Gerecke (Ed.), 2006 - Süßwasserfauna Von Mitteleuropa, Vol. 7, 2-1 Chelicerata - Araneae, Acari I

Süßwasserfauna von Mitteleuropa
Reinhard Gerecke Editor

Ilse Bartsch · Kees Davids · Ralf Deichsel
Antonio Di Sabatino · Grzegorz Gabryś · Reinhard Gerecke
Terence Gledhill · Peter Jäger · Joanna Makol · Harry Smit
Henk van der Hammen · Gerd Weigmann
Andreas Wohltmann · Eberhard Wurst
Süßwasserfauna
von Mitteleuropa,
Vol. 7/2-1 Chelicerata:
Araneae/Acari I
Gerecke (ed.)
Chelicerata: Araneae, Acari I
Begründet von A. Brauer
Herausgegeben von J. Schwoerbel (†) und P. Zwick
Stellvertretender Herausgeber für Band 7: R. Gerecke
Band 7/2-1
To the memory of Jürgen Schwoerbel (1930–2002) who took the initiative for the new
edition of the Brauer series and coordinated the first steps of the the production of this
volume.
Ilse Bartsch Kees Davids Ralf Deichsel Antonio Di Sabatino Grzegorz Gabryś
Reinhard Gerecke Terence Gledhill Peter Jäger Joanna Mąkol Harry Smit
Henk van der Hammen Gerd Weigmann Andreas Wohltmann Eberhard Wurst
Chelicerata: Araneae, Acari I

Süßwasserfauna von Mitteleuropa 7/2-1
Ilse Bartsch Peter Jäger
Frankfurt am Main, Deutschland Frankfurt, Deutschland
Kees Davids Joanna Makol

Amsterdam, Niederlande Wroclaw, Polen
Ralf Deichsel Harry Smit

Berlin, Deutschland Amsterdam, Niederlande
Antonio Di Sabatino Henk van der Hammen

Coppito L’Aquila, Italien Amsterdam, Niederlande
Grzegorz Gabryś Gerd Weigmann

Zielona Góra, Polen Berlin, Deutschland
Reinhard Gerecke Andreas Wohltmann

Tübingen, Deutschland Bremen, Deutschland
Terence Gledhill Eberhard Wurst

Kendal, Großbritannien Stuttgart, Deutschland

ISBN 978-3-662-55957-4 ISBN 978-3-662-55958-1 (eBook)
https://doi.org/10.1007/978-3-662-55958-1
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Die Anschrift der Gesellschaft ist: Heidelberger Platz 3, 14197 Berlin, Germany
1. Order Araneae
By Peter Jäger
2. Order Acari – General Introduction and Key to major groups

By Reinhard Gerecke, Gerd Weigmann, Andreas Wohltmann and Eberhard Wurst
3. Acari: Limnic Acaridida (Astigmata)

By Eberhard Wurst
4. Acari: Limnic Oribatida

By Gerd Weigmann and Ralph Deichsel
5. Acari: Halacaridae
By Ilse Bartsch
6. Acari: Terrestrial Parasitengona inhabiting transient biotopes

By Andreas Wohltmann, Grzegorz Gabryś and Joanna Ma˛kol
7. Acari: Hydrachnidia I
By Kees Davids (†), Antonio Di Sabatino, Reinhard Gerecke, Terence Gledhill,
Harry Smit and Henk van der Hammen (†)
Verzeichnis der Autoren
Dr. Ilse Bartsch Dr. Peter Jäger

Forschungsinstitut Senckenberg, c/o DESY Forschungsinstitut Senckenberg
Notkestraße 85 Senckenberganlage 25
D- 22607 Hamburg, Germany D- 60325 Frankfurt am Main, Germany
email: bartsch@meeresforschung.de email: peter.jaeger@senckenberg.de
Dr. Kees Davids (†) Dr. Joanna Ma˛kol
Vakgroep Aquatische Oecologie Department of Zoology and Ecology
Universiteit van Amsterdam Agricultural University of Wrocław
Kruislaan 320 Kozuchowska 5b
NL- 1098 SM Amsterdam, The Netherlands PL- 51-631 Wrocław, Poland
email: makol@ozi.ar.wroc.pl
Ralf Deichsel
Laboratory for Soil Zoology & Ecology Drs Harry Smit
Grunewaldstraße 34 Zoological Museum, University of
D- 12165 Berlin, Germany Amsterdam
email: flar@zedat.fu-berlin.de Plantage Middenlaan 64
NL- 1018 DH Amsterdam, The Netherlands
Dr. Antonio Di Sabatino
email: smit.h@wolmail.nl
Dipartimento di Scienze Ambientali
Università dell’Aquila Dr. Henk van der Hammen (†)
Via Vetoio 20, Amsterdam, The Netherlands
I- 67100 L’Aquila, Italy
Prof. Dr. Gerd Weigmann
e-mail: adisab@univaq.it
Laboratory for Soil Zoology & Ecology
Prof. Dr. Grzegorz Gabryś Grunewaldstraße 34
Department of Biology D- 12165 Berlin, Germany
Institute of Biotechnology & Environmental email: weigmann@zedat.fu-berlin.de
Sciences
Dr. Andreas Wohltmann
University of Zielona Góra
Institut für Ökologie und Evolutionsbiologie
Monte Cassino 21B
Universität Bremen, Fachbereich 2
PL- 65-561 Zielona Góra, Poland
Postfach 33 04 40
e-mail: g.gabrys@ibos.uz.zgora.pl
D- 28334 Bremen, Germany
Dr. Reinhard Gerecke email: wohltman@uni-bremen.de
Biesingerstraße 11
Dr. Eberhard Wurst
D- 72070 Tübingen, Germany
Fachgebiet Parasitologie (220 B)
email: reinhard.gerecke@uni-tuebingen.de
(Institut für Zoologie)
CBiol MIBiol FLS Terence Gledhill Universität Hohenheim
Freshwater Biological Association Emil-Wolff-Straße 34
Windermere Laboratory D- 70599 Stuttgart, Germany
The Ferry House, Far Sawrey email: wurst@uni-hohenheim.de
GB Ambleside Cumbria, LA 22 OLP,
United Kingdom
email: allgledhills@hotmail.com
Contents
1. Order Araneae (Peter Jäger)

General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
General introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Key to spider families occurring on fresh water sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Family Argyronetidae T HORELL , 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Genus Argyroneta L ATREILLE , 1804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Glossary Araneae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2. Order Acarri – General Introduction and key to major groups

(Reinhard Gerecke, Gerd Weigmann, Andreas Wohltmann and Eberhard Wurst)
Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Morphology and terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Sampling and sample processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Collecting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Treatment of samples . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Classification of undissected material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Rearing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Preservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Dissection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Mounting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
The major groups of mites found in aquatic habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Key to life-cycle stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Key to larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Key to nymphs and adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
General Glossary Acari . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3. Acari: Limnic Acaridida (Astigmata) (Eberhard Wurst)

General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Chaetotaxy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Ontogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Special Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Key to the adults of limnic Acaridida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Family Acaridae L ATREILLE , 1802 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Genus Schwiebea O UDEMANS , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Genus Naiadacarus FASHING , 1974 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Histiostomatidae B ERLESE , 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
VIII Contents
4. Acari: Limnic Oribatida (Gerd Weigmann and Ralf Deichsel)

General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Special Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Key to families and genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Family Mucronothridae K UNST, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Genus Mucronothrus T RÄGÅRDH , 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Family Trhypochthoniellidae K NÜLLE , 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Genus Trhypochthoniellus W ILLMANN , 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Family Malaconothridae B ERLESE , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Genus Malaconothrus B ERLESE , 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Genus Trimalaconothrus B ERLESE , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Family Hydrozetidae G RANDJEAN , 1954 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Genus Hydrozetes B ERLESE , 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Family Limnozetidae G RANDJEAN , 1954 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Genus Limnozetes H ULL , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Family Mycobatidae G RANDJEAN , 1954 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Genus Punctoribates B ERLESE , 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Family Zetomimidae S HALDYBINA , 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Genus Heterozetes W ILLMANN , 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Genus Zetomimus H ULL , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
5. Acari: Halacaridae (Ilse Bartsch)

General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
External morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Juvenile stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Anatomy and ultrastructure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Relations to the environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Feeding and food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Enemies, epibionts and competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Reproduction and development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Origin and phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Family Halacaridae M URRAY, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Key to genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Subfamily Copidognathinae B ARTSCH , 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Genus Copidognathus T ROUESSART, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Subfamily Halacarinae K.V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Genus Caspihalacarus K.V IETS , 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Genus Halacarellus K.V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
Genus Lobohalacarus K.V IETS , 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Genus Porohalacarus T HOR , 1922 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Genus Troglohalacarus K.V IETS , 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Subfamily Limnohalacarinae K.V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Genus Limnohalacarus WALTER , 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Genus Parasoldanellonyx K.V IETS , 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Genus Soldanellonyx WALTER , 1917 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Genus Stygohalacarus V IETS , 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Contents IX
Subfamily Lohmannellinae V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

Genus Lohmannella T ROUESSART, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Subfamily Porolohmannellinae K.V IETS , 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Genus Porolohmannella K.V IETS , 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Subfamily Rhombognathinae K.V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Genus Isobactrus N EWELL , 1947 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Subfamily Ropohalacarinae B ARTSCH , 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Genus Ropohalacarus B ARTSCH , 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
6. Acari: Terrestrial Parasitengona inhabiting transient biotopes

(Andreas Wohltmann, Grzegorz Gabryś and Joanna Ma˛kol)
General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Biology and life history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Habitat specificity and adaptive characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
Phylogeny and taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Special Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Key to major subgroups of Parasitengona . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Superfamily Calyptostomatoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Family Calyptostomatidae O UDEMANS , 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Genus Calyptostoma C AMBRIDGE , 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Superfamily Erythraeoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Key to families, subfamilies and genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
Family Erythraeidae R OBINEAU -D ESVOIDY, 1828 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Genus Abrolophus B ERLESE , 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Genus Charletonia O UDEMANS , 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Genus Erythraeus L ATREILLE , 1806 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Genus Leptus L ATREILLE , 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Superfamily Trombidioidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Key to Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Family Johnstonianidae T HOR , 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Genus Centrotrombidium K RAMER , 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Genus Diplothrombium B ERLESE , 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Genus Johnstoniania G EORGE , 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Family Microtrombidiidae T HOR , 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
Genus Atractothrombium F EIDER , 1952 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
Genus Camerotrombidium T HOR , 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Genus Campylothrombium K RAUSSE , 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Genus Echinothrombium W OMERSLEY, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . 208
Genus Enemothrombium B ERLESE , 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Genus Microtrombidium H ALLER , 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Genus Valgothrombium W ILLMANN , 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Family Tanaupodidae T HOR , 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
Genus Lassenia N EWELL, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
Genus Tanaupodus H ALLER, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Family Trombidiidae L EACH , 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Genus Allothrombium B ERLESE , 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
Genus Paratrombium B RUYANT, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Genus Podothrombium B ERLESE , 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
Genus Trombidium FABRICIUS , 1775 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
X Contents
7. Acari: Hydrachnidia I
(Kees Davids [†], Antonio Di Sabatino, Reinhard Gerecke, Terence Gledhill,
Harry Smit and Henk van der Hammen [†])
General part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
External morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
Food and feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
Sperm transfer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259
Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
Larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
Deutonymphs and adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266
Life-histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Life span . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270
Adaptations and Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Deformities, diseases and epibionts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Standing waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Running waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Springs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
Interstitial waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Temporary pools and semi-aquatic habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Rare and threatened species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Special Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
Special part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Area treated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Key to superfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Superfamily Stygothrombioidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Family Stygothrombiidae (T HOR , 1935) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291
Genus Stygothrombium K. V IETS , 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291
Superfamily Hydrovolzioidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294
Key to families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
Family Acherontacaridae C OOK , 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
Genus Acherontacarus K. V IETS , 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
Family Hydrovolziidae T HOR , 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
Genus Hydrovolzia M ONTI , 1905 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
Superfamily Eylaoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Key to families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Family Limnocharidae G RUBE , 1859 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
Genus Limnochares L ATREILLE , 1796 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
Family Piersigiidae O UDEMANS , 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
Genus Piersigia P ROTZ 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Family Eylaidae L EACH , 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
Genus Eylais L ATREILLE, 1796 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
Superfamily Hydrachnoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
Family Hydrachnidae L EACH , 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
Genus Hydrachna M ÜLLER , 1776 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
Index Araneae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Index Acari . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
1. Order Araneae 1
1. Order Araneae
Peter Jäger
General part
Diagnosis
Small to large in size (body length 0.4100 mm); body divided into two parts, these called
prosoma and opisthosoma, these connected by a thin petiolus prosoma including segments
I–VI with chelicerae, palps, and four pairs of walking legs, with dorsal shield and ventral
sternum; chelicerae two-segmented with basal limb large, with hairs and usually teeth, the
secondary limb as heavily sclerotised fang with opening of a poison gland on its tip; palps
with six, legs with seven segments (additional metatarsus between tarsus and tibia); palpal
coxae used as mouthparts together with the unpaired labium; opisthosoma consisting of
the fused body segments VIIXVII, only in Mesothelae slightly segmented; ventral opistho-
soma with 2–4 openings for book-lungs, a genital opening and a tracheal spiracle; plesio-
typically two pair of spinnerets on each of the segments X and XI, these shifted in all groups
but Mesothelae to the end of the opisthosoma; number of spinnerets reduced usually to six,
in some taxa to four or two.
Sexual reproduction. Sperm transfer indirect by using the modified tarsi of male pedipalps
as transfer organ (gonopod); postembryonal development with 5–10 stages, all of which
with complete number of appendages and moulting inbetween.
General introduction
True spiders (Arachnida: Araneae) are distributed roughly worldwide and live in almost all
kinds of terrestrial habitats from the sea shore to mountain ranges up to altitudes of nearly
7000 metres. Only one of 38000 currently known species the water spider Argyroneta
aquatica (C LERCK , 1757) is secondarily adapted to aquatic life in fresh water. Individuals
live, hunt, feed, moult and mate under water, by using a web as a diving bell filled with air.
All other species are not capable to perform their whole life cycle under water. Inspite, there
are several species from various families, which occur on fresh water sites. Among them are
forms whose ecological needs do not allow a life in places without a certain humidity or
even open water. Some wolf spiders for example (members of the family Lycosidae) hunt on
the water surface and survive floodings while hiding in their retreats between or under
stones. Others build their catching webs exclusively close to the water surface (e.g. Ther-
idiosoma gemmosum L. K OCH , 1877). Representatives of one family (Desidae) are hunting
in the tidal zones of the sea shore. They are able to rest during the high tide in their retreats
under water. Beside these specialised forms there are ubiquists, which may settle in every
kind of habitat (e.g. some Linyphiidae). Between these two extremes spiders developed all
sorts of partitioning.
Morphological features of the spider body are explained in Figure 1-1 c, e, g–h, j–k. For
the special terminology see in the glossary at the end of this chapter.

I. Bartsch et al., Süßwasserfauna von Mitteleuropa, Vol. 7/2-1
Chelicerata: Araneae/Acari I, Süßwasserfauna von Mitteleuropa,
https://doi.org/10.1007/978-3-662-55958-1_1
2 Special part
Special part
Area treated
The following key includes all spider families which are likely to be found in the Palearctic
directly on the water surface, in vegetation standing in the water or in the vegetation belt
close to the shore line. The present key is not only valid for Central Europe, but may also be
of help in most of the eastern parts of the Palearctic region. However, some tropical and
subtropical elements remain excluded, especially species occurring in China and Japan. For
information on taxonomy and biology of species representing further genera or families
which in Central Europe may appear in such habitats by accident, see N ENTWIG et al.
(2003) and for information on the distribution ranges within Germany see S TAUDT
(2005). Only short information is given on ten of the eleven included families, including
a list of species occuring at fresh water habitats (for compilations see B ELLMANN 2001,
N ENTWIG et al. 2003). Only Argyroneta aquatica is treated in a separate paragraph in de-
tail.
Key to spider families occurring on fresh water sites

1 Each tarsus with two distal claws (1-1 d), sometimes hidden by claw tufts of adhesive hairs
(1-1 g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Each tarsus with three distal claws, median claw in several species tiny and hardly recognizable
(1-1 j, 1-2 c, l, 1-3 m, q, , 1-4 e–f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Anterior median eyes very large (1-1 a), eyes in three rows (4 - 2 - 2), posterior eyes forming a
rectangle (1-1 b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salticidae
Small to medium-sized spiders with stout legs; hunting in vegetation and on the ground, building
retreats between leaves, behind bark and under stones. E.g. Heliophanus patagiatus T HORELL ,
1875, Marpissa radiata (G RUBE , 1859), Mendoza canestrinii (N INNI , 1868), Sitticus caricis
(W ESTRING , 1861), S. distinguendus (S IMON , 1868), S. floricola (C.L. K OCH , 1837).
– Eyes subequal and in two rows (4 - 4) (1-1 c, e–f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Legs laterigrade, i.e. especially legs I + II tilted to their retrolateral side and being longer and in
most species distinctly stronger than legs III + IV, tibia and metatarsus I + II with double rows of
short and stout spines (1-1 c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thomisidae
Medium-sized spiders with a crab-like habit; hunting in vegetation and on the ground; females
building only retreats for their egg-sacs. E.g. Heriaeus graminicola (D OLESCHALL , 1852) (1-1 c),
Xysticus ulmi (H AHN , 1831).
– Legs prograde, not tilted to their retrolateral side, tibia and metatarsus I + II only with single spines
(1-1 e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clubionidae
Medium-sized spiders with a velvet-like appearance; hunting in vegetation and on the ground;
building retreats between leaves and behind bark, some species building a special kind of retreat
by folding a lanceolate grass leave twice. E.g. Clubiona lutescens W ESTRING , 1851; C. phrag-
mitis C.L. K OCH , 1843; C. reclusa O.P.-C AMBRIDGE , 1863; C. stagnatilis K ULCZYNSKI , 1897.
4 Lateral eyes separated by more than two diameters of an anterior lateral eye, posterior eye row
strongly recurved, forming a trapezoid (1-1 h–j, k–l). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Lateral eyes touching each other or separated by about one diameter of the anterior lateral eye,
posterior eye row only slightly recurved, straight or procurved (1-2 a–b, g, n–o, 1-3 d, h–i, p, u–t,
1-3 a–b, h–j, n–p). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 Posterior median eyes enlarged, clypeus short, i.e. about one diameter of anterior eyes in frontal
view (1-1h), anterior eyes and in many cases posterior median eyes reaching the frontal margin of
the dorsal shield of prosoma in dorsal view (1-1 i). . . . . . . . . . . . . . . . . . . . . . . Lycosidae
Medium-sized to large spiders; hunting on the ground and on the water surface; females carry-
ing their egg-sac attached to the spinnerets and later, the offspring on their opisthosoma; some
species being able to dive (Pirata spp.) or to stay in their retreat under water during a period of
flooding (e.g. Arctosa cinerea (FABRICIUS , 1777) in river gravels, or Pardosa purbeckensis
F.O.P.-C AMBRIDGE , 1895 in salt water habitats on coast lines). E.g. Arctosa cinerea (F ABRICIUS ,
1. Order Araneae 3
1777), A. leopardus (S UNDEVALL, 1833), A. maculata (H AHN , 1822), A. stigmosa (T HORELL ,

1875), Pardosa agricola (T HORELL , 1856), P. saturatior S IMON , 1937, P. wagleri (H AHN ,
1822), Pirata hygrophilus T HORELL 1872, P. insularis E MERTON , 1885, P. knorri (S COPOLI ,
1763), P. latitans (B LACKWALL , 1841), P. piraticus (C LERCK , 1757), P. piscatorius (C LERCK ,
1757), P. tenuitarsis S IMON , 1876.
– Posterior median eyes not enlarged, clypeus wide, i.e. about three diameters of the anterior eyes in
frontal view (1-1 k), anterior eyes and posterior eyes never reaching frontal margin of the dorsal
shield of prosoma in dorsal view (1-1 l). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pisauridae
Large spiders; juveniles of Dolomedes spp. hunting in vegetation of swampy areas like meadows
around springs; adults of Dolomedes sitting at the shore line and holding the fore-legs on the
water surface using the latter as a ‘fluid spider web’ in means of getting information of move-
ments of potential prey; hunting on the water surface or even under water; Pisaura mirabilis
(C LERCK , 1757) hunting in all stages in vegetation; females of all species carrying their egg-sac in
the chelicerae. Dolomedes fimbriatus (C LERCK , 1757) , D. plantarius (C LERCK , 1757); ubiquist:
Pisaura mirabilis (C LERCK , 1757).
6 Tarsi with two rows of trichobothria (1–2 c), leg IIIIV with long and dense hairs (1–2 d), spina-
tion of legs differing between forelegs and hindlegs, i.e. leg I+II almost without spines, leg III+IV
with many spines (1-2 e–f). . . . . . . Argyronetidae (Argyroneta aquatica), see next paragraph.
– Tarsi without trichobothria (1-3 m, q, 1-4 e–f), all legs with similar length of hairs and spination
(1-2 j–k, 1-3 k–l). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Gnathocoxae elongated (2–3 times as long as wide), proximal part laterally bent (1-2 p: arrow),
i.e. lateral outline concave (1-2 h: arrow, 1-3 a,c), male paracymbium elongated, as half as cym-
bium length (1-2 m, q) or complex (1-3 b, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Gnathocoxae not elongated (1–2 times as long as wide), proximal part directed to median line
(1-3 g, n, 1-4 c: arrow) or sub-rectangular (1-4 k), male paracymbium smaller (1-3 r, v–w,
1-4 l, q) or absent (1-4 k) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Chelicerae elongated and diverging (1-2 g, n), basal femora dorsally with one or more rows of
trichobothria (1-2 i), male paracymbium elongated with one knob-like protrusion (1-2 m, q: ar-
rows) and parallel to cymbium length axis, females without sclerotized epigyne, i.e. no folds or
copulatory opening visible (1-2 r). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnathidae
Medium-sized spiders; orb-weavers (except for hunting adults of Pachygnatha spp.); especially
Tetragnatha spp. abundant in vegetation close to water sites or in reed-belts; sitting in a char-
acteristic posture, stretching the legs forewards and backwards, imitating a stalk of vegetation.
E.g. Pachygnatha clercki S UNDEVALL , 1823; Tetragnatha dearmata T HORELL , 1873; T. extensa
(L INNAEUS , 1758); T. montana S IMON , 1874; T. nigrita L ENDL , 1886; T. reimoseri (R OSCA ,
1939); T. striata L. K OCH , 1862.
– Chelicerae neither elongated nor diverging, femora without trichobothria, male paracymbium
complex and transversal to cymbium length axis (1-3 b, e), females with (partly sclerotized) epi-
gyne, i.e. folds and copulatory opening visible (1-3 f). . . . . . . . . . Metinae (Tetragnathidae)
Medium-sized to large spiders; spinning orb-webs with open hub; living in humid places, some-
times close to streams (Metellina spp.), or in caves and similar habitats (Meta spp.). E.g. Meta
bourneti S IMON , 1922; M. menardi (L ATREILLE , 1804); Metellina merianae (S COPOLI , 1763).
9 Sternum with two pits close to labium (openings of sternal glands) (1-3 g), small spiders (1.5–3.0
mm body length) with spheroidal opisthosoma (1-3 j), this with silver-shimmery pattern. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Theridiosomatidae
Tiny to small spiders; spinning a small 3-dimensional orbweb directly above the water surface of
stagnant water sites. Theridiosoma gemmosum L. K OCH , 1877 (1-3 g–m).
– Sternum without pits (1-3 n), small to medium-sized spiders. . . . . . . . . . . . . . . . . . . . . . 10
10 Chelicerae laterally with stridulatory files (in some species weak) (1-3 o: arrow), male para-
cymbium sickle shaped (1-3 r, v–w), paracymbium separated from cymbium by a membraneous
part (1-3 w: arrow), males of many smaller species with cephalic lobes (1-3 o) and tibial apophysis
(1-3 r: arrow) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linyphiidae
Tiny to medium-sized spiders; spinning sheet-webs in the vegetation or on the ground; several
species of this largest Palearctic spider family are pioneer-settlers (Erigone spp.), various stages
being able to balloon and thus, to land on or close to open water surfaces. E.g. Hypomma
4 Special part
bituberculatum (W IDER , 1834), Kaestneria dorsalis (W IDER , 1834), Microlinyphia impigra

(O.P.-C AMBRIDGE , 1871), M. pusilla (S UNDEVALL , 1830), ubiquists: Erigone atra B LACKWALL ,
1833, E. dentipalpis (W IDER , 1834), Linyphia triangularis (C LERCK , 1757).
– Chelicerae laterally without stridulatory files, male paracymbium absent (1-4 g) or knob-like
(1-4 l, q: arrows). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Anterior eyes reaching in most species not the frontal margin of the dorsal shield of prosoma in a
dorsal view (1-4 a), clypeus mostly wide (1-4 b), males without a paracymbium, but with long
bristles covering the bulbus (1-4 g), females mostly with simple epigynal pit without scapus (ex-
ception: few species with craniad scapus), epigynal structures mostly separated from epigastric
furrow (exception: 1-4 d), ventral tarsus IV with serrated bristles, these in a higher density
than dorsal hairs (1-4 e, exception: males of some species, 1-4 f). . . . . . . . . . . . Theridiidae
Small to medium-sized spiders; spinning 3-dimensional webs in the vegetation, some of them
with gum-foots; prey being wrapped with sticky silk pulled out of the spinnerets with the ser-
rated bristles of the tarsi of the fourth pair of legs. E.g. Crustulina sticta (O.P.-C AMBRIDGE ,
1861), Enoplognatha caricis (F ICKERT, 1876), Robertus arundineti (O.P.-C AMBRIDGE ,
1871), Rugathodes instabilis (O.P.-C AMBRIDGE , 1871), Theonoe minutissima (O.P.-C AM-
BRIDGE , 1879), Theridion hemerobium S IMON , 1914.
– Anterior eyes reaching frontal margin of the dorsal shield of prosoma in dorsal view, eyes in most
species integrated in a characteristic silhouette (1-4 h, n), clypeus mostly short (1-4 j, o–p, excep-
tion: males of few species, 1-4 i), male paracymbium knob-like (1-4 l, q: arrows), females with
caudad scapus (1-4 r) or epigyne (here: copulatory openings) connected with epigastric furrow by
folds (1-4 m), ventral tarsus IV without serrated bristles. . . . . . . . . . . . . . . . . . . Araneidae
Medium-sized to large spiders; spinning orb-webs in vegetation along the shore line or in reed-
belts. E.g. Hypsosinga heri (H AHN , 1831), Larinia bonneti S PASSKY, 1939, L. elegans S PASSKY,
1939, Larinioides cornutus (C LERCK , 1757), L. ixobolus (T HORELL , 1873), L. sclopetarius
(C LERCK , 1757), Singa nitidula C.L. K OCH , 1844.
Family Argyronetidae T HORELL , 1870

The family contains 12 genera with 149 species (P LATNICK 2004). It was formerly placed as
subfamily in the Agelenidae C.L. K OCH , 1837. L EHTINEN (1967) considered it a subfamily
of Dictynidae O.P.-C AMBRIDGE 1871, F ORSTER (1970) elevated it to family rank. S ELDEN
(2002) divides Argyronetidae (sub Cybaeidae) into two subgroups, the Argyronetinae
T HORELL , 1870 and ‘other cybaeids’ (= Cybaeinae B ANKS , 1892). Together with Argyro-
neta aquatica, he placed the extinct Vectaraneus yulei S ELDEN , 2001 (upper Eocene) and an
extinct unnamed Argyroneta species (Miocene) in Argyronetinae and considered all three
taxa derived members of this family.
N.B.: Argyronetidae and thus Argyronetinae are the valid names for the family and sub-
family (see International Code of Zoological Nomenclature [ICZN 1999: Article 23.3], i.e.
Argyronetidae is not a nomen oblitum as indirectly indicated in P LATNICK 2004).
Genus Argyroneta L ATREILLE , 1804

1804 Argyroneta L ATREILLE , Histoire naturelle générale et particulière des Crustacés et des
Insectes. Paris. 7: 134.
Four species were originally described in this genus: A. antiqua von H EYDEN , 1859, A.
aquatica, A. longipes H EER , 1865 and Argyroneta palustris R ISSO , 1826. The extinct
A. antiqua and A. longipes were regarded by S ELDEN (2002) as species incertae sedis within
araneomorph spiders. The status of Argyroneta palustris R ISSO , 1826, described from
France, is not yet clear. In the subsequent paragraph only the extant water-dwelling species
Argyroneta aquatica is treated.
1. Order Araneae 5
Argyroneta aquatica (C LERCK , 1757) (1-2 a–f, 1-5 a–l)

1757 Araneus aquaticus C LERCK , Svenska spindlar, uti sina hufvud-slågter indelte samt under nå-
gra och sextio särskildte arter beskrefne och med illuminerade figurer uplyste. Stockholm:
143, pl. 6, fig. 8.
1758 Aranea aquatica L INNAEUS , Systema naturae per regna tria naturae, secundum classes, or-
dines, genera, species cum characteribus differentiis, synonymis, locis. Editio decima, refor-
mata. Stockholm: 623.
1761 Aranea urinatoria P ODA , Insecta Musei Graecensis, quae in ordines, genera et species juxta
systema naturae Caroli Linnaei. Graz: 123.
1775 Aranea aquatica FABRICIUS , Systema entomologiae, sistens insectorum classes, ordines, gen-
era, species, adiectis, synonymis, locis descriptionibus observationibus. Flensburg and Leip-
zig: 436.
1776 Aranea amphibia M ÜLLER , Zoologicae danicae prodromus, seu animalium daniae et norve-
giae indigenarum, characteres, nomina et synonyma imprimis popularium. Copenhagen:
194.
1804 Argyroneta aquatica L ATREILLE , Histoire naturelle générale et particulière des Crustacés et
des Insectes. Paris. 7: 134.
1837 Clubiona fallax WALCKENAER , Histoire naturelle des insectes. Aptères. Paris. 1: 603.
2002 Argyroneta aquatica japonica O NO , Bulletin of the National Science Museum, Series A
(Zoology), 28 (1): 53.
2002 Argyroneta aquatica aquatica O NO , Bulletin of the National Science Museum, Series A
(Zoology), 28 (1): 53.
For further citations see P LATNICK (2004).
This unique species is known to science since C AROLUS C LERCK (1757: pl 6, fig. 8) illu-
strated a male in his book on Swedish spiders. Even tiny details such as genitalia or the
eye arrangements were included in the excellent colour illustrations. Since the original
description only few (three) synonyms were described despite the wide geographical range
(P LATNICK 2004, see below).
O NO (2002) described a subspecies from Japan: Argyroneta aquatica japonica O NO , 2002.
Both subspecies are treated here as one ecological unit, as no differentiating data are present
for A. a. japonica. Examination of European material in comparison with original draw-
ings of both subspecies revealed difficulties to distinguish both forms. Therefore it is re-
ferred in the taxonomic part only to the nominal form.
Information of the following paragraphs was compiled from B ELLMANN (2001), C ROME
(1951), G ROTHENDIECK & K RAUS (1994), O NO (2002), R OBERTS (1995) and own obser-
vations.
De sc ri pti on : Medium-sized spider, males being larger than females both in body size and
leg span; body length (male): 10–20 mm, (female): 8–12 mm; chelicerae with 2 promarginal
and 3 retromarginal teeth, in males 2 distal teeth more separated from the 3 basal teeth
(1-5 e–f); spinnerets slightly conical; tracheal spiracle close to epigastric furrow (1-5 l);
opisthosoma and sternum in living specimens with a quicksilver-like appearance when sub-
merged due to adherent aircover; prosoma and legs brown to dark-brown, opisthosoma
grey to greyish-black; body and legs densely covered by specialised hairs, these longer on leg
III and IV.
Male copulatory organ (1-5 a–d): Distal cymbium elongated (cymbium length / width distal
retrolateral tibial apophysis = 4.75–6.25) and with several spines, some of which variable
in position; subtegulum clearly visible in ventral view; embolus arising from tegulum in
10-o’clock-position, running a semi-circle, its distal part covered by functional conductor
(retrolateral part of tegulum), embolus occupying two thirds of the functional conductor,
6 References
this latter with curved tip, which is resting close to soft membrane of tarsus-tibia-joint;
palpal tibia with broad and blunt retrolateral tibial apophysis.
Female genitalia (1-5 g–k): Epigyne simple, consisting of two short sclerotised rims at the
copulatory openings, the former variable in shape; slit sense organs may be present in an
anterio-lateral position to the copulatory openings; internal duct system with short intro-
mittent ducts and spherical spermathecae; fertilisation ducts arising from ventral position,
running in a semi-circle to the dorsally situated uterus externus.
H a bit at : Fresh water with only little current and rich underwater vegetation.
B io lo gy : Argyroneta aquatica is the only spider which is able to live almost its whole life-
time under water. Spiders construct a silken diving bell, which is filled with air. For this
purpose the spider comes to the water surface, stick the opisthosoma out and take
some air hold by the hindlegs and carry it to the retreat, where it is released. In different
kinds of diving bells the spiders moult, filling the palps with sperms, mate, lay eggs or hi-
bernate. Prey is caught in the water usually outside the retreat, when a prey animal touches
a thread of the capturing web. In rare cases A. aquatica was observed taking prey above the
water surface. Consumption takes place in the air bubble, as the extraoral digestion does
not work in the water. Adults are found all-the-year round. In some cases snail-shells were
documented as hibernation sites. Argyroneta aquatica swims upside down using the legs as
paddles (elongated hairs of legs II and IV).
The poison of A. aquatica is not harmless to humans: a bite causes pain and numbness in the
bitten parts. However, symptoms will disappear at the latest after 14 days without leaving
any damage.
Di stri bu ti on: Palearctic; country records according to YAGINUMA (1986) and B ONNET
(1955): Austria, Belarussia, Belgium, Bulgaria, Czech Republic, China, Denmark, Finland,
France, Germany, Great Britain, Greece, Japan, Korea, Ireland, The Netherlands, Norway,
Poland, Romania, Russia, Serbia, Slowakia, Sweden, Switzerland.
In Germany – and probably also in other countries – the water spider is endangered mostly
by eutrophication. For this reason, several former records (some about 100 years ago) had
to be cancelled from recent distribution maps.
References
B ELLMANN , H. (2001): Kosmos Atlas Spinnentiere Europas. 2nd ed. Franckh-Kosmos. Stuttgart:
1–304.
B ONNET, P. (1955): Bibliographia Araneorum. Analyse méthodique de toute la litérature ara-
néologique jusqu’en 1939. Tome II. Systématique des Araignées. Part 1. Douladoure. Toulouse:
1–918.
C LERCK , C. (1757): Svenska spindlar, uti sina hufvud-slågter indelte samt under några och sextio särs-
kildte arter beskrefne och med illuminerade figurer uplyste. Stockholm. 154 pp. [in Swedish and
Latin]
C ROME , W. (1951): Die Wasserspinne. Die neue Brehm-Bücherei. Leipzig: 1–47.
F ORSTER R.R. (1970): The spiders of New Zealand. Part III. Otago Museum Bulletin 3: 1–184. Du-
nedin.
G ROTHENDIECK , K. & K RAUS , O. (1994): Die Wasserspinne Argyroneta aquatica: Verwandtschaft
und Spezialisation (Arachnida, Araneae, Agelenidae). Verh. naturw. Ver. Hamburg (NF) 34:
259–273.
ICZN (1999): International Code of Zoological Nomenclature. Fourth edition. London: 1–306.
L EHTINEN , P.T. (1967): Classification of the cribellate spiders and some allied families, with notes on
the evolution of the suborder Araneomorpha. Ann. Zool. Fenn. 4: 199–468.
N ENTWIG , W., H ÄNGGI , A., K ROPF, C. & B LICK , T. (2003): Central european spiders - determination
key. Version 8.12.2003 — http://www.araneae.unibe.ch/index.html
1. Order Araneae 7
O NO , H. (2002): New and remarkable spiders of the families Liphistiidae, Argyronetidae, Pisauridae,
Theridiidae and Araneidae (Arachnida) from Japan. Bull. National Sci. Mus. Series A (Zoology) 28
(1): 51–60. Tokyo.
P LATNICK , N.I. (2004): The world spider catalog, Version 5.0, http://research.amnh.org/entomology/
spiders/catalog/
R OBERTS , M.J. (1995): Collins Field Guide: Spiders of Britain & Northern Europe. HarperCollins.
London: 1–383.
S ELDEN , P.A. (2002): Missing links between Argyroneta and Cybaeidae revealed by fossil spiders. Jour-
nal of Arachnology 30 (Proceedings of the 15th International Congress of Arachnology): 189–200.
Lawrence.
S TAUDT, A. (2005). Nachweiskarten der Spinnentiere in Deutschland (Arachnida: Araneae, Opiliones,
Pseudoscorpiones). http://www.spiderling.de.vu/
YAGINUMA , T. (1986): Spiders of Japan in colour. New edition Hoikusha. Higashiosaka: 1–305. (in
Japanese)
Glossary
bristle derivate of cuticle, stiff (!) „hair“, thinner than a (!) „spine“
bulb ventral part of palpal (!) „tarsus“ of adult male spiders, hairless
chelicera appendage of the first prosomal segment of Chelicerata, in spiders two-segmen-
ted, with the distal segment acting as fang against the toothed basal segment.
claw tufts dense brush of fine hairs at the tip of (!) „tarsus“, enabling the spider to walk on
smooth surfaces
claw sclerotised structure at the tip of (!) „tarsus“
conductor part of male (!) „bulb“, which guides the (!) „embolus“ during copulation
and insertion, may consist of analogous structures
cymbium dorsal part of palpal (!) „tarsus“ of adult male spiders, bearing hairs (versus
ventral hairless part = bulb)
dorsal shield sclerotised usually domed plate on the dorsal side of (!) „prosoma“
of prosoma (= [err.] „carapace“)
embolus part of male (!) „bulb“, which is inserted during copulation for transferring
sperm
epigastric furrow posterior margin of the body segment VIII
epigyne sclerotised structure in front of genital opening of adult females, for anchoring
the male genital organ via apopyhses
fertilisation duct inner parts of (!) „epigyne“, where male sperm is transferred from (!) „sper-
mathecae“ to the (!) „ uterus externus“
gnathocoxa coxa of (!) „palps“, usually with serrated anterior margin (serrula), used during
chewing prey and cutting threads (= [err.] „maxilla“)
hair derivate of the cuticle, in most cases innervated and working as sensillum, e.g.
(!) „trichobothrium“
leg appendage of segments III–VI, consisting of seven segments (coxa, trochanter,
femur, patella, tibia, metatarsus, tarsus).
metatarsus see „leg“, is lacking in palp
opisthosoma hind part of the body, consists of fused segments VII–XVII (= [err.] „abdomen“)
palp second pair of appendage, consisting of six segments (metatarsus missing), see
„leg“
paracymbium basal, retrolateral apophysis of the male (!) „cymbium“, having a characteristic
shape in different families
pedipalp see „palp“
8 Glossary
prosoma front part of the body, consists of fused segments I–VI (= [err.] „cephalothorax“)
scapus nail-like structure of the (!) „epigyne“, male apophysis may be fixed with sca-
pus during copulation
spermatheca inner parts of (!) „epigyne“, where male sperm is deposited
spine derivate of cuticle, stiff and thickened (!) „hair“, with well developed base,
thicker than a (!) „bristle“
spinneret reduced paired appendages of (!) „opisthosoma“ segments X and XI, situated
at the posterior end of (!) „opisthosoma“, one- to three-jointed, bearing spi-
gots, which are connected by fine tubes with the silk glands
sternum sclerotised ventral shield of (!) „prosoma“
stridulatory files fine ridges, which are used in combination with peg teeth or spines to produce
sounds either in defensive or mating behaviour
subtegulum basal sclerotised part of the (!) „bulb“
tarsus see „leg“
tegulum second basal sclerotised part of the (!) „bulb“
tibia see „leg“
tracheal spiracle opening of the tracheal system, situated on ventral (!) „opisthosoma“
trichobothrium fine (!) „hair“ on appendages with special base, arising at a right angle, for
receiving air vibrations, arranged in characteristic patterns in different families
uterus externus duct connecting the ovar and the uterus internus with the genital opening, eggs
are fertilised most likely in this part, while being laid
1. Order Araneae 9
Fig. 1-1: a–b, Salticidae: Marpissa radiata, eye arrangement of female (a, frontal; b, dorsal), c–d,
Thomisidae: Heriaeus graminicola, female (c, habitus, dorsal; d, leg claws, prolateral), e–g, Clubio-
nidae: Clubiona stagnatilis, female (e, habitus, dorsal; f, eye arrangement, dorsal; g, leg claws and claw
tuft, prolateral), h–j, Lycosidae: Pirata hygrophilus (h–i, eye arrangement of male, h, frontal, i, dorsal;
j, leg claws, prolateral), k–l, Pisauridae: Dolomedes fimbriatus, eye arrangement of female (k, frontal;
l, dorsal).
10 Figures
Fig. 1-2: a–f, Argyronetidae: Argyroneta aquatica, female (a–b, eye arrangement, a, dorsal, b, frontal;
c, tarsus of leg I with claws and trichobothria, retrolateral; d, coxa to patella of leg IV showing dense
and long hairs, prolateral; e–f, patella, dorsal, e, leg IV, f, leg I), g–r, Tetragnathidae: g–i, l–m, Pachy-
gnatha clercki, g–i, l female, m, male (g, eye arrangement and chelicerae, frontal; h, gnathocoxae and
labium, ventral; i, basal part of femur IV showing dorsal trichobothria, prolateral; l, leg claws, pro-
lateral; m, male palpus, retrolateral); j–k, n–r, Tetragnatha extensa, j–k, n–p, r, female, q, male (e–f,
patella, dorsal, j, leg I, k, leg IV; n, eye arrangement and chelicerae, frontal; o, prosoma, dorsal; p,
gnathocoxae and labium, ventral; q, male palpus, retrolateral; r, epigyne, ventral).
1. Order Araneae 11
Fig. 1-3: a–f, Metinae: a–b, Meta menardi, male (a, gnathocoxae and labium, ventral; b, male palpus,
dorsal); c–f Metellina merianae, c–d, f, female, e male (c, gnathocoxae and labium, ventral; d, eye
arrangement, dorsal; e, male palpus, dorsal; f, epigyne, ventral), g–m, Theridiosomatidae: Theridio-
soma gemmosum, female (g, gnathocoxae, labium and sternum, ventral; h–i, eye arrangement, h, fron-
tal, i, dorsal; j, habitus, lateral; k–l, patella, dorsal, k, leg IV, l, leg I; m, tarsus of leg I with leg claws,
prolateral), n–w, Linyphiidae: n–s, Hypomma cornutum (B LACKWALL 1833), n, p–q, s, female, o, r,
male (n, gnathocoxae, labium and sternum, ventral; o, habitus, lateral; p, eye arrangement and che-
licerae, frontal; q, distal tip of tarsus IV with leg claws, prolateral; r, male palpus, retrolateral; s, epi-
gyne, ventral), t–u, Microlinyphia impigra, eye arrangement of female (t, dorsal; u, frontal), v–w,
Neriene clathrata (S UNDEVALL 1830), male palpus (v, dorsal, w, retrolateral, enlarged).
12 Figures
Fig. 1-4: a–g, Theridiidae: a–e, Rugathodes bellicosus (S IMON , 1873), female (a–b, eye arrangement,
a, dorsal, b, frontal; c, gnathocoxae and labium; d, epigyne, ventral; e, tarsus of leg IV, prolateral), f–g,
Rugathodes instabilis, male (f, tarsus of leg IV, prolateral; g, male palpus, retrolateral), h–r, Araneidae:
h–m, Hypsosinga heri, h–i, k–l, male, j, m, female (h–j, eye arrangement, h, dorsal, i–j, frontal;
k, gnathocoxae and labium; l, male palpus, retrolateral; m, epigyne, ventral), n–o, r, Larinioides
ixobolus, female (n–o, eye arrangement, n, dorsal, o, frontal; r, epigyne, ventral), p–q, Larinioides
sclopetarius, male (p, eye arrangement, frontal; q, male palpus, retrolateral).
1. Order Araneae 13
Fig. 1-5: Argyronetidae: Argyroneta aquatica, a–e, male, f–k, female (a–d, male palp, a, ventral,
b–c, retrolateral, c, spines omitted; d, tegulum, ventral; e–f, chelicera, ventral; g–l, female genitalia,
g–h, j epigyne, ventral; i, k, left half of internal duct system, dorsal; l, opisthosoma showing tracheal
spiracle close to epigastric furrow, ventral).
14
2. Order Acari - General introduction and key to the

major groups
Reinhard Gerecke, Gerd Weigmann, Andreas Wohltmann & Eberhard Wurst
Diagnosis
Small to minute in size (body length in general 0.2-2.0 mm, in extreme cases 0.1-30 mm).
The ancestral arachnid tagmata, the prosoma and the opisthosoma, not apparent as main
body divisions (2-1). Opisthosoma completely fused to prosoma; body secondarily divided
into: (1) the gn at hos om a bearing as appendages the chelicerae and palps, formed in dif-
ferent kinds by the palp coxae and further sclerotized elements of the palp segment; (2) the
i dio som a formed by fusion of the leg-bearing region (called p od osom a) and the opistho-
soma. Often, leg-bearing segments grouped in two parts, with a large area (t ra nsv e rsal
fu rrow) separating the segments of legs I and II (p ro po doso m a) from those of legs III
and IV (me ta pod oso m a). If metapodosoma and opisthosoma form a functional unit,
then the proterosoma (gnathosoma + propodosoma) separated from the hy ste ro som a
(metapodosoma + opisthosoma), thus forming a third level of body subdivision. Genital
opening in anterior part of opisthosoma, anal opening in posterior part, both ventrally.
Respiratory system evolved in different ways within the main taxa. Postembryonal devel-
opment plesiotypically including five free-living stages (plus the non-hatching prelarva):
larva with three pairs of legs only (2-3), three nymphal stages (2-2, 2-4 g-h) and the adult
(2-4 a-f) with the typical arachnid pattern of four pairs of walking legs. Developmental
stages relinquished or realized under particular conditions; e.g. in Parasitengona two of
the nymphal stages transformed into pupa-like resting stages without appendages (2-2).
Fig. 2-1: Mite ground plan and basic terminology - for details see chapters „Diagnosis“ and „General
introduction“.

https://doi.org/10.1007/978-3-662-55958-1_2
2. Order Acari - General introduction 15
Introduction
Like other Arachnida, mites are primarily terrestrial, and therefore aquatic mite taxa are
derivatives from terrestrial ancestors. The small body size, name-giving for this group of
animals (Greek: so ajaqı̀ = the mite, from ajaqiaı̂oy = tiny), developed in several groups to
an extreme extent. Species so small may appear „terrestrial“ only at a first glance: often
they are bound to elevated moisture and important steps in mite phylogeny may have taken
place in hygric microhabitats. Mites demonstrate a range of morphological and physiolo-
gical adaptations not found in other orders of Chelicerata. They can be herbivores, carni-
vores or parasites; they disperse by active migration, passive transport or phoresy, in or
without combination with parasitism; they reproduce bisexually or parthenogenetically,
and they have radiated successfully in both terrestrial and aquatic habitats.
The suborder A na ct in otr ic hi da, with the Opilioacarida, Holothyrida, Gamasida (2-4 a)
and Ixodida (ticks, 2-3 a), only includes a few hygrophilic species, mainly in the Gamasida,
but not one aquatic taxon. In contrast, in the other suborder, the A c ti not ri ch id a, many
different clades include hygrophilous species regularly found in aquatic fauna samples, and
adaptations to submerged life evolved several times independently. An exploitation of
freshwater and marine biotopes, with subsequent adaptive radiation, took place only in
two clades: the Halacaroidea, a morphologically homogenous monotypical superfamily
with ecological gravitation in marine habitats, and the Hydrachnidia, a subtaxon of the
phalanx Parasitengona including about 50 families whose representatives are restricted
mostly to freshwater habitats. Both, halacarid mites and hydrachnid mites developed si-
milar morphological and physiological adaptations to a submerged life (e.g. closed spira-
cles) but display different strategies in their life cycles: halacarid mites are quite homoge-
neous in morphology and particularly minute in size, and their eggs and early stages can be
distributed passively both by water currents and attached to larger animals. Some, or all
stages in the life cycle of their freshwater representatives are capable of colonizing extreme
aquatic habitats such as high mountain pools with extreme changes in water level. Here
they coexist with microcrustaceans, nematodes, oligochaetes, flight active insects, and
hygrophilous representatives of „terrestrial“ mite groups, „true freshwater mites“ (Hy-
drachnidia) being absent. A prerequisite for the evolutionary success of the Hydrachnidia
was probably the presence of a parasitic-phoretic stage in the life cycle of their stem group,
the Parasitengona. Parasitism on flying insects enable the Hydrachnidia to migrate hori-
zontally and thus to compensate for the disappearance of habitats due to summer drought
or due to changes over a longer time scale. This is obviously necessary for long-term co-
lonization success of inland waters.
Due to the particular radiation of these two mite groups (resulting in about 1000 known
species of Halacaroidea and 5000 of Hydrachnidia), the presence of representatives of
other mite taxa in aquatic habitats has often been neglected. However, in many habitats
such as mountain springs, temporary lowland pools or the hyporheic interstitial, species of
Trombidiidae, Oribatida or Acaridida may be found to be dominant in terms of population
density. Many of them display well developed adaptations to a submerged life, sometimes
during all life stages.
Morphology and terminology

The basic morphological ground plan of the mite body is explained in Fig. 2-1. Several
systems of morphological terminology have developed in terrestrial and aquatic acarolo-
gical research, particularly in North American, West European and East European scien-
tific traditions. Therefore, a detailed explanation of the morphological terminology and a
16 Sampling and sample processing
specific glossary for each of the mite groups treated in this work are provided in the appro-
priate chapters.
Sampling and sample processing

Collecting
Due to the minute size of mites, they require a special treatment both during field work and
in the laboratory (e.g., B ARR 1973, S MITH & C OOK 1991, for methodological discussion
with particular regard to spring habitats, see G ERECKE & D I S ABATINO in press). If stan-
dard techniques such as Surber sampling, dredging, hand netting or drift sampling are ap-
plied (for details see S CHWOERBEL 1994), adult Hydrachnidia can be collected adequately
only if the m e sh si z e does not exceed 250 lm. For collecting their larval stages as well as
the smaller representatives of mite groups such as halacarids, a mesh size of 100 lm is
necessary. Many limnological sampling methods have, however, two major disadvantages:
first, they do not adequately represent species with a patchy distribution - species adapted
to particular life conditions, accumulating in particular microhabitats at the edge of water
bodies, may be easily overlooked or underestimated. Second, especially many species of
Hydrachnidia as specialized predators and/or parasites, may be present in population den-
sities which are lower by an order of magnitude compared with other elements of the meio-
fauna or macrofauna (G ERECKE et al. 2005). Consequently, such species, often character-
istic elements of the studied habitat, are hardly represented in collections applying area- or
time-limited standard methods. For a representative investigation of the acarological di-
versity, in te nsi ve h an d-n et ti ng is obligatory in all microhabitats. The sediment should
be dispersed locally, tufts of plant material vigorously shaken and the material washed and
filtered through a net. This method does not allow for quantification, but differences in the
relative abundance between the species can be studied.
Since most Hydrachnidia (like many limnic insects) are aerial during particular phases, the
use of Ma la ise t rap s and swe e p nets helps to estimate population densities and both
instruments are indispensable tools to investigate population development and phenology.
Moreover, soil probing and hand collecting during the dry phases of temporary waters al-
lows the detection of resistant stages and the estimation of survival strategies in such ha-
bitats. In the following, we give a more detailed survey on sampling techniques and their
application in various habitat types:
Lentic habitats:
1. D ip -ne ts are often used by hand for sampling in the littoral vegetation, but this tech-
nique is also useful in scubadiving in vegetation deeper than 1-2 m.
2. Fra m e- ne ts operate like a dip-net but have a square, round or triangular opening. Col-
lecting is accomplished by pulling a frame-net for a certain distance through the sediment to
a depth of 2.0-2.5 cm. Each sample taken represents a certain area. Samples are passed
through a sieve and are further sorted by eye (T EN W INKEL 1987). Vertical haul: nets
are lowered to the bottom and hauled vertically (B AGGE et al. 1996).
3. G rab s are suitable for estimating the density of the organisms living on the bottom.
Organisms swimming close to the bottom will also be collected (K. V IETS 1930).
4. Un de rwa te r t rap s can attract mites to a light source or to some kind of bait. L i gh t
tr aps may work on the base of electric (P IECZYNSKI 1962, 1969; B AGGE et al. 1996) or
chemoluminescent light sources (B ARR 1979a, see there for more practical details) placed in
the centre of a polyethylene container with a wire-basket like entrance. Many Hydrachni-
dia are attracted in particular to yellow-orange light. However, the technique works selec-
tively: a few species, e.g. Hygrobates nigromaculatus and H. trigonicus, show no positive
phototaxis (D AVIDS et al. 1994). C ONROY (1973) proposed an underwater trap working
with bait instead of a light source.
Lotic habitats:
1. Ex tra c ti ng a nd wa shi ng of submerged substrata (plants, detritus, stones). The ma-
terial is at best transferred under water into a net and then put into a white tray filled up to a
few cm of clear water. Regular collection of a constant amount of determinate material (e.g.
a certain volume of mosses) may provide quantitative indications (B ADER 1977). Larger,
immovable structures such as trunks of trees and large rocks are brushed in situ and the
material collected in a hand net.
2. K ic k sam p li ng : In a gravelly stream habitat a net is held vertically in the stream, with
the mouth opening upstream. The collector stands upstream from the net and vigorously
disturbs the substratum to a depth of 15-20 cm, so that detritus and animals are washed
into the net. The net is then withdrawn from the water and the sediment collected distrib-
uted over the bottom of a white tray (B ARR 1973).
4. Additional species with a patchy distribution may be detected by drift netting. In contrast
to other stream invertebrates, Hydrachnidia drift with maximum abundance in daylight
(S CHMIDT 1969).
5. D EWEZ & WAUTHY (1984) worked with an artificial substratum kept submerged for at
least eight weeks. At the end of the experiment, mites were extracted and counted.
Interstitial habitats:
1. The „K a ra m an -C hap pu is“ d ig gi ng is in most cases the technique best suitable for
collecting the biocoenosis in the hyporheic interstitial (C HAPPUIS 1942). A hole is dug in the
exposed gravels of a stream, deep enough to penetrate the water table. The interstitial water
percolating into the hole is removed using a small bucket or bowl and washed through a net
or sieves to eliminate turbidity.
2. The use of a so called „Bo u Ro uc h“ pump is an often applied alternative (V IGNA TA-
GLIANTI et al. 1969; B OULTON et al. 1992). This technique has the advantage of being ap-
plicable in the absence of gravel banks, pumping the sample directly from the interstitial
below the bed of the stream. However, in general, in small streams without sand or gravel
banks, the sediments below the running water do not provide habitats suitable for colo-
nization by hyporheobiont specialists. In general, as compared with digging, pumping pro-
duces lower numbers of specimens and species and a lower amount of hygrophilous ripi-
coles, but a higher percentage of species confined to hyporheic habitats (G ERECKE et al.
2005).
Semiterrestrial habitats:
For quantitative collecting hygric populations of A c ari di da, O ri ba tid a and t e rre stri al
Pa ra sit en go na, it is recommended that substrate (algae, moss cushions, detritus) be
brought into the laboratory. Mites can be easily extracted from the substrate using a
heat-driven dynamic extraction method such as a Berlese- or MacFadyen-apparatus, or
the mites can be sorted from the samples directly. However, these methods work selectively
and are not suitable for extracting halacarids.
Treatment of samples
Due to their bright colours and/or active behaviour, most deutonymphs and adults of the
Parasitengona may be sorted on the spot from the collected material if light is favourable
and a flat white pan or tray is used. Mites become active a short time after the sample is
poured into the pan and can easily be sorted by eye and picked up with a pipette. Samples
with much detritus and silt should be washed carefully through a set of two sieves. By stir-
ring clean water through the upper sieve (mesh size about 1.5 mm.), the mites will accu-
mulate in the lower sieve (mesh size 100 lm). Typical lotic species do not move actively
when the sample is removed from the habitat. However, under the influence of rising tem-
perature and decreasing oxygen availability, Hydrachnidia leave the collected substrata to
swim or crawl in the open water. Mites living in moss growing at the air-water interface or
in cascades often cling tightly. To extract them, a clump of moss is picked apart in a tray of
water, strand by strand, with a pair of forceps, carefully searching for mites, or the sample is
stirred vigorously until the mites become detached. Sorting in the field is of particular im-
portance if the remaining sample is to be preserved in a fixative such as formalin or alcohol
(see below). However, not only representatives of the Halacaridae, Acaridida and many
groups of Oribatida which are small in size and less agile, but also larvae and small nymphs
of Parasitengona can be sorted only in the laboratory under a stereo microscope, preferably
from the living material. Living samples are best transported at a low and stable tempera-
ture, bathed by, but not submerged in water (in order to avoid mechanical damage), and in
contact with a large volume of air for oxygenation. Most mites are rather resistant against
many kinds of stress during sampling and transport and are found still alive in the sample
when many of the other invertebrates have died. Also in the laboratory, sorting should be
processed through stacked sieves, the top one with a mesh size of 1000 lm, and the bottom
one with 60–100 lm, being rinsed with a strong water jet.
In the laboratory, mites can be extracted from bottom mud by placing the sample for 10
minutes in a concentrated magnesium sulphate solution. The sample is stirred occasio-
nally to allow animals to float to the top and the floating mites are subsequently decanted
(E FFORD 1965). Further flotation methods, which make use of the specific weight of the
animals work with solutions of sodium chloride, sugar or silicate. P ROCTOR (2001) got
good results with a kerosene-flotation technique. FAIRCHILD et al. (1987) were successful
in extracting Hydrachnidia from moss samples by inducing them to respond to a tempera-
ture gradient. Preserved samples with little detritus can be stained with Bengal red.
Classification of undissected material

The mastery of dissection and slide mounting techniques (see below) is an important pre-
requisite not only for the inexperienced student. Confronted with material from little
known habitat types or unstudied geographical areas, even a specialist who is an old
hand at classifying mites is obliged to produce series of preparations in order to investigate
diagnostic characters and their variability range. However, once a certain experience and
familiarity with a local fauna is built up, most of the recorded mites may be recognized
without preparative work. For instance, large parts of collections from habitats regularly
investigated in the course of long term monitoring projects may be sorted under the stereo-
microscope to species level. For the solution of problems requiring stronger magnifications
(recognition of sexes, separation of similar sister species) it is in general sufficient to ex-
amine selected character states under the light microscope, temporarily placing the undis-
sected mites on a concavity slide or, in order to bring the specimen into a particular po-
sition, on a slide prepared by the application of a viscous medium such as H OYER’S fluid
(see below). In this manner classifying mites from aquatic habitats may become as routine
as the study of other groups of benthic invertebrates.
However, specimens which are conspicuous e.g. due to particular dimensions, coloration
patterns or aberrant characters merit special attention and should be dissected and slide
mounted. Water mites tend to form monstrosities, often probably due to developmental
disturbance at the proto- and tritonymphal resting stages, but there are also very rare, often
enigmatic species of particular faunistic interest which appear only on rare occasions.
Furthermore, selected specimens of the more problematic species should be regularly slide
mounted at regular intervals in order to control the correct taxonomic attribution. In any
case, slides as well as material classified without slide mounting should be carefully label-
led and conserved for later reinvestigations. For long term preservation in vials, both
K OENIKE’S fluid (see below) or 96 % glycerine are suitable media (B ARR 1973).
Rearing
The most satisfactory method of obtaining larval material for study is from eggs laid by
identified females in the laboratory. Depending on the scope of the study, mites are isolated
in couples or as single females in small vials or petri dishes together with substrata suitable
for spermatophore and/or egg deposition (e.g. plant stems or pieces of filter paper). Tem-
perature is best at a stable level similar to that of the species’ natural habitat. As most spe-
cies are adapted to starvation, in most cases feeding is not necessary for successful repro-
duction, at least if females are ovigerous at the start of the experiment. However, eggs
should be kept separate from parents in order to avoid feeding by parents on their progeny.
If the whole life cycle of a parasitengone species is to be studied, potential hosts are exposed
to the freshly hatched mite larvae in small aquaterraria. Completing a parasitic phase de-
mands much ingenuity and effort, no general rules can be given. For the resolution of ques-
tions concerning the host range of a selected species, it is recommended to interrupt the
experiment when larvae have attached to their hosts and prepare them on microscopical
slides (M ARTIN 2000, 2003). A rather simple method for investigating parasite-host rela-
tions under natural conditions is the exposure of sediments which include both a mite po-
pulation and their presumptive hosts in plastic bags in the field (G ERECKE & M ARTIN
2006). However a good knowledge of the fauna of the site is necessary because attribution
of larvae and adults cannot be made with 100 % certainty. Bibliography concerning the
wide field of specific requirements for successful reproduction is cited in the special part.
Preservation
Mites of the „terrestrial“ groups (acaridids, oribatids, terrestrial Parasitengona), also ha-
lacarids can be preserved in ethanol (75 %). Some drops of glycerine may be added to pre-
vent the sample from drying out if the ethanol accidently evaporates (see K RANTZ 1978).
Formalin should be avoided, at least the animals should be transferred to ethanol as quickly
as possible. For histological studies animals should be preserved with glutar aldehyde or
Bouin’s solution and subsequently be transferred to ethanol. If a sample dries up comple-
tely, water with a drop of washing-up liquid or some vinegar should be added. After soak-
ing, the mites are transferred to ethanol. For Hydrachnidia required for dissection and light
microscope studies, preservatives containing formalin or alcohol, are particularly disad-
vantageous. These substances fix the tissues in such a way that they cannot be properly
cleared in either basic or acid corrosives. In general, clearing with lactic acid leads to ac-
ceptable results in all mite groups.
In the case of strongly sclerotized species like many oribatids, a preceding exposure to hot
(90 C) 10 % KOH is advantageous; combination of KOH followed by lactic acid may also
allow clearing of mites preserved in formalin or alcohol. Water mite workers traditionally
use as a preservative a glycerine - acetic acid - water mixture commonly referred to as
„Koenike’s fl ui d“. In the first days following fixation, this liquid may cause a shrinking
of the animal and gas bubbles may fill parts of its body. A few days later, the animal extends
again, the gas disappears and it becomes clear due to tissue breakdown. At the end of
this process, all important characters of the skin as well as the sclerotized parts are clearly
visible.
Molecular aspects which have gained increasing importance during the past years can be
studied best on specimens preserved in special buffer solutions. Indications that molecular
DNA studies are possible also with material preserved in Koenike’s fluid are based on a very
small sample size and gave worse results compared with ethanol-fixed material (R EY et al.
2004).
Contact with any type of preservative nearly always causes the loss of most of the coloura-
tion, namely in Parasitengona where most pigments are distributed in the internal organs or
in lipids below the cuticle. Only colours located in, or originating from, sclerotized struc-
tures and some basic contrast patterns (e.g., the whitish contents of the excretory organ)
may remain visible after years of preservation. Therefore, it is useful to document the col-
ouration of interesting specimens before they are transferred to the preservative. A little
studied, but promising alternative could be freeze-drying. However, specimens treated
in this manner are mostly useless for further detailed microscopical investigation.
For the tiny Ac a rid id a with their delicate cuticular structures only two methods of treat-
ment are recommended: if the mites are alive, the specimens should be mounted directly in
Hoyer‘s fluid; in doing so the mites stretch their legs and the natural shape can be preserved.
In order to avoid compression of large specimens the coverslip should be supported at the
edges by wax or modelling clay. For a more permanent preparation, after drying the cover-
slip should be ringed with Glyceel (BDH Chemicals Ltd, Poole, England) or Glyptal (not
available in Europe, see T RAVIS 1968). If mites are not subject to immediate slide prepara-
tion they should be stored in ethanol (at least 70 %). Clearing agents are not necessary and
should not be applied.
In contrast, many O r ib ati da have a dark and poorly transparent cuticle. Lactic acid
(90 %) can be used to clear the body of the mites by maceration of the inner organs, which
allows better examination of the cuticular characters. Temporary mounts in a cavity slide
filled with glycerine or lactic acid (under a coverslip) is preferred. If the coverslip covers
only a third or half of the cavity, the mite can be manipulated using mounted fine needles.
Such a preparation allows the specimens to be turned and thus be observed from all per-
spectives.
H a la ca ri ds, again a group of rather tiny and often poorly sclerotized mites, can often be
slide mounted without preceding treatment. Dark specimens may be cleared in pepsin or
lactic acid for some minutes, exceptionally up to three days. If the body content of a speci-
men is very hard, it helps to apply heat up to 50 C or to add acetic acid. If the animals are
very fragile or the integument cleared too strongly, they can be stained with chlorazol black
for microscopic examination.
Dissection
As mentioned above, dissection and preparation are not obligatory steps if the study of
water mites is included into large scale limnological work. However, they are important
for the documentation of little known or newly detected species and during studies on po-
pulation variability. The dis se ct io n of body parts is best done in a drop of glycerol or
K OENIKE’S fluid on a slide with a central cavity under a stereo microscope. Tools such as
hooked minuten pins, fine needles (insect pins number 4 _ 1) and micro-scalpels can be used.
First the gnathosoma and the legs of one side are cut off with a sharpened needle and re-
moved to the edge of the drop in such a way that the drop becomes enlarged and the objects
are lying in a semicircle, so the legs cannot be mistaken one for the other. The body content
is then gently squeezed out with a blunt needle. If the body content is too dark to allow
transmission of light, or if there are important details to observe both on the dorsal and
ventral surface, it is necessary to separate the dorsum from the venter. The mite is laid down
on its side and cut with a micro scalpel into a ventral and a dorsal part. Next, the body
contents can be removed. If the specimen has been preserved in Koenike’s fluid it will be
very easy to remove all soft parts leaving only the exoskeleton.
If the body parts need further clearing, e.g. in alcohol-preserved specimens, this can be done
in lactic acid: the objects are placed in a drop of lactic acid on a corner of a slide and heated
with a spirit flame. The flame must be moved along the edge of the drop till the lactic acid
starts to smell. In this way the drop remains. A further possibility is the use of André’s fl ui d
(see below) which has the advantage that specimens may be left in it for several hours with-
out damage to the integument. However, submersion of lightly sclerotized specimens for
more than 48 hours tends to weaken everything, e.g. leg joints, and it obscures shields. A
possible problem is that slightly sclerotized specimens may become so transparent that they
are virtually impossible to find in the preparation dish. Another disadvantage is that any
colour pattern disappears almost immediately. A basic corrosive, 10 % Pot ass iu m H y -
dr oxi de (KOH), is advised for use only with heavily sclerotized mites, KOH should be
used at room temperature and the specimens left in the solution for only one to two hours
(unsclerotized mites less than 15 minutes). A short contact with KOH does not destroy the
colour pattern of the mite as is visible in preserved specimens.
Attention should be paid to the fact that representatives of different genera may need dif-
ferent treatment. For instance, in Hydrachnidia of the genus Hygrobates the gnathosoma is
fused to the first coxae and cannot be isolated. Therefore, chelicerae can be removed only
after cutting the idiosoma, pulling them out from inside the body. Species of Eylais can be
determined best with the gnathosoma in situ (vs. visible from its ventral surface).
Mounting
The dissected parts of the mite should be mounted in a rather restricted area in the centre of
the slide, so that they are easily found in the microscope preparation, at best following a
stable scheme. If the idiosoma is dissected horizontally into two parts, dorsum and venter
should be placed side by side, both with their external surface directed to the observer. If the
idiosoma is not separated, the optimum position should be selected depending on the taxon
in study. In many soft bodied species it is recommended to orientate the dorsal surface to the
observer because in this way the often minute dorsal structures are visible, but also, by
transparency, those of the venter. Detached legs are best located laterally from the idioso-
ma, the genital skeleton or the eggs posteriorly, the gnathosoma and its appendages ante-
riorly. It is important that both palps are orientated in the same direction to assure both
lateral and medial views - naturally the same is true also if both legs of the same segment are
dissected.
An overview of mounting media and their refraction indices for small arthropods can be
found in K OOMEN & V ON VAUPEL K LEIN (1995) and S TOCK & V ON VAUPEL K LEIN
(1996). In recent years, in most acarological laboratories aqueous mounting media
have replaced oil soluble resins such as balsam, primarily because specimens may be
mounted without preliminary fixing operations as required for resin mounts. Furthermore,
they have better optical properties. A disadvantage is that they are hygroscopic and often
tend to liquefy or to dry up, depending on the humidity in the room where they are kept
(U PTON 1993). The application of a sealant ring may slow down, but not completely inter-
rupt, such processes.
Gl yc e rin e – je ll y (a mixture of glycerine, gelatine, distilled water and a little carbolic
acid) is probably the best medium for pe rm a ne nt pre pa ra ti ons (K. V IETS 1923,
1936, H EVERS 1985). In museum collections, slides mounted in this medium have remained
in good condition over many decades. The study objects retain shape without shrinking and
also the fine integumental structures do not undergo alterations. The unique phenomena of
deterioration frequently observed in older collection material are the forming of fine, dark
droplets covering the body structures, or the precipitation of crystals. To avoid the potential
risk of mustiness, preparations should be kept in a dry climate. An important advantage is
that mounts can be opened again by carefully heating, then cleaned and reorganized. Speci-
mens preserved in an aqueous medium can be transferred directly to glycerine-jelly if it is
liquefied by heating to about 50 C. Prior to mounting cleared specimens the lactic acid or
other clearing agents have to be removed. For each mounting session, only an adequate
portion of glycerine jelly should be heated (heated too often, the medium becomes useless).
A droplet is placed in the centre of a slide and extended to an area equal in surface to the
coverslip. During the process of solidifying with lowering temperature, the dissected parts
of the mite can be placed and orientated. Action may be taken on the cooling speed, e.g.
placing the slide on artificial ice for accelerating, or keeping it warm if more time is needed
for a complicated mounting procedure. However, after cooling down, the objects can be
rearranged with warm needles. Air-bubbles and dust can be taken away with a micropip-
ette. After hardening of the medium, the preparation is closed with a coverslip bearing a
thin layer of liquefied medium on its bottom side. Excess of glycerine- jelly after placing the
coverslip can be removed with a micropipette. It is important to keep the layer of mounting
medium as thin as possible: a too large distance between the object and the coverslip would
hamper the use of high microscope magnifications. An equally simple and effective sug-
gestion was published by J ÄNTSCH (2002 - for more details see there): mites are mounted
on the coverslip instead of the slide. Due to their own weight, dissected parts sink down and
are placed as close as possible to the future upper side. After hardening of the medium, the
coverslip, still upside-down, is picked up with a thin drop of liquefied medium which hangs
in the centre of a slide. When the preparation mounted in this way is kept upside down until
completely hardened, gravitation will guarantee that the objects remain in the desired po-
sition. The mount can be sealed after half an hour with a rim of lacquer. Afterwards the
slides must be labelled and should be stored in slide drawers or boxes so that the slides lie
flat and, if possible, upside down.
A technique for making particularly perfect preparations and of particular aesthetic value,
with the help of a turn-table is described by H EVERS (1985, see there for details).
H oy e r’s Fl ui d is a medium that has become popular because it is not only very easy to
use, but it has particularly suitable optical properties (K RAUS 1984). It is viscous and tends
to harden by evaporation at room temperature; as in the case of glycerine-jelly, the objects
may be transferred directly from water or Koenike’s fluid. The medium has the capacity of
clearing the mounted material as time goes by. After slide-making, preparations can be
completely hardened in about 48 hours by heating to about 50 C. The possible problem
that during this heating turbulence may cause displacement of the objects can be avoided if
one or two days before mounting, slides are prepared with flat droplets of the medium
(G ERECKE 1991). For mounting, only small quantities of additional liquid medium are
added to the surface of these hardened droplets. Here, the objects may be arranged perfectly
in the locally liquefied surroundings and will not change their orientation during the fol-
lowing short process of drying. These preparations can be sealed with a coverslip imme-
diately or after hardening and a final control. Such preparations may be opened again very
easily by adding water (acceleration by warming). Hoyer’s fluid is recommended if large
numbers of specimens are to be investigated in a short time. Furthermore, Hoyer’s fluid has
been used with good results for tiny water mite larvae. Living or preserved larvae are placed
directly into the media and a cover slip added. However, this medium should not be used for
mounting valuable material such as type specimens or representatives of rare and uncom-
mon species. Its properties in long-term conservation are unknown. Both liquefying or
complete drying up have been observed, often accompanied by deformation of the objects
by compression and/or torsion.
Po ly vi ny l la c top he no l, another medium easy to handle, should also not be used for
producing type material or voucher specimens, because the integument of the objects soft-
ens and it is almost impossible to remount specimens without deforming the integument.
Glycerol (sealed with a wax ring) is a useful medium for short and medium term prepara-
tions.
B ARR (1973) and C OOK (1974) advise and describe a double coverslip method. The idio-
soma of mites mounted between two coverslips can be observed from both sides and there-
fore this method is economic with dissection time. In order to reduce the risk of breaking or
loss of information (the double coverslip may detach from its base), the „coverslip sand-
wich“ may be introduced into appropriate frames (for example aluminium frames).
Formulae:
Koenike’s fluid Alternative mixture (B ARR 1973,
(modified) C OOK 1974):
Glacial Acetic Acid 2 parts by volume 1 part
Glycerine (96 %) 5 parts „ „
5 parts
„ „
Distilled water 3 parts 4 parts
To avoid strong maceration, e.g. for Eylais spp., the use of the alternative mixture is re-
commended.
André’s fluid (Acetic Corrosive)
50 ml Glacial Acetic Acid
50 g Chloral Hydrate
50 ml Distilled water
Fig. 2-2: Pupa-like resting stages of aturid mites (Hydrachnidia) in ventral view: a, protonymph, char-
acterized by one pair of acetabula and detaching pieces of the larval exuvia (dorsal shield and left coxal
plates, with the larval „urstigma“ visible); b, tritonymph, characterized by four pairs of minute acet-
abula (in curved lines in the posterior part) and detaching pieces of the deutonymphal skin (coxae with
leg remnants, „provisory genital organ“). Note the adult developing inside the nymphal integument,
with folded legs and numerous acetabula arranged parallel to the posterior margin, typical for species
of the subfamily Aturinae.
Fig. 2-3: Mite larvae (a-b ventral view): a, Ixodida; b, Acaridida (Schwiebea sp.); c, Oribatida, dorsal
view (Hydrozetes lemnae); d-e, Halacaridae (Lobohalacarus weberi); d, ventral, e, dorsal view (with-
out legs); f, terrestrial Parasitengona (Abrolophus quisquiliarus); g, Hydrachnidia (Sperchon squamo-
sus); h, Hydrachnidia, dorsal view of gnathosoma (Stygothrombium sp., only the basal leg segments
given).
Fig. 2-4: Mites (a-f adults, ventral view): a, Gamasida; b, Astigmata (Schwiebea sp.); c, Oribatida
(Hydrozetes lemnae); d, Halacaridae (Lobohalacarus weberi); e, terrestrial Parasitengona (Abrolo-
phus quisquiliarus); f, Hydrachnidia (Sperchon mutilus); g-h, deutonymphs; g Hydrachnidia (Sperch-
on mutilus - given at the same magnification as adult in f); h, Acaridida (Schwiebea sp.).
28 Key to major groups
Hoyer’s fluid
50 g Distilled water
30 g Gum arabic, clear crystals
20 g Glycerine
The ingredients should be mixed in the order listed above. Each substance should be fully
dissolved and mixed before the next is added hence each step may take several days. The
procedure is accelerated by heating in a water bath (K RAUS 1984).
Lundblad’s Solution
To get rid of oil droplets prior to mounting in glycerine-jelly, L UNDBLAD used this solution:
112,5 g Phenol
12.5 ml Distilled water
Use for about 5 minutes. It is necessary to follow the process under the microscope because
specimens and appendages become more-or-less colourless (work in a cavity slide).
The major groups of mites found in aquatic habitats

Key to life-cycle stages
1 Mouth opening and mouth parts, body sclerites, eyes and legs absent (but exuviae of appendages
of the preceding stage may be attached to the outer surface; also, sclerotized structures and eyes of
the next stage may be seen through the integument, 2-2); animals spherical or egg-shaped, covered
by a thick, often sculptured integument; the only external organ is a smooth or slightly sclerotized
field generally with one or two pairs of minute, cup-shaped osmoregulatory organs („acetabula“,
absent in Erythraeoidea) located in the centre of the venter (future genital area); a longish zone
with a smooth surface, may indicate the future mouth-part area („pharyngeal organ“). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . resting proto- and tritonymphs of Parasitengona
– Body with appendages, eyes often present; sclerites present or absent . . . . . . . . . . . . . . . . . 2
2 Three pairs of legs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . larvae
(„mite-larvae-like“ wingless insects are generally distinguished by the presence of articulated
antennae, in particular cases, also by mouthparts transformed into a sucking apparatus, or
by the presence of appendages on the ventral abdomen modified for jumping).
– Four pairs of legs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . active nymphs and adults
Key to larvae
1 Mouth opening on an elongated projection ventrally densely covered by barbed hooks (2-3 a) or a
bifurcated appendage („tritosternum“) located at the ventral base of the gnathosoma (as given in
2-4 a for an adult); coxae of legs movable, spherical or cylinder-shaped. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ixodida and Gamasida
(terrestrial, accidentally in aquatic habitats, not treated in this work)
– Mouth opening not on an elongated projection densely covered by barbed hooks, no bifurcated
appendage („tritosternum“) located at the ventral base of the gnathosoma; coxae of legs fused to
the body surface in the form of coxal plates, not movable. . . . . . . . . . . . . . . . . . . . . . . . 2
2 Palps with one or two movable segments only (2-3 b); one pair of rod-like Claparède organs (2-3 b)
or ring-shaped osmoregulatory organs (3-34) between leg I and II; prodorsum never bearing
trichobothria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acaridida (page 38)
– Palps with more than two movable segments (e.g. 2-3 f, 2-3 g – but note that Oribatid larvae may
have their palps retracted into their rostrum); osmoregulatory organs between leg I and II rod-like
(„Claparède organs“), cup-shaped (2-3 g) or invisible (2-3 f), generally paired, rarely supernumer-
ary; trichobothria present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Gnathosoma (including palps and chelicerae) more or less covered by the rostrum of the pro-
dorsum, not or hardly visible in dorsal view (2-3 c); prodorsum with or without trichobothria
in dorsolateral position; anal opening in the form of a longitudinal slit; often species with a re-
markable contrast in colour between the appendages (dark) and body (clear). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oribatida (page 89)
– Palps and chelicerae visible in dorsal view (2-3 h), not covered by a prodorsal rostrum, anal open-
ing in the form of a round pore (2-3 g), a longitudinal slit (2-3 d), or absent (2-3 f); colour var-
ious. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Gnathosoma snout-like (cone-shaped, equally tapering from the base to the tip, as given in 2-5 a
for an adult); palp tarsus distally with two conspicuous, long setae; palp tibia without claw;
chelicerae elongate, distally with small movable digit; two pairs of trichobothria; active at terres-
trial conditions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bdellidae (Eupodina)
(several species present in semi-aquatic habitats, not treated in this work)
– Gnathosoma not snout-like as described above; palp tarsus various, but without two conspicuous,
long distal setae; palp tibia with or without claw; chelicerae various; trichobothria present or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Idiosoma dorsally covered by one single hemispherical shield (as given in 2-5 c for an adult); em-
podium of legs branched into adhesive setules; palp tibia with claw; trichobothria absent from
idiosoma, body red in colour, specimens often found floating on water surface. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homocaligidae (Eleutherengona)
(restricted to aquatic habitats, in Central Europe only Homocaligus scapularis (C.L.K OCH ,
1838), not treated in this work)
– Dorsal shield, if present, plain or only little curved; empodium of legs claw-like or absent; palp
tibia with or without claw; trichobothria present or absent. . . . . . . . . . . . . . . . . . . . . . . . 6
6 Idiosoma with only three pairs of ventral setae (2-3 d - no anal setae) and at most seven pairs of
dorsal setae; palp four-segmented; legs with five movable segments; free living. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halacaroidea (page 113)
– Idiosoma with more than four pairs of ventral setae (2-3 f-g - at least one on each coxa and two
pairs of anal setae) and more than seven pairs of dorsal setae; number of palp segments various,
often five; legs with five or six movable segments; mostly parasitic on arthropods, but may be
found during the preparasitic phase in high population densities in sediment samples. . . . . . 7
7 Anal opening present or absent (2-3 f); without a sclerite or flanked by a pair of sclerites. Osmor-
egulatory organs never stalked, coxa II with one or more setae; legs I/II short or long (occasionally
more than double the length of the idiosoma); number of setae on palp genu various: one (Trom-
bidioidea), one to two (Erythraeoidea) or many (Calyptostomatidae). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . terrestrial and hygric Parasitengona (page 158)
– Anal opening present, usually within an undivided sclerite (2-3 g); if flanked by paired sclerites
(Stygothrombidioidea) then osmoregulatory organs stalked and reaching well beyond the lateral
body sides; if anal sclerite absent (Hydryphantoidea: Wandesia) then coxa II without setae; length
of legs I/II never more than double the length of the idiosoma; palp genu with two setae. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aquatic Parasitengona: Hydrachnidia (page 241)
Key to nymphs and adults

In the following key, we use adult character states also found in the related nymphal stages. In general,
besides the genital area being less developed and with the gonopore closed, nymphs differ from adults
in their smaller size, lower numbers of setae on the appendages, lower numbers of osmoregulatory
organs („acetabula“, in Acaridida called „genital papillae“ or „ring organs“) and relatively large areas
of unsclerotized integument (compare 2-4 f-g).
1 Gnathosoma with minute palps consisting of one to two movable segments (2-4 b), or completely
reduced; if a gnathosoma and mouth opening are absent, mouthparts replaced by a longish ap-
pendage („palposoma“, 3–13) with two long distal sensory setae („solenidia“) or by a group of
sensory setae between the insertions of the first legs; furthermore, nymphs (3-13) with a reduced
gnathosoma bear an adhesive organ or a clasping apparatus with cuticular plates ventrally at the
posterior end ; stigmata and trichobothria always absent; only one claw on the pretarsus; males
ventrally with an external copulatory organ (aedeagus). . . . . . . . . . . . . Acaridida (page 38)
30 Key to major groups
– Gnathosoma well developed, palps with three to five movable segments (2-4 a, c-g); ventral body
surface without any adhesive organ or clasping apparatus; with or without stigmata and tricho-
bothria; one or two claws on the pretarsus; males only in exceptional cases with an external co-
pulatory organ. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Coxae of legs movable, spherical or cylindrical - thus, legs with seven movable segments (2-4 a); a
pair of stigmata on sclerotized plates laterally near the third or fourth legs (2-4 a); leg pretarsus
always with well developed paired claws and an unpaired membranous pulvillus; chelicera with
well developed pincers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Coxae of legs fused to the ventral body area in the form of sclerotized plates on which the leg
trochanters insert - thus, legs maximally with six movable segments (2-4 c-g); stigmata, if present,
in the anterior area of the idiosoma or in the gnathosoma, often not clearly visible; leg claws and
chelicera of variable shape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Gnathosoma with an elongated extension (“hypostome”) on its ventral side densely covered by
barbed hooks (as given in 2-3 a for the larva); stigmata on a porose plate laterally near coxa IV;
palp tarsus without a fork-like seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ixodida
(ticks - occasionally found in aquatic environments frequented by cattle as drinking places;
however, purely terrestrial, not treated in this work).
– Gnathosoma without a hooked, elongated stiletto-like extension; stigmata on a plate near the
third legs and associated with a longish sclerite more or less extending anteriorly („peritrema“
= tracheal groove, 2-4 a); palp tarsus at the inner margin with a fork-like seta. . . . Gamasida
(several hygrophilous species regularly found in samples of riverine substrata from all kinds of
inland waters, and also in cave waters; however, no aquatic species known; not treated in this
work).
4 Chelicerae and pedipalps normally not visible in a dorsal view, protected by a projection („ros-
trum“) of the prodorsum (4-1); with or without trichobothria dorsolaterally on the prodorsum; (in
some nonaquatic families, posterior dorsum segmented by transverse sclerotized plates); adults
characterized by genital and anal openings similar in dimensions, both protected by “window
shutter”-like paired plates (2-4 c), usually heavily sclerotized, with a yellow to dark brown cu-
ticula (sometimes with a mirror-like reflecting surface, sometimes whitish due to reflecting plas-
tron structures); nymphs characterized by a genital field without sclerotized plates, in the form of
longitudinal slits, mostly with a soft integument and unpigmented, but sometimes bearing dis-
tinctly contrasting brownish or reddish plates and appendages. . . . . . . Oribatida (page 89)
– Chelicerae and palps generally visible in a dorsal view, not protected by a projection („rostrum“)
of the prodorsum (2-4 i, exceptions are made in species of Isobactrus, halacarids with the gnatho-
somal base not visible in a dorsal view, of Calyptostoma, terrestrial Parasitengona, as well as a few
Hydrachnidia taxa with gnathosoma and its appendages being retractable into the idiosoma); gen-
erally with extended membranous areas (2-4 e-g, i) or, if heavily sclerotized, plate surfaces not
darkened, at most opaque due to dense porosity (2-4 d, 7-18 b, 7-19); colour variable, from shiny
white to intensely red, blue, green or brown; dark coloration, if present, always originating from
internal organs, not from sclerite pigmentation; dorsum variable, but never with several transverse
dorsal shields; adults with a genital pore far larger than the anal opening, “window shutter”-like
plates may be present flanking the gonopore, but never the anal pore (2-4 f); if nymphs have ap-
pendages with dark colour, sclerotized structures similar in colour are generally found on the idio-
soma. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Idiosoma dorsally covered with a single hemispherical shield (2-5 c), empodium consisting of one
to five adhesive hairs (2-5 b); palp tibia with claw; trichobothria and genital acetabula absent on
idiosoma, body of red coloration, specimens often found floating on water surface. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homocaligidae (Eleutherengona)
(restricted to aquatic habitats, in Central Europe only Homocaligus scapularis (C. L. K OCH ,
1838) , not treated in this work)
– Idiosoma dorsally without a single hemispherical shield, empodium absent or claw-like; palp tibia
with or without claw; trichobothria and genital acetabula present or absent. . . . . . . . . . . . 6
6 Palp tarsus distally with two conspicuous, long setae, gnathosoma snout-like (2-5 a); chelicerae
elongate, distally with small movable digit; palp tibia without claw; two pairs of trichobothria;
reddish body coloration, active at terrestrial conditions. . . . . . . . . . . . Bdellidae (Eupodina)
(several species present in semi-aquatic habitats, not treated in this work)
– Palp tarsus various, but without two conspicuous, long setae distally, gnathosoma not snout-like;
chelicerae various; palp tibia with or without claw; trichobothria present or absent. . . . . . . 7
7 Leg and idiosoma surface with a dense cover of setae („hypertrichous“, 2-4 e); osmoregulatory
organs („acetabula“) present or absent; if present always arranged in the genital area, at most three
pairs; idiosoma normally with a soft integument, often red in colour; legs always lacking swim-
ming setae. . . . . . . . . . . . . . . . . . . . . . . . Trombidia (terrestrial Parasitengona) (page 158)
– Leg and idiosoma surface generally with setae arranged in distinct patterns (2-4 d, f, g, i); idiosoma
never densely covered with setae; osmoregulatory organs, integument and colour various; legs may
bear swimming setae (2-4 h). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Palp bearing at most four movable segments (5-1 c); osmoregulatory organs („acetabula“) in three
pairs in the genital field, minute, knob-shaped; two pairs of legs directed anteriorly and two pairs
posteriorly (2-4 d); mostly minute in dimension and with little apparent coloration, idiosoma in
general with four dorsal and four ventral plates in a cruciform arrangement (5-1 a-b, but fusion of
these plates possible: 2-4 d); without regularly arranged idiosomal glands associated with sensillae;
legs without swimming hairs (vs. fine hairs extending in length over more than 50 % of the next,
following segment).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halacaroidea (page 113)
– Palp typically with five segments (7-14 - see there 7-14 f for a striking exception); osmoregulatory
organs („acetabula“) in several taxa not restricted to the genital field, often enlarged or numerous
c
a
Fig. 2-5: Mites occasionally found in water: a, Bdellidae, Eupodina (Molgus sp. - details: tips of leg,
palp and chelicera); b-c, Homocaligidae, Eleutherengona (Homocaligus scapularis), b, dorsal view.
(detail: leg claws with adhesive hairs), c, lateral view.
32 References
(7-15); in general, legs in a circular arrangement or restricted to the anteriormost part of the idio-
soma (2-4 f- g, i - exceptions are found in members of the Hydrovolzioidea, 7-12 c, with leg ar-
rangement similar to Halacaroidea); dimensions, coloration, and idiosoma sclerites variable, with
numerous pairs of regularly arranged idiosomal glands associated with sensillae (7-1); legs varia-
ble in shape, in many taxa with swimming setae (2-4 h). . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachnidia (aquatic Parasitengona) (page 241)
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V IETS , K. (1936): Wassermilben oder Hydracarina (Hydrachnellae und Halacaridae). In: D AHL , F.
(ed.), Tierwelt Deutschlands 31: 1-288; 32: 289-574. G. Fischer, Jena.
V IGNA TAGLIANTI , A., C OTTARELLI , V. & A RGANO , R. (1969): Messa a punto di metodiche per la
raccolta della fauna interstiziale e freatica. Archivio Botanico e Biogeografico Italiano 45 (4):
375-380.
General Glossary Acari

(further terms used only in particular groups of mites are listed in the glossaries of the respective
chapters)
acetabulum Osmoregulatory organ of actinotrichid mites: Plesiotypically situated in the genital
field, three pairs in adults, two pairs in nymphs, in larvae and some juvenile
Oribatida only one pair (called „urstigma“ or „Claparède organ“); in larvae of Hy-
drachnidia and in all stages of Acaridida in the interspace between leg I and II;
secondarily, acetabula can be numerous and spread all over the idiosoma (= genital
papillae, [German] Genitalnapf)
ambulacrum clinging organ at the tips of legs, made up of an unpaired (often reduced) empodium
and paired or unpaired claws; claws may be provided with clawlets (claw-like sub-
divisions of claws) and a claw blade (knife-like extension of the inner claw surface);
in some taxa, the empodium may be claw like and occasionally the only functional
claw.
anal opening posterior opening of the digestive system. The rectum in some mites may be sepa-
rated from the anterior intestine and then functions as an excretory system, often
also called „anal pore“ or „excretory pore“. (= anus, [German] Exkretionsporus)
anus see „anal opening“
apodeme internal extension of a cuticular sclerite, in general functioning as muscle attach-
ment site
apoderma see „resting stage“
calyptostase see „resting stage“
camerostome see „gnathosoma“
capitulum see „gnathosoma“
34 Glossary
chelicera appendage of the first prosomal segment of Chelicerata, in mites at least

basally embedded into the gnathosoma; plesiotypically three-segmented,
with the two distal segments acting as scissors („chelate“), but in many
taxa two-segmented, with the third segment reduced to a membranous
extension, or all segments fused to a stilleto-like penetrating appendage
(= [err.] „maxilla“)
Claparède organ see „acetabulum“
claw, clawlet, claw blade see „ambulacrum“
condyle heavily sclerotized articulation point between leg segments
cornea see „lateral eyes“
coxa basal leg segment, in all mite taxa treated here (Actinotrichida) fused to
the idiosoma surface and functioning as a basal plate on which the tro-
chanter inserts as the first movable leg segment; neighbouring coxae
may fuse and form „coxal plates“ („epimeral plates“), or all coxae
may be fused into a continuous (!) „ventral shield“ (= „epimeron“)
coxal plate see „coxa“
cuticle as in all arthropods, consisting of an internal procuticula with „pore chan-
nels“ (rarely differentiated in endo- and exocuticula), and an external epi-
cuticula; a cerotegument (cement- or wax layer hampering dehydration) is
present in representatives of terrestrial mite groups - probably for this rea-
son, aquatic Oribatida evolved particular respiratory surfaces (singular:
„plastron“); in many Hydrachnidia, when pore channels become enlarged
in order to allow respiratory activity of tracheoles, the procuticula has a
reticular structure
dorsal furrow see „dorsal shield“
dorsal shield large sclerotized shield covering the whole dorsal idiosoma; in most cases
separated from the (!) „ventral shield“ by a ring shaped membranous
area called „dorsal furrow“
epimeron see „coxae“
excretory pore see „anal opening“
eye capsule see „lateral eyes“
femur see „leg“, „palp“
frontale plesiotypically a sclerotized ring in the anterodorsal area of idiosoma sur-
rounding an unpaired, weakly pigmented „frontal eye“ (occasionally
called „median eye“ or „frontal organ“); in many clades, this eye being
reduced, a more or less extended sclerite („frontal plate“, „frontal shield“)
is formed in this area, occasionally including also surrounding (!) dor-
salia.
frontal eye, fr. organ see „frontale“
frontal plate, fr. shield see „frontale“
genital bay see „genital field“
genital field area surrounding the opening of the reproductive system („gonopore“),
plesiotypically on the anterior ventral surface of the opisthosoma, with
anterior („pregenital“) and posterior („postgenital“) sclerites delimiting
the slit-shaped gonopore; occasionally, a depressed area surrounding
the male gonopore is called „genital pit“
genu see „leg“, „palp“
glandularium defence organ typical for, and a synapomorphy of, Hydrachnidia (see
there)
gnathosoma anterior body section, formed by the fusion of palp coxae, an unpaired
sternal sclerite, and a dorsal extension, bearing as appendages the cheli-
cerae and palps; partly, a caudal extension of the gnathosomal base, ex-
tending medially between the anterior coxal plates is called „anchoral pro-
cess“; the bay formed by the medial margins of first coxae which embrace
the gnathosomal base is called „gnathosomal bay“; if the attachment site
for the gnathosomal base at the anterior opening of the idiosoma is formed
as a collar-like sclerotized ring or tube, it is called „camerostome“; if the
anterior part of the gnathosoma (anterior from the palp insertions) is
narrowed and projecting, it is called „rostrum“. (= capitulum; infracapi-
tulum [referring to the gnathosoma without appendages]; [German, err.]
Maxillarorgan)
gnathosomal bay see „gnathosoma“
gonopore see „genital field“
hypostome see „infracapitulum“
hysterosoma posterior secondary body section, probably formed by segments of legs III
and IV and the opisthosoma (in Oribatida often covered with (!) noto-
gaster plate); anterior body section (!) proterosoma
idiosoma body section posterior to gnathosoma, including segments of legs and all
segments following caudally, as well as elements belonging to the mouth
part-bearing segments I and II (! „frontal eye, lateral eyes“)
infracapitulum referring to the gnathosomal base (= gnathosoma without appendages);
the sclerotized ventral part of this structure is called „subcapitulum“,
its median longitudinal channel „hypostome“
integument see „cuticle“
lateral eyes paired light sensitive organs at the anterior edge of the idiosoma, generally
consisting of paired lenses („corneae“) and retinulae, often enclosed in a
sclerotized eye capsule
lateralia see „plate“
leg appendage of podosomal segments, often numbered „I-L, II-L, III-L,
IV-L“, in most mite species treated here, consisting of a coxal plate fused
to the idiosomal surface and originally six movable segments („trochanter,
basifemur, telofemur, genu, tibia, tarsus“, often numbered: I-IV-L-1-6);
tarsus with (!) „ambulacrum“; water mites with particular adaptations
for swimming bear „swimming setae“ (see chapter Hydrachnidia)
lyrifissure see „slit organ“
median eye see „frontale“
metapodosoma the posterior part of the podosoma, bearing legs III and IV, separated from
the propodosoma, by the (!) „transversal furrow“
notogaster plate dorsal shield of posterior body part in Oribatida
opisthosoma in the ground plan of chelicerata a posterior tagma behind the last leg-
bearing segment of the “prosoma”; in mites secondarily fused with the
four leg-bearing segments of the (!) „prosoma“ forming the (!) „idio-
soma“
palp appendage of segment II of all chelicerata, generally involved in the uptake
of food (grasping, tasting), occasionally also in spermatophore deposition
or transfer; in actinotrichid mites plesiotypically consisting of five free seg-
ments („trochanter, femur, genu, tibia, tarsus“, often simply numbered as
„P-1“ to „P-5“); pincer-like palps are called „chelate“ if P-5 is acting
against a pointed dorsal projection or „antagonistic bristle“ of P-4, or
„subchelate“ (= „uncate“) if P-5 is acting against a ventrodistal projection
of P-4. (= pedipalp)
pedipalp see „palp“
36 Glossary
pharynx anterior, dilatable part of the digestive tract, in mites located in the gnatho-
soma
plastron non-compressible thin air layer on hydrophobic surface areas (! respir-
atory system)
plate idiosoma sclerite of minor (then sometimes called „platelets“) or medium
dimensions, generally functioning as muscle attachment sites and plesio-
typically arranged in a characteristic pattern reflecting the original body
segmentation („dorsalia“, lateralia, (!) „frontale“); if numerous plates
fuse to larger structures, covering extended parts of the body, they are
called „shields“
podosoma body area of the leg-bearing segments
pore channels see „cuticle“
prodorsum anterior dorsal part of idiosoma, probably the dorsal plate of the „pro-
podosoma“ with legs I and II
propodosoma the anterior part of the podosoma, bearing legs I and II (posterior part of
the podosoma: metapodosoma)
prosoma in the ground plan of chelicerata a tagma including segments I to VI
(mouth part and leg segments); in mites its anterior part forming a sepa-
rate body division (!) „gnathosoma“, its posterior part including the seg-
ments of the legs („podosoma“) may be fused with the (!) „opisthoso-
ma“ forming the (!) „idiosoma“
proterosoma anterior secondary body section, probably formed by (!) gnathosoma
and segments of legs I and II (in Oribatida often covered with (!) pro-
dorsum); posterior body section (!) hysterosoma
resting stage inactive prelarva („schadonophan“) of all mites, and protonymph („nym-
phophan“) and tritonymph („teleiophan“) of Parasitengona; the term
„apoderma“ is used in such quiescent stages for the internal cuticle,
not exposed to the surrounding environment. (= „calyptostase“)
respiratory system in mites consisting of tracheae and tracheoles plesiotypically communicat-
ing with the environment via „stigmata“; in water dwelling or hygric
mites, these stigmata are closed and the unsclerotized parts of integument
take over respiratory function, often facilitated by porosity of sclerites and
development of fine peripheral „tracheoles“; occasionally completely re-
duced
rostrum a narrowed anterior projection, (1) of the (!) „gnathosoma“ (apically
bearing the mouth opening); or (2) of the (!) „prodorsum“ (in Oribatida,
dorsally covering the gnathosomal base)
shield see „plate“, „dorsal shield“, „ventral shield“
slit organ mechanoreceptive organ typically found in all arachnids, consisting of an
innervated slit-shaped opening in the sclerotized cuticle; organs composed
of several slits are called „lyriform organs“ or „lyrifissures“, a term some-
times incorrectly used also for the single slits found in mites; in the ground
pattern of Hydrachnidia, four pairs are found on the dorsum, and one in
the posteroventral region; in other mite groups, different patterns are
found
stigma see „respiratory system“
subcapitulum see „infracapitulum“
suture strongly sclerotized line visible on a plate or shield originating from the
fusion of several separate sclerotized elements, indicating their former
contours
tarsus see „leg“, „palp“
tibia see „leg“, „palp“
tracheae, tracheoles see „respiratory apparatus“

transversal furrow area between the segments of leg-II and leg-III, often distinctly separating
the anterior leg segments („propodosoma“) from the posterior ones („meta-
podosoma“) by a membranous intersegmental zone
trochanter see „leg“, „palp“
urstigma see „acetabulum“
ventral shield a continuous sclerotized plate including coxae, genital field, and ventralia,
occasionally also the anal opening; in Hydrachnidia generally found in com-
bination with a (!) dorsal shield. In higher Oribatida, this term is used for
a plate surrounding the anal and genital openings, generally fused with the
coxae
38 General part
3. Acari: Limnic Acaridida (Astigmata)

Eberhard Wurst
General part
Diagnosis
Mostly weakly sclerotized non-predatory mites with size range (adults) from 200 lm to
1800 lm. Movable coxae lacking in all instars; palps with maximally two movable po-
domeres; chelicerae in most families chelate, never sword-like or styletto-like; oviporus
and copulatory opening two spatially separate body openings; males always with aedea-
gus; regularly two oil gland openings dorsally; life cycle comprising: egg (vivipary occurs),
larva, protonymph, deutonymph (if present mostly facultative), tritonymph, adults; deu-
tonymph missing (e. g. in all stationary parasites) or modified into an instar specialized for
phoresy.
Fig. 3-1: a, idiosoma (dorsum) of a female acarid mite. OG = oil gland opening, PS = propodosomal
shield. All other symbols designate setae and cupules; b, venter of a female acarid mite (legs partly
omitted). AS = anal slit, OP = oviporus. All other symbols designate setae and cupules.

https://doi.org/10.1007/978-3-662-55958-1_3
3. Acari: Limnic Acaridida (Astigmata) 39
Introduction
The suborder Acaridida (also Astigmata) contains a large number of mites with represen-
tatives of all ecological adaptations from free-living to parasitic. An estimated number of
species is approximately 1000 for central Europe. Most species are rather small, with an
idiosoma length below 500 lm. Many species gained medical and economical importance
e. g. as pathogens for humans and animals, as storage pests, and as important compounds in
the decomposition process of organic material. As the name indicates the Astigmata lack
stigmata and a tracheal system. Other striking characters include: (1) Lack of the coxa
(according to traditional theories the coxae have fused with the ventral body wall, which
led to the term “coxal field” meaning the ventral parts of the body wall, that are more or
less completely encompassed by the leg apodemes); (2) the males possess an aedeagus; (3) a
single claw at the praetarsus (if not completely reduced as in most stationary parasites). (4)
the palps have maximally two movable podomeres and the chelicerae are chelate (most
important exception: Histiostomatidae).
The habitus of the Acaridida resembles very much the habitus of the Oribatida (especially
the juveniles of both taxa are very similar), but Acaridida never have trichobothria. Most
Acaridida are only weakly sclerotized, which gives them a whitish translucent appearance.
On the idiosoma well defined areas of sclerotized cuticle (=shields) regularly occur dorsally
behind the gnathosoma (propodosomal shield) (3-1 a), at the opisthosomal end of both
sexes (opisthosomal shield) (3-5 a), and ventrally on the coxal fields. The degree of
body sclerotization and the size of the shields increase during ontogenesis.
Sperm transfer is direct i. e. the males insert their aedeagus in a specific body opening (co-
pulatory opening) of the female which leads to a spermatheca that has connections with the
paired ovaries (3-7 b, 3-11 b, 3-19 b, 3-25 b). The copulatory opening is situated mostly at
the caudal end of the opisthosoma (3-5 b), but can shift to the opisthosomal dorsum (e. g.
among Histiostomatidae). The area around the copulatory opening is called bursa copu-
latrix and may possess cuticular sculpturing for the attachment of the aedeagus. The co-
pulatory opening is spatially separate from the oviporus that opens ventrally between leg
pairs III and IV (3-1 b, 3-7 a). Copulation may occur at two different positions that are
species-specific: (1) in the retroconjugate mode the sexes point in opposite directions the
male riding on the female or at least placing its opisthosoma on the female dorsum. (2) In
the proconjugate mode the male riding on the female is oriented like the female partner.
This mating behaviour led to remarkable sexual dimorphism in Acaridida. In species that
copulate in the retroconjugate mode males possess two adhesive discs (3-4) lateral to the
anal slit that connect with the female dorsal cuticle. Males of proconjugate species often
have enlaged legs I and II that help in clasping the partner. Praecopulatory guarding of
female tritonymphs by the males is widespread. In these cases the respective position during
copulation is already applied to the female tritonymph. In cases with praecopulatory guard-
ing the copulation with the newly hatched female is a rather short procedure and is rarely
observed.
Most free-living Acaridida are saprophagous, some are specialized fungivores. Most spe-
cies are confined to humid to wet or aquatic environments. Two different organs related to
osmoregulation and water uptake occur in the Acaridida: the genital papillae and Clapa-
rède organs, respectively, and the supracoxal glands. The genital papillae of adults and
nymphs are bulbous to fingerlike cuticular protuberances that are hidden under the genital
folds (3-4, 3-6 b, c, 3-7 a). They are filled with the extensions of one or several cells. The
cells contain an extensive network of vesicular tubules or numerous vesicles as well as nu-
merous mitochondria and their apical plasmalemma is deeply folded. Apically the cells are
covered by a cap of specialized cuticle. Ultrastructurally very similar to the genital papillae
are the larval Claparède organs which originate between leg I and II and frequently possess
40 General part
Fig. 3-2:
Nomenclature of leg chaetae in Acaridida
The nomenclature was developed by G RANDJEAN (1939). It is a system that is based on the (relative)
position of chaetae.
Setae
The leg setation is very similar for all legs. The G RANDJEAN theory combines particular setae in groups
in which the setae are arranged in transversal rows.
Tarsus I and II (3-2 a-b):
dorsal apically: row f (medial) – d (central) – e (lateral)
dorsolateral: aa (only in leg I)
dorsomedial: ba
ventral apically: row (beginning with the most medial seta) q – v – s (central) – u – p
ventral in the middle la (medial) – wa (central) – ra (lateral)
Tibia I and II (3-2 a-b):
ventral distally: row gT (medial) – hT (lateral)
Genu I and II (3-2 a-b):
dorsomedial at basis: cG
ventrolateral: mG
Femur I and II (3-2 a-b):
ventral: vF
Trochanter I and II (3-2 a-b):
ventromedial: pR
The legs III and IV (3-2 c-d) “emerge” in the G RANDJEAN theory from legs I and II by rotation of 180
degree; medial becomes lateral and vice versa.
Modifications compared with legs I and II are:
Setae ba and aa always lacking.
The row la, wa, ra of legs I and II comprises only two setae (r, w).
Tibia: ventrolaterally: kT
Genu: ventrolaterally: nG (only in leg III)
Femur: ventrally: wF (only in leg IV)
Trochanter: ventrolaterally: sR (only in leg III)
Solenidia and famulus

Tarsus I and II (3-2 a-b):
All solenidia are dorsal. The most prominent solenidion x1 is situated in the proximal half of the tar-
sus, solenidion x2 is lateral, and solenidion x3 is apical. The famulus is very near to solenidion x1. On
tarsus II only solenidion x1, no famulus e.
Tibia I–IV (3-2 a-d):
Solenidion u distally in the middle of tibia.
Genu I–III (3-2 a-d):
Solenidia r1,2 apically in the middle of genu (in genu II and III only solenidion r1).
Appearance of chaetae in ontogeny

From larva to deutonymph the number of solenidia x on tarsus I increases from 1 to 3.
In tarsus IV of protonymph setae e, f, s are lacking. Tibia, genu, femur, of the same leg without setae. All
trochanters in protonymph and larva lack setae.
Rules for modifications of leg chaetome

Chaetae preferably shift in axial direction.
The following chaetae are preferably reduced: aa, r2, nG, u, v.
42 Chaetotaxy
a long stalk. With the genital papillae/Claparède organs the mites take up fluid water but
they also appear to play an important role in osmoregulation (FASHING 1988; W ITALINSKI
et al. 1990). In the non-deutonymphal instars of histiostomatids the genital papillae/Cla-
parède organs are modified into flat disc-like structures (ring organs, 3-34) that are situated
uncovered on the venter. The ring organ cells possess deep infoldings basally and contain
many mitochondria (W ITALINSKI et al. 2002). The ring organ size varies among histiosto-
matid species and can intraspecifically be adapted to the osmolarity of the environment.
The supracoxal glands open dorsally to trochanter I. They consist of a branched cuticular
invagination ending in large cells densely filled with mitochondria and membrane infold-
ings. The supracoxal gland produces a brane that flows to the mouth opening. Depending
on the relative humidity of the ambient air, water vapour condenses into the brane (W HAR-
TON & F URUMIZO 1977, W URST 1993). Genital papillae/Claparède organs are reduced in
all species living in environments with little or no access to fluid water whereas the supra-
coxal glands apparently are lost in all aquatic species.
Chaetotaxy
For the determination of species and their developmental stages, the configuration of idio-
somal and leg chaetae is essential (G RANDJEAN 1939, G RIFFITHS et al. 1990). In the Acar-
idida the set of chaetae (=chaetome) shows limited variation i. e. there exists a maximal
number of chaetae that is modified in the different taxa (by reduction or shift of chaetae).
Up to now three types of chaetae have been found in Acaridida: (1) Solenidia: these are
chemosensitive chaetae which occur only on the legs and palps. They are usually stout
to bulbous chaetae, if they are long and slender they can be distinguished from the setae
(see below) by their blunt tip. The surface structure often is striated or punctured (3-3 c).
Besides shape and position the solenidia can be distinguished from the other chaetae by
their interaction with polarized light: the solenidia are the only chaetae lacking birefin-
gence. (2) Famulus: a chemosensitive chaeta whose shape can vary considerably. It occurs
only on tarsus I and is always close to solenidion x1 (3-3 c). (3) Setae: these are mechan-
osensitive chaetae which can be filiform, spine-like or more or less foliate. Setae are indi-
cated by Latin letters, solenidia and famuli by Greek letters. Four pairs of easily visible
cuticular strain detectors (= cupules ia, im, ip, ih) occur on the idiosoma. The nomenclature
for idiosomal setae was developed by G RIFFITHS et al. (1990) and is nowadays widely ac-
cepted. The idea behind the nomenclature is that the idiosoma consists of a series of body
segments that bear the setae (e. g. c, d, e, ps, ad). Schematic representations of the general
arrangement of chaetae is given in Figs 3-1 and 3-2.
Fig. 3-3: a, Schwiebea talpa, female; proterosoma, frontal view. ba = seta on tarsus I, C = chelicera, cG
3
= seta on genu I, Cl = claw, d = seta on tarsus I, d2 = seta on proximal palp podomere, e = seta on tarsus I,
om1 = solenidion x1 on distal palp podomere, P = palp, ph = solenidion u on tibia I, si = solenidion r on
genu I (SEM). Scale bar: 20 lm; b, Schwiebea talpa, tritonymph, gnathosoma, ventral view. Inset: right
distal palp podomere (detail). Cl = claw, d1 = seta on distal palp podomere, d2 = seta on proximal palp
podomere, e = seta on tarsus I, om1 = solenidion x1 on distal palp podomere, ph = solenidion u on tibia
I, si = solenidion r on genu I, subc = subcapitular seta, v = seta on proximal palp podomere, 1, 2, 3 =
solenidia on distal palp podomere (SEM). Scale bar: 10 lm; c, Schwiebea eurynympha, female, leg I,
tarsus. ep = famulus e, ba = seta, om1,2 = solenidion x1,2 (SEM). Scale bar: 5 lm; d, Schwiebea eur-
ynympha, deutonymph, so-called sucker plate at ventral hind end. Asterisk = soft walled rim that en-
circles the adhesive plate, AS = anal slit, g = seta, GO = genital opening, 4a = seta (SEM). Scale bar: 10
lm; e, Hormosianeotus sp., female, gnathosoma, dorsal view. C = chelicera, G = gnathosoma, om1,2 =
solenidia on distal palp podomere, vi = seta, arrowhead = tip of chelicerae (SEM). Scale bar: 20 lm; f,
Hormosianeotus sp., female, gnathosoma, dorsolateral view. C = chelicera, d = seta d1 on distal palp
podomere, om1,2 = solenidia on distal palp podomere, vi = seta, 1, 2, 3 = solenidia on distal palp po-
domere, arrow = detail shown in inset, arrowhead = tip of chelicerae (SEM). Scale bar: 10 lm.
a b
c d
e f
44 Chaetotaxy
Ontogeny
The life cycle of acaridid mites includes: egg, larva, two or three nymphs, and adult. The
second nymph (deutonymph) is an instar specialized for phoresy, its occurrence in the life
cycle is normally facultative. The larva, protonymph, and tritonymph closely resemble the
adult female. The life stages can easily be identified by leg and body chaetome and the
occurrence of particular organs:
L a rva : Three pairs of legs; only one solenidion on tarsus I.
Pr oto ny m ph: Four pairs of legs (as all following instars); one pair of genital papillae in a
cuticular slit between legs IV; two solenidia on tarsus I.
De ut ony m ph : Morphologically completely different from other instars; two pairs of gen-
ital papillae in a cuticular slit between legs IV; three solenidia on tarsus I.
Tri to ny m ph: Two pairs of genital papillae in a cuticular slit between legs IV; three so-
lenidia on tarsus I.
A dul t: External sexual organs (oviporus, copulatory opening, bursa copulatrix, aedea-
gus); two pairs of genital papillae under the genital folds; three solenidia on tarsus I.
Due to its phoretic role the deutonymph deviates extremely in morphology from the other
life stages which has led to the special term “hypopus” (3-13, 3-14 b, 3-20, 3-26, 3-31, 3-35
c). It is dorsoventrally flattened and more strongly sclerotized (especially the dorsum) than
the other instars. Instead of a gnathosoma and a mouth opening the deutonymph bears a
short movable rod bearing solenidia and setae („palposoma“) (3-13), but even this rod
might be lacking in some species. Ventrally at the hind end an attachment organ occurs
its form depending on the transport host which is used by the species: if arthropods are
used for transportation, a so-called sucker plate occurs which consists of a soft-walled cu-
ticular rim surrounding setae modified into flat discs or bulbs („conoids“) (3-3 d). In mam-
mal-associated nidicoles an ensemble of plates and modified setae enables the hypopus to
clasp to the host hairs. Under favourable living conditions the mite develops directly from
the protonymph into a tritonymph, adverse environmental factors (desiccation, food de-
pletion, high population densities) and genetic disposition trigger the formation of the deu-
tonymph („facultative hypopody“). The deutonymph stage is lost in some free-living spe-
cies and in all stationary parasites.
Among the Acaridida only a few taxa can be regarded as truly aquatic, living permanently
completely submerged in water. Some taxa (especially the Histiostomatidae) may occur
both in wet and aquatic habitat. Fresh water mites are only found in the families Acaridae
(Schwiebea, Naiadacarus) and Histiostomatidae. While Histiostomatidae and Naiada-
carus have been so far recorded only from surface water, various species of Schwiebea
have in addition been found in samples of ground water.
Special glossary
aedeagus penis
bursa copulatrix the area around the copulatory opening of female Acaridida; it may pos-
sess cuticular structures or special sculpturing for the attachment of the
aedeagus.
conoid seta with more or less spherical shape.
ductus conjunctivus a duct occuring always pairwise and connecting the receptaculum seminis
at each side with one ovary; lined with cuticle; in Acaridida.
epimerite the leg apodeme that originates posterior to the leg insertion.
facultative hypopody the deutonymph (=hypopus) may lack in the life cycle of an individual
acaridid mite. In Acaridida the deutonymph serves for dispersal by
phoresy; its formation is induced by a mixture of environmental and ge-

netic factors.
famulus a chemosensory chaeta occuring in Actinotrichida.
hypopus the heteromorphic deutonymph in Acaridida (see facultative hypopody);
originally a genus name.
inseminatory canal the duct connecting the copulatory opening and the receptaculum seminis
in Acaridida; always lined with cuticle.
palposoma a rodlike movable appendage occuring instead the gnathosoma in Acar-
idid deutonymphs; it bears two solenidia and setae.
praecopulatory guarding immature females are clasped by males until the female hatches and co-
pulation occurs.
proconjugate mating sexes are oriented in the same sense; in Acaridida.
retroconjugate mating sexes point at opposite directions; in Acaridida.
solenidion a chemosensory chaeta occuring in Actinotrichida.
supracoxal gland gland that serves in water supply in Acaridida; it opens dorsal to trochan-
ter I and secretes a brane that flows on the cuticular surface to the mouth
opening; depending on the relative humidity of the ambient air, water va-
pour condenses into the brane on its way to the mouth.
Special part
Area treated
The following key includes taxa occuring in central Europe (Belgium, The Netherlands, Germany,
Switzerland, Austria, Italian Alps, Czechia, Slowakia, Poland) which have been recorded from aquatic
habitats.
Key to the adults of limnic Acaridida

1 Chelicerae chelate (3-3 a, b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acaridae 2
– Chelicerae non-chelate and shaped like a paddle or a rake (3-3 e, f) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Histiostomatidae (page 55)
2 Legs longer than 50 % of idiosomal length (3-33 a, b), dorsal surface clearly corrugated or mod-
erately wrinkled; in water filled tree holes . . . . . . . . . . . . . . . . . . . . Naiadacarus (page 55)
– Legs shorter than 50 % of idiosomal length, dorsal surface smooth; in ground water (Schwie-
bea). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 With one solenidion on genu I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– With two solenidia on genu I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 Idiosoma length 375 – 660 lm; propodosomal shield with incision at hind margin; solenidion x1 of
leg I club shaped with bulbous tip (3-3 c); seta nG of leg III present . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schwiebea eurynympha (page 48)
– Idiosoma length 270 – 355 lm; propodosomal shield without incision at hind margin; solenidion
x1 of leg I club shaped, gradually tapering towards its base; seta nG of leg III missing . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schwiebea talpa (page 46)
5 Seta nG of leg III present; sac of receptaculum seminis wrinkled (3-25 b) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schwiebea cavernicola (page 53)
– Seta nG of leg III missing; sac of receptaculum seminis different . . . . . . . . . . . . . . . . . . . . 6
6 Seta la of tarsus I and II long and thin (3-30 b) . . . . . . . . . Schwiebea-nova-group (page 55)
– Seta la of tarsus I and II spinelike. . . . . . . . . . . . . . . . . . . Schwiebea schmuttereri (page 50)
46 Special part
Family Acaridae L ATREILLE , 1802

Genus Schwiebea O UDEMANS , 1916
1916 Schwiebea O UDEMANS, Ent. Ber., Amst. 4 (88): 264.
1936 Megninietta J ACOT, Annls Ent. Soc. Amer. 29: 631.
1941 Troupeauia Z ACHVATKIN, Fauna of U.S.S.R. Arachnoidea, 6, 1: 298.
Typus generis: Schwiebea talpa O UDEMANS , 1916, by monotypy.
Diagnosis
Retroconjugate mites with facultative hypopody; leg length of non-hypopal instars less
than 50 % of idiosomal length. Id io som a l c ha e tom e : seta scx of non-hypopal instars
a short spine; female: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b, g, 4a; male: as given for
female but additionally ps1, ps2, ps3; the following deviations of this configuration of char-
acters were found in some species: occurrence of ps3 also in females, tritonymphs and pro-
tonymphs; lack of e1 and d2; in individuals of one lineage of generations d1 as well as c1 and
d1 can occur. L eg c ha et om e (non-hypopal instars) (3-3 c, 3-8): Seta aa always lacking;
setae d, f, ra long and thin, never clearly lanceolate (exception: the seta d in the male leg IV
that is modified into an attachment structure, 3-8 e); in species related to S. nova seta la is
long and thin; seta e a strong spine (exception: the seta e in the male leg IV that is modified
into an attachment structure, 3-8 e); seta ba spinelike closely situated to solenidion x1
(3-3 c).
Mites of the genus Schwiebea do not conspicuously deviate in morphology and behaviour
from other genera of the Acaridae. Constitutive for the genus is mainly the idiosomal chae-
tome of the non-hypopal instars i. e. a particular pattern of presence and form of idiosomal
setae. Males are polymorpic. They may possess strongly enlarged legs III with strong claw
(heteromorphic males) or their legs III may be similar to the female (homeomorphic males).
Among the heteromorphic males variability exists with respect to the relative size of leg III
and the proportions of its parts. The two extremes are termed weak-legged and strong-
legged heteromorphic males (W URST 2002).
Most species of the genus Schwiebea live in rotting plant material, typically under the bark
of dead wood, where they feed on the decaying plant tissue and fungal materials (hyphae,
spores). In some rare cases Schwiebea species have also been found at tulip bulbs, in sap or
in the decaying fruit bodies of higher fungi (W URST 2002), or parasitizing in the skin, gills,
and gut of trouts (Salmo trutta fario) (FAIN & F ERRANDO 1990). Also in some aquatic
situations Schwiebea species were detected: swimming pool, aquarium, aqueduct, and
in ground water (FAIN 1977, FAIN & PAGANI 1989, O KABE & O’C ONNOR 2000). In
his revision of the genus Schwiebea W URST (2002), however, showed that all the species
found in the ground water in central Europe also occur epigaeic in decaying plant materials.
1. Schwiebea talpa O UDEMANS , 1916 (3-3 a-b, 3-4–3-9)

1916 Schwiebea talpa O UDEMANS, Ent. Ber., Amst. 4 (88): 265.
1957 Schwiebea cavernicola T ÜRK & T ÜRK, In: S TAMMER , H.-J. (ed.), Beiträge zur Systematik und
Ökologie mitteleuropäischer Acarina, 1: 126–129 (erroneous attribution).
Gn ath oso m a of acarid type; terminal palp podomere with two button-like and one short
rod-shaped (x1) solenidion and one short dorsal seta d1; proximal palp podomere with one
short dorsal seta d2 and one short ventral seta v; palp coxa with ramified laterocoxal seta;
two long filiform subcapitular setae (subc) ventrally (3-3 a, b).
A dul ts:
Ma le (h om e om or phi c ): Length of idiosoma approximately 270 lm; body form ovoid;

colour: beige; idiosomal chaetome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b, 4a, g, ps1,
ps2 , ps3, ia, im, ip, ih; all idiosomal setae smooth. Dor sum (3-5 a): Propodosomal shield
without indentation at posterior margin; setae e1 and d2 short (approximately 10 lm); with
opisthosomal shield between h1; anterior margin of opisthosomal shield convex and irre-
gularly corrugated. Ve nte r (3-4): Epimeres I medially fused into a sternum; sternum and
epimeres II and IV covered by lobed sclerotized areas, epimeres III only in their distal region
covered by a sclerotized patch; on each side a sclerotized band running along the border
between proterosoma and hysterosoma; at its posterior margin each band connected proxi-
mally with a shorter narrow apodeme running transversally behind the larger one; aedea-
gus strongly sclerotized, hidden under genital folds forming a triangle (3-6); one pair of
adanal copulatory dics; setae ps3 short and fine, ps1 and ps2 short spines. L e gs (compare
with 3-8 a-e): Leg I: Solenidion x1 club shaped gradually tapering towards its base; sole-
nidion x2 rod shaped and slender inserting in close vicinity to x1; solenidion x3 rod shaped
inserting anterior to seta f; famulus e button-like inserting in close vicinity to x1 and x2;
solenidion u long and rod shaped tapering distally; solenidion r rod shaped approximately
as long as genu I; seta ba spiniform inserting near the complex of x1, x2, and e; setae u, v, p,
q, s, wa, la short spines; setae e, hT, gT, cG spiniform; seta mG short and thin; setae d, f, ra
long and thin, with weakly broadened distal end; setae vF and pR filiform; leg II: Solenidion
r rod shaped, shorter than genu. Except for the regularly lacking chaetae (x2, x3, e) as given
for leg I; leg III: Solenidion u rod shaped, longer than tibia III; solenidion r rod shaped
shorter than genu III; setae d, f, r, long and thin; setae e and kT strong spines; setae u,
v, p, q, s, w short spines; seta sR short and filiform; leg IV (3-8 e): Solenidion u rod shaped
approximately as long as tibia, but clearly thicker than solenidion u of tibia III; seta f long
and thin; setae u, v, p, q, s, r, w, kT spiniform; seta wF short and thin.
Fe m a le : Length of idiosoma 300 – 355 lm; body ovoid, elongate; colour: beige; idiosomal
chaetome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b, 4a, g, ia, im, ip, ih. Dor sum (3-5 b):
propodosomal shield without indentation at posterior margin; setae d2 and e1 short (ap-
proximately 10 lm); copulatory opening (3-5 b) a bare pore without supporting structures
situated at opisthosomal end; receptaculum seminis (3-7 b) with bulb shaped basal part; at
the transition between basal part and main sac of receptaculum seminis two weakly conical
ductus conjunctivi lead into the receptaculum seminis; inseminatory canal short connecting
receptaculum seminis and copulatory opening on the shortest distance; copulatory opening
and receptaculum seminis with all appendages missing in some females. Ven te r (3-7 a):
Epimeres I medially fused into a sternum; sternum and epimeres II and IV covered by lobed
sclerotized areas, epimeres III only proximally and distally covered by small sclerotized
patches; on each side a sclerotized band running along the border between proterosoma
and hysterosoma; at its posterior margin each band connected proximally with a shorter
narrow apodeme running transversally behind the larger one; oviporus between legs III and
IV. Le g s (3-8 a-d): Legs I, II and III as given for the male; leg IV: Solenidion u rod shaped,
thin and shorter than tibia; setae d and f long and thin; setae e, u, v, p, q, s, r, w, kT spinelike;
seta wF short and thin.
De ut ony m ph : Unknown.
H a bit at : In a variety of substrates: under bark of rotten wood, in tree-stumps, leave litter,
sphagnum moss, humous soil, peat soil, ground water and mole dens (T ÜRK & T ÜRK 1957;
FAIN 1977, 1982). In the literature one finds many different actual or alleged habitats of S.
talpa. I am not able, however, to assess whether the species mentioned actually belong to S.
talpa. For a compilation of references, see FAIN (1977).
48 Special part
B io lo gy : Males are exceedingly rare in the central European populations (FAIN 1976,
1977; W URST 2002). The rare occurrence of males is accompanied by the absence of
the copulatory opening and all appendages (inseminatory canal, receptaculum seminis)
in numerous females of all populations. For food, this species apparently prefers faeces
of other consuments (e.g. larvae of mycetophilids) (W URST 2002, W URST & F RANK
1998). The development of deutonymphs in S. talpa was never observed despite extensive
rearing experiments (W URST 2002, W URST & F RANK 1998). FAIN (1977), after investigat-
ing material from several localities concludes that this species apparently lacks a hypopus.
In their work on S. cavernicola (= S. talpa), T ÜRK & T ÜRK 1957 describe a hypopus, yet
in the text they do not refer to rearing experiments which could justify such a reference.
B UGROV (1996) reports on the presence of adults of S. talpa under the elytra of a carabid
beetle (Carabus jankovskii). This finding, suggesting a phoretic behaviour, represents a
„pure accident for that species“ according to S. B UGROV (W URST 2002).
Distribution: Austria, Belgium, Finland, France, Germany, Hungaria, Poland, Romania,
Slovakia, Spain (Tenerife), Sweden.
2. Schwiebea eurynympha (O UDEMANS , 1911) (3-3 c-d, 3-9–3-15)

1911 Tyroglyphus eurynympha O UDEMANS, Ent. Ber., Amst. 3 (62): 189.
1923 Schwiebea talpa V ITZTHUM, Arch. Naturgesch. 89: 151–156 (erroneous attribution).
1932 Schwiebea ipidis V ITZTHUM, Zool. Jb. (Syst., Ökol., Geogr.) 63: 697.
1982 Schwiebea danielopoli FAIN, Acarologia 23 (4): 362–366.
Gn ath oso m a as given for S. talpa.
A dul ts:
Ma le ( he te ro m orph ic ): Length of idiosoma 425 – 600 lm; body elliptical, opisthoso-
mal end blunt; colour: white; idiosomal chaetome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a,
3b, 4a, g, ps1, ps2, ps3, ia, im, ip, ih; all idiosomal setae smooth. Dor sum (3-12 a-c): Pos-
terior margin of propodosomal shield with an incision of variable depth, ranging from a
short indentation to an incision that can reach the level of the sockets of setae ve (recogniz-
able by the small “hooks” at the lateral margin of propodosomal shield); the margin of the
incision is corrugated to denticulate. Opisthosomal shield convex with irregularly corru-
gated and denticulate margin, reaching approximately the level of setae e2 anteriorly; cu-
pules ip always situated outside the opisthosomal shield; setae d2 and e1 short and thin; seta
h1 inserting within the opisthosomal shield and shorter and thinner than seta h3 (examples
for the variability of the relative length of setae h1–3 in Figs. 3-12 a-c). Ve nt e r (3-9): Epi-
meres I fused medially forming a sternum; sternum and epimeres II covered by sclerotized
lobed areas; epimeres III and IV are accompanied by sclerotizations with an irregular pos-
terior border and broadening distally; apodemes III and IV never fused; on each side a
sclerotized band running along the border between proterosoma and hysterosoma; at
its posterior margin each band connected proximally with a shorter narrow apodeme run-
ning transversally behind the larger one; strongly sclerotized aedeagus hidden under genital
folds forming a triangle (3-10); one pair of adanal copulatory discs; a narrow sclerotized
band with irregular border accompanying the margin of the opisthosomal end between the
sockets of h2; it is the ventral continuation of the opisthosomal shield and encompasses the
setae h3 and ps1 but never reaches setae ps2 and the adanal copulatory discs; setae h2 and h3
inserting at the opisthosomal margin, h2 being shorter and thinner than h3; setae ps1–3 short
and thin. L e gs (3-14 a; for legs I, II, IV compare with 3-8 a-b, d-e): Leg I (compare with 3-3
c): Solenidion x1 club shaped, apically bulbous; solenidion x2 rod shaped and slender in-
serting in close vicinity to x1; solenidion x3 rod shaped inserting anterior to seta f; famulus e
button-like inserting in close vicinity to x1 and x2; solenidion u long and rod shaped taper-
ing distally; solenidion r rod shaped longer than genu I; seta ba spiniform inserting near the
complex of x1, x2, and e; setae u, v, p, q, s, wa, la, cG, mG short spines; setae e, hT, gT
spiniform; setae d, f, ra long and thin, with weakly broadened distal end; setae vF and pR
filiform; leg II: Solenidion r rod shaped, its length measuring approximately the length of
genu. Except for the regularly lacking chaetae (x2, x3, e) as given for leg I; leg III (3-14 a):
Tarsus with large claw; solenidion u rod shaped longer than tibia III; solenidion r rod
shaped shorter than genu III; setae d, f, r, w long and thin, w remarkably shorter than
the other three setae; setae p, q, u, v flat and buttonlike; setae e, s, kT, nG strong spines,
setae s, kT, nG with blunt apex; seta sR filiform; the relative size of leg III varying among the
heteromorphic males of one population; leg IV: Solenidion u thumb-like and stout, shorter
than tibia; seta f long and thin; setae u, v, p, q, s, r, w, kT spiniform, seta p very small and
short; seta wF short and filiform.
Ma le (h om e om o rph ic ): Length of idiosoma 375 – 425 lm; body in shape and colour
like the heteromorphic male, leg III, however, formed like female leg III, only seta p remark-
ably shorter and smaller than in the female.
Rem ar k : abnorm heteromorphic males can occur (idiosomal length approximately 415 lm):
one leg III is of the homeomorphic type, the other leg III with respect to the tarsus can be
regarded as a mixture of homeomorphic and heteromorphic type: setae d, e, f, r, s correspond
to the heteromorphic type, w is spiniform and corresponds to the homeomorphic type; p, q, u,
v are small to medium-sized spines, both legs are approximately equal in length.
Fe m a le : Length of idiosoma 395– 660 lm; body elongate; colour: white; idiosomal chae-
tome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b, 4a, g, ia, im, ip, ih; all idiosomal setae
smooth. Do rsum (3-12 d): Posterior margin of propodosomal shield with an incision of
variable depth, ranging from a short indentation to an incision that can reach the level of
the sockets of setae ve (recognizable by the small “hooks” at the lateral margin of propo-
dosomal shield); the margin of the incision corrugated to denticulate; a narrow, irregularly
shaped area consisting of cuticular bumps („wart patch“) running transversally immedi-
ately behind setae h1; the length of this field of bumps in transversal direction approxi-
mately half the distance between setae h1; bursa copulatrix a low protrusion at the opistho-
somal end, always strongly sclerotized and of intensive brown colour (even in newly
hatched females); copulatory opening localized at the ventral face of bursa copulatrix;
opisthosomal end sclerotized to a variable degree; the sclerotization originating from
the bursa copulatrix occasionally extending dorsally beyond the setae h1 or protruding
to the ventral face (if the sclerotization reaching near the setae h1, a distinctive border
to the unsclerotized cuticle apparent and the sclerotization becoming a true opisthosomal
shield); receptaculum seminis (3-11 b) consisting of a thin cuticular sac with a basal
strongly sclerotized and therefore intensively brown coloured invagination; inseminatory
canal running into this invagination and connecting with the sac at the end of the invagi-
nation; ductus conjunctivi originating approximately in the middle of this invagination;
inseminatory canal not coiling but connecting receptaculum seminis and copulatory open-
ing in a straight line; setae d2 and e1 short and thin. Ve n te r (3-11 a): Epimeres I merging
into a sternum medially; sternum and epimeres II covered by sclerotized lobed areas, epi-
meres III and IV accompanied by sclerotizations with an irregular posterior border and
broadening distally; the extent of sclerotizations of all coxal fields differing individually,
correlated with the degree of the opisthosomal sclerotization; apodemes III and IV of one
side often connected; on each side a sclerotized band running along the border between
proterosoma and hysterosoma; at its posterior margin each band connecting proximally
with a shorter narrow apodeme running transversally behind the larger one; oviporus be-
tween legs III and IV. L e g s: Leg I (3-3 c) and II: As given for the male; leg III (compare with
3-8 c): Solenidion u rod shaped, longer than tibia III; solenidion r rod shaped, shorter than
genu III; setae d, f, r long and thin; setae e, p, q, u, v, s, w, kT, nG strong spines; seta sR
50 Special part
filiform; leg IV (compare with 3-8 d): Solenidion u rod shaped, shorter than tibia IV; setae d
and f long and thin; setae e, p, q, u, v, s, r, w, kT strong spines, seta wF short and thin.
De ut ony m ph : Length of idiosoma 220 – 310 lm; body oval; colour: intensively brown;
idiosomal chaetome: scx, vi, si, se, c1, c2, c3, cp, d1, d2, e1, e2, f2, h1, h2, h3, 1a, 3a, 3b, 4a, g,
ia, im, ip, ih; all filiform idiosomal setae smooth. Do rsu m (3-14 b): Proterosoma tapering
to an acute tip anteriorly; length of hysterosomal setae measures 1/5 to 1/2 of the distance to
the respective posterior seta. Ve nte r (3-13): Palposoma extending beyond the propodo-
soma; epimeres I fused medially forming a sternum; coxal fields I and II covered by a con-
tinuous sclerotized area; lateral margins of this common area straight, running approxi-
mately parallel to the epimeres II; epimeres II slightly bent; epimerites II inconspicuous;
posterior margin of the common area straight and smooth; epimeres II not in touch medi-
ally; sternum not reaching the hind end of epimeres II; coxal fields III and IV covered by a
continouous sclerotized area; posterior margin of this field shaped irregularly; behind this
field another sclerotized area following around the genital opening with smooth anterior
margin; this field often connected with the anterior field by narrow lateral extensions; all
these sclerotizations together with the sucker plate surrounded by a large sclerotized area
separated from the medial sclerotizations by soft cuticle; laterally this area forming a thin
rim; sucker-plate (3-3 d) only slightly broader than long and relatively small (length ap-
proximately 1/8 of idiosomal length) clearly removed from caudal end; seta 1a short
and thin, seta 3a filiform, 3b an oval and 4a a longish conoid. L eg s: Leg I (3-15 a): So-
lenidion x1 club-shaped, slender; solenidia x2,3 rod shaped; famulus e a tiny spine; sole-
nidion u rod shaped at least as long as tarsus; solenidion r rod shaped, longer than genu;
seta e spoon shaped; setae f, p, q, ra, la leaflike, seta wa a slender spine; setae hT, gT and mG
long and strong spines; seta cG a short spine; setae d, vF, pR filiform; leg II (3-15 b): So-
lenidion u shorter than tarsus, solenidion r approximately as long as genu; seta e rod
shaped, seta ba an elongated spine. With respect to the regularly lacking chaetae (x2,
x3, e) otherwise as given for leg I; leg III (3-15 c): Solenidion u rod shaped, longer
than tarsus; setae e, p, q, s, r, w leaflike, setae d, f, nG, sR filiform; seta kT sword like
with pectinations; leg IV (3-15 d): Except the missing seta sR and seta wF, as given for
leg III but solenidion u shorter than tibia; setae d, f, r, w, wF filiform, seta w with pectina-
tions.
H a bit at : Under humid to wet decaying bark and wood, the deutonymphs also in humid
leaf litter; in the female gallery of Ips laricis (Coleoptera: Ipidae, V ITZTHUM 1923); under
the bark and in old holes by scolytids as well as in spruce stumps (C OOREMAN 1963); in
ground water (FAIN 1982).
B io lo gy : The deutonymphs were never detected attached to arthropods. O UDEMANS
(1911) and W URST (2002) report the deutonymph from decaying leaves. This is not ne-
cessarily another habitat of this species since the deutonymphs might be very mobile or
the hypopi could have been brought to this site by a larger arthropod.
Di stri bu ti on: Austria, Belgium, Germany, Hungaria, Romania, Slovakia.
3. Schwiebea schmuttereri T ÜRK & TÜRK , 1957 (3-16–3-21)

1957 Schwiebea schmuttereri T ÜRK & T ÜRK , Systematik und Ökologie der Tyroglyphiden Mit-
teleuropas. In: S TAMMER , H.-J. (ed.), Beiträge zur Systematik und Ökologie mitteleuro-
päischer Acarina, 1: 154.
1979 Schwiebea ruienensis FAIN & WAUTHY, Bull. Ann. Soc. r. belge Ent. 115: 174.
A dul ts:
Ma le ( he te ro m orph ic ): Length of idiosoma 255 – 320 lm; body elliptical, opisthoso-
mal end blunt; colour: white; idiosomal chaetome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a,
3b, 4a, g, ps1, ps2 , ps3, ia, im, ip, ih; all idiosomal setae smooth. D orsu m (3-17 a): Pro-
podosomal shield corrugated or with weak indentation at posterior margin; opisthosomal
shield convex with irregularly corrugated margin, reaching approximately the level of setae
e2 anteriorly; cupules ip always at the margin or outside the opisthosomal shield; seta h1
inserting posterior to ip within of opisthosomal shield. Ve nt er (3-16): Epimeres I fused
medially forming a sternum; sternum and epimeres II covered by sclerotized lobed areas;
a sclerotized area between epimeres II connecting the sclerotizations of coxal fields I and II;
epimeres III and IV accompanied by sclerotizations with an irregular posterior border and
broadening distally; apodemes III and IV and the accompanying sclerotizations frequently
fused at their proximal end; on each side a relatively wide sclerotized band running along
the border between proterosoma and hysterosoma; at its posterior margin each band con-
nected proximally with a shorter narrow apodeme running transversally behind the larger
one; aedeagus strongly sclerotized, hidden under genital folds forming a triangle (3-18); one
pair of adanal copulatory discs; opisthosomal shield continuing ventrally as sclerotization
of the opisthosomal end without clear margin; setae h2 and h3 inserting at the posterior
opisthosomal margin; length of h1 and h2 approximately equal, h3 being approximately
twice as long as h1 or h2; setae ps1,2 short and spinelike, ps3 thin and clearly longer
than the other ps setae. L e gs (3-17 b-c; for legs I, II, IV compare with 3-8 a-b, d-e):
Leg I: Solenidion x1 club shaped gradually tapering towards its base; solenidion x2 rod
shaped and slender inserting in close vicinity to x1; solenidion x3 rod shaped inserting an-
terior to seta f; famulus e button-like inserting in close vicinity to x1 and x2; solenidion u
long and rod shaped tapering distally; solenidion r1,2 rod shaped, r1 clearly longer than r2;
seta ba spiniform inserting near the complex of x1, x2, and e; setae u, v, p, q, s, wa, la, hT,
cG, short spines; setae e, gT spiniform; setae d, f, ra long and thin, with weakly broadened
distal end; setae mG, vF, pR filiform; leg II: Solenidion r rod shaped, approximately as long
as the genu; except for the regularly lacking chaetae (x2, x3, e) as given for leg I; leg III (3-17
b-c): Relative size of leg III varying among the heteromorphic males of a population; tarsus
with large claw; solenidion u rod shaped longer than tibia III; solenidion r rod shaped
approximately as long as genu III; seta e a very large spine, setae d, f, r, w long and
thin, w and r approximately equal in length; setae p, q, u, v, s short spines, v and s approxi-
mately equal in size; seta kT spiniform, sR filiform; leg IV: Solenidion u thick, shorter than
tibia; seta f long and thin; setae u, v, p, q, s, r, w, kT spiniform, setae r and w slender; seta wF
short and thin.
Ma le (h om e om o rph ic ): Length of idiosoma 215 – 260 lm; body in shape and colour
like the heteromorphic male, leg III, however, formed like female leg III.
Fe m a le : Length of idiosoma 315 – 365 lm; body elongate; colour: white; idiosomal chae-
smooth. Do rsu m : Propodosomal shield lobed or with weak indentation at posterior mar-
gin; opisthosomal end sclerotized without clear margin; anteriorly this sclerotization reach-
ing beyond the level of setae h1; copulatory opening a bare pore at the opisthosomal end
without supporting structures; receptaculum seminis (3-19 b) consisting of a thin cuticular
sac apically divided into five or six small fields by weakly sclerotized strips; connection with
the inseminatory canal via a jar-shaped structure; in this structure inserting the ductus con-
junctivi; diameter of the inseminatory canal small but the canal widening into a funnel
immediately before reaching the bursa copulatrix; the inseminatory canal long and forming
coils; setae h3 longer than setae h1,2. Ve nt er (3-19 a): Epimeres I merging into a sternum
medially; sternum and epimeres II covered by sclerotized lobed areas, in some individuals
the sclerotization of coxal fields I connected with the sclerotization of coxal fields II; epi-
52 Special part
meres III and IV accompanied by sclerotizations that broaden distally; epimeres IV occa-
sionally accompanied by a very broad band of sclerotized cuticle with constant width; these
sclerotizations with an irregular posterior margin; apodemes and the accompanying scler-
otizations III and IV always connected; on each side a sclerotized band running along the
border between proterosoma and hysterosoma; at its posterior margin each band con-
nected proximally with a shorter narrow apodeme running transversally behind the larger
one; opisthosomal end sclerotized without sharp margin; oviporus between legs III and IV.
L e gs: Leg I and II as given for the male; leg III (compare with 3-8 c): Solenidion u rod
shaped and very long, clearly longer than tibia III; solenidion r rod shaped, approximately
as long as genu III; setae d, f, r long and thin; setae e, p, q, u, v, s, w, kT spiniform; seta sR
filiform; leg IV (compare with 3-8 d): Solenidion u rod shaped, shorter than tibia IV; setae d
and f long and thin; setae e, p, q, u, v, s, r, w, kT spiniform, seta wF filiform.
D e u t on y mp h : Length of idiosoma 150 – 170 lm; body oval; colour: brown; idiosomal
chaetome: scx, vi, si, se, c1, c2, c3, cp, d1, d2, e1, e2, f2, h1, h2, h3, 1a, 3a, 3b, 4a, g, ia, im, ip, ih;
all filiform idiosomal setae smooth. D or s um : Hysterosomal setae of different length. Ve n-
t e r (3-20): Palposoma extending beyond the propodosoma; epimeres I fused medially form-
ing a sternum; coxal fields I and II covered by a continuous sclerotized area; epimeres II bent
and running in its posterior portion approximately parallel to the longitudinal body axis;
epimerites II only weakly bent, almost straight; epimeres II ending medially clearly anterior
to the posterior margin of the common sclerotized area, epimeres II not in touch medially;
sternum only slightly longer than 50 % of the length of the common sclerotized area; coxal
fields III and anterior part of coxal fields IV covered by a continouous sclerotized area;
anterior margin of this area with deep incision extending to the insertion of setae 3a, pos-
terior margin of this area with irregular course; posterior parts of coxal fields IV sclerotized
and separated from the anterior sclerotization by soft cuticle, the anterior margin of these
sclerotized areas smooth; the sclerotizations encompassing setae 4a and g; flanking the gen-
ital opening and sending along the insertion of leg IV a thin extension at each side to the
anterior sclerotization; all these sclerotizations and the sucker plate surrounded by a large
area with increasing sclerotization towards the body margin; this area forming a thin rim at
the body margin; sucker-plate only slightly broader than long, length approximately 20 %
of idiosomal length; only weakly removed from caudal end; seta 3a filiform, seta 1a a tiny,
seta 3b a small oval, and 4a a spherical conoid. L e g s : Leg I (3-21 a): Solenidion x1,2 club-
shaped, slender; solenidia x3 rod shaped; famulus e a tiny spine; solenidion u rod shaped,
longer than tarsus; solenidion r rod shaped, approximately as long as genu; seta e spoon
shaped; setae f, p, q, ra, la leaflike, seta wa, hT, gT slender spines; seta cG a short spine; setae
d, mG, vF, pR filiform; leg II (3-21 b): Solenidion u clearly shorter than in leg I, solenidion r
shorter than genu, seta e leaflike, setae ba long and thin, seta gT clearly longer than in leg I;
with respect to the regularly lacking chaetae (x2, x3, e) otherwise as given for leg I; leg III (3-
21 c): Solenidion u rod shaped, approximately as long as tarsus; setae e, f, p, q, s, r, w leaf-
like, setae d, sR filiform; seta kT a long spine with pectinations; leg IV (3-21 d): Except the
missing seta sR and seta wF, as given for leg III but solenidion u shorter than tibia; setae d, f,
r, w, wF filiform, seta w with a pectination, seta kT smooth.
H a bit at : Widespread under humid to wet bark of decaying wood and in rotting wood.
FAIN (1982) reports it also from ground water.
B io lo gy : Apparently a generalist that feeds on a large variety of rotting plant material. The
deutonymphs were found on Lithobius forficatus K OCH (Chilopoda: Lithobiomorpha:
Lithobiidae), Platynus assimilis PAYKULL , Carabus violaceus L INNAEUS (Coleoptera: Car-
abidae), Lasius brunneus L ATREILLE (Hymenoptera: Formicidae) (T ÜRK & T ÜRK 1957;
W URST 2002).
Di stri bu ti on: Austria, Belgium, Finland, Germany, Netherlands, Poland, Romania, Slo-
vakia.
4. Schwiebea cavernicola V ITZTHUM , 1932 (3-22–3-27)

1932 Schwiebea cavernicola V ITZTHUM, Zool. Jahrb. Syst. 63: 697.
1957 Schwiebea menzeli T ÜRK & T ÜRK, Systematik und Ökologie der Tyroglyphiden Mittel-
europas. In: S TAMMER , H.-J. (ed.), Beiträge zur Systematik und Ökologie mitteleuropäischer
Acarina, 1: 137.
1982 Schwiebea aquatilis FAIN, Acarologia 23 (4): 370.
A dul ts:
Ma le (he t ero m orp hi c) : Length of idiosoma 335 – 565 lm; body elliptic, opisthosomal
end blunt; colour: white; idiosomal chaetome: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b,
4a, g, ps1, ps2 , ps3, ia, im, ip, ih; all idiosomal setae smooth. Do rsu m (3-23 a): Posterior
margin of propodosomal shield with an incision of variable depth, ranging from 30 % of
the shield length to nearly the level of the sockets of setae ve (recognizable by the small
“hooks” at the lateral margin of propodosomal shield); the margin of the incision mostly
irregular and frequently with unclear margin; opisthosomal shield convex with irregularly
corrugated margin, anteriorly reaching approximately the level of setae e2; setae h1 insert-
ing outside the opisthosomal shield at the opisthosomal margin; cupule ip situated outside
the opisthosomal shield. Ve nt er (3-22): Epimeres I fused medially forming a sternum; ster-
num and epimeres II covered by sclerotized lobed areas; epimeres III and IV accompanied
by sclerotizations with an irregular posterior border and broadening distally; on each side a
sclerotized band running along the border between proterosoma and hysterosoma; at its
posterior margin each band connected proximally with a shorter narrow apodeme running
transversally behind the larger one; aedeagus strongly sclerotized, hidden under genital
folds forming a triangle (3-24); one pair of adanal copulatory discs; opisthosomal shield
continuing ventrally as narrow sclerotization of the opisthosomal end and encompassing
the setae h3; setae h2 and h3 inserting at the posterior opisthosomal margin; length of h2 and
h3 approximately equal, h1 mostly being shorter than h2 and h3; setae ps1,2 spinelike and
slender, ps3 thin and only slightly longer than the other ps setae. L e g s (3-23 b, c; for legs I,
II, IV compare with 3-8 a, b, d, e): Leg I: Solenidion x1 club shaped, apically bulbous;
solenidion x2 rod shaped and slender inserting in close vicinity to x1; solenidion x3 rod
shaped inserting anterior to seta f; famulus e button-like inserting in close vicinity to
x1 and x2; solenidion u long and rod shaped, tapering distally; solenidion r1,2 rod shaped
approximately equal in length; seta ba spiniform inserting near the complex of x1, x2, and
e; setae u, v, p, q, s, wa, la, hT, cG, mG short spines; seta gT a slender, seta e a large spine;
setae d, f, ra long and thin, with weakly broadened distal end; setae mG, vF, pR filiform;
leg II: Solenidion r rod shaped, shorter than genu; with respect to the regularly lacking
chaetae (x2, x3, e) as given for leg I; leg III (3-23 b, c): Tarsus with large claw; solenidion
u rod shaped longer than tibia III; solenidion r rod shaped shorter than genu III; seta e a
very large spine, setae d, f, r, w long and thin, w clearly shorter than r; setae p, q, u, v, s short
unequal spines, v and s approximately equal in size; seta kT and nG spiniform, sR filiform;
leg IV: Solenidion u thick, shorter than tibia; seta f long and thin; setae u, v, p, q, s, r, w, kT
spiniform; seta wF short and thin.
Ma le ( hom e om o rph ic ): Length of idiosoma approximately 340 lm; body in shape and
colour like the heteromorphic male, leg III, however, formed like female leg III.
R em a rk: Abnorm heteromorphic males can occur (idiosomal length approximately
325 lm): One leg III is of the homeomorphic type, the other leg III of the heteromorphic
type; both legs are approximately equal in length.
Fe m a le : Length of idiosoma: 380 – 640 lm; body elliptic; colour: white; idiosomal chae-
54 Special part
smooth. D orsu m : Posterior margin of propodosomal shield with an incision of variable

depth, ranging from a short indentation to nearly the level of the sockets of setae ve (re-
cognizable by the small “hooks” at the lateral margin of propodosomal shield); the margin
of the incision mostly irregular and frequently with unclear margin; bursa copulatrix a
wart-like protrusion at the opisthosomal end; receptaculum seminis (3-25 b) a spherical,
thin walled strongly wrinkled sac, at its basis a thin ring like sclerotization; ductus con-
junctivi inserting at this ring; inseminatory canal inserting in the center of the area marked
by the ring; inseminatory canal relatively short, connecting receptaculum seminis and co-
pulatory opening without coiling; setae h2,3 approximately equal in length, seta h1 shorter
than setae h2,3. Ve nte r (3–25 a) : Epimeres I merging into a sternum medially; sternum
and epimeres II covered by sclerotized lobed areas of variable extension (3-25 a showing a
female with weak sclerotization, the sclerotization equaling in extreme cases the scleroti-
zation of the males); epimeres III and IV accompanied by sclerotizations with irregular pos-
terior margin that broaden distally; on each side a sclerotized band running along the bor-
der between proterosoma and hysterosoma; at its posterior margin each band connected
proximally with a shorter narrow apodeme running transversally behind the larger one;
oviporus between legs III and IV. L e gs: Leg I and II: as given for the male; leg III (compare
with 3-8 c): Solenidion u rod shaped and very long, clearly longer than tibia III; solenidion r
rod shaped, shorter than genu III; setae d, f, r long and thin; setae e, p, q, u, v, s, w, kT, nG
spiniform; seta sR filiform; leg IV(compare with 3-8 d): Solenidion u rod shaped, approxi-
mately as long as tibia IV; setae d and f long and thin; setae e, p, q, u, v, s, r, w, kT spiniform,
seta wF filiform.
De ut ony m ph : Length of idiosoma 180 – 210 lm; body ovoid; colour: beige; idiosomal
chaetome: scx, vi, si, se, c1, c2, c3, cp, d1, d2, e1, e2, f2, h1, h2, h3, 1a, 3a, 3b, 4a, g, ia, im, ip,
ih; all filiform idiosomal setae smooth. Do rsum : Entire dorsum covered with small pits; all
setae at least 6 lm. Ve nte r (3-26): Palposoma extending beyond the propodosoma; the
first setae posterior to the solenidia clearly longer than the width of the palposoma at
the level of their insertion; epimeres I fused medially forming a sternum; coxal fields I
and II covered by a continuous sclerotized area; epimeres II only slightly bent, almost
straight; epimerites II not visible; lateral edge of the sclerotized area running approximately
parallel to the epimeres II; epimeres II ending proximally immediately anterior to the pos-
terior boundary of the sclerotized area, the epimeres do not touch medially; sternum ex-
tending beyond half length of the sclerotized area; coxal fields III and anterior part of coxal
fields IV covered by a continouous sclerotized area; anterior margin of this area with deep
incision almost reaching the posterior margin of this sclerotization; posterior margin of this
area almost smooth; posterior parts of coxal fields IV sclerotized and separated from the
anterior sclerotization by soft cuticle, the anterior margin of these sclerotized areas smooth;
the sclerotizations encompassing setae 4a and g and flanking the genital opening; all these
sclerotizations and the sucker plate surrounded by a large sclerotized area with increasing
sclerotization towards the body side; this area forming a thin rim at body margin; sucker-
plate broader than long, length approximately 15 % of idiosomal length; clearly removed
from caudal end; seta 3a filiform, seta 1a and 3b oval conoids, seta 4a a heart shaped con-
oid. L e g s: Leg I (3-27 a): Solenidion x1 slender with bulbous apex; solenidion x2 club-
shaped and slender; solenidion x3 rod shaped; famulus e a small slender spine; solenidion
u rod shaped longer than tarsus; solenidion r rod shaped shorter than genu; seta e spoon
shaped; setae f, p, q, ra, la leaflike, setae wa, hT, gT, cG, mG slender spines; setae d, vF, pR
filiform; leg II (3-27 b): Solenidion x1 a thick club; solenidion u clearly shorter than in leg I;
seta e rod shaped, seta ba a long spine; except for the regularly lacking chaetae (x2, x3, e)
otherwise as given for leg I; leg III (3-27 c): Solenidion u rod shaped, approximately as long
as tibia; setae e, f, p, q, s, r, w leaflike; seta kT a long spine; seta nG spiniform; setae d, sR
filiform; leg IV (3-27 d): Except the lacking seta sR and seta wF, as given for leg III but
solenidion u shorter than tibia; setae d, r, wF filiform, seta w sword like with a pectination.
H a bit at : Widespread under humid to wet bark of decaying wood and in rotting wood and
at tulip bulbs. Also in aquatic habitats like fish ponds, sewage ponds (V ITZTHUM 1932) and
ground water (FAIN 1982). J AMES P. E NRIGHT (Kasel, pers. comm.) found S. cavernicola in
ground water in Aschaffenburg (Germany) (13.5 m depth, temperature approximately
11 C).
B io lo gy : The deutonymphs were found on Cryptops parisii (Chilopoda: Scolopendromor-
pha: Cryptopidae) and other Myriapoda, larvae of Cerambycidae, larvae of Melanotus
castanipes PAYKULL , Dolopius marginatus L . (Coleoptera: Elateridae) and Myrmica laevi-
nodis N YLANDER (Hymenoptera: Formicidae) (T ÜRK & T ÜRK 1957; W URST 2002).
Di stri bu ti on: Austria, Croatia, Hungaria, Germany.
5. Schwiebea-nova-group (3-28–3-32)
1982 Schwiebea subterranea FAIN, Acarologia 23 (4): 366–370.
Besides other characters (like the very similar deutonymphs) members of the Schwiebea-
nova-group can most easily be separated from the other Schwiebea species by the long and
filiform setae la of leg I and II in the non-hypopal instars (3-30 b). Schwiebea subterranea,
described from ground water in Vienna, belongs without doubt to this group (W URST
2002). The Schwiebea-nova-group comprises a large number of very closely related species
that need taxonomic revision. Under these circumstances the description by FAIN (1982)
allows no certain determination. In order to show the general features of the Schwiebea-
nova-group Figs. 3-28–3-32 present a very common but unidentified species of this species
group collected under the bark of decaying wood.
Genus Naiadacarus FASHING , 1974 (3-33)

1974 Naiadacarus FASHING, Acarologia 16 (1): 166–181.
Typus generis: Naiadacarus arboricola FASHING , 1974.
Di ag no sis: Retroconjugate mites with facultative hypopody; leg length of non-hypopal
instars more than 50 % of idiosomal length; idiosomal chaetome: Seta scx of non-hypopal
instars a short spine; female: scx, vi, se, cp, d2, e1, e2, h1, h2, h3, 1a, 3a, 3b, g, 4a; male: As
given for female but additionally ps1, ps2, ps3; idiosomal cuticle with sculpturing consisting
of narrow corrugations and flat bumps dorsally.
A comparison with the diagnosis for the genus Schwiebea suggests that Naiadacarus tax-
onomically is a “Schwiebea with long legs”. Species of Naiadacarus occur in water-filled
tree holes (3-33) and in pitcher plants (Nepenthes) (FASHING 1974, FASHING & C HUA
2002), feeding on fungi, and decaying plant and insect tissues. They are recorded from
North America and Southeast Asia but occur also in central Europe: the author found
non-hypopal instars of a new species in Austria in a water filled hole in a tree stump.
Family Histiostomatidae B ERLESE , 1897 (3-3 e, f, 3-34–3-35)

Syn.: Anoetidae O UDEMANS , 1904
Di ag no sis: Proconjugate mites with mostly facultative hypopody (3-35); gnathosoma
highly modified and transformed into an apparatus to collect small particles from aqueous
media; chelicerae transformed into paddle- to rake-like instruments; the two distal palp
segments fused and at least two of the three palp solenidia strongly elongated; additional
soft walled appendages at the rostral end of the gnathosoma regularly occurring (3-3 e, f);
non-hypopal instars with disc-shaped osmoregulatory organs (= ring organs) (3-34) ven-
trally.
56 References
The Histiostomatidae is a very large and polymorphic family with world wide distribution.
The minimal condition for feeding is a thin water film covering the habitat of the histios-
tomatids. Many species collect their food from the uppermost water layer where the con-
centration of food particles is maximal due to interfacial effects. Other species normally live
completely submerged and utilize particles that are actively removed from solid surfaces by
the mites. In both cases the chelicerae are quickly moved back and forth creating a vortex
that transports the food to the preoral cavity (W URST & K OVAC 2003).
Despite their significance in species number and individual density, the taxonomic classi-
fication of the Histiostomatidae is far behind that of other mite taxa. The most recent com-
prehensive review of central European species is the beautiful book of S CHEUCHER (1957),
including identification keys and species descriptions.
Acknowledgement
I am very grateful to Dr. T. Romig (Stuttgart) who critically read the manuscript and im-
proved it stylistically.
References
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W ITALINSKI , W., S ZLENDAK , E. & B OCZEK , J. (1990): Anatomy and ultrastructure of the reproductive
systems of Acarus siro (Acari: Acaridae). Expl. appl. Acarol. 10: 1–31.
W URST, E. (1993): Investigations on the anatomy and the behaviour of the fur mite Listrophorus leuck-
arti (Acari: Listrophoridae). Stuttg. Beitr. Naturk. (Ser. A; Biologie) 503: 1–68.
W URST, E. (2002): Beitrag zur Revision der mitteleuropäischen Arten der Gattung Schwiebea O UDE-
MANS , 1916 (Acari: Acaridae); Verlag Grauer, Beuren, Stuttgart: 1–128 + plates.
W URST, E. & F RANK , B. (1998): Contributions to a revision of the genus Schwiebea (Acari: Acaridae).
I. Redescription of Schwiebea talpa and Schwiebea nesbitti. Stuttg. Beitr. Naturk. (Ser. A; Biologie)
579: 1–52.
W URST, E. & K OVAC , D. (2003): Description and biology of the mite Tensiostoma veliaphilum n. gen.
n. sp. from the water surface of bamboo phytotelmata of Southeast Asia (Arachnida, Acari, Astig-
mata, Histiostomatidae). Senckenberg. biol. 82 (1/2): 63–98.
58 Figures
Fig. 3-4: Schwiebea talpa, male, ventral view (left legs partly omitted). A = aedeagus, AD = adanal
copulatory discs, AS = anal slit, GP = genital papillae. All other symbols designate setae and cupules.
Scale bar: 40 lm.
Fig. 3-5: a, Schwiebea talpa, male, dorsal view. OS = opisthosomal shield, PS = propodosomal shield.
All other symbols designate setae and cupules. Scale bar: 40 lm; b, Schwiebea talpa, female, dorsal
view. BC = bursa copulatrix, PS = propodosomal shield. All other symbols designate setae and cupules.
Scale bar: 40 lm.
60 Figures
a b
c d
Fig. 3-6: Schwiebea talpa, male, aedeagus. Optical sections (light micrographs, Nomarski DIC) taken
through different levels beginning with the superficial level in Fig. a. A = aedeagus, AS = anal slit, g =
seta, GF = genital folds, GP = genital papillae, 4a = seta, arrowhead points at tip of aedeagus. Scale bar:
10 lm.
Fig. 3-7: Schwiebea talpa, female; a, ventral view (left legs partly omitted). AS = anal slit, GP = genital
papillae, OP = oviporus. All other symbols designate setae and cupules. Scale bar: 40 lm; b, recep-
taculum seminis with appendages. BC = bursa copulatrix, DC = ductus conjunctivus, IC = insemina-
tory canal, RS = receptaculum seminis. Scale bar: 5 lm.
62 Figures
Fig. 3-8: Schwiebea talpa, legs. a, female leg I; b, female leg II; c, female leg III; d, female leg IV. Legs
are shown in dorsal view, the inset shows the tarsus from the opposite side; e, male tarsus IV. Setae d
and e are modified into mushroom shaped copulatory adhesive devices. Scale bars: 15 lm.
Fig. 3-9: Schwiebea eurynympha, heteromorphic male, ventral view (left legs partly omitted). Scale
bar: 40 lm.
64 Figures
a b
c d
Fig. 3-10: Schwiebea eurynympha, male, aedeagus. Optical sections (light micrographs, Nomarski
DIC) taken through different levels beginning with the superficial level in Fig. a. A = aedeagus, AS
= anal slit, g = seta, FP = fecal pellet, GF = genital folds, GP = genital papillae, 4a = seta, arrowhead
points at tip of aedeagus. Scale bar: 20 lm.
Fig. 3-11: Schwiebea eurynympha, female. a, ventral view (left legs partly omitted). Scale bar: 40 lm;
b, receptaculum seminis with appendages. BC = bursa copulatrix, DC = ductus conjunctivus, IC =
inseminatory canal, RS = receptaculum seminis. Scale bar: 10 lm.
66 Figures
Fig. 3-12: Schwiebea eurynympha, opisthosomal end. a, male, dorsal view; b-c, male, ventral view,
showing two extreme variations of setae h1–3; d, female, dorsal view. WP = wart patch. Scale bars:
a, 40 lm, b-c, 30 lm, d, 40 lm.
Fig. 3-13: Schwiebea eurynympha, deutonymph, ventral view (left legs partly omitted). PS = palpo-
soma. Scale bar: 30 lm.
68 Figures
Fig. 3-14: a, Schwiebea eurynympha, heteromorphic male, leg III, dorsal view. The inset shows the
tarsus from the opposite side. Scale bar: 15 lm; b, Schwiebea eurynympha, deutonymph, dorsal view.
Scale bar: 30 lm.
Fig. 3-15: Schwiebea eurynympha, deutonymph. a, leg I; b, leg II; c, leg III; d, leg IV. Legs are shown in
dorsal view, the inset shows the apex of tarsus from the opposite side. Scale bars: 15 lm.
70 Figures
Fig. 3-16: Schwiebea schmuttereri, heteromorphic male, ventral view (left legs partly omitted). Scale
bar: 30 lm.
Fig. 3-17: Schwiebea schmuttereri, heteromorphic male. a, opisthosomal end, dorsal view. OS =
opisthosomal shield; b, leg III, dorsal view; c, leg III, showing the tarsus from the opposite side as
given in Fig. b. Scale bars: a, 30 lm, b-c, 10 lm.
72 Figures
Figs. 3-18: Schwiebea schmuttereri, male, aedeagus. Optical sections (light micrographs, Nomarski
DIC) taken through different levels beginning with the superficial level in Fig. a. A = aedeagus, AS =
anal slit, g = seta, GF = genital folds, GP = genital papillae, 4a = seta, arrowhead points at tip of ae-
deagus. Scale bar: 10 lm.
Fig. 3-19: Schwiebea schmuttereri, female; a, ventral view (left legs partly omitted). Scale bar: 30 lm;
b, receptaculum seminis with appendages. BC = bursa copulatrix, DC = ductus conjunctivus, IC =
74 Figures
Fig. 3-20: Schwiebea schmuttereri, deutonymph, ventral view (left legs partly omitted). Scale bar:
20 lm.
Figs. 3-21: Schwiebea schmuttereri, deutonymph. a, leg I; b, leg II; c, leg III; d, leg IV. Legs are shown in
dorsal view. Scale bars: a-b, 20 lm, c-d, 10 lm.
76 Figures
Fig. 3-22: Schwiebea cavernicola, heteromorphic male, ventral view (left legs partly omitted). Scale
bar: 40 lm.
Fig. 3-23 : Schwiebea cavernicola, heteromorphic male. a, opisthosomal end, dorsal view; b, leg III,
dorsal view; c, leg III showing the tarsus from the opposite side as given in Fig. b. Scale bars: a, 40 lm,
b-c, 20 lm.
78 Figures
a b
c d
Fig. 3-24: Schwiebea cavernicola, male, aedeagus. Optical sections (light micrographs, Nomarski
DIC) taken through different levels beginning with the superficial level in Fig. a. A = aedeagus, AS
= anal slit, g = seta, GF = genital folds, GP = genital papillae, 4a = seta, arrowhead points at tip
of aedeagus. Scale bar: 20 lm.
Fig. 3-25: Schwiebea cavernicola, female. a, ventral view (left legs partly omitted). Scale bar: 40 lm; b,
female, receptaculum seminis with appendages. BC = bursa copulatrix, DC = ductus conjunctivus, IC =
80 Figures
Fig. 3-26: Schwiebea cavernicola, deutonymph, ventral view (left legs partly omitted).
Scale bar: 30 lm.
Fig. 3-27: Schwiebea schmuttereri, deutonymph. a, leg I; b, leg II; c, leg III; e, leg IV. Legs are shown in
dorsal view. Scale bars: a-b, 15 lm, c-d, 10 lm.
82 Figures
Fig. 3-28: Schwiebea sp. (nova-group), heteromorphic male, ventral view (left legs partly omitted). In
some species of the nova-group seta hT of leg I and II is lacking. Scale bar: 30 lm.
Fig. 3-29: Schwiebea sp. (nova-group), heteromorphic male. a, opisthosomal end, dorsal view; b, leg
III, dorsal view; c, leg III showing the tarsus from the opposite side as given in Fig. b. Scale bars: a,
30 lm, b-c, 10 lm.
84 Figures
Fig. 3-30: Schwiebea sp. (nova-group), female. a, ventral view (left legs partly omitted). In some
species of the nova-group seta hT of leg I and II is lacking; b, tarsus I, medial view. Scale bars:
a, 40 lm, b, 15 lm.
Fig. 3-31: Schwiebea sp. (nova-group), deutonymph, ventral view (left legs partly omitted). Scale bar:
30 lm.
86 Figures
Fig. 3-32: Schwiebea sp. (nova-group), deutonymph. a, leg I; b, leg II; c, leg III; d, leg IV. Legs are
shown in dorsal view. ). In some species of the nova-group seta hT of leg I and II is lacking. Scale
bars: a-b, 15 lm, c-d, 10 lm.
Figs. 3-33: Naiadacarus sp. (light micrographs). a, male; b, female. Scale bars: 100 lm.
Fig. 3-34: Idiosoma (venter) of a histiosto-

matid female (legs partly omitted). AS = anal
slit, OP = oviporus, RO = osmoregulatory ring
organs.
88 Figures
Fig. 3-35: Hormosianoetus sp.. a, male (light

micrograph); b, female (light micrograph);
c, deutonymph (light micrograph). The genus
Hormosianoetus contains many aquatic species.
The deutonymphs typically use diptera for
transport. Scale bars: a-b, 50 lm, c, 40 lm.
4. Acari: Limnic Oribatida 89
4. Acari: Limnic Oribatida

Gerd Weigmann and Ralf Deichsel
General part
In Europe (excluding Caucasus and Macaronesian Islands) S CHATZ (2005) calculates in
total about 2100 species of described oribatid mite species, worldwide about 10.000 spe-
cies. In Germany about 520 species are determined and evaluated (without several syno-
nyms; after W EIGMANN 2006). The considered geographical region of this chapter is Cen-
tral Europe, i. e. Germany and the surrounding countries; other Eastern European countries
are excluded, as well as the mediterranean region, if not mentioned especially.
This chapter covers 17 oribatid mite species, namely species that live exclusively or reg-
ularly in fresh water habitats. Some of the species that prefer very wet habitats, especially
bogs, wet meadows or fen woodlands, can be found from time to time submersed as well as
terrestrially. These species were only exceptionally included. Terrestrial oribatid mite spe-
cies can also occasionally be found in aquatic samples, if they were flushed into the water by
rain or flooding or they fell from trees etc. We recommend the use of general textbooks for
the determination of those species not included here, in cases of doubt or for studying the
fauna of amphibious habitats.
General Morphology
Most adult oribatid mites have a compact and well sclerotized body. From a dorsal view,
two body shields are visible: anteriorly the prodorsum, posteriorly the notogaster. The pro-
dorsum covers the propodosoma, which represents the segments of legs I and II (W EIG-
MANN 2001). The gnathosoma is normally covered by a “rostrum”, which is in Oribatida
the anterior part of the prodorsum. The mouthparts can be retracted into a rostral cavity; a
ventral shield formed by parts of the pedipalpal coxae and the sternite (4-2 c-e) may serve as
a hinged lid to protect the retracted mouthparts. The propodosoma and the gnathosoma
form the proterosoma. All remaining posterior body segments form the hysterosoma,
which includes the segments of legs III and IV (together called the metapodosoma) and
the segments of the opisthosoma. In most oribatid mites, the adults have an undivided no-
togaster, the dorsal plate of the hysterosoma. However, in some species the notogaster is
subdivided by transversal sutures into 2-4 notogastral shields (4-2 a).
N ot og ast e r: The basic notogastral setation scheme of Oribatida consists of 16 pairs of
setae. Originally, these notogastral setae are arranged in six transversal rows, indicating six
hysterosomal segments. In many cases, one or more pairs of setae are reduced. Exception-
ally, the notogastral setae number can also be increased. The number and configuration of
setae (chaetotaxy) are essential taxonomic and systematic characters. Some higher oribatid
mite taxa have a set of 4 pairs of areae porosae (4-1 a, d: several aggregated glandular
pores) or sacculi (4-1 e: glandular pores in an epidermal cavity with one orifice) on the
notogaster. Sometimes these areae porosae or sacculi are hardly visible. The notogaster
occasionally has a pair of opisthosomal glands with lateral openings (4-1 a).
Pr opo dos oma ( 4- 1 a, c ): A typical Oribatida character is a pair of sensory setae at the
posterio-lateral margin of the prodorsum, called sensilli (ss), arising from bothridia (bot),
together representing trichobothria. The sensilli occur in very different but specific shapes,
such as filiform, fusiform or claviform, with long or short stalks. The sensilli most probably
have a mechano-receptive function. A typical set of prodorsal setae exists: the interlamellar
setae (in) between the bothridia, the lamellar setae (le) in the middle of the prodorsum, the

https://doi.org/10.1007/978-3-662-55958-1_4
90 General part
Fig. 4-1: a-c General morphology of oribatid mites. a, dorsal view; b, ventral view; c, lateral view on
anterior body part; d, area porosa; e, sacculus.
Aa-A3 - row of areae porosae; ag – aggenital setae; AN – anal plate; an – anal setae; bot - bothridium;
csp - cuspis; epi – epimer (= coxa); G – genital plate; gla - glandular opening; in - interlamellar seta; le -
lamellar seta; lam - lamella; NG – notogaster; ptc – pedotectum; ptm - pteromorph; ro - rostral seta; ss -
sensillus; trl - translamella; tut – tutorium; VP – ventral shield. Legs are detached.
rostral setae (ro) near the rostral edge. Besides the bothridiae, 1-2 pairs of exobothridial
setae (ex) are often present laterally. A pair of ridges or blades, the lamellae (lam), are often
situated on the prodorsum, mostly reaching from the bothridia to the lamellar setae. The
lamella may end anteriorly as a free tip, called cuspis (csp), bearing the lamellar seta on each
side. If the lamellae are reduced to ridges, they are called costulae; in some Oribatida the
lamellae are lacking completely.
Oribatid mites use different methods to protect their appendages against predation. The
Phthiracaroidea and others fold the prodorsum towards the ventral hysterosoma and shut
the retracted legs in the gap (4-2 b). In other taxa, a series of lateral blades protects at least
the bases of the legs: the lamellar blade (lam), the tutorium (tut), the pedotecta (ptc) at legs I
and II, the discidium with the custodium between legs III and IV (4-1 c). If present, the
anterio-lateral blade- or wing-like projections (pteromorphs: ptm) at the shoulder edge
of the notogaster serve the same purpose (4-1 c).
Ve n t r a l a s p e c t (4- 1 b ): The coxae of all legs are immobile; they form so-called epi-
meres, part of a ventral shield. The border between the proterosoma and hysterosoma
is marked by a transverse suture between epimeres I+II and epimeres III+IV in “primitive”
Oribatida, but are totally fused in “higher” Oribatida. In the opithosomal region, the mites
show genital and anal openings; both are protected by well-sclerotized plates, which can be
opened like window shutters. In “higher” Oribatida, these genital plates and anal plates are
surrounded by a ventral shield, which is fused with the epimeres of legs IV and is separated
from the notogaster by a soft ligament (4-1 b). In “primitive” Oribatida, such a compact
ventral shield is not present; the genital and anal plates are integrated into a soft ventral
field, often flanked by aggenital shields (beside the genital plates) and adanal shields (beside
the anal plates).
Ju ve ni le s are not as well sclerotized as the adults. The sclerotized plates on their body
surface don’t match the adult’s scheme in most cases. Different types can be found: from no
shields and wrinkled cuticula to large shields, covering most of the body surface.
The morphological terminology follows H UNT et al. (1998).
Special Glossary
areae porosae cuticular areas with pore canals (glandular openings); the a. p. of notogaster are
mostly organized in 4 pairs (= “octotaxic system”)
bothridium (pl. bothridia) cup-like depression laterally on the prodorsum in which the sen-
sillus inserts, a trichobothrium as part of a sense organ (old synonym: pseudos-
tigma)
cuspis (pl. cuspides) free distal end of lamella on prodorsum (or sensu lato of other
lamellar structures)
custodium (pl. custodia) an acute anterior extension (cuspis) of the discidium,
discidium (pl. discidia) a cuticular blade between the bases of legs III and IV
epimeres ventral areas of the podosoma segments: obviously homologous with coxae of
the legs, which are fused with the sternal area
exobothridial setae seta at the lateral side of bothridium in most Oribatida; in species without bo-
thridia situated laterally from interlamellar setae
interlamellar setae seta between the bothridia in Oribatida
lamella (pl. lamellae) a pair of longitudinal blade-like cuticular structures on the pro-
dorsum of many adult Oribatida. The proximal end is close to the bothridia,
the distal end bears the lamellar setae, mostly. The blades protect the distal parts
of legs I when these are retracted. Some Oribatida have a translamella, a con-
necting structure between both lamellae in the distal region. Costulae (= lamellar
92 General part
costulae) are rib-like ridges instead of blades and are regarded as homologous
structures of lamellae
notogaster the dorso-lateral shield of the hysterosoma; it may be one single shield or it may
be subdivided by transversal scissures into 2-4 parts.
pedotectum (pl. pedotecta) scale-like blades behind the insertion of legs I or II (old synonym
tectopedium II and III)
prodorsum dorso-lateral shield of the propodosoma with some special structures (bothridi-
um, lamellae; the rostral, lamellar, interlamellar and exobothridial setae; etc.).
Fig. 4-2: a, Brachychthonoid mite with notogaster sutures; b, Ptychoid mite (Phthiracaridae) with
“jackknifing” mechanism; c, stenarthric infracapitulum of Mucronothrus (sternite with setae h sepa-
rated from pedipalp coxae); d, anarthric infracapitulum of Malaconothrus; e, diarthric infracapitulum
of Trhypochthoniellus. Legs are detached.
The anterior part is the so-called rostrum, which mostly covers the gnathosoma
with the chelicerae and pedipalps. Mostly the prodorsum is distinctly separated
from the notogaster, exceptionally both may be fused (synonym: aspis)
pteromorph (pl. pteromorphs) large blade-like structure of the humeral region of the noto-
gaster in many derivative Oribatida, often bent downwards; it may be immo-
vable or movable, and it serves as protection for the posterior legs (homologous
structures are smaller “humeral blades”)
rostrum anterior part of the prodorsum which mostly covers the gnathosoma in Oriba-
tida (syn. rostral tectum); in other groups of mites the term is used differently
sacculus (pl. sacculi) invaginated porose organ; the sacculi of the notogaster are homo-
logous with the notogastral areae porosae.
sensillum (pl. sensilli) setal sense organ of the bothriotrich type; in most Oribatida inserted
in the bothridium in posterior region of the prodorsum (old synonyms: bothri-
dial seta; pseudostigmatic organ)
translamella transverse ridge or tectum that connects the paired prodorsal lamellae
tutorium (pl. tutoria) a paired ridge or blade on the lateral prodorsum in many derivative
Oribatida, often with an anterior cuspis (old synonym: tectopedium I)
Special part
Area treated
Europe.
Key to families and genera

This determination key almost completely ignores terrestrial species. To avoid wrong determination of
species from terrestrial or semiaquatic origins, the descriptions and figures given for the aquatic species
must be checked carefully. Special attention must be payed to the prodorsum structures, the arrange-
ment of the genital and anal plates and the presence of areae porosae or pteromorphs.
1 Ventral shield complete, surrounding the genital and anal plates; ventral shield well separated
from the entire notogaster shield; all epimeres fused with each other and with the ventral shield
(4-1 b) (“Higher Oribatida”) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Other combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Transverse scissures divide the notogaster into two to four shields (4-2 a). . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (some non-aquatic Oribatida)
– Notogaster entire, without transverse scissures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Prodorsum movably articulated with hysterosoma (4-2 b); legs inserted in soft ventral epimeral
region. (Legs can be withdrawn into a cavity and the prodorsum then closed over them like a lid)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . (non-aquatic: Phthiracaroidea and Mesoplophoridae)
– Prodorsum not movably articulated with hysterosoma, i.e. prodorsum fixed to notogaster or con-
nected to it by a weak ligament; with distinct sclerotized epimeral parts I+II and III+IV. Without
complete ventral plate surrounding the genital and anal plates (cf. 4-3 e, h).. . . . . . . . . . . . 4
4 Without bothridia and sensilli also in adults; only shorter exobothridial setae outside the large
interlamellar setae (cf. 4-3 a, d, f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– With bothridia (in adults) and sensilli, at the side of the sensilli with shorter exobothridial setae,
interlamellar setae are also present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Rostral tip narrow, protruding nose-like; rostral and lamellar setae near each other (4-3 a); bo-
thridium shaped like an open funnel, sensillus filiform (4-3 b); legs with one claw (15-20 pairs of
genital setae; infracapitulum stenartric (4-2 c); body length 580-800 lm) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mucronothridae: Mucronothrus (page 94)
94 Special part
– Rostrum rounded; rostral and lamellar setae not near each other; sensillus smooth and fusiform;
legs with 3 claws (with 10-18 pairs of genital setae; infracapitulum diarthric (4-2 e); two pairs of
adanal setae; one pair of anal setae; genu of legs I and II each with three setae and one solenidium;
body length 435-580 lm) (4-3 c-e) . . . . . Trhypochthoniellidae: Trhypochthoniellus (page 96)
6 Legs with one claw (4-3 f-h) . . . . . . . . . . . . . . . Malaconothridae: Malaconothrus (page 97)
– Legs with three claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Cerotegument of notogaster with porose plaques; three pairs of adanal setae; 5-12 pairs of genital
setae (infracapitulum anarthric, without hinge: 4-2 d) . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malaconothridae: Trimalaconothrus (page 98)
– Cerotegument of notogaster without porose plaques, meshlike structures sometimes in anterior
part; two pairs of adanal setae; 10-18 pairs of genital setae (infracapitulum diarthric, 4-2 e) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trhypochthoniellidae: Trhypochthoniellus (page 96)
8 Notogaster with pteromorphs, i. e. larger blades at the shoulders, often bent downwards
(4-1 a,c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Notogaster without pteromorphs at the shoulders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Notogaster anteriorly with lens-like convex clear spot (“lenticulus”: 4-5 a); lamellar costulae
straight, anteriorly convergent, but at a good distance from each other . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrozetidae: Hydrozetes (page 101)
– Notogaster anteriorly without a clear spot or other character combinations . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . diverse mainly terrestrial Oribatida
10 Pteromorphs narrow, ribbonlike or triangular, posteriorly distinctly curved towards the notogaster
border (4-7 a-d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnozetidae: Limnozetes (page 106)
– Pteromorphs broad at their base, posteriorly gradually curving into notogaster border. . . . 11
11 Between pteromorphs, the anterior border of the notogaster forms a relatively broad roof-like
bridge, covering the posterior part of the prodorsum and mostly also the bothridia (4-7 f)
(Pteromorphs movable, hinge line often hardly visible) . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mycobatidae: Punctoribates (page 108)
– Between immovable pteromorphs, the anterior border of the notogaster as a very small ribbon, not
covering the posterior part of the prodorsum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Notogaster without visible setae (at least the insertion points present) and without areae porosae;
legs I with one claw, legs II-IV with three claws; lamellae slightly converging, with cuspides; sensilli
long and claviform (4-8 d) (Body black-brown, longer than 600 lm) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zetomimidae: Heterozetes (page 109)
– Notogaster with 10 pairs of fine setae and with four pairs of areae porosae; legs I-II with one claw,
legs III-IV with three claws, the lateral claws very thin; cuspis long, translamella visible only as a
fine line; sensilli slender and claviform, with granulation (4-8 a) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zetomimidae: Zetomimus (page 109)
Family Mucronothridae K UNST, 1971

1971 Mucronothridae K UNST, In: D ANIEL M., C ERNY V. (eds), Klı́č zvı́řeny ČSSR. Česk. Akad.
Věd, Praha: 554.
Genus Mucronothrus T RÄGÅRDH , 1931

1931 Mucronothrus T RÄGÅRDH, Zool. Faroes 49: 31
Typus generis: Mucronothrus rostratus T RÄGÅRDH , 1931 by original designation; syn. of
Malaconothrus nasalis W ILLMANN , 1929.
Di ag no sis o f the ad ul ts: The family has one genus, thus the diagnosis of the adults is
the same for both (see family key). Anterior edge of rostrum nasiform, narrow, bent
downwards; rostral setae close together on a nasiform tip. Bothridium open funnel-
like, sensillus filiform, formed like normal seta. N o tog as te r: Elongated with almost par-
allel sides, 15 pairs of fine setae, partly very long (three pairs only visible from the ventral
Fig. 4-3: a-b, Mucronothrus nasalis: a, dorsal view; b, bothridial region; c-e, Trhypochthoniellus lon-
gisetus: c, dorsal view T. l. forma typica; d, prodorsum of T. l. forma setosus in dorsal view; e, ventral
view; f-h, Malaconothrus monodactylus: f, dorsal view; g, lateral view; h, ventral view. Legs are de-
tached.
96 Special part
aspect). Ve n tra l re gi on : Genital, adanal and anal plates in a longitudinal gap of the no-
togaster shield, which is bent towards the ventral side, two pairs of adanal setae, two pairs
of anal setae, no aggenital plates and setae epimeres III and IV each medially fused. Infra-
capitulum with stenathric articulation. Each tarsus with one claw.
Di ag no sis o f th e j uv en il e s: Facies similar to that of the adults with less sclerotization;
genital plates less developed.
Di stri bu ti on: A genus with two species; M. nasalis is the single species in Europe (Skan-
dinavia, Germany, Austria, Czechia, Pyrenean Mountains, GB), occuring also in Nearctic,
Neotropic, Australia and New Zealand. M. willmanni N ORTON et al., 1996, is known
from the Nearctic, mainly Canada.
Mucronothrus nasalis (W ILLMANN , 1929) (4-2 c, 4-3 a-b)

1929 Malaconothrus (?) nasalis W ILLMANN, Bergens Mus. Arb. Nat. Rekke 5: 4.
1931 Mucronothrus rostratus T RÄGÅRDH, Zool. Faroes 49: 31.
1933 Mucronothrus nasalis: W ILLMANN, Z. Morph. Ökol. Tiere 27: 374.
1996 Mucronothrus nasalis: N ORTON et al., Can. J. Zool. 74: 928.
1997 Mucronothrus nasalis: W EIGMANN, Spixiana 20: 208.
A dul ts: Body length 580-800 lm; colour pale yellow. Pro do rsum : interlamellar setae
about 260 lm long, anterior exobothridial setae about 15 lm, posterior exobothridial setae
vestigial, sensillus about 35 lm long. Ve ntra l re gi on : normally 15-20 pairs of genital
setae (exceptionally 13-22).
Ju ve ni le s: As described in the genus diagnosis. Body length: larvae about 275 lm, pro-
tonymph about 330 lm, deutonymph about 435 lm, tritonymph about 530 lm (after
N ORTON et al. 1996).
H a bit at : In mosses and cold spring seepage of wet mountain, alpine and subarctic regions.
B io lo gy : Feeding on higher plant debris, fungal hyphae and diatoms. Parthenogenetic.
Di stri bu ti on: Cosmopolitan; Central European records from Germany, Czechia and
Austria.
Family Trhypochthoniellidae K NÜLLE, 1957

1957 Trhypochthoniellidae K NÜLLE , Mitt. Zool. Mus. Berlin 33: 150
Genus Trhypochthoniellus W ILLMANN , 1928

1928 Trhypochthoniellus W ILLMANN, Zool. Anz. 76: 2.
Typus generis: Trhypochthoniellus setosus W ILLMANN , 1928 by monotypy; syn. of T. long-
isetus (Berlese, 1904).
Di ag no sis of t he a du lt s: The family has one genus, thus the diagnosis of the adults is the
same for both (see family key). Notogaster: oval in shape, without transverse suture, cu-
ticula pale brown, structured partly netlike anteriorly, lateral and ventral region; middle
and posterior region coarsely pustulated; 14 pairs of smooth notogastral setae (f2 vestigial
or missing), most notogastral setae long (about 70-90 lm), c2 is the shortest (about 30-40
lm). Ve nt ral reg i on: Genital, adanal and anal plates in a longitudinal gap of the noto-
gaster shield, which is bent towards the ventral side, 10-18 pairs of genital setae, two pairs
of adanal setae, one pair of anal setae, no aggenital setae, epimeres II normally with one
pair of setae. Gna tho som a : Infracapitulum with diarthric articulation; legs: Each tarsus
with three claws, several short and stout setae close to the claws, genu of legs I and II each
with three setae and one solenidium.
Di ag no sis o f th e j uv en il e s: Facies similar to that of the adults with less sclerotization;

genital plates less developed.
Di stri bu ti on: A genus with worldwide distribution, with six species according to S UBIAS
(2004); several described synonyms; the genus needs to be revised. T. longisetus with re-
gionally distributed subspecies in different continents.
Trhypochthoniellus longisetus (B ERLESE , 1904) (4-2 e, 4-3 c-e)

1904 Trhypochthonius longisetus B ERLESE, Redia 2: 27.
1905 Nothrus crassus WARBURTON & P EARCE, Proc. Zool. Soc. London 2: 568.
1919 Camisia excavata W ILLMANN, Abh. Naturw. Ver. Bremen 24: 552.
1922 Tripochthonius trichosus S CHWEIZER, Verh. Naturf. Ges. Basel 33: 75.
1928 Trhypochthoniellus setosus W ILLMANN, Zool. Anz. 76: 3.
1997 Trhypochthoniellus longisetus W EIGMANN, Spixiana 20: 212.
1998 Trhypochthoniellus crassus S ENICZAK et al., Zool. Anz. 237: 87.
This parthenogenetic species occurs in two forms, partly within the same population: forma
typica with well developed fusiform sensilli, forma setosus without sensilli (W EIGMANN
1997), therefore the genus was included twice in the introduction key of families and gen-
era.
A dul ts: Body length 435-580 lm. Pro dor sum : Rostrum rounded, long interlamellar se-
tae (100-130 lm) close to the short exobothridial setae in T.l. forma setosus, close to the
bothridia in T.l. forma typica; rostral and lamellar setae of moderate length; se ns il lu s
smooth fusiform with pointed tip in T.l. forma typica, absent in T.l. forma setosus; speci-
mens with diminished sensilli or asymmetrical forms were found in some populations
(W EIGMANN 1997).
Ju ve ni le s: As described in the genus diagnosis. Body length: Larvae about 205 lm, pro-
tonymph about 280 lm, deutonymph about 365 lm, tritonymph about 455 lm. (after
S ENICZAK et al. 1998).
H a bit at : In floating Sphagnum mosses in wet mires and bogs; occasionally in wet mea-
dows and lakes.
B io lo gy : Parthenogenetic.
Di stri bu ti on: Worldwide (except Antarctica); widespread in Europe.
Family Malaconothridae B ERLESE , 1916

1916 Malaconothridae B ERLESE , Redia 12: 175
Genus Malaconothrus B ERLESE , 1904

1904 Malaconothrus B ERLESE, Redia 2: 24.
Typus generis: Nothrus monodactylus M ICHAEL , 1888; designated by B AKER & W HARTON
(1952).
Di ag no sis of th e ad ul ts: Without revision of the genus the differentiations between
genus and species characters are incertain; characters of M. monodactylus are described
below.
Di stri bu ti on: A genus with worldwide distribution, with 56 described species in two
subgenera according to S UBIAS (2004); the genus needs to be revised (all studied European
species are synonyms, as noted below).
98 Special part
Malaconothrus monodactylus (M ICHAEL , 1888) (4-2 d, 4-3 f-h)

1888 Nothrus monodactylus M ICHAEL, British Oribatidae II: 528.
1904 Lohomannia (sic!) (Malaconothrus) egregia B ERLESE, Redia 2: 24.
1952 Malaconothrus gracilis VAN DER H AMMEN, Zool. Verh. Leiden 17: 27.
1952 Malaconothrus processus VAN DER H AMMEN, Zool. Verh. Leiden 17: 25.
1952 Malaconothrus punctulatus VAN DER H AMMEN, Zool. Verh. Leiden 17: 27.
1957 Malaconothrus gracilis K NÜLLE, Mitt. Zool. Mus. Berlin 33: 156.
A dul ts: Body length 375-455 lm. Pro dor sum : Rostrum rounded, long interlamellar se-
tae (25-100 lm) and short exobothridial setae (mostly less than 25 lm) closely associated;
no bothridia and no sensilli. N oto ga ste r: Elongated with almost parallel sides, cuticula
pale brown, cerotegument formed by porose plaques; 15 pairs of notogastral setae. Ve n -
tr al r eg io n: Genital, adanal and anal plates in a longitudinal gap of the notogaster shield,
which is bent towards the ventral side, four to six pairs of genital setae, three pairs of adanal
setae, one pair of anal setae, no aggenital setae; epimeres II normally with one pair of setae.
Gn ath oso m a: Infracapitulum with anathric articulation. Legs: Each tarsus with one
claw.
Ju ve ni le s: The juveniles are not described adequately, as far as we know.
S im i la r sp e c ie s : K NÜLLE (1957) argued that all Malaconothrus spp. described from cen-
tral Europe belong to a single species. We come to the same conclusion (details in W EIG-
MANN 2006). Additional European species only described from the British isles (L UXTON
1987) need to be revised.
H a bit at : In reeds, bogs, wet meadows, fen woodlands.
Di stri bu ti on: Holarctic and Neotropic; Central European records from Germany, Po-
land, Denmark and Austria.
Genus Trimalaconothrus B ERLESE, 1916

1916 Trimalaconothrus B ERLESE, Redia 12: 336
Typus generis: Malaconothrus (Trimalaconothrus) indusiatus B ERLESE , 1916, by original
designation.
Di ag no sis of the a du lt s: P rod ors um : Connected to the notogaster by a soft ligament,
neither bothridia nor sensilla, long interlamellar setae close to the short exobothridial setae,
rostrum rounded; N ot og ast er : Without transverse suture, slightly flattened, 15 pairs of
notogastral setae. Ven tra l re g io n: genital, adanal and anal plates in a longitudinal gap of
the notogaster shield, which is bent towards the ventral side, without aggenital setae beside
the genital plates, 5-12 pairs of genital setae, one pair of anal and three pairs of adanal
setae; infracapitulum with anarthric articulation; epimeres II normally with one pair of
setae; L e gs: all with three claws.
Di ag no sis o f th e j uv en il e s: Facies similar to that of the adults with less sclerotization,
the posterior part of notogaster with scerotized integument and mostly with pits, before this
a transverse region with transverse folds, anterior part soft; genital plates less developed
(after S ENICZAK 1993).
Di stri bu ti on: A genus with worldwide distribution, with 78 described species in two
subgenera according to Subias (2004); the genus needs to be revised. Eight species in Cen-
tral Europe (W EIGMANN 2006).
Fig. 4-4: a-b, Trimalaconothrus maior: a, dorsal view; b, detail of notogastral ornamentation; c-e,
Trimalaconothrus foveolatus: c, dorsal view; d, anogenital region in ventral view; e, detail of noto-
gastral ornamentation; f, Trimalaconothrus sculptus: in dorsal view; g-h, Trimalaconothrus vietsi: g,
dorsal view; h, ventral view. Legs are detached. Figures f and h after K NÜLLE (1957).
100 Special part
Key to the European species (adults)

1 Notogaster dilated posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Notogaster not dilated, slender-oval or with parallel sides (5-6 pairs of genital setae) . . . . . 4
2 Notogaster ornamentation without foveolae (pitlike structures) but with punctuated cuticula
. . . . . . Trimalaconothrus spp. (predominantly terrestrial species from wet meadows or fens)
– Notogaster ornamentation with foveolae and punctuated cuticula . . . . . . . . . . . . . . . . . . . 3
3 Foveolae on notogaster quite large, without clear borders (distance between foveolae more or less
equal to the diameter of the foveolae); posterior part of notogaster with slight dents, anteriorly
with weak longitudinal ridges, distance between notogastral setae c1-d1 shorter than d1-e1; body
length 520-615 lm (4-4 a-b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. maior (page 100)
– Foveolae on notogaster smaller and at larger distances; notogaster without dents, distances
between notogastral setae c1-d1 and d1-e1 approximately the same; body length 380-440 lm
(4-4 c-e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. foveolatus (page 100)
4 Strong setae of notogaster and prodorsum ciliated, notogaster with longitudinal and transverse
ridges; 6 pairs of genital setae, one pair of long anal setae, length of genital plates equally to length
of adanal plates (4-4 f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. sculptus (page 101)
– All setae fine and smooth, notogaster without ridges; 5 pairs of genital setae, anal setae reduced,
genital plates longer than adanal plates (4-4 g-h) . . . . . . . . . . . . . . . . . . T. vietsi (page 101)
1. Trimalaconothrus foveolatus W ILLMANN, 1931 (4-4 c-e)

1931 Trimalaconothrus foveolatus W ILLMANN, Tierwelt Deutschlands 22: 106.
1957 Trimalaconothrus foveolatus K NÜLLE, Mitt. Zool. Mus. Berlin 33: 162.
A dul ts: Body length 380-440 lm. N oto ga ste r: Posteriorly dilated; cuticula pale brown,
punctuated with small foveolae (distance larger than their diameter); all setae smooth, 3
pairs of setae elongated (e2, h2, h3). Ve n tra l re g io n: Mostly 7 pairs of genital setae, anal
setae minute; normal epimeral setae formula 3-1-2-3.
Ju ve ni le s: General characters as indicated above in genus diagnosis. Body length: larvae
about 255 lm, protonymph about 295 lm, deutonymph about 345 lm, tritonymph about
420 lm (after S ENICZAK 1993); differential diagnosis to juveniles of T. maior incertain.
H a bit at : In floating Sphagnum moss in bogs.
Di stri bu ti on: Germany, Belgium, Poland and Austria.
2. Trimalaconothrus maior (B ERLESE , 1910) (4-4 a-b)

1910 Malaconothrus maior B ERLESE, Redia 6, 223.
1921 Malaconothrus novus S ELLNICK, Schr. Naturf. Ges. Danzig (NF) 15: 76.
1957 Trimalaconothrus novus K NÜLLE, Mitt. Zool. Mus. Berlin 33: 160.
A dul ts: Body length 520-615 lm. N oto ga ste r: Posteriorly dilated, cuticula pale brown,
punctuated with large foveolae (distance = diameter); all setae smooth, 3 pairs of setae
elongated (e2, h2, h3). Ven tra l re gi on: mostly 7-9 pairs of genital setae, anal setae re-
duced; epimeral setae formula 3-1-(2)3-3.
Ju ve ni le s: General characters as indicated above in genus diagnosis. Body length: Larvae
about 300 lm, protonymph about 375 lm, deutonymph about 455 lm, tritonymph about
610 lm (after S ENICZAK 1993).
Di stri bu ti on: Nearly cosmopolitan; Central European records from Denmark, Austria,
Poland and Germany.
3. Trimalaconothrus sculptus K NÜLLE , 1957 (4-4 f)

1957 Trimalaconothrus sculptus K NÜLLE, Mitt. Zool. Mus. Berlin 33: 163.
A dul ts: Body length about 390 lm. No to ga ste r: With almost parallel sides, remarkable
longitudinal and transverse ridges; cuticula pale brown, strong setae with short ciliation.
Ve nt ral r eg io n: Mostly 6 pairs of genital setae, one pair of long anal setae.
H a bit at : In Sphagnum and other mosses in bogs.
B io lo gy : Parthenogenetic (?).
Di stri bu ti on: Only recorded from Germany.
4. Trimalaconothrus vietsi (W ILLMANN , 1925) (4-4 g-h)

1925 Malaconothrus (Trimalaconothrus) vietsi W ILLMANN, Jahresb. Ent. Ver. Bremen 13: 8.
1957 Trimalaconothrus vietsi K NÜLLE, Mitt. Zool. Mus. Berlin 33: 165.
A dul ts: Body length 325-345 lm. N o tog a ste r: Cuticula pale brown, all setae smooth
and fine, almost parallel sides. Ve n tra l re g io n: 5 pairs of genital setae, anal setae re-
duced.
Juveniles are not described as far as we know.
B io lo gy : Parthenogenetic (?).
Di stri bu ti on: Only recorded from Germany.
Family Hydrozetidae G RANDJEAN, 1954

1954 Hydrozetidae G RANDJEAN , Bull. Soc. Zool. France 78: 436.
Genus Hydrozetes B ERLESE , 1902

1902 Hydrozetes B ERLESE, Zool. Anz. 25: 698.
Typus generis: Notaspis lacustris M ICHAEL , 1882, by original designation.
Di ag no sis o f the ad ul ts: Prod ors um : Rostrum rounded, tiny interlamellar setae, con-
verging lamellae flanked by less converging costulae (small ridges), no translamella. N o -
to ga ste r: Oval shaped, cuticula brown, no areae porosae or pteromorphs present, with 13
or more pairs of setae, with a photosensitive organ (lenticulus) at the anterior edge of the
notogaster, granulated stripes in shallow furrows at the lateral edges of the notogaster.
Ve nt ral re gi on: Genital and anal plates embedded in an undivided ventral shield,
one pair of aggenital setae, 6 or 7 pairs of genital setae with notable variation (4-9),
two pairs of anal and three pairs of adanal setae. L e gs: I-III in all species with a single
prominent claw. Living specimens with shining silvery stripes of a plastron on the noto-
gaster, the ventral plate and prodorsum.
Di ag no sis of th e j uve n il es : Opaque light-yellowish cuticula. N ot og ast er : Elongated
and rounded with a tuberculate sculpture, 13 pairs of notogastral setae, some of the pos-
terior ones elongated, normal setae often serrate. Pro dor sum : With smooth cuticula, fili-
form sensillus and short exobothridial setae. In the larval stage not with elongated noto-
gastral setae.
Di stri bu ti on: A single genus in the family with worldwide distribution, with 27 de-
scribed species in 3 subgenera according to S UBIAS (2004); the genus needs to be revised.
102 Special part
Fig. 4-5: a-b, g-h, Hydrozetes lacustris: a, H.l. forma parisiensis, dorsal view; b, H.l. forma typica,
dorsal view; g, tarsus of leg IV; h, leg IV; c-f, Hydrozetes lemnae: c, dorsal view; f, tarsus of leg IV; d,
Hydrozetes confervae: dorsal view of anterior body; e, i, Hydrozetes thienemanni: e, dorsal view of
anterior body; i, femur of leg IV. Legs are detached. Figures h and i after WALGRAM (1975). Abbrevia-
tion: len – Lenticulus.

1 Leg IV with normal claw and thin accessory claw, in contrast to normal setae the accessory claw is
bent distally (4-5 f); setae near the claws mostly pointed, sensillus clavate, 13 pairs of notogastral
setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– No accessory claw on leg IV (4-5g), setae near the claws with a few blunt ends, sensillus like a seta,
with a tiny rounded head or missing, 13 or more pairs of notogastral setae (4-5 a) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lacustris (page 103)
2 Femur of leg IV with three setae (4-5 i), notogaster with more elongated granulae in the marginal
plastron furrows (4-5 e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. thienemanni (page 105)
– Femur of leg IV with two setae (4-5 h), notogaster with round granulae in the marginal plastron
furrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Notogastral setae long, da-setae at least as long as the distance between da and dm, setae near the
claws short (10-20 % of claw length) (4-5 c) . . . . . . . . . . . . . . . . . . . H. lemnae (page 105)
– Notogastral setae short, da-setae shorter than the distance between da and dm, setae near the
claws long (30-40 % of claw length) (4-5 d) . . . . . . . . . . . . . . . . . H. confervae (page 103)
Key to the European species (nymphs)

1 Two pairs of elongated posterior notogastral setae (4-6 d) . . . . . H. thienemanni (page 105)
– More than two pairs of elongated posterior notogastral setae.. . . . . . . . . . . . . . . . . . . . . . 2
2 All notogastral setae of the anterior notogaster very short and thin (5 15 lm), except the 5 most
posterior pairs (4-6 a-b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. confervae (page 103)
– At least the lateral notogastral setae of the anterior notogaster longer than 15 lm . . . . . . . 3
3 More than three pairs of elongated posterior notogastral setae (4-6 e) H. lacustris (page 103)
– Three pairs of elongated posterior notogastral setae (4-6 c). . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lacustris or H. lemnae (page 103, 105)
1. Hydrozetes confervae (S CHRANK , 1781) (4-5 d, 4-6 a-b)

1781 Acarus confervae S CHRANK, Enumeratio Insectorum Austriae indigenorum 511.
1910 Hydrozetes terrestris B ERLESE, Redia 6: 266.
A dul ts: Body length 490-570 lm. Prodorsum: Rounded rostrum, sensillus clavate, well-
developed. N o tog a ste r: Lateral longitudinal groves with round granules, 13 pairs of
short notogastral setae. Ve ntr al re gi on: Most individuals with 6 pairs of genital setae;
L e gs: Femur of leg IV with 2 setae, leg IV with thin accessory claw; setae closest to the claw
pointed and comparatively long (30-40 % of claw length).
Ju ve ni le s: General characters as indicated above in genus diagnosis. Nymphs with 3 pairs
of elongated setae posteriorly. Protonymphs have 1 pair, deutonymphs 3 pairs and trito-
nymphs 5 pairs of genital setae.
H a bit at : Lakes, ponds, also in slowly flowing water; abundant especially on duckweed
(Lemna).
B io lo gy : Bisexual. All adult Hydrozetidae use a plastron (a thin air layer in connection
with the tracheae) for respiration. The plastron covers only parts of the cuticula. The ju-
veniles possess no plastron and use cutaneous respiration. H. confervae was observed feed-
ing on tissue of duckweed but seems not to be specialised on this food.
Di stri bu ti on: Holarctic; Central European records from Poland, Germany and Austria.
2. Hydrozetes lacustris (M ICHAEL , 1882) (4-5 a-b, g-h, 4-6)

1882 Notaspis lacustris M ICHAEL, J. R. microsc. Soc. 2: 12.
1948 Hydrozetes parisiensis G RANDJEAN, Bull. Mus. nat. Hist. natur., 2e série 20: 328.
Two morphotypes of this species can be distinguished: Specimens of H. l. forma parisiensis
have an increased number of notogastral setae, generally 15-17 pairs. H. l. forma typica
show the normal number of 13 pairs of notogastral setae. Populations with intermediate
setae numbers found in Germany were the reason for synonymizing Hydrozetes parisiensis
(D EICHSEL 2005). The two morphotypes seem to prefer different habitats. To verify this
observation the distinction of both forms in ecological studies is helpful.
A dul ts: Body length 480-560 lm. P rod orsu m : Sensillus tiny or like a small seta, often
broken. Notogaster: Lateral longitudinal groves with round granules. Ve ntr al re gi on:
104 Special part
Fig. 4-6: a-b, Hydrozetes confervae: a, dorsal view, tritonymph; b, ventral view, tritonymph; c, Hy-
drozetes lemnae: dorsal view, tritonymph; d, Hydrozetes thienemanni: dorsal view, tritonymph;
e, Hydrozetes lacustris forma parisiensis: dorsal view, tritonymph. Legs are detached.
Most individuals with seven pairs of genital setae. L eg s: Femur of leg IV with two setae; leg
IV without accessory claw, setae closest to the claw short and blunt.
Ju ve ni le s: General characters as indicated above in genus diagnosis. Nymphs with three
to seven pairs of elongated setae posteriorly. In contrast to the other Hydrozetes-species
protonymphs have two pairs, deutonymphs four pairs and tritonymphs six pairs of genital
setae.
H a bit at : H. lacustris forma typica prefers floating Sphagnum moss in bogs and algae in
strongly acidic waters. Forma parisiensis was found in lakes and ponds; mainly on sub-
merse vegetation, but also on detritus.
B io lo gy : Parthenogenetic. For details on respiration, see H. confervae.
Di stri bu ti on: Holarctic: Central European records from Poland, Germany, Denmark,
Austria and France.
3. Hydrozetes lemnae (C OGGI , 1899) (4-5 c, f, 4-6 c)

1899 Notaspis lemnae C OGGI, In: C ANESTRINI , Prospett. Acarofauna Ital. 8: 916.
A dul ts: Body length 470-520 lm. P rod orsu m : Rostrum rounded, sensillus clavate, well-
developed. N ot og ast e r: Lateral longitudinal groves with round granules, 13 pairs of long
notogastral setae; Ven tra l re g io n: Most individuals with six pairs of genital setae. L e gs :
Femur of leg IV with two setae, leg IV with thin accessory claw, setae closest to the claw
pointed and comparatively short (10-20 % of claw length).
Ju ve ni le s: General characters as indicated above in genus diagnosis. Nymphs with three
pairs of elongated setae posteriorly. Length of the centro-dorsal notogaster setae is very
variable. Genital setae like H. confervae.
H a bit at : Lakes, ponds, also in slowly flowing water; mainly on submerse vegetation, but
also on detritus.
B io lo gy : Parthenogenetic. For details on respiration see H. confervae. This species is
known to feed on duckweed in South America (F ERNANDEZ and ATHIAS -B INCHE
1986). In Germany it lives primarily on submersed vegetation, where it must consume
also other ressources.
Di stri bu ti on: Cosmopolitan; Central European records from Poland, Austria, Denmark
and Germany.
4. Hydrozetes thienemanni S TRENZKE , 1943 (4-5 e, i, 4-6 d)

1943 Hydrozetes thienemanni S TRENZKE, Arch. Hydrobiol. 40: 57.
A dul ts: Body length 570-660 lm. P rod orsu m : Sensillus clavate, well-developed. N o -
to ga ste r: Lateral longitudinal groves with elongated granules, 13 pairs of long notogas-
tral setae. L e g s: Femur of leg IV with three setae, leg IV with thin accessory claw, setae
closest to the claw pointed.
Ju ve ni le s: General characters as indicated above in genus diagnosis. Nymphs with two
pairs of elongated setae posteriorly. Genital setae like H. confervae.
H a bit at : Lakes, ponds, also in slowly flowing water; mainly on submerse vegetation, but
also on detritus.
B io lo gy : Bisexual. For details on respiration see H. confervae.
Di stri bu ti on: Holarctic. Central European records from Poland, Germany, Austria,
Denmark and France.
106 Special part
Family Limnozetidae G RANDJEAN, 1954

1954 Limnozetidae G RANDJEAN , Bull. Soc. Zool. France 78: 436
Genus Limnozetes H ULL , 1916

1916 Limnozetes H ULL, Trans. Nat. Hist. Soc. Northumberland, N.S. 4: 391
Typus generis: Acarus ciliatus S CHRANK , 1803; designated by H ULL (1916).
Di ag no sis of t he ad ul ts: Pro do rsum : Sensillus like a drumstick, setiform or reduced;
tutorium and lamellae ribbonlike, lamellar setae on slim cuspides, genal process present at
lateral edge (4-7 b, d). N oto ga st er: oval shaped, anterior edge of notogaster entire, cu-
ticula light-brown , no areae porosae or sacculi, pteromorphs slender or triangular. Ve n -
tr al r eg io n: Genital and anal plates without sutures and embedded in an undivided ven-
tral shield, one pair of aggenital setae, six or seven pairs of genital setae, two pairs of anal
and three pairs of adanal setae. L e gs: with three claws.
Di ag no sis of th e ju ve n il es : The European species were not described adequately yet,
as far as we know. For North American species see B EHAN -P ELLETIER 1989.
Di stri bu ti on: Two genera in the family; Limnozetes with worldwide distribution in
aquatic habitats, with 14 described species according to S UBIAS (2004). Three described
species occur in Europe; L. amnicus from North America has been found in Spain, recently,
and might occur also in Central Europe.

1 Notogaster ornamentation with irregular lines anteriorly and irregular foveolae in the middle;
pteromorphs broadly triangular, approximately as long as wide in lateral view; anterior notogaster
edge hardly protruded (4-7 c-d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. rugosus (page 108)
– Anterior part of notogaster almost smooth with weak small foveolae, ornamentation of the no-
togaster close to the pteromorphs with blurred stripes; pteromorphs narrowly triangular, wider
than long in lateral view; incision (seen in dorsal view) behind the pteromorphs clear; anterior
notogaster edge protruded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Tutorium with anterior cusp, lamellar width about 10 lm, tarsus of leg III with 13 setae (4-7 a-b)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. ciliatus (page 106)
– Tutorial cusp absent, lamellar width about 6 lm, tarsus of leg III with 14 setae (4-7 e)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. amnicus (page 108)
1. Limnozetes ciliatus (S CHRANK , 1803) (4-7 a-b)

1803 Acarus ciliatus S CHRANK, Fauna boica 3,1: 214.
1880 Oribata sphagni M ICHAEL, J. R. microsc. Soc. 3: 179.
A dul ts: Body length: 270-330 lm; Pro dors um : Rostral seta close to tutorium, rostrum
anteriorly with two longitudinal ridges, Sensillus with capitate head, but often broken;
N ot og ast e r: Pteromorphs narrowly triangular; L e gs : Genu IV with 2 setae.
H a bit at : In floating Sphagnum in bogs.
Di stri bu ti on: Holarctic; Central European records from Poland, Germany and Austria.
Fig. 4-7: a-b, Limnozetes ciliatus: a, dorsal view; b, lateral view of anterior body; c-d, Limnozetes
rugosus: c, dorsal view; d, lateral view of anterior body; e, Limnozetes amnicus: dorsal view of anterior
body (after B EHAN -P ELLETIER 1989); f-g, Punctoribates sellnicki: f, dorsal view; g, lateral view of
prodorsum. Legs are detached.
108 Special part
2. Limnozetes rugosus (S ELLNICK , 1923) (4-7 c-d)

1923 Ceratozetes rugosus S ELLNICK, Acari (Lötzen) 1: 2.
A dul ts: Body length 350-365 lm. Prod ors um : Rostral seta in front of tutorium, rostrum
anteriorly with two joining longitudinal ridges, se nsi l lus with large capitate head. N o -
to ga ste r: Pteromorphs broadly triangular. L e g s: Genu IV without setae.
H a bit at : In floating Sphagnum in bogs and in submersed plants in oligotrophic Lakes.
Di stri bu ti on: Central and Northern Europe; Central European records from Denmark,
Netherlands, Germany, Austria and Poland.
3. Limnozetes amnicus B EHAN -P ELLETIER , 1989 (4-7 e)

1989 Limnozetes amnicus B EHAN -P ELLETIER , Can. Entomol. 121: 496.
A dul ts: Body length 345-390 lm. Pro do rsum : Rostrum rounded with a pair of conver-
ging ridges, sensillus well developed with large clavate to capitate head and minute barbs.
N ot og ast e r: Pteromorphs narrowly triangular. L eg s: Genu IV with two setae.
H a bit at : In cool, fast-flowing streams and creeks.
Di stri bu ti on: Described from North America; one European record in Peñalara, Spain
(S UBIAS & G IL -M ARTIN 1995).
Family Mycobatidae G RANDJEAN, 1954

1954 Mycobatidae G RANDJEAN , Bull. Soc. Zool. France 78: 439.
Genus Punctoribates B ERLESE , 1908

1908 Punctoribates B ERLESE, Redia 5: 6.
Typus generis: Oribates punctum C. L. K OCH , 1839, by original designation.
Di ag no sis of th e ad ul ts: Pr odo rsu m : Rostrum rounded, sensillus clavate ; lamellae
connected by a translamella, lamellar setae on cuspides. N o tog as te r: oval shaped, ante-
rior edge extended like a roof over the prodorsum connecting the pteromorphs, ptero-
morphs bent downwards, areae porosae present. Ve nt ra l re gi on: Genital and anal plates
without sutures and embedded in an undivided ventral shield, one pair of aggenital setae,
six or seven pairs of genital setae, two pairs of anal and three pairs of adanal setae. L e gs :
with three claws.
Di ag no sis of th e ju ve n il es : The European species were not described adequately yet,
as far as we know.
Di stri bu ti on: Genus with worldwide distribution mostly in terrestrial habitats, with 20
described species according to S UBIAS (2004). Three described species occur in Central Eur-
ope (W EIGMANN 2006).
Punctoribates sellnicki W ILLMANN , 1928 (4-7 f-g)

1928 Punctoribates sellnicki W ILLMANN, Abh. naturw. Ver. Bremen 27: 157.
A dul ts: Body length 320-380 lm. N o tog as te r: anterior edge with a convexly expanded
margin, without indentation; notogastral setae minute. Pr odo rsu m : rostral setae clearly
visible, lamellar setae long, clearly projecting over rostrum tip, lamella and translamella
well developed, distance between interlamellar setae narrower than between lamellar setae;
cuspides as long as the translamella, narrow; clavate sensilli reaching the translamella.
S im i la r s pe ci e s: Two predominantly terrestrial species: Punctoribates hexagonus B ER-

LESE , 1908: the bridge between the pteromorphs has two points with a concave median
indentation; with shorter cuspides. Punctoribates punctum (C. L. K OCH , 1839): the ante-
rior notogaster margin protrudes more than in sellnicki; the lamellar setae are very thin and
hardly project over the rostrum.
H a bit at : In floating Sphagnum in bogs; wet meadows and reeds.
Di stri bu ti on: Europe; Central European records from Denmark, Netherlands, former
Czechoslovakia, Poland, Germany, Austria and Belgium.
Family Zetomimidae S HALDYBINA , 1966

1966 Zetomimidae S HALDYBINA , Probl. poch. zool. 1966: 153.
Genus Heterozetes W ILLMANN , 1917

1917 Heterozetes W ILLMANN, Schr. phys.-ökon. Ges. Königsberg 58: 10.
Typus generis: Ceratozetes (Heterozetes) palustris W ILLMANN , 1917, by monotypy.
Di ag no sis of th e ad ul ts: N o tog as te r: Oval in shape, without transverse suture, cu-
ticula dark brown with smooth surface, 10 or 11 pairs of tiny notogastral setae, no sacculi
or areae porosae present, pteromorphs present. Ve nt ral re gi on : Genital and anal plates
embedded in an ventral undivided shield, six pairs of genital setae in a longitudinal row,
three pairs of adanal setae, two pairs of anal setae, one pair of aggenital setae. L e gs: Tarsus
of leg I with a single claw, the other tarsi (legs II-IV) with three claws.
Di ag no sis of th e ju ve n il es : Cuticula light yellowish, soft; Rounded body shape;
nymphs with very long serrate setae and anterior rostrum edge with many teeth; very
fine microtuberculation all over the body. Nymph of H. palustris first figured by H AARLØV
(1957).
Di stri bu ti on: Genus with 5 species in holarctic, oriental and neotropical region; one
European species according to S UBIAS (2004).
Heterozetes palustris (W ILLMANN , 1917)

1917 Ceratozetes palustris W ILLMANN, Schr. phys.-ökon. Ges. Königsberg 58: 10.
A dul ts: Body length 665-700 lm. Pro do rsum : Lamellae with short cuspides, without
translamella; sensillus fusiform and granulated; broad tutorium with a long free tip; ros-
trum with upright central ridge.
Ju ve ni le s: As described in the diagnosis of the genus.
H a bit at : In reeds and fens (“sphagnophob, acidophob” after P EUS 1932).
B io lo gy : Bisexual.
Di stri bu ti on: Palaearctic; Central European records from Denmark, Poland, Austria and
Germany.
Genus Zetomimus H ULL , 1916

1916 Zetomimus H ULL, Trans. Nat. Hist. Soc. Northumberland, N.S. 4: 395.
Typus generis: Oribata furcata P EARCE & WARBURTON , 1906, by original designation.
Di ag no sis o f th e a du lt s: P rod orsu m : Rostrum rounded with two small incisions,
long cuspides. N ot og ast er : oval in shape, without transverse suture, cuticula brown
with smooth surface, 10 pairs of notogastral setae, four pairs of areae porosae present,
pteromorphs present. Ve nt ral re gi on : Genital and anal plates embedded in a ventral un-
110 Special part
Fig. 4-8: a-c, Zetomimus furcatus: a, dorsal view; b, ventral view, tritonymph; c, dorsal view, trito-
nymph; d-f, Heterozetes palustris: d, dorsal view; e, ventral view, tritonymph; f, dorsal view, trito-
nymph. Legs are detached in most figures.
divided shield, six pairs of genital setae in a longitudinal row, three pairs of adanal setae,
two pairs of anal setae, one pair of aggenital setae. L eg s: tarsi of legs I and II with a single
claw, the tarsi of legs III-IV with one prominent middle claw and two thin lateral claws.
Di ag no sis of the j uv en il e s: Cuticula light yellowish, soft, with tuberculate structure;
oval body shape with typical T-like structure with dense microtuberculation and central
tubercle on the posterior body. Juveniles of Z. furcatus first figured by S HALDYBINA (1969).
Di stri bu ti on: Genus with 12 species in holarctic and neotropical region; two European
species according to S UBIAS (2004); one of these is a species inquirenda from GB, Z. boot-
hianus (H ULL , 1915).
Zetomimus furcatus (P EARCE & WARBURTON , 1906)

1906 Oribata furcata P EARCE & WARBURTON, Proc. Zool. Soc. London 1905: 564.
A dul ts: Body length 440-500 lm. P rodo rsu m : translamella like a line, sensillus clavate
and granulated, broad tutorium with a long free tip, anterior rostrum with two longitudinal
ridges.
Ju ve ni le s: As described in the diagnosis of the genus.
H a bit at : Reeds, fens, wet meadows, fen woodlands (“sphagnophob, acidophob” after
Peus 1932).
B io lo gy : Bisexual.
Di stri bu ti on: Palaearctic and Neotropic; Central European records from Austria, Den-
mark, Switzerland, Poland, former Czechoslovakia and Germany.
Acknowledgement
We thank Dr. D AVID RUSSELL , Görlitz, for linguistic correction of the manuscript.
References
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5. Acari: Halacaroidea 113
5. Acari: Halacaroidea
Ilse Bartsch
General part
The Halacaroidea include only a single family, the Halacaridae. They are exclusively
benthic organisms and form part of the meiofauna with a body length between 150-
2000 lm. Halacarids are primarily a marine taxon, as indicated by their name (Greek:
:ga ky = the sea; so ajaqi = the mite). In addition to more than 1000 marine species there
are about 60 species, which occur exclusively, mainly or at least regularly in fresh water.
K ARL V IETS (1927) established the subfamily Limnohalacarinae for the genera adapted to
life in freshwater, and K. V IETS (1933) introduced the term Porohalacaridae to include all
freshwater species with external genital acetabula, in contrast to the marine Halacaridae
with internal acetabula. However, the group of freshwater species proved to be very het-
erogeneous. On the one hand they show strong discrepancies on subfamily and genus level
and on the other hand they share features with various marine subfamilies and genera,
which cannot be considered as pure convergences (N EWELL 1947, 1959, 1984; B ARTSCH
1973; P ETROVA 1974, 1981). According to current nomenclature, freshwater halacarids
are partly grouped into limnic subfamilies. However, some species are assigned to marine
subfamilies and in single cases, freshwater species have even been assigned to marine gen-
era. With a growing interest for the meiofauna we should expect to find forms to be placed
between marine and limnic groups.
Halacarids differ from other mites in freshwater by a combination of the following fea-
tures: 1) cuticular plates present, but never strongly sclerotized; 2) units consisting of epi-
meres I and II as well as III and IV usually separated by a gap, such that leg pairs I and II
always are directed forward, leg pairs III and IV backward, disconnected from the front
legs; 3) body with a maximum of five pairs of gland pores and dorsally with no more than
seven pairs of idiosomal setae; 4) palps with four segments; 5) legs of adults with six seg-
ments.
External morphology
Id io soma (5-1 a-b): The idiosoma bears plates. Dorsally, we usually find four plates: the
unpaired anterior dorsal plate (AD), the pair of ocular plates (OC) and the posterior dorsal
plate (posterodorsal plate) (PD). Two or more plates can be fused, and plate OC can be
largely reduced. The plates are separated by areas of striated cuticle. Some species possess
additional small platelets in the area of the striated cuticle.
The most frequent combination of the v e ntr al pl a te s includes an unpaired anterior epi-
meral plate (AE), a pair of posterior epimeral plates (PE) and a genital plate (GP). The body
usually terminates with the anal plate (AP) consisting of the anal pore and the anal sclerites.
In many species, the genital and the anal plate are fused, forming the genitoanal plate (GA).
In a few species all ventral plates are fused forming the ventral shield. In other species the
epimeral plates, each assignable to one of the legs, are separated from the genital plate. The
anal plate can be missing.
The do rsa l p la te s of many species are ornamented, foveate or reticulate. Many limnic
species of the marine genus Copidognathus are characterized by porose areolae and rosette
pores (5-1 g). Rosette pores have been described in strongly sclerotized marine species.
They consist of a central porus, the ostium (O), which extends to the depth of the plate,
Updated and authorized translation of the German contribution submitted in June 2001

https://doi.org/10.1007/978-3-662-55958-1_5
114 General part
Fig. 5-1: Halacarid mite; a, dorsal, b, idiosoma, ventral; c, gnathosoma, lateral; d, gnathosoma, ven-
tral; e, leg I, medial; f, tarsus I, lateral; g, integument with rosette pores, schematically (after N EWELL
1947 and C ROWE & C AMARA 1973, modified). a-f, Porolohmannella violacea; g, Copidognathus.
Abbreviations: gac genital acetabula pc porus canaliculus
A anus ge genu PD posterodorsal plate
AD anterior dorsal plate glp gland pore PE posterior epimeral plate
AE anterior epimeral plate GnB gnathosoma base PE-ds dorsal seta on the PE
bfe basifemur GO genital opening pgs perigenital seta
ca canaliculus gsc genital sclerite Prk procuticle
Ch chelicera Kr claw rs rostral setae
co cornea mxs maxillary setae so solenidion
ds dorsal seta O ostium ta tarsus
Epd epidermis OC ocular plate ti tibia
Epk epicuticle P-1, P-2, P-3, P-4 tfe telofemur
FB bipectinate seta first to fourth palp segment tr trochanter
GA genitoanal plate pas parambulacral setae
the procuticle, where it widens to form the alveolus. It is surrounded by fine canaliculi (ca)
(N EWELL 1947; C ROWE & C AMARA 1973). In species with a thin cuticle, as in brackish
water species, the canaliculi seem to reach the epicuticle, and ostia and alveoli are reduced
to foveae. The ornamentation of the ventral plates is usually weaker than that of the dorsal
plates.
The number of se ta e a nd g la nd po re s is low, compared with many other mite groups.
Halacarid mites have at most five pairs of gland pores (glp) and seven pairs of tiny dorsal
idiosomal setae (ds); the posteriormost pair of setae, the adanal setae, is inserted close to, or
on the, anal cone. Anal setae are unknown. The gland pores are located dorsally, rarely
marginally. The OC often bear one or two corneae (co) and one pore canaliculus (pc)
(‘slit organ’). A second pair of pore canaliculi is located on the anal plate besides the
anus. The AE usually bears three pairs of setae, the PE each bear dorsally one seta (PE-
ds), marginally and ventrally two or three setae in total. The AE of Copidognathus species
additionally bears a pair of epimeral pores (ep). The genital opening is covered by genital
sclerites (gsc) and is surrounded by the GP. In species with external genital acetabula (gac)
these are arranged on the genital sclerites or directly on the genital plate. The perigenital
setae are located in front, or to both sides, of the GO (pgs), in females 2-15 pairs, in males
always more. The subgenital setae (genital setae) (sgs), small setae or spines on the genital
sclerites, are not present in all species. The female ovipositor, or the male spermatopositor,
is visible through the transparent genital plate. Females and males have a similar appear-
ance. Differences exist primarily in the genital region and the females of many species are
slightly larger than males.
G n a t h o s o m a (5-1 c-d): The gnathosoma consists of the gnathosomal base, the rostrum
and two pairs of appendages, the palps and the chelicerae. The rostrum forms a dorsally
open trough, which may be covered basally by the tectum. The gnathosoma bears two pairs
of maxillary setae and two pairs of rostral setae (mxs, rs). The palps of the European fresh-
water halacarids have four segments, called P-1 to P-4. The basal segment (P-1) is always
short, cylindrical. The second segment (P-2) is usually longer, sometimes also wider than
the other segments, dorsally it bears one or two setae. The P-3 is short and has a spine or
spine-like seta in the majority of species. The P-4 always bears several setae. The chelicerae
distally possess a claw, with a finely serrated dorsal edge. The claw can make up 1/4 of the
length of the chelicera, but may also be minute and less than 1/10 of the chelicera. In some
species the chelicerae are styletiform.
L e gs: The legs of the adults always have six segments (5-1 a, e), these are: trochanter,
basifemur, telofemur, genu, tibia and tarsus. Telofemora and tibiae are usually longer
than the other segments. Number, shape and arrangement of leg setae in the majority
of species are only slightly variable. Many species bear spines on leg I. Often the tibiae
ventrally bear elongate setae, which are either bipectinate (FB) or wide and somewhat ser-
rate. Tarsi I and II (5-1 f) carry three dorsal setae plus one solenidion (so); its position (dor-
solateral, dorsal or dorsomedial) has taxonomic implication. Distally, the tarsi possess
small setae, the parambulacral setae (pas), which are hollow (eupathid) and blunt on tarsi
I and II, but seti- or spiniform on tarsi III and IV. The dorsal fossa, into which the claws can
be retracted, may be flanked by membranes. Each of tarsi I to IV carry a pair of claws; the
small empodium (central sclerite, median claw) often terminates with a bidentate claw.
Between the end of the tarsus and the empodium a rod-like sclerite (carpite) may occur.
The claws carry pectines.
Juvenile stages
From the egg hatches a six-legged la rv a, with each of its legs five-segmented. Plates AD,
OC and PD are present but smaller than in adults, a genital plate is missing. The number of
116 General part
dorsal idiosomal setae and gland pores is equal to that of adults, the number of ventral setae
on the idiosoma and on the legs is still incomplete. The gnathosoma is smaller but its shape
and chaetotaxy equal those of adults.
The p roto ny m ph already has four leg pairs, legs I-III have six segments. The genital plate
is present, the genital acetabula are either external, situated on the genital plate, or internal,
close to the future genital opening. The number of setae is still incomplete.
The de ut ony m ph has eight legs, all with six segments. The dorsal and ventral plates of the
idiosoma are somewhat smaller than those of the adults. The genital acetabula are either
scattered on the genital plate or internal close to the genital slit; in the latter case there are
only two pairs of acetabula. The plate carries perigenital setae in addition to the genital
acetabula.
The tri to ny mp h resembles the deutonymph, but there are differences in the arrangement
of the genital acetabula. The number of leg setae is usually equal to that of the adults.
Anatomy and ultrastructure

A study on the structure of the inner organs is available for the marine species Thalassar-
achna basteri (J OHNSTON , 1836) (T HOMAE 1926). The structure of the integument and
some glands are described by C ROWE & C AMARA (1973), A LBERTI & S TORCH (1977)
and A LBERTI (1980, 1981). The idiosoma is filled by the large caeca of the midgut and
the sexual organs. The intestines are closed. Stored end-products (concrements), often re-
cognizable in the median axis of the idiosoma as white bar-shaped or spherical objects, are
eliminated through the anus. The structure of the integument of halacarids is the same as in
other mites. The cuticle is divided into pro- and epicuticle, with a further subdivision of the
procuticle into endo- and exocuticle, which may be visible. The integument of limnic spe-
cies is hydrophilic, whereas among marine taxa, species with a hydrophobic cuticle are
known.
Relations to the environment

Habitat
Halacarids are benthic organisms, unable to swim and they are bound to a substratum.
Their radius of activity is small. The freshwater species colonise the ground water, the hy-
porheic zone, sand filters or active carbon filters, the flocculent detritus of lentic or slowly
running waters, the spaces between moss leaves (Sphagnum, Fontinalis) and epiflora and
-fauna („aufwuchs“), e.g. bryozoans and colonies of the mussel Dreissena. Halacarids may
be also found in the interstitial system of vascular plant roots and inside structures of larger
animals, e.g. in the gill chamber of crustaceans.
European halacarids are free-living. Parasitic species or life stages are unknown. Specimens
found in the gill chambers of crustaceans only use the microcavernes offered. Halacarids
are unknown to cause damage to their hosts. Limnohalacarus-species have been found re-
peatedly in aquaria with ornamental fish (cichlids). Whether there is a relationship between
fish and halacarid is unknown.
Halacarids are absent from ephemeral waters, or from habitats that are regularly subject to
drastic environmental change such as shifting or silting-up of substrata, continuously de-
creasing oxygen or strongly fluctuating physical or chemical parameters. Nevertheless, the
halacarid species so far examined proved to be very tolerant towards chemical and physical
environmental conditions. For example a number of species live in boggy, acetic waterbo-
dies as well as in brackish, alkaline lakes and channels near the sea; they survive freezing but
also a rise in temperature of several hours caused by strong exposure to the sun. They tol-
erate a short-term decrease in oxygen of one or two hours, they even survive drying of
colonised substrata for hours or days, depending on the remaining humidity. Such tolerance
is usually higher than in other arthropods occurring in the same habitat, such as copepods
and ostracods. Halacarids are by no means rare: e.g. up to 500 specimens of Porohalacarus
alpinus brachypeltatus were extracted from a 50 ccm Sphagnum sample of a former peat
bog (B ARTSCH 1987), and up to 20 % of the bottom fauna captured by a 200 lm mesh in
the Danube river near Vienna consisted of Caspihalacarus hyrcanus (B ARTSCH & PANESAR
2000).
Feeding and food

The mouthparts of halacarids, the chelicerae, are positioned in the trough-shaped rostrum.
They can only be moved backward and forward, but not in a chewing motion. By repeated
back- and forward pushing the integument or cell walls of the prey organisms are pierced
and the contents is digested extraorally and then sucked up as known from other Arachni-
dae. There are only few observations concerning the food selection of the freshwater
halacarids, for example those of T ESCHNER (1963). Often only the colouring of particles
in the intestine and caeca of the midgut can provide information on the consumed food.
Small fragments of prey organisms, such as oligochaete setae, are rarely present. In some
species the first pair of legs is different from the following pairs; spines on these legs indicate
a predatory lifestyle. This assumption could be confirmed by the observation of marine
species (L OHMANN 1889, 1893; K IRCHNER 1969; B ARTSCH 1974; K RANTZ 1970;
M AC Q UITTY 1984).
The majority of halacarids are classified as predators and scavengers most likely feeding on
small metazoans and protozoans. Bacteria, fungi, algal and plant cells are certainly a food
source for several species. An intensively green coloured content of the intestine points to-
wards the ingestion of plant cells.
Enemies, epibionts and competition

Populations of halacarids are decimated by predators, such as carnivorous invertebrates,
including other mites, and small fish. External or internal parasites are unknown in hala-
carids. Fungi that were observed in dead halacarids certainly colonised the mite after their
death. Epizoans, in particular suctoria and peritrich ciliates are often found. Many marine
halacarids are covered by algae, especially densely packed diatoms attached to the hala-
carid mite can completely cover the idiosoma. Diatoms can be found to colonise also fresh-
water halacarids, but in lower densities. The idiosoma and legs of some species may be
partially covered by a bacterial film.
The presence of halacarids in a habitat might be less attributable to optimal environmental
conditions for the mites, but rather to low competition pressure. As compared to potential
competitors, in particular other arthropods, colonization and survival of halacarids is fa-
vored by their low degree of specialization. Though tolerating fluctuations of environmen-
tal conditions, drastic change of the environment and destruction of the colonised substrata
might have the greatest influence on the continued existence of halacarid populations.
Reproduction and development

Halacarids are o vi pa rou s. The European freshwater species pass through one larval stage
and one, two or three nymphal stages (proto-, deuto-, and tritonymph) before moulting to
the adult stage. Depending on the species, both sexes occur either in equal numbers, or
males are almost unknown. A population of Porohalacarus alpinus brachypeltatus con-
tained only one male among 2500 adults (B ARTSCH 1987). The majority of marine species
118 General part
and many of the limnic species are bisexual. Parthenogenetic reproduction is presumably
restricted to a few species.
As far as known, limnic genera produce only one generation per year. Gonad maturation of
the ground water mite Lobohalacarus weberi is triggered by the descent of temperatures in
the winter (T ESCHNER 1963). The reproductive cycle of some marine species appears to be
influenced by a change of both temperature and day length (B ARTSCH 1972). Eggs are often
deposited in springtime. A strict intrapopulational synchronisation of egg deposition has
not been found in limnic species so far.
Sperm is transferred indirectly by spermatophores. These are deposited by the males on the
colonised substratum and are absorbed by the genital opening of the females. Pocket-
shaped spermathecae have been mentioned for some marine species (N EWELL 1984;
A BÉ 1990a, b; B ARTSCH 1996b). In the majority of limnic species only one to five eggs
can be recognized inside the idiosoma; the period of maturation and egg deposition cor-
respondingly takes several weeks. The number of eggs per female is estimated a rather low
10-20.
Eggs are deposited into the colonised substrata: into the sediment, between “aufwuchs”
(epibionts) and underground, or into leaf axils of mosses. The genus Limnohalacarus is
an exception in which the cocoons are attached to, and carried on, the tibiae of the fourth
pair of legs. Each cocoon contains one embryo. Up to nine cocoons per leg have been ob-
served.
Larvae with six legs hatch from the eggs, or the cocoons respectively. After a few days, or
weeks, the larvae pupate. After a short pupal period, the first nymphal stage emerges from
the larval exuvia. Depending on the genus one or two further nymphal stages follow, each
after a further short pupal period, or directly the adult stage. Moulting of adults has not
been observed.
The development duration of individual stages is temperature-dependent. Development of
larvae takes from one week up to two months. The following nymphal stages mostly have a
longer development duration and the life span of adults is at least six to ten months before
they die after sperm or egg deposition.
Origin and phylogeny

All halacarids, also freshwater groups, are probably descendents of animals which have
evolved in brackish or sea water. The majority of the halacarids live in marine environments
where they colonised various benthic habitats and developed adaptations and different life-
form types. In a following step, originating from various clades, halacarids moved into
freshwater habitats (B ARTSCH 1982). Several of the extant limnic genera presumably al-
ready colonised the freshwater in the Mesozoic, probably even earlier (B ARTSCH 1996a),
such as Lobohalacarus, Ropohalacarus, Porohalacarus and Porolohmannella. Species of
these genera are characterised by their distribution across several continents. It is believed
that these species remained relatively unchanged over 50 million years. In contrast, fresh-
water species of other genera such as Caspihalacarus and Troglohalacarus have probably
evolved in the Tertiary, descending from species which survived the freshening of their mar-
ine habitats and adapted to a life in freshwater. In addition to species belonging to genera
exclusively or mainly living in freshwater habitats, we can also find members of otherwise
marine genera in the freshwater. These are Halacarellus, Copidognathus, Lohmannella and
Isobactrus, which include both species colonising habitats with salinities between marine
and fresh water, and species restricted to fresh or slightly brackish water. These limnic spe-
cies have probably become adapted to life in fresh water within the last one to five million
years.
Special part
Family Halacaridae M URRAY, 1877
1877 Halacaridae Murray, South Kensington Museum Science Handbooks: 95.
The identification keys refer to adults of the taxa found in fresh water. Genera diagnostics
were assembled with regard to characters of European limnic species. Seven subfamilies of
the Family Halacaridae are currently accepted: Halacarinae K.V IETS , 1927; Limnohalacar-
inae K.V IETS , 1927; Lohmannellinae K.V IETS , 1927; Rhombognathinae K.V IETS , 1927;
Porolohmannellinae K.V IETS , 1933; Copidognathinae B ARTSCH, 1983 and Ropohalacar-
inae B ARTSCH , 1989. However, neither identification keys nor descriptions are given at
subfamily level, because the definition of subfamilies hitherto described and the taxonomic
assignment of marine and limnic genera into the system of halacarids (see A BÉ 1998) re-
quires revision.
Species descriptions refer to adults. In addition, characters of juveniles are mentioned if
data are available. In addition to references to original descriptions, further relevant lit-
erature is listed. A bibliography can be found in K.V IETS (1956) and K.O. V IETS
(1987). An overview of freshwater species worldwide is given in B ARTSCH (1996a).
For ex pl an ati on s o f a bb re vi at io ns see Figs 5-1, 5-2 and 5-8.
The scale bars added to the figures of idiosoma correspond to 100 lm.
Area treated
In this survey are included all species recorded from European freshwaters.
Key to genera
1 Idiosoma with intensive green colouring. Ventral plates small, epimeral plates I and II in the me-
dian separated (5-27 b-c). Gnathosoma hidden under the idiosoma in dorsal view. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isobactrus (page 152)
– Idiosoma with yellowish, brown or slightly pink colouring, rarely slightly green. Epimeral plate I
and II always fused. In dorsal view, rostrum and palps distinctly visible . . . . . . . . . . . . . . . 2
2 Dorsal plates AD and PD fused to a shield; OC tiny (5-28 a). . . . Ropohalacarus (page 154)
– Dorsal plates AD, OC and PD separated; OC large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Palps attached dorsally to gnathosoma, distance between the P-1 smaller than width of P-1 . .
.................................................................. 4
– Palps attached laterally to gnathosoma, distance between the P-1 larger than width of P-1. . .
.................................................................. 9
4 Rostrum slender, at least twice as long as wide (5-25 e, 5-26 d), in ventral view often with parallel
margins. P-3 without or with a small spine-like extension . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Rostrum triangular in ventral view. P-3 with a strong spine (5-15 e, 5-18 c-d).. . . . . . . . . . 6
5 Plates PE with at least four setae, one dorsally, three ventrally (5-23 a-b). P-2 bears two setae, P-4 a
basal spur (5-23 c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lohmannella (page 146)
– Plates PE with three setae, a dorsal, marginal and ventral one (5-26 a-b). P-2 with one seta, P-4
with three slender basal setae but without spur (5-26 c) . . . . . . Porolohmannella (page 150)
6 External genital acetabula arranged along the lateral margins of the GP, or the part corresponding
to the GA, and anteriorly reaching beyond the GO (5-13 b, 5-14 b). GO in a terminal position.
Anal sclerites small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnohalacarus (page 134)
– External genital acetabula in the distolateral part of the GP (5-16 b, 5-18 b, 5-19 b), not reaching
beyond the GO. GO in a ventral position. Anal sclerites large. . . . . . . . . . . . . . . . . . . . . 7
7 All genua and tibiae with broad bipectinate spines (5-21 c). Claw pectines on tarsus I with their
broad tines arranged in the shape of a J. . . . . . . . . . . . . . . . . . . . Stygohalacarus (page 144)
120 Special part
– Setae shaped differently, on genua II-IV slender or spine-like, but not strongly bipectinate. Large
tines of claw pectines on tarsus I arranged in the shape of an umbrella. . . . . . . . . . . . . . . . 8
8 Second segment of the palps (P-2) with one seta (5-16 d). Genu I short, joint-like, its length less
than the height of the adjacent segments and less than 1/3 of the length of the tibiae (5-16 g). Tibia I
club-shaped.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parasoldanellonyx (page 137)
– Second segment of the palps (P-2) with two setae or short spines (5-18 c). Genu I not joint-like and
at least half as long as tibia I. Tibia I cylindrical. . . . . . . . . . . . . . Soldanellonyx (page 140)
9 Third segment of the palps (P-3) without spine (5-5 e).. . . . . . . . Copidognathus (page 120)
– Third segment of the palps (P-3) medially with a spine (5-6 d, 5-7 c, 5-8 d, 5-12 a). . . . . . 10
10 Tarsus I with one ventromedial seta and the pair of apical parambulacral setae (5-9 f). P-2 with one
seta each in the basal and the distal half (5-9 c). . . . . . . . . . . . . . Porohalacarus (page 130)
– Tarsus I with a bristle-like or spine-like ventromedial seta, two or more small sensorial setae and
the apical pair of parambulacral setae (5-6 f; 5-7 e, 5-8 f, 5-12 c). P-2 with only one seta.. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Ventral plates fused (5-8 b). Anterior margin of AD projecting in a frontal spine (5-8 a-b). Genu I
longer than telofemur I (5-8 e) . . . . . . . . . . . . . . . . . . . . . . . . . Lobohalacarus (page 128)
– Ventral plates separated. Anterior margin of AD mostly rounded or truncated, rarely forming a
frontal spine. Genu I shorter than telofemur I.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Genital acetabula of females inside the genital opening, in ventral view visible through the trans-
parent genital sclerites.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halacarellus (page 126)
– Genital acetabula of females on the genital plate or the genital sclerites (5-6 b). . . . . . . . . 13
13 Telofemur I ventrally with two short spines (5-12 b) . . . . . . . . . Troglohalacarus (page 133)
– Telofemur I ventrally with one slender bristle (5-6 e). . . . . . . . . . Caspihalacarus (page 125)
Subfamily Copidognathinae B ARTSCH, 1983

1983 B ARTSCH , Zool. Jb. Syst., 110: 186.
Genus Copidognathus T ROUESSART, 1888

1888 Copidognathus T ROUESSART, C. r. Acad. Sci. Paris 107: 754.
1947 Copidognathus N EWELL , Bull. Bingham oceanogr. Coll. 10: 129.
1983 Copidognathus B ARTSCH , Zool. Jb. Syst. 110: 187.
Typus generis: Copidognathus glyptoderma T ROUESSART, 1888, by original designation.
Di ag no sis: Dorsum with the plates AD, OC and PD, six pairs of idiosomal setae and up to
four pairs of gland pores. GP and AP always fused to form the GA. AE carries three pairs of
setae, PE one dorsal seta and three ventral setae, GA of females three, rarely four pairs of
setae and the genital sclerites a pair of sgs. The number of pgs of males (4-40 pairs) is always
higher than in females of the same species; the genital sclerites carry three to four pairs of
sgs. Female and male with one pair of internal genital acetabula, visible in the inside of the
genital opening. Palps four-segmented, attached laterally. P-2 carries one dorsal seta, P-3
none, P-4 three basal setae. One pair of maxillary setae on the base of the gnathosoma, one
pair on the rostrum. Genua shorter than telofemora or tibiae. Genua I and II each with four
setae, genua III and IV each with three setae. Number of dorsal/ventral setae on the tibiae I-
IV: 4/3, 4/3, 3/2, 3/2. One of the ventral setae always smooth, the others often bipectinate.
The claws carry an accessory process and usually also a pecten. Empodium small, mostly
bidentate.
R em a rks : One larval stage and only one nymphal stage, the protonymph, occurs during
ontogeny. The genus Copidognathus is primarily a marine taxon, and makes up almost one
third of the marine halacarids with respect to number of species. So far only five species
have been collected in freshwater habitats, one in waters in proximity to the coast in the
Northeast of America, four in Southeastern Europe, in Italy, Croatia, Macedonia and coun-
tries of the Black Sea region. Three of the European species, Copidognathus dactyloporus,
C. hephaestios and C. tectiporus, differ only slightly in body length, ornamentation of the
plates, shape of epimeral pores, and length: height ratio of telofemora. Studies of large
amounts of material are required for a strict differentiation or synonymisation of the species.
Key to species
1 Sculpturing of dorsal plates consisting of small foveae. Plate OC without eye pigment. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Copidognathus profundus (page 123)
– Plate AD with at least three round or ovate porose areolae, PD with at least two porose costae;
these with rosette pores. OC with distinct eye pigment.. . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Telofemur I slender, 2.2 times as long as high. Length of idiosoma 4 370 lm. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Copidognathus dactyloporus (page 121)
– Telofemur I 1.7-1.8 times as long as high. Idiosoma shorter, up to 360 lm. . . . . . . . . . . . . 3
3 Diameter of the epimeral pore 13-15 lm. . . . . . . . . . Copidognathus hephaestios (page 122)
– Diameter of the epimeral pore 20-25 lm. . . . . . . . . . . Copidognathus tectiporus (page 123)
1. Copidognathus dactyloporus B ENFATTI , M ARI & M ORSELLI , 1989 (5-2 a-e)

1989 Copidognathus dactyloporus B ENFATTI et al., Boll. Zool. 56: 100.
De sc ri pti on : Length of idiosoma 373-407 lm. Porose areolae of dorsal plates with ro-
sette pores. AD with three rounded porose areolae. Anteriorly broadly truncated PD with
two pairs of porose costae; medial costae three to four rosette pores wide. Ventral plates
with pore groups; in the central area of AE 8-15 canaliculi per pore group. Diameter of
epimeral pore 18-20 lm; pore surrounded by 9-15 teeth (5-2 c). Female with six, male with
16-25 pgs. Rostrum short, not reaching the end of the P-2. Telofemur I slender, length/
height ratio 2.2. Tibiae I and II each ventrally with two bipectinate setae and one smooth
bristle, tibiae III and IV each with a bipectinate seta and a smooth bristle. Tarsus III dorsally
with four, tarsus IV with three setae. Pecten of posterior legs with many small tines.
Fig. 5-2: Copidognathus dactyloporus; a, idiosoma, dorsal; b, idiosoma with gnathosoma, ventral,
male; c, epimeral pore; d, genitoanal plate, female; e, leg I, medial (ep, epimeral pore) (after B ENFATTI
et al. 1989).
122 Special part
Pr ea du lt sta g es : Plate PD of protonymph and larva distinctly shorter than in adults,

anteriorly pointed-ovate. Plate GP of the nymphs large. Tibiae I and II each ventrally
with one bipectinate and one smooth bristle. Number and arrangement of setae on tibiae
III and IV corresponding to adults.
H a bit at a nd B i ol og y: In springs, between fine and coarse sand, vascular plants and
mosses. Salinity at the only known collecting site 0.3 ‰.
Di stri bu ti on: Northern Italy, near Verona.
2. Copidognathus hephaestios B ENFATTI , M ARI & M ORSELLI , 1992 (5-3 a-e)

1992 Copidognathus hephaestios B ENFATTI et al., Boll. Zool. 59: 107.
De sc ri pti on : Length of idiosoma 324-360 lm. With rosette pores in the porose areolae.
AD with three rounded porose areolae. OC with strong eye pigment. PD truncated ante-
riorly, with two pairs of porose costae, width of the medial costae three rosette pores. In-
tegument outside porose costae foveate. Marginal areas of ventral plates with rosette pores,
median areas with pore groups arranged in polygons, the AE with one to seven canaliculi
per pore group. Diameter of epimeral pores 13-15 lm; pores surrounded by 6-10 teeth (5-3
c). Anterior margin of GA of females and males truncated. Females with six, males with 20-
24 pgs. Rostrum short, does not reach end of P-2. Telofemur I 1.8 times longer than high.
Tibiae I and II each with two bipectinate setae and one smooth bristle; tibiae III and IV each
with a bipectinate and a smooth bristle. Tarsus III dorsally with four, tarsus IV with three
setae. Pectines of the posterior legs with many small tines.
Pr ea du lt st ag e s: Plate PD distinctly smaller than in adults, anteriorly pointed-ovate.
Tibiae I and II ventrally with only one bipectinate and one smooth bristle.
H a bit at an d B i ol og y: Between sand and debris in lakes with Myriophyllum and Phrag-
mites.
Di stri bu ti on: Central Italy, north of Rome.
Fig. 5-3: Copidognathus hephaestios; a, idiosoma, dorsal; b, idiosoma with gnathosoma, ventral,
male; c, epimeral pore; d, genitoanal plate, female; e, leg I, medial (after B ENFATTI et al. 1992).
Fig. 5-4: Copidognathus profundus; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, part of the AE
with epimeral pore; d, gnathosoma, lateral; e, tarsus I, lateral; f, leg I, medial.
3. Copidognathus profundus (K.V IETS ) (5-4 a-f)

1936 Copidognathopsis tectiporus profundus K.V IETS, Arch. Hydrobiol. 29: 391.
1996 Copidognathus profundus B ARTSCH , J. nat. Hist. 30: 80.
1999 Copidognathus profundus B ARTSCH, Mitt. hamb. zool. Mus. Inst. 96: 174.
De sc ri pti on : Length of idiosoma 312-335 lm. Dorsal plates with small foveae, ventral
plates faintly reticulate; porose areolae absent. Rostrum short. Telofemur I with length/
height ratio of 2.0. Tibiae I and II each ventrally with two bipectinate setae and one smooth
bristle, tibiae III and IV each with one bipectinate seta and one smooth bristle. Tarsi III and
IV each with three dorsal setae, with the basal seta being thicker than the following, paired
setae. Claws slender. Pectines reduced.
H a bit at a nd B io lo gy : Only found in a lake at a a depth of 80-100 m.
Di stri bu ti on: Macedonia, Lake Ohrid.
4. Copidognathus tectiporus (K.V IETS, 1935) (5-5 a-h)

1935 Rhombognathopsis tectiporus K.V IETS , Zool. Anz. 109: 36.
1936 Copidognathopsis tectiporus K.V IETS, Arch. Hydrobiol. 29: 390.
1999 Copidognathus tectiporus B ARTSCH, Mitt. hamb. zool. Mus. Inst. 96: 17.
De sc ri pti on : Length of idiosoma 278-360 lm. Porose areolae with rosette pores. Ante-
rior portion of AD and OC each with a spot of strong eye pigment. AD with four porose
areolae, three ovate and one transversely elongated on the posterior margin of the plate. PD
with two pairs of porose costae; width of medial costae three to five rosette pores. Ventral
plates with shallow porose foveae. Diameter of epimeral pores 20-25 lm, pores surrounded
by 6-12 teeth. Females with six, males with approximately 20 pgs. Rostrum slightly shorter
than base of gnathosoma. Length/height ratio of telofemur I 1.8. Tibiae I and II each ven-
124 Special part
Fig. 5-5: Copidognathus tectiporus; a, dorsal; b, idiosoma, ventral, female; c, part of the AE with
epimeral pore; d, genitoanal plate, male; e, gnathosoma, lateral; f, gnathosoma, ventral; g, tarsus
I, lateral; h, leg I, medial.
trally with two bipectinate setae and one smooth bristle, tibiae III and IV with one bipec-
tinate seta and one smooth bristle each. Tarsus III dorsally with four, tarsus IV with three
setae. Claws with fine pectines.
Pr ea du lt s ta ge s: Plate PD distinctly shorter than in adults, anteriorly ovate. Tibiae I and
II each ventrally with one bipectinate seta and one smooth bristle.
H a bit at an d B io log y : In fresh and brackish water in proximity to the coast, here mainly
in regions with at least periodically very low salinity. Copidognathus tectiporus inhabits the
mud between mosses and vascular plants.
Di stri bu ti on: Macedonia and Black Sea region (Bulgaria, Ukraine, Turkey).
Subfamily Halacarinae K.V IETS , 1927

1927 Halacarinae K.V IETS , Z. wiss. Zool. 130: 90.
Genus Caspihalacarus K.V IETS , 1928

1928 Caspihalacarus K.V IETS , Abh. naturw. Ver. Bremen 27: 71.
Typus generis: Caspihalacarus hyrcanus K.V IETS , 1928, by original designation.
Di ag no sis: In addition to AD, OC and PD further platelets in the striated integument.
Idiosoma dorsally with ds-1 to ds-6. Three pairs of setae on each of AE and PE, on the PE
one dorsally and two ventrally. Genital and anal plates fused. Females with two pairs of
large and one pair of small genital acetabula in the area of the GO; GA with two pairs of
pgs. Males with two pairs of large genital acetabula located posteriorly, outside of the GO;
GO surrounded by numerous pgs. Palps with four segments, attached laterally. P-2 dorsally
with one seta, P-3 medially with a spine, P-4 basally with three setae. Slender rostrum with
two pairs of maxillary setae. Genua distinctly shorter than telofemora and tibiae. Leg I with
smooth bristles. Tarsus I ventromedially with a spine-like bristle and several eupathid ven-
tral setae; tarsus II with eupathid ventral setae. Solenidion located dorsolaterally on tarsus
I, dorsomedially on tarsus II. Claws with pectines. Median claw present, but small.
R em a rks : Two nymph stages occur during ontogeny. Only one species known.
5. Caspihalacarus hyrcanus K.V IETS , 1928 (5-6 a-f)

1928 Caspihalacarus hyrcanus K.V IETS, Abh. naturw. Ver. Bremen 27: 73.
1989 Caspihalacarus hyrcanus B ARTSCH, Acarologia 30: 225.
1998 Caspihalacarus hyrcanus B ARTSCH, Mitt. hamb. zool. Mus. Inst. 95: 148.
2000 Caspihalacarus hyrcanus B ARTSCH & PANESAR , Natur Mus., Frankfurt 130: 258.
D e s c r i p t i o n : Length of idiosoma 447-540 lm. With three small, but intensively coloured
eye pigment spots, otherwise colour of the idiosoma yellowish, intestinal content with
slightly brown spots. Dorsal plates reticulate. OC rounded, with a cornea each. PD very
wide. Ventral plates AE, PE and GA separated. Genital sclerites of females with three pairs
of large genital acetabula; ovipositor reaches anterior margin of GA. GA of males charac-
terised by almost 100 pgs surrounding the GO. Spermatopositor visible through the plate.
Behind the GO two pairs of large exterior genital acetabula. Rostrum shorter than base of
gnathosoma. Palps extend beyond the rostrum. First pair of legs only slightly wider than the
following legs. Genu I ventrally with three bristles, one of the bristles conspicuously long.
Number of ventral bristles on tibiae I-IV: 6, 4, 2, 2. Claws of tarsi with pecten.
Pr ea du lt sta ge s: Deutonymph with two pairs, and protonymph with one pair of genital
acetabula. Genu I of deutonymph with three ventral bristles, one of these conspicuously
long. Tibia I of the deutonymph with two pairs of ventral bristles.
Habit at an d Bi ol ogy: Caspihalacarus hyrcanus was first found in the Caspian Sea, in
brackish water of 12.5 ‰ salinity. The majority of findings are from rivers. Most likely this
species inhabits unsorted sediments, well aerated and rich in organic material. Owing to the
spines present on the first pair of legs and setae of oligochaetes found in the intestine, C.
hyrcanus is classified as predatory and carnivorous.
Di stri bu ti on: Caspian Sea, Black Sea, and the rivers flowing into the Black Sea (Danube,
Dnjepr, Donets). In the Danube River just before Vienna (B ARTSCH & PANESAR 2000).
These findings are from an area once flooded by the Paratethys. Recently, Caspihalacarus
hyrcanus was reported from the Rhine in France, Germany and the Netherlands, where it is
regarded as a new colonist (BIJ DE VAATE et al. 2002, M ARTENS et. al. 2006). Juveniles were
126 Special part
Fig. 5-6: Caspihalacarus hyrcanus; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma, ventral; d,
idiosoma, ventral, male; e, leg I, medial; f, tarsus I, lateral.
reported from Wales (G REEN 1956), but a reexamination and identification is recom-
mended.
Genus Halacarellus K.V IETS , 1927

1927 Halacarellus K.V IETS , Z. wiss. Zool. 130: 91.
1933 Halacarellus S CHULZ , Schr. naturw. Ver. Schlesw. Holst. 20: 98.
1997 Halacarellus B ARTSCH , J. nat. Hist. 31: 1229.
Typus generis: Halacarus balticus L OHMANN , 1889, by original designation.
Di ag no sis: Dorsum with AD, OC, PD and six pairs of dorsal idiosomal setae. Gland
pores always located on the plates, not in the striated integument. AE with three pairs
of setae, PE with one dorsal and two to three ventral setae. GA of females with two to
three pairs of perigenital setae, GA of males with distinctly more pgs. Genital acetabula
small, in females always internal, in males mostly internal, rarely external and situated on
the posterior part of the genital sclerites. Gnathosoma slender. Palps with four segments,
attached laterally. P-2 dorsally with one seta, P-3 with a medial spine, P-4 basally with three
setae. The two pairs of maxillary setae usually on the rostrum. Genua always shorter than
telofemora or tibiae. Tibia I ventrally with smooth bristles or spines, tarsus I ventromedially
with a bristle or a spine, ventrally with two eupathid setae, tarsus II ventrally with 0-1 setae,
tarsi III and IV ventrally without setae. Solenidion of tarsus I on the dorsolateral margin of
the fossa, of tarsus II on the dorsomedial margin. Tarsi end with paired claws. Median claw
present but small.
R em a rks : In addition to the larva, two nymph stages occur. The majority of Halacarellus
species lives in marine habitats, most often in substrata rich in microcavernes. Only one
species has so far been exclusively reported from fresh or strongly freshened brackish water.
6. Halacarellus phreaticus P ETROVA , 1972 (5-7 a-e)

1972 Halacarellus phreaticus P ETROVA , Acarologia 13: 36.
1998 Halacarellus phreaticus B ARTSCH, Mitt. hamb. zool. Mus. Inst. 95: 164.
De sc ri pti on : Length of idiosoma 475-495 lm. Idiosoma mainly colourless. AD trun-
cated anteriorly. PD ovate. Ventral plates separated. Genital plate of female with two pairs
of setae. Palp segments P-3 and P-4 short. P-3 with a strong spine. Tectum truncated. Leg I
distinctly longer and wider than the following legs. Basifemur I with one, telofemur with
Fig. 5-7: Halacarellus phreaticus; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, gnathosoma,
ventral; d, tarsus I, lateral; e, leg I, medial.
128 Special part
three ventral spines. Genu I with one pair, tibia I with three pairs of spines. Tibia II ventrally
with three, tibiae III and IV with four bristles each.
Pr ea du lt s ta ge s: Plates smaller than in adults and number of setae and bristles on the legs
incomplete in comparison to adults. Basifemur I ventrally with a short spine in all stages.
S im i la r spe c ie s: Many of the Halacarellus species living in coarse sands of the North
Atlantic coastal regions also move into freshened habitats, along the groundwater horizon
(H. subterraneus S CHULZ , 1933) or in surface sediments (H. capuzinus (L OHMANN ,
1893)). These species are not described here.
H a bit at an d B io lo gy : Ground water in proximity to the coast, up to 120 km inland
(P ETROVA 1972; M ORSELLI & M ARI 1986; B ARTSCH 1998); in strongly freshened brackish
or fresh water.
Di stri bu ti on: Mediterranean (Italy) and Black Sea region (Bulgaria, Ukraine).
Genus Lobohalacarus K.V IETS , 1939

1939b Lobohalacarus K.V IETS , Abh. naturw. Ver. Bremen 31: 506.
Typus generis: Walterella weberi R OMIJN & K.V IETS, 1924, by original designation.
Di ag no sis: Idiosoma usually with frontal spine. Dorsum with seven pairs of idiosomal
setae, the 7th pair (the ads) located on the margin of the anal conus. Elongated OC with two
setae. Ventral plates fused to a shield. In the area of the AE three pairs of setae and one pair
of epimeral pores, in the area of the PE one dorsal, marginal and ventral seta. Females with
one to three external genital acetabula arranged on the genital sclerites. Gnathosoma slen-
der. Palps with four segments, attached laterally to the rostrum. One of the maxillary setae
on the base of the gnathosoma, one on the rostrum. P-2 with one seta, P-3 with one spine,
P-4 basally with three setae. Leg I slightly more robust than leg II. Telofemora and genua of
the legs almost equal in length. Genu and tibia I with ventral spines. Tarsus I with enlarged
lateral fossa membrane, a dorsolateral solenidion, three dorsal setae, a ventromedial spine,
a pair of ventral setae and the paired pas. Including the solenidion, tarsi II and III dorsally
each bear four, tarsus IV three setae, ventrally one seta each. Solenidion on tarsus II in
dorsomedial position. All tarsi with paired pas. Paired claws large. Median claw with upper
and lower tooth.
R em a rks : Two nymph stages occur during ontogeny. Only one species occurs in European
waters.
7. Lobohalacarus weberi (R OMIJN & K.V IETS , 1924) (5-8 a-f)

1924 Walterella weberi R OMIJN & K.V IETS, Arch. Naturgesch. 90: 217.
1939 Lobohalacarus weberi K.V IETS , Abh. naturw. Ver. Bremen 31: 506.
1947 Lobohalacarus weberi quadriporus WALTER , Verh. naturf. Ges. Basel 58: 236.
1963 Lobohalacarus weberi quadriporus T ESCHNER , Arch. Hydrobiol. 59: 73.
1975 Lobohalacarus weberi quadriporus B ARTSCH , Gewäss. Abwäss. 57/58: 35.
1989 Lobohalacarus weberi B ARTSCH , Acarologia 30: 223.
1995 Lobohalacarus weberi B ARTSCH , Ann. Cape Prov. Mus. (nat. Hist.) 19: 177.
De sc ri pti on : Length of idiosoma 300-450 lm. Idiosoma light-coloured, eye pigment
only slightly developed or absent. Dorsal plates with small foveae. Mostly two to three
pairs of external genital acetabula on the posterior part of the genital sclerites, rarely
one acetabulum per side. Genital opening of female surrounded by four to five pairs of
perigenital setae. Rostrum extends to the end of the P-2. Genu I ventrally mostly with
Fig. 5-8: Lobohalacarus weberi; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma, lateral; d,
gnathosoma, ventral; e, leg I, medial; f, tarsus I, lateral (ep, epimeral pore; St, frontal spine).
one spine and one bristle. Tibia I ventrally with a pair of spines and a pair of bristles. Tibiae
II-IV with 2, 1-2, 2 bipectinate setae.
Pr ea du lt sta ge s: Nymphs and larvae with separated ventral plates. Juvenile stages can
be identified as L. weberi on account of the frontal spine, the genua being as long as the
telofemora and the flap-like membrane on tarsus I.
S im i la r sp ec i es : Due to the number of genital acetabula, two subspecies had been se-
parated, L. weberi quadriporus with two pairs of genital acetabula and L. weberi weberi,
with three pairs of genital acetabula. However, the number of genital acetabula has proven
to be variable.
H a bit at an d B io lo gy : In subterranean as well as surface interstitial systems: regularly in
the ground water, in hyporheic habitats, in sandfilters of waterworks and in surface waters
within the detritus and between mosses. The majority of females from the ground water
have only two pairs of genital acetabula, those from surface waters have three pairs.
130 Special part
Although males have been mentioned (T ESCHNER 1963; H USMANN & T ESCHNER 1970)
they have not been described so far.
Di stri bu ti on: Europe. Also reported from North Africa and North America, and one
subspecies each from Japan and the South Atlantic (I MAMURA 1970; B ARTSCH 1995).
Genus Porohalacarus T HOR, 1922

1922 Porohalacarus T HOR , Nyt Mag. Naturvidenskab. 61: 91.
Typus generis: Halacarus alpinus T HOR, 1910, by original designation.
Di ag no sis: Dorsum with the plates AD, OC and PD and (five? to) six pairs of idiosomal
setae. Gland pores not pronounced. AE with three pairs of setae, PE with two to three setae.
GP of females with 5-15 pairs of pgs; the three to eight pairs of genital acetabula on the
genital sclerites. Males with 20-80 pgs loosely arranged around the GO. Genital acetabula
on or behind the genital sclerites. Gnathosoma elongated. Rostrum short. The two pairs of
maxillary setae inserted on the rostrum. The palps with four segments are attached later-
ally. P-2 dorsally with two setae, P-3 with a spine, P-4 basally with three setae. Legs slender.
Genua I and II shorter than telofemora and tibiae of these legs. Tibia I ventrally with three
to five bristles, tibia II with three and tibiae III and IV with two bristles each. Number of
ventral setae on tarsi I-IV (without pas): 1, 0, 0, 0; number of dorsal setae, including the
solenidia: 4, 4, 4, 3. Solenidion of tarsus I dorsolaterally, of tarsus II dorsomedially located.
Parambulacral setae of all tarsi paired. Claws large, empodium small, bidentate.
R em a rks : During ontogeny one larval and two nymphal stages (proto- and deutonymph)
occur.
Key to species and subspecies

1 Plate OC as long or longer than AD. Plate PD distinctly widened at the level where the third pair of
legs is attached. Ventral plates fused to a ventral shield. . . Porohalacarus gallicus (page 132)
– Plate OC shorter than AD. PD barely widened at the level where the third pair of legs is attached.
Ventral plates separated.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Length of idiosoma 300-370 lm. Distance between GO and anterior margin of GP of females
corresponds to almost length of the GO.. . . . . . . . Porohalacarus alpinus alpinus (page 130)
– Length of idiosoma 250-270 lm. Distance between GO and anterior margin of GA smaller than
length of GO.. . . . . . . . . . . . . . . . . . . . . . Porohalacarus alpinus brachypeltatus (page 132)
8. Porohalacarus alpinus (T HOR , 1910)

1910 Halacarus alpinus T HOR , Zool. Anz. 36: 348.
1922 Porohalacarus alpinus T HOR , Nyt Mag. Naturvid. 61: 110.
1973 Porohalacarus alpinus B ARTSCH, Entomol. Tidskr. 74: 116.
8a. Porohalacarus alpinus alpinus (T HOR , 1910) (5-9 a-f)

De sc ri pti on : Length of idiosoma 300-370 lm. Dark spots of eye pigment underneath AD
and OC. Anterior margin of the idiosoma broadly truncated. Dorsal plates reticulated. OC
shorter than AD; each with cornea. PD anteriorly truncated. Structure of the ventral plates
finely porose. PE each with one seta dorsally, marginally and ventrally. GP and AP of fe-
males separated; distance between GO and the anterior margin of GP corresponds to about
length of the GO. GP with mostly five pairs of pgs, genital sclerites each with four to eight
external genital acetabula. Rostrum short, only slightly extending beyond the middle of the
P-2. Legs slender. Tibia I ventrally with four to five bristles, tibia II with three bristles and
tibiae III and IV each with two bristles. Pectines with fine tines.
Fig. 5-9: Porohalacarus alpinus alpinus; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma, lateral;
d, gnathosoma, ventral; e, leg I, medial; f, tarsus I, lateral.
Pr ea du lt st ag e s: Outline of plates and their structure similar but smaller than in females.
Genital plate of the deutonymph with two pairs of pgs; the four to seven pairs of genital
acetabula arranged on the posterior part of the plate; plates of the protonymph with two
to three pairs of external genital acetabula.
S im i la r spe c ie s: The specimen which L OHMANN (1893) reports as Halacarus hydrach-
noides, most likely a deutonymph, is similar to P. alpinus; a reliable assignment to one or
the other subspecies is not possible.
H a bit at a nd B io lo gy : In oligo- and mesotrophic waterbodies, in channels and lakes, in
continental as well as in slightly brackish coastal lakes and ditches. It was found between
mosses in springs, epibionts („Aufwuchs“) on Phragmites, colonies of the mussel Dreisse-
na, and in aquaria. Also reported from charcoal filters in waterworks (H USMANN 1982).
Males have not been described so far.
Di stri bu ti on: Europe; nearshore areas of the Turkish Black Sea, Africa, North America
and Australia.
132 Special part
Fig. 5-10: Porohalacarus alpinus brachypeltatus; a, idiosoma, ventral, female; b, idiosoma, ventral, male.
8b. Porohalacarus alpinus brachypeltatus K.V IETS, 1927 (5-10 a, b)

1927 Porohalacarus alpinus brachypeltatus K.V IETS , Verh. int. Ver. Limnol. 3: 469.
1987 Porohalacarus alpinus brachypeltatus B ARTSCH , Arch. Hydrobiol. 111: 85.
De sc ri pti on : Length of idiosoma 250-270 lm. With dark pigment spots. Dorsal plates
reticulate, distinctly emphasising the longitudinal direction. PD anteriorly broadly trun-
cated. PE usually with two setae, one each marginally and ventrally. GP and AP of females
separated, GP with five pairs of pgs; distance between GO and anterior margin of GP smal-
ler than length of GO; elongated genital sclerites mostly with four genital acetabula. GP
and AP of males contiguous, GP with 10 pairs of pgs. Each with five to six genital acetabula
on two humps posterior to the GO. Rostrum short, extends just beyond the middle of the
P-2. Legs slender. Number of ventral bristles on tibiae I-IV: 4, 3, 2, 2.
H a bit at an d B i ol og y: Only found in acetic peatbogs. It may be an ecotype rather than a
subspecies. The body contents often have a greenish colour, indicating at least occasional
phytophagy. Males are very rare; in a population from a peat-cutting there was only one
male among 2500 adults (B ARTSCH 1981, 1987).
Di stri bu ti on: Belgium, Germany, Hungary, Italy.
9. Porohalacarus gallicus (M IGOT, 1926) (5-11 a-e)

1926 Walterella gallicus M IGOT, Bull. Soc. zool. Fr., Paris 51: 376.
1939 Lobohalacarus gallicus K.V IETS , Abh. naturw. Ver. Bremen 31: 508.
1965 Lobohalacarus gallicus A NGELIER , Annls Limnol. 1: 216.
1996 Porohalacarus gallicus B ARTSCH , J. nat. Hist. 30: 75.
1954 Lobohalacarus dolgarae G REEN , Proc. zool. Soc. London 124: 669.
1996 Porohalacarus dolgarae B ARTSCH , J. nat. Hist. 30: 75.
De sc ri pti on : Length of idiosoma 403-500 lm. Anterior margin of idiosoma with small
frontal spine or truncated. Dorsal plates finely foveate. OC large, longer than AD, with
small cornea and two setae. PD long and ovate. Ventral plates fused to a ventral shield.
GO of females surrounded by approximately 30 pgs, males with 70-80 pgs. External genital
acetabula on the genital sclerites. Rostrum short, extending to the middle of the P-2. Legs
slender. Leg I not larger than leg II. Tibia I ventrally with three bipectinate setae; tibiae II to
IV correspondingly with three, two and two ventral bristles, one of each of these bipecti-
Fig. 5-11: Porohalacarus gallicus; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, gnathosoma,
ventral; d, leg I, medial; e, tarsus I, lateral.
nate. Number of ventral setae on tarsi I to IV: 1, 0, 0, 0; number of dorsal setae (including
the solenidia): 4, 4, 4, 3. Solenidia of tarsi I and II short and baculiform. Claws on tarsus I
without distinct pecten, claws of legs III and IV with distinct tines.
S im i la r sp ec i es : Due to the number of setae on P-2, tarsi I and II and the shape of genua I,
B ARTSCH (1996a) assigned this species to the genus Porohalacarus. E UGENE A NGELIER
(1965) synonymised Lobohalacarus dolgarae (G REEN, 1954) with L. gallicus. This sugges-
tion was accepted by G LEDHILL & V IETS (1976) and B ARTSCH (1996a).
H a bit at a nd Bi ol og y: In sediments of lakes, caves and riverine sands, in lowlands as
well as up 2200 m altitude.
Di stri bu ti on: England, France, Spain.
Genus Troglohalacarus K.V IETS , 1937

1937 Troglohalacarus K.V IETS , Arch. Hydrobiol. 31: 561.
Typus generis: Troglohalacarus dentipes K.V IETS , 1937, by original designation.
Di ag no sis: Genital acetabula partly outside, partly inside the GO. Gnathosoma short,
four-segmented palps attached laterally. The two pairs of maxillary setae located on the
rostrum. P-2 with one seta, P-3 with one spine, P-4 basally with three setae. Genua shorter
than telofemora and tibiae. Tarsus I ventrally with one spine and two setae. Solenidion of
tarsus I located dorsolaterally, of tarsus II dorsomedially. Tarsi with claws and claw-like
empodium.
134 Special part
R em a rks : Up to now only one specimen is known. Many characters match those of
Halacarellus and the examination of more material may show that Troglohalacarus is
to be classified as Halacarellus.
10. Troglohalacarus dentipes K.V IETS , 1937 (5-12 a-c)

1937 Troglohalacarus dentipes K.V IETS , Arch. Hydrobiol. 31: 562.
1989 Troglohalacarus dentipes B ARTSCH , Acarologia 30: 225.
De sc ri pti on : Length of idiosoma 280 lm. OC without cornea and eye pigment. Rostrum
1.6 times longer than wide, triangular in dorsal view. First pair of legs slightly more robust
than the following. Setae on the legs short, ventrally bristle- or spine-like. Telofemur I ven-
trally with two small, compressed spines, telofemora II-IV each with one short spine. Tibia I
ventrally with three short spines and two bristles. Including the short solenidia, tarsi I to IV
carry dorsally 4, 4, 3 and 3 setae, the two distal setae are located on the large fossa mem-
branes.
H a bit at a nd B io lo gy : Reported from a cave.
Di stri bu ti on: Northeast Spain, near San Sebastian.
Subfamily Limnohalacarinae K.V IETS , 1927

1927 Limnohalacarinae K.V IETS , Z. wiss. Zool. 130: 95.
Genus Limnohalacarus WALTER , 1917

1917 Limnohalacarus WALTER , Rev. suisse Zool. 25: 412.
Typus generis: Halacarus wackeri WALTER , 1914, by original designation.
Di ag no sis: Dorsal plates AD, OC and PD reticulated. Internal concrements, if present, as
a rounded to egg-shaped body. Dorsum with two to three idiosomal setae, one pair minute
adanal setae (sometimes in ventral position) and five pairs of gland pores, the first on the
anterior margin of the AD, the second in the lateral area of the body margin, the third in the
distal area of the OC and the fourth and fifth in the striated integument next to the PD. Pore
canaliculi in the distal part of the OC. OC often divided transversely. Ventral plates finely
punctated, in some species fused to a shield. AE with three pairs of setae, PE mostly with
one seta dorsally, laterally and ventrally. External genital acetabula arranged on the lateral
margin of the GA (or the part that corresponds to the GA). Females with 3-10 pairs of pgs in
Fig. 5-12: Troglohalacarus dentipes; a, palp, medial; b, leg I, medial; c, tarsus I, lateral.
the area of the GP. Males always with larger number of pgs. Genital sclerites large, sgs
usually present. Anal sclerites small. Base of the gnathosoma wide, rostrum triangular
and according to the given species either broad or slender. The basal pair of maxillary setae
located on the base of the gnathosoma. Palps with four segments, attached dorsally. P-2
large, higher than the other palp segments. Basal seta of P-2 short, distal seta long. P-3
ventrally with a robust spine, P-4 with six setae and a spine. Segments of legs slender,
with long bristles. Genua I and II shorter than telofemora and tibiae of these legs. Tibia
III with two bipectinate setae, located one behind the other. Number of ventral setae on
tarsi I-IV: 1, 0, 0, 0; number of dorsal setae (including the solenidia): 4, 4, 4, 3. Solenidion
on tarsus I as well as on tarsus II located dorsolaterally. Tarsi with paired pas. Claws large,
empodium without tine. Claw pectines of tarsus I often different from those of the follow-
ing tarsi. Tines on claws II-IV always distinct.
R em a rks : Females often with egg cocoons attached to the tibiae IV. Limnohalacarus-spe-
cies therefore appear to perform brood care. A key to all species described so far is given by
B ARTSCH (1999).
Key to species
1 Ocular plate without subdivision (5-14 a). Ventral plates fused to a ventral shield. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnohalacarus mamillatus (page 136)
– Ocular plate with subdivision (5-13 a, 5-15 a). Ventral plates AE, PE and GP separated. . . . 2
2 Anterior ocular plate wider than long. Claw pecten of tarsus I J-shaped, with large tines (5-15 g).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnohalacarus wackeri (page 136)
– Anterior ocular plate rounded. Claw of tarsus I with small tines apically and a process basally
(5-13 d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnohalacarus inopinatus (page 135)
11. Limnohalacarus inopinatus FAIN & L AMBRECHTS , 1987 (5-13 a-d)

1987 Limnohalacarus inopinatus FAIN & L AMBRECHTS , Bull. Annls Soc. r. belge Ent. 123: 113.
De sc ri pti on : Length of idiosoma 285-308 lm. OC with subdivision, anterior part
rounded, posterior part triangular. PD long and ovate. Ventral plates separated. GA of
female with six to seven pairs of genital acetabula and three pairs of pgs; genital sclerites
Fig. 5-13: Limnohalacarus inopinatus; a, idiosoma with gnathosoma, dorsal; b, idiosoma, ventral,
female; c, palp; d, tibia and tarsus I, medial (after FAIN & L AMBRECHTS 1987).
136 Special part
Fig. 5-14: Limnohalacarus mamillatus; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, palp;
d, tarsus I (after FAIN & L AMBRECHTS 1987).
with two pairs of sgs. P-2 conspicuously wide in dorsal view. Tibiae I, II and IV each with
one bipectinate seta, tibia III with two bipectinate setae. Claws of all legs basally with a
process bearing two to three tines. Claws on tarsus I with few short tines, claws of the
following tarsi with numerous tines.
H a bit at a nd B io lo gy : The only finding is from aquaria with cichlids.
Di stri bu ti on: Belgium (Antwerp).
12. Limnohalacarus mamillatus FAIN & L AMBRECHTS , 1987 (5-14 a-d)

1987 Limnohalacarus mamillatus FAIN & L AMBRECHTS , Bull. Annls Soc. r. belge Ent. 123: 109.
De sc ri pti on : Length of idiosoma 295-310 lm. OC without subdivision, with eye pig-
ment. PD long and ovate. Ventral plates fused to a shield. Genital region of females
with five to six pairs of genital acetabula and three pairs of pgs, genital sclerites with
only one pair of sgs. P-2 not widened. Bristles on tibiae I and II not bipectinate, tibia
III with bipectinate setae. Claws of all tarsi with long tines.
H a bit at a nd B io lo gy : The only finding is from aquaria with cichlids.
Di stri bu ti on: Belgium (Antwerp).
13. Limnohalacarus wackeri (WALTER , 1914) (5-15 a-g)

1914 Halacarus wackeri WALTER , Arch. Hydrobiol. 9: 281.
1917 Limnohalacarus wackeri WALTER , Rev. suisse Zool. 25: 412.
1989 Limnohalacarus wackeri B ARTSCH , Acarologia 30: 229.
De sc ri pti on : Length of idiosoma 320-420 lm. Dorsal plates evenly reticulated. OC sub-
divided; anterior part wider than long, with cornea and eye pigment, posterior part small,
triangular. PD elongated; anteriorly rounded. Ventral plates separated. GA of females with
six to nine pairs of pgs; the large genital sclerites with four pairs of sgs. Males with 50-60
pgs. Males and females each with six to eight pairs of genital acetabula. Distal pairs of
maxillary setae very broad, moved dorsally. Tarsi I to IV and tibiae I and II each with ba-
siventral spines. Ventral bristles on tibiae I, II and IV long and slender, tibia III with two
long, slender and two very short, broadly bipectinated setae. Pectines of claws on tarsus I J-
shaped, the tines equal in length.
Fig. 5-15: Limnohalacarus wackeri; a, dorsal, posterior legs with cocoons; b, idiosoma, ventral,
female; c, genitoanal plate, male; d, gnathosoma without palps, dorsal; e, gnathosoma, lateral;
f, leg I, medial; g, tarsus I, lateral.
Pr ea du lt sta ge s: Deutonymphs with four pairs of pgs and five to six pairs of genital
acetabula, these arranged along the lateral margin of the plate.
H a bit at an d B i ol og y: Primarily an inhabitant of standing or slowly flowing waters, the
findings are from the bottom sediment also found in the gill chambers of crayfish.
Di stri bu ti on: Europe.
Parasoldanellonyx K. V IETS , 1929

1929 Parasoldanellonyx K. V IETS , Zool. Anz. 86: 28.
1989 Parasoldanellonyx B ARTSCH , Acarologia 30: 233.
Typus generis: Soldanellonyx parviscutatus WALTER , 1917, by original designation.
138 Special part
Di ag no sis: Dorsal plates reticulate; separated by large areas of striated integument. This
integument with papilliform thickenings. Small platelets present next to the dorsal plates
AD, OC and PD. Dorsum with five pairs of gland pores, five pairs of idiosomal setae and
one pair of adanal setae. First pair of gland pores on the anterior end of the AD, the others
in the striated integument. Ventral plates finely porose; AE, PE and GP separated. AE with
three pairs of setae and one pair of epimeral pores; PE usually with three setae, one each
dorsally, marginally and ventrally. In females the GP is separated from the AP. Genital ace-
tabula of females and males on the GP, distolateral of the large GO. GP of females with 4-10
pairs of pgs; on the genital sclerites three pairs of sgs. Males with more than 50 pgs, ar-
ranged on both sides of the GO. Palps with four segments, attached dorsally. Maxillary
setae in the distal half of the rostrum; one pair moved dorsally. P-1 very short, P-2
high and much larger and longer than the other palp segments. P-2 with a distodorsal
seta, P-3 with a spine, P-4 with four setae, two distal bristles and a spine-like process. Che-
licerae stylet-like. Telofemur and tibia I club-shaped, genu I spherical. Ventral bristles on
tibiae I-IV slender, not bipectinated. Number of dorsal setae on tarsi I to IV (including the
solenidion): 4, 4, 4(-3), 3; number of ventral setae 1, 0, 0, 0. Solenidion of tarsus I inserted
dorsolaterally, of tarsus II inserted dorsomedially. Tips of tarsi with paired pas. Paired claws
large; claws with pectines; tines of tarsus I arranged in umbrella-shape, and on the follow-
ing tarsi in J-shape. Empodium small, without tines.
R em a r k: Parasoldanellonyx can be distinguished from the similar genus Soldanellonyx by
the shape of the first pair of legs and the smaller size of the dorsal plates.
Key to species
1 Plate PD narrow, anteriorly ovate (5-16 a). Rostrum triangular in ventral view (5-16 c). Plate OC
with eye pigment. . . . . . . . . . . . . . . . . . . . . . . . Parasoldanellonyx parviscutatus (page 139)
– Plate PD wide (5-17 a). Apex of rostrum trunk-like (5-17 d). Plate OC without eye pigment.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parasoldanellonyx typhlops (page 138)
14. Parasoldanellonyx parviscutatus (WALTER , 1917) (5-16 a-g)

1917 Soldanellonyx parviscutatus WALTER , Rev. suisse Zool. 25: 420.
1919 Soldanellonyx parviscutatus WALTER , Rev. suisse Zool. 27: 241.
1929 Soldanellonyx (Parasoldanellonyx) parviscutatus K.V IETS , Zool. Anz. 86: 28.
1943 Parasoldanellonyx parviscutatus K.V IETS , Zool. Anz. 143: 244.
1955 Parasoldanellonyx parviscutatus S CHWOERBEL , Zool. Anz. 155: 148.
1959 Parasoldanellonyx parviscutatus K.V IETS , Veröff. Inst. Meeresforsch. Bremerhaven 6: 490.
De sc ri pti on : Length of idiosoma 480-566 lm. Idiosoma broad. With yellow to red spots
due to this colour in midgut caeca. Anterior end of the idiosoma truncated. AD wider than
long; the pair of gland pores in the anterolateral corners. OC rounded, often with eye pig-
ment. PD narrow, anteriorly ovate. Ventral plates separated. Females with three to nine
pairs of genital acetabula, 6-10 pairs of pgs and three pairs of sgs. Males with three to
five pairs of genital acetabula and almost 80 pgs; on the genital sclerites three pairs of
sgs. Genital acetabula, as well as the epimeral pores, twinned. Rostrum triangular in ventral
view. First pair of legs conspicuously differing in shape from the posterior legs. Telofemur I
dorsally and dorsolaterally together with six to nine short bristles. Number of slender ven-
tral bristles of tibiae I to IV: 3, 2-3, 3, 2-3. Claws of tarsus I with basal process; claws of the
following legs sickle-shaped, without such process.
H a bit at a nd B i olo gy : In subterranean as well as surface waters. Populations always
contain both females and males.
Di stri bu ti on: Europe, Siberia and North America.
Fig. 5-16: Parasoldanellonyx parviscutatus; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma

base and rostrum, ventral; d, gnathosoma, lateral; e, genitoanal plate, male; f, tarsus I, lateral;
g, leg I, medial (ep, epimeral pore).
15. Parasoldanellonyx typhlops K.V IETS , 1933 (5-17 a-e)

1933 Parasoldanellonyx typhlops K.V IETS , Zool. Anz. 102: 284.
1935 Parasoldanellonyx typhlops K.V IETS , Verh. int. Ver. Limnol. 7: 86.
1936 Parasoldanellonyx typhlops belgicus K.V IETS , Bull. Mus. r. Hist. nat. Belg. 12 (28): 7.
1948 Parasoldanellonyx typhlops S ZALAY, Fragm. faun. hung. 4: 76.
1959 Parasoldanellonyx typhlops K.V IETS , Veröff. Inst. Meeresforsch. Bremerhaven 6: 494.
De sc ri pti on : Length of idiosoma 370 lm. AD only slightly wider than long. OC
rounded, without eye pigment. PD anteriorly broad, rounded. Genital plates of females
140 Special part
Fig. 5-17: Parasoldanellonyx typhlops; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, genitoanal
plate, male; d, gnathosoma base and rostrum, ventral; e, leg I, medial.
and males with three pairs of genital acetabula. Males with about 70 pgs. Tip of rostrum
trunk-like.
H a bit at a nd B io lo gy : Groundwater and wells.
Di stri bu ti on: Europe. A subspecies is recorded from Japan (I MAMURA, 1957), a variety
from Lake Baikal (S OKOLOV & J ANKOVSKAJA, 1970).
Genus Soldanellonyx WALTER , 1917

1917 Soldanellonyx WALTER , Rev. suisse Zool. 25: 414.
1989 Soldanellonyx B ARTSCH , Acarologia 30: 233.
Typus generis: Soldanellonyx chappuisi WALTER , 1917, by original designation.
Di ag no sis: Dorsal plates reticulate, ventral plates finely punctate. Dorsum with five pairs
of gland pores, four to five pairs of idiosomal setae and a pair of adanal setae on the anal
plate. First pair of gland pores in the lateral margin of the AD, second to fifth pair in the
striated integument. AE with three pairs of setae; epimeral pores small, often not visible. PE
usually with three setae. GP separated from AP. External genital acetabula in the distolat-
eral area of the GP. Females with four pairs of pgs on the GP. Base of gnathosoma broad,
rostrum triangular in dorsal view. One pair of maxillary setae on the base, one pair on the
rostrum. Palps with four segments, dorsally attached. P-2 dorsally with two setae or short
bristles, P-3 ventrally with a spine, P-4 basally with three setae, apically with three setae and
one spine-like process. Genua shorter than tibiae. Number of ventral setae on tarsi I-IV: 1,
0, 0, 0; number of dorsal setae (including the dorsolaterally inserted solenidia): 4, 4, 4, 3.
Solenidion on both tarsi I and II located dorsolaterally. Claws with pectines; tines on tarsus
I arranged in an umbrella shape.
Key to species
1 Legs I and II similar in length. Telofemur I only slightly longer than high (length less than 1.5 times
the height). All tibiae with two ventral bristles, at least one of which is bipectinate. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Soldanellonyx monardi (page 142)
– Leg I distinctly longer than leg II. Telofemur I at least twice as long as high. Tibiae with more than
two ventral bristles, which are serrated but not bipectinate. . . . . . . . . . . . . . . . . . . . . . . 2
2 Idiosoma broad. Spine on P-3 on, or distally from, the middle of the segment, without socket.
Spines on leg I blunt . . . . . . . . . . . . . . . . . . . . . . . . . . Soldanellonyx chappuisi (page 141)
– Idiosoma slender. Spine on P-3 inserted in the basal half of the segment, on a socket. Spines on leg I
tapered. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Soldanellonyx visurgis (page 143)
16. Soldanellonyx chappuisi WALTER , 1917 (5-18 a-f)

1917 Soldanellonyx chappuisi WALTER , Rev. suisse Zool. 25: 414.
1959 Soldanellonyx chappuisi K.V IETS , Veröff. Inst. Meeresforsch. Bremerhaven 6: 488.
1975 Soldanellonyx chappuisi B ARTSCH , Gewäss. Abwäss. 57/58: 32.
Fig. 5-18: Soldanellonyx chappuisi; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma, lateral; d,
gnathosoma, ventral; e, leg I, ventromedial; f, tarsus I, lateral.
142 Special part
De sc ri pti on : Length of idiosoma 365-452 lm. Of yellow-brownish colour, due to the

visible body contents. Without eye pigment spots. AD almost square, anterior and posterior
margin truncated. Plates coarsely reticulate. Epimeral pores not visible. GP of females with
three to four pairs of pgs and four to eight pairs of genital acetabula, each acetabulum
surrounded by two to four little ‘warts’. Genital sclerites with three sgs each. Palps
long in relation to the rostrum. The two pairs of maxillary setae equal in shape. Rostrum
triangular in ventral view. P-3 with strong spine, which is located slightly posterior to the
middle of the segment. Leg I considerably longer and stronger than the following legs. The
third and fourth segment of leg I each with two long, slightly dentate ventral spines, the fifth
segment with four such spines. Number of short ventral spines on tibiae II to IV: 3, 3, 4;
number of short dorsomedial spines: 2, 1, 1. Tines of pectines of tarsus I large, on the fol-
lowing legs small.
S im i la r sp e c ie s : P ETROVA (1975) mentions the subspecies Soldanellonyx chappuisi tra-
cicus; but the differences with the nominate form, such as the number of genital acetabula
and the position of the bristles on P-2, do not require a taxonomic separation.
H a bit at a nd B i ol og y: Ground water, larger depths of lakes and streams near springs.
Lives in lowlands as well as at 2000 m altitude. Males are said to be abundant (H USMANN
& T ESCHNER 1970), but have not been described so far.
Di stri bu ti on: Europe; North America, Korea and Japan.
17. Soldanellonyx monardi WALTER , 1919 (5-19 a-f)

1919 Soldanellonyx monardi WALTER, Rev. suisse Zool. 27: 238.
1975 Soldanellonyx monardi B ARTSCH , Gewäss. Abwäss. 57/58: 28.
De sc ri pti on : Length of idiosoma 267-378 lm. Idiosoma with reddish-brown colour due
to the visible intestinal content, additionally with each small, brown eye pigment spot be-
low AD and OC. AD slightly longer than wide, truncated anteriorly. OC without corneae.
PD ovate. Dorsal plates reticulate. AE with epimeral pores. GP of females with three to five
pairs of setae and 4-10 pairs of genital acetabula. Subgenital setae often absent. Gnatho-
soma short. The pair of maxillary setae located dorsally slightly more enlarged than the pair
located basally. P-2 large, basally with a short bristle, distally a long seta. Legs I and II, with
the exception of the claws, similarly shaped. Telofemur I ventrally with a thin seta, genu I
with a pair of bristles, tibia I with a pair of spines. Tibiae II-IV each with a pair of ventral
bristles, of which the ventromedial is always distinctly bipectinate. Claws on leg I with tines
arranged in an umbrella shape; on the following legs the tines are large and arranged along
the claws.
R em a r k: S OKOLOV (1952) mentions the occurrence of 11-12 genital acetabula in males.
S im i li ar sp ec i es : Two subspecies of Soldanellonyx monardi were reported from Japan
and one from Java (B ARTSCH 1996a).
H a bit at a nd B i ol og y: Ubiquist, distributed in surface and subterranean waters, lentic
and lotic waterbodies, in slightly brackish coastal waters and dystrophic bog ponds, in the
lowlands as well as in the mountains; colonizing sediment, mud, aquatic plants and auf-
wuchs.
Di stri bu ti on: Europe; East Africa, North America, Hawaii, Japan and the Falkland
Islands (I MAMURA 1981; P UGH & D ARTNALL 1994).
Fig. 5-19: Soldanellonyx monardi; a, dorsal; b, ventral, female; c, gnathosoma, ventral; d, gnathoso-
ma, lateral; e, leg I, medial; f, tarsus I, lateral.
18. Soldanellonyx visurgis K.V IETS , 1959 (5-20 a-e)

1959 Soldanellonyx visurgis K.V IETS , Veröff. Inst. Meeresforsch. Bremerhaven 6: 485.
De sc ri pti on : Length of idiosoma 252-351 lm. Dorsal plates weakly reticulate. Posterior
margin of AD and anterior margin of PD truncated or broadly rounded. Females with three
to five genital acetabula on both sides of the GO. Each acetabulum surrounded by little
warts. P-2 dorsally with two short bristles, the basal bristle in the basal third of the segment.
The two pairs of maxillary setae slender. P-3 in lateral view slender, twice as long as high,
the spine located on a small socket in the basal half of the segment. Leg I distinctly longer
than leg II. Ventral and dorsal spines slightly dentated; bipectinate setae missing. Telofemur
and genu I each with two ventral spines, tibia I with four spines. On genua of the following
legs one spine each, on tibia II two and on tibiae III and IV each three spines. Tines on claws
144 Special part
Fig. 5-20: Soldanellonyx visurgis; a, idiosoma, dorsal; b, gnathosoma, lateral; c, idiosoma, ventral,
female; d, leg I, medial; e, tarsus I, lateral.
I arranged in an umbrella shape, tines of the following claws slightly more slender and
extending to the base of the claws.
H a bit at a nd B io lo gy : Littoral sediments and mosses.
Di stri bu ti on: Germany, Spain, Italy, Montenegro, Bulgaria. Also reported from North
America (B ARTSCH 1982).
Genus Stygohalacarus K.V IETS , 1934

1934 Stygohalacarus K.V IETS , Zool. Anz. 105: 137.
1934 Stygohalacarus K.V IETS , Zool. Anz. 106: 120.
1989 Stygohalacarus B ARTSCH , Acarologia 30: 231.
Typus generis: Stygohalacarus scupiensis K.V IETS , 1934, by original designation.
Di ag no sis: Dorsal plates AD, OC and PD reticulate. The first of the five gland pores
located on the lateral margin of the AD, the second marginally between AE and PE. In-
cluding the ads on the anal cone, six pairs of dorsal idiosomal setae. AE with small epimeral
pores and three pairs of setae, PE with each a dorsal, marginal and ventral seta. GP and AP
separated. External genital acetabula of males located in the distolateral part of the GP.
Palps with four segments, attached dorsally. P-2 wide. On P-2 dorsally two setae, on
P-3 one spine, on P-4 basally three setae, laterally one seta, distally two bristles and a spine-
like process. Genua I and II shorter than telofemora and tibiae of these legs. Fossa mem-
branes missing. Number of dorsal setae on tarsi I-IV (including the solenidia): 4, 4, 4, 3;
number of ventral setae: 1, 0, 0, 0. Solenidion on tarsi I and II located dorsolaterally. Claws
large, with pectines. Empodium without claw.
R em a rk: According to B ARTSCH (1989) Stygohalacarus is close to the genus Soldanello-
nyx but not to Limnohalacarus. The following characters are supportive of a relationship
with Soldanellonyx and against one with Limnohalacarus: 1) the arrangement of the geni-
tal acetabula on the GP (in the distolateral region instead of along the lateral margin of the
GP and extending beyond the GO); 2) the location of the GO (ventrally instead of termin-
ally) and 3) the shape of the anal sclerites (large instead of small).
19. Stygohalacarus scupiensis K.V IETS , 1934 (5-21 a-d)

1934 Limnohalacarus (Stygohalacarus) scupiensis K.V IETS , Zool. Anz. 105: 138.
1934 Stygohalacarus scupiensis K.V IETS , Zool. Anz. 106: 120.
1989 Stygohalacarus scupiensis B ARTSCH , Acarologia 30: 231.
De sc ri pti on : Length of idiosoma 400 lm. Corneae and eye pigment missing. The third,
fourth and fifth pair of gland pores located in the striated cuticle between PD and PE. Males
with approximately five pairs of genital acetabula in the distolateral region of the GP. Nu-
merous pgs arranged loosely on the GP. Genua I and II each with a pair of fan-shaped,
pectinate bristles, genua III and IV each only with one such bristle. On tibiae I-III ventrally
each three bristles, tibia IV with four bristles of which the distal pair is fan-shaped. Claws
with strong pectines; tines on the first pair of legs not as numerous but wider than those on
the following.
H a bit at a nd B io lo gy : Recorded from a well.
Di stri bu ti on: So far only known from Skopje, Macedonia.
Fig. 5-21: Stygohalacarus scupiensis; a, idiosoma, dorsal; b, gnathosoma, lateral; c, leg I, medial; d,
tarsus I, lateral.
146 Special part
Subfamily Lohmannellinae K.V IETS , 1927

1927 Lohmannellinae K.V IETS , Z. wiss. Zool. 130: 95.
Genus Lohmannella T ROUESSART, 1901

1901 Lohmannella T ROUESSART, Bull. Soc. amis sci. nat. Rouen, (sér. 4) 14: 250.
Typus generis: Leptognathus falcatus H ODGE , 1863, by original designation.
Di ag no sis: Idiosoma flat, broad. AD large, often hexagonal. Dorsum with the plates AD,
OC and PD, up to five pairs of gland pores and, including the ads, up to six pairs of idio-
somal setae. AE with three to four pairs of setae, PE with four to five pairs, of these one or
two dorsally, three ventrally. GP and AP fused, to form the GA. Genital acetabula partly
inside the genital opening, and partly on the genital sclerites. Females with two to about 10
pairs of pgs; in males many pgs arranged around the GO and none to three pairs outlying.
Gnathosoma with broad base and slender rostrum, the latter with parallel margins. Palps
attached dorsally, extending beyond the rostrum. P-2 bearing two setae, P-3 a spinelike
process, P-4 basally two to three setae and a spine. Legs slender. Genua shorter than tel-
ofemora and tibiae. Ventral bristles of the tibiae elongate, mostly bipectinate. Tarsus I ven-
trally with none to four bristles. Solenidion on tarsus I inserted dorsolaterally, on tarsus II
dorsally or dorsomedially. Tarsi with claws. Empodium without claw.
R em a rks : During ontogeny a larval stage and two nymph stages occur. The species of the
genus Lohmannella are primarily marine. Their appearance differs from members of the
very similar, exclusively freshwater genus Porolohmannella by the following features: 1)
the number of setae on the PE (four to five instead of three); 2) the shape of the bipectinate
setae (elongated instead of short); 3) the number of setae on P-2 (two instead of one) and 4)
the spine on P-4 (present as opposed to absent).
The marine species of the genus Lohmannella are known to be variable not only in body
size, but also in the number of leg setae. The descriptions of the limnic species are based on
only a few specimens, and the variability is unknown. Three of the four European species,
namely L. curvimandibulata, L. cvetkovi and L. stammeri, are very similar and a revision is
necessary to clarify the species status.
Key to species
1 Plate OC rhomboid (5-22 a). Tarsus I ventrally without setae or bristles. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lohmannella andrei (page 146)
– Plate OC drop-shaped (5-23 a), posterior corners pointed. Tarsus I ventrally with one or two bris-
tles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Telofemur I ventrally with one bristle. . . . . . . . . Lohmannella curvimandibulata (page 147)
– Telofemur I ventrally with two bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Plate GA anteriorly broad, truncated. The basal pair of maxillary setae located in the middle of the
rostrum. Genu I with four ventral bristles. . . . . . . . . . . . . Lohmannella cvetkovi (page 148)
– Plate GA ovate, anteriorly rounded. The two pairs of maxillary setae located in the anterior third
of the rostrum. Genu I with three ventral bristles . . . . . . . Lohmannella stammeri (page 149)
20. Lohmannella andrei (A NGELIER , 1951) (5-22 a-d)

1951 Porolohmannella andrei A NGELIER, Bull. Mus. natn. Hist. nat, sér. 2 23: 505.
1996 Lohmannella andrei B ARTSCH , J. nat. Hist. 30: 80.
De sc ri pti on : Length of idiosoma 360 lm. Yellow-orange colouring, without eye pig-
ment. AD large, longer than OC. Anterior margin of the PD broadly rounded. Length
of AE equal to GP. The males with about 30 pgs closely surrounding the GO; one pair
Fig. 5-22: Lohmannella andrei; a, idiosoma, dorsal; b, idiosoma, ventral, male; c, gnathosoma, lateral;
d, leg I, medial (after E.A NGELIER 1951).
of pgs outlying. Each genital sclerite with two genital acetabula. Gnathosoma long. P-2
curved. Telofemora I and II each with two ventral bristles, genua I and II each with
one pair of ventral bristles, tibia I with three long, serrated ventral bristles. Tarsi without
ventral bristles. Paired claws slender.
H a bit at a nd B io lo gy : River sediments.
Di stri bu ti on: France.
21. Lohmannella curvimandibulata (P ETROVA , 1969) (5-23 a-d)

1969 Porolohmannella curvimandibulata P ETROVA , Arch. Hydrobiol. 66: 103.
1996 Lohmannella curvimandibulata B ARTSCH , J. nat. Hist. 30: 81.
De sc ri pti on : Length of idiosoma 390-420 lm. Dorsal plates reticulate. AD rectangular.
OC drop-shaped, posterior angle pointed. Genital sclerites of females each with three sgs.
Anterior margin of the male GA ovate; two pairs of the pgs outlying, and more than 40 pgs
closely arranged around the GO. P-2 strongly curved. Telofemur I ventrally with a long,
almost smooth bristle, dorsally five setae, none of which is distinctly bipectinate. Genu I
ventrally with two pairs of bipectinate setae, tibia I with three pairs of such setae. Tarsus I
ventrally with two bipectinate setae, dorsally with three setae. According to P ETROVA
(1969) with external genital acetabula located on the genital sclerites.
H a bit at a nd B io lo gy : Recorded from wells.
Di stri bu ti on: Bulgaria.
148 Special part
Fig. 5-23: Lohmannella curvimandibulata; a, idiosoma, dorsal; b, idiosoma, ventral, male; c, rostrum
and palp, lateral; d, leg I, ventromedial (after P ETROVA 1969).
22. Lohmannella cvetkovi (P ETROVA , 1965) (5-24 a-e)

1965 Porolohmannella cvetkovi P ETROVA , Fauna Trakiya 2: 338.
1969 Porolohmannella cvetkovi P ETROVA , Arch. Hydrobiol. 66: 100.
1996 Lohmannella cvetkovi B ARTSCH , J. nat. Hist. 30: 81.
De sc ri pti on : Length of idiosoma 318-324 lm. Dorsal plates reticulate. OC drop-shaped,
posteriorly pointed. Outline of PD almost rectangular. First pair of gland pores on the la-
teral margin of the AD on the level of the ds-1. One pair of gland pores on the distal margin
of the PD. Anterior margin of the GA in males and females broad, almost truncated. Fe-
males with two pairs of pgs anterior to the GO. Males with about 40 pgs closely arranged
around the GO and with two pairs outlying. P-2 in lateral view slightly curved. The basal
pair of maxillary setae located in the middle of the rostrum. Telofemur I ventrally with two
bipectinate setae, dorsally three pennate setae and one smooth bristle. Genu I ventrally with
two bipectinate setae, tibia I with three pairs of bristles. Tarsus I dorsally with three setae,
ventrally with two bipectinate setae. According to P ETROVA (1969) with external genital
acetabula, two pairs in females, three pairs in males.
H a bit at a nd B io lo gy : Recorded from springs.
Di stri bu ti on: Bulgaria.
Fig. 5-24: Lohmannella cvetkovi; a, idiosoma, dorsal; b, idiosoma, ventral, female; c, idiosoma, ven-
tral, male; d, rostrum and palp, lateral; e, leg I, ventral (after P ETROVA 1969).
23. Lohmannella stammeri K.V IETS, 1939 (5-25 a-f)

1939 Lohmannella stammeri K.V IETS , Arch. Hydrobiol. 35: 625.
De sc ri pti on : Idiosomal length 290-310 lm. Dorsal plates almost smooth. Setae ds-1 very
long. OC inversely drop-shaped, posteriorly pointed; anterior region with gland pores and
seta. Anterior margin of PD broad, truncated. One pair of gland pores just behind the level
of insertion of the fourth pair of legs, one pair close to the posterior margin. AE with four
pairs of setae. GA of females rounded anteriorly; with five to seven pairs of pgs. GA of
males anteriorly ovate, about 30 pgs arranged closely around the GO and three pairs pe-
ripherally. Genital acetabula of females and males inside the genital opening, the three pairs
only slightly visible through the genital sclerites. P-2 curved, the two setae located in the
distal half. The two pairs of maxillary setae located in the anterior third of the rostrum.
Legs slender. Telofemur I bears ventrally two bipectinate setae, genu I three bipectinate
setae, tibia I two pairs of bipectinate setae and tarsus I one or two bipectinate setae. Num-
ber of ventral bipectinate setae on telofemur to tarsus II: 2, 2, 4, 2; bipectinate setae on
telofemur to tarsus III: 1-2, 1, 3, 2; and on leg IV: 2, 1, 2-3, 1. Claws slender, almost smooth.
With small carpite between end of tarsus and empodium. Genital acetabula located inside
the genital opening, only slightly visible through the genital sclerites.
H a bit at a nd B io lo gy : Reported from groundwater and a cave near the coast.
Di stri bu ti on: Italy, Montenegro (P EŠIĆ 2004).
150 Special part
Fig. 5-25: Lohmannella stammeri; a, dorsal; b, idiosoma, ventral, female; c, genitoanal plate, male;
d, leg I, medial; e, gnathosoma, ventral; f, gnathosoma, lateral.
Subfamily Porolohmannellinae K.V IETS , 1933

1933 Porolohmannellinae K.V IETS, Zool. Anz. 102: 284.
Genus Porolohmannella K.V IETS , 1933

1933 Porolohmannella K.V IETS , Zool. Anz. 102: 283.
Typus generis: Leptognathus violacea K RAMER , 1879, by original designation.
Di ag no sis: Idiosoma broad, flat. Dorsal plates reticulate. AD hexagonal. Dorsum with
five pairs of idiosomal setae, one pair of adanal setae and four pairs of gland pores. Ventral
plates large, AE bears three pairs of setae, PE dorsally, marginally and ventrally each with
one seta. Genital and anal plate fused. Females with two to three pairs of pgs. External
genital acetabula located on the genital sclerites. Gnathosoma, in particular the rostrum,
slender. Palps with four segments, attached dorsally. P-2 with only one seta, located distally.
P-3 with flap-like process. P-4 basally with three thin setae. Legs slender. Genua distinctly
shorter than tibiae. Ventral setae short, often bipectinate. Tarsi I to III with each four and
tarsus IV with three dorsal setae, including the solenidia. Ventral setae are missing on all
tarsi. Solenidion of tarsus I and II inserted dorsolaterally. Claws small. Empodium with a
tiny tooth.
24. Porolohmannella violacea (K RAMER , 1879) (5-26 a-f)

1879 Leptognathus violacea K RAMER , Arch. Naturgesch. 45: 147.
1933 Porolohmannella violacea K.V IETS , Zool. Anz. 102: 283.
1989 Porolohmannella violacea B ARTSCH , Acarologia 30: 235.
Fig. 5-26: Porolohmannella violacea; a, dorsal; b, idiosoma, ventral, female; c, gnathosoma, lateral; d,
gnathosoma, ventral; e, tarsus I, lateral; f, leg I, medial.
152 Special part
De sc ri pti on : Length of idiosoma 300-550 lm. Idiosoma very flat; colouring often
slightly pink, with eye pigment spot beneath AD and OC. Anterior margin truncated
or with small frontal spine. Dorsal plates delicately reticulate. AD fused with the AE, which
extends dorsad. OC with gland pore, cornea and two setae. PD large, with two pairs of
gland pores in the lateral margin, two pairs of setae and the adanal setae. Ventral plates AE,
PE and GA separated. GA of females large, with three pairs of pgs. Genital sclerites each
with two genital acetabula. Anal plate located ventrally. Rostrum long and slender, palps
slender, not curved. Legs slender. The four pairs of legs almost equal-shaped. Tibiae I to II
ventrally each bear four bristles, tibia IV three bristles, some of them bipectinate, others
slender and smooth. Claws small.
H a bit at a nd Bi ol og y: Primarily in lentic surface waters; in mud and between vascular
plants, mosses and algae; also reported from brackish coastal waters. Males have not been
described so far.
Di stri bu ti on: Europe; Greenland, North America and Japan (L ETTEVALL 1962;
B ARTSCH 1982; A BÉ 1990c).
Subfamily Rhombognathinae K.V IETS , 1927

1927 Rhombognathinae K.V IETS , Z. wiss. Zool. 130: 89.
Genus Isobactrus N EWELL , 1947

1947 Isobactrus N EWELL , Bull. Bingham oceanogr. Coll. 10, 21: 68.
Typus generis: Aletes setosus L OHMANN , 1889, by original designation.
Di ag no sis: Idiosoma intensive green, due to colouring of the gut caeca. OC mostly with
two gland pores. Adanal setae missing. Epimeral plates small, separated by broad areas of
striated cuticle. EI and EII not fused in the median in European species. Genital plate of
females mostly small, that of males slightly larger. Females with three pairs of pgs, males
with 40-90 pgs. Gnathosoma always short, in dorsal view mostly hidden under the idio-
soma. Rostrum short; the two pairs of maxillary setae on the rostrum. Palps short, barely
reaching beyond the rostrum. Palps with four, rarely with three segments. P-2 with a long
seta. Legs shorter than the idiosoma. Legs I and II similar in shape. Genua always shorter
than telofemora and tibiae. Tibiae I and II ventrally each with one pair of bristles, tibiae III
and IV with one or two bristles; often at least one of the bristles bipectinate. Tarsi I to III
dorsally mostly with four setae, including the solenidia, tarsus IV mostly with three setae.
Solenidia always located dorsolaterally. Tip of the tarsi with pair of pas. Tarsi with paired
claws. A carpite is always present between tip and empodium of the tarsus. Empodium
without tine.
R em a rk: Four juvenile stages exist, larva, proto-, deuto- and tritonymph.
25. Isobactrus uniscutatus (K.V IETS , 1939) (5-27 a-g)

1939 Rhombognathus uniscutatus K.V IETS , Arch. Naturgesch. (NF) 8: 545.
1953 Isobactrus uniscutatus N EWELL , Syst. Zool. 2: 128.
1972 Isobactrus uniscutatus B ARTSCH, Abh. Verh. naturwiss. Ver. Hamburg (NF) 16: 191.
De sc ri pti on : Length of idiosoma 353-439 lm. Idiosoma dark green, legs colourless. Eye
pigment present. Idiosoma anteriorly and posteriorly ovate. AD and PD fused to form a
dorsal shield; OC separated, with a cornea. Epimeral plates I to IV fused at the body mar-
gin, but separated by large areas of striated cuticle on the ventral side. Genital plate of
females largely reduced, the three pairs of pgs located in the striated cuticle. Genital plate
of males with approximately 40 pgs. Genital acetabula of females inside the GO, in males
Fig. 5-27: Isobactrus uniscutatus; a, dorsal; b, idiosoma, ventral, female; c, idiosoma, ventral, male; d,
gnathosoma, ventral; e, gnathosoma, lateral; f, Tarsus I, lateral; g, leg I, medial.
two pairs located outside, behind the GO. One of the dorsal setae of tarsi I-IV long, dis-
tinctly longer than the length of the tarsi. Claws smooth.
Pr ea du lt sta ge s: Plates AD and PD separated in juveniles. All developmental stages are
assignable to I. uniscutatus because of the smooth claws and the long seta on the tarsi.
Habit at a nd Bi ol ogy: Isobactrus uniscutatus demonstrates mass occurance in brackish
water regions of the North Atlantic and the North Sea, still abundant upstream in oligoha-
line and adjacent limnic regions. In the river Elbe this species occurs near Hamburg (Blan-
kenese). On the coast I. uniscutatus inhabits the Blidingia (green algae) zone, in oligohaline
and limnic areas it lives among the moss Cinclidotus.
Di stri bu ti on: Only found near the coast, in regions of the North Atlantic and the North
Sea from France to Great Britain and Germany, in the Mediterranean from the Adria (Croa-
tia). Recently also reported from Australia and there classified as neozoan (B ARTSCH &
G WYTHER 2004).
154 Special part
Subfamily Ropohalacarinae B ARTSCH, 1989

1989 Ropohalacarinae B ARTSCH, Acarologia 30: 227
Genus Ropohalacarus B ARTSCH , 1989

1989 Ropohalacarus B ARTSCH , Acarologia 30: 227.
Typus generis: Porohalacarus uniscutatus B ARTSCH , 1982, by original designation.
Di ag no sis: AD and OC present, OC often reduced. Gland pores reduced. Four to five
pairs of dorsal idiosomal setae. AE with three pairs of setae, PE with 2-3 setae. Genital
acetabula on the genital sclerites. GA of females with three pairs of pgs. Palps with
Fig. 5-28: Ropohalacarus uniscutatus; a, overall view, dorsal; b, idiosoma, ventral, female; c, gnatho-
soma, ventral; d, palp, medial; e, leg I, medial; f, tarsus I, lateral.
four segments, attached dorsolaterally. P-2 with two setae, P-3 with a spine, P-4 with a very
broad seta. First and second pair of legs of similar shape. Genua of all legs shorter than
telofemora and tibiae. Solenidion on both tarsus I and tarsus II located dorsolaterally. Pair-
ed claws large, empodium small.
26. Ropohalacarus uniscutatus (B ARTSCH, 1982) (5-28 a-f)

1982 Porohalacarus uniscutatus B ARTSCH , Gewäss. Abwäss. 68/69: 43.
1989 Ropohalacarus uniscutatus B ARTSCH, Acarologia 30: 227.
De sc ri pti on : Length of idiosoma 254-300 lm. Idiosoma pale. AD and PD fused to a
shield; OC largely reduced. Dorsum with four to five pairs of idiosomal setae. Adanal setae
not seen. AE, PE and GA fused to a shield. PE with two to three setae. The three pairs of pgs
of females arranged closely around the GO. Genital sclerites with three pairs of external
genital acetabula. Ovipositor short. Gnathosoma short. Rostrum shorter than half the
length of the gnathosomal base, not reaching the end of the P-2. P-4 with broad bristle.
Tibiae I and II each with three, tibiae III and IV each with two slightly bipectinate setae.
Tarsi I and II dorsally each with three setae and one club-shaped solenidion, ventrally each
with one seta. On tarsus III dorsally four, ventrally no setae, on tarsus IV dorsally three,
ventrally no. Tips of tarsi I, II and IV each with paired pas; the tip of tarsus III only with the
lateral pas. Empodium with a small tine. Claws with a few small tines on their pectines.
S im i la r s pe ci e s: At low magnification similar to Porohalacarus alpinus, but in contrast
to the latter, there are no conspicuous eye spots.
H a bit at a nd B io lo gy : Probably exclusively in groundwater.
Di stri bu ti on: Northern Germany. Also reported from North America (B ARTSCH 1982;
S TRAYER 1988).
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158 General part
6. Acari: Terrestrial Parasitengona inhabiting transient

biotopes
Andreas Wohltmann, Grzegorz Gabryś and Joanna Ma˛kol
General part
Introduction
The Parasitengona (Acari: Prostigmata) constitute one of the most diverse taxa among the
Acari with respect to species richness as well as with respect to inhabited biotopes and
realised life styles. Many of the Parasitengona species are conspicuous due to their relatively
large size and their often bright red colouration. Hitherto, about 9,000 species of Parasi-
tengona have been described, with the water mites (Hydrachnidia, chapter 7) making up
the main portion. The terrestrial Parasitengona (Trombidia, about 4,000 species) are com-
posed of the Erythraeoidea, the monogeneric Calyptostomatoidea and the Trombidioidea
including chiggers (Trombiculidae s.l.) (6-1).
Terrestrial Parasitengona colonised a wide range of biotope types from xeric to hygric
habitats and from lowland areas to mountains. A high percentage of ubiquistic opportu-
nists inhabiting particular alluvial floodplains is evident in terrestrial Parasitengona
(W OHLTMANN 2005), similar to other arthropods of the terrestrial mesofauna in tempora-
rily aquatic biotopes (W EIGMANN & W OHLGEMUTH -VON R EICHE 1999).
Because of their similar morphology and life style, the larvae of terrestrial Parasitengona
may at first glance easily be misidentified as water mite larvae. To help to reduce this prob-
lem, this chapter will thus focus on parasitengone species usually associated with amphi-
bious biotopes in Central Europe. We restrict keys to the identification of higher taxa and
only provide detailed keys at species level for those taxa limited to amphibious biotopes or
with high abundance in such biotopes.
Biology and life history

Like in water mites, the life cycle of Trombidia usually includes the typical sequence of
immobile, parasitic and predatory instars (6-2). The few exceptions from this pattern
in Trombidia concern an additional moulting of postlarval instars or a deviation of life
styles of particular instars (e.g. reduction of parasitism in the larva). Possibly with the ex-
ception of some Balaustium spp. (Erythraeoidea), parasitengone mites are obligatory di-
oecious species, without apparent sex chromosomes (N ORTON et al. 1993). Sperm is trans-
ferred indirectly via stalked spermatophores; the particular design of spermatophores as
well as the presence or absence of signalling structures and mating behaviour differ between
genera (W ITTE 1984, 1991a). Eggs are usually deposited in one to three clutches; egg size as
well as egg number per female varies between species and genera (W OHLTMANN 1999a).
Eggs of Trombidia are predominantly of a reddish colouration and do not possess a gela-
tinous sheath as in water mites. However, in Erythraeoidea a lipid-protein cover which
probably improves desiccation resistance provides the eggs with a brown - black colour.
Phototactic and geotactic responses guide larval search for hosts; there is no evidence for a
specific distance recognition of hosts. Usually the host instar to be parasitized is recognised
by the larva (as in most primitive Hydrachnidia). However, some hygric Trombidioidea
display preparasitic attendance to host pupae as also known for hygrobatoid water mites.
Host range varies between species; however, the most common host resource of water mites
(Chironomidae) is only rarely parasitized by terrestrial Parasitengona. Overlapping host
resources between water mites and Trombidia usually concern tipulid Diptera, which

https://doi.org/10.1007/978-3-662-55958-1_6
6. Acari: Terrestrial Parasitengona inhabiting transient biotopes 159
Fig. 6-1: Phylogenetic system of Parasitengona. Traditional grouping documented above, the Trom-
bidia as well as the Trombidioidea will become paraphyletic. For supporting characters see references
(numbers at squares): 1) W ELBOURN 1984, 2) W ELBOURN 1991, 3) W ITTE 1991a, b, 4) Z HANG 1994,
5) W ITTE 1995, 6) Z HANG 1995, 7) S ÖLLER et al. 2001, 8) W OHLTMANN 2000. Black squares indicate
well-supported taxa, grey squares indicate ambiguous groupings, white squares means no apomor-
phies known. (Derived from W OHLTMANN 2000).
are often parasitized by hydryphantoid water mites and also by terrestrial Parasitengona
(e.g. Calyptostoma, Johnstonianidae, and some species of Microtrombidiidae). In some
cases water mites and terrestrial Parasitengona occur together on the same host individual,
but use different body parts of the host (W OHLTMANN 2000, 2001). The duration of the
parasitic phase differs between species and ranges from three to 14 days; usually the host
survives. The parasitic larva allows - via the host - fast colonisation even of patchily dis-
tributed habitats and is considered the main dispersal agent in Parasitengona. The ances-
trally active instars proto- and tritonymph are immobile in Parasitengona. Before entering a
calyptostatic stage specimens retreat into the soil or crevices in stones or organic substrates,
and are thus difficult to detect in the field. The original cause for the evolution of calypto-
static nymphs may have been to ease synchronisation of the simultaneous abundance of
instars. Later on, the calyptostatic protonymph in particular allowed considerable orga-
nismal reorganisation and facilitated the evolution of heteromorphic parasitic larvae
and predatory postlarval instars (W OHLTMANN et al. 2001). The active postlarval instars
- deutonymph and adult - prey on insect eggs and larvae, and for a few species predation on
insect imagines and mites has additionally been observed (e.g. Allothrombium, Erythraeus,
Johnstoniana). The deutonymph constitutes the primary growth stage in many Trombidia.
However, in a number of species the larva is able to produce additional cuticle while feeding
on the host (without intermittent moult: neosomy) and contributes most to growth during
ontogeny. Species of Trombidium, Eutrombidium and, in the water mites, Eylais and Hy-
drachna are examples for such change in larval growth capacity. The annual abundance of
the larvae and their specific hosts as well as of males and females during the mating period
160 General part
are synchronised in a species by means of obligatory diapause of a particular instar. The

diapausing instar may be an immobile (egg, protonymph) or an active (deutonymph, adult)
instar. The diapausing instar is fixed within a species, but varies considerably even within
genera. In the species so far investigated, induction of diapause is not dependent on envir-
onmental cues but obligatorily coupled to a particular ontogenetic phase. The duration of
diapause is controlled by an internal clock and moreover dependent on low environmental
temperature, the termination of diapause is triggered after a chilling period by a raise in
environmental temperature and, sometimes, day-length. This results in semi- to univoltine
(Calyptostoma, most Trombidioidea) or strictly univoltine (most Erythraeoidea, some
Trombidioidea) life cycles with seasonally limited abundance of instars.
Comprehensive reviews about host associations, life cycles and life history evolution of
terrestrial Parasitengona have been published by W ELBOURN (1983), Z HANG (1998)
and W OHLTMANN (2000).
Habitat specificity and adaptive characters

It is still difficult to determine which species of Trombidia are actually restricted to am-
phibious biotopes. This problem is certainly not limited to Trombidia, but is present in
other groups as well. Often hypotheses on habitat specificity are derived solely from sta-
tistical correlation of species to sampled locations, and often sample size or biased sampling
of different biotope types severely limits statistical analysis. Moreover, species with a wide
Palaearctic distribution may have different habitat preferences depending on latitude and
availability of particular biotope types. In the Trombidia, only few taxa can be considered
as restricted to amphibious biotopes based on the distribution in the field as well as on
experimental investigations into physiological properties and host relationships. For other
species the distribution in the field strongly suggests such habitat specificity, however, no
data are available for a causal reasoning. Finally, a number of opportunistic species colo-
nises temporarily flooded localities but also inhabits permanent terrestrial biotopes. Such
species are obviously not restricted to transient habitats. In general, the percentage of spe-
Fig. 6-2: Life cycle of

Parasitengona. Gener-
alised life cycle with the
example of a species
diapausing as adult
(e.g. Eutrombidium
trigonum). Annual
abundance of instars
indicated by corre-
sponding segments
around the disk show-
ing months of the year;
shaded area indicates
diapause. Calyptostatic
instars (proto- and tri-
tonymph) remain in the
cuticle of the preceding
instar. (Derived from
W OHLTMANN 2000).
cies with habitat binding towards amphibious biotopes is highest in biotopes at the marine
coast and in temporarily flooded woodland ponds, whereas on alluvial floodplains ubiquis-
tic species are dominant (W OHLTMANN 2005). With regard to the latter, evolutionary adap-
tations not related to hygric biotopes allowed colonisation of temporary aquatic biotopes
without further adaptive changes in these groups. Such adaptations are e.g. optimised pro-
tection against water flux in order to increase desiccation resistance in Erythraeoidea, pro-
longed and dense body setation to avoid water contact in edaphic Trombidioidea. Irrespec-
tive of the evolutionary origin of character patterns enabling colonisation of amphibious
biotopes, Trombidia and water mites may co-occur in temporary waters. Sometimes even
direct interference between water mites and terrestrial Parasitengona has been observed,
e.g. feeding of adult Johnstoniana errans (Trombidia: Johnstonianidae) on Acerbitas
barbigera K. V IETS, 1911 (Hydrachnidia: Hydryphantidae – for the synonymy of the genus
Acerbitas see Ö ZDIKMEN 2006) adults just after drying up of the temporary pond. The
main difference between water mites and Trombidia concerns the timing of activity of post-
larval instars. Although deutonymphs and adults of some water mite genera (e.g. Acerbitas,
Hydryphantes) are able to move under terrestrial conditions, nutrition, mating and ovipo-
sition take place under aquatic conditions only. In Trombidia, by contrast, feeding and
reproduction is restricted to terrestrial periods although in some species the active instars
are able to survive aquatic periods for considerable time.
Phylogeny and taxonomy

The monophyly of Parasitengona is convincingly supported by a number of apomorphic
characters, such as the switch from predatory to parasitic life style of the larva, or the trans-
formation of ancestrally active proto- and tritonymphs into calyptostatic (immobile) in-
stars. The sister group of Parasitengona within the Prostigmata are the Anystidae or pos-
sibly a subgroup of the Anystidae (W ITTE 1991b, L INDQUIST 1996, N ORTON et al. 1993).
Although the phylogenetic relationships within the Parasitengona have drawn considerable
attention and some proposals based on a reliable phylogenetic methodology are published
(W ITTE 1991b, 1995, W ELBOURN 1984, 1991, Z HANG 1994, 1995, S ÖLLER et al. 2001,
W OHLTMANN in press), the phylogenetic system (6-1) is not completely resolved. While the
monophyly of Hydrachnidia, Calyptostomatoidea and Erythraeoidea, as well as the rela-
tionships within the Erythraeoidea, are well supported by a number of often complex char-
acters (W ITTE 1995), the monophyly of the Trombidioidea is not convincingly supported
and some subgroups of the Trombidioidea may very well be closer related to water mites
than to any of the terrestrial taxa of Parasitengona (W OHLTMANN 2000, in press).
The actual state of taxonomy is rather deficient in terrestrial Parasitengona, mostly because
of the heteromorphic larval and postlarval instars of which only few have been correlated
through rearing experiments. Moreover, many descriptions published in the last century do
not allow unambiguous identification, and redescriptions of species and re-evaluation of
genera and families are urgently needed. Thus, the key provided below is artificial and still
provisional, and restricted to properly described species often found in amphibious bio-
topes in Central Europe. Taxonomic groups not reported from this area are excluded;
only the active instars are taken into account. As for other Acari, identification of terrestrial
Parasitengona is preceded by clearing and mounting of specimens. In general, the termi-
nology (6-3, 6-4) follows G RANDJEAN (1947); specific terms are explained in figures. Fig-
ures of larvae always show freshly emerged specimens but one should keep in mind that
parasitengone larvae may undergo considerable enlargement during their parasitic phase.
Only sclerites keep their size (legs, scutum etc.) whereas the prefolded ’smooth’ cuticle ex-
tends and thereby often changes positions of body setae.
162 General part
Fig. 6-3: Morphology and taxonomic terminology of larval Parasitengona (Microtrombidium pusil-
lum shown, after G ABRYŚ & W OHLTMANN 2001):
a, idiosoma, upper half: body in dorsal view, lower half: body in ventral view; prodorsal sclerite (=scu-
tum) bearing setae S (sensilla of trichobothrium; S1 and S2 in taxa having two pairs of trichobothria)
AM (anterior median seta, often a single unpaired seta in Trombiculidae, obviously homologous to S1
and thus missing in taxa having two pairs of trichobothria), AL (anterior lateral seta), PL (posterior
lateral seta). Usually the scutum is restricted to the dorsum, but in several microtrombidiid genera its
lateral edges extend to the ventral side of idiosoma (stolascutum). Eyes usually placed on an ocular
plate. Further dorsal setae usually arranged in rows (cn - hn); setae c1 often on fused c1 plates (= scu-
tellum). Cx I-III: The coxae of legs I-III form non-movable plates (epimeral plates or epimera), which
bear setae 1a, b on coxa I (additionally a small supracoxal seta at the distal margin in dorsal position of
coxa I may be present); seta 2b on coxa II and seta 3a on coxa III (3b usually located outside coxa III).
Co: Claparède’s organ, located between coxae I and II (absent in Erythraeoidea). Anal pore: may be
surrounded by paired or unpaired sclerites. Psn: pseudanal setae;
b, segmentation and setation pattern of leg I - legs of larvae consist of coxa (not shown, cx in A);
trochanter (tr); femur (fe), which may be completely or partially divided into basifemur and telofemur;
genu (ge), tibia (ti) and tarsus (ta) including pretarsus. Beside non-specialized setae (usually with setules
or barbs) specialised setae occur mostly at the dorsal side of leg segments. Solenidia appear usually
transversely striated under the microscope, they occur on tarsus (x), tibia (u), genu (r) and sometimes
on the femur (h) of legs I-III. On genu and tibia small setae (j) may occur, which are tiny in Micro-
trombidiidae and Trombidiidae, but somewhat larger in Johnstonianidae. Another small seta of a dif-
ferent type (like solenidia not birefringent) occurs only on the tarsus I and occasionally II, and is named
famulus (e). Eupathidia (f) constitute a type of comparably large specialised setae, which differ from
solenidia in that they do not possess the inner spiral structure; they are restricted to tarsi I-III;
c, tarsus of leg I - only the specialised setae are shown, with the typical position of eupathidia. The
pretarsus is not counted as an independent leg segment, it consists of inner (=anterior) and outer (=pos-
terior) claws and empodium. Arrangement of claws and empodium may differ between legs I-III.
Claws may be empodium-like in appearance or modified in another way, the empodium may look
claw-like;
d, gnathosoma (ventral view) - the palp consists of a small trochanter (the palp coxae contribute to the
formation of the subcapitulum and are not discernible as individual segments), a femur, genu, tibia and
tarsus. Tibia and tarsus form a thumb-claw complex, the odontus (a prominent spine-like seta often
called ’tibial claw’ mostly bifid and in dorsal position at the distal end of the tibia) usually extend over
the palp tarsus. Setae on palp femur and palp genu, if present, are sometimes tiny and difficult to detect.
Specialised setae (solenidia x, eupathidia f) are restricted to the palp tarsus. Chelicerae always with
two segments; the tip of the claw-like digitus fixus is usually visible at the anterior mouth opening
whereas the basal segment of the chelicera remains internal. At its anterior end the gnathosoma is
dorsally not completely closed and sometimes equipped with horseshoe-like sheathes and sclerites (ste-
phanostome). Setae on the gnathosoma are termed hypostomal, subcapitular or tritorostral setae (bs)
and oral setae (os, often minute). Additionally, a pair of adoral setae may occur.
164 General part
Fig. 6-4: Morphology and taxonomic terminology of postlarval Parasitengona (Centrotrombidium

schneideri [adult] shown, after W OHLTMANN et al. 2004):
a, idiosoma, upper half: body in dorsal view; lower half: body in ventral view. Always with four pairs
of walking legs (leg I-IV), tarsi and in particular tarsus I usually enlarged. Legs with 7 segments, femur
always completely divided into basi- and telofemur. Coxae of legs I-IV form epimera and are not mo-
vable against the body. Width and length of legs, leg segments and sometimes even the body are often
used in taxonomic keys;
b, prodorsal sclerite - scutum with a median thickening called crista metopica. The latter usually pro-
minent and well sclerotized, the rest of scutum often inconspicuous. One or two pairs of trichobothria,
sensillae usually setiform. An anterior projection (naso) may or may not be present. One or two pairs of
eyes, usually on common sclerite, eyes flat or situated on more or less distinct movable stalks. A median
eye may be present at the anterior part of crista metopica;
c, dorsal body setae - posterior dorsal setae constitute a major key character in postlarval Parasiten-
gona. Only minor differences can be observed in the shape of setae in deutonymphs and adults. They
consist of a basal sclerite and the seta, which may be spine-like, setiform with setules or modified in
another way;
d, chelicera - consisting of a blade-like digitus fixus and the basal segment; the digitus mobilis of other
Acari is always absent. In Calyptostomatoidea and Erythraeoidea the chelicera is styliform, a small
digitus fixus is present in Calyptostomatoidea whereas in Erythraeoidea it is absent and the chelicera
consists of the basal segment only. Stigmata located dorsally at level of distal half of the basal segments
of chelicerae, usually badly visible in undissected specimens;
e, palp - coxa forms part of the gnathosoma and is not discernible. Femur always undivided. Palp tibia
usually with a prominent distal spine (odontus), which forms a thumb-claw complex with the palp
tarsus. Specialized setae (x, n) are restricted to the palp tarsus. In particular the expression and ar-
rangement of setae on the palp tibia are important characters in taxonomic keys;
f, genital opening - visible but closed in deutonymphs. Surrounded by two pairs of sclerites (epi- and
centrovalves). Usually three pairs of genital acetabula in adults (and tritonymphs), two pairs in deu-
tonymphs, (one pair in the protonymph). However, there are several exceptions from this rule; e.g.:
multiple acetabula occur in many Hydrachnidia, acetabula are reduced or modified in Erythraeoidea,
only two pairs of acetabula are present in adults of Calyptostoma, but three in deutonymphs of Allo-
thrombium;
g, anal pore - surrounded by paired sclerites or one unpaired sclerite.
166 Special glossary
Special glossary
The general body organization of terrestrial Parasitengona and special characters used in the key are
shown for larvae (6-3) and adults (6-4) separately.
basidont prominent spine situated ventrolaterally on palp tibia
conala serrate spine-like seta at palp tibia, sometimes palp genu of some Erythraeidae
crista metopica strongly sclerotized area of the scutum, usually elongate
ctenidium row of spine-like setae located dorsally at palp tibia
empodium median cuticular extension at the pretarsus between the lateral claws, may be claw-
like
eupathidium see „specialized setae“
famulus see „specialized setae“
glandularium defense organ typical for Hydrachnidia; consisting of a round plate bearing a sen-
sory seta and a glandular opening (see chapter 7)
hypertrichy dense setation of idiosoma and/or legs, often velvet-like
hypostomal seta seta located on the ventral side of gnathosoma
lassenia organ paired plates with a glandular opening present anterior to coxae III in Tanaupodi-
dae
lophotrix modified setae located ventrally at distal tarsus III of some microtrombidiid larvae
microseta see „specialized setae“
naso triangular median extension of the scutum
ocular plate sclerite carrying the lateral eyes
odontus prominent spine at the distal end of palp tibia, resembling a claw, may be divided in
two to three tips in its distal part (= tibial claw)
paradont spine located dorso-medially of odontus on palp tibia
pseudoradula a group of delicate setae on medial side of palp tibia (e.g. Valgothrombium spp.)
radula a group of bare, spine-like setae on medial side of palp tibia, ventrally to ctenidia
scobala normal (not specialized) seta
scopa modified seta located dorsally at distal tarsus III of some microtrombidiid larvae
scutellum unpaired dorsal sclerite just behind the scutum as a result of fused c1 plates, usually
carrying one pair of normal setae
scutum unpaired dorsal sclerite near anterior margin of idiosoma. Includes a sensillary area
with trichobothria and (usually in postlarval instars) strongly sclerotized parts (cris-
ta metopica) - see „dorsal shield“
sensilla receptive structure of the trichobothrium, usually long and hair-like
sensillary area area on the scutum where the trichobothria are located
smilum modified claw at tarsus III of some microtrombidiid larvae
solenidion see „specialized setae“
specialized setae setae at body or legs, which differ in structure from normal setae and probably serve
reception of particular (chemical, sensorial) signals - most obvious are sensillae
(sound receptors), eupathidia and solenidia (probably chemical receptors), micro-
setae and famuli (small spine-like setae of unknown function)
stephanostome horseshoe-like cuticle at distal end of gnathosoma, present in Microtrombidiidae
stolascutum a particular type of scutum, extended antero-laterally and forming flaps, which are
visible from ventral view, present in microtrombidiid larvae
subcuticular mesh a net of proteinous fibrils below the epidermis, serving as stabilizing element, not
directly connected to the cuticle; present in postlarval Calyptostoma and some
Trombidioidea
supracoxal seta small seta located dorsally on coxa of leg I and/or of palp
urnula tubular invagination surrounded by ring-like sclerites located on the anterior
half of dorsal idiosoma, present in some Balaustiinae, function unknown
vertex narrowed, well sclerotized plate at anterior most part of aspidosoma, usually
bearing several setae
Abbreviation Explanation
f eupathidium = specialized smooth seta on legs, in particular on tarsus
e famulus = specialized smooth seta on tarsus of legs I-II, usually small
z accompanion seta = smaller seta close to dorsal eupathidium on tarsus leg I
x omega / tarsala = solenidion localized on tarsus
u phi / tibiala = solenidion localized on tibia
r sigma / genuala = solenidion localized on genu
h theta / femorala = solenidion localized on (telo-) femur
j microseta / vestigiala = small seta on tibia and genu of walking legs
I first walking leg
II second walking leg
III third walking leg
IV fourth walking leg
AL second pair of non-sensillary setae on scutum in larvae
AM first pair of non-sensillary setae on scutum in larvae
bFe basifemur
bs hypostomala, subcapitular seta, tritorostral seta; located ventrally on gnathosoma
cs adoral setae, usually small spine-like setae located anterior and dorsal at the gnatho-
soma, (= or)
Cx coxa = epimeral plate
DS dorsal setae
Fe femur
Ge genu
GOP genital opening
IP index pedibus; the total length of all legs (including coxae)
N non-specialized setae on palp and legs
or oral seta, located anteriorly at gnathosoma
PaTa palp tarsus
pDS posterodorsal setae on idiosoma
pDS I, pDS II posterodorsal setae of first and second type, respectively
PL third pair of non-sensillary setae on scutum in larvae
Psn pseudanal setae, usually 2 pairs of setae lateraly to anal opening, never located on anal
plates
S sensilla on crista metopica in active postlarval forms and on scutum in larvae
Ta tarsus
tFe telofemur
Ti tibia
Tr trochanter
168 Special part
Special part
Area treated
The systematic part includes all taxa reported from Central Europe. In this chapter this area
is defined as marked out by the territory of the following eight countries: Austria, Czech
Republic, Germany, Hungary, Poland, Slovakia, Switzerland, and The Netherlands.
Key to the major subgroups of Parasitengona

1 Three pairs of locomotory legs (larva) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Four pairs of locomotory legs (deutonymph or adult) . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Acetabulum situated ventrally between coxae I and II, anal opening present (6-5 a, c), legs of
about the same length as the idiosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Acetabulum and anal opening absent, legs usually long (4 twice the length of idiosoma) (6-5 b)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythraeoidea (page 169)
3 Palp genu with two simple setae (6-5 d). Anal pore usually surrounded by an undivided sclerite or
unsclerotized; if paired sclerites present (Stygothrombiidae) then acetabulum stalked and extend-
ed well beyond the lateral body sides; if sclerite absent (Wandesia) then coxa II without setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachnidia (chapter 7, page 241)
– Palp genu without setae or with one seta. Anus usually without surrounding sclerites (6-5 a), if
sclerites present then as pair of valves. Acetabulum never stalked. Coxa II with one or more
setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Coxal fields I-III with few setae (510, usually one to three) (6-5 a). Distinct, well developed
scutum present, ocular plate without setae. . . . . . . . . . . . . . . . . Trombidioidea (page 178)
– Coxal fields I-III hypertrichous (4 15 setae) (6-5 c). Scutum reduced to sensillary area bearing a
pair of trichobothria, ocular plate with two setae . . . . . . . . Calyptostomatoidea (page 168)
5 Idiosoma with glandularia (sclerites bearing a seta and the opening of a gland) (6-6 a), only few
body setae, prolonged leg-setae („swimming setae“) may be present, aquatic . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachnidia (chapter 7, page 241)
– Idiosoma without glandularia, hypertrichous, no extremely prolonged leg setae, terrestrial . 6
6 Chelicerae styliform (6-6 b), retractable, movable digit absent or inconspicuous . . . . . . . . 7
– Chelicerae usually blade-like, not retractable, with distinct movable digit (6-6 c) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidioidea (page 178)
7 Two pairs of prodorsal trichobothria (6-6 d) on distinct scutum (usually with crista metopica).
Chelicera not segmented, movable digit of chelicera absent. Ocular plate without setae, genital
acetabula not visible, legs usually long (4 twice the length of idiosoma) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythraeoidea (page 169)
– One pair of prodorsal trichobothria (6-6 e), crista metopica and scutum reduced. Chelicera with
small movable digit. Ocular plate with two setae, two pairs of distinct genital acetabula, legs
short, general habitus as in larva (6-5 c) . . . . . . . . . . . . . . Calyptostomatoidea (page 168)
Superfamily Calyptostomatoidea
Monogeneric, with only one species described from Central Europe.
Family Calyptostomatidae O UDEMANS , 1923

1923 Calyptostomatidae, O UDEMANS, Tijds Entom. 66: 77
Genus Calyptostoma C AMBRIDGE , 1875

1875 Calyptostoma C AMBRIDGE, Ann. Mag. Nat. hist., London 16 (96): 384.
Typus generis: Calyptostoma hardii C AMBRIDGE , 1875, by monotypy (= Acarus velutinus
O. F. M ÜLLER , 1776).
Di ag no sis: Adult and Deutonymph: Comparatively large, idiosoma length of adults

1000-3000 lm, width up to 2000 lm. Colouration bright orange to brownish. Scutum
reduced to sensillary area carrying only one pair of sensillae. Laterally, two pairs of red
eyes (6-6 e). Gnathosoma completely retracted except for feeding or prior to moulting.
Palp four-segmented, chelicera elongate, styliform, consisting of small digitus and chelic-
eral base. Idiosoma covered with strong setae. Cuticular folds form scale-like patterns on
the idiosoma. Two pairs of acetabula present in both postlarval instars. Larva: In general
appearance (6-5 c) similar to postlarval active instars, but gnathosoma not retractable.
Chelicera not styliform, digitus blade-like. Scutum small, almost reduced to sensillary
area; one pair of sensillae. Acetabulum between coxae I and II.
R em a r k: V ISTORIN -T HEIS (1976) united all European species as Calyptostoma velutinus;
however, doubts about the validity of this view have recently arisen. Although no morpho-
logical differences could be found in the larvae, the differing attachment sites of parasitizing
Calyptostoma captured in May and September, respectively, and the different development
times of the subsequent protonymphs indicate that two species may be present. Moreover,
some differences in idiosomal setae of postlarval Calyptostoma collected in Norway were
found (M A˛ KOL , unpublished), thus at least two species possibly also occur in Central
Europe.
1. Calyptostoma velutinus (O. F. M ÜLLER, 1776) (6-5 c; 6-6 e)

1776 Acarus velutinus O. F. M ÜLLER, Zoologiae Danicae Prodromus, Hafniae (Copenhagen),
p. 187, n. 2226.
1875 Calyptostoma hardii C AMBRIDGE , Ann. Mag. Nat. hist., London 16 (96): 384.
For further synonymy see V ISTORIN -T HEIS (1976).
Di ag no sis: As for the genus.
S im i la r sp e ci e s: see above.
Habit at an d Bi ol ogy: Calyptostoma velutinus larvae parasitize species of Tipulinae and
Limoniinae. Parasitic larvae usually appear May-June (attached to the dorsal surface of the
first thorax segment of the host), at some locations a second larval peak occurred in Sep-
tember (with the larvae attached to the first abdominal segments) (W OHLTMANN et al.
1999). After locating a suitable host pupa, the larvae stay nearby and await ecdysis of
the host imago (preparasitic attendance); parasitism on tipulid larvae was observed
only once (W OHLTMANN et al. 1999) and seems to be exceptional. During hatching of
the host imago, Calyptostoma larvae attach and start parasitism. Adults, deutonymphs
and host searching larvae occur in the litter layer throughout the year. Postlarval active
instars feed on dipteran larvae, whereby the protractible gnathosoma facilitates piercing
and ingestion of prey hidden in small crevices. All active instars including the larvae may
survive for some months without food. Calyptostoma velutinus displays a semivoltine life
cycle. Abundant in amphibious (limnic) biotopes.
Di stri bu ti on: Central Europe.
Superfamily Erythraeoidea
European Erythraeoidea are usually fast-moving long-legged hunters on substrate surfaces;
edaphic lifestyles as observed in several Trombidioidea are unknown. Most species are of a
reddish body colour, but iridescent colouration (some Erythraeinae) or with white spots on
dark background colouration (Leptus trimaculatus (R OSSI, 1794)) also occur. Dry and xe-
ric biotopes are most typical of Erythraeoidea, but some species occur also at hygric lo-
calities and very few seem confined to amphibious biotope types. Charletonia cardinalis
(C. L. K OCH, 1837), Erythraeus rupestris (L INNAEUS , 1758) and some Abrolophus and
170 Special part
Leptus species were found on alluvial floodplains, other Leptus (e.g. L. beroni FAIN , 1992)
inhabit summer dry temporary pools in forested areas, some Abrolophus spp. (A. rubipes
(B ERLESE & T ROUESSART, 1889), A. passerini (B ERLESE , 1904)) are obviously restricted to
amphibious localities at the rocky shores of marine coasts. However, no physiological cau-
Fig. 6-5: Larvae of Parasitengona, ventral view: a, Trombidium brevimanum, details: anal pore, palp
genu, Claparède’s organ (after W OHLTMANN 1999b); b, Leptus trimaculatus (after W ENDT et al.
1992); c, Calyptostoma velutinus, details: coxae, Claparède’s organ, anal pore; d, Euthyas truncata,
detail: anal pore; (scale bars = 50 lm, enlarged details not to scale).
ses for habitat restriction towards amphibious habitats have been found, with the exception
of those Abrolophus spp. which inhabit the littoral fringe of marine coasts and which dis-
play adaptive transformation of the seasonal abundance of instars and increased hemo-
lymph osmolality (W ENDT & W ITTE 1985, B ÜCKING et al. 1998). Even Charletonia car-
dinalis and Erythraeus rupestris (both possibly confined to hygric biotopes) display desic-
cation resistance like their close relatives inhabiting very dry places (W OHLTMANN 1998,
2005). Thus it seems that many of the erythraeoid species found at amphibious locations
Fig. 6-6: Parasitengona adults, dorsal view: a, Euthyas truncata, detail: glandularium; b, d, Abrolo-
phus quisquiliarius, b, detail: chelicera; d, details: scutum, eyes; c, Trombidium brevimanum, detail:
chelicera (after W OHLTMANN 1999b); e, Calyptostoma velutinus, details: scutum, eyes; (scale bars =
50 lm).
172 Special part
are opportunistic invaders rather than especially adapted to the particular conditions of this
type of transient biotope. With the exception of Balaustium spp. and some Abrolophus spp.
which have predatory larvae, erythraeoid larvae parasitize arthropods. All European Er-
ythraeidae hitherto investigated display strictly univoltine life cycles with the abundance of
larvae as well as of other ontogenetic instars being restricted to particular periods of the
year. The annual abundance of larvae differs between species depending on the hibernating
instar. Larvae firmly attach to the host by means of a sticky secretion, a rapid mode of host
encounter enabling erythraeoid larvae to use even fast moving hosts. Preparasitic atten-
dance is unknown. Species inhabiting amphibious localities survive winter flooding in
the diapausing instar. As far as immobile instars (egg, prelarva, proto- and tritonymph)
are concerned they seem not to be affected by flooding and develop normally when sub-
merged in water. Species diapausing as deutonymph or adult instars retreat into crevices
(e.g. hollow shafts of dead grass and reed), which remain air-filled during flooding. When
individuals were kept submerged in water, air covers were absent or only a very small air
cover around the body was observed, which diminished within a few days. Survival of deu-
tonymphs and adults kept in water does not exceed one month in most species (20 C)
(W OHLTMANN 2005).
Key to families, subfamilies and genera

(M ar k e d w ith a s te r is k*: groups containing taxa regularly found in amphibious biotopes).
Ex clude d: Andrevella parkeri (A NDRÉ, 1930); Eatoniana georgei O UDEMANS, 1941; Forania men-
tonensis (A NDRÉ, 1929); Hauptmannia O UDEMANS, 1941; Rudaemannia H AITLINGER , 2000. The
incomplete descriptions available for Andrevella parkeri, Eatoniana georgei and Forania mentonensis
do not allow unambiguous identification.Within the subfamily Abrolophinae, the state and validity of
the genera Abrolophus (based on postlarval stages), Hauptmannia (based on larvae) and Rudaemannia
(based on larvae) cannot be resolved until a thorough revision including correlation of adults and lar-
vae through laboratory rearing is available. Provisionally, species described within Hauptmannia and
Rudaemannia are treated as members of Abrolophus based on laboratory rearing of Abrolophus quis-
quiliarus (H ERMANN , 1804), Rudaemannia rudaensis (H AITLINGER , 1986) and Hauptmannia long-
icollis (O UDEMANS , 1910) (W OHLTMANN 2000, 2001). A deficient state of taxonomical knowledge is
pronounced within European Erythraeidae, and revisions are urgently needed for nearly every genus
and family. Very few species are known from larval and postlarval instars; moreover many of the de-
scriptions do not allow unambiguous identification of species. For this reason, the following key allows
identification only of genera but not of species.
Larvae
1 Legs without trichobothria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythraeidae 3
– At least one trichobothrium-like structure present on genu, tibia or tarsus of leg I (6-7 a) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smarididae 2
2 Two pairs of eyes, odontus without prominent lateral extension . . . . . . . . . . . . Smaridinae
– One pair of eyes, odontus at its basis with a prominent spine-like extension laterally. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirstiosomatinae
3 Two eyes on each side of prodorsum (6-7 b) . . . . . . . . . . . . . . . . . . . . . . . Erythraeinae 8
– Only one eye on each side of prodorsum (6-7 c). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Genu II, III each with 11-12 non-specialized setae (N), coxae II, III each with two setae . . 5
– Genu II, III with less than 11 N (usually nine) (6-7 g), coxae II, III usually with one seta . . 6
5 Pretarsal claws I, II each with distinct terminal claw-like hook . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callidosoma W OMERSLEY, 1934
– At least one pretarsal claw on each leg I, II, without terminal claw-like hook (6-7 d). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charletonia* (page 177)
6 Scutum roughly triangular, cheliceral base prolonged anteriorly (6-7 e), flask-like . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptus*(page 178)
– Scutum roughly rectangular or circular (6-7 f-g), cheliceral base short, normal . . . . . . . . . 7
7 Palp trochanter without setae, genu II, III each with one solenidion (r), odontus without pro-
minent tooth, scutum without crista metopica (6-7 g) . . . . . . . . . . Abrolophus* (page 177)
– Palp trochanter with one to two setae, genu II, III without r, odontus with prominent tooth, crista
metopica present (6-7 f). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balaustium H EYDEN, 1826
8 Empodium claw-like, one claw may be setulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Empodium with setules, claws never setulose . . . . . . . . . . . . . . . Phanolophus A NDRÉ 1927
In Central Europe only Phanolophus oedipodarum (F RAUENFELD , 1868)
9 Palp femur and coxae II and III with one seta each . . . . . . . . . . . . Erythraeus* (page 178)
– Palp femur with two setae, coxae II and III with more than two setae . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Curteria S OUTHCOTT, 1961
Deutonymphs and Adults

1 Only chelicerae retractable, dorsal setae usually setiform. . . . . . . . . . . . . . . Erythraeidae 3
– Complete gnathosoma retractable, dorsal setae inflated (6-8 a) . . . . . . . . . . . Smarididae 2
2 Two pairs of eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smaridinae
– One pair of eyes (6-8 a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirstiosomatinae
3 One eye on each side of prodorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Two eyes on each side of prodorsum (6-8 b-c) . . . . . . . . . . . . . . . . . . . . . . Erythraeinae 4
4 Conalae on palp absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Conalae on palp tibia, sometimes also on palp genu (6-8 d) . . . . . . Erythraeus* (page 178)
5 Dorsal setae leaf-like, setulose (6-8 e), scutum with distinct naso (6-8 f) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phanolophus A NDRÉ 1927
In Central Europe only Phanolophus oedipodarum (F RAUENFELD , 1868)
– Dorsal setae setiform or plumose, scutum without naso . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Dorsal setae 5 40 lm, plumose (6-8 g). Anterior sensillae more or less equal to posterior sen-
sillae, length always less than 85 lm. Non-sensillary setae on scutum short, maximum 60 lm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kamertonia G ABRYŚ, 2000
In Central Europe only Kamertonia polonica G ABRYŚ, 2000
– Dorsal setae 4 50 lm, not plumose (6-8 h). Anterior sensillae (4 120 lm) always shorter than
posterior sensillae (4150 lm). Non-sensillary setae on scutum 4 70 lm long . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Curteria S OUTHCOTT, 1961
7 Eyes behind middle of scutum, idiosoma setae nude, blade-like, expanded or with few setules,
usually orange to reddish in life, palp setae nude or setulose . . . . . . . . . . . . . . . . . . . . . . 8
– Eyes anterior to middle of scutum (6-8 i), idiosoma setae densely covered with setules, usually
dark brown in life (white spots may occur in some species), palp setae heavily setulose (6-8 j)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptus* (page 178)
8 Dorsum of idiosoma without ring-like sclerites around tubular invaginations (urnulae) posterior
to eye lenses, crista metopica present, odontus without prominent tooth. . . . . . . . . . . . . . 9
– Dorsum of idiosoma with urnulae, crista metopica absent (6-9 a), odontus with prominent tooth
(6-9 b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balaustium H EYDEN, 1826
9 Palp tarsus enlarged (6-9 c), extending beyond the odontus, setae on palp femur of approximately
the same length, palp trochanter with one to two setae at least twice as long as other setae . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Palp tarsus small, palp femur with at least one seta twice as long as others (6-9 d), all setae on palp
trochanter of approximately the same length, habitus as in 6-9 f . . Abrolophus* (page 177)
10 Palp tarsus spherical (6-9 c), scutum between trichobothria wider than crista metopica (6-9 e),
idiosoma setae setiform with setules . . . . . . . . . . . . . . . . . . . . . . Charletonia* (page 177)
– Palp tarsus nearly cylindrical, scutum as wide as crista metopica, idiosoma setae blade-like . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callidosoma W OMERSLEY, 1934
174 Special part
Fig. 6-7: Erythraeoidea larvae, dorsal view: a, Hirstiosoma ampulligera, detail: tibia I with trichobo-
thria; b, Erythraeus (Zaracarus) rupestris (leggs omitted); c-d, Charletonia cardinalis; c, details: an-
terior part of idiosoma, gnathosoma; d, detail: tarsus II (lateral); e, Leptus trimaculatus, details: an-
terior part of idiosoma, gnathosoma (after W ENDT et al. 1992); f, Balaustium sp., details: anterior part
of idiosoma, gnathosoma; g, Abrolophus quisquiliarius, details: palp tarsus, genu III; (scale bars =
50 lm).
Fig. 6-8: Erythraeoidea adults, dorsal view: a, Hirstiosoma ampulligera, details: scutum, eyes, dorsal
body seta; b-c, Erythraeus (Zaracarus) rupestris; c, details: scutum, eyes; d, Erythraeus phalangoides,
detail: palp (lateral); e-f, Phanolophus oedipodarum; e, detail: body setae; f, detail: scutum; g, Kamer-
tonia polonica, detail: body setae; h, Curteria episcopalis, detail: body setae; i-j, Leptus trimaculatus,
deutonymph; i, detail: scutum, eyes; j, detail: palp tarsus and tibia; (scale bars = 50 lm).
176 Special part
Fig. 6-9: Erythraeoidea adults, dorsal view: a-b, Balaustium murorum ; a, details: scutum, eyes, ur-
nulae; b, detail: palp (lateral); c, e, Charletonia cardinalis; c, detail: palp (lateral); e, details: scutum,
eyes; d, f, Abrolophus quisquilarius; d, detail: palp (lateral); (scale bars = 50 lm).
Family Erythraeidae R OBINEAU- D ESVOIDY, 1828

1828 Erythraeidae R OBINEAU- D ESVOIDY, Rech. sur L’Org. vertébrale Crust. Arachn. Ins.: 144
Genus Abrolophus B ERLESE , 1891 (6-6 b, d; 6-7 g; 6-9 d, f)

1891 Abrolophus B ERLESE, Acari, Myriopoda et Scorpiones hucusque in Italia reperta 59, 1: 2.
Typus generis: Trombidium quisquiliarum H ERMANN, 1804, by original designation.
In clud ed: Hauptmannia O UDEMANS, 1910 with type species Achorolophus longicollis
O UDEMANS , 1910; Rudaemannia H AITLINGER, 2000 with type species Hauptmannia ru-
daensis H AITLINGER, 1996.
Di ag no sis: Adult and Deutonymph: Small to medium-sized. Colouration reddish to oran-
ge. Idiosoma setae setiform. Crista metopica elongate with two pairs of trichobothria, a
pair of eyes on ocular sclerites situated laterally and behind the middle of crista metopica
(6-9 f). Palp tarsus normal, not or only slightly extending over odontus (6-9 d). Legs com-
parably short, less than twice the length of idiosoma. Larva: Comparably small. Legs less
than twice the length of idiosoma (6-7 g). Posterior part of idiosoma covered with rows of
barbed setae, each inserted on a plate-like sclerite. Scutum roughly pentagonal, bearing two
pairs of trichobothria plus two pairs of non-specialized setae. Lateral to scutum a pair of
ocular sclerites, each with one eye. Gnathosoma distinctly separated from idiosoma. Coxa
of legs I-III with one seta each. Tarsus I-III each with two lateral claws and claw-like em-
podium.
H a b it at an d B i o lo gy : Abrolophus spp. inhabit meadows including alluvial plains, but
are also present at the rocky shores of marine coasts. The life cycle is univoltine. Adults and
deutonymphs are active on the surface of the litter layer and vegetation. Larvae are either
predatory or parasitize on Thysanoptera (W ELBOURN 1983, W OHLTMANN 2000, 2001).
Di stri bu ti on: Worldwide.
Genus Charletonia O UDEMANS, 1910 (6-7 c-d; 6-9 c, e)

1910b Charletonia O UDEMANS, Ent. Ber. (Amst.) 3 (53): 73.
1910 Sphaerolophus B ERLESE, Redia 6: 349.
Typus generis: Erythraeus singularis O UDEMANS , 1910, by original designation.
Di ag no sis: Adult and Deutonymph: Large to medium-sized. Colouration reddish to
brownish. Idiosoma setae setiform. Crista metopica elongate with two pairs of trichobo-
thria, a pair of eyes on ocular sclerites situated laterally and behind the middle of crista
metopica (6-9 e). Palp tarsus enlarged, globular (6-9 c). Legs long, more than twice the
length of idiosoma. Larva: Comparably large. Legs long, more than twice the length of
idiosoma. Posterior part of idiosoma covered with rows of barbed setae, each inserted
on a plate-like sclerite. Scutum roughly pentagonal, bearing two pairs of trichobothria
plus three pairs of non-specialized setae (6-7 c). Lateral to scutum a pair of ocular sclerites,
each with one eye. Gnathosoma distinctly separated from idiosoma. Coxa of legs I with
one, II-III with two setae each, at least one pretarsal claw II-III setulose and without hook
(6-7 d).
Habit at a nd Bio lo gy: Charletonia spp. inhabit a wide range of biotope types including
alluvial plains and other amphibious biotopes (W OHLTMANN 2005). The life cycle is
strictly univoltine. Adults and deutonymphs are fast moving hunters on the surface of
the litter layer and vegetation, and prey on insect eggs and larvae. Larvae parasitize on
Delphacidae and other arthropods (W ELBOURN 1983, W OHLTMANN 2000).
178 Special part
Genus Erythraeus L ATREILLE, 1806 (6-7 b; 6-8 b-d)

1806 Erythraeus L ATREILLE, Genera crustaceorum et insectorum 1: 146.
1910a Bochartia O UDEMANS, Ent. Ber. (Amst.) 3 (52): 49.
1995 Erythraeus (Zaracarus) S OUTHCOTT, Acarologia 36 (3): 223.
Typus generis: Acarus phalangoides D E G EER , 1778, by original designation.
Di ag no sis: Adult and Deutonymph: Large. Colouration reddish to brownish. Idiosoma
setae setiform. Crista metopica elongate with two pairs of trichobothria, two pairs of eyes
laterally on ocular sclerites (6-8 c). On palp tibia and sometimes palp genu series of serrate,
stout and conical spines (conalae) (6-8 d). Legs long, more than twice the length of idio-
soma (6-8 b). Larva: Comparably large. Legs long, more than twice the length of idiosoma.
Posterior part of idiosoma covered with rows of barbed setae, each inserted on a plate-like
sclerite. Scutum bearing two pairs of trichobothria plus two pairs of non-specialized setae
(6-7 b). Lateral to scutum a pair of ocular sclerites, each with two eyes. Gnathosoma dis-
tinctly separate from idiosoma. Coxae of legs I-III with one seta each. Palp tarsus with
prominent odontus.
H a bit at an d B io lo gy : Erythraeus spp. inhabit a wide range of biotope types including
alluvial plains and other amphibious biotopes (W OHLTMANN 2005). The life cycle is
strictly univoltine. Adults and deutonymphs are fast moving hunters on the surface of
the litter layer, and prey on less mobile insects and on ants. Larvae usually parasitic on
aphids (W OHLTMANN 2000).
Genus Leptus L ATREILLE, 1796 (6-5 b; 6-7 e; 6-8 i-j)

1796 Leptus L ATREILLE, Précis de caractères génériques des Insectes (Brive) 5: 177.
1877 Ritteria K RAMER, Arch. Naturgesch. 43 (1): 228.
1899 Achorolophus B ERLESE, Acari, Myriopoda et Scorpiones hucusque in Italia reperta 59, 1: 2.
Typus generis: Acarus phalangii D E G EER , 1778, by original designation.
Di ag no sis: Adult and Deutonymph: Medium-sized to large. Colouration brownish, also
black with white spots may occur. Idiosoma setae strong, heavily setulose. Crista metopica
elongate with two pairs of trichobothria, a pair of eyes on ocular sclerites situated laterally
to the anterior half of crista metopica (6-8 i). Legs robust, always less than twice the length
of idiosoma. Larva: Large to medium-sized. Legs long, more than twice the length of idio-
soma (6-5 b). Posterior part of idiosoma covered with rows of barbed setae, each inserted
on a plate-like sclerite. Scutum triangular, bearing two pairs of trichobothria plus two pairs
of non-specialized setae; laterally with a pair of ocular sclerites, each with one eye (6-7 e).
Gnathosoma distinctly separated from idiosoma. Cheliceral base elongate, flask-like. Co-
xae of legs I-III with one seta each; one setulose pretarsal claw I-III.
Habit at a nd Bio lo gy: Leptus spp. occur in a wide range of biotopes including amphi-
bious ones (W OHLTMANN 2005). The life cycle is univoltine. Adults and deutonymphs in-
habit the litter layer and prey on insect eggs and larvae. Larvae parasitize on a wide range of
insects and arachnids (W ELBOURN 1983, W OHLTMANN 2000).
Superfamily Trombidioidea
Trombidioidea constitute the largest subgroup of terrestrial Parasitengona. Despite their
bright red body colouration and abundance in a variety of biotope types (ranging from
xeric to hygric) these mites are often overlooked due to their cryptic life style. Many species
inhabit the litter layer or display an almost complete edaphic life style; members of the
Microtrombidiidae and Trombidiidae are capable of active digging even in hard soil sub-
strates. An obvious habitat binding to extremely hygric biotopes is known for representa-
tives of Johnstonianidae (N EWELL 1957, W ENDT 1995, W OHLTMANN et al. 1999, 2005),
and frequently more than one species of Johnstonianidae, and even of the genus Johnstoni-
ana, can be found in a single locality (W OHLTMANN 2001). Habitat binding to hygric bio-
topes is not only evident from the failure to find Johnstonianidae in non-hygric biotopes
(R OBAUX 1970), but also from experiments on physiological properties, which allow to
describe the present causes for the apparent habitat restriction. All hitherto investigated
Johnstonianidae display strong hydrophilic behaviour in the deutonymph and adult instar
(W ENDT 1995). Moreover, desiccation resistance of mobile (W ENDT 1995) and immobile
(W OHLTMANN 1998) instars is extremely low which prevents species from invading typical
terrestrial localities with fluctuating humidity. In adaptation to semiaquatic environments,
the hemolymph osmolality in Johnstonianidae is considerably lower than in other terres-
trial Parasitengona (W ENDT 1996), sometimes even lower than measured in plesiotypic
water mites (O LOMSKI 1991). Johnstonianid larvae display a comparably high host-spe-
cificity. Species using flying hosts parasitize hydrophilic Tipulidae; they search host pupae
in the litter layer and await ecdysis of host imagines before starting parasitism. Attachment
to the host is rather loose and often larvae detach when captured, e.g., with a sweep net.
Johnstoniana parva parasitizes other Trombidioidea (other Johnstonianidae in particular)
and displays no preparasitic attendance, J. rapax finally lost parasitism and the larva
switched to a predatory life style. The life styles of J. rapax and J. parva larvae may be
regarded as alternative tactics which reduce the risk of being dispersed into unsuitable ha-
bitats via flying hosts, however, there is a trade-off with reduced dispersal abilities. Because
of their low desiccation resistance and rather low temperature optimum for development
( 15 C) Johnstonianidae are not present on alluvial floodplains with short vegetation and
temporarily high temperature, but occur frequently at temporary woodland ponds. The
diapausing eggs easily withstand several months of exposure to aquatic conditions without
any observable reduction of viability. During winter rehydration of the biotope, adults and
deutonymphs of J. errans were found aggregated in crevices of submerged rotting wood
(W OHLTMANN 1996).
Of the Central European Microtrombidiidae, Valgothrombium spp., Echinothrombium
rhodinum (C. L. K OCH , 1837), and Atractothrombium sylvaticum (C. L. K OCH , 1835)
are obviously restricted to amphibious biotopes. This is mainly derived from reported sam-
ple sites (summarised in G ABRYŚ 1996, G ABRYŚ & W OHLTMANN 2001, G ABRYŚ et al.
2005). Moreover the finding of specimens in the soil of flooded areas, as well as the quick
appearance of species on terrestrial ’islands’ which emerge at lowering water levels strongly
indicate that these species do not recolonise amphibious biotopes each year but remain in
situ during flooding (W OHLTMANN 2005). When deutonymphs and adults of Microtrom-
bidiidae and Trombidiidae are submerged in water, the dense setation prevents direct con-
tact of water with the cuticle; an air film around the body persists for about three weeks
(5 C, 100 % oxygen saturation). However, even A. sylvaticum deutonymphs reared from
protonymphs at submerged conditions (and thus without air cover) survived aquatic con-
ditions (20 C, 100 % oxygen saturation) without any food uptake for more than two
months (W OHLTMANN 2005). Like in Johnstonianidae the hemolymph osmolality is de-
creased in V. valgum (G EORGE , 1909c) (W OHLTMANN & W ENDT 1996), but values typical
for terrestrial arthropods have been measured in A. sylvaticum (W ENDT, cited after
W OHLTMANN 2000). Moreover, desiccation resistance of immobile instars in E. rhodinum
and A. sylvaticum is like that of Microtrombidium pusillum (H ERMANN , 1804) and other
terrestrial Parasitengona (W OHLTMANN 1998, 2000). Thus the actual causes for the hy-
pothesised habitat binding of these species remain unknown and probably differ from those
found in Johnstonianidae. Valgothrombium valgum, Enemothrombium bifoliosum (C A-
180 Special part
NESTRINI , 1884), and A. sylvaticum co-occur with Johnstonianidae (and serve as hosts for
J. parva larvae), however, A. sylvaticum inhabits also alluvial floodplains (together with
Echinothrombium rhodinum) and has even been found in salt marshes. Other species of
Microtrombidiidae (e.g. Microtrombidium pusillum, Dactylothrombium pulcherrimum
(H ALLER , 1882), Camerotrombidium spp., Campylothrombium clavatum (G EORGE ,
1909c)) frequently found at amphibious locations probably inhabit primarily adjacent ter-
restrial areas.
Of the Trombidiidae, Podothrombium filipes and Paratrombium insulare have been col-
lected on temporarily flooded meadows and in reed belts respectively (M A˛ KOL 2000a,
2000b; W OHLTMANN 2005). Trombidium and Allothrombium spp. usually inhabit perma-
nent terrestrial areas with fluctuating humidity. However, in the Lower Oder Valley T. ho-
losericeum, T. brevimanum and A. fuliginosum are also frequent on alluvial plains and
evidently survive winter flooding in the soil (W OHLTMANN 2005). When submerged in
water the dense setation preserves an air film around the body. Because this air film is
not balanced to the weight of specimens but will drive them to the water surface it should
not be termed a ’plastron’. Moreover, since specimens survive for long time after dimin-
ishing of the air cover, it has probably no crucial function for oxygen uptake at aquatic
conditions. In laboratory tests, eggs and protonymphs of T. brevimanum developed nor-
mally when exposed to aquatic conditions, deutonymphs reared from protonymphs kept
submerged in tap water survived for up to 136 days (20 C, 100 % oxygen saturation)
(W OHLTMANN , 2005).
Of the remaining Trombidioidea, species of the Tanaupodidae have been reported from
subaquatic biotopes in North America (N EWELL 1957) and Europe (R OBAUX 1967). The
data set is rather limited and any hypotheses about habitat specificity and its causes seem
speculative at the moment. However, because prerequisites such as a synchronised
life style with winter diapause, an edaphic life style and a dense body setation enable other
species to temporarily tolerate aquatic conditions without further adaptive changes, one
should expect the discovery of some more species inhabiting amphibious biotopes.
Key to families
Ma r ked w ith as ter i s k*: groups containing taxa regularly found in amphibious biotopes.
Larvae
1 With two dorsal sclerites (scutum and scutellum) (6-10 a), tibia III without solenidion u . . 2
– With one dorsal sclerite (scutum present; scutellum absent), tibia III with u (6-10 b) . . . . . 4
2 Femur I with six non-specialized setae (N), genu II/III usually each with two N (6-10 a) . . 3
– Femur I with five N, genu II/III each with three to four N . . . . . Trombidiidae* (page 220)
3 Femur II with four N, genu II without microseta j . . . . . . . . . . . . . . . . . . Neothrombiidae
– Femur II with five N, microseta j present on genu II (6-10a) Microtrombidiidae* (page 195)
4 Femur I without solenidia (h), anal sclerite absent or present . . . . . . . . . . . . . . . . . . . . . . 5
– Femur I with at least one solenidion h, anal sclerite usually absent. . . . . . . . . . . . . . . . . . 6
5 Anal sclerite present, coxa III with two to three setae (6-10 c), scutum with two pairs of tricho-
bothria and two pairs of non-specialized setae (AL, PL); parasitic on insects . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanaupodidae* (page 217)
– Anal sclerite absent, coxa III with one seta, scutum with one pair of trichobothria, paired AL, PL
setae and usually a single unpaired anterior seta (AM); parasitic on vertebrates. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombiculidae
6 Leg claws absent (6-10 d); pretarsus with claw-like empodium. Femora II/III each with two or
more solenidia h . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Legs with paired claws, empodium absent. Femora II/III each with one or no solenidia h (6-10 b),
at least one solenidion on tarsus II enlarged and club-shaped. . Johnstonianidae* (page 181)
7 Femur I with seven non-specialized setae (N), coxae I-III with reticulate pattern (6-10 d) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotrombidiidae
– Femur I with six N, coxae I-III without reticulate pattern . . . . . . . . . . . . . . . Trombellidae

1 Pregenital tubercle anteriorly of genital opening present (6-11 a), dorsal setae setiform or spine-
like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanaupodidae* (page 217)
– Pregenital tubercle absent, dorsal setae variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Dorsal idiosoma setae distinctly trifurcate (6-11 b), no spine-like setae on palp tibia arranged in
rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotrombidiidae
– Dorsal idiosoma setae variable, but usually not trifurcate. If trifurcate, then spine-like setae on
palp tibia arranged in rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsum with circular structures (6-11 c), scutum narrow, poorly sclerotized or absent . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombellidae
– Dorsum without circular structures, scutum variable. . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 All dorsal idiosoma setae spine-like, nude or with few barbs. Crista metopica with one or two
pairs of trichobothria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Idiosoma setae usually with numerous setules (additionally some spine-like setae may occur).
Crista metopica with one pair of trichobothria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 Crista metopica usually bearing two pairs of trichobothria; if only one pair present then sensilla
of trichobothrium not flagelliform. Palp tibia with a large ventro-lateral spine (6-11 d). At least
one club-shaped solenidion (x) on tarsus II . . . . . . . . . . . . . . Johnstonianidae* (page 181)
– Only one pair of trichobothria present on crista metopica, sensillae flagelliform. Palp tibia with
one spine in dorsal position (deutonymphs; adults of Podothrombium hispanicum), additional
spines may occur in ventral position (deutonymphs) or several spines of different thickness in
dorsal (adults excluding P. hispanicum) and ventral position (adults) (6-11 e). Solenidia on tarsus
II not club-shaped . . . . . . . . . . . . . . . Trombidiidae* in part (Podothrombium) (page 220)
6 Palp tibia with two to three distinct lateral spines in tandem (6-11 f), idiosoma attenuated be-
tween leg II/III (6-11 g), eyes often absent . . . . . . . . . . . . . . . . . . . . . . . . . Trombiculidae
– Palp tibia different, idiosoma not attenuated between legs II/III, eyes present or absent . . . 7
7 Anterior end of scutum with one to two setae, eyes absent or incorporated into the scutum . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neothrombiidae
– Anterior end of scutum usually with more than two setae, usually two pairs of eyes which are not
incorporated into the scutum (6-11 h) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Trichobothria in posterior position on crista metopica (6-11 h). Palp tibia with at least one pro-
minent spine-like seta (paradont) adjacent to palp tibial claw (odontus), other spine-like setae on
tibial claw usually arranged in rows (ctenidia) (6-11 i), eyes usually not on distinct stalks, general
habitus as in 6-11 k . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microtrombidiidae* (page 195)
– Trichobothria in posterior or mid position on the crista metopica. Setae on palp tibia neither
spine-like nor arranged in rows (6-11 j), eyes on prominent stalks . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidiidae* in part, (page 220)
Family Johnstonianidae T HOR , 1935

1935 Johnstonianidae T HOR , Zool. Anz. 109: 387.
Di ag no sis: Adult and deutonymph: Small- to large-sized. Colouration reddish to brown.
Idiosoma cuticle almost smooth without large sclerotized areas. Posterior dorsal idiosoma
setae simple (smooth or with few small barbs), inserted on prominent sclerites (6-4 c; 6-15
g-i). Prodorsal shield (scutum) with projecting naso. At level of coxa IV genital opening,
surrounded by paired sclerites (two pairs in adults: epi- and centrovalves; one pair in deu-
tonymphs) of similar shape (6-4 f). Anal pore posterior to genital opening, surrounded by a
pair of elongate sclerites (6-4 g). Palp tibia with a prominent spine (basidont) ventrally to
the odontus (6-4 e; 6-11 d; 6-13 c, e). In addition to other solenidia at least one club-shaped
182 Special part
solenidion present on tarsus II. Larva: Medium- to large-sized. Without scutellum; scutum
roughly triangular with projecting naso (6-10 b; 6-12 b-f). Posterior idiosoma setae simple
(smooth or with few small barbs), often inserted on prominent platelets (6-10 b; 6-12 e, f).
Anal pore present, without surrounding sclerites. Coxae I-II-III with two-one-one setae;
seta 1a on medially extended part of coxa I (pars medialis). Femur of legs I-III completely
or partially divided into basi- and telofemur, at least one solenidion present on telofemur I
(6-10 b). Famuli present on tarsi I and II. Tarsus II with at least one club-shaped solenidion
(6-15 f). Tarsi I-III with two claws but without empodium (6-10 b). Supracoxal setae on
palp and leg I absent.
Fig. 6-10: Trombidioidea larvae: a, Microtrombidium pusillum (dorsal), details: pretarsus, femur II,
tibia III (after G ABRYŚ & W OHLTMANN 2001); b, Johnstoniana eximia (dorsal), details: femur III, tibia
III (after W OHLTMANN et al. 1994); c, Lassenia lasseni (ventral), details: lassenia organ, anal sclerite
(after N EWELL 1957); d, Neotrombidium tricuspidum (ventral), detail: coxa I (after S INGER 1971);
(scale bars = 50 lm).
Fig. 6-11: Trombidioidea adults: a, Tanaupodus passimpilosus (ventral), details: genital opening,
pregenital tubercle; b, Neotrombidium furcigerum (dorsal), detail: body seta (after T HOR & W ILL-
MANN 1947); c, Trombella lusitanica (left dorsal, right ventral view) (after R OBAUX 1967); d, Diplo-
thrombium carpaticum, detail: palp tibia and tarsus (lateral) (after W OHLTMANN et al. 2004); e, Po-
dothrombium filipes, detail: palp tibia and tarsus (lateral); f, Morelacarus sp., detail deutonymph: palp
tibia and tarsus (lateral); g, Neotrombicula autumnalis deutonymph (dorsal), h, Atractothrombium
sylvaticum (dorsal), details: crista metopica, eyes (after G ABRYŚ et al. 2005); i, k, Enemothrombium
bifoliosum (dorsal); i, detail palp, lateral (after W OHLTMANN & G ABRYŚ in press); j, Trombidium
holosericeum (lateral), detail: palp tibia and tarsus (after M A̧KOL & W OHLTMANN 2000); (scale
bars = 50 lm).
184 Special part
H a bit at a nd B i ol og y: Larvae originally parasitize nematoceran insects, some species

switched to other hosts (mites, water beetle pupae) or even abandoned parasitism. All
members of the family are confined to amphibious (limnic) biotopes.
Key to genera
(all species of Johnstonianidae are obviously confined to amphibious biotopes)
Larvae
1 Microseta j present on genu I/II (6-12 a), scutum with two pairs of trichobothria plus two pairs
of non-specialized setae (6-12 b) . . . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana (page 191)
– Microseta j absent on genu I/II, scutum with one pair of trichobothria . . . . . . . . . . . . . . 2
2 Scutum with one pair of flagelliform trichobothria plus three pairs of non-specialized setae
(6-12 c), (telo-) femur I with two solenidia h, habitus as in 6-12 f . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplothrombium (page 188)
– Scutum with one pair of globose trichobothria plus one pair of non-specialized setae (6-12 d),
(telo-) femur I with one h (6-12 e) . . . . . . . . . . . . . . . . . . . Centrotrombidium (page 184)

1 Sensillae of prodorsal trichobothria flagelliform (6-13 a, d) . . . . . . . . . . . . . . . . . . . . . . . 2
– Sensillae of prodorsal trichobothria short and globose (6-13 b), habitus as in 6-2 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrotrombidium (page 184)
2 Scutum expanded, roughly triangular (6-13 a), basidont bidentate (6-13 c), no cuticular exten-
sions near base of setae on palp tibia, habitus as in 6-13 f . . . . . . Johnstoniana (page 191)
– Scutum linear or nearly linear (6-13 d), basidont unidentate; in adult sharkfin-like cuticular ex-
tensions distally to setal bases, at outer side of palp tibia (6-13 e) habitus as in 6-13 g. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplothrombium (page 188)
Genus Centrotrombidium K RAMER , 1896

1896 Centrotrombidium K RAMER, Zool. Anz. XIX 515: 445.
1957 Evansiella V ERCAMMEN -G RANDJEAN, Ann. Mag. Nat. Hist. 10: 283.
Typus generis: Centrotrombidium schneideri K RAMER , 1896, by original designation.
Di ag no sis: Adult and deutonymph: Relatively small in size, body length of adults about
500 lm, width about 350 lm. Idiosoma ovoid (6-4 a), densely covered with spine-like setae
located on rounded, slightly asymmetrical tubercles (6-4 c). Scutum with crista metopica
elongate with anteriorly projecting naso, widened at bases of trichobothria (6-4 b). One
pair of trichobothria, sensillae distally globular, moreover one pair of non-specialized setae
anteriorly to trichobothria. One to two pairs of eyes laterally to scutum, each pair situated
on small protrusions of a common ocular sclerite. Palp tibia with unidentate basidont (6-4
e). Three (adult) or two pairs (deutonymph) of genital acetabula. Larva: Posterior part of
idiosoma covered with rows of barbed setae, each inserted on a plate-like sclerite (6-12 e).
Scutum roughly triangular with projecting naso, bearing one pair of trichobothria with
distally globular sensillae and only one pair of non-specialized setae (6-12 d). Lateral
to scutum a pair of ocular sclerites, each with one to two eyes. Legs I-III with partially
divided femora. Tarsus I without small accompanying seta z. Palp tarsus with one promi-
nent spatulate seta distally.
Key to species
Ex clude d: Centrotrombidium motasi (F EIDER, 1945) (nomen dubium).
Larvae
1 Seta 2 b on coxa II bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Seta 2 b on coxa II simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Seta 3 b on coxa III bifurcate, one seta on palp femur Centrotrombidium schneideri (page 185)
– Seta 3 b simple, three setae on palp femur . . . . . Centrotrombidium blackwellae (page 185)
3 One seta on palp femur . . . . . . . . . . . . . . . . . . . Centrotrombidium romaniense (page 185)
– Three setae on palp femur . . . . . . . . . . . . . . . . . . Centrotrombidium culicoides (page 185)

Only one species described in Central Europe: Centrotrombidium schneideri (page 185)
1. Centrotrombidium blackwellae B AKER, 1999

1999 Centrotrombidium blackwellae B AKER, Syst. Parasitol. 44: 42.
D e s c r i p t i o n : B AKER (1999); postlarval instars unknown. Larva: Three setae on palp fe-
mur, seta 1 b on coxa I simple, seta 2 b on coxa II bifurcate, seta 3 b on coxa III simple.
H a bit at an d B io lo gy : Larvae parasitic on adults of Ceratopogonidae. Confined to am-
phibious (limnic) biotopes.
Di stri bu ti on: Scotland.
2. Centrotrombidium culicoides (V ERCAMMEN -G RANDJEAN , 1957)

1957 Evansiella culicoides V ERCAMMEN -G RANDJEAN, Ann. Mag. Nat. Hist. 10: 283.
1973 Centrotrombidium culicoides V ERCAMMEN -G RANDJEAN & F EIDER, Riv. Parassitol. 34 (2):
125.
D e s c r i p t i o n : V ERCAMMEN -G RANDJEAN (1957); postlarval instars unknown. Larva:
Three setae on palp femur, seta 1 b on coxa I simple, seta 2 b on coxa II simple, seta 3
b on coxa III simple.
H a bit at an d B io lo gy : Larvae parasitic on adults of Ceratopogonidae. Confined to am-
phibious (limnic) biotopes.
3. Centrotrombidium romaniense V ERCAMMEN -G RANDJEAN & F EIDER, 1973

1973 Centrotrombidium romaniense V ERCAMMEN -G RANDJEAN & F EIDER, Riv. Parassitol. 34 (2):
122.
D e s c r i p t i o n : V ERCAMMEN -G RANDJEAN & F EIDER (1973); postlarval instars unknown.
Larva: One seta on palp femur, seta 1 b on coxa I simple, seta 2 b on coxa II simple, seta 3 b
on coxa III simple.
Di stri bu ti on: Romania.
4. Centrotrombidium schneideri K RAMER, 1896 (6-4; 6-12 d-e; 6-13 b)

1896 Centrotrombidium schneideri K RAMER, Zool. Anz. 19 (515): 445.
1922 Diplothrombium walteri S CHWEIZER, Verhdl. Naturf. Ges. Basel 33: 85.
D e s c r i p t i o n : K RAMER (1896) (adult); W OHLTMANN & W ENDT (1996) (larva), W OHLT-
MANN et al. (2004) (active instars). Adult and Deutonymph: No other species is described
for postlarval instars in the genus. Larva: One seta on palp femur, seta 1 b on coxa I with
three setules, seta 2 b on coxa II bifurcate, seta 3 b on coxa III bifurcate.
186 Special part
Fig. 6-12: Johnstonianidae larvae dorsal view: a, Johnstoniana rapax, detail: tarsus-genu leg II; b,
Johnstoniana eximia, detail: scutum; c, f, Diplothrombium longipalpe; c, detail: scutum; d-e, Centro-
trombidium schneideri; d, detail: scutum; (a-f after W OHLTMANN et al. 2004; scale bars = 50 lm).
Fig. 6-13: Johnstonianidae adults dorsal view: a, c, Johnstoniana errans; a, details: scutum, eyes; c,
detail palp (lateral); b, Centrotrombidium schneideri, details: scutum, eyes; d-e, Diplothrombium
longipalpe; d, details: scutum, eyes; e, detail: palp (lateral); f, Johnstoniana eximia (left: dorsal, right:
ventral); g, Diplothrombium wittei (left: dorsal, right: ventral); (a-g after W OHLTMANN et al. 2004;
scale bars = 50 lm).
188 Special part
H a bit at and B io log y : Obviously uni- to semivoltine. Eggs develop into larvae without
intermittent diapause. Larva abundant in late June. In laboratory experiments larvae para-
sitize adult Ceratopogonidae after preparasitic attendance. Confined to amphibious
(limnic) biotopes.
Di stri bu ti on: Belgium, Czech Republic, Germany, Norway, Switzerland.
Genus Diplothrombium B ERLESE , 1910

1910 Diplothrombium B ERLESE, Redia 6: 357.
Typus generis: Trombidium longipalpe B ERLESE , 1887, by original designation.
Di ag no sis: Adult and Deutonymph: Medium in size, body elongate (6-13 g), length about
1000-1500 lm, width about 600-1000 lm. Idiosoma covered with spine-like setae, each
seta located on a sclerite (6-14 a-c). Scutum with anteriorly projecting naso and elongate
crista metopica, widened at bases of trichobothria(6-13 d). Two pairs of trichobothria with
setiform sensillae. Additionally two to six non-specialized setae laterally to crista, close to
the level of posterior sensillae. Two pairs of eyes laterally to scutum, each pair situated on
small protrusions of a common ocular sclerite. Palp tibia with unidentate basidont (6-13 e).
Sharkfin-like cuticular extensions present distally to the bases of setae on the palp tibia (6-
13 e) and tibiae of legs in adults (absent in deutonymphs). Three pairs (adult) or two pairs
(deutonymph) of genital acetabula. Larva: Idiosoma behind the level of scutum covered
with rows of barbed setae, each seta inserted on a plate-like sclerite (6-12 f). Scutum
roughly triangular with projecting naso. One pair of trichobothria with setiform sensillae
(6-12 c). Laterally on scutum two pairs of prominent setae and additionally one pair of
small setae in anterior position. Laterally to scutum two pairs of eyes, each located on
an ocular sclerite. Legs I-III with partially divided femora. Tarsi I-III each bearing two
claws, the anterior one in all cases distinctly larger than the posterior one. Tarsus I without
small accompanying seta z. Subterminal eupathidium f prominent, curved backwards.
Key to species
Ex clude d: D. moldavicum F EIDER, 1959 (nomen dubium).
Larvae
1 More than six setae on rows C, D . . . . . . . . . . . . . Diplothrombium longipalpe (page 190)
– Six setae on rows C, D . . . . . . . . . . . . . . . . . . . . . ........................... 2
2 Lateral seta (1b) on coxa I setulose . . . . . . . . . . . . . Diplothrombium ludwinae (page 190)
– Lateral seta (1b) on coxa I bifid . . . . . . . . . . . . . . . ........................... 3
3 Seta (2b) on coxa II simple . . . . . . . . . . . . . . . . . . Diplothrombium zbigniewi (page 191)
– Seta (2b) on coxa II setulose . . . . . . . . . . . . . . . . . . . Diplothrombium rackae (page 190)

1 Idiosoma setae short (40 lm in adults), on prominent sclerites densely covering the dorsal side of
idiosoma and forming a polygonal network (6-14 a) Diplothrombium carpaticum (page 190)
– Idiosoma setae 50 lm or longer in adults, setal sclerites less well sclerotized (6-14 b-c) . . . 2
2 Idiosoma setae (65-70 lm in adults) inserted in cylindrical sclerites (6-14 b), basidont located
halfway between base of tibia and base of odontus (6-14 d) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplothrombium wittei (page 191)
– Idiosoma setae (55-65 lm in adults) inserted in truncated cone-shaped sclerites (6-14 c), basidont
in distal half of tibia close to odontus (6-14 e) . . . . . Diplothrombium longipalpe (page 190)
Fig. 6-14: Diplothrombium adults, dorsal view: a, D. carpaticum, detail: setae; b, d, D. wittei;
b, detail: setae; d, detail: palp (lateral); c, e, D. longipalpe; c, detail: setae; e, detail: palp (lateral);
(a-e after W OHLTMANN et al. 2004; scale bars = 50 lm).
190 Special part
1. Diplothrombium carpaticum (ŠS TORKAN , 1938) (6-11 d; 6-14 a)

1938 Johnstoniana carpatica ŠS TORKAN, Zool. Anz. 121 (5-6): 110.
1947 Diplothrombium carpaticum T HOR & W ILLMANN, Das Tierreich 71b: 223.
D e s c r i p t i o n : ŠS TORKAN (1938), W OHLTMANN et al. (2004) (adult); larva unknown.
Adult: scutum with two pairs of non-specialized setae at the level of posterior tricho-
bothria. The remaining part of idiosoma covered with numerous smooth, spine-like setae,
directed backwards. Setae inserted on broad, slightly elevated sclerites of irregular shape
(approximately 25 lm in diameter), which are strongly sclerotized at the margins. Palp
tarsus with three eupathidia, ratio of palp tibia length to distance between bases basi-
dont/odontus = 4.5.
Di stri bu ti on: Poland, Slovakia, Ukraine.
2. Diplothrombium longipalpe (B ERLESE , 1887) (6-12 c, f; 6-13 d-e; 6-14 c, e)

1887 Trombidium longipalpe B ERLESE, Acari, Myriopoda et Scorpiones hucusque in Italia reperta.
XLII No. 2.
1910 Diplothrombium longipalpe B ERLESE, Redia 6: 357.
1950a Diplothrombium bujori F EIDER, An. Acad. R.P.R. 3 (15): 576.
D e s c r i p t i o n : B ERLESE (1887) (adult), W OHLTMANN et al. (2004) (active instars). Adult:
Colour in life dark red to red-brown. Scutum with two pairs of trichobothria with smooth
sensillae, additionally two pairs of non-specialized smooth setae (64-75 lm). Idiosoma cov-
ered with numerous smooth, spine-like setae, directed backwards, inserted on rounded,
slightly asymmetrical tubercles (25-30 lm in diameter). Venter covered with numerous
spine-like setae situated on rounded, slightly asymmetrical tubercles (15-20 lm in diame-
ter). Palp tarsus with seven to ten eupathidia, ratio length palp tibia to distance between
bases basidont/odontus = 5.8 to 8.0. Larva: Colour in life reddish. Posterior to scutum 38-
44 smooth setae arranged in rows (with eight or more setae in dorsal rows C, D), each seta
placed on an ovoid sclerite (50-55 lm long, 35 lm wide). Coxa I with bifurcate seta 1b (75-
90 lm) and another bifurcate seta 1a (50-65 lm) on pars medialis. Coxa II with simple seta
2b (75-85 lm), between coxae I-II the Claparède’s organ. Coxa III with simple seta 3b (45-
55 lm); seta 3a simple, smooth (30-35 lm) and located outside the coxa. Posterior to coxa
III approximately 50 smooth setae inserted on small platelets (about 15 lm in diameter).
Genu I with 9-11 solenidia.
H a bit at an d B i olo gy : Probably uni- to semivoltine. Larva abundant in late June, para-
sitic on adult Ptychopteridae and Dixidae (Diptera). Confined to amphibious (limnic) bio-
topes.
Di stri bu ti on: Germany, Great Britain, Hungary, Italy, Norway, Poland, Romania, Spain,
Switzerland.
3. Diplothrombium ludwinae H AITLINGER , 1993

1993 Diplothrombium ludwinae H AITLINGER, Tijdschr. Entomol. 136: 11.
D e s c r i p t i o n : H AITLINGER (1993), W OHLTMANN et al. (2004) (larva); postlarval instars
unknown. Larva: With seven to eight solenidia on genu I, seta 1b on coxa I setulose, seta 2b
on coxa II setulose. Dorsal idiosoma with each six setae in dorsal rows C, D.
Di stri bu ti on: Poland.
4. Diplothrombium rackae W OHLTMANN , M A˛ KOL & G ABRYŚ , 2004

2004 Diplothrombium rackae W OHLTMANN et al., Ann. Zool. 54 (3): 612.
D e s c r i p t i o n : W OHLTMANN et al. (2004) (larva); postlarval instars unknown. Larva:

With six to nine solenidia on genu I, seta 1b on coxa I bifid, seta 2b on coxa II setulose.
Dorsal idiosoma with each six setae in dorsal rows C, D.
H a bit at a nd B io lo gy : Larvae parasitize Limoniinae.
Di stri bu ti on: Austria, Germany.
5. Diplothrombium wittei W OHLTMANN , M A˛ KOL & G ABRYŚ , 2004

(6-13 g; 6-14 b, d)
2004 Diplothrombium wittei W OHLTMANN et al., Ann. Zool. 54 (3): 614.
D e s c r i p t i o n : W OHLTMANN et al. (2004) (adult); larva unknown. Adult: Scutum with
two pairs of trichobothria and two pairs of non-specialized smooth setae placed close
to posterior trichobothrium. Idiosoma with numerous smooth, spine-like setae, directed
backwards, inserted on elevated platelets, the shape of which varies from rounded, slightly
asymmetrical in the anterior part of idiosoma to cylindrical (15-20 lm in diameter) in the
posterior part. Palp tarsus with three to four eupathidia, ratio length palp tibia to distance
between bases basidont/odontus = 2.0 to 2.3.
H a bit at a nd B io lo gy : Confined to amphibious (limnic) biotopes.
Di stri bu ti on: Germany.
6. Diplothrombium zbigniewi H AITLINGER , 2001

2001a Diplothrombium zbigniewi H AITLINGER, Syst. Appl. Acarol. 6: 179.
D e s c r i p t i o n : H AITLINGER (2001a), W OHLTMANN et al. (2004) (larva); postlarval instars
unknown. Larva: With six solenidia on genu I, seta 1b on coxa I bifid, seta 2b on coxa II
simple. Dorsal idiosoma with each six setae in dorsal rows C, D.
Genus Johnstoniania G EORGE , 1909

1909b Johnstoniana G EORGE, Naturalist 631: 281.
1911a Rohaultia O UDEMANS, Ent. Ber. (Amst.) 3 (57): 121.
1911b Musitania O UDEMANS, Ent. Ber. (Amst.) 3 (62): 186.
Typus generis: Rhyncholophus errans J OHNSTON , 1852, by original designation.
Di ag no sis: Adult and Deutonymph: Medium to large in size, body ovoid, elongate (6-13
f). Idiosoma covered with spine-like setae, each located on a sclerite (6-15 g-i). Scutum
triangular, with crista metopica and anteriorly projecting naso. Two pairs of trichobothria
with setiform sensillae, additionally 4-36 non-specialized setae present (6-13 a). Two pairs
of eyes laterally to scutum, each pair situated on a small protrusion of a common ocular
sclerite. Palp tibia with bidentate basidont (6-13 c). Three pairs (adult) or two pairs (deu-
tonymph) of genital acetabula. Larva: Posterior part of idiosoma covered with rows of
barbed setae, each inserted on a plate-like sclerite (6-10 b). Scutum roughly triangular
with projecting naso, two pairs of trichobothria with setiform sensillae, additionally
two pairs of non-specialized setae (6-12 b). Laterally to scutum two pairs of eyes, each
pair located on an ocular sclerite. Legs I-III with completely divided femora. Tarsus I
with one small accompanying seta z near dorsal eupathidium (6-15 a, b).
Key to species
Ex clude d: J. helvetica (C OOREMAN , 1959) (nomen dubium).
192 Special part
Larvae
1 Tarsus I with one solenidion (x) and two eupathidia (n) (6-15 a); width and length of scutum less
than 140 lm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Tarsus I with more than four x and more than 10 n (6-15 b); width and length of scutum more
than 145 lm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Odontus of palp tibia slightly curved, length of tibial claw 1/5 of total tibia length (6-15 d),
digitus mobilis of chelicera with one small distal tooth (6-15 c) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana eximia (page 194)
– Odontus of palp tibia prominent, strongly curved, length of tibial claw 1/2 of total tibia length
(6-15 e), digitus mobilis of chelicera with one small distal and one prominent proximal tooth
(6-15 l)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana parva (page 194)
3 Tarsus II with two to four club-shaped solenidia (x) (6-15 f) . Johnstoniana rapax (page 195)
– Tarsus II with one club-shaped x . . . . . . . . . . . . . . . . . . . Johnstoniana errans (page 192)

1 Idiosoma setae on prominent tubercles (6-15 g-h). More than six setae on scutum in addition to
posterior trichobothria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Idiosoma setae on flat ovoid sclerites (6-15 i). Less than seven setae on scutum in addition to
posterior trichobothria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Idiosoma setae on truncated cone-shaped tubercles (6-15 g), tarsus II with two to three club-
shaped solenidia (x) dorsally . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana rapax (page 195)
– Idiosoma setae on prominent cylindrical tubercles (6-15 h), tarsus II with a single club-shaped x
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana errans (page 192)
3 About half of setae on tarsus I setulose, with many setules (6-15 j) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnstoniana eximia (page 194)
– All setae on tarsus I smooth (6-15 k) . . . . . . . . . . . . . . . . . Johnstoniana parva (page 194)
1. Johnstoniana errans (J OHNSTON , 1852) (6-13 a, c; 6-15 b, h)

1852 Rhyncholophus errans J OHNSTON, B.N.C. vol. III (3): 113.
1909b Johnstoniana errans G EORGE, Naturalist 631: 281.
1911a Rohaultia biungulum O UDEMANS, Ent. Ber. (Amst.) 3 (57): 122.
1911b Musitania verrucipes O UDEMANS, Ent. Ber. (Amst.) 3 (62): 186.
1916 Diplothrombium eximium var. insignius B ERLESE, Redia 12: 22.
1922 Diplothrombium longipes S CHWEIZER, Verhdl. Naturf. Ges. Basel 33: 85.
1947 Johnstoniana insignia T HOR & W ILLMANN, Das Tierreich 71b: 226.
1949 Johnstoniana errans C OOREMAN, Bull. Inst. Roy. Sci. Nat. Belg. 25 (2): 1 [part.].
1967 Johnstoniana franzi R OBAUX, Mém. Mus. Nat. Hist. Nat. Paris, (n.s.), Sér. A, Zool. 46 (1): 24.
1970 Johnstoniana errans R OBAUX, Acarologia, 12 (2): 339.
Rejected attribution: Non Johnstoniana errans sensu T HOR & W ILLMANN (1947), D ANIEL
(1956).
D e s c r i p t i o n : B ERLESE (1916), C OOREMAN (1949) (adult), R OBAUX (1970) (larva),
W OHLTMANN et al. (2004) (active instars). Adult and Deutonymph: With cylindrical basal
sclerites of body setae, no setulose setae on tarsus I, one club-shaped solenidion on tarsus II,
one eupathidium on basifemur IV, ratio length palp tibia to distance between bases basi-
dont/odontus = 5.4 to 5.9. Large species, adults reach 3200 lm length, 2000 lm width.
Larva: With 6-20 solenidia and 15-20 eupathidia on tarsus I, one club-shaped solenidion
and three eupathidia on tarsus II, four to five solenidia on genu II, two to three solenidia on
genu III, two distal teeth on cheliceral blade.
Fig. 6-15: Johnstoniana larvae and adults: a, c, d, J. eximia (larva); a, detail: tarsus I; c, detail: chelicera
tip (after W OHLTMANN et al. 1994, W ENDT et al. 1994); d, detail: palp tibia; b, J. errans (larva), detail
tarsus I; e, l, J. parva (larva); e, detail: palp tibia (after W ENDT et al. 1994); l, chelicera tip (after
W OHLTMANN et al. 1994, W ENDT et al. 1994); f, J. rapax (larva), detail: tarsus II (after W ENDT &
EGGERS 1996); g, J. rapax (adult), detail: body setae; h, J. errans (adult), detail: body setae; i-j, J. eximia
(adult); i, detail: body setae; j, detail: tarsus I; k, J. parva (adult), detail: tarsus I; (a, b, d, g-k after
W OHLTMANN et al. 2004; scale bars = 50 lm).
194 Special part
H a bit at a nd B io lo gy : Uni- to semivoltine. Postlarval instars occur throughout the year,

mostly in the moss and litter layer (W OHLTMANN 1996). Eggs display obligatory diapause
during winter, postlarval instars are also able to survive throughout the winter. Abundance
of larvae restricted to a few weeks in May/June (data from northern Germany). Larvae
parasitize adult Tipula spp. (Diptera: Tipulidae) and display preparasitic attendance on
tipulid larvae/pupae. Confined to amphibious (limnic) biotopes in forested areas.
Di stri bu ti on: Austria, Belgium, Czech Republic, France, Germany, Great Britain, Hun-
gary, Italy, Poland, Romania, Slovakia, Spain, Switzerland, The Netherlands.
2. Johnstoniana eximia (B ERLESE , 1910) (6-10 b; 6-12 b; 6-15 a, c-d, i-j)

1910 Diplothrombium eximium B ERLESE, Redia 6: 357.
1947 Johnstoniana errans T HOR & W ILLMANN, Das Tierreich 71b: 225 [part.].
1951 Johnstoniana tuberculatua S CHWEIZER (lapsus calami), Ergeb. Wiss. Unters. Schweiz. Nat.
Parks (n.f.) III 23 (2): 83.
1956 Johnstoniana errans D ANIEL 1956, Zool. Zhurn. 35 (10): 1463.
D e s c r i p t i o n : T HOR & W ILLMANN (1947) (adult), W OHLTMANN et al. (1994, 2004) (ac-
tive instars, as J. tuberculata). Adult and Deutonymph: With flat basal sclerites of body
setae, setulose setae present on tarsus I, one club-shaped solenidion on tarsus II, eupathi-
dium on basifemur IV absent, ratio length palp tibia to distance between bases basidont /
odontus = 5.8 to 8.7. Medium sized species, adults reach 1500 lm length, 1000 lm width.
Larva: With one solenidion and two eupathidia on tarsus I, one club-shaped solenidion and
one eupathidium on tarsus II, each two solenidia on genu II and III, one distal tooth on
cheliceral blade.
H a bit at an d B io lo gy : Uni- to semivoltine. Postlarval instars occur throughout the year
and display a hidden life style in the moss and litter layer (W OHLTMANN et al.1994). Eggs
with obligatory diapause during winter, postlarval instars are also able to survive winter.
Abundance of larvae restricted to a few weeks in May/June (data from Northern Germany).
Larvae parasitize imagines of Limonia spp. (Diptera: Tipulidae) and display preparasitic
attendance on host larvae/pupae (W OHLTMANN et al. 1994). Confined to amphibious (lim-
nic) biotopes including shores of creeks and ponds.
Di stri bu ti on: Austria, Germany, Italy, Poland, Romania, Switzerland.
3. Johnstoniana parva W ENDT, W OHLTMANN , E GGERS & O TTO , 1994 (6-15 e, k-l)
1994 Johnstoniana parva W ENDT et al., Acarologia 35 (1): 52.
D e s c r i p t i o n : W ENDT et al. (1994), W OHLTMANN et al. (2004) (active instars). Adult and
Deutonymph: With flat basal sclerites of body setae, no setulose setae on tarsus I, one club-
shaped solenidion on tarsus II, eupathidium on basifemur IV absent, ratio length palp tibia
to distance between bases basidont/odontus = 4.9 to 6.3. Medium sized species, adults
reach 1500 lm length, 1000 lm width. Larva: With one solenidion and two eupathidia
on tarsus I, one club-shaped solenidion and one eupathidium on tarsus II, one solenidion
on genu II, one to two solenidia on genu III, two distal teeth on cheliceral blade.
H a bit at a nd Bi ol og y: Uni- to semivoltine. Postlarval instars abundant throughout the
year. Eggs deposited in autumn display diapause and larvae hatch in May/June of the fol-
lowing year. After hibernation, females may also deposit eggs in spring; these eggs do not
diapause and larvae hatch in June/July. Larvae parasitize postlarval instars of Johnstonia-
nidae and Microtrombidiidae (Acari: Parasitengona) (W ENDT et al. 1994). Confined to
amphibious (limnic) biotopes.
Di stri bu ti on: Germany, Poland.
4. Johnstoniana rapax W ENDT & E GGERS , 1996 (6-15 f-g)

1996 Johnstoniana rapax W ENDT & E GGERS , Mitt. Hamb. Zool. Mus. Inst. 91: 114.
D e s c r i p t i o n : W ENDT & E GGERS (1996), W OHLTMANN et al. (2004) (active instars).
Adult and Deutonymph: With rounded, slightly asymmetrical basal sclerites of body setae,
no setulose setae on tarsus I, two or more club-shaped solenidia on tarsus II, one or two
eupathidia on basifemur IV, ratio length palp tibia to distance between bases basidont/
odontus = 3.8 to 4.6. Large species, adults reach 2500 lm length, 2000 lm width. Larva:
With five to eight solenidia and 18-25 eupathidia on tarsus I, two to four club-shaped so-
lenidia and one or two eupathidia on tarsus II, four to six solenidia on genu II, three to five
solenidia on genu III, two distal teeth on cheliceral blade.
H a bit at an d B io lo gy : Strictly univoltine with diapausing eggs as the only hibernating
instar. Emergence of larvae restricted to a few weeks in May/June, deutonymphs in June/
July, adults in July to September (data from northern Germany). Larvae not parasitic but
predatory on small arthropods (E GGERS 1995, W OHLTMANN et al. 1999, 2004). Confined
to amphibious (limnic) biotopes.
Di stri bu ti on: Finland, Germany, Poland.
Family Microtrombidiidae T HOR , 1935

1935 Microtrombidiidae T HOR , Zool. Anz. 110: 47.
Di ag no sis: Adult and Deutonymph: Small- to medium-sized. Colouration reddish. Cu-
ticle covered with dense setation (6-11 k), setae setiform or modified (6-18; 6-19). Prodor-
sal shield narrow, almost reduced to elongate crista metopica, area sensilligera located at
the posterior part of crista metopica (6-19 b, d). Palp with characteristic tibia carrying
dorsally one or two rows of spines (ctenidia) (6-11 i; 6-19 c, e; 6-20 c, d, g), further
spine-like setae situated ventro-laterally (basidont) or behind (paradont) the odontus.
Two pairs of eyes laterally to crista metopica, not distinctly stalked. Adults with three,
deutonymphs with two pairs of genital acetabula, genital valves covered with numerous
setae. Larva: Small to medium-sized with scutum, scutellum (6-3 a) and sometimes addi-
tional unpaired medial sclerites. Scutum usually large, its anterior margins often bent ven-
trally (6-3 a; 6-16 f). Posterior idiosoma setae simple (smooth or with few small barbs)
arranged in rows and inserted on prominent platelets (6-3 a). Anal pore present, without
surrounding sclerites. Coxae I-II-III with two-two-one setae (6-3 a; 6-16 e, f). Femora of
legs undivided. Famuli and eupathidia present on tarsi I (6-3 b) and II. Pretarsus III often
modified (6-16 a; 6-17 e, f). Supracoxal setae on palp and leg I absent.
H a bit at a nd B iol og y : Larvae originally parasitize dipteran insects, some species
switched to other hosts (Orthoptera, Coleoptera). Deutonymphs and adults of most species
feed on eggs of Diptera and display an edaphic life style; presence on soil surface is almost
restricted to the mating period. Some species confined to amphibious biotopes are the do-
minant terrestrial Parasitengona in this habitat.
Key to genera
Ma r ked w ith an as te r is k*: genera containing species regularly found in amphibious biotopes
Ex clude d: Cercothrombium loricatum M ETHLAGL , 1928; Ettmuelleria kasjani H AITLINGER , 1998;
Kamitrombidium stellae H AITLINGER , 2001b; Stirlitrombidium dominikae H AITLINGER , 1998; Tri-
chotrombidium muscarum (R ILEY, 1878) (larva); Willmannella hilmari H AITLINGER , 1998; Willman-
nella racovitzai (F EIDER , 1948) (larva). For these taxa the available descriptions of larval forms do not
allow unambiguous determination. Postlarval instars of Willmannella racovitzai and Trichotrombi-
dium muscarum are identifiable, but of the remaining species only the larva is known. The following
196 Special part
key tentatively includes Eutrombidium into Microtrombidiidae and does not follow the traditional
ranking of Eutrombidiidae as family and as sister taxon of Microtrombidiidae, because phylogenetic
evaluation of larval characters suggest a closer relationship of Eutrombidium and related genera to the
higher Microtrombidiidae (W OHLTMANN, in press). The definitive ranking of Eutrombidium and re-
lated genera is still under discussion (M A˛ KOL & G ABRYŚ 2005).
Larvae
1 Pretarsus I-III each with paired claws and claw-like empodium, tarsus III with lophotrix and
scopa, inner claw of pretarsus III modified (smilum) (6-16 a), scutellum without median ridge 2
– Pretarsus I/II without empodium, tarsus III with empodium and two claws, lophotrix and scopa
may be present on tarsus III, inner claw of pretarsus III normal or modified, scutellum with me-
dian ridge (6-16 b, 6-17 g) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 Lophotrix elongate and distally bifurcate (6-16 a), habitus as in 6-3. . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microtrombidium* (page 211)
– Lophotrix not distally bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Stephanostome smooth, may be inconspicuous, without marginal teeth (6-16 c) . . . . . . . . 4
– Stephanostome distinct, with about 30 marginal teeth (6-16 d) . . . . . . . . . . . . . . . . . . . . 7
4 Setae on coxae (1b, 2b, 3b) and hypostomal (bs) setae at gnathosoma (6-16 c) stout, the last two
pairs of posterior dorsal setae (h1+2) on idiosoma prolonged (6-16 e) . . . . . . Eutrombidiinae
In Central Europe only Eutrombidium trigonum (H ERMANN , 1804)
– Setae on coxae I-III setiform, hypostomal seta bs with finger-like projections (6-16 d), only one
pair of posterior dorsal setae on idiosoma (h2) prolonged (6-16 f) . . . . . . . . . . . . . . . . . . 5
5 Scutellum with four to eight (usually six) setae. . . . . . . . . Sucidothrombium F EIDER, 1973.
In Central Europe only Sucidothrombium sucidum (L. K OCH , 1879)
– Scutellum with two setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Stephanostome conspicuous, without protruding lamellae (6-16 g), solenidion (r) on genu I-III
long, extending over tibia (6-16 h) . . . . . . . . . . . . . . . . . Dactylothrombium F EIDER, 1952
In Europe only Dactylothrombium pulcherrimum (H ALLER , 1882)
– Stephanostome inconspicuous, dorsally with protruding lamellae (6-17 a), r on genu I-III not
extending over tibia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinothrombium* (page 208)
7 Sclerites d1 clearly separated (6-16 h, 6-17a, e), median and lateral dorsal idiosoma setae of about
the same size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Sclerites d1 close together with parallel median sides, lateral dorsal setae on idiosoma much smal-
ler than median ones (6-17 b). . . . . . . . . . . . . . . . . . . . . . Camerotrombidium* (page 204)
8 At least one solenidion u in proximal half of tibia I, famulus (e) on tarsus I proximally to x
(6-17 c).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Both solenidia (u ) on tibia I in distal half, famulus (e) on tarsus I distally to solenidion (x) (6-17 d)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campylothrombium* (page 207)
9 Lophotrix small, with few setules (6-17 e), seta 1 a on coxa I smooth . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractothrombium* (page 202)
– Lophotrix prominent, inner branch with numerous long setules (6-17 f), seta 1a barbed . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platytrombidium T HOR , 1936
10 Tarsus III without lophotrix or scopa, pretarsus III with two normal claws and empodium (6-17 g)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium* (page 213)
– Tarsus III modified with lophotrix, scopa and smilum (6-17 e) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enemothrombium* (page 209)
Fig. 6-16: Microtrombidiidae larvae: a, Microtrombidium pusillum, detail: tarsus III (after G ABRYŚ &
W OHLTMANN 2001); b, Enemothrombium bifoliosum (dorsal) (after W OHLTMANN & G ABRYŚ in
press); c, Eutrombidium trigonum, detail: gnathosoma ventral; d, Atractothrombium sylvaticum, de-
tail: gnathosoma ventral (after G ABRYŚ et al. 2005); e, Eutrombidium trigonum (ventral); f, Campy-
lothrombium clavatum (ventral), detail: hypostomal setae bs; g-h, Dactylothrombium pulcherrimum
(after W OHLTMANN & G ABRYŚ 2003); g, detail: gnathosoma ventral; h, dorsal view; (scale bars = 50 lm).
198 Special part
Fig. 6-17: Microtrombidiidae larvae: a, Echinothrombium rhodinum (dorsal), detail: gnathosoma

ventral (after G ABRYŚ & W OHLTMANN 2003); b, Camerotrombidium pexatum (dorsal), detail: tarsus
III (after W OHLTMANN et al. 2003); c, e, Atractothrombium sylvaticum (dorsal) tibia I (after G ABRYŚ et
al. 2005); c, detail tibia I; e, detail: tarsus III; d, Campylothrombium clavatum, detail: tibia I; f, Pla-
tytrombidium fasciatum, detail: tarsus III (after G ABRYŚ et al. 2005); g, Valgothrombium valgum (dor-
sal) (after W OHLTMANN & W ENDT 1996); (scale bars = 50 lm).

1 No shield on posterior dorsum of idiosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– One distinct large sclerite dorsally at the posterior end of idiosoma . . . . . . . Eutrombidiinae
In Central Europe only Eutrombidium trigonum (H ERMANN , 1804)
2 Two distinctly different types of posterior dorsal setae . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Posterior dorsal setae may vary in size but belong to one single type . . . . . . . . . . . . . . . 10
3 Setiform posterior dorsal setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– At least one type of posterior dorsal setae setiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 One type of idiosoma setae spherical or slightly elongate with numerous tubercles in distal half.
The other type of setae longer, without tubercles and deeply incised apically (6-18 a) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enemothrombium* (page 209)
– Setae of different shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Larger posterior dorsal setae inflated, with two chambers; smaller posterior dorsal setae trumpet-
shaped (6-18 b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Willmannella F EIDER , 1952
– Larger posterior dorsal setae without chambers, divided into two parts, both inflated, proximally
setulose, distally bare; smaller posterior dorsal setae in the form of multi-rayed star with cylind-
rical extension distally (6-18 c). . . . . . . . . . . . . . . . . . . Trichotrombidium K OBULEJ , 1951
In Central Europe only Trichotrombidium muscarum (R ILEY, 1878)
6 Both types of posterior dorsal setae setiform; first type short with thin stem; second type long with
stout stem (6-18 d) . . . . . . . . . . . . . . . . . . . . . . . . . . Dimorphothrombium F EIDER , 1952
– First type of posterior dorsal setae setiform; second type of different shape . . . . . . . . . . . . 7
7 At least two distinct spines (basidonts) ventrally to odontus on palp tibia; second type of posterior
dorsal setae covered with very long setules along the complete length, with three prongs distally
(6-18 e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trischidothrombium F EIDER , 1952
In Central Europe only Trischidothrombium discrepans (W ILLMANN , 1950)
– One basidont; second type of posterior dorsal setae of different shape . . . . . . . . . . . . . . . 8
8 Eyes on prominent stalks; second type of posterior dorsal setae covered with setules along the
complete length, with thin, long stem and two to four short lobes distally (6-18 f) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mirabilithrombium G ABRYŚ , 1999
In Central Europe only Mirabilithrombium mirabilis (W ILLMANN , 1950)
– Eyes sessile; second type of posterior dorsal setae of different shape . . . . . . . . . . . . . . . . . 9
9 Some dorsal idiosoma setae blade-like, much longer than surrounding setiform setae (6-18 g);
brush-like setae absent from idiosoma . . . . . . . . . . . . . . . . . Echinothrombium* (page 208)
– Some dorsal idiosoma setae brush-like (6-18 h), blade-like setae absent.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milandanielia G ABRYŚ , 1999
In Central Europe only Milandanielia intermedia (F EIDER , 1950)
10 Dorsal idiosoma setae expanded distally, with lobate, digitate or leaf-like processes (6-19 a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylothrombium F EIDER, 1952
In Central Europe only Dactylothrombium pulcherrimum (H ALLER , 1882)
– Dorsal idiosoma setae of different shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Anterior border of crista metopica widened (vertex) (6-19 b), in adults palp tibia with two rows
of spines (ctenidia) (6-19 c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
– Anterior border of crista metopica knob-like (6-19 d), palp tibia with one row of spines (ctenidia)
(6-19 e), dorsal idiosoma setae setiform with fine setules (6-19 f). . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium* (page 213)
12 Dorsal idiosoma setae setiform (6-19 g). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
– Dorsal idiosoma setae inflated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
13 Legs I and IV longer than idiosoma; length/width ratio of tarsus I in adults 4 3.5, in deuto-
nymphs 4 2.7; number of spinisetae in distal and in proximal ctenidium 4 10 . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dromeothrombium B ERLESE , 1912
In Central Europe only Dromeothrombium coiffaiti R OBAUX , 1966a
– Legs I and IV shorter than idiosoma; length/width ratio of tarsus I in adults 5 2.5, in deuto-
nymphs 5 2.0; number of spinisetae in distal and in proximal ctenidium 5 10 . . . . . . . 14
200 Special part
Fig. 6-18: Microtrombidiidae, body setae of adults: a, Enemothrombium bifoliosum (after W OHLT-
MANN & G ABRYŚ in press); b, Willmannella sp. (after G ABRYŚ 1999); c, Trichotrombidium muscarum
(after G ABRYŚ 1999); d, Dimorphothrombium sp. (after G ABRYŚ 1999); e, Trischidothrombium sp.
(after G ABRYŚ 1999); f, Mirabilithrombium sp. (after G ABRYŚ 1999); g, Echinothrombium rhodinum
(after G ABRYŚ & W OHLTMANN 2003); h, Milandanielia intermedia (after G ABRYŚ 1999); (scale bars =
50 lm).
Fig. 6-19: Microtrombidiidae adults: a, Dactylothrombium pulcherrimum, detail: body setae (after
W OHLTMANN & G ABRYŚ 2003); b, c, g, Microtrombidium pusillum (after G ABRYŚ & W OHLTMANN
2001); b, detail: scutum, eyes; c, detail: palp; g, body setae; d-f, Valgothrombium valgum; d, detail:
scutum, eyes; e, detail: palp; f, detail: body seta; h, l, Campylothrombium clavatum; h, detail: brush-
like leg setae; l, body setae; i, Atractothrombium sylvaticum, detail: body setae (after G ABRYŚ et al.
2005); j, Platytrombidium fasciatum, detail: body setae (after G ABRYŚ et al. 2005); k, Camerotrom-
bidium pexatum, detail: body setae; (scale bars = 50 lm).
202 Special part
14 Length/width ratio of tarsus I both in adults and deutonymphs 5 1.5; basidont slender; one distal
solenidion on palp tarsus . . . . . . . . . . . . . . . . . . . . . . . . . . Microtrombidium* (page 211)
– Length/width ratio of tarsus I in adults 4 _ 2.0, in deutonymphs 4 _ 1.5; basidont stout; at least three
distal solenidia on palp tarsus. . . . . . . . . . . . . . . . . . . . . . Sucidothrombium F EIDER, 1973
In Central Europe only Sucidothrombium sucidum (L. K OCH , 1879)
15 Dorsal idiosoma setae without chambers, fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
– Dorsal idiosoma setae with at least one chamber, some setae on palp femur, palp genu, and basal
segments of legs I-IV brush-like (6-19 h) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 Dorsal idiosoma setae less than 2.5 times longer than wide, body colour in life reddish, uniform
(6-19 i) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractothrombium* (page 202)
– Dorsal idiosoma setae more than three times longer than wide, body colour in life may be red
with white transversal stripes (6-19 j) . . . . . . . . . . . . . . . . . . . . . . . . . . . Platytrombidium
17 Dorsal idiosoma setae inflated (spherical, spheroidal or calyciform) with an apical opening lead-
ing to a chamber; inserted on distinct papillae, usually with protruding denticles forming a collar
around the setal stem, papillae at least 1/3 of the whole seta length (6-19 k)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camerotrombidium* (page 204)
– Dorsal idiosoma setae thick, clavate and with inflated tip, with two closed chambers. Distally
covered with tubercles, papillae without protruding denticles (6-19 l)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campylothrombium* (page 207)
Genus Atractothrombium F EIDER , 1952

1952 Atractothrombium F EIDER, Bul. Şti. Sect. Sti. Biol. Agr. Geol. Geogr. Acad. Repub. Pop. Rom.
4 (3): 609.
1945 Foliotrombidium W OMERSLEY, Rec. S. Aust. Mus. 8 (2): 342 [part.].
Typus generis: Microtrombidium fusicomum B ERLESE , 1910, by original designation.
Di ag no sis: Adult: Medium sized, length about 1000 lm, width about 800 lm. Medial
surface of palp tibia with two, very often overlapping ctenidia and a radula. Radula con-
spicuous, spine-like setae usually strong and numerous. Lateral surface of palp tibia with
one well developed, strong basidont. Palp tarsus cylindrical, elongate. Aspidosoma trian-
gular in outline. Crista metopica typical, with anterior and posterior processes and well
sclerotized circular or oval sensillary area bearing two filiform sensillary setae. Accessory
posterior process usually present. Double eyes sessile or with tendency to form a very short
stalk, situated at sides of crista metopica, more or less at the level of mid-point of anterior
process (6-11 h). Posterior dorsal setae of same type, short, stout, fusiform, less than three
times longer than wide, all covered with very numerous, delicate, short setulae (6-20 a, b).
The latter sometimes arranged in regular rows. Legs without lamellae, much shorter than
idiosoma. Tarsus I dilated, rounded distally, longer than wide, always longer than tibia I.
Deutonymph: Medial surface of palp tibia with one ctenidium. Number of spine-like setae
forming radula reduced. Lateral surface of palp tibia with one basidont. Idiosoma setae
similar to that in adults. Two pairs of genital acetabula. Larva: Gnathosoma with stepha-
nostome and internal horseshoe-like sclerite bearing 35-40 prominent lateral teeth (6-16 d),
palp with minute setae on femur and genu. Tritorostral (bs) setae distally with finger-like
projections (6-16 d). Dorsum of idiosoma with stolascutum bearing 3 2 peripheral setae
plus two trichobothria (6-17 e). Scutellum with two c1 setae. C2 plates slightly enlarged,
oval in shape, d1 plates much larger than c2 and almost round in shape. Setae on coxa I
smooth or bifid, setae on coxae II and III with three to four small setules. Tarsus III with
modified inner claw (smilum), trifid outer claw, scopa, and a setulose lophotrix, which is
smaller than the smilum (6-17 e).
Key to species
Ex clude d: Atractothrombium schweizeri (S OUTHCOTT, 1994) syn. Microtrombidium simulans var.
minor S CHWEIZER, 1922 (Switzerland, description insufficient for proper interpretation). For com-
ments see G ABRYŚ (1999) and G ABRYŚ et al. (2005).
Larvae
Only species described for the area: Atractothrombium sylvaticum* (page 204)

1 Posterior idiosoma setae regularly fusiform, all pointed apically (6-20 a) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractothrombium sylvaticum* (page 204)
– Posterior idiosoma setae irregularly fusiform, the majority of setae rounded apically (6-20 b) 2
2 Basidont long (60-70 lm), situated at the palp tarsus base (6-20 c) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractothrombium fusicomum* (page 203)
– Basidont short (c. 30 lm), situated in the middle between palp tarsus and odontus bases (6-20 d)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractothrombium rafalskii* (page 203)
1. Atractothrombium fusicomum (B ERLESE , 1910) (6-20 c)

1910 Microtrombidium (Enemothrombium) fusicomum B ERLESE, Redia 6: 360.
D e s c r i p t i o n : R OBAUX (1967 - deutonymph), G ABRYŚ (1996 - adult). Adult: Distal cte-
nidium of palp tibia composed of four to six strong spinisetae (+ paradont); proximal cte-
nidium consisting of five to nine slightly weaker and thinner spinisetae. Lateral surface of
palp tibia at palp tarsus base with a long basidont (60-70 lm). Apex of palp tarsus with five
solenidia, lateral side with one. Posterior idiosoma setae of one type, very short (15-20 lm),
irregularly fusiform, slightly pointed or rounded apically, relatively broad at base, all cov-
ered with a lot of short setulae. Legs much shorter than idiosoma; tarsus I (200-280 lm
long, 105-180 lm wide) oval-rounded, length/width ratio always less than 2/1; tibia I short
(165-190 lm). Deutonymph: Generally smaller. Body length about 650 lm, width about
400 lm. Distal ctenidium of palp tibia consists of four spinisetae (+ paradont), proximal
ctenidium absent. Posterior idiosoma setae shorter (12 lm). Legs shorter, tarsus I c. 155 lm
long, c. 90 lm wide, tibia I c. 90 lm long. Larva unknown.
H a bit at a nd B io lo gy : Hygrophilous (G ABRYŚ 1996).
Di stri bu ti on: Europe (accurate localities can not be given due to numerous misidenti-
fications of the species).
2. Atractothrombium rafalskii (G ABRYŚ, 1997) (6-20 b, d)

1997 Platytrombidium rafalskii G ABRYŚ, Genus (Wrocław) 8 (1): 41.
D e s c r i p t i o n : G ABRYŚ (1997) Adult: Medium in size. Lateral surface of palp tibia with a
specific basidont, short (27-32 lm), stout, apically blunt and situated in the middle between
palp tarsus and odontus bases. Distal ctenidium of medial surface of palp tibia composed of
a stout paradont and four to seven strong spinisetae, proximal ctenidium consisting of five
to six considerably weaker and thinner spinisetae; radula composed of six to 11 relatively
short spine-like setae. Apex of palp tarsus with five, and lateral side with one solenidion.
Posterior idiosoma setae of one type and variable length, based on distinct papillae, very
short, the stem never exceeding 20 lm, irregularly fusiform, some rounded and some very
slightly and broadly pointed apically, somewhat narrower at base, all covered with numer-
ous, short setulae. Tarsus I oval-rounded, length/width ratio always less than 1.75 (1.48-
1.73). Deutonymph and Larva unknown.
204 Special part
H a bit at a nd B io lo gy : Hygrophilous (G ABRYŚ 1997).

3. Atractothrombium sylvaticum (C. L. K OCH , 1835) (6-11 h; 6-16 d; 6-17 c, e;

6-19 i, 6-20 a)
1835 Trombidium sylvaticum C. L. K OCH, Deutschlands Crustaceen, Myriapoden und Arachni-
den, Fasc. 1, No. 2.
1910 Microtrombidium (Enemothrombium) simulans B ERLESE, Redia 6: 358.
D e s c r i p t i o n : R OBAUX (1966a), G ABRYŚ (1996), G ABRYŚ et al. (2005). Adult: Distal cte-
nidium of palp tibia composed of four to seven strong spinisetae (+ paradont), the proximal
one of seven to nine thinner and more slender spinisetae. Lateral surface of palp tibia with
long (60-90 lm) basidont at the base of palp tarsus. Apex of palp tarsus with five solenidia,
lateral side with one. Posterior idiosoma setae short (c. 25 lm), regularly fusiform, narrow-
ing basally and distally, pointed apically, covered with delicate setulae. Legs much shorter
than idiosoma. Tarsus I oval-elongate (290-385 lm long, 130-190 lm wide); length/width
ratio always more than 2:1; tibia I short (200-280 lm). Deutonymph: Medial surface of
palp tibia has only one ctenidium of four spinisetae (+ paradont). Tarsus I of same propor-
tions as in adult (205-230 lm long, 100-110 lm wide); tibia I 220-240 lm long. Larva: See
generic diagnosis.
H a bit at a nd B io log y : Uni- to semivoltine with adults as diapausing instars. Spermato-
phore deposition and oviposition in spring. Postlarval active instars feed on dipteran eggs,
larvae parasitize nematoceran and brachyceran Diptera in July-September. Restricted to
amphibious biotopes, often the most abundant species of Trombidioidea in inundation
areas and salt marshes.
fications).
Genus Camerotrombidium T HOR , 1936

1877 Ottonia K RAMER, Arch. Naturg. 43 (1): 227, erroneously placed in Microtrombidium by
T HOR & W ILLMANN (1947: 358) [with wrong page „233“].
1910 Microtrombidium (Enemothrombium) B ERLESE, Redia 6: 358 [part.].
1936 Camerotrombidium T HOR, Zool. Anz. 114 (1-2): 31 [part.].
1952 Oicothrombium F EIDER, Bul. Şti. Sect. Sti. Biol. Agr. Geol. Geogr. Acad. Repub. Pop. Rom.
4(3): 621.
Typus generis: Trombidium pexatum C. L. K OCH , 1837, by original designation.
Di ag no sis: Adult: Medium sized, length about 1200 lm, width about 850 lm. Medial
surface of palp tibia with two ctenidia of spinisetae and radula. Distal ctenidium consisting
of much stronger setae than proximal one. Radula composed of very strong and long spine-
like setae. Lateral surface of palp tibia with at least one basidont. Aspidosoma triangular-
oval in outline. Crista metopica well developed with anterior and posterior processes and
distinct accessory posterior process. Double eyes sessile, or with tendency to form a very
short immobile stalk based on a distinct ocular plate, situated at sides of crista metopica,
more or less at the level of mid-point of anterior process. Posterior dorsal setae uniform,
relatively short, sometimes differing in size but always similar in structure: based on distinct
papillae (at least 1/3 of the whole seta length), spherical, spheroidal, oval or calyciform and
elongate, with only one, open, empty chamber inside, covered with numerous and delicate
setulae; sometimes the delicate membrane present, but never in equatorial plane (6-19 k).
Papillae very long and stout, occasionally even longer than setae stems. Distal part of pa-
pillae with or without pointed denticles. Legs without lamellar processes, much shorter
than idiosoma. Tarsus I dilated, longer than wide, longer than tibia I. Deutonymph: Medial
surface of palp tibia with one ctenidium. Number of spine-like setae forming radula re-
duced. Lateral surface of palp tibia at least with one basidont. Idiosoma setae similar
to those in adults. Two pairs of genital acetabula. Larva: Gnathosoma with stephanostome
and internal horseshoe-like sclerite bearing 35-40 prominent lateral teeth, palp with minute
setae on femur and genu. Tritorostral (bs) setae with finger-like projections. Dorsum of
idiosoma with stolascutum bearing three pairs of peripheral setae plus two trichobothria.
Scutellum with two c1 setae. Plates of d1 enlarged and parallel-sided at their inner margins
(6-17 b). Posterolateral dorsal setae much smaller than the median ones. Tarsus III with
scopa, smilum, smooth trifid outer claw and a prominent setulose lophotrix (6-17 b). Ca-
merotrombidium larvae differ from other microtrombidiids in the irregular ornamentation
of the scutellum (visible only in some embedding media), the special form of d1-sclerites (in
unfed specimens one might get the impression of fused d1-sclerites), and the smaller size of
postero-lateral body setae in comparison with medial setae.
Key to species
Ex clude d: Camerotrombidium fusiforme W ILLMANN , 1956 (insufficient description based on a sin-
gle deutonymph from Czech Republic); Camerotrombidium sigthori T HOR & W ILLMANN , 1947
(nomen novum for Ottonia purpurea T HOR, 1900 non Trombidium purpureum C. L. K OCH ,
1837, known only from postlarval instars from Germany and Norway, description insufficient).
Larvae
1 Solenidion (r) on genu III longer than half the length of accompanying normal setae (6-20 e)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camerotrombidium rasum* (page 206)
– Solenidion (r) on genu III about half the length of accompanying normal setae (6-20 f)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camerotrombidium pexatum* (page 205)

1 Palp tibia with three to five stout basidonts (adults; in deutonymphs one basidont or basidont
absent) (6-20 g), idiosoma setae strongly elongate (6-20 h)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camerotrombidium pexatum* (page 205)
– Palp tibia with only one basidont (adult and deutonymph), posterior idiosoma setae spherical
................................................................. 2
2 Distal part of papillae with denticles (6-20 i) . . . . . . Camerotrombidium rasum* (page 206)
– Distal part of papillae without denticles (6-20 j). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camerotrombidium sanguineum* (page 206)
1. Camerotrombidium pexatum (C. L. K OCH , 1837) (6-17 b; 6-19 k; 6-20 f-h)

1837 Trombidium pexatum C. L. K OCH, Deutschlands Crustaceen, Myriapoden und Arachniden,
Fasc. 15, No. 11.
1909a Ottonia conifera G EORGE, Naturalist (London) 628: 194.
1910 Microtrombidium (Enemothrombium) calicigerum B ERLESE, Redia 6: 359.
D e s c r i p t i o n : G ABRYŚ (1996) (adult), W OHLTMANN et al. (2003) (all active instars).
Adult: Medial surface of palp tibia with two ctenidia. Distal ctenidium composed of
six to seven spinisetae (+ paradont), proximal one of nine to 12 more delicate spinisetae.
Lateral surface of palp tibia with three to five strong basidonts, situated at palp tarsus base.
Dorsal posterior idiosoma setae variable in length (20–55 lm, usually 25–40 lm excluding
papillae); most of them elongate, funnelform or calyciform, with straight or rounded sides,
with chamber inside, covered densely with numerous, small setulae; apical part in some
206 Special part
setae with a membraneous structure which traverses the chamber. Setal stems based on
stout papillae, the length of which is about 1/3 – 1/4 length of entire seta; the distal parts
of papillae with numerous protruding pointed denticles which make the cross sections of
papillae look star-like. Deutonymph: Medial surface of palp tibia with one ctenidium, la-
teral surface with or without one basidont. Larva: differing from C. rasum larvae only in
the relatively short solenidion on genu III.
H a bit at a nd B io lo gy : Uni- to semivoltine. Postlarval instars feed on insect eggs, larvae
parasitize brachyceran Diptera in June to August. Abundant in proximity of waters in
forested areas.
fications).
2. Camerotrombidium rasum (B ERLESE , 1910) (6-19 k; 6-20 e, i)

1910 Microtrombidium (Enemothrombium) rasum B ERLESE, Redia 6: 360.
D e s c r i p t i o n : R OBAUX (1967), G ABRYŚ (1996) (postlarval instars), W OHLTMANN et al.
(2003) (all active instars). Adult: Medial surface of palp tibia with two ctenidia. Distal
ctenidium composed of six to seven spinisetae (+ paradont), proximal one of four to
nine more delicate spinisetae. Lateral surface of palp tibia with one long narrow basidont,
situated at palp tarsus base. Dorsal posterior idiosoma setae various in length (16–47 lm),
usually 18–34 lm (excluding papillae); most of them spherical or spheroidal with chamber
inside, covered densely with numerous, small setulae all over; some setae slightly elongate,
with all kinds of transitional forms between spherical and elongate; membraneous struc-
ture which traverses the chamber absent. All setal stems based on stout papillae, the length
of which is about 1/2 length of entire seta; the distal parts of papillae with numerous pro-
truding pointed denticles which make the cross sections of papillae look star-like. Deuto-
nymph: Medial surface of palp tibia with one ctenidium composed of five to eight spinisetae
(+ paradont). Lateral surface of palp tibia with one basidont. Larva: differing from C. pex-
atum larvae only in the relatively long solenidion on genu III.
H a bit at a nd B io lo gy : Uni- to semivoltine. Postlarval instars feed on insect eggs, larvae
parasitize brachyceran Diptera in June to August. Abundant on temporarily flooded mea-
dows.
fications).
3. Camerotrombidium sanguineum (C. L. K OCH , 1837) (6-20 j)

1837 Trombidium sanguineum C. L. K OCH, Deutschlands Crustaceen, Myriapoden und Arachni-
1910 Microtrombidium (Enemothrombium) subrasum B ERLESE, Redia 6: 362.
1914 Ottonia ignota G EORGE, Naturalist (London): 306.
D e s c r i p t i o n : R OBAUX (1967a) (for deutonymph and adult, metric data based on Spanish
specimens only). Adult: Idiosoma length 1230-1345 lm, width 865-960 lm. Palps long and
slender. Medial surface of palp tibia with two ctenidia; distal ctenidium composed of six
spinisetae (+ paradont), proximal one consists of six spinisetae. Radula formed of eight to
ten longer and slender spine-like setae. Lateral surface of palp tibia with only one basidont.
Palp tarsus with five, rarely six, solenidia apically. Anterior part of crista metopica broad,
sensillary area with two smooth sensillary setae, vertex with about ten long, barbed setae,
posterior process delicate and narrow. Double eyes based on very short stalks situated at the
level of anterior part of crista metopica; anterior lenses somewhat bigger than posterior
ones. Posterior idiosoma setae longer than aspidosomal ones (31-46 lm and 18-37 lm
respectively, excluding papillae). In general dorsal setae spherical, covered with minute se-
tules all over the surface except for their distal part. All setae placed on distinct papillae of
the same length as setae. Distal part of papillae in form of a circular rim without denticles.
All legs much shorter than idiosoma. Tarsus I oval-elongate (315-325 lm long, 145-155 lm
wide), tibia I short (220-240 lm). Length/width ratio of tarsus I: 2.1 to 2.2. Deutonymph:
Generally smaller. Idiosoma length 480 lm, width 430 lm. Medial surface of palp tibia
with only one ctenidium consisting of four spinisetae (+ paradont), radula composed of
three spine-like setae. Lateral surface with one basidont. Palp tarsus with three solenidia
terminally. Dorsal setae similar to those in adult but shorter (12-28 lm without papillae).
Tarsus I 160 lm long, 85 lm wide. Tibia I 100 lm long. Length/width ratio of tarsus I: 1.9.
Larva unknown.
H a bit at a nd B io lo gy : Hygrophilous (R OBAUX 1967).
Di stri bu ti on: Germany, Italy, Spain.
Genus Campylothrombium K RAUSSE , 1916

1916b Microtrombidium (Campylothrombium) K RAUSSE, Zool. Anz. 47(4): 98.
Typus generis: Microtrombidium (Campylothrombium) langhofferi K RAUSSE , 1916a, subse-
quent designation by V ITZTHUM (1929: 65).
Di ag no sis: Adult: Medium to large, length up to 2000 lm, width up to 1600 lm. Palps
strong, with prominent odontus and paradont. Medial surface of palp tibia with two over-
lapping ctenidia of spinisetae and a radula. Distal ctenidium consisting of stronger spini-
setae. Lateral surface of palp tibia with at least one long and strong basidont. Aspidosoma
triangular in outline. Crista metopica typical, consisting of anterior and posterior processes
and relatively small, oval sensillary area bearing two filiform sensillary setae. Double eyes
placed on ocular plates, sessile, or with tendency to form a very short immobile stalk, si-
tuated at sides of crista metopica, more or less at the level of mid-point of anterior process.
Posterior dorsal setae uniform in size and structure, clavate, two-chambered, straight or
slightly bent, with thick stem covered with relatively scarce, long setulae (6-19 l). Stem
dilated distally forming a closed chamber; second chamber, narrow and elongate, separated
from the first one, filling the remaining (proximal) part of the stem. Setulae covering the
stem sometimes substituted with small tubercles situated laterally. Legs much shorter than
idiosoma, with clavate setae (6-19 h) on all articles. Tarsus I dilated, elongate, rounded
distally, longer than wide, longer than tibia I. Deutonymph: Medial surface of palp tibia
with one ctenidium. Number of spine-like setae forming radula reduced. Lateral surface of
palp tibia with at least one basidont. Idiosoma setae similar to those in adults. Two pairs of
genital acetabula. Larva: Gnathosoma with stephanostome and internal horseshoe-like
sclerite bearing about 30 prominent lateral teeth, palp with minute setae on femur and
genu. Tritorostral (bs) setae with finger-like projections (6-16 f). Dorsum of idiosoma
with stolascutum bearing three pairs of peripheral setae plus two trichobothria. Scutellum
with two c1 setae. Plates of d1 enlarged, ovoid. Other dorsal setae on much smaller platelets.
Tarsus III with scopa, smilum, smooth trifid outer claw and a prominent setulose lophotrix.
Only one species is known from Central Europe.
1. Campylothrombium clavatum (G EORGE , 1909c) (6-16 f; 6-17 d; 6-19 h, l)

1909c Ottonia clavata G EORGE, Naturalist (London) 635: 423.
1910 Microtrombidium (Enemothrombium) densipapillum B ERLESE, Redia 6: 360.
1910 Microtrombidium (Enemothrombium) densipapillum var. boreale B ERLESE, Redia 6: 360.
208 Special part
1916a Microtrombidium (Campylothrombium) langhofferi K RAUSSE, Zool. Anz. 47 (4): 97.

1924 Microtrombidium (Enemothrombium) berlesei A NDRÉ, Bull. Mus. Hist. nat. Paris 30 (4): 274.
1925 Microtrombidium (Campylothrombium) antonii A NDRÉ, Bull. Mus. Hist. nat. Paris 31: 87.
1932 Microthrombidium (Enemothrombium) kervillei A NDRÉ, Extr. Bull. Soc. Zool. Franc. 57:
190.
1932 Microthrombidium (Enemothrombium) kervillei var. diversipalpis A NDRÉ, Extr. Bull. Soc.
Zool. Franc. 57: 193.
1955a Campylothrombium metlaglianum [sic!] F EIDER, Fauna R. P. R. 5 (1): 128.
1955a Campylothrombium dobrogiacum F EIDER, Fauna R. P. R. 5 (1): 128.
De sc ri pti on : Redescriptions published by R OBAUX (1974) (larvae), R OBAUX (1966b)
and G ABRYŚ (1996, 1999) (postlarval instars).
H a bit at a nd B i ol og y: Uni- to semivoltine with obligate egg diapause. Larvae abundant
in May, parasitize brachyceran Diptera. Present in woodland soils, also in the area of tem-
porary waters.
Genus Echinothrombium W OMERSLEY, 1937

1937 Echinothrombium W OMERSLEY, Rec. S. Aust. Mus. 6 (1): 89 [part.].
1950b Oxithrombium F EIDER, Acad. Repub. Pop. Rom., Filiala Iaşi, Stud. cerc. Şti. 1(2): 846 [part.].
Typus generis: Ottonia spinosa C ANESTRINI , 1885 [orig. „Ottonia spinosum C ANESTRINI ,
1877“], by original designation.
Di ag no sis: Adult: Body length 1000–1600 lm, width 450–950 lm. Medial surface of
palp tibia with two ctenidia and a radula. Distal ctenidium consists of much stronger spi-
nisetae than proximal one. Lateral surface with one weak, slender basidont. Aspidosoma
triangular in outline. Crista metopica typical with anterior and posterior processes and
circular sensillary area bearing two filiform sensillary setae. Accessory posterior process
present but not distinct. Double eyes sessile, placed at sides of crista metopica at the level
of mid-point of anterior process. Posterior dorsal setae of two types (6-18 g; 6-20 k, l). The
longer setae bare, except for the base which is delicately setulose, flattened, blade-like or
reminding of a sword (ensiform), pointed apically, smooth or with a few short distal pro-
cesses. The shorter setae setiform, short, thin, with strong setulae covering all the stem
except for the extremely apical part. Legs without lamellae, much shorter than idiosoma.
Tarsus I dilated, longer than tibia I. Deutonymph: Medial surface of palp tibia with one
ctenidium. Number of spine-like setae forming radula reduced. Lateral surface of palp tibia
with one basidont. Idiosoma setae similar to that in adults. Two pairs of genital acetabula.
Larva: Gnathosoma with stephanostome and inconspicuous internal horseshoe-like sclerite
without lateral teeth (6-17 a). Sclerite ensheathed by a cuticular membrane, which is dor-
sally extended. Palp with minute setae on femur and genu. Dorsum of idiosoma with sto-
lascutum bearing three pairs of peripheral setae plus two trichobothria. Scutellum with two
c1 setae. Remaining dorsal setae on plates or platelets, c2 and d1 plates enlarged (6-17 a).
Double eyes present, anterior lens prominent, about twice as wide as posterior one. Coxa
III directed posteriorly, tibia III distally enlarged, tarsus III with scopa, smilum distally bi-
fid, smooth trifid and ciliated outer claw and setulose, comparatively small lophotrix with
four branches in its proximal half.
Key to species
Larvae
Only species described in the larval instar: Echinothrombium rhodinum* (page 209)

1 Longer setae on posterior dorsal idiosoma provided with small spines which distally form a
crown-like pattern (6-20 k) . . . . . . . . . . . . . . . . Echinothrombium rhodinum* (page 209)
– Longer setae on dorsal idiosoma distally smooth (6-20 l) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinothrombium spinosum (C ANESTRINI , 1885)
1. Echinothrombium rhodinum (C. L. K OCH , 1837) (6-17 a; 6-18 g; 6-20 k)

1837 Trombidium rhodinum C. L. K OCH, Deutschlands Crustaceen, Myriapoden und Arachni-
1922 Microtrombidium berlesei S CHWEIZER, Verhdl. Naturf. Ges. Basel 33: 90.
1950 Echinothrombium rhodinum var. schweizeri A NDRÉ, Bull. Mus. Hist. nat. Paris, 2e sér. 22
(5): 575.
D e s c r i p t i o n : G ABRYŚ (1996) (adult), G ABRYŚ & W OHLTMANN (2003) (larva and postlar-
val instars). Adult: Distal ctenidium of palp tibia with three to six (+ paradont) strong spi-
nisetae; proximal ctenidium placed immediately behind the distal one with four to seven
thinner and weaker spinisetae. Lateral surface of palp tibia with long and strong basidont
placed close to the base of palp tarsus. Terminal part of palp tarsus with four to five so-
lenidia, lateral side with one. Dorsal posterior idiosoma setae of two types. The longer ones
(50-110 lm) characteristic: straight or curved, rigid, only slightly setulose at base (maxi-
mum up to a third of total length), the remaining part bare, slightly bent distally and
pointed apically; at some distance from the apex two short pointed processes directed op-
posite to the curvature of the seta stem, always shorter than the distance between their bases
and apex of the seta; surface between the processes and the seta stem covered with short
(more or less of the processes’ length) very delicate setulae. Shorter setae (25-40 lm)
straight, setulose all over, similar to setae of Microtrombidium with slightly bent apex.
Generally, longer setae lower in number (about one per 10 short ones) and placed mainly
in posterior part of idiosoma. Legs shorter than idiosoma; tarsus I oval-elongate, about
twice as long as wide (210-280 lm long, 130-160 lm wide); tibia I short (125-175
lm). Deutonymph: Medial surface of palp tibia with only one ctenidium. Lateral surface
of palp tibia with one basidont. Dorsal posterior idiosoma setae similar to those in adult but
slightly shorter. Legs shorter than in adult. Larva: See generic diagnosis.
H a bit at a nd B i olo gy : Regularly present in inundation areas. Life cycle probably uni- to
semivoltine, controlled by obligatory egg diapause (G ABRYŚ & W OHLTMANN 2003).
Genus Enemothrombium B ERLESE , 1910

1947 Eutrichothrombium T HOR & W ILLMANN, Das Tierreich 71a: 385 [part.].
1952 Mastothrombium F EIDER, Bul. Şti. Sect. Sti. Biol. Agr. Geol. Geogr. Acad. Repub. Pop. Rom.
4 (3): 618 [part.].
1974 Feiderium (Parafeiderium) V ERCAMMEN -G RANDJEAN & C OCHRANE [on base of ‘Feiderium’
as nomen nudum], J. Kansas Entomol. Soc. 47 (1): 67.
1994 Parafeiderium sensu S OUTHCOTT, Zoologica, Stuttgart 144: 91.
Typus generis: Trombidium bifoliosum C ANESTRINI , 1884 [orig. „T. E. sanguineum K.“,
changed by B ERLESE (1912: 174) on the ground of misidentification for „M. E. bifoliosum
(C AN .)“].
Di ag no sis: Adult: Medial surface of palp tibia with only one, relatively long ctenidium,
consisting of at least ten spinisetae. Lateral surface of palp tibia without basidont (6-11 i).
210 Special part
Fig. 6-20: Microtrombidiidae adults and larvae: a, Atractothrombium sylvaticum, detail adult: body
setae (after G ABRYŚ 1996); b, Atractothrombium rafalskii, detail adult: body setae (after G ABRYŚ
1997); c, Atractothrombium fusicomum, detail adult: palp (after G ABRYŚ 1996); d, Atractothrombium
rafalskii, detail adult: palp (after G ABRYŚ 1997); e, i, Camerotrombidium rasum; e, detail larva: genu
leg III (after W OHLTMANN et al. 2003); i, detail adult: body setae; f-h, Camerotrombidium pexatum; f,
detail larva: genu leg III (after W OHLTMANN et al. 2003); g, detail adult: palp (after G ABRYŚ 1996); h,
detail adult: body setae (after G ABRYŚ 1996); i, Camerotrombidium rasum, detail adult: body setae
(after G ABRYŚ 1996); j, Camerotrombidium sanguineum, detail adult: body setae (after R OBAUX
1967); k, Echinothrombium rhodinum, detail adult: body setae (after G ABRYŚ 1996); l, Echinothrom-
bium spinosum, detail adult: body setae (after G ABRYŚ 1996); m, Valgothrombium alpinum, detail
deutonymph: body setae (after G ABRYŚ 1996); n, Valgothrombium confusum, detail deutonymph:
body setae (after G ABRYŚ 1996); o, Valgothrombium major, detail adult: body setae (after G ABRYŚ
1996); p, Valgothrombium valgum, detail adult: body setae (after G ABRYŚ 1996); (scale bars = 50 lm).
Aspidosoma triangular-oval in outline (6-11 k). Posterior process of crista metopica rudi-
mentary or absent. Sensillary area round, bearing two filiform, short sensillary setae. Dou-
ble eyes sessile, situated at sides of crista metopica at the level of mid-point of anterior
process. Anterior lenses slightly larger than posterior ones, both placed on distinct ocular
plates. Posterior dorsal setae of two types (6-18 a): First type short or elongate, spherical or
clavate, more or less incised apically, with irregular surface covered with small but distinct
tubercles. Second type usually longer (in some cases very slim and elongate) than the former
ones, club-shaped, without typical spherical forms, without tubercles but with deeper in-
cision in apical part, and numerous minute setulae, arranged in several longitudinal rows
extending over the whole seta. Legs with very delicate lamellae, not sharply pointed, rather
rounded distally. All legs much shorter than idiosoma. Tarsus I dilated, elliptical, longer
than tibia I. Deutonymph: Medial surface of palp tibia with one ctenidium consisting of six
to eight spinisetae. Number of spine-like setae forming radula reduced. Lateral surface of
palp tibia without basidont. Idiosoma setae similar to that in adults. Two pairs of genital
acetabula. Larva: Scutellum provided with a median ridge (6-16 b). Scutum not bent lat-
erally. Posterior dorsal setae serrate, arranged in rows c2-3, d1-3, e1-3, f1-3, h1-2 on plate-like
sclerites, the median d, e, f plates much larger than the lateral ones. Tarsus I and II without
empodium, tarsus III with empodium and two claws, one of them modified into a smilum;
lophotrix and scobala present. Palp femur and genu without setae. Tritorostral setae mi-
nute, spine-like.
Only one species described for the area:
1. Enemothrombium bifoliosum (C ANESTRINI , 1844) (6-11 i, k; 6-16 b; 6-18 a)

1884 Trombidium bifoliosum C ANESTRINI, Atti. Ist. Veneto Sci. Lett. Art. 2 ser. 6: 693.
1951 Enemothrombium folliculum S CHWEIZER, Ergebn. Wiss. Unters. Schweiz. Nat. Parks (n.f.)
III 23 (2): 109.
1955a Mastothrombium oltenicum F EIDER, Fauna R. P. R. 5 (1): 121.
1955 Eutrichothrombium itale C OOREMAN, Bull. Inst. r. Sci. nat. Belg. 31 (48): 7.
D e s c r i p t i o n : R OBAUX ( 1967 – as ‘Mastothrombium oltenicum’) and G ABRYŚ (1996)
(postlarval instars); W OHLTMANN & G ABRYŚ (in press) (larva and postlarval instars).
See diagnosis of the genus.
H a bit at a nd B i ol ogy : Abundant in inundation areas. Larvae display preparasitic atten-
dance on the pupae of their dipteran hosts, abundant in June-July. Several species of Ce-
ratopogonidae have been found as hosts of larvae of the North American E. culicoidium
(V ERCAMMEN -G RANDJEAN & C OCHRANE , 1974); a comparable host range is assumed for
E. bifoliosum.
Genus Microtrombidium H ALLER , 1882

1882 Microtrombidium H ALLER, Jh. Ver. vaterl. Naturk. Württemb. 38: 322.
1955b Elahistothrombium F EIDER, An. Ştiinţ. Univ. „Al. I. Cuza“ Iaşi (ser. nouă), secţ. II (Şti. Nat.) 1
(1-2): 91.
1955b Physetothrombium F EIDER, An. Ştiinţ. Univ. „Al. I. Cuza“ Iaşi (ser. nouă), secţ. II (Şti. Nat.) 1
(1-2): 96.
1978 Megophthrombium M ULLEN & V ERCAMMEN -G RANDJEAN, Internat. J. Acarol. 4 (1): 39.
1993 Phlebotrombium FAIN & B AKER, Bull. Annls Soc. r. belge Ent. 129: 327.
Typus generis: Trombidium pusillum H ERMANN , 1804 (= Microtrombidium purpureum
H ALLER , 1882 non C. L. K OCH , 1837), subsequent designation by B ERLESE (1912: 124).
212 Special part
Di ag no sis: Adult: Small to medium sized, body length 750-1350 lm, width 475-900 lm.
Medial surface of palp tibia with two ctenidia and a radula (6-19 c). Lateral surface with
one slender basidont. Palp tarsus with one solenidion distally. Aspidosoma triangular in
outline. Crista metopica typical, with anterior and posterior processes and circular or
oval sensillary area, bearing two filiform sensillary setae (6-19 b). Double eyes sessile
or with tendency to form a very short immobile stalk, situated at sides of crista metopica,
more or less halfway anterior process. Posterior dorsal setae setiform (6-19 g), always of
one type and more or less uniform in size, straight, thin, pointed, covered with setulae of
various types (delicate or strong). Legs without lamellar processes, always shorter than
idiosoma and with dilated tarsi. Tarsus I broad, swollen, more or less oval, longer than
wide and much longer than tibia I. Length/width ratio of tarsus I 5 1.5. Deutonymph:
Medial surface of palp tibia with one ctenidium. Number of spine-like setae forming radula
reduced. Idiosoma setae similar to that in adults. Two pairs of genital acetabula. Length/
width ratio of tarsus I 5 1.5. Larva: Gnathosoma with stephanostome and internal horse-
shoe-like sclerite without lateral teeth (6-3 d). Palp tarsus with a prominent solenidion
(length about 10 lm), no setae on palp femur and genu. Dorsum of idiosoma with stolas-
cutum with three pairs of peripheral setae and two trichobothria, and scutellum with two c1
setae. Remaining dorsal setae on plates or platelets, c2 and d1 plates enlarged (6-3 a). Dou-
ble eyes present, anterior lens prominent, about twice as wide as posterior one. Coxa III
directed posteriorly (6-3 a), tibia III enlarged distally, tarsus III with scopa, smilum bifid
distally, outer claw ciliated, lophotrix bifid distally (6-16 a) with both branches covered
with small setules.
R em a rks : Only one species from Central Europe, M. pusillum, is adequately described
(see below). Thirteen further species recorded from this area need revision before proper
identification is possible. These are: Microtrombidium basiliense S CHWEIZER & B ADER ,
1963 (Switzerland), Microtrombidium bispinum S CHWEIZER & B ADER , 1963 (Switzer-
land), Microtrombidium botschi S CHWEIZER & B ADER , 1963 (Switzerland), Microtrom-
bidium ferociforme (T RÄGÅRDH , 1904) (West Africa, Switzerland ?, Poland ?), Microtrom-
bidium medium S CHWEIZER & B ADER , 1963 (Switzerland), Microtrombidium mihelcici
Z HANG , 1992 (Austria), Microtrombidium minor S CHWEIZER , 1922 (Switzerland), Micro-
trombidium parvulum S CHWEIZER , 1922 (Switzerland), Microtrombidium plurispinum
S CHWEIZER , 1951 (Switzerland), Microtrombidium raeticum S CHWEIZER & B ADER ,
1963 (Switzerland), Microtrombidium spiniferum (T HOR , 1900) (Norway, Austria), Mi-
crotrombidium subferociforme S CHWEIZER & B ADER , 1963 (Switzerland), Microtrombi-
dium tenuipalpe S CHWEIZER & B ADER , 1963 (Switzerland). For further discussion see
G ABRYŚ (1999).
1. Microtrombidium pusillum (H ERMANN , 1804) (6-3; 6-10 a, 6-16 a; 6-19 b-c, g)

1804 Trombidium pusillum H ERMANN, Mémoire aptérologique: 27.
1835 Trombidium puniceum C. L. K OCH, Deutschlands Crustaceen, Myriapoden und Arachniden,
Fasc. 1. No. 1.
1882 Microtrombidium purpureum: H ALLER, Jh. Ver. vaterl. Naturk. Württemb. 38: 322 [treated
properly by B ERLESE (1912: 135) as misidentified Microtrombidium pusillum (H ERMANN ,
1804) non Trombidium purpureum C. L. K OCH ].
1912 Microtrombidium platychirum B ERLESE, Redia 8: 162 [non Microtrombidium platychirum
R OBAUX , 1967].
1914 Microthrombidium parvum O UDEMANS, Arch. Naturg., Abt. 79A 9: 129.
1967 Microthrombidium marcandrei R OBAUX, Mém. Mus. Nat. Hist. Nat. Paris, (n.s.), Sér. A,
Zool. 46 (1): 49.
D e s c r i p t i o n : G ABRYŚ & W OHLTMANN (2001) (postlarval and larval instars), see diagno-
sis of the genus.
H a bit at an d B io lo gy : Uni- to semivoltine, adults seem to diapause before reproduction.
Postlarval active instars feed on dipteran larvae. Larvae parasitize imagines of Diptera in
July. Often found in the area of temporary flooded soils.
Genus Valgothrombium W ILLMANN , 1940

1940 Valgothrombium W ILLMANN, Zool. Anz. 131 (9-10): 250.
1994 Feiderella S OUTHCOTT, Zoologica, Stuttgart 144: 77.
1999 Parafeiderium sensu B AKER, Syst. Parasitol. 44: 41 [part.].
Typus generis: Ottonia valga G EORGE , 1909c, by original designation.
Di ag no sis: Adult: Small to medium sized; length 1000-2400 lm, width 400-1200 lm.
Palps with slender and elongate palp tibia (6-19 a). Medial surface of palp tibia with only
one ctenidium. Typical radula of long, smooth spine-like setae absent; instead a few weak,
sometimes setulose setae. Lateral surface of palp tibia without basidont. Palp tarsus short,
conical. Aspidosoma triangular or slightly rounded in outline. Apex of anterior part of
crista metopica with a characteristic knob-like, striated „nose“, always with two processes
curved in posterior direction (6-19 b). Posterior process absent or rudimentary, of variable
shape and size, without connection to sensillary area; the latter with two smooth sensillary
setae. Double eyes sessile, situated at the level of posterior part of crista metopica, usually
very close to sensillary area. Ocular plates and lenses relatively large. Posterior dorsal setae
uniform within a species (6-19 f), in different species different types of setae occur: straight
and pointed, fusiform, clavate, delicately setulose, often incised apically (6-20 m-p). Legs
without lamellae. Tarsus I dilated, longer than wide and longer than tibia I. Deutonymph:
Medial surface of palp tibia with one ctenidium. Number of weak, sometimes setulose setae
reduced. Lateral surface of palp tibia without basidont. Idiosoma setae similar to that in
adults. Larva: With scutum and scutellum, both provided with a median ridge (6-17 g).
Scutum not bent laterally. Posterior dorsal setae serrate, arranged in rows c2-3, d1-3,
e1-3, f1-3, h1-2, on plate-like sclerites equal in size. Posterior solenidion of genu I and sole-
nidion of genu II located in a notch of anterior margin of trochanters I and II, respectively.
Tarsus I and II without empodium, tarsus III with two claws and empodium, lophotrix and
scobala absent (6-17 g). Palp femur and genu without setae. Gnathosomal setae minute,
spine-like.
Key to species
Ex clude d: Valgothrombium dubiosum S CHWEIZER, 1951 (Switzerland, characters given in original
description allow to state that it is a deutonymph; description not sufficient for proper interpretation);
Valgothrombium moravicum (W ILLMANN , 1954) (Czech Republic, described from a single female as
subspecies of the Mexican species Platytrombidium homocomum (B ERLESE, 1918b), description in-
sufficient for proper interpretation). For further discussion see G ABRYŚ (1999, 2002).
Larvae
1 Total length of legs (including coxa, excluding claws): leg I 4 210 lm, II 4 190 lm, III 4 210 lm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium stuarti* (page 216)
– Total length of legs (including coxa, excluding claws): leg I 5 200 lm, II 5 180 lm, III 5 200 lm,
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium valgum* (page 216)
214 Special part

1 Posterior dorsal idiosoma setae (pDS) incised apically (6-20 m) . . . . . . . . . . . . . . . . . . . . 2
– pDS pointed apically (6-20 p). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 pDS densely covered with long setules (6-20 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– pDS covered with very delicate short setules (6-20 n) Valgothrombium confusum (page 214)
3 pDS cylindrical, broadest part at 1/3 length from base (excl. papillae) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium longipes (page 215)
– pDS clavate, broadest part at 2/3 length from base (excl. papillae) (6-20 m) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valgothrombium alpinum (page 214)
4 pDS slender, thin, conical, broadest at base (6-20 o) . . . . Valgothrombium major (page 215)
– pDS setae fusiform, broadest at half length (6-20 p) . . . Valgothrombium valgum (page 216)
1. Valgothrombium alpinum W ILLMANN , 1940 (6-20 m)

1940 Valgothrombium alpinum W ILLMANN, Zool. Anz. 131 (9-10): 253
D e s c r i p t i o n : W ILLMANN (1940) (adult?), G ABRYŚ (1996) (deutonymph). Adult: Idioso-
ma length about 2400 lm. Anterior part of crista metopica broad with longitudinal cavity
running along its axis and reaching sensillary area; apex of crista metopica forming a short,
well sclerotized „nose“ extending only slightly behind the vertex; setae (about twelve) on
and along crista metopica (parallel to the longitudinal cavity) arranged at regular intervals;
posterior part of crista metopica with triangular-oval sensillary area bearing two long,
smooth sensillary setae; posterior process absent. Double, sessile eyes placed just at the
sensillary area (anterior lenses at the level of posterior end of crista metopica rod, posterior
lenses at the level of sensillary area). Anterior lenses distant from anterior edges of the ocu-
lar plates, posterior lenses close to posterior edges of the plates. Dorsal posterior idiosoma
setae (45-50 lm) based on short papillae, clavate, slightly incised apically, broadest at half
length from the base, covered with delicate, relatively long setules. Deutonymph: Idiosoma
length 1300 lm, width 850 lm. Medial surface of palp tibia with one ctenidium of six
spinisetae (+ paradont); pseudoradula composed of two setae. Palp tarsus with one
very long and smooth seta terminally on lateral side. Crista metopica in anterior part
with a sclerotized „nose“ and two lateral processes curved in posterior direction. Double
eyes slightly farther from crista metopica, posterior lenses at the level of oval sensillary
area. Dorsal posterior idiosoma setae similar to those in adults but slightly shorter (30-
40 lm), broadest at 2/3 from base excluding papillae, all covered with delicate, relatively
long setules; distal part of papillae has several, pointed denticles. Legs much shorter than
idiosoma; tarsus I about 180 lm long, 100 lm wide, tibia I about 120 lm long. Larva
unknown.
H a bit at an d B io lo gy : Confined to amphibious (limnic) biotopes, exclusively in moun-
tains.
Di stri bu ti on: Austria, Poland.
2. Valgothrombium confusum (B ERLESE , 1910) (6-20 n)

1910 Microtrombidium (Enemothrombium) confusum B ERLESE, Redia 6: 362
D e s c r i p t i o n : R OBAUX ( 1967) (adult and deutonymph), G ABRYŚ (1996) (deutonymph).
Adult: Palps long and slender. Medial surface of palp tibia with single ctenidium of six
to eight spinisetae running along dorsal edge from behind the paradont; instead of radula
two long, setulose setae forming pseudoradula. Terminal part of the short palp tarsus with
two smooth specialized setae: one long, the other short; one solenidion on lateral side in
proximal half. Anterior part of crista metopica extending beyond vertex and forming a
short, well sclerotized „nose“ with two lateral processes curved in posterior direction;
the „nose“ without setae; posterior part of crista metopica with triangular-oval sensillary
area bearing two long, smooth sensillary setae; posterior process absent. Double sessile eyes
placed just at sensillary area (anterior lenses at the level of posterior end of crista metopica
rod, posterior lenses at the level of sensillary area); anterior lenses bigger than the posterior
ones. Dorsal posterior idiosoma setae similar in structure and size (30-45 lm), almost cy-
lindrical or clavate, sometimes curved, delicately incised apically, all scarcely covered with
very delicate, short setules. Legs much shorter than idiosoma; tarsus I usually twice as long
as wide (220-240 lm long, 100-120 lm wide); tibia I short (130-170 lm). Deutonymph:
Idiosoma length 350-950 lm, width 310-650 lm. Medial surface of palp tibia with one
ctenidium consisting of two to six spinisetae; pseudoradula reduced to two setae (in a speci-
men from Poland one almost bare, the other completely bare) or absent. Distal part of palp
tarsus laterally with one very long seta, medially with two very short and smooth specia-
lized setae; solenidion on lateral side present. Aspidosoma similar to that in adult, but an-
terior end of crista metopica not reaching the vertex; sensillary area with two symmetrical,
sclerotized, S-shaped appendages laterally. Posterior idiosoma setae almost cylindrical or
clavate, various in shape, occasionally curved, incised apically, all covered with very de-
licate, short setules; all setae on short papillae the distal parts of which have several pointed
denticles. Legs shorter, tarsus I 120-155 lm long, 75-95 lm wide, tibia I short (75-85 lm).
Larva unknown.
H a bit at a nd B io lo gy : Confined to amphibious (limnic) biotopes in forested areas.
Di stri bu ti on: South-West and Central Europe.
3. Valgothrombium longipes F RANKE , 1942

1942 Valgothrombium longipes F RANKE, Zool. Anz. 137 (7/8): 166.
D e s c r i p t i o n : F RANKE (1942) (adult?), G ABRYŚ (1996) (adult). Adult: Medial surface of
palp tibia with single ctenidium of seven to eight spinisetae situated along dorsal edge just
behind paradont; instead of radula two to three long, slightly setulose setae. Palp tarsus
short, in the distal part laterally with one very long and medially with two very short
and smooth specialized setae, and one lateral solenidion in the proximal part. Anterior
part of crista metopica extending beyond the vertex and forming a relatively long
„nose“ covered with conspicuous stripes; anteriorly from the „nose“ some very small setae
similar to idiosomal ones, but much shorter (after F RANKE 1942, not observed in a speci-
men from north-east Poland); posterior part of crista metopica dilated to form a triangular-
oval or oval sensillary area with two long and smooth non-sensillary setae; posterior pro-
cess reduced to a rudimentary crescent-shaped sclerite without connection with the sensil-
lary area of crista metopica. Double, sessile eyes situated just at sensillary area (anterior
lenses at the level of posterior end of crista metopica rod, posterior lenses at the level of
sensillary area); anterior lenses slightly bigger than posterior ones. Posterior idiosoma setae
uniform (25-40 lm), almost cylindrical, broadest at 1/3 from base (excluding papilla), dis-
tinctly incised apically, all densely covered with long setulae; all setae on short papillae the
distal parts of which have several, pointed denticles. Legs slightly shorter than idiosoma but
relatively long; legs I almost as long as idiosoma; tarsus I 155-170 lm long, 110-120 lm
wide, tibia I very short (65-80 lm). Deutonymph and Larva unknown.
Di stri bu ti on: Poland. Locality record from Austria (R OBAUX 1967) incorrect.
4. Valgothrombium major (H ALBERT, 1920) (6-20 o)

1920 Microtrombidium pusillum var. major H ALBERT, Proc. R. Irish Acad. 35B (7): 146.
216 Special part
D e s c r i p t i o n : H ALBERT (1920), R OBAUX (1967) (adult and deutonymph), G ABRYŚ (1996)

(adult). Adult: Body length 1000-2000 lm, width 450-1300 lm. Palps relatively strong.
Medial surface of palp tibia with single ctenidium of five to nine spinisetae located along
the dorsal edge just behind paradont; instead of radula three to four long, setulose, almost
bare or completely bare setae forming a pseudoradula. Distal part of the short palp tarsus
with three to four smooth specialized setae: one long and two (three according to R OBAUX
1967) short; lateral side of proximal part bearing one solenidion. Anterior part of crista
metopica almost reaching the vertex and forming a short, well sclerotized, striated „nose“
with two lateral processes curved in posterior direction; the „nose“ without setae; crista
metopica posteriorly gradually dilating into the oval-triangular or rhomboidal sensillary
area which bears two long, smooth sensillary setae; typical posterior process absent, only a
rudimentary, broad and narrowing sclerite directed posteriorly, without connection with
sensillary area of crista metopica. Double sessile eyes situated close to crista metopica at the
level of its posterior half but anterior to the level of sensillary area; anterior lenses bigger
than posterior ones. Dorsal posterior idiosoma setae uniform (25-40 lm) (35-50 lm ac-
cording to H ALBERT 1920), slender, thin, conical, narrowing apically and more or less
pointed, without incision, all covered with very delicate setulae; all setae on short papillae
the distal parts of which have several pointed denticles. Legs much shorter than idiosoma;
tarsus I usually about twice as long as wide (145-205 lm long, 70-120 lm wide); tibia I
short (95-140 lm). Deutonymph: Generally smaller than adult: Idiosoma length 800 lm,
width 530 lm. Tarsus I 145 lm long, 65 lm wide; tibia I 80 lm long. Larva unknown.
Di stri bu ti on: Europe (accurate localities cannot be given due to numerous misidentifi-
cations, particularly confusion with Valgothrombium valgum).
5. Valgothrombium stuarti (B AKER , 1999)

1999 Parafeiderium stuarti B AKER, Syst. Parasitol. 44: 38.
D e s c r i p t i o n : B AKER (1999). Adult and Deutonymph unknown. Larva: Scutum 75-85 lm
long, 67-81 lm wide, AM setae 38-46 lm, AL setae 40-48 lm, PL setae 63-71 lm long.
Pseudanal setae 32-39 lm long.
H a bit at an d B io lo gy : Parasitic on Ceratopogonidae. Confined to amphibious (limnic)
biotopes.
6. Valgothrombium valgum (G EORGE , 1909) (6-17 g; 6-19 e-f; 6-20 p)

1909c Ottonia valga G EORGE, Naturalist (London) 635: 423.
D e s c r i p t i o n : W OHLTMANN & W ENDT (1996) (larva), G ABRYŚ (1996) (adult). Adult:
Palps relatively strong. Palp tibia slender, medial surface with a single ctenidium of five
to seven short spinisetae dorsally (eight to nine according to T HOR & W ILLMANN
1947); pseudoradula composed of three to four smooth setae. Palp tarsus short and thick,
at the apex with one very long lateral and two very short and smooth specialized setae; one
solenidion laterally in the proximal part. Anterior part of crista metopica reaching almost
the vertex and forming a short, well sclerotized, striated „nose“ with two lateral processes
curved in posterior direction; vertex developed in form of a well sclerotized, oval plate with
three („several“ according to T HOR & W ILLMANN 1947) setulose non-sensillary setae in
shape different from idiosomal ones; posteriorly dilating into an oval sensillary area with
two long and smooth sensillary setae; typical posterior process absent, only a rudimentary,
oval sclerite with no connection to sensillary area of crista metopica. Double sessile eyes
placed slightly anterior of sensillary area, close to crista metopica; lenses more or less of the
same size, posterior lenses at the level of sensillary area. Dorsal posterior idiosoma setae
uniform, short (15-20 lm) (25-30 lm according to H ALBERT 1915), fusiform, pointed api-
cally, dilated at half length, all covered densely with setules. Legs shorter than idiosoma;
tarsus I usually about twice as long as wide (120-160 lm long, 65-90 lm wide); tibia I short
(65-95 lm). Deutonymph not described. Larva: Scutum 54-56 lm long, 55 lm wide, AM
setae 32-35 lm, AL setae 31 lm, PL setae 54 lm long. Pseudanal setae 24 lm long.
H a bit at a nd B i ol og y: Uni- to semivoltine. Confined to amphibious biotopes with re-
gular inundations. Postlarval active instars in the litter layer, preying on dipteran larvae.
Larvae parasitic on Ceratopogonidae with preparasitic attendance on host pupae.
Di stri bu ti on: Europe (accurate localities cannot be given due to numerous misidentifi-
cations, particularly with Valgothrombium major).
Family Tanaupodidae T HOR , 1935

1935 Tanaupodidae T HOR, Zool. Anz. 109: 108.
Di ag no sis: Adult and Deutonymph: Small- to medium-sized. Colouration reddish to
brown. Idiosoma cuticle almost smooth without large sclerotized areas. Posterior dorsal
idiosoma setae simple, spine-like, inserted on prominent sclerites (6-21 d-f). Prodorsal
shield (scutum) with (6-21 e) or without (6-21 c) projecting naso. At level of coxa IV genital
opening, surrounded by paired sclerites of similar shape. Two pairs of sessile eyes laterally
to crista metopica on ocular sclerites. Anterior to genital opening an unpaired sclerite (6-11
a). Palp tibia with one or more prominent spines laterally to the odontus. Larva: Medium-
to large-sized, without scutellum; scutum roughly rectangular without projecting naso.
Posterior idiosoma setae simple (smooth or with few small barbs), inserted on prominent
platelets (6-21 b). Anal pore surrounded by paired sclerites bearing one or two pairs of setae
(6-21 a). Odontus on palp tibia small.
H a bit at a nd B io lo gy : All species included are rather rare, their biology remains almost
unknown. Rhinothrombium is exclusively known from montane areas. Species of Tanau-
podus inhabit lowlands and seem confined to hygric biotopes.
Key to genera
Marked with an asterisk*: genera containing species regularly found in amphibious biotopes
Larvae
1 Anal sclerite with two pairs of setae, „lassenia organ“ present (6-21 a), palp odontus claw-like,
bifid; lateral claws of legs I-III simple . . . . . . . . . . . . . . . . . . . . . . . Lassenia* (page 203)
– Anal sclerite with one pair of setae, „lassenia organ“ absent, palp odontus setiform, simple; la-
teral claws of legs I-III bifurcate, deeply incised (6-21 b) . . . . . Polydiscia M ETHLAGL , 1928
In Central Europe only Polydiscia squamata M ETHLAGL , 1928

1 Scutum without protruding naso (6-21 c), idiosoma cuticle papillate (6-21 d) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanaupodus* (page 203)
– Scutum with naso (6-21 e), idiosoma cuticle striate (6-21 f) Rhinothrombium B ERLESE , 1910
In Central Europe only Rhinothrombium nemoricola (B ERLESE , 1886)
Genus Lassenia N EWELL, 1957 (6-10 c; 6-21 a)

1957 Lassenia N EWELL, Pacif. Sci. 11: 447.
Typus generis: Lassenia lasseni N EWELL, 1957, by original designation.
218 Special part
Fig. 6-21: Tanaupodidae: a, Lassenia lasseni, detail larva: lassenia organ (after N EWELL 1957); b,
Polydiscia squamata, detail larva: claws (after W OHLTMANN 2004); c-d, Tanaupodus passimpilosus,
adult; c, detail: scutum, eyes; d, detail: body setae; e-f, Rhinothrombium nemoricola, adult; e, detail:
scutum, eyes; f, detail: body setae; (scale bars = 50 lm).
D iagnos i s : Postlarval instars known only for American species. Larva: Gnathosoma with three pairs
of subcapitular setae, dorsally a pair of supracoxal setae. Palp tibia with strong uni- or bidentate odon-
tus. Scutum with two pairs of trichobothria plus two pairs of non-specialized setae; anterior pair of
trichobothria may be separated from the rest of scutum. Laterally one pair of eyes on common ocular
sclerite. Posterior to scutum rows of setae set on prominent plates. Number of setae on coxae I-II-III
two-two-two (rarely three). One pair of distinct plates with a glandular opening („lassenia organs“)
present anterior to coxa III (6-21 a). Anal sclerite present, with two pairs of setae (6-10 c). Leg femora
undivided, tarsus I-III with lateral claws (simple) and median empodium.
Only one species is known from Central Europe.
1. Lassenia xymenae H AITLINGER , 1995

1995 Lassenia xymenae H AITLINGER, Wiad. Parazytol. 41: 471.
R em a rk: No postlarval Lassenia has ever been found in Europe. The description of post-
larval Lassenia from North America does not allow affiliation to any postlarval European
Tanaupodidae. On the other hand no lassenia organ has been mentioned in the description
of L. xymenae. Regarding its habitat preference, L. xymenae possibly constitutes the larva
of Tanaupodus; the generic affiliation needs confirmation.
Genus Tanaupodus H ALLER, 1882

1882 Tanaupodus H ALLER, Jh. Ver. Vaterl. Naturk. Württemberg 38: 323.
Typus generis: Tanaupodus steudelii H ALLER, 1882, by original designation.
Di ag no sis: Adult: Idiosoma slightly elongate, length about 1000 lm, width about
500 lm. Palp tibia with prominent odontus and lateral paradont. Scutum with prominent
crista metopica, anteriorly thickened, without naso. Two pairs of lateral eyes (6-21 c). In-
tegument surface tuberculate, small spine-like setae placed on circular platelets. Genital
opening with three pairs of acetabula, anterior to genital opening a circular small ‘prege-
nital sclerite’ (pregenital tubercle) (6-11 a). Deutonymph and Larva unknown.
Key to species
Re mar k: The only characters known to distinguish the European species are listed in the key. It can-
not be excluded that these differences are within the variability of a single species and thus only one
species might be present in the geographical area.
Adults
1 Setae on dorsum in longitudinal rows; areoles at legs roundish Tanaupodus steudelii (page 220)
– Setae on dorsum irregularly arranged; areoles polygonal . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Tips of dorsal idiosoma setae pointed . . . . . . . . . . . . Tanaupodus passimpilosus (page 220)
– Tips of dorsal idiosoma setae bifid . . . . . . . . . . . . . . . . Tanaupodus bifurcatus (page 219)
1. Tanaupodus bifurcatus C ARL, 1966

1966 Tanaupodus bifurcatus C ARL, Rev. Ecol. Biol. Sol. 3 (2): 329.
D e s c r i p t i o n : C ARL (1966), R OBAUX (1967) (adult).
Di stri bu ti on: Norway, Portugal, Spain, Switzerland.
220 Special part
2. Tanaupodus passimpilosus B ERLESE, 1910 (6-11 a; 6-21 c-d)

1910 Tanaupodus passimpilosus B ERLESE, Redia 6: 354.
D e s c r i p t i o n : C OOREMAN (1943) (female), R OBAUX (1967) (deutonymph).
Di stri bu ti on: Austria, Belgium, France, Hungary, Italy, Norway, Poland, Romania,
Spain, Switzerland.
3. Tanaupodus steudelii H ALLER, 1882

1882 Tanaupodus steudelii H ALLER, Jh. Ver. vaterl. Naturk. Württemb. 38: 325.
1943 Tanaupodus areolatus C OOREMAN, Bull. Mus. r. Hist. nat. Belg. 19 (64): 12.
D e s c r i p t i o n : C OOREMAN (1943) (as Tanaupodus areolatus).
Di stri bu ti on: Belgium, Germany, Poland, Switzerland.
Family Trombidiidae L EACH , 1815

1815 Trombidiidae L EACH , Trans. Linn. Soc. London 11: 387.
Di ag no sis: Adult and Deutonymph: Relatively large (length of idiosoma in some Central
European species up to 5000 lm). Colour in life red, purple or brownish. Body trapezoidal
in outline, oval or elongate. Sensillary area of crista metopica situated in the anterior (6-23
b, c) or middle (6-23 d) part of the sclerite. One pair of trichobothria. Double eyes on ped-
uncles (6-23 b-d). Idiosoma densely covered with setae (neotrichous state). Chelicerae with
movable and fixed digit, not retractable (6-6 c). Palps robust (6-11 e, j). Larva: Idiosoma
oval, with two dorsal sclerites (scutum and scutellum) (6-22 c, e). Scutum with four pairs of
setae: three pairs of non-sensillary setae (AM, AL, PL) and one pair of trichobothria (S).
Scutellum with one (c1) or two (c1-c2) pairs of setae. One pair of eyes on each side of idio-
soma, at the level of posterior edge of scutum. Urstigma between coxae I and II. Anus
surrounded by membraneous valves (6-5 a). Palp genu with one seta or lacking setae.
Coxae I-III with one to two setae (6-5 a; 6-22 f). Femur I with five setae, genua II-III
with three to four setae. Tarsi I-III terminated with paired claws and claw-like empodium.
In some species inner claw on tarsus III may be reduced.
Key to genera
Ma r ked w ith an as te r is k*: genera containing species regularly found in amphibious biotopes.
Larvae
1 Coxa II with one seta, tarsus leg I with one to several ventral eupathidia f, (6-22 a), palp femur
and genu each with a prominent seta, hypostomal seta (bs) long, simple (6-22 b), scutellum less
than half the width of scutum (6-22 c). . . . . . . . . . . . . . . . . . Podothrombium* (page 228)
– Coxa II usually with two setae, tarsus I without ventral eupathidia f (6-22 d), setae on palp femur
and genu, if present, not prominent but small, other characters variable . . . . . . . . . . . . . . 2
2 Genu II/III each with two solenidia (r), palp femur without setae, scutum distinctly wider than
scutellum, pretarsus III with normal claws or inner claw small and directed outwards . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium* (page 221)
– Genu II/III each with one r, palp femur usually with one seta, scutellum of about the same width
as scutum (6-22 e), inner claw at pretarsus III small and directed outwards (6-22 e-g) . . . . 3
3 Seta 1a on coxa I highly modified, odontus at palp tibia bent towards tarsus, hypostomal seta (bs)
long with fine setules (6-22 f). . . . . . . . . . . . . . . . . . . . . . . . . Paratrombium* (page 226)
– Seta 1a on coxa I simple, odontus not bent towards palp tarsus, hypostomal seta with few setules
or with finger-like projections. . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium* (page 231)

1 Palp tibia with spine-like setae, dorsal idiosoma setae simple (6-23 a) or with few setules, eyes on
small protuberances (6-23 b) . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium* (page 228)
– Palp tibia without spine-like setae, dorsal idiosoma setae heavily setulose or modified in another
way, eyes on distinct movable eye stalks (6-23 c-d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 No prominent multisetulose (aliform) setae at tip of tarsi I-IV, prodorsal trichobothria located in
the anterior half of the crista metopica, crista metopica linear (6-23 c), posterior idiosoma setae
various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Pretarsal claws on legs I-IV subtended by a pair of prominent multisetulose (aliform) setae (6-23
f), prodorsal trichobothria located approximately in the centre of crista metopica, crista metopica
widened at level of trichobothria (6-23 d), posterior idiosoma setae setiform (6-23 e) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium* (page 221)
3 Posterior idiosoma setae with simple stem, setiform or with modified stem, but never markedly
thickened and distally truncated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Posterior idiosoma setae (pDS) with markedly thickened stem; majority of pDS with rounded or
truncated top (6-23 h) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium* (page 231)
4 Body not distinctly elongated, idiosoma oval or widened in the anterior part, not incised at the
level of coxae III/IV; pDS with simple stem densely covered with setules (6-23 g); plane of eye
lenses in adults forming a sharp angle with one of the peduncle sides, posterior dorsal setae seti-
form, palp tibia distally with long nude setae. . . . . . . . . . . . . . Paratrombium* (page 226)
– Body elongated, idiosoma incised at level of coxae III/IV, pDS with simple or modified (e.g. long-
itudinally grooved) stem sparsely covered with setules, plane of eye lenses in adults not as de-
scribed above, palp tibia distally without long, nude setae. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dolichothrombium F EIDER , 1945
Genus Allothrombium B ERLESE , 1903

1903 Allothrombium B ERLESE, Redia 1 (2): 251.
1909 Metathrombium O UDEMANS, Ent. Ber. (Amst.) 3 (50): 16 [part.].
1980 Aphithrombium C HILDERS & V ERCAMMEN -G RANDJEAN, J. Kans. Entomol. Soc. 53 (4): 721.
1986 Nippotrombium S OUTHCOTT, Aust. J. Zool. suppl. ser. 123: 40.
Typus generis: Trombidium fuliginosum H ERMANN , 1804, by original designation.
Di ag no sis: Adult and Deutonymph: Body oval or slightly widened at aspidosoma/
opisthosoma border. Sensillary area of crista metopica elongated, in the shape of an inverse
triangle, with bothridia adjoining the triangle base (prodorsal trichobothria located ap-
proximately in the middle of scutum) (6-23 d). Posterior dorsal setae setiform, covered
with setules and set on button-shaped bases (6-23 e; 6-24 c-i). Three pairs of genital acet-
abula. Palp tibia without spine-like setae. Tibia I shorter than tarsus I. Tarsi I-IV distally
with aliform setae (false empodium) (6-23 f). Larva: Scutum longer than wide (in unen-
gorged larvae covering less than half of dorsum), narrowing towards the anterior part.
Hypostomalae with thickened stem and digit-like terminations. Inner edge of cheliceral
claw relatively short and distinctly curved. Palp trochanter, femur and genu without setae.
Leg genua II-III each with two solenidia. Posterior claw on tarsus III reduced or normally
developed (6-24 a, b).
Key to species
Marked with an asterisk*: species regularly found in amphibious biotopes.
Ex clude d: Insufficient descriptions led to exlcusion of the following species: Allothrombium angu-
stiforme W ILLMANN , 1951 (Austria, only deutonymph described); A. fuligineum O UDEMANS , 1905
(The Netherlands, Czech Republic and Germany; only postlarval instars described); A. ignotum
W ILLMANN , 1956 (Czech Republic, known only from larvae - generic affiliation doubtful); A. minu-
222 Special part
Fig. 6-22: Trombidiidae, larvae: a-c, Podothrombium filipes; a, detail: tarsus I; b, detail: gnathosoma
ventral, palp; c, detail: scutum, eyes, scutellum; d, g, Trombidium holosericeum (after M A̧KOL &
W OHLTMANN 2000); d, detail: tarsus I; g, detail: tarsus III; e, f, Paratrombium insulare; e, dorsal
view, details: genu II–III; f, ventral view, details: gnathosoma and coxa with seta 1a; (scale bars =
50 lm).
tissimum W ILLMANN , 1956 (Czech Republic, only deutonymph described); A. pulvinum E WING , 1917
(United States, reported also from Great Britain, Hungary, Spain, Iran and China, no sufficient back-
ground, both with respect to morphology and zoogeography for distinction between A. pulvinum and A.
fuliginosum); A. subtile D ANIEL , 1955 (Czech Republic, description probably based on a deutonymph);
A. tuberculatum W ILLMANN , 1953 (Austria, based on a single, probably anomalous specimen);
A. wolffi K RAUSSE , 1925 (Italy, Austria, Cyprus, and Czech Republic, larvae unknown).
Fig. 6-23: Trombidiidae, postlarval instars: a-b, Podothrombium filipes; a, detail adult: body seta; b,
detail adult: scutum, eyes; c, h, i, Trombidium holosericeum, adult (after M A̧KOL & W OHLTMANN
2000); c, detail: scutum, eyes; h, detail: body setae; i, detail: palp; d-f, Allothrombium fuliginosum,
adult; d, detail: scutum, eyes; e, detail: body setae; f, detail: tip of tarsus III; g, Paratrombium insulare,
detail deutonymph: body seta; (scale bars = 50 lm).
224 Special part
Larvae
1 Tarsus III with two normally developed claws and claw-like empodium (6-24 a) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium fuliginosum* (page 226)
– Inner claw on tarsus III absent or very small (6-24 b) . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Dorsal side of idiosoma with 20 setae (excluding setae on scutum and scutellum) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium neapolitanum O UDEMANS , 1910
– Dorsal side of idiosoma with more than 30 setae (excluding setae on scutum and scutellum)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium wolmari H AITLINGER , 2000
Adults
Characters and metric data specified in the key apply only to adults unless stated otherwise.
1 Idiosoma relatively small (5 2000 lm), elongated, without well-marked widening behind the
aspidosoma-opisthosoma border; crista metopica distinctly narrow, especially in the posterior
part; posterior dorsal setae (pDS) relatively short (5 85 lm), divided into two types (longer/
shorter pDS) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Idiosoma small, medium-sized or large (4 _ 1500-5000 lm), usually with well-marked widening
behind the aspidosoma-opisthosoma border; crista metopica not distinctly narrowed, pDS short
or long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Length pDS 20-80 lm; longer pDS (50-80 lm) with slightly thickened stem, shorter pDS (20-40
lm) with more delicate stem; both types covered with setules of similar length (6-24 c) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium gracile B ERLESE , 1910
– Length pDS 30-50 lm; longer pDS with slightly thickened stem and relatively short setules, short-
er pDS with slightly longer setules, especially in the basal part of the stem (6-24 d) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium incarnatum O UDEMANS , 1905
3 Eyes short, asymmetrical, distinctly narrowed at the base, with much more developed anterior
part of the peduncle and anterior lens; idiosoma small or medium-sized (5 3500 lm); pDS
5100 lm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Eyes not distinctly narrowed at the base; idiosoma medium-sized or large (2000-5000 lm); pDS
4 70 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 Length pDS length 5 70 lm, with relatively long and scattered setules curved towards the setal
stem termination; setules arising close to the top of seta reaching setal stem termination; stem
forming a delicate zigzag-shaped line [the latter character can be misleading and caused by un-
even distribution of the dye within setal stem - however, it was observed in all specimens repre-
senting A. meridionale deposited in Berlese Acaroteca] (6-24 e) . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium meridionale B ERLESE , 1910
– Length pDS 60-95 lm, with almost straight setules, distinctly longer at the base of setal stem;
setules arising close to the top of seta not reaching setal stem termination; stem straight or slightly
bent (6-24 f) . . . . . . . . . . . . . . . . . . . . . . Allothrombium molliculum (C. L. K OCH , 1837)
5 Length pDS 70-115 lm, setules arising close to the tip of pDS reaching the termination of stem;
tarsus I cylindrical or widened distally; idiosoma medium-sized or large (2000-5000 lm) . . 6
– Length pDS 4 100 lm, with thickened, apically narrowing stem covered with relatively short
setules (setules arising close to the tip of pDS not reaching the termination of stem); tarsus I cy-
lindrical, never widened distally (6-24 g); idiosoma large (ca. 5000 lm) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium pergrande B ERLESE , 1903
6 Setae pDS with slightly thickened stem; crista wide, outer lines of middle part of crista convex;
eyes robust, relatively wide; subcuticular mesh with rounded ‘perforations’ (6-24 h); idiosoma
length 3000-4900 lm . . . . . . . . . . . . . . . . . . . Allothrombium adustum O UDEMANS , 1905
– Setae pDS with relatively thin stem, outer lines of middle part of crista slightly concave; eyes only
slightly narrowed at the base and gradually widening towards the lenses; subcuticular mesh with
irregular (oval or polygonal) ‘perforations’ (6-24 i); idiosoma length 2000-4100 lm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothrombium fuliginosum (page 226)
Fig. 6-24: Allothrombium: a, b, larvae; a, Allothrombium fuliginosum, detail: tarsus III; b, Allothrom-
bium wolmari, detail: tarsus III; c–i, adults, detail posterior dorsal body setae; c, Allothrombium
gracile; d, Allothrombium incarnatum; e, Allothrombium meridionale; f, Allothrombium molliculum;
g, Allothrombium pergrande; h, Allothrombium adustum; i, Allothrombium fuliginosum; (d, f, h, i:
after M A̧KOL 2005) (scale bars = 50 lm).
226 Special part
1. Allothrombium fuliginosum (H ERMANN , 1804) (6-23 d-f; 6-24 a, i)

1758 ? Acarus gymnopterorum L INNAEUS, Systema naturae. 10th ed.
1778 Allothrombium aphidis D E G EER, Mémoires pour Servir a l’Histoire des Insectes.’ Vol. VIII:
121.
1804 Trombidium fuliginosum H ERMANN, Mémoire aptérologique. Strasbourg: 23.
1837 ? Trombidium erythrellum C. L. K OCH, D.C.M.A., fasc. 15.
1860 Trombidium holosericeum: PAGENSTECHER, Beiträge zur Anatomie der Milben. Heft I: 1.
1900 Trombidium fuliginosum var. norvegicum T HOR, Christiania 2: 7.
1932 Allothrombium fuliginosum var. clypeata A NDRÉ, Extr. Bull. Soc. Zool. Franc. 57: 196.
1994a Allothrombium reinholdi H AITLINGER, Mitt. Schweiz. Ent. Ges. Bull. Soc. Ent. Suisse 67:
406.
1996 Allothrombium lechi H AITLINGER, Wiad. Parazytol. 42 (4): 455.
De sc ri pti on : For active instars: M A˛ KOL (2005). Adult: Idiosoma length: 2400-4100 lm
(females), 2350-2700 lm (males); width: 1450-2700 lm (females), 1500-2000 lm (males).
Posterior dorsal setae (pDS) length 70-115 lm, with relatively thin stem. Outer lines of
middle part of crista slightly concave; eyes only slightly narrowed at the base and gradually
widening towards the lenses; subcuticular mesh with irregular (oval or polygonal) ‘perfora-
tions’. Deutonymph: All linear measurements shorter than in adults. Larva: R OBAUX
(1974).
H a bit at a nd B io lo gy : Uni- or semivoltine. Postlarval instars active on soil surfaces and
on plants, preying on a range of insects, preferably on aphids. Larvae abundant from June
to August, parasitic on aphids. Widespread, also present in inundation areas.
Di stri bu ti on: Austria, Czech Republic, Finland, France, Germany, Great Britain (Eng-
land, Scotland), Greece, Hungary, Italy, Norway, Poland, Romania, Russia, Spain, Switzer-
land, Slovenia, The Netherlands, Turkey, Algeria, Tunisia.
Genus Paratrombium B RUYANT, 1910

1910 Paratrombium B RUYANT, Zool. Anz. 35 11: 350.
nec Parathrombium B ERLESE , 1918a: Redia 13: 94 (= Calothrombium B ERLESE , 1918a).
nec Paratrombium A NDRÉ 1928b: Bull. Soc. Zool. Fr. 53: 514 (= Trombidium FABRICIUS ,
1775).
Typus generis: Paratrombium egregium B RUYANT, 1910 by original designation.
Di ag no sis: Adult and Deutonymph: Body trapezoidal (slightly widened at aspidosoma/
opisthosoma border) or oval in outline. Sensillary area of crista metopica always in the
anterior part of the sclerite (anterior process of crista always shorter than the posterior
one). Plane of eye lenses in adults forming a sharp angle with lateral side of peduncle. Pos-
terior dorsal setae setiform, with slightly thickened stem, narrowing apically, covered with
setules and set on trunk-shaped bases (6-25 a-d). Three (in adults) or two (in deutonymphs)
pairs of genital acetabula. Palp tibia without spine-like setae. Tibia I always shorter than
tarsus I. Larva: Scutum large (in unengorged larvae covering about half of idiosoma dor-
sum), almost as wide as long (6-22 e). Hypostomalae bipectinate, brush-like (6-22 f). Inner
edge of cheliceral claw relatively long and only slightly curved. Palp trochanter and palp
genu without setae; one spine-like seta on palp femur. Palp tibial claw distinctly curved
towards palp tarsus. Medial coxala I (1a) pectinate (comb-like) (6-2 f), with several digita-
tions. Lateral coxala (1b) with slightly thickened stem. Coxa II with two setae. Genu II-III
each with one solenidion. Tarsus I with one subterminal eupathidium. Posterior claw on
tarsus III reduced or normally developed.
R em a rk: The genus probably includes also some species affiliated to Caenothrombium
O UDEMANS , 1927 (representatives of the genus are known in Europe from Spain, France,
Romania, Cyprus, but not from Central Europe – M A˛ KOL 2000c). With the only exception
of P. insulare and P. megalochirum, no information is available on the biology of Central
European species.
Key to species
Ex clude d: Paratrombium purpureum (C. L. K OCH , 1837) (Germany, Czech Republic, only deuto-
nymph described, description insufficient).
Larvae
1 Length of scutum 4 _ 200 lm . . . . . . . . . . . . . . . . . . . Paratrombium insulare* (page 228)
– Length of scutum 5 200 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . Paratrombium egregium B RUYANT, 1910 or P. megalochirum (B ERLESE , 1910)

1 Posterior dorsal setae (pDS) with relatively short setules arising from the setal stem; the entire seta
(stem plus setules) conical or rectangular in shape; setal stem not thickened . . . . . . . . . . . 2
– Posterior dorsal setae (pDS) with relatively long setules arising from the setal stem; the entire seta
(stem plus setules) oval in shape; setal stem slightly thickened . . . . . . . . . . . . . . . . . . . . . 3
2 Length pDS 45-50 lm; setules arising from the setal stem longer in the proximal part of the seta,
shorter in the distal part; the entire seta conical in shape (6-25 a) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratrombium setosulum (B ERLESE , 1885)
– Length pDS 30-35 lm; proximal and distal setules arising from the setal stem similar in size; the
entire seta conical-rectangular in shape (6-25 b) Paratrombium klugkisti (O UDEMANS , 1917)
3 Live specimens carmine in colour (dye is retained in setae of dissected specimens) (6-25 c) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratrombium insulare* (page 228)
– Live specimens very dark, purple in colour (dye is retained in setae of dissected specimens) (6-25 d)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratrombium megalochirum (B ERLESE , 1910)
Fig. 6-25: Paratrombium adults, detail posterior dorsal body setae: a, Paratrombium setosulum;
b, Paratrombium klugkisti; c, Paratrombium insulare; d, Paratrombium megalochirum; (b, d: after
M A̧KOL 2005) (scale bars = 50 lm).
228 Special part
1. Paratrombium insulare (B ERLESE , 1910) (6-22 e-f; 6-23 g; 6-25 c)

1910 Trombidium insulare B ERLESE, Redia 6: 364.
1947 Dinothrombium insulare T HOR & W ILLMANN, Das Tierreich 71b: 447.
D e s c r i p t i o n : M A˛ KOL (2000b) (larva and postlarval stages, based on material originating
from Finland). Adult: Idiosoma length: 1600-2300 lm (females), 1400-1900 lm (males);
width: 1200-1800 lm (females), 1000-1400 lm (males). Posterior dorsal setae with only
slightly thickened stem, covered with long setules giving the impression of the seta being
oval in shape; length of setae in females 25-32 lm, in males 22-30 lm. Deutonymph: All
linear measurements except for the length of dorsal posterior idiosoma setae shorter than in
adults; dorsal setae on posterior idiosoma 23-32 lm. Larva: (M A˛ KOL (2000b) See diagnosis
of the genus.
H a bit at a nd B i ol og y: In amphibious areas, often salt marshes. Larvae parasitic on Hy-
menoptera.
Di stri bu ti on: Germany, Greece (Corfu), Finland, Norway, Algeria.
Genus Podothrombium B ERLESE , 1910

1910 Podothrombium B ERLESE, Redia 6: 354.
Typus generis: Trombidium filipes C. L. K OCH , 1837, by original designation.
Di ag no sis: Adult: Body oval or slightly widened at aspidosoma/opisthosoma border.
Crista metopica linear. Widened sensillary area located in the anterior or mid part of sclerite
(6-23 b). Posterior dorsal setae simple, sparsely covered with setules or nude, set on button-
shaped, usually asymmetrical bases (6-23 a). Three pairs of genital acetabula. Palp tibia
with modified, spine-like setae forming dorsal and ventral combs on the inner side of
the segment (6-11 e). Leg tibia I shorter, of similar length or longer than tarsus I. Deuto-
nymph: Two pairs of genital acetabula. Dorsal and ventral combs on palp tibia with fewer
setae than in adults; on the dorsal comb behind the odontus one seta. Tibia I always shorter
than tarsus I. Other characters as in adults. Larva: Scutum longer than wide, triangular in
outline, with rounded margins (in unengorged larvae covering less than half of idiosoma
dorsum) (6-22 c). Anterior part of the sclerite distinctly constricted, forming an anterior
process. Scutellum much narrower than scutum. Hypostomalae smooth or with fine barbs
(6-22 b). No setae on palp trochanter. One prominent seta on palp femur and palp genu.
Palp tibial claw single. Coxa II with one seta. Numerous ventral eupathidia on tarsus I (6-
22 a). Inner claw on tarsus III normally developed.
R em a rk: The actual number of species is questionable because of possible intraspecific
geographical variation of characters and pronounced sexual dimorphism (M A˛ KOL & M AR-
USIK 1999). The latter applies mainly to the tibia I / tarsus I ratio. In some species the female
tibia I is shorter, while the male tibia I is longer than tarsus I. In other species both male and
female tibia is either shorter or longer than tarsus I. The problem of double systematics,
common within terrestrial Parasitengona and consisting in constructing separate classifi-
cation systems for larvae and postlarval stages, is even more complicated in relation to
Podothrombium. This is caused by differences between deutonymphs and adults, which
are manifested not only in quantitative characters (e.g. chaetotaxy of palp tibia, tibia I
always shorter than tarsus I in deutonymphs) but also in qualitative characters (e.g. shape
of idiosoma setae).
Key to species
Ex clude d: Insufficient descriptions led to exclusion of Podothrombium curtipalpe (T HOR , 1900)
(Norway, in Central Europe recorded from Austria and Czech Republic, known from postlarval
stages); P. dariae H AITLINGER , 1995 (Poland, based on a single larva); P. dearmatum M IHELČIČ , 1958
(Austria, only postlarval stages known); P. dubiosum (S CHWEIZER , 1951) (Switzerland, only deuto-
nymph described); P. montanum B ERLESE , 1910 (Italy, in Central Europe recorded from Austria, Czech
Republic, Germany and Switzerland, only deutonymph described); P. multispinosum W ILLMANN ,
1951 (Austria, known from postlarval stages only); P. subnudum B ERLESE , 1910 (Italy, Switzerland?,
only deutonymph described).
Larvae
1 Up to ten setae in row C, at the level of scutellum (without setae on scutellum); setae arranged
almost regularly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– More than 14 setae in row C, at the level of scutellum (without setae on scutellum); setae ar-
ranged irregularly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2 Four setae in row C . . . . . . . . . . . . . . . . . Podothrombium exiguum FAIN & R IPKA , 1998
– Six to ten setae in row C (6-26 a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Number of setae on ventral side of the body, behind coxae III 5 25; total length of legs I-III
including coxae (IP) 5 1400 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Number of setae on ventral side of the body, behind coxae III 4 _ 25; IP 4 _ 1400 lm . . . . . 5
4 Number of ventral eupathidia on tarsus I 5 13 (6-26 b); number of dorsal setae, including setae
on scutellum 4 _ 40 . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium filipes* (page 231)
– Number of ventral eupathidia on tarsus I 4 _ 13; number of dorsal setae, including setae on scu-
tellum 5 40 . . . . . . . . . . . . . . . . . . . . Podothrombium pannonicum FAIN & R IPKA , 1998
5 Total length of legs I-III including coxae (IP) 5 1650 lm; number of dorsal setae, including setae
on scutellum 4 _ 40 . . . . . . . . . . . . . . . . . . . . . . Podothrombium verae H AITLINGER , 1995
– IP 4_ 1650 lm; number of dorsal setae, including setae on scutellum 5 40 . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium tymoni H AITLINGER , 1994
6 Total number of dorsal and ventral setae, including setae on scutellum 5 100 . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium kordulae H AITLINGER , 1995
– Total number of dorsal and ventral setae, including setae on scutellum 4 _ 110. . . . . . . . . . 7
7 Total length of legs I-III including coxae (IP) 5 1300 lm; on tarsus I about 10-16 ventral
eupathidia and more than 30 normal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium piriforme R OBAUX & S CHIESS , 1982
– IP 4 1400 lm; on tarsus I about five to six ventral eupathidia and less than 20 normal setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium karlovaicus H AITLINGER , 2003
Adults
The characters observed in deutonymphs depart from the characters pertaining to adults and specified
in the following key. In deutonymphs (except for P. hispanicum, in which both deutonymphs and
adults have one spine), the number of spine-like setae constituting the dorsal and ventral comb on
palp tibia is lower than in adults: one seta is usually observed in the dorsal comb behind the palp tibial
claw (odontus) and one to three setae in the ventral comb; tibia I never exceeds tarsus I in length (see
also the comments in species descriptions).
1 Posterior dorsal setae (pDS) length 5 _ 40 lm, nude or with one or two short barbs, narrowing
apically; spine-like setae forming the ventral comb on palp tibia relatively short and stout; in both
sexes tibia I shorter than tarsus I (6-26 c-d) . . . . Podothrombium bicolor (H ERMANN , 1804)
– Length pDS 4 40 lm, with barbs; narrowing apically or bifurcate at the end; spine-like setae
forming the ventral comb on palp tibia slender, sometimes with ‘splinters’ on both sides; tibia I
shorter or longer than tarsus I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Length pDS 5 _ 65 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Length pDS 4 65 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3 Tibia I shorter than tarsus I (females) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Tibia I longer than tarsus I (males) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
230 Special part
Fig. 6-26: Podothrombium: a, b, Podothrombium filipes, larva; a, details: dorsal shields and setae
(idiosoma borders not shown); b, detail: tarsus I; c-k, postlarval instars, detail: posterior dorsal
body setae; c, Podothrombium bicolor, adult; d, Podothrombium bicolor, deutonymph); e, Podothrom-
bium filipes, adult, female; f, Podothrombium spinosum, adult; g, Podothrombium filipes, adult, male;
h, Podothrombium peragile, adult); i, Podothrombium hispanicum, adult); j, Podothrombium strandi,
adult; k, Podothrombium macrocarpum, adult; (c-k: after M A̧KOL 2005) (scale bars = 50 lm).
4 Setae pDS almost straight, with several barbs and fork-like termination (6-26 e) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium filipes* (females) (page 231)
– Setae pDS curved, pointed, with four to six barbs along the distal half of the stem (6-26 f) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium spinosum F EIDER , 1955
5 Setae pDS pointed, with several barbs along the outer edge of the upper two thirds of the stem (6-
26 g) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium filipes* (males) (page 231)
– Setae pDS pointed or with fork-like termination, very few barbs arising from the stem (6-26 h)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium peragile B ERLESE , 1910
6 One robust, spine-like seta on palp tibia in the dorsal comb, behind the odontus; crista metopica
relatively narrow and slender; posterior process of crista metopica more than twice as long as the
anterior one; eye peduncles square-outlined; tibia I as long as tarsus I (6-26 i) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium hispanicum R OBAUX , 1967
– More than one robust, spine-like seta in the dorsal comb; posterior process of crista metopica
about 1.5 times as long as the anterior one; eye peduncles square-outlined or narrowed at the
base; tibia I and tarsus I different in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 pDS slightly curved, narrowing apically, with three to five short barbs along the distal half of the
stem; on palp tibia two to four spine-like setae in the dorsal comb, behind the odontus; in both
sexes tibia I longer than tarsus I (6-26 j) . . . . . . . . Podothrombium strandi B ERLESE , 1910
– pDS slightly curved or straight, with several barbs, seta termination asymmetrical (the most dis-
tally placed setula not reaching the tip of setal stem); on palp tibia three to six spine-like setae in
the dorsal comb, behind the odontus; in females tibia I shorter than tarsus I (6-26 k) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podothrombium macrocarpum B ERLESE , 1910
1. Podothrombium filipes (C. L. K OCH , 1837) (6-11 e; 6-22 a-c; 6-26 a-b, e, g)
1837 Trombidium filipes C. L. K OCH , D.C.M.A., fasc. 15.
1910 Podothrombium magnum B ERLESE, Redia 6: 355.
1963 Podothrombium bicolor var. cisalpinum: S CHWEIZER & B ADER, Mem. Soc. Helv. Sci. Nat. 84
(2): 363.
1994b Podothrombium proti H AITLINGER, Linzer biol. Beitr. 26 (1): 534.
2000 Podothrombium roari H AITLINGER, Ent. Fenn. 11: 188.
D e s c r i p t i o n : M A˛ KOL (2005) for active instars. Adult: Idiosoma length: 1900-3100 lm
(females), 1400-1600 lm (males); width: 1050-1700 lm (females), 800-900 lm (males).
Posterior dorsal setae: in females (length 45-65 lm) stout, almost straight, with several
barbs and fork-like termination; in males (length 40-50 lm) curved, pointed, with several
barbs along the outer edge of the distal two thirds of the stem. In females: tibia I shorter
than tarsus I, in males: tibia I longer than tarsus I. Deutonymph: Posterior dorsal setae
barbed in the distal half of the stem, usually with asymmetrical fork-like termination.
All linear measurements shorter than in adults. Larva: M A˛ KOL (2000a).
H a bit at a nd B io lo gy : Uni- to semivoltine. Postlarval active instars prey on aphids as
well as on dipteran larvae and pupae. Larvae abundant from June to August, parasitic on
aphids. Often found at humid to wet locations.
Di stri bu ti on: Austria, Czech Republic, France, Germany, Great Britain, Hungary, Italy,
Norway, Poland, Slovenia, Switzerland, Turkey.
Genus Trombidium FABRICIUS , 1775

1758 Acarus L INNAEUS, Syst. Nat. 10th ed.: 617 [part].
1775 Trombidium FABRICIUS, Syst. Entomol.
1795 ?Atomus L ATREILLE, Mag. Encycl. J. Sci. 4, 18, 28.
1909 Metathrombium O UDEMANS, Ent. Ber. 3 (50): 16 [part.].
232 Special part
1910 Sericothrombium B ERLESE, Redia 6: 365.

1951 Teresothrombium F EIDER , Acad. R. P. R.: 12.
Typus generis: Acarus holosericeus L INNAEUS , 1758 (= Trombidium holosericeum (L IN-
NAEUS , 1758), by subsequent designation (L ATREILLE , 1810)).
Di ag no sis: Adult and Deutonymph: Idiosoma trapezoidal in outline, widened at aspi-

dosoma/opisthosoma border. Posterior edge of idiosoma with distinct incision. Sensillary
area of crista metopica always in anterior part of the sclerite (anterior process of crista
always shorter than the posterior one) (6-23 c). Posterior dorsal setae with thickened
stem (widened distally, with parallel sides or exceptionally narrowing apically), covered
with setules and set on trunk-shaped bases (6-27 d-g; 6-28). Three (in adults) or two
(in deutonymphs) pairs of genital acetabula. Palp tibia without spine-like setae (6-23 i).
Tibia I always shorter than tarsus I. Larva: Scutum large (in unengorged larvae covering
about half of idiosoma dorsum), almost as wide as long. Hypostomalae either in the shape
of a calyx, with several digitations or slender, narrowing apically and covered with setules
(6-5 a; 6-27 a, b). Inner edge of cheliceral claw relatively short and curved. Palp trochanter
and genu without setae; one spine-like seta on palp femur. Medial coxala I (1a) simple.
Coxa II with two setae. Genua II-III each with one solenidion. Tarsus I with one sub-
terminal eupathidium (6-22 d). Inner claw on tarsus III reduced in length and directed out-
wardly (6-22 g).
Key to species
Ex clude d: Insufficient descriptions led to exclusion of following species: Trombidium fturum
S CHWEIZER , 1951 (Switzerland, Spain, known only from deutonymph); T. fuornum S CHWEIZER ,
1951 (Switzerland, France, known only from deutonymph); T. hungaricum K OBULEJ , 1957 (Hungary,
known from larva and postlarval instars, larva excluded from key due to the poor description);
T. monoeciportuense (A NDRÉ , 1928a) (Monaco, Czech Republic, known only from postlarval stages);
T. pygiacum C. L. K OCH , 1837 (Germany, known only from adults); T. raeticum (S CHWEIZER &
B ADER , 1963) (Switzerland, known only from adults, generic affiliation doubtful); T. susteri F EIDER ,
1956 (Hungary); T. toldti (M ETHLAGL , 1928) (Austria, known only from larvae).
Larvae
1 Hypostomalae (bs) with finger-like or brush-like digitations (6-27 a) . . . . . . . . . . . . . . . . 2
– Hypostomalae simple, setulose, with slightly thickened and narrowing apically stem (6-27 b) 7
2 Setae c3 placed anteriorly to scutellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . Trombidium rhopalicus (V ERCAMMEN -G RANDJEAN & P OPP, 1967)
– Setae c3 levelled with scutellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Hypostomalae (bs) stout, distally brush-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Hypostomalae (bs) stout, with finger-like digitations (6-27 a) . . . . . . . . . . . . . . . . . . . . . 5
4 Anterior non-specialized setae on scutum (AM) nude, digitations of palp tibial claw (odontus)
equal in length; scutellum trapezoidal in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium mediterraneum (B ERLESE , 1910)
– Anterior non-specialized setae on scutum (AM) with distinct barbs; digitations of palp tibial claw
(odontus) unequal in length (6-27 c); scutellum crescent-like in shape. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium dacicum (F EIDER , 1950)
5 Scutellum with delicate incision in the anterior border . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium latum C. L. K OCH , 1837
– Anterior edge of scutellum in the form of a straight line . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Setae SL (c1) of scutellum situated at half length of the sclerite; hypostomalae (bs) with about 14
digitations . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium holosericeum (L INNAEUS , 1758)
– Setae SL (c1) of scutellum situated just behind half length of the sclerite; hypostomalae with about
11-12 digitations. . . . . . . . . . . . . . . . . . . . . . . . Trombidium geniculatum (F EIDER , 1955)
7 Setae SL (c1) of scutellum situated at about 1/3 length of the sclerite . . . . . . . . . . . . . . . . . .

.... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium brevimanum* (page 236)
– Setae SL (c1) of scutellum situated at about half length of the sclerite . . . . . . . . . . . . . . . . . .
.... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium breei S OUTHCOTT, 1986
Fig. 6-27: Trombidium: a-c, larvae; a, Trombidium holosericeum, detail: gnathosoma ventral;
b, Trombidium brevimanum, detail: gnathosoma ventral; c, Trombidium dacicum, detail: odontus
of palp tibia; d-g, adults, detail posterior dorsal body setae; d, Trombidium heterotrichum; e, Trom-
bidium dacicum; f, Trombidium kneissli; g, Trombidium holosericeum; (d-g: after M A̧KOL 2005) (scale
bars = 50 lm).
234 Special part

Metric data given in the following key constitute an auxiliary character, apply to adults (unless stated
otherwise) and pertain to the extreme forms (e.g. the longest setae were taken into account in case of
pDS I and the shortest – in case of pDS II); in deutonymphs all linear measurements are relatively
smaller.
1 Posterior dorsal setae slightly widened in the distal part, with parallel sides or narrowing apically;
crown-like structure around the setal stem base moderately or distinctly developed . . . . . . 2
– The majority of posterior dorsal setae markedly widened distally, sometimes with well developed
chamber in the distal part; crown-like structure around the setal stem base moderately developed
or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2 Longer posterior dorsal setae, pDS I (60-90 lm) with parallel sides or slightly widened in the
distal part; shorter posterior dorsal setae, pDS II (40-55 lm) narrowing apically; crown-like
structure around the setal stem base consisting of distinct long and rough setules (6-27 d) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium heterotrichum (B ERLESE , 1910)
– All posterior dorsal setae (pDS I and pDS II) with parallel sides or slightly widened in the distal
part, never narrowing apically; setules around the setal stem base distinct or moderately devel-
oped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 All pDS with truncated top; pDS II with distinct tubercles; setules around the setal stem base well
developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– At least some pDS with rounded top; setules around the setal stem base moderately developed
................................................................. 5
4 Setae pDS I (80-110 lm) and pDS II (35-55 lm) similar in shape, slender, all slightly widened in
the distal part, sometimes with obliquely truncated top covered with tubercle-like protrusions;
setules forming the crown-like structure around the setal stem base distinctly elongated (6-27 e)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium dacicum (F EIDER , 1950)
– Setae pDS I (50-80 lm), pDS II (25-50 lm) with distinct tubercle-like protrusions; crown-like
structure around the setal stem base well developed but not distinctly elongated (6-27 f) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium kneissli* (page 236)
5 Setae pDS I and pDS II different in shape; the differences pertain especially to extreme forms: pDS
I (55-110 lm) club-shaped, pDS II (25-55 lm) truncated, both widened in the distal part and
covered with tubercle-like protrusions; crown-like structure around the setal stem base moder-
ately developed (6-27 g); male and female genital sclerites similar in size . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium holosericeum (L INNAEUS , 1758)
– Gradual transition of pDS I into pDS II; differences between setae pertain to extreme forms;
genital sclerites of males may be larger than in females . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 The longest pDS 5 120 lm; pDS I (65-115 lm) club-shaped, pDS II (20-55 lm) almost with
parallel sides; crown-like structure around the setal stem base moderately developed (6-28 a);
genital sclerites in males much larger than in females . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium geniculatum (F EIDER , 1955)
– The longest pDS 4 120 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Setae pDS I (105-155 lm) club-shaped, with parallel sides or slightly widened distally, sometimes
with oblique termination and few tubercles; pDS II (30-50 lm) gradually widening towards the
end, with distinct tubercles; crown-like structure around the setal stem base moderately devel-
oped (6-28 b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium monticola M A˛ KOL , 2005
– Setae pDS I (75-125 lm) only slightly and gradually widening towards the distal part, pDS II (30-
60 lm) with parallel sides; setae on aspidosoma markedly longer; distinct crown-like structure
consisting of rough setules around the setal stem base (6-28 c); in live specimens white-dusted
spots on idiosoma dorsum; male and female genital sclerites comparable in size . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium latum C. L. K OCH , 1837
8 Setae pDS almost symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– At least some pDS distinctly asymmetrical in the distal part . . . . . . . . . . . . . . . . . . . . . 11
9 Setae pDS I and pDS II different in shape; pDS I (75-90 lm) with rounded top in the form of bulb;
pDS II (35-50 lm) with distinctly widened top, truncated at the end; pDS II termination seen
from above in the form of a 10-13-pointed (sometimes slightly asymmetrical) star (6-28 d) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trombidium bulbisetum* (page 236)
– Gradual transition of pDS I into pDS II. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Setae pDS I and pDS II almost symmetrical; setules arising at the setal stem base more robust and
longer than the remaining ones (6-28 e) . . . . . . . Trombidium hungaricum K OBULEJ , 1957
– Setae pDS I and pDS II similar in shape, with the termination in the form of inverted cone, slightly
depressed at the top; pDS termination seen from above in the form of a 10-11-pointed star with-
out distinct protrusions (6-28 f) . . . . . . . . . . . . . . . Trombidium conisetum M A˛ KOL , 2005
Fig. 6-28: Trombidium adults, detail posterior dorsal body setae: a, Trombidium geniculatum; b,
Trombidium monticola; c, Trombidium latum; d, Trombidium bulbisetum; e, Trombidium hungari-
cum; f, Trombidium conisetum; g, Trombidium mediterraneum; h, Trombidium brevimanum; i, Trom-
bidium rimosum; (b-d, f, h, i: after M A̧KOL 2005) (scale bars = 50 lm).
236 Special part
11 Setae pDS I (50-75 lm) somewhat widened distally, with rounded, usually symmetrical termi-
nation, pDS II (25-45 lm) with irregular, distinctly frayed termination formed by several lobes/
petals (6-28 g) . . . . . . . . . . . . . . . . . . . . . . Trombidium mediterraneum (B ERLESE , 1910)
– Setae pDS widened distally, termination compact, not frayed, usually asymmetrical, covered with
several tubercles, pDS II widened and flattened at top, sometimes slightly frayed, but without
distinct lobes/petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Setae pDS I (35-60 lm) gradually widening towards the top, pDS II (20-35 lm) with widened and
slightly flattened, sometimes frayed, top (6-28 h); live specimens purple in colour (dye retains also
in setae of dissected specimens) . . . . . . . . . . . . . . . Trombidium brevimanum* (page 236)
– Setae pDS I (40-75 lm) (when flattened) with distinct triangle-like top, widening in the distal
part, pDS II (20-40 lm) not so distinctly widened, with tubercle-like protrusions at the top;
the proximal part of setal stem almost parallelsided, densely covered with setules (6-28 i),
live specimens reddish to orange in colour . . . . . . Trombidium rimosum C. L. K OCH , 1837
1. Trombidium brevimanum (B ERLESE , 1910) (6-5 a; 6-6 c; 6-27 b; 6-28 h)

1910 Sericothrombium brevimanum B ERLESE, Redia 6: 365.
1996 Trombidium rowmundi H AITLINGER, Wiad. Parazytol. 42 (4): 457.
D e s c r i p t i o n : M A˛ KOL (2005) for active instars. Adult: Idiosoma length 1800-2900 lm
(females), 1650-2350 lm (males); width 1300-2100 lm (females), 1200-1650 lm (males).
Posterior dorsal setae: pDS I (35-60 lm) gradually widening towards the top, pDS II (20-35
lm) with widened and slightly flattened top; live specimens purple in colour (dye is retained
in setae of dissected specimens). Deutonymph: All linear measurements shorter than in
adults. Larva: W OHLTMANN (1999b: redescription with data on biology and ecology).
H a bit at a nd B i ol og y: Uni- to semivoltine. Larvae abundant from June to August, para-
sitizing Argiopidae (Araneae). Often found in amphibious biotopes like inundation areas,
although not restricted to such biotopes.
Di stri bu ti on: Finland, Germany, Great Britain (England), Italy, Norway, Poland, Spain,
Switzerland.
2. Trombidium bulbisetum M A˛ KOL , 2005 (6-28 d)

2005 Trombidium bulbisetum M A˛ KOL , Trombidiidae of Poland: 1 (ns): 64
D e s c r i p t i o n : M A˛ KOL (2005) for adult and deutonymph. Adult: Only females known;
idiosoma length: 2700-4100 lm; width: 2000-2700 lm. Posterior dorsal setae pDS I
and pDS II different in shape; pDS I (length 75-90 lm) with rounded top in the form
of bulb; pDS II (length 35-50 lm) with distinctly widened top, truncated at the end;
pDS II termination seen from above in the form of a 10-13-pointed (sometimes slightly
asymmetrical) star. Deutonymph: All linear measurements shorter than in adults. Larva
unknown.
H a bit at a nd B io lo gy : Life cycle unknown; hygrophilous, found in forested and in open
habitats.
3. Trombidium kneissli (K RAUSSE , 1915) (6-27 f)

1915 Sericothrombium Kneissli K RAUSSE, Arch. für Naturg. A. 7 (81): 128.
1951 Trombidium holosericeum alpinum S CHWEIZER, Ergeb. Wiss. Unters. Schweiz. Nat. Parks
(n.f.) III 23 (2): 113.
D e s c r i p t i o n : M A˛ KOL (2005) for adult and deutonymph. Adult: Similar to Trombidium
holosericeum. Idiosoma length 2200-2650 lm (females), 1650-2300 lm (males); width
1650-2150 lm (females), 1200-1600 lm (males). Posterior dorsal setae pDS I (50-80 lm)
slightly widened distally, pDS II (25-50 lm) with distinct tubercle-like protrusions. Crown-
like structure around the setal stem base well developed. Deutonymph: All linear measure-
ments shorter than in adults. Larva unknown.
H a bit at a nd B io lo gy : Life cycle unknown. Hygrophilous.
Di stri bu ti on: Austria, Bosnia and Herzegovina, Czech Republic, France, Germany,
Hungary, Poland.
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7. Hydrachnidia 241
7. Acari: Hydrachnidia
Kees Davids [†], Antonio Di Sabatino, Reinhard Gerecke, Terence Gledhill, Harry Smit
and Henk van der Hammen [†]
General part
Diagnosis
Plesiotypically soft bodied mites, in many clades with more or less secondary sclerotization.
At the larval stage characterized by (1) the anal opening usually surrounded by an undi-
vided sclerite (probably apotypical: paired sclerites in Stygothrombidioidea, reduced scler-
ites in the subfamily Wandesiinae [Hydryphantoidea]); (2) Legs I/II never more than double
the length of the idiosoma; (3) P-3 (genu) with two setae. Dimensions, coloration, and idio-
soma sclerites variable in deutonymphs and adults, typically with (1) legs in a circular ar-
rangement or restricted to the anteriormost part of the idiosoma; (2) palp five-segmented;
(3) plesiotypically with 16 pairs of regularly arranged idiosomal glands associated with
sensillae (apotypically with number of glands often reduced, rarely increased); (4) legs va-
riable in shape, in many taxa with swimming setae.
Introduction
The members of the Suborder Parasitengona comprise highly specialized terrestrial as well
as secondarily aquatic species. They are characterized by a complex life cycle with parasitic
larvae and predatory post-larval stages. Prelarva, protonymph and tritonymph are immo-
bile transformation stages (calyptostases) (see also the preceeding chapter).
In the literature, the „true water mites“ have been variously referred to as Hydracarina, Hy-
drachnellae, Hydrachnidia or Hydrachnida. The old term „Hy d r ac a ri na“ included Hy-
drachnellae and the family Halacaridae. However, as Halacarid mites are, in morphology
and life-history, very different from the Hydrachnidia (see chapter 5 of this book), „Hydra-
carina“ is obviously a polyphyletic grouping. The name „Hydrachnidia“, introduced by VAN
DER H OEVEN (1849), has been generally accepted during the past decades. The proposal of
S CHWOERBEL (1986b), to incorporate the Hydrovolzioidea, a basic superfamily of Hydrach-
nidia, into the Halacaroidea is not supported by synapomorphies, but is based on external
similarities clearly developed convergently in the two phylogenetically distant clades.
C OOK (1974) established a system grouping the families into eight s up e rf a m ilie s (Stygo-
thrombioidea, Hydrovolzioidea, Eylaoidea, Hydrachnoidea, Hydryphantoidea, Lebertioi-
dea, Hygrobatoidea and Arrenuroidea). T UZOVSKIJ (1987) proposed to split the division of
Eylaoidea into three superfamilies Eylaoidea, Limnocharoidea and Piersigioidea, the intro-
duction of Wandesioidea (formerly Hydryphantidae Wandesiinae) and Pontarachnoidea
(formerly Hygrobatidae Pontarachninae), and to synonymize Arrenuroidea with Hygroba-
toidea. He considered the Stygothrombioidea as a separate phalanx, the Stygothrombidia.
PANESAR (2003) provided evidence for the synonymy of Hygrobatoidea and Arrenuroidea.
H ARVEY (1998) proposed to place the family Sperchontidae in its own superfamily, the
Sperchontoidea. However, in numerous details questions are still unresolved and further
research is needed, mainly concerning the morphology of immature stages and molecular
systematics, before a new subdivision of the Hydrachnidia into superfamilies may be phy-
logenetically corroborated and generally accepted. In this basic textbook, we therefore fol-
low the arrangement of superfamilies and families proposed by C OOK (1974). In some as-
pects, this „conservative“ system is evidently artificial, but it works well in separating the
enormous diversity of European water mites into more „digestible“ units.

https://doi.org/10.1007/978-3-662-55958-1_7
242 General part
In the following, we give a world-wide survey of families and subfamilies. Superfamilies are
given in the (approximatively phylogenetic) sequence in which they are traditionally trea-
ted (also in this work), families and subfamilies are in alphabetical order. Taxa not occur-
ring in the area covered are placed in parentheses (based on K. O. V IETS 1987 and updated
from C OOK 1992, C OOK, S MITH & H ARVEY 2000, G ERECKE 2004, G ERECKE et al. 1999,
G OLDSCHMIDT 2004, G OLDSCHMIDT & G ERECKE 2003, M ITCHELL 2003, PANESAR 2004,
S CHWOERBEL 1984, I.M. S MITH 1989, 1990a, b, I.M. S MITH & C OOK 1997, T UZOVSKIJ
et al. 2001, VALDECASAS 2001, and W ILES 1996). Following Ö ZDIKMEN (2006) the pre-
occupied name Thyadinae is replaced by Euthyadinae. The synonymization of Anisitsielli-
dae with Limnesiidae (PANESAR 2004) is not followed here. Consequently subfamilies
introduced there in Limnesiidae, are here placed in Anisitsiellidae.
Walter mite families and subfamilies - an updated worldwide survey.
Superfamily Family Subfamily
Stygothrombioidea Stygothrombiidae
Hydrovolzioidea Acherontacaridae
Hydrovolziidae
Eylaioidea (Apheviderulicidae)
Limnocharidae Limnocharinae
(Rhyncholimnocharinae)
Eylaidae
Piersigiidae Piersigiinae
(Stygolimnocharinae)
Hydrachnoidea Hydrachnidae
Hydryphantoidea (Ctenothyadidae)
Hydrodromidae
Hydryphantidae (Ankelothyadinae)
(Chimerathyadinae)
(Cyclothyadinae)
Diplodontinae
(Eupatrellinae)
Hydryphantinae
(Mamersinae)
Protziinae
Pseudohydryphantinae
Tartarothyadinae
Euthyadinae
Wandesiinae
(Rhynchohydracaridae) (Clathrosperchontinae
(Rhynchohydracarinae)
(Santiagoacarinae)
(Teratothyadidae)
(Thermacaridae)
(Zelandothyadidae)
Lebertioidea (Acucapitidae)
Anisitsiellidae Anisitsiellinae
(Bandakiinae)
(Bharatoniinae)
Nilotoniinae
Bandakiopsidae
Lebertiidae
Oxidae
Rutripalpidae
Sperchontidae Sperchontinae
(Apeltosperchontinae)
7. Hydrachnidia 243

(Stygotoniidae)
Teutoniidae
Torrenticolidae (Neoatractidinae)
(Testudacarinae)
Torrenticolinae
Hygrobatoidea (Astacocrotonidae)
Aturidae Albiinae
Aturinae
Axonopsinae
(Notoaturinae)
(Zelandospinae)
Feltriidae
(Ferradasiidae)
Frontipodopsidae
Hygrobatidae Hygrobatinae
(Rhynchaturinae)
Lethaxonidae
Limnesiidae (Epallagopodinae)
Kawamuracarinae
Limnesiinae
(Mixdeinae)
(Mixolimnesiinae)
(Neomamersiinae)
(Neotorrenticolinae)
(Nicalimnesiinae)
(Protolimnesiinae)
(Psammolimnesiinae)
(Rheolimnesiinae)
(Stygolimnesiinae)
(Tyrrelliinae)
(Omartacaridae)
Pionidae Foreliinae
Huitfeldtiinae
(Najadicolinae)
Pioninae
(Schminkeinae)
Tiphyinae
Pontarachnidae
Unionicolidae (Encentridophorinae)
(Omanohydracarinae)
Pionatacinae
(Pollicipalpinae)
Unionicolinae
Wettinidae
Arrenuroidea Acalyptonotidae
(Amoenacaridae)
(Arenohydracaridae)
Arrenuridae
Athienemanniidae (Africasiinae)
Athiennemanniinae
(Notomundamellinae)
(Plaumanniinae)
(Stygameracarinae)
Bogatiidae Bogatiinae
(Horreolaninae)
244 General part

Chappuisididae Chappuisidinae
(Tsushimacarinae)
(Gretacaridae)
(Harpagopalpidae)
Hungarohydracaridae Hungarohydracarinae
Balcanohydracarinae
(Cubanohydrocarinae)
(Kantacaridae)
Krendrowskiidae
(Laversiidae)
Mideidae
Momoniidae (Cladomomoniinae)
(Cyclomomoniinae)
Momonidinae
Momoniinae
Stygomomoniinae
Mideopsidae Mideopsinae
(Mideopsellinae)
(Phreatomideopsinae)
Neoacaridae
(Nipponacaridae)
Nudomideopsidae
(Uchidastygacaridae) Morimotacarinae
Uchidastygacarinae
External morphology
Dimensions of adult Hydrachnidia range from 200 up to 10.000 lm in length, most species
are between 500 and 2.000 lm in length. As in other acarines, the body plan comprises two
tagmata, an antero-ventrally situated gnathosoma and a posterior idiosoma (2-1). Hy-
drachnidia exhibit a broad spectrum of body pigmentation. Red and other bright c ol ou rs
are a striking feature of many species of the group. Representatives of most plesiomorphic
taxa are red (Hydrachnoidea, Eylaoidea, Hydryphantoidea) similar to terrestrial parasiten-
gones, due to the presence of pigment distributed throughout the body. In Limnochares
aquatica L INNAEUS , 1758, M EYER & K ABBE (1994) observed 16 different carotenoids.
Members of other taxa show various colours, ranging from red through orange, yellow,
green to blue. Colours may originate not only from pigment granules, but also from
accumulation of particular compounds in internal organs, interference at particular inte-
gumental structures, or a combination of all three factors. The in te gu m en t consists of,
endo-, exo-, and epicuticle covered by a wax layer and is described in detail by S CHMIDT
(1935), C ASSAGNE-MÉJEAN (1966a), R EDMOND (1972) and A LBERTI & C OONS (2001).
Setae and spines can be discerned as integumental structures. A seta can be hollow or solid,
if stiff it is called a spine. Swimming setae (2-4 h) are long and slender, mostly attached in
rows to the anterior surface of leg segments and often similar in length to, or longer than,
the following segment.
The id ios om a is plesiotypically mainly unsclerotized, exhibits no external segmentation
and is round or ovoid and somewhat flattened dorsoventrally in outline (2-4 f-g, i). Where
the idiosoma is covered with dorsal and ventral shields these are separated by the so called
„dorsal furrow“ (visible e.g. in 7-7 c). The ground plan of the organization of the idiosoma
is given in 7-1 and 7-2. Two pairs of l at er al e ye s, typically located close together on each
side and often enclosed in bean-shaped capsules, may be reduced or even absent in inter-
7. Hydrachnidia 245
Fig. 7-1: Hydrachnidia, plesiotypical idiosoma organization, dorsal view (after D I S ABATINO et al.
2002, modified). Terminology following L UNDBLAD 1927 (in parentheses: following Z AHVATKIN ,
after T UZOVSKIJ 1987).
Explanation of abbreviations:
T UZOVSKIJ 1987 L UNDBLAD 1927
- Fr: Frontale
Fch: frontalis chelicerarum Dgl-1: Dorsoglandulare 1 (Antenniforme)
Vi: verticalis interna Dgl-2: Dorsoglandulare 2
Oe: occipitalis externa Dgl-3: Dorsoglandulare 3
Hi: humeralis interna Dgl-4: Dorsoglandulare 4
Sci: scapularis interna Dgl-5: Dorsoglandulare 5
Li: lumbalis interna Dgl-6: Dorsoglandulare 6
Si: sacralis interna Dgl-7: Dorsoglandulare 7 (= Vgl-4, Ventroglandulare 4)
Ve: verticalis externa Lgl-1: Lateroglandulare 1
He: humeralis externa Lgl-2: Lateroglandulare 2
Sce: scapularis externa Lgl-3: Lateroglandulare 3
Le: lumbalis externa Lgl-4: Lateroglandulare 4
Oi: occipitalis interna Postoc: Postoculare
Fp: frontalis pedipalporum Preoc.: Preoculare
- Dc-1-4: Dorsocentralia 1-4
- Dl-1-4: Dorsolateralia 1-4
246 General part
Fig. 7-2: Hydrachnidia, plesiotypical idiosoma organization, ventral view (after D I S ABATINO et al.
2002, modified). Terminology following L UNDBLAD 1927 (in parentheses, following Z AHVATKIN , after
T UZOVSKIJ 1987).
Ac-1-3: Acetabulum 1-3
An: Anus
Genpl: Genital plate
Postgen: Postgenitale
Pregen: Pregenitale
T UZOVSKIJ 1987 L UNDBLAD 1927
- Cx-1-4: first to fourth coxae
Hv: humeralis ventralis Cxgl-2: Coxoglandulare 2 (Cxgl-1 and -3 not existing)
Sce: scapularis externa Cxgl-4: Coxoglandulare 4
Si: sacralis interna Dgl-7: Dorsoglandulare 7 (= Vgl-4: Ventroglandulare 4)
Pi: praeanalis interna Vgl-1: Ventroglandulare 1
Se: sacralis externa Vgl-2: Ventroglandulare 2
Ci: caudalis interna Vgl-3: Ventroglandulare 3
- V-1-4: Ventralia 1-4
7. Hydrachnidia 247
stitial species. Two pairs of prominent setae are situated anterior (pr eo cu la ri a), and pos-
terior (p ost oc ul ari a) to the lateral eyes. Their basal sclerites may be expanded and fused
with neighbouring plates to form the frontal plate („fr ont al e“, e.g. 7-13 f-g). In the more
early derivative genera a single m e di an ey e („frontal organ“) is situated on the surface
between the lateral eyes. It may or may not contain pigment and it may or may not lie on a
frontal plate - in the first case often together with the pre- and postocularia. Water mites are
unique compared to other mites in having g la nd ul ari a (defense glands, see below, „anat-
omy“) on the idiosoma. Longitudinal series of such paired glandularia, m u sc le at tac h -
m e nt scl e rit e s („dorsalia“, „ventralia“) and sl it org an s (slender sac-like structures,
which open through slits in the integument) can be observed. The slit organs are sometimes
incorrectly named „lyrifissures“, a term referring to multiple slit organs which are not pre-
sent in mites. The sclerite plates and glandularia generally have a name related to their
location, hence the terms „dorsocentralia, dorsolateralia, ventralia“ for muscle attach-
ments and „dorsoglandularia, lateroglandularia, ventroglandularia, coxo- or epimeroglan-
dularia“ for glandularia. They are often individually characterized by numbers, counting
from the anterior to the posterior body margin. W ILES (1997), in a study on the homology
of elements at the water mite idiosoma surface postulated that plesiotypically five pairs of
slit organs and 16 pairs of glandularia are present (dorsally six pairs of dorso- and four pairs
of lateroglandularia, ventrally four pairs of ventro-, and two pairs of coxoglandularia).
G ERECKE (1996b) compared the terminology of plates and glands proposed by different
authors and recommended that of L UNDBLAD (1927c). In this system (7-1, 7-2, right hand),
the most anterior located pair of glandularia (dorsoglandularia 1) which often display par-
ticular characteristics, are called „pr ea nt en ni for m ia“, posteriorly followed by dorso-
glandularia 2-7. S MITH et al. (2001) follow this system, except for dorsoglandularia 7
sensu L UNDBLAD named by these authors „lateroglandularia 5“. In a series of papers,
B ADER (e.g. 1975, 1982a) examined the segmentation of water mites and developed his
own scheme for identifying plates and glands, based on a hypothetical archetypal acarine
morphology. As he decided not to include the most anterior located pairs of glandularia
(dorsoglandularia 1 and 2, called by Bader „p ra e- and po sta nte n ni form i a“) in the
numbering, dorsoglandularia 1 (+ n) sensu B ADER correspond to dorsoglandularia 3 (+
n) sensu L UNDBLAD . Furthermore, he misidentified two additional pairs of coxoglandular-
ia, which are universally absent. The only two glandularia associated with the coxae are
plesiotypically associated with coxae two and four and are therefore named by W ILES
(1997) as coxoglandularia 2 and 4. T UZOVSKIJ (1987), following Z AHVATKIN (1952) gives
a comparative setal terminology reflecting the segmental structures of the idiosoma, this
being equally applicable for all developmental instars (7-1, 7-2 - see also G ERECKE &
T UZOVSKIJ 2001, T UZOVSKIJ et al. 2001). However, in some genera it is exceedingly diffi-
cult to work out the homologies of the glandularia. In figures 7-1 and 7-2 we allow the
comparison of the two most important, basically different terminology schemes; in the
following chapters of this work we follow the system of L UNDBLAD .
The c o xa e are the first basal segments of the legs. In water mites, as in all members of
Actinotrichida, they are incorporated in the ventral surface and immovable. They may
be enlarged to such an extent that they cover the major part of the venter. In general,
the first and second coxae, and the third and fourth coxae of both sides are fused to
form the anterior and the posterior c o xal gr oup respectively (7-2). The boundary be-
tween these fused coxae is called a suture. Occasionally the anterior coxal group may
fuse medially (three coxal groups, e.g. 2-4 f-g) or all of the coxae may be fused to form
a single group („co xal sh ie ld“, 7-12 f-h). The anterior pair of coxae embraces the gnatho-
somal bay.
The ge n ita l f ie l d is located posterior to the coxae, often in an indentation („g en it al
ba y“) bordered by the second coxal group (7-2). It consists of the gonopore (the opening
248 General part
of the reproductive system), often flanked by sclerotized plates: Movable paired sclerites
which may cover the gonopore and acetabula when closed, are called „g e ni ta l fl ap s“ (7-
15 f-g). If the acetabula are not arranged between the sclerites flanking the gonopore, but
on their surface proper, these sclerites are called „g en it al pl at es“ (7-15 c, i- k, m-n). Such
plates are generally rather immovable, often fused to a ring enclosing the gonopore (7-15 i),
but in females they remain often paired (7-15 k) and in some groups they can rotate slightly
against their longitudinal axis. Additional small sclerites called „p re -“ and „po stg e ni -
ta li a“ are situated at the anterior and posterior margin of the gonopore, in species with
movable genital flaps acting as their hinge joint. The e xc re tor y p ore (anal pore), often
surrounded by a sclerotized ring, is situated posterior to the genital field. If a posterior
extension of the body is present, this part of the idiosoma is called a „cauda“. Such a cauda
is particularly developed in males of the Arrenurinae (7-12 f), being more or less distinctly
set off from the remainder of the body, often in combination with a small projection (pe-
tiole) located posteriorly.
The four pairs of l e gs (7-13 h-k) are inserted laterally and have six movable segments being
designated from the insertion of the leg as segment 1 (trochanter), -2 (basifemur), -3 (tel-
ofemur), -4 (genu), -5 (tibia) and -6 (tarsus). In water mite literature legs are often num-
bered from anterior to posterior with Roman numerals, and the segments by Arabic nu-
merals. For instance, in this system the tibia of the third leg is referred to as III-leg-5. The leg
tarsi usually terminate in two claws („a m bul a cru m“), a claw-like em p od ium is devel-
oped only in the plesiomorphic Stygothrombioidea (7-16 c), in other groups only remnants
of this structure are found. The ambulacrum can be folded back into a dorsal recess in the
tarsus. Subdivisions of the claws resulting in the development of additional dorsal, ventral
or lateral tips are called clawlets and when these are absent, the claws are said to be simple.
The concave base of the claw may be enlarged („cl a w bl ad e“). The legs are always furn-
ished with setae, which may be long and slender swimming setae (7-13 i). The position of
individual setae on the legs is highly diverse within taxa. The main promotor-remotor
movement is at the coxa-trochanter joint, forming heavily sclerotized articulation condyles.
Swimming by Limnochares americana L UNDBLAD , 1941 was analyzed by B ARR & S MITH
(1979). Leg shape and length, of species of the family Hydryphantidae variously adapted to
swimming or crawling, are compared by M ITCHELL (1957b).
The g n a t h o s o m a (7-3) bears, in the centre of the distal surface, the mouth opening, which
may be directed dorsally, ventrally or (in most cases), anteriorly. In general, the mouth
opening is flanked by two pairs of fine setae. Frilled membranes or collars of fine hair-
like extensions have evolved in several taxa which help to closely attach to the prey during
extraoral digestion (7-14 g-h, 7-25). The paired c he l ic e rae (7-3 c, 7-14 c, k, p) lie dorsally
in longitudinal grooves, flanked on both sides by sockets for the palp insertions. The che-
liceral structure is designed for piercing the integument of prey organisms and consists ty-
pically of a cylindrical basal segment bearing a movable terminal claw. The paired palps
have both tactile and raptorial functions. Typically, they consist of five segments (trochan-
ter, femur, genu, tibia and tarsus), which flex ventrally. In water mite literature they are
generally referred to as „P-1 to P-5“. Absolute and relative measurements of palp segments
are often important for species discrimination ( see 7-3 d for details). The condition of a
palp in which the dorso-distal end of the fourth segment extends beyond the insertion of the
fifth segment is known as a c he l at e palp, a pincer-like appendage (7-3 a-b, key 3 b, i). If P-
4 is ventrodistally expanded to oppose P-5 (7-14 o), then the palp is called su bc he l at e o r
un c ate . The posteriorly directed, paired or unpaired gnathosomal apodemes (7-3 a, 7-14
h: subcutaneous sclerotized projections of the exoskeleton functioning as muscle attach-
ments) have two major functions: (1) they allow for the retraction or changes in the direc-
tion of the mouth organ as a whole. For this purpose, muscles extending between the
gnathosomal base and idiosomal sclerites work antagonistically against changes in haemo-
7. Hydrachnidia 249
Fig. 7-3: a-d, Hydrachnidia, gnathosoma and its appendages in lateral view, example of Thyopsis
cancellata (P ROTZ , 1896) (after D I S ABATINO et al. 2002, modified); a, gnathosoma and left palp;
b, right palp laterally (indicating numbering of segments and traditional segment terminology); c, che-
licera; d, palp indicating important measurement distances (L = length, H = height); e, Hydrachnidia,
schematical lateral view; dashed lines indicate hypothetical segment borders (for abbreviations see
Figs 7-1-2, frontal eye posterodorsally, paired lateral eyes posterolaterally from Dgl-2; setae Ce re-
duced in most taxa, developed only in postlarval instars of Limnochares and Pontarachnidae; after
T UZOVSKIJ 1987).
250 General part
lymph pressure. In some taxa a long protrusable membranous tube connecting the gnatho-
somal base to the anterior idiosoma margin („camerostome“) has evolved which allows for
a tentacle-like activity, or complete retraction into the body, of the whole gnathosoma. (2)
the gnathosoma functions to support the two pairs of oral appendages, and the caudal
apodemes provide attachment sites for the muscles associated with these feeding organs.
As a rule the gnathosoma is narrowed distally from the palp insertions and drawn into an
anterior snout-like projection termed „ro stru m“ (7-14 d). The most comprehensive com-
parative studies of water mite mouthparts were published by M OTAŞ (1928f), M ITCHELL
(1955, 1962), C ASSAGNE -M ÉJEAN (1966a) and Z HAVORONKOVA (1990, 1992, 1993).
Anatomy
A survey of the anatomical organization of a typical water mite is given in Figs 7-4 d-e.
N e ura l Sy ste m a nd S e nse O rg a ns: As in all Acari, ganglia are concentrated in a cir-
cumoesophageal ganglion mass. From here dorsal nerve trunks lead to the anterior sense
organs and mouthparts, and ventral ones lead to the legs and genital region (S CHMIDT
1935). There are typically two pairs of lateral eyes (B AKER 1988) functioning as ocelli:
the mites can detect the intensity and, as evidenced for some taxa (e.g. Unionicola), the
wave-length of incident light. The function of the unpaired median eye or frontal organ,
which is found only in some plesiomorphic taxa, is uncertain. The main ta c ti le re c e p t o r s
on the body and its appendages are specialized setae. Also each g la nd ul ari um is asso-
ciated with a sensory seta. A secretion of unknown composition stored in the gland sacculus
is ejected through a narrow pore as the seta is stimulated. This substance quickly modifies
into a sticky gel after it comes in contact with water and may be a deterrent that acts both
against vertebrate and invertebrate predators (K ERFOOT 1982). Consequently, visual pre-
dation by fishes has selected for enhanced conspicuousness and increased distastefulness in
water mites and not for reduced conspicuousness by increased transparency as e.g in cla-
docerans. In males of Arrenuridae secretions of certain glandularia are obviously modified
and stick the bodies of male and female together during sperm transfer (7-8 b). The five
pairs of slit organs are thought to be p ropr io re ce pt ors (K RANTZ 1978). A variety of
specialized ch em o re ce p tor s is located on the distal segments of the palps (B AKER
1996) and legs. They may be setiform or shortened to peg-like or cup-shaped structures.
The most striking of these organs are the sol en id ia inserted on the three terminal segments
of legs and palps.
A li m en ta ry C an al a nd D ig es tiv e S y ste m : Digestion starts preorally within the prey.
The chelicerae tear a hole in the prey, oral glands (often three pairs) pass secretions into the
wound and then the pharyngeal pump draws in fluid nutrient via the mouth (or buccal
cavity) into the tubular oesophagus. Food then enters the lobed midgut (7-4 d-e) where
digestion and absorption is completed. Undigested material accumulates in cells of the mid-
gut (B ADER 1954, M ITCHELL 1970). Greenish or brownish lobes of the midgut occupy
nearly the entire body. The hindgut is reduced; in a number of genera a small median di-
verticulum can be observed, which is possibly a remnant of the connection between the
midgut and the anal opening. All lobes of the midgut end blindly, and there is no connection
between the gut and the excretory pore (anal pore) (B ADER 1938, 1954).
R es pir at ory S yst e m: Plesiotypically many groups of water mites have paired stigmata,
located in the gnathosoma between the bases of the chelicerae (hence the outdated taxo-
nomic term „Prostigmata“). These stigmata are closed and oxygen uptake functions by
cuticular respiration. In stream-dwelling water mites, which are generally minor in size
and often flattened, the stigmata lead to tracheal trunks, anastomosing into tracheolar tu-
bules. Each trachea has a portion of its length lying directly under the cuticle and one or
both ends of the trachea turn into the body to supply some organs. So, certain areas of the
7. Hydrachnidia 251
ang anterior gut
dvm dorsoventral muscles
ejc ejaculatory complex
exl excretory organ lobes
exp excretory pore
gla glandularia
gop gonopore
ley lateral eyes
mgl midgut lobes
sgl salivary glands
tel testes lobes
Fig. 7-4: a-b, Thyopsis cancellata, ejaculatory complex (after G ERECKE 1996, terminology following
B ARR 1972); a, anterior view; b, lateral view; c, Hydrachna conjecta, spermatophore with sperm sac
(after D AVIDS 1979); d-e, Piona coccinea (K OCH , 1836), schematical anatomy of the idiosoma, d, dor-
sal view, e, lateral view (modified after H ALÍK 1929a).
252 General part
cuticle serve as a respiratory surface. In several standing-water-dwelling species of various

genera the presence of a more complicated respiratory system has been demonstrated, with
numerous tracheoles anastomosing on either side of the brain. As no link between this sys-
tem and the anterior stigmata could be found, it is obviously a closed system (W ILES 1984).
The thick (30-40 lm) integument of Arrenurus spp. has a regular pattern of pits, each lying
under a layer of thin cuticle (2-3 lm thick). In most of the pits a tracheal loop extends and
turns around one to three times (M ITCHELL 1972, R EDMOND 1972, P OPP 1991a,b).
C irc ul a tory Sy st em : As typical for most small arthropods, the circulatory system is la-
cunar, no heart or vessel systems are found. The organ systems are bathed by haemolymph,
which is circulated by movements of the body musculature. The haemolymph not only
functions in the transport of nutrients, hormones, and waste products but also plays an
important mechanical role in supporting tissues and in transferring energy in the form
of hydrostatic pressure.
E xc re to ry S y ste m : The excretory organ consists of a large thin-walled excretory tubule,
apparently derived from the primitive hindgut, and is situated dorsally, in close contact
with the midgut. Waste products are absorbed from haemolymph, and stored in the tubules
as insoluble, whitish or yellowish crystals of unknown chemical composition. When filled,
the excretory tubule may be visible through the dorsal integument as a ’T’- or ’Y’- shaped
structure (7-4 d-e). These shapes indicate an originally paired organization, as is found in
some Hygrobates spp. (S CHMIDT 1935). The excretory tubule connects ventrally with the
excretory pore. C la pa rè de o rg an s („urstigmata“) situated ventrally between legs I and II
in larvae (2-3 f) together with a ce ta bu la (sometimes also called „genital papillae“) of
nymphs and adults are considered to be important sites of osmoregulation. They are water
and/or ion transporting organs for maintaining water balance. All these organs have essen-
tially the same ultrastructure and thus the same function (A LBERTI 1977, 1979). Plesioty-
pically, acetabula are found in adults as three pairs flanking the gonopore, each located free
in the soft integument on a small sclerite ring or stalk (7-1). In many species however, and
obviously as a consequence of numerous independent evolutionary pathways, the aceta-
bula are more numerous. They can be found externally on the plates flanking the genital
opening (7-15 a, c), widely scattered in the soft integument of the venter (7-15 b), or even of
the whole idiosoma (7-15 d). Hydrovolzioidea are unusual in that acetabula are only lo-
cated on the coxae (A LBERTI & B ADER 1990, B ENFATTI & G ERECKE 1999, T UZOVSKIJ et al.
2001); acetabula both in the genital field and on coxae are found in species of the neotro-
pical limnesiid genus Neotyrrellia (G OLDSCHMIDT et al. 1999).
R ep rod uc ti ve Sy ste m : Males have multilobed paired or unpaired testes (7-4 d, see also
B ÜCKING 2001). Each lobule empties via a sperm duct that unites with others in a bran-
ching system, leading ultimately to a pair of long, wide, seminal vesicles lying midway in the
body, which join in an unpaired vas deferens before entering the e ja cu la to ry c om pl e x
(„genital skeleton“). B ARR (1972) described this complex for representatives of many gen-
era and families. It consists of an intricate series of membranous chambers attached to a
sclerotized framework (7-4 a-b) and is situated just inside the genital opening, immediately
above the genital field. The ejaculatory complex functions as a syringe-like organ for com-
pacting masses of spermatozoa, assembling spermatophores (7-4 c), and expelling them
from the genital tract through the gonopore. The structure of the spe rm a top ho re cluster
is described by B ÜCKING (2001) for Piona carnea (K OCH , 1836). The u lt ras tru ct ure o f
sp erm a to z oa was studied by A LBERTI (1991), B ÜCKING (2001) and C ASSAGNE-MÉJEAN
(1966a). In females, a pair of partly fused strongly branched ovaries is spread out between
all organ systems. Eggs are transported through oviducts, which terminate in a single duct
leading to the genital chamber and the gonopore. Spermatozoa may be stored in a re ce p -
ta c ul um s em i ni s, a blind sac of the genital chamber not always present. Females appar-
ently have the ability to store sperm for a long period of time (R IESSEN 1982). Detailed
7. Hydrachnidia 253
descriptions of the female reproductive system are given by S CHMIDT (1935) and M EYER
(1985) for Hydrodroma despiciens (M ÜLLER , 1776). K EYL (1957) and S OKOLOW (1962)
report on the karyology of water mites.
Development
As compared with the plesiotypical life cycle of mites, Parasitengona show a reduction in
the number of free living stages (see also chapter 6). In his historical overview H ENKING
(1882) observed the following stages in water mites: (1) Egg, (2) Schadonophan (inactive),
(3) Larva (active), (4+5) Nymphochrysalis and Nymphophan (inactive), (6) Deutonymph
(active), (7+8) Teleiochrysalis and Teleiophan (inactive), (9) Adult (active). The inactive
stages develop only a few organs such as skin and some acetabula. Stages named with
a term ending in „-phan“ differ from „-chrysalis“-stages in the presence of a fully developed
skin („apoderma“). When this characteristic, often particularly sculptured skin can be dis-
tinguished, the nymphochrysalis has passed to the nymphophan stage. This concept found
its way into water mite literature and is, though rare, still in use.
G RANDJEAN (1938) showed discrepancies in the scheme of H ENKING and he opined that
the stages of Parasitengona could be identified with the original six stages found in other
taxa of the Acari, namely prelarva (= „schadonophan“), larva, protonymph (= „nympho-
chrysalis“), deutonymph, tritonymph (= „teleiochrysalis“) and adult. In the terminology of
G RANDJEAN , the interval between two moults is called a „stase“, inactive stases (in water
mites prelarva, proto- and tritonymph), are called „calyptostases“. As an exception adults
of certain Limnocharinae may undergo supernumerary moults (B ÖTTGER 1972a).
E g gs are generally deposited in clutches of various shapes surrounded by a gelatinous ma-
trix and attached to a substrate (e.g., C ASSAGNE -M ÉJEAN 1966a), but deposition of single
eggs has been observed in small species of several genera (M ARTIN pers. comm.). The pro-
tective envelope of the eggs displays rapid swelling in water (S OKOLOW 1925c, 1977). Hy-
drachna females have evolved an extrusable ovipositor and deposit their eggs, which lack
such an envelope, into the air cavities of water plants, thus providing an oxygen supply and
protection. Generally, females of unionicoline species use stylet like setae situated along the
midline of the genital field for depositing eggs in the tissues of sponges, mussels or gastro-
pods. Within the egg, a minute, non-hatching prelarva is formed. Further details of em-
bryonic development are described by M EYER (1985, 1989) for Hydrodroma despiciens.
One single case of vivipary is reported from Ignacarus salarius G ERECKE , 1999, a species
only known from a hypersaline spring in Spain.
The six legged la rv a (2-3 f) emerges from the thin prelarval skin and is dissimilar to the
other free-living stages. It is nearly always parasitic (7-5 a-d, 7-9). The short gnathosoma is
built up as in deutonymphs and adults, but in newly hatched larvae proportionally larger
than the idiosoma and with its appendages adapted to attaching to, and piercing through,
the host cuticle. Two pairs of lateral eyes are situated in the dorsal integument of the idio-
soma, arranged close together or enclosed in a capsule on each side.The ventral side of the
idiosoma bears three pairs of coxal plates, variously fused to each other and occasionally
merged to a coxal shield. A pair of u rsti g m ata (in rare cases doubled or multiplicated,
acting as acetabula) is generally situated in the interspace between coxal plates I and II.
Three pairs of legs insert laterally on the coxal plates. In the Hydryphantoidea and Eylaoi-
dea they are six-segmented, those of all other groups have two segments fused. The seg-
ments have characteristic complements of setae and solenidia which are found more stable
than in deutonymphs and adults and thus useful for resolving taxonomic problems (T U-
ZOVSKIJ 1987). Larvae of certain more derivative groups are adapted for swimming using
elongate and plumose setae on the three terminal leg segments. S PARING (1959), P RASAD &
C OOK (1972), WAINSTEIN (1976c, 1980), I.M. S MITH (1982, 1984), M ARTIN (2000, 2003)
254 General part
and G ERECKE & M ARTIN (2006) give reviews of larval morphotypes with diagnoses and
keys for a wide variety of species. The size, shape and position of sclerites and appendages
in combination with the chaetotaxy provide useful taxonomic characters. Occasionally,
species recognition is more easy with larvae than with postlarval stages (WAINSTEIN
1980). For reasons of limits in space and time, the larval stage is not treated in this
work, but available references to special bibliography are given for each taxon.
As mentioned before, pro to- and tri ton ym p h are inactive „pupal“ stages (2-2). They are
both poor in particular morphological characters, but attribution to genera and species is
often possible thanks to visible elements of the preceeding or the following stage. Sclerites
and appendages of the preceeding stage may remain attached to the outer surface, devel-
oping structures of the following stage become visible through the „pupal“ skin. After en-
gorging on host fluids the larva transforms by a moult, but without leaving the larval exu-
via, into a protonymph (2-2 a). This may happen when the larva is still attached, or after
leaving the host and continuing development in suitable microhabitats.
The deu ton ymp h (2-4 g) develops within the skin of the protonymph (2-2 a) and for
hatching it must rupture two skinlayers, that of the larva and that of the protonymph.
The predacious deutonymph is very different from the larva and morphologically similar
to the adult (compare 2-4 f, 2-4 g). As it is sexually immature, there is a so called „p ro vi -
si ona l g e ni ta l fi e ld“ without a gonopore, and the genital plates bear an incomplete
number of acetabula, in many cases just two pairs. Furthermore, it differs from adults
in generally minor dimensions, lower numbers of setae and less extended idiosoma scler-
otization - in several cases (e.g. Arrenurus), species completely sclerotized at the adult stage
may have deutonmyphs with a smooth skin. With rare exceptions, no sexual differences are
found at the deutonymphal stage - in any case, no secondary sexual characters are devel-
oped. T UZOVSKIJ (1990) gives a survey of the morphology of deutonymphs in several taxa.
In the work presented here, references to deutonymphal descriptions are given for each
taxon. In many cases, with some experience, it will be possible to attribute deutonymphs
to genera and species using our keys for adults.
The predacious deutonymph transforms following a growth period into the motionless and
inactive t rit ony m ph al st ag e (2-2 b). The morphological change from deutonymph to
adult is much less extreme than that from larva to deutonymph. As far as is known, most
adults feed on the same prey as deutonymphs, but in exceptional cases (Eylais hamata K OE-
NIKE , 1897, see G ERECKE 1991), differences in the shape of mouth parts suggest a different
diet of these two stages.
Food and feeding

L a rva e : Immediately after hatching, the larva starts seeking out an appropriate host
(usually an insect, in exceptional cases an amphibian, there are no records of parasitism
on any other groups of vertebrates or invertebrates, see below) and becomes an ectopar-
asite. After attachment of a larva to its host a feeding device is produced, the sty l osto m e,
extending into the tissues of the host. Two main types are recognized: a simple tubule or
blind sack (7-5 d) and a ramified tubular complex (7-5 e-f) (B.P. S MITH 2003).
The simp le t ubu le or bli nd sack (7-5 d) is reported for larvae of species of Arrenurus
(ÅBRO 1979; R EDMOND & H OCHBERG 1981) and Unionicola (H EVERS 1978a), but with-
out doubt, many other species will develop a similar type of stylostome. This closed type
differs from those of larval trombidiid mites, parasitic on mammals and birds, where the
axial canal is kept open to the liquified host tissue and appears to function as a true sucking
tubule (V OIGT 1970). Most studies on tubular stylostomes were made by ÅBRO (1979,
1982, 1984, 1992) on Arrenurus larvae. The pedipalp claws grasp the membranous cuticle
of the host and the distal sabre-like cheliceral segments tear it. Chelicerae and palps co-
7. Hydrachnidia 255
operate to anchor the larva. During puncturing, the chelicerae move alternately, each in
turn forming a sliding support for the other. The larva is presumed to inject venom
into the wound, in which a subcuticular vesicle appears, serving as a buffer zone smoothing
the way for the nascent stylostome and protecting it from direct defensive reactions of the
host. Epidermal cells, adjoining the primary vesicle, undergo lysis and fuse with the vesicle,
transforming it into an epidermal abcess, whose cell juice acts as a pool of nutrient for the
larval mite. Subsequently, the larva ejects a liquid that rapidly gels and forms a slender
resilient blind sac. The stylostome increases in size, elongating and remodelling as larval
engorgement proceeds and may grow up to a length of 300 to more than 1.000 lm (H EVERS
1978a; R EDMOND & H OCHBERG 1981; ÅBRO 1979). The fully formed stylostome of a
Fig. 7-5: (a-c) larvae of Hydrachna skorikowi P IERSIG , 1900 attached to, and engorging on, a leg of
their insect host (by courtesy of M ICHAEL S TEVENS , Düsseldorf); d, fragment of body wall of a zy-
gopteran imago, cemented to it a fully formed stylostome, whose blind end is a smooth dilatation
(after ÅBRO 1982); e-f stylostomes of Hydrachna conjecta in two different development stages; e,
length of larva 640 lm; f, length of larva 1700 lm (after D AVIDS 1973a).
256 General part
given Arrenurus sp. is constant in shape, recognized to be a product of the mite, not the host
(L ANCIANI & S MITH 1989, ÅBRO 1992). Uptake of host’s tissue fluid seems to take place
mostly through the wall of the terminal bulb which is subject to dilatation and contraction
by pharyngeal movement during active feeding (ÅBRO 1984, 1992). Within three hours
after anchorage, larvae exhibit clear signs of expansion (ÅBRO 1982). The stylostome re-
mains in the host after the larva drops off (see also „parasitism“ and „life history“).
In contrast to the tubular type, r am i fi ed sty lo sto m es (7-5 e-f) are inconsistent in form.
They are characteristic for larvae of the genus Hydrachna, parasitizing water bugs (D AVIDS
1973a) and water beetles (B LUNCK 1923). This type is also found in various terrestrial ar-
thropods parasitized by the larvae of the terrestrial Trombidium holosericum L INNAEUS ,
1758 (P FLUGFELDER 1970). D AVIDS (1973a) described the structure and development of
the stylostome of Hydrachna conjecta K OENIKE , 1895. The main channel ramifies into
numerous smaller channels. While the larva is growing the stylostome increases in size
and it is shown that the bore of the tubes increases with the growth of the stylostome
as a whole. It appears that stylostomes associated with mite larvae of the same length
in summer remain smaller (short parasitic phase) with fewer ramifications than those pro-
duced in winter. It seems likely that the stylostomes of Hydrachna spp. result from coag-
gulation of the haemolymph as a reaction to the saliva of the mite larva (P FLUGFELDER
1970, D AVIDS 1973a). Minute papillae give the stylostome a frothy appearance (D AVIDS
1973a). There is a strong similarity to the papillae on the terminal bulb of the arrenurid
stylostome (ÅBRO 1979). Most stylostomes seem to thwart the ability of the host to recog-
nize them as foreign bodies (ÅBRO 1982). The host’s defence appears as a two-step reaction:
1. initial haemolymph clotting and deposition of a melanin sleeve around the stylostome at
the penetration site, 2. occasional subsequent melanization and cellular encapsulation of
the stylostome. But, these defence reactions appear to be casual and ineffective in many
cases. However, when a Hydrachna conjecta larva attaches to the corixid Sigara falleni
(F IEBER ) the bug’s haemolymph reacts so strongly to the saliva that the mites’ mouthparts
become obstructed, the attachment site becomes necrotic and subsequently the attached
larva dies and shrivels (D AVIDS 1973a). The same effect was found by R EILLY & M C C AR-
THY (1991) with Sigara distincta (F IEBER ) as host.
De ut ony m ph s a nd a du lt s: Both active postlarval instars are predators (B ÖTTGER

1970a), parasitism (see below) is an exception. In a number of species of the genera Lim-
nesia and Piona cannibalism has been observed. In general, food categories are crustacean
zooplankton, ostracods, insect larvae, insect eggs and occasionally mite eggs. It is clear that
water mites are neither plant feeders nor scavengers: only very freshly dead invertebrates
may become part of the diet of certain species (G ERECKE & I.M. S MITH 1994).
When P ROCTOR & P RITCHARD (1989) surveyed the feeding of water mites, available in-
formation was mostly restricted to stagnant water species. Additional information for
stream dwelling taxa is available from M ARTIN (2005). U CHIDA (1932) reported that
the diet of Hydrodroma despiciens consists of dead aquatic insects, but W ILES (1982)
proved that this species predates upon chironomid eggs. In addition to H. despiciens,
egg predation is also known in many other taxa: Hydrachna spp. feeding on corixid
eggs (D AVIDS 1973a, S TEVENS & G REVEN 1999) (7-6), Acerbitas spp. (the name introduced
by Ö ZDIKMEN (2006) replacing the preoccupied genus name Thyas) on mosquito eggs
(M ULLEN 1975b), Hydryphantes spp. on dragonfly eggs (L ANCIANI 1979c), Limnesia
maculata (M ÜLLER , 1776) on trichoptera eggs (Z HAVORONKOVA 1993) and some species
of Hydryphantidae and Hydrovolzia placophora on water mite eggs (M ARTIN 1998,
2005). Egg predators are probably attracted to the surface chemicals of the gelatinous coat-
ing of the eggs (W ILES 1982). Eylais spp. feed on cladocerans and, more rarely, on ostracods
(B ÖTTGER 1962, S CHEIDEGGER 1999). Certain Limnesia and Piona spp. have a wide food
spectrum, they take microcrustaceans (cladocerans, copepods, ostracods) and insect larvae
7. Hydrachnidia 257
Fig. 7-6: Adult Hydrachna skorikowi feeding on eggs of Sigara striata (by courtesy of M ICHAEL
S TEVENS , Düsseldorf).
(ephemeropterans, chironomids, mosquitoes) or even trichopteran eggs. Others are more

restricted to chironomid and blackfly larvae (Sperchon, Lebertia and Hygrobates spp.,
M WANGO et al. 1995, M ARTIN 2005) or ostracods (Arrenurus spp.; Chelomideopsis an-
nemiae R OMIJN , 1920, M ARTIN 2005). Water mites feeding on zooplankton mostly prefer
cladocerans to copepods. Rotifers are selected by the Australian Physolimnesia australis
(H ALÍK , 1940), which predates on Brachionus spp. (P ROCTOR 1997a).
The pattern of prey selection for adults and nymphs appears to depend primarily on the
escape response of the prey. Water currents produced by a moving prey influence the pre-
dator’s behaviour through mechano-reception (M ITCHELL 1962). This is probably the most
important mode of detection. Contact chemoreception is possibly used to determine the
palatability of an already captured prey item (P ROCTOR & P RITCHARD 1990a). C la do -
c e ran s are heavily preyed upon by water mites. Their small size and the lack of a fast
escape reaction leaves them extremely vulnerable (PAVELJEVA & Z ANKAI 1971, S CHEIDEG-
GER 1999). O th e r m ic roc ru sta ce a are preyed upon to a by far lower degree due to their
effective protective (o stra c oda) or escape (c o pe po da) behaviour. D AVIDS et al. (1981a)
observed that an ostracod colliding with, and embraced by the legs of a Piona species, im-
mediately shut its valves and slipped away out of the embrace like a wet piece of soap.
However, representatives of several groups have evolved special tactics to overwhelm mem-
bers of these groups: Arrenurus spp., which have ostracods as their preferred food, try to get
hold of the ostracod’s antenna by means of the uncate palp scissors or to re-open closed
valves with the help of its first two pairs of legs (7-7). Overpowering takes several minutes
after which the antenna is pierced by the chelicerae and the prey’s body fluids sucked out
(M OTAŞ 1928f, B ÖTTGER 1962, D AVIDS et al. 1981a). The long-legged Unionicola cras-
sipes (M ÜLLER , 1776) prefers Cyclops as food and shows two types of specialized catching
behaviour. In the first case, observed also in Neumania spp., the mite supports itself on the
bottom by means of its two pairs of hindlegs, while the two front pairs are raised from the
substratum, held outstretched and facing towards vibrations in the water column (7-7 a).
The mite’s body, palps and legs form a kind of basket, and when a prey enters this basket
and touches the legs or palps, they all contract immediately (P ROCTOR & P RITCHARD
1990a). The other method is used when the mite is swimming. It moves in little jerks
through the water and after several strokes it sinks, the legs are spread out like the ribs
of an umbrella and contract whenever they touch a prey. A prey embraced by the long
legs covered with spines and bristles has great trouble to free itself (H EVERS 1980, D AVIDS
et al. 1981a). Another „sit - and - wait predator“ is Sperchonopsis verrucosa (P ROTZ ,
1896), which waits for dipteran larvae to crawl underneath it (U LLRICH 1978). C hi ro-
no m id la rva e are a substantial part of the diet in benthic species, e.g. for the spring-
and stream-dwelling water mite species of the genera Sperchon, Lebertia and Torrenticola.
258 General part
Fig. 7-7: a, Unionicola crassipes in net-stance posture, orienting to the vibrations of a swimming
Daphnia (after P ROCTOR & P RITCHARD 1990); b, Arrenurus maculator (M ÜLLER , 1776) feeding
on an ostracod (after M OTAŞ 1928f); c, Limnochares aquatica attacking a tube-dwelling chironomid
larva (inspired by Z HAVORONKOVA 1990)
7. Hydrachnidia 259
Several taxa which in view of their small dimensions are potentially suitable as prey (Oli-
gochaeta and other Hydrachnidia, larvulae of Ephemeroptera, Trichoptera and some dip-
teran families) were ignored by nearly all observed mites (M ARTIN 2005). Most chirono-
mid larvae are tube-dwelling where they are less vulnerable (PATERSON 1970), but Lim-
nochares aquatica is able to attack them by piercing the head capsule projecting from the
tube (7-7 c, B ÖTTGER 1972a; Z HAVORONKOVA 1990). T EN W INKEL & D AVIDS (1985a)
report that bioturbation affects the food availability for water mites: benthic feeding bream
(Abramis brama L INNAEUS ) suck sediment into the mouth and select the large chironomids
for food. Small chironomid larvae such as Cladotanytarsus spp. pass through the gills of
bream, but being disturbed, they leave their sturdy tubes of sand grains. By these actions
their prey vulnerability is greatly increased in favour of water mites of the genus Hygro-
bates. The authors estimated that Hygrobates reduced the density of the chironomid Cla-
dotanytarsus sp. by more than 50 %, the numerical decline closely matched the population
food-requirements of the water mite between November and May (T EN W INKEL et al.
1989). G LIWICZ & B IESIADKA (1975) and M ATVEEV & M ARTINEZ (1990) suggest that
Piona sp. in North America can cause a summer depression in Daphnia populations of
about 50 %. Other studies indicate that water mite predation has a significant impact
on prey populations such as corixids (D AVIDS & S CHOOTS 1975, P IECZY N ~ SKY 1976)
and chironomids (PATERSON 1970).
In a number of water mite species, all active instars affect the same insect population; larvae
as parasites, deutonymphs and adults as predators (D AVIDS 1973a, M ULLEN 1975b, L AN-
CIANI 1978). There is evidence that in Hydrachna conjecta the decrease in fitness of a cor-
ixid population is lowered more by egg predation by deutonymphs and adults than in poor
reproduction caused by parasitic larvae (D AVIDS 1991). Most invertebrate predators are
considered to be size-dependent. Consequently, size differences of predators will involve
differences in the food niche. Also feeding time is directly proportional to the size of
the prey item (R IESSEN 1982a). On the other hand, temperature has a great influence
on the predation rate in invertebrate predators (B UTLER & B URNS 1991) and as a conse-
quence on egg production. B UTLER (1993) analyzed the predation rate of Piona exigua
K.V IETS , 1949 at different temperatures, prey densities, prey size, and prey species. In Un-
ionicola crassipes searching time was considerably longer at 12 C compared with 24 C.
The same temperature effect holds true for the handling time (attack time + feeding time)
(D AVIDS et al. 1981a ).
Terrestrial Parasitengona and their derivative water mites are predominantly red and it has
been observed by several authors that water mites are seldom preyed upon. Fish, when
mouthing a mite, obviously experience noxious chemicals (R IESSEN 1985). Red mites
are particularly distasteful and fish quickly learn to reject them as potential prey (K ERFOOT
1982). However, water mites are not distasteful to all fish, but their contribution to the food
of these fish is small and the effect of fish predation on mite populations is negligible (P IEC-
ZY N~ SKI 1976; P IECZY N~ SKI & P REJS 1970). A strong positive correlation was noted by
E RIKSSON et al. (1980) between lakes that contained fish and Piona carnea. In lakes with-
out fish, P. carnea populations were kept at low densities because of the predation pressure
on P. carnea by aquatic insects.
Sperm transfer
Among water mites all possible modes of sperm transfer can be observed from substrate
deposition of spermatophores with complete dissociation between male and female to in-
direct or direct copulation. Indirect copulation is generally done via an intromittent organ
(generally a modified leg). P ROCTOR (1992a) synthesized studies of sperm transfer and
spermatophore morphology, W ITTE & D ÖRING (1999) give a clear insight into the con-
260 General part
straints and pathways in the evolution of water mite sperm transfer. One of the most strik-
ing features within the Hydrachnidia is that indirect and direct copulation have evolved
convergently in several groups (S CHWOERBEL 1962d, W ITTE 1991, P ROCTOR 1991a,
1992a) and species belonging to the same genus may differ in the mode of direct or indirect
sperm transfer (WAINSTEIN 1976c).
S ubst ra tum - de po sit ed spe rm a to pho re s and subsequent pick-up by the female are
found, in addition to the Acari, also in Pseudoscorpiones, Scorpionida, Symphyla and Col-
lembola. It is conceivable that this phenomenon evolved polyphyletically in land-living an-
imals inhabiting biotopes with a high rate of humidity so that spermatophores did not de-
siccate. In water mites it is a remnant from terrestrial ancestors, little suited to aquatic life
and can only be maintained by those which move in close contact with the substratum.
Males deposit spermatophores for subsequent retrieval by conspecific females. Attraction
by means of diffusing pheromones will have favoured the evolution of fields of sperma-
tophores to which several males may contribute, as is found in many water mite groups
(W ITTE 1991). A single male of Piona carnea may contribute 14-18 per cluster (B ÜCKING
2001). In other cases the presence of females appears to stimulate males to deposit sper-
matophores. In some Eylais spp. males actively assist females to find the spermatophores,
by leading them in a circular dance over the deposited spermatophores (L ANCIANI 1972).
A transfer with no contact between the sexes has been observed for a number of
species from earlier derivative taxa: Hydrachnoidea (D AVIDS & B ELIER 1979), Eylaoidea
(L ANCIANI 1972), Hydryphantoidea (M ITCHELL 1958b, M ULLEN 1977, W ILES 1982;
M EYER 1985), Lebertioidea (E FFORD 1966) and Hygrobatoidea (H EVERS 1978, P ROCTOR
1992a).
As adaptations to in di rec t spe rm t ran sfe r, in several taxa of the Hygrobatoidea the
tarsi of the third pair of legs are modified into spermatophore-transfer structures (gono-
pods). The spermatophores are carried between the tips of the tarsal claws of the flexed
third legs (K.V IETS 1914f, U CHIDA 1932, M ITCHELL 1957c, B ÖTTGER 1962, P ROCTOR
1997a; B ÜCKING 2001). In Piona spp., the male uses specially modified appendages (usual-
ly the fourth pair of legs) to hold the female in a characteristic posture so that spermato-
phores can be transferred to the gonopore of the female by simply extending his third pair
of legs (7-8 b-c). Mating can last up to more than three hours. (B ÜCKING 2001). Excep-
tionally, in Unionicola intermedia (K OENIKE , 1882) the fourth instead of the third leg is a
gonopod (H EVERS 1978). The most complex modes of indirect sperm transfer occur in the
Arrenurinae (7-8 a). In males the posterior part of the idiosoma is modified and forms a
cauda with characteristic, species specific modifications in shape and setation. A receptive
female mounts a male, positioning herself so that her gonopore rests over the back edge of
the male cauda. Secretions from glandularia on the cauda of the male cement the pair to-
gether and the male then deposits one or two stalked spermatophores on the substratum. In
some species, the male then moves his body down and forward to bring the gonopore of the
female into contact with the spermatophore. Next the whole process starts again, lasting up
to three to four hours in which time about six spermatophores have been transferred. In the
intervals between spermatophore transfer, the couple is well synchronized in leg move-
ments and able to perform fast crawling and effective swimming. In the last phase the
male pushes the female away with his fourth pair of legs (L UNDBLAD 1929c, B ÖTTGER
& S CHALLER 1961, B ÖTTGER 1962, C ASSAGNE -M ÉJEAN 1966a, P ROCTOR & S MITH
1994). Males with a petiole, a cup-shaped posterior appendage of the cauda, use this organ
to scoop up spermatophores and insert these into the gonopore of the female (B ÖTTGER
1965). Also, in representatives of other superfamilies, modifications of male legs (probably
functioning as grasping organs during indirect sperm transfer) have evolved. Examples are
the second leg of Nilotonia species of the subgenus Dartia (Lebertioidea), and the fourth leg
of species of Ignacarus and Diplodontus (Hydryphantoidea) and Acherontacarus (Hydro-
7. Hydrachnidia 261
Fig 7-8: Spermatophore transfer in water mites; a, Arrenurus globator spermatophore transfer (after
L UNDBLAD 1929); b-c Piona; b, male extracting a spermatophore from its genital pit with the help of
the III-L (modified after H ALÍK 1929); c, male using III-L for spermatophore transfer to the female
gonopore (modified after B ÖTTGER & S CHALLER 1961); d, Midea orbiculata, copulation (the male is
smaller than the female - after L UNDBLAD 1929b).
262 General part
volzioidea). Sexually dimorphic palps, as often found in taxonomically unrelated taxa,

probably indicate that these appendages in males are also involved in allocation and/or
transfer of spermatophores.
Di re ct spe rm a top hor e t ran sfe r by alignment of the genital openings developed inde-
pendently in many clades of water mites (7-8 d). Males may have the gonopore reduced in
diameter and transversely slit shaped (some species of Hydrachna, WAINSTEIN 1976c), pro-
truding from the venter on a cone-shaped projection (some species of Eylais, B ÖTTGER
1962, L ANCIANI 1972), or are equipped with wing-like appendages flanking the gonopore
to facilitate transfer as in Midea orbiculata (M ÜLLER , 1776) (L UNDBLAD 1929b). The
genital plates of Rutripalpus limicola S OKOLOV, 1934 males are fused forming a penis-
like hinged tube unique in water mites (G ERECKE & T UZOVSKIJ 2001). L EIMANN
(1991), W ITTE & D ÖRING (1999) and B ÜCKING (2001) described spermatophore charac-
teristics. In stalked spermatophores, the stalk is mostly continued into two membranes
clasping the sperm sac (7-4 c). During a few days hundreds of spermatophores can be pro-
duced in Hydrachna conjecta, whereby deposition takes a few seconds (D AVIDS & B ELIER
1979). An Unionicola sp. produces up to 35 spermatophores a day (H EVERS 1978).
Parasitism
Larvae
The first illustration of an attached water mite larva (apparently a Hydrachna sp. on a Nepa
sp.) was given by S WAMMERDAM (1637-1680) (published by B OERHAVE in 1737) (7-9 a), a
follower of the „ovism theory“. He was amazed that an „egg“ was able to increase in size in
that way. It is a remarkable fact that nearly all parasitic associations of water mite larvae
detected since then have been established with insect hosts (for a critical review discarding a
series of unsupported older data see I.M.S MITH & O LIVER 1986, further information in
B.P. S MITH 1988 and R OLFF 2001). L INDQUIST (1975) suggests that mite-insect associa-
tions may be as old as 100 million years. This is ample time for co-evolution of the mites
and for gaining a strategic advantage in their relationships. No such relationships have
evolved with any other group of limnic invertebrates such as turbellarians, rotiferans, gas-
trotrichs, nematodes, crustaceans or molluscs (see below for the particular interrelations
between postlarval water mites and sponges, bivalves and gastropods). The fact that aqua-
tic vertebrates are also generally ignored, highlights the particular significance of the phore-
tic aspect in the relationship between water mite larvae and their hosts. Selection of parti-
cular insect hosts guarantees not only larval feeding and engorgement, but also for aerial
transport and the return to a microhabitat suitable for feeding and reproduction of post-
larval stages. In view of the well investigated documented ichthyological parasitology, fish
can be generally excluded from the spectrum of potential hosts of water mite larvae. Ex-
ceptions are made only by the occurrence of larvae of four water-mite genera on fry and
fingerlings of several species, as well as transforming unionicolid larvae encapsulated in the
walls of the pharynx and oesophagus of adult whitefish, both reported by B YKHOVSKAYA -
PAVLOVSKAYA (1962). These observations have never been corroborated by other authors.
A remarkable further case of parasitism on vertebrates has been detected recently: larval
Thermacarus andinus M ARTIN & S CHWOERBEL , 2002 from thermal springs in Bolivia
parasitize the amphibian Bufo spinulosus W IEGMANN , 1835 (S CHWOERBEL 1987). In a
small number of species from widely divergent groups, the larvae are suppressed in their
parasitism (B.P. S MITH 1998, see below).
Generally, only insect imagines are infested, i.e. species living in the water column at the
adult stage, or aerial insects with aquatic larval stages. In a few exceptions, parasitism on
immature insects is reported: Limnochares aquatica larvae attach to Hemiptera nymphs
and initiate engorgement, affecting growth and instar length of the host. When the host
7. Hydrachnidia 263
Fig. 7-9: Parasitism of water mite larvae; a, Hydrachna sp., larva (g) on Nepa (S WAMMERDAM in
B OERHAVE 1737); b, Stictochironomus sp. infested with larvae of Unionicola aculeata (K OENIKE ,
1890); c, Arrenurus cuspidifer P IERSIG , 1894 larvae under the wing sheaths of a naiad of Ischnura
sp. (after C ASSAGNE -M ÉJEAN 1966a).
moults, mite larvae detach, retain their mobility and re-attach to the next instar to continue
engorgement (B ÖTTGER 1972a, B.P. S MITH 1989c). Larvae of several Stygothrombium spe-
cies initiate their parasitic phase on late larval instars of stone flies (M OTAŞ 1928). Arre-
nurus globator (M ÜLLER , 1776) larvae have a low host specificity and may parasitize larvae
of the beetle genus Dytiscus (B ÖTTGER 1962, B ÖTTGER & M ARTIN 2003). Probably as an
exceptional case, R ENZ et al. (2004) report on the nearly total loss of a simuliid population
264 General part
following intense feeding by Sperchon setiger T HOR , 1898 larvae on host pupae. Normally,
larvae of this species show preparasitic attendance and feed on the adult insect only. Re-
presentatives of nearly all insect groups with at least one aquatic developmental stage can
become hosts of water mite larvae. The only orders for which no data are published are
ephemeropterans (probably not suitable due to their short adult life and extra moult at the
adult stage) and lepidopterans (a group with maximum diversity of aquatic species in little
studied parts of the globe). Water beetles are of particular significance for several early
derivative clades such as Hydrachnidae, Eylaidae, Piersigiidae, Apheviderulicidae, Acher-
ontacaridae and Hydryphantidae (B ENFATTI & G ERECKE 1998, G ERECKE et al. 1999, T U-
ZOVSKIJ et al. 2001, M ORENO et al. 2004, Z AWAL 2002, 2003). But from a quantitative
point of view, dipterans, especially chironomid midges, have by far the largest importance
as hosts of water mites (I.M. S MITH & O LIVER 1986).
After hatching, larvae of all species so far studied must locate a host within one or two
weeks, otherwise they will perish (E LLIS -A DAM & D AVIDS 1970, S TECHMANN 1978a). Lar-
vae of Hydrovolziidae, most Eylaoidea and Hydryphantoidea reach the water surface by
positive phototaxis, break through the surface film and there run in search for potential
hosts (U LLRICH 1978, B ÖTTGER 1962, M EYER 1985). The movements of the larvae appear
random in general, but Limnochares aquatica larvae respond to shadows (B ÖTTGER
1972a) and for a few species, it is evident that their larvae can detect potential hosts
over a short distance, a water-borne chemical cue is most likely (E LLIS -A DAM & D AVIDS
1970, B.P.S MITH & M C I VER 1984a). Larvae of Protzia, Acerbitas and Euthyas species are
able to jump several centimeters (L UNDBLAD 1927c, M ULLEN 1977, U LLRICH 1978), other
hydryphantoid larvae attack dipterans when they emerge from their pupal stage or when
ovipositing. Eylais larvae enter the subelytral space of water bugs and beetles when they
come to the surface to replenish air supplies. The apotypic adaptation of searching for hosts
not at the surface, but within the water column, evidently developed several times inde-
pendently in Hydrovolzioidea, Hydrachnoidea, Lebertioidea, Hygrobatoidea and Arrenur-
oidea. Within a mix of dipteran pupae representing various families, Nilotonia longipora
(WALTER , 1925) larvae are able to select, and attach exclusively to, ceratopogonid pupae,
probably attracted by chemical cues (G ERECKE & I.M. S MITH 1996). In contrast, U LLRICH
(1978) showed that simuliid pupae were discovered by Sperchon setiger larvae only by
accidental encounter. The respiratory filaments of simuliid pupae differ in morphology
in the diverse species. As a consequence, entry into the pupal case for water mite larvae
is, in certain simuliids, more obstructed than in others. Also, the way of attachment of
the simuliid case to the substratum can play a part in obstructing entry. Therefore G LED-
HILL et al. (1982) suggested that pupal morphology may be the most important factor in the
selection of host simuliids by Sperchon larvae.
Larval water mites regularly parasitize 10-50 % and sometimes up to 70-100 % of the in-
dividuals in natural populations of their host insects (M EYER 1985). However, at least in
the species belonging to early derivative families which often produce very high numbers of
eggs, the probability of finding a host and then successfully returning to water is apparently
low. M EYER (1985) calculated the host-finding success of Hydrodroma despiciens larvae as
about one percent only. B RUCE P. S MITH & M C I VER (1984a) observed in Arrenurus larvae
in the single step from pupae to imaginal mosquitoes, a transfer loss of more than 50 %.
Ceratopogonid midges ecdyse in ten seconds - so Arrenurus larvae have little time to select
attachment sites before the fly attempts to dislodge them (M ITCHELL 1998). On odonates,
however, arrenurid larvae have 10 to 20 minutes for loading and exhibit precise loading
behaviour as they attach to the host (M ITCHELL 1969). Larval Unionicola spp., Atractides
spp. and Hygrobates spp. eliminate this risk by piercing the integument of the pupa into the
cuticle of the developing imago and are dragged through the pupal skin when the imago
emerges (B ÖTTGER 1972b, U LLRICH 1978, H EVERS 1980). Heavy loads can prevent moult-
7. Hydrachnidia 265
ing because the midge lacks the strength to pull so many mites through its cuticle. The
engorgement period is very different from taxon to taxon, and within one species highly
dependant on temperature. For instance, Hydrachna cruenta needs 58 days at 15 C, but
only 7 days at 25 C (H ARRIS & H ARRISON 1974).
Larvae of many species exhibit a low hos t s el e ct iv it y (generally representatives of one
insect family are preferred by a certain mite species), but a high selectivity for attachment
sites on the host’s body (B ÖTTGER 1962, C ASSAGNE -M ÉJEAN 1966a, E FFORD 1966, D AVIDS
1973a, H EVERS 1978a, S TECHMANN 1980, B ENNETT & S CUDDER 1997, M ARTIN 1999,
2004, M ARTIN & S TUR 2005). These preferences illustrate another adaptive process that
has permitted mites to coevolve successfully with their hosts. Re so urc e pa rti on in g is
possible because the mites remain segregated by the precise site of attachment. Only the
larvae of some early derivative taxa (e.g. Hydrovolzia placophora and certain species of
Hydryphantidae) exploit a wide range of host species from different insect orders and at-
tach rather indiscriminately to various parts of the body of the host (E FFORD 1966, M UL-
LEN 1977, U LLRICH 1978, M ARTIN 2000). Most other species are more selective and will
not respond to insects outside of their normal observed spectrum (C ASSAGNE -M ÉJEAN
1966a, S TECHMANN 1978a). In general, larvae of Hydryphantoidea, Lebertioidea and Ar-
renuroidea parasitizing Diptera prefer thoracic sites (S TECHMANN 1978a), while Hygro-
batoidea larvae exclusively select the abdomen (7-9 b). Larvae of Eylais and some Hydrach-
na spp. select „terrestrial“ attachment sites bathed by air supplies under the elytra of aqua-
tic bugs and beetles. The differential parasitism of related host species in nature is not only
the result of actual preference by the mite, but also of par asi te av oi da nc e behaviour and
host location in its micro-habitat (S MITH & M C I VER 1984b). Zygopteran naiads rid them-
selves of attendant mites by more active crawling and by vigorously rubbing the body with
the legs or tip of the abdomen (F ORBES & B AKER 1990). However, this anti-mite grooming
activity places them at greater risk of predation from visually hunting fish (B AKER & B.P.
S MITH 1997). Mesoveliid bugs limit infestations by catching and eating the larval mites
(L ANCIANI 1985). Female mosquitoes often bear more parasitic larvae of the genus Arre-
nurus than males do (S TECHMANN 1980; B.P. S MITH & M C I VER 1984a, 1984b). However,
laboratory studies failed to demonstrate sexual bias in host selection (B.P. S MITH 1988).
Water mite larvae are not randomly distributed over the individuals of their host popula-
tions, but often show a highly ’clumped’ or ov e rdi spe rse d d ist rib ut ion. This may in-
dicate several waves of infection or that the parasite kills heavily infected hosts. The at-
tachment frequency of freshly attached parasitic larvae mostly fits a negative binomial dis-
tribution. The calculated parameters of the truncated form of the negative binomