .Dermanyssus, 3, 4, 5,

.
eos,: Haemolaelaps, .3.8 ,,:.. 8, 9, 10, 12. 14. 86, 120,

.
eos, Steatonyssus, 114 :,: ; ;.- 121, 122..: 12,3,..198, 200.
epigynial plate, 24 ; ’,: :;. . ,. 261, 262, 268 ... ,

epipharynx, _1_6, 18, 19 . Gamasina, 1. ; :.

.
.

equilibrium center, -20 ’..:’. ; .. Gamasolaelaps, 198

eruditus, Ornithonyssus, 87 Gamasus, .113, .126, 141
esophagus, 18 . Geneiadol.aelaps, 196

.
.
.
Eubrachylaelaps, 16, 31, 4.8.,; 196 genital plate, 24,.’2 6
Eugynolaelaps, 196 : .
genital pore, 23. 25
Euhaemogamasus, 129 genitoventral plate, 24
’

Eulaelaps, 25. 126. 139, 196, 20.0 geographical distribution, 199, 262
euryalis, Spinturnix, 186. geomydis, Hirstionyssus, 103
eusoricis, Hirstionyssus, 103 geomys, Haemolaelaps, 37, 261
evers:manni, Hirstionyssus, 103 georgicus, Hirstionyssus, 103
Eviphididae, 27 geotaxis, 20
.
evotomydis, Dermanyssus, 122 giganteus, Echinolaelaps, 74
ewingi, Entonyssus, 144 giganteus, Laelaps, 60
ewingi, Haemolaelaps, 36 Gigantolaelaps, 31, 50, 196
ewingia, Spinturnix, 186 gigas, Myonyssus, 124, 125
exceptipnalis, Laelaps, 62 gilmorei, Gigantolaelaps, 51
excretory system, 19 glasgowi, Haemolaelaps, 9. 10,
eyes, 20 13, 14, y!_, 199, 200, 261,
"eye spots, " 21 262
glasmacheri, Entonyssus, 145
fahrenhoizi, Haemolaelaps, 37 glutinimargo, Periglischrus, 190
farrieri. Hemilaelaps. 139 glutinbsus, Ichoronyssus, 91
Pednzzioidea. 2 gnathosoma, 1, 15. 197
feeding, 11 gnathosomal base ring, 15
female. 24 Gneidolaelaps, 38, 112
festinus, Laelaps, 58, 59 gordonensis, Sauronyssus, 96
filaria, 82, 84, 85 goyanensis, Gigantolaelaps, 51
finlay soni, Laelaps, 62 grandis, Echinolaelaps. 74
Flavionyssus, 156, 171 grandis, Rhinoecius, 165
flavus, Ichoronyssus, 91 granulbs.us, Ichoronyssus, 91
Fonsecaonyssus, 81 gravity, response to, 9, 10
food, 2, 198 grenieri, Laelaps, 62
forsythi, Ichoronyssus, 89,90,91 griffithi, Pneumonyssus, 151
fowl pox. 82 griseisciuri. Hepatozoon,: 4
foxi, Pneumonyssus, 151 grossus, Spinturnix, 186
fragilis, Entonyssus, 145 grubei, Laelaps, 64
freemani, Haemolaelaps, 34. ~
guanin, 20
’
funnel, Berlese, 263 Gymnolaelaps. 198
furmani, Ischyropoda, 138.

galagus, Haemolaelaps, 37. haematophagus, Ichoronyssus, 92,

galagus. Ornithonyssus, f37 94
322

Haemogamasidae, 2, 25, 28, 126, Hirstionyssus, 72. 79, 99

193 hirsutosimilis. Haemogamasus,
Haemogamasus, 126, 129, 200 132

-
,
,

haemogregarine .4. 96, 98 hirsutus. Haemogamasus, iQi^ZS,

Haemolaelaps, 31. 32, 127, 139, 129
141. 200 hirundinis, Dermanyssus; 122,:.:
haemorrhagic fever, 5 123 :.:.. ,
haemorrhagic septicemia,. 4, 98 hirundinis, Sternostoma, 167 .

haemorrhagicus, Haemolaelaps, hoogstraali, Ichoronyssus, 92

34 hollisteri, Eubrachylaelap.s, 48
Halarachne. 148 . : holoventral plate,. 25
Halarachnidae, 2, 25. 29, 147 hopkinsi, Raillietia, 141
halichoeri, Halarachne. 148 horridus. Haemogamasus, 19. ’20,
halli, Entonyssus, .144
’

.. 132 :

.
.
.
.
.
,
hamertoni, Ophiopneumicola, -.146 hosts. 199,’ 215 .

Hamertonia, 144. 146 ;-. ... host specificity, 199. 261 ’.

.
-
.
hapaloti, Laelaps, 62 hubbardi. Patrinyssus, 10 9.’
harperi, Haemogamasus, 132; human habitation, mites in, 5

’
’
.
.
hasei, Ichoronyssu’s, 89, 92 :, .
humidity, response’to, 8, .10
hawaiiensis, Laelaps,. 66 .;... , hutsoni, Sternosto’ma, 167 : . ’"
heart, 19 .".,..V; ., hydrophilus. Pseudomonas,. 4,’ 98’
heat, response to, 7, 10 : .... Hyletastinae, 27 . ...

.
... .
.
.
helminths, 3, 4 ...
.
.. Hyperlaelaps.- ^7, ’67
.
.
.
Hemilaelaps Ewing; 138, .139 Hypoaspidinae. 27, 196
Hemilaelaps Hull, J.26. : ., hypoaspidine. 197
Hepatozoon, 4, 73 Hypoaspis, 32. 127. 19 6;’ 197 :
hermanni, Ichoronyssus, .,9.5 hypodermis, 17
heterodontus, Entonyssus, 145-
~~-... hypopharyngeal processes,’ 16’’’’
,
Heterolaelaps, 31, 3.2,, ,55 hypopharynx;’.18’ ’

.
.
heteromydis, Steptolaelaps, .56. hypostome, 15 r ’.
’

.
heteromys, Steptolaelaps, 56. hypostomal processes, ’18, 19’’
heterotarsus, Manitberionyssus,. hypostracum. 1_7
142,14_3 . ,:
hystrici, Haemolaelaps; 39_
.
.....
.
Heterozercon, 192.
’
Heterozerconid.ae, 2,.28;"^&1 .
ianzai. Spinturnix. 185 ’’
hilaris, Laelaps, 57, 56. 63. 69 Ichoronyssus, 80, 85, 89, 200,
hilaroides, Laelaps, 62 . . 261, 262
.
.
.
hilli. Hirstionyssus, 104 .
icteridius, Paraneonyssus, n7
himantopus, Rhinonyssus, 19., 158 idiosoma. 15, 16 ’ ’
.

hindgut. 18 iheringi,’ Ornithonyssus, 87

hipposider.us, Periglischrus, .190 iheringi, Periglischrus; 19-1
Hirstesia. 79, 110 ". imphalensis, Hemilaelaps, 139, ’

hirsti, Echinolaelaps, 73. 74. IAO_- . .

.
.
.
hirsti, Haemolaelaps, 39 Inchorony’ssus, 90
hirsti, Hirstionyssus, 104 incomptus, Hirstionyssus, 104’
hirsti. Laelaps, 62 indicatoris, Rhirtonyssoides, 172
hirsti, Ornithonyssus, 87 Indogynium, 19 6
hirsti, Paraneonyssus, 177 ingricus. Myonyssus, 125
 323

mops. Haemolaelaps, 47 labuschagnei, Haemolaelaps, 40

insculptus, Haemolaelaps, 40. lacertarum, Karyolysus, 96
insemination,. 13 lacertinus, Saurdnyssus, 97
integument, 16
intermedius, Neonyssus, 161
Laelaps, 4, 17, 21, 32,
138, 200. 262
5^7 112,

internal mites, 6 Laelaptidae, 2, 13, 16, 23, 24,

interruptus, Periglischrus, 190. 25, 29, 196, 198, 267
191 Laelaptinae, 30, 193, 196, 200
inversus, Haemolaelaps, 40 laelaptine, 197
iowae, Spinturnix, 185 lagonostictae, Sternostoma, 167
Iphiopsidae, 27 lagotisinus, Mesolaelaps, 54
Iphis, 34 lamborni, Laelaps, 63
isabellinus, Hirstionyssus. _104 lanii, Ptilonyssus. 180
Ischnolaelaps., 32 laniorum, Sternostoma, 167
Ischyropoda, 126, 137 lanius, Hirstionyssus, 102
ivanovi, Haemogamasus, 133 Larinyssus, 157, 160
Ixobiodes. 138’ larva, 11, 23
ixobrychi, Neonyssus, 163 lateralis, Spinturnix, 183, 187
Ixodides. 1 latiscutatus, Hirstionyssus, 100,
Ixodorhynchidae. 2, 16, 29, 138 105
Ixodorhynchus, 138 lativentralis, Laelaps, 63
latus, Spelaeorhynchus, 142 ’

jacksoni, Spinolaelaps, 95 lavellanus, Spinturnix, 187

jamesoni, Eubrachylaelaps, 49 lavieri, Laelaps, 63
jamesoni, Myonyssus, 125 lavoipierrei, Laelaps, 64^
japonicus, Haemogamasus, 133 lawrencei, Haemolaelaps. 40.
japuibensis, Ptilonyssus, 180 lawrencei, Spinturnix, 187
javensis, Spinturnix, 186 Leiognathus, 81
javensis, Steatonyssus, 114 Leiostaspis, 183
jettmari, Laelaps. 62 lemmi, Laelaps, 6jl
joaquimi, Steatonyssus, 113, 114 lemni, Laelaps. 64
johnstoni, Hirstionyssus, 105 leopoldi, Rhinophaga. 154
jordani, Spinturnix, 187 lepidopeltis, Ichoronyssus, 93
Lepronyssoides, 79. 110
Karyolysus, .96 Lepronyssus, 89
keegani. Haemogamasus, 133 leprosus, Ichoronyssus, 89, 93
keegam, Laelaps, 63 levin sent, Sommatericola, 157,
,
kenyaensis, Hirstesia, 111 159
kenyaensis, Spinturnix, 187 liberiensis, Haemogamasus, 134
kerosene, control with, 5 liberiensis, Hirstionyssus, 105
kitanoi, Haemogamasus, 133 liberiensis, Laelaps, 64
kochi, Ichoronyssus, 92 life cycles, _1()
kochi, Laelaps, 67 life stages. 2J5
kolenati, Ichoronyssus, 92 light, response to, 9, 10
Kplenationyssus, 117 limnocoracis, Rallinyssus, 160
kolpakovae, Eulaelaps. 128 lindane, 6
kolpakovae. Laelaps, 63 lindbergi, Indogynium, 196
kusumotoi, Haemogamasus, 133 liomydis, Steptolaelaps, 57
324

Liponysella, 80, 112 martini, Eubrachylaelaps, 50

Liponyssoides, 78, 119. 120 mating, 11, 12, JL3_
Uponyssoides, Haemogamasus, mattogrossehsis, Gigantolaelaps,
I

126, ^134. 135. ^1

.
liponyssoides, Ixodorhyncbus, 138 matogrosso, Lepronyssoides, 110
Liponyssus, 81, 143 maur’itanicus, Haemolaelaps, 41
Liroaspina, 1 .’;’.. mawsoni, Haernogamasus, 135
Liroaspoidea, 1. maximus, Gigantolaelaps, 52
Litomosoides, 84, 85, 200 mazzai, Laelaps, 64

.
lobatus, Ichoronyssus, 91_ meddal, Sternostonia, 167
longevity, 14 medical importance. 3_
longimanus. Macronyssus, 81 megaventralis, Haemolaelaps, 34^
longipes, Haemolaelaps, 40 Megisthanoidea, 2
longiseta, Pililaelaps, 196 melittophagi, Ptilonyssoides, 170;
longisetosus. Ichoronyssus. 93 melloi, Neonyssus, 163
longiventris, Laelaps. 64: .’ melomys. My s olael ap s, 7 6
Longolaelaps, 32, 7_1 .: meprai, Ornithonyssus, 88
longulus, Longolaelaps, 7.1 meridensis, Periglischrus, 191
Lopbotes, 82 meridianus, Hirstionyssus, 105
lophuromius, Haemolaelaps, 41_ meridionalis, Radfordilaelaps. 53,
’
:
lung mites, 3 54 :

,
lutzi, Ornithonyssus. 8J Meristaspis, 183
luzonensis, Ophlomegistus, 192, Mesolaelaps, 31, 54 ;.,.,.. .
’
193 mesopicus, Haemolaelaps, 41 ,,.
metapodaltol:ates,’24, 25’...^^. ..^.

macaci, Pneumonyssua, ’1’Sl’ : metapodoSbma. 15

macedonicus, Hirstionyssus, 105 metasternal plate, .25 .. ..:. ,, :

macroglossi, Spinturniic,-. 187 metasternal pore, 25 ’

,^ ... i

Macrolaelaps, T2 ;-: ;.:; metasternal’seta. 2^ .

, .,, .;’
Macronyssidae, 78 ’. :
^ michaeli, Haemogamasus, 130 ,
Macronyssinae, 24, 78, 80, :124

~
Microgynioidea, 1 ... ;:., ,

.
Macronyssus, 78, 80, 8J_ micromydis, Laelaps, 58,. j[5 ’...,
macroventralis. Haemolaelaps, 41 microspiriosus, Mysolaelaps,. ,.7.6;.
madagascariensis, Liponysella, microti, Haemogamasus, 131
112 ;, microti, Haemolaelaps, j58 ,;
.
magellanica. Orthohalarachne. 150 microti, Laelaps, 65
’
. .

magnistigmatus. Neolaelaps, 111, microti, Tetragonyssus,’.^7 _ ,

.

,1-12. Microtilaelaps, 58 ’ .
. . .
.
’
male, 25 midgut, J_8, 19’
’

malpighian tubules. 20 . .. miniopteri, Neospinplaelaps, 98 :

malucus, Pellonyssus. 117 minutus, fehinonyssus, 159

mandschuricus, Haemogamasus,
134, 136
manguinhosi, Laelaps.. 64
. miroungae,’ Halarachhe, 149
mitchelli, Australolaelaps, 98, 99_
miticides, 5
Manisicola, 143 . mohrae, Haemolaelaps. 4, 37, 38 .

Manisilaelaps, 194 .. mohrae, Ichoronyssus, 93 .

.
Manitherionyssus, 142, 143 molestus, Haemolaelaps, 34 .^ ,

naarsupialis, Haemolaelaips, ’32, 33 montanus, Myonyssus,. 125 , . ,,

 325

monodi, Ophionyssus, 97 Myrmolaelaps. 198 .

Monogynaspida, 1. 27 Mysolaelaps, 32. 76

montanus, Brevisterna, 136 mystromys, Haemolaelaps, 42
monteiroi, Ornithonyssus, 88

-
morlani, Brevisterna, 137 nagayoi, Ornithonyssus, 10, 84
morlani, Haemolaelaps, 41 namrui, Haemolaelaps, 42
morsitans, Ornithonyssus, 86 nasutus, Echinonyssus. 109
’
mosquensis, Laelaps, 65 natalensis. Haemolaelaps, 42
"

mossambicensis, Pneumonyssus, natalensis, Steatonyssus, 115

’’
’

’
: "

’
152 natricis, Opbionyssus, 4. 6, 7, 9,
:
motacillae, Ptilohyssus, 180 .10, 11, 12, 14,. 31, 22, 97_,
mounting medium, ’Ber’lese’s, 2’64, 144, 261,.,267, 269
:
265 ’’’ ’
natricis, Ophiopneumicola, 146 .

mounting medium, Doetschman’s, navajasi, Tinaminyssus, .161, 170

266 navasi, Laelaps, _66
mounting medium, Ewing’s, 265 nectarinia, gternostoma. 167
mounting medium, "Ga’ter’s. 265 neivai, Dasyponys.sus, 143
mounting medium, Hoyer’s. 264, nematodes, 4, 200
266 :;." ^. : Neoicho.ronyssus, 79, 108
mounting medium," King, Bradley Neolaelaps, 56,. 80, .111
:
and McNeel’ES, 2B6
’
’
Neoliponyssus, 96

.
.

.
mountijig mediuto’, MethoceUul’ose, Neonyssus, 25, 27,, 157, 161, 174
;

’"
:" i
’ ’

:’"
286 -" :’
’
Neonyssoides, 161
mounting medium, polyvinyl alco- Neoparalaelaps. 30, 53
hol, S64; 266 ’ Neospinolaelaps, 79^ 98
mounting methods, 264 neotis, Vitznyssus, 175
mouthparts, 1 : .
neotomae, Hirstionyssus. 106,
.
.
mulleri, Spin’turnix, ’184 : 199
.
multispinosus, Laelaps, 65, 199,
’ "" ~- nervous system, ^0 ’
.
’
i
’ " ’

’
262 nests, .2. 198 .
.
.
Mungosiicola; 195 Newcastle^ disease, 3, 83
muricola, Echinolaelaps. 74 ngami, Hirstionyssus. 106
murinus, Haemolaelaps, 41 nicotine .sulphate, control with, 6
murinus, Spinturnfai:, 183, 188 nidi, Haemogamasus, 130, 131
murinus, Steatonyssus, 113 nidiformes, Haemogamasus, 1.34
muris, Hepatozodn, 73 nitzschi, Rhinonyssus, 174, 176
muris, Laelaps, 4, 57, j65, 66 nitzschi, Vitznyssus.’ 173, 174,
muris, Liponyssoides, 120 175 ,
musculature, 18 noctulae, Hirstionyssus, 100
musculi, Hirstionyssus, 105 nova-guinea. Scissuralaelaps, 196
musculi, Parasitus,: 10() nova-guineae, Rhinonyssoides,
musculi, Ste’atonyssus, 113, 115
’
173
Myolaelaps, 58 novae-hollandae, Spinturnix. 188
Myohyssinae, 77, 124 novikovae, Laelaps. 66
Myonyssoides, 194’ novus, Eulaelaps, 128
Myonyssus, 77, 124 nucifragae, Neonyssus, 161, 163
myoti, Spinturnix, 183 nudus, Paraneonyssus, 177
myrmecophagus, Sauronyssus, 97 nudus. Ptilonyssus, 177, ’180
326

nuttalli, Laelaps, 8, .10. -13, 14, .

oudemansi, Spintumix, 188
^6. 199, 200,.261 Oudemansiella, 96

-
nyassae, Steatonyssus, 115 ;
ovalis; ..Haemogamasus, 130
nyctinomi, Ichoronyssus, 93 ovary, 23^ ’ ’

nyctinomius, Ichoronyssus, 93;’.. 94 oviduct, 23^ ’

obsoletus. Hirstionyssus, 106 Pachylaeiaps, 198

occidentalis, Haemogamasus, 135 Pachylaelaptidae, ’196
occidentalis, Hirstionyssus,’ 103, pachyptilae, Haemolaelaps, 43
, 106 .;...
.; pachypus, Hirstionyssus, 107
.
.
.
.
occidentalis, Steatonyssus, 115 pachypus, Laelaps, 58, 67, 69
oculatus, Haemolaelap’s, 34 pacificus, Ornithonyssus, 8j2
odor» response to, 8. &, 20. pallidus. Echinolaelaps. 74
’

olaioi,. Ptilonyssus,. 181

’

. . palpal claw," 17 ’ "’

.
olfaction, 20 .
palps, 1, 16-
.
oliffi,. Haemolaelaps, 42, papiohis, Rhinophaga, 154
.
omahonyi, Spintumix, 188
. paradoxus, Tympanospinctus, 195
omnitectus, Haemolaelaps; 42 Paralaelaps, 53
ondatrae, Ornithonyssus. 88 ’. .
Parameglstidae, 2, 192
onychomydis, Haeinogam-asus, Paramelaelaps, 63
129, 130 .. paranensis, Trav’anyssus, 171
.
Onychopalpida, 1 Paraneonyssus, 156, 1’76
Ophidilaelaps, 139. ’. .
parapodal plates, 24, 25
.
Ophiomegistus, -16, 27, 192, 197 Parasitoidea, 1.-2, 4, 27, 28
Ophionyssus, 24, 25, 77. .78, 97^ Parasitus, 100
Ophiopneumicola, 144,: 145 .: .
parthenogenesis, 11. 22
;
opisthosoma, 15 parvanalis, Laelaps,’^5
oraniensis, Laelaps, 67 parvispinosus, Mysblaelaps, 76

.
orblcularis, Larinyssus, 25, 160 parvulus, Laelaps; 68
orcadensis, Macronyssus, 81 passeri, Pellonyssus, 10, 116
oregonensis, Hae-mogamasus, 130’ ’"
Pasteurella. 4, 84
Oribatei,. 200 patavinus, Ornithonyssus, 82
oribatoides, Eulaelaps, 128 patersoni, Haemolaelaps, 43
orientalis; Ptilonyssus, 181 : pathogenicity, 3
orioli, Ptilonyssus; 181 Patrinyssus, 79, 109
Ornithonyssus, 17, 24, 77. 80,’ 81, pauli, Hirstionyssus. 107
200, 269 paulistanensis, Laelaps. 6S
Orthohalarachne. 148, 149 pavlovskyi, Laelaps, 59’. 68
Oryctolaelaps, 31, 71-. pedalis. Eulaelaps, 129
.
oryzomydis, Laelaps, 67 pedipalps, I, 15
otariae, Orthohalarachne, .151
. Pellonyssus, 77, 78, 116
oti, Dermanyssus. 123 pereirai, Lepronyssoides, 110
oti, Rhinoecius, 164 ... periblepharus, Steatonyssus. 113
.
otomys, Hirstionyssus, 100, 106 Periglischrus,- 183, 189.’262
:
oudemansi, Gigantolaelaps, 52 peritremal plate, 24
’
oudenaansi, Haenaogamasus, 135’ peritremalia, 25
.
oudemansi, Heterozercon, 192 peritreme, 22, 23
oudemansi, Radfor-diella. 117, 118 peruvianus,’ Gigantolaelaps, 52
 32?

pestis,’ Pasteur.el’la’, 8.4 : Pteropus, 183 " ’!’

’ ’

phacochoeri, Pneumonyssbides, py’cnonoti, Ptilonyssus, 181

.; ’ 155 pyrethrum, 5, 6

’
.

.
pharynx, 1, 18, 19
phoeniculi, Haemolaelaps, 43 "Q" fever, 54: "
’

;.’" ’ ’

phyllastrephi, Ptilonyssus, 181 quadrldentatuB, ’Ichoronyssus, 93

Phytoseiidae, 27, 194 quadrisetatus.
:
Haemogamasus. ’
’’ "

’
’
" ’;

piger, Hemilaelaps, 140 135

Pililaelaps, 196 queenslandica, Scissuralaelaps, ’

_17, 77

’
pilus dentilis, 196

’
.
’
pipistrellae, Hirstionyssus, 100 quintus, Dermanyssus, "123
pipistrelli, Steatonyssus, 114, 115
pipistrellus, Splnturnix, 188 rabelloi, Flavionyssus, 171
~~

’
pitymidis, Laelaps. 68 Rad, 196
:
"’’’"
plecotinus, Spinturnix, 188 radfordi, Haemolaelaps, 43

’
ploceanus, Paraneonyssus, 178 Radfordiella, 80, 117
;’
Pneumonyssoides, 148, 154 Radfordilaelaps, 31, 53
Pneumonyssus. 148, 151 Raillietia. 141 : :
Pneumophi.onyssus, 143, 144, 147 Raillietidae. 2. 3, 10, 28, MO
’

Pneumotuber, 151 Rallinyssus, 157, 160 "’’

podosoma, 15 Rattilaelaps, 58
poliocephali, Rhinonyssus, 159 raz’umovae. Haemolaelaps,’ 43
polychaeta, Haemogamasus. 133 receptaculum serninis, 23
pontiger, Haemogamasus, 135 rectal sac, 18
populations, 199 reedi, Pellonyssus, 117
pores, VS. 21. 19 reflexa. Orthohalarachne, 150
poultry mites, 5 reidi. Haemogamasus, 130
praecursor, Spelaeorhynchus, 142 reithrodont.is, Haemolaelaps, 32,
presternal plates, 25 43
procavians, Pneumonyssus, 153 reproductive system, 22
prognephilus, Dermanyssus. 123 respiratory system, 21
propheticus, Eulaelaps. 129 reticulata, Davisi.ella, 196
propodosoma, 15 reticulatus, Haemolaelaps, 32, 44
proteronymph, 23 rhabdomys, Haemolaelaps, 44
protonymph, 11. 23 Rhinacarus Castro, 169
Protgnyssus, 118 Rhinacarus Nehring, 148
protosternum, 16 rhinitis, 3
protoxera, Ugandolaelaps, 195 Rhinixodes, 148
Protozoa, 3 Rninoecius, 157, 164, 262
psalidoprocnei, Ptilonyssus, 181 rhinolethrum, Rhinonyssus, 157,
Pseudolaelaps, 196 159, 261
Pseudomonas, 98 rhinolophi, Ornithonyssus, 88, 89
pseudoscorpions, 199 Rhinonyssidae, 2, 14, 25, 29, ’155
psi. Spinturnix, 189 Rhinonyssoides, 156, 172, 174"
Psilognathus, 94 Rhinonyssus, 22, 156, 157, 173,
Ptilonyssoides, 156, 161, 169 174
Ptilonyssus, 21, 22, 156, 174, Rhinophaga, 148. 153
178, 262 rhodesiensis, Haemolaelaps, 44 ;
328

rickettsia, 3, 73 Sejus, 69
rickettsialpox, 4, 84 semidesertus, Haemolaelaps, 45
rileyi, Entonyssus, 145 semilunaris, Spinturnix, 189
robustipes. Ichoronyssus, 89, 90., semitectus, Laelaps, 69
92, 94, 95 serdjukovae. Haemogamasus, 136
robustipes, Laelaps, 68 serini, Paraneonyssus, 178
Rochanyssus, 156, 171 .,,;,;,:.: Serpehticola, 9T_
rodhaini, Pneumonyssus, 153 serpentium, Ophionyssus, 97
rose-innesi, Ornithonyssus, 88 serraoi, Neonyssus, 163
rosmari, Orthohalarachne, 150 setae. j_7
rossicus, Myonyssus, 125 !
setiger, Laelaps,; 67^
rotenoi?e, 5 sh’ib’atai, Myonyssus, 125
rothschildi, Mysolaelaps. 76 sicula; Laelaps, 69
rotundus, Eubrachylaelaps, .SO signiodoni, Haemolaelaps, 38
roubaudi, Laelaps, 6Q_ . simicola, Pneumonyssus, 151

.
ruandae, Ptilonyssus, 182 similis, Pellonyssus, 117
Ruandanyssus. 157, 165 simillimus, Laelaps. 69
sinusitis, 3
sairae, Ptilonyssus, 182 sminthopsis, Laelaps, 69
salivary glands, j_9 ; snake mites, 6, 9
salivary stylets, 16, 19 .’..’ Sommatericola, 157
sangsteri, Hae molaelaps, ;.’4’4^ soricis. Hirstionyssus, Turk,
sanguineus, Allodermanyssus. 4, 1945.. 107 ’^i^ ’:

.
5, .10, ,12, 14, 118, 119 soricis, Hirstionyssus, Zemska, ;
sanguineus, Echinolaelaps, 72, 75 : ;"-. ’,.’.’.
1955. 103

.
sanguineus, Ornithonyssus, 83,. 84 soricis, Laelaps. 69 .i-,:.i
sanguisugus, Echinolaelaps, 72’,’ 75 souzai, Rhinonyssoides, 173 ,
santos-diasi, Hirstionyssus, 107 "spaltorgane, " 21
santos-diasi, Pneumonyssus, 153 spatulatum’,’ Sternostoma, 168
saurarum, Sauronyssus, 96 spatuliformis. Haemolaelaps, ^5
Sauronyssus, 78, 80, £H> special senses, 20^
scalopi, Haemolaelaps,. 38 spegazzinii, Haemolaelaps, 45
scapularis, Haemolaelaps, 44 Spelaeprhynchidae, 2. 28, 141
Shizogynildae, 196. ; . Spelaeorhynchus, 142-
schoutedeni, Hainertonia, 147 spencei, Geneiaddlaelaps, 196
schoutedeni, Neonyssus,. "163 sperm, 22
schoutedeni, Pn^umonysBus, 153 spermatodactyl, 26
Scissuralaelaps, 196 ’i . ... : .
spermatophore, 1’3, 22
sciureus, Haemolaelaps, .45 .. ;-^ , spinifer, Laelaps, 70^
sciurinus, Hirstionyssus,, :_107 spinlger, Ischyropoda, 137
sciuropteri, Haemogamasus, 130 spiniger, Laelaps. 1^5.:
scotophili, Spinturnix, 189 spinitarsus, Haemolaelaps, ^5
scotornis, Vitznyssus, 173, 174, Spinolaelaps, 80. 95
175 ;.; spinosulus, Haemolaelaps, 45
sculpturatus, Echinolaelaps, 75. spinosus, Eubrachylaelaps, 50
scutatus, De.rimanyssus, 123 spinosus, Ichoronyssus, 94
scutatus, Ichoronyssus, 89, 90 spinosus.. Myonyssoides, 195
segmentation,. 15 spinosus..’ Steatonyssus, 113, 115
 329

Spinturnicidae, 2, 16, 29, 182, tectum, 15

.
195, 267 tenuiscutatus, Ornithonyssus, 84
Spinturnix. 183. 190, 200, 262 terpsiphonei, Ptilonyssus, 182
Spirochaeta, 84 terpsiphonei, Ruandanyssus, 165
spirochaetosis, 4 testis, 2ji
spreo, Haemolaelaps, ^46 Tetragonyssus, 65, 6,6, 67, 124
squamosus, Rhinonyssoides, 173 texensis, Ichoronyssus, 89, 94
stabularis, Eulaelaps. 126,. 127, thalacdmys, Mesolaelaps, 55_
128, 200. 261 thamnomys, Laelaps, 70
staffordi, Hirstionyssus, 7.7, 107 thienponti, Sternostoma, 168
stammeri, Pneumonyssus, 153 thompsoni. Laelaps, 70_
Steatonyssus, 77, 78, 113,.. 2 62 thori, Laelaps, T0_
stegemani, Haemolaelaps, ,38 tinae, Orni1,honyssus, 88
Steptolaelaps. 31, .56. 72 Tinaminyssus, 157, J61, 170
sternalis, Haemogamasus., 130 Toxaphene, 6
sternalis, Haemolaelaps. jl6 trachea, 21_ .

sternalis, Hirstesia, 110, 111 tracheacolum. Sternostoma, 6,

sternal plate, 24, 26 168
Sterno.taelaps, 13^6 Trachytoidea. 1
Sternostoma. 19, 22. 155, 165 transiliensis, Hirstionyssus, 108
Sternostomum, 157 transovarial transmission, 4
stigma, 21. 22 transvaalensis, Hirstesia, 1.11 .

stigmata, 1 transvaalensis, Laelaps, T0_ .

strandtmanni, Haemolaelaps, 38 transversus, Spinturnix, 188

strandtmanni, Periglischrus. 191 trappi, Tinaminyssus, 161 . .. .

strandtmanni, Rhinonyssoides, 1.73 traubi, Haemolaelaps, 47 ; ’.

strandtmanni, Sternostoma, 168 Travanyssus, 156. 171
stresemanni, Ptilonyssus, 182 travassosfilhoi, Paraneonyssus,
sturnicola, Sternostoma, 168 178
subterraneus, Haemolaelaps, 46 treronis, Neonyssus, 163
sudanensis, Steatonyssus, 116 triacanthus, Hirstionyssus, 108
sudanicus, Haemolaelaps, 46 triangulus, Hemilaelaps, 140
sulfur, control with, 6 tricholabiatus, Bolivilaelaps, 48_
sulphenone, control with, 6 Tricholaelaps. 32, _55, 70
superans, Steatonyssus, 116 Trichosurolaelaps, 30. 56
sylstrai., Aetholaelaps, 76 Trigynaspida, 2, 27, 192~
sylvaticus, Haemogamasus, 136 triscutatus, Ptilonyssoides, 169,
sylviarum, Ornithonyssus, 3, 5, 170
6, 9, 10, 11, 12, 78, 80, tritonymph, 23
81, 8J?, 198, 261, 262 tritosternum, 16
trouessarti, Rhinonyssoides, 172
tachyoryctes, Haemolaelaps, 46 trypanosomes, 4, 100, 122
talpae, Hirstionyssus, 100 tularemia, 39
tanneri, Hemilaelaps, 140 tularensis, Pasteurella, 4
taprobanius, Laelaps, 70 Tur, 29, 118
taterae, Haemolaelaps, 47 turdi, Sternostoma, 169
tateronis, Haemolaelaps, 47 turkestanicus, Laelaps, 70
technaui, Sternostoma, 168 twitchelli, Haemogamasus, 130
330

Tympanospinctus, 195 virginxanus, Haernolaelaps, 38

typhus group rickettsia, 73 viruses. 3 . .

typhus; muririe, 4. 67. 84 VUznyssus, 156, ^73

tytohis, Rhinoecius, 165 vitzthumi, Echinolaelaps, 73
vitzthumi, Entonyssus, 145
ugandae, Mungosicola, 195 vitzthumi, Eulaelaps, 129
ugandanus, Echinolaelaps, 75 vitzthurni, Gigantolaelaps, 50

.
Ugandolaelaps, 1.95 :; v:ttzthumji., Ornithonyssus, 89

.
.
uncinatus, Liponyssus, 89. viizthuBiii,;, Vitanyssus, 175, 176.
uniscutatus, Tur, 118 volgensis, Laelaps, 71

.
Uroppdina, 1’.’. ’:

’
:.
,1. .

Uropodoidea, 1. .;: .’ , ,
walkerae. Spintumix, 189
utahensis, Brevisterna, 9, 10, 136 waterstoni, Rhinonyssus, 159
uterus, 23 wernecki, Neoichoronyssus. 108

’
.

.
.
werneri, Rochanyssus, 171
validipes, Echinonyssus, 1,10 wetmorei, ,;L.aelap£i. 71

.
vansoBiereni, Echinolaelaps,,’7.5 wolffsohni. Gigantolaelaps, 52
vargasi,;Periglischrus, 191 .

variabilis, Ophionyssus, 98 yaoundensis, Echinolaelaps, 75

vasa deterentia, 22 . ,", .

venezblanus, Ichoronyssus, .94, -95 zachyatklHi, Haemogaraasus, 136

ventralpla.te.-24 ,. :.. zaiophi, Orthohalarachne, 151
.
versteegi, GigaAtolaelaps, 52 zeiebori, Spintumix, 18,3, 189
vespertUioniS) Spintumix, 183 :. zenaidurae, Neonyssus, 164
vespertilionis,. Steatonyssus, 11.3 Zerconoidea, 1. - . .

.
viator, Pellonyssus, 117 :. . zulu, .Haernolaelaps, 47
.
.
viduae. PtUonyssus, 182 zumpti, Andreacarus, .7,5
viduus, Spintumix, 189. zurnpti. Laelaps,, ’n_.
.

villosissimus. Haernolaelaps, 47 zumpti.. Paraneonyssus, .178

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FIGURE 41. Ophionyssus natricis. female: _A, ventral and dorsal views ;
B, chelicera; C, tritosternum. (’From Camin, 1953.)

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FIGURE 77. Rallinyssus caudistigmus: 1_5, female tarsus I; 16, fe-
male chelae; 1T_. female dorsnna; 18. female venter; IJJ, male venter;
20, male chelae; 31, female gnathosoma. (From Strandtmann, 194B.>

-55-
FIGURE 78. Meonyssus genaidurae. ^, female chela; 2, male chela;
3, tarsus I of female; 4. female genital, plate; E^, female dorsum; Q.
male ventral plate; 7, male anal plate; Q, female venter; j). per-itreme
and associated plate; 10, male venter; 11^ female gnathosoma, dorsal
view; ^_2, female palpal tarsus; 13, female gnathoso.ma, ventral view,
(From Crossley, 1952.)

-56-
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 FIGURE 83. Tinamj.nyssus trappi, female: dorsum FIGURE 84. Flavionyssus rabelloi. female; dorsum
and chela.(After Peretra and Castro, 1949.) and chela. (After Castro, 1948.)

^URE 86. Travanyssus paranensis: female dorsum. FIGURE 85. Rochanyssus werneri: female dorsum.
ter Castro, 1948.1 (After Castro, 1948.)
-61-
FIGURE 88. yj.tznyssus sp., female: 1^ venter; 2, dorsum; ^, chela;
4, gnathosoma and tectum; 5, stigma and peritreme. (Original,)

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-69-
 CONTRIBUTIONS OF THE INSTITUTE OF ACAROLOGY

Conrad E. Yunker. Editor

Price

No. 1 Arachnida. Vol. IV, No. 2. Fauna of the U.S. S. R.

Ixodid Ticks (Ixodidae). by B. I. Pomerantzev. (A
translation from the Russian,) Edited by G. Anastos ;
translated by A. Elbl. April. 1957. Second print-
ing, 1959. $10.OO^
No. 2 The Ticks orIxodidesoftheU.S.S.R., by G. Anastos.
Public Health Service Publication No. 548. August,
1957, 397 pp. .
$ 2.25*

No. 3 Guide to the Families of Mites, by E. W. Baker,

J. H. Camin, F. Cunliffe, T. A. Woolley and C. E.
Yunker. June. 1958, 242 pp. $ 5.00**

No. 4 A Manual of Mesostigmatid Mites Parasitic on Ver-

tebrates, by R. W. Strandtmann and G. W, Wharton.
December, 1958, pp. 330 + xi + 96 plate-figures. $ 7.50**

Contribution No. 1, second printing, will be available, early 1959,

from: American Institute of Biological Sciences, 2000 P Street, N. W.,
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Contribution No. 2 is available from: Superintendent of Documents,

U. S. Government Printing Office, Washington 25, D. C. Checks
or money orders should be made payable to "Superintendent of Docu-
ments. "

Contributions No. 3 and No. 4 are available from: The Institute of

Acarology, Department of Zoology, University of Maryland, College
Park, Maryland. Orders accompanied by remittances will be sent
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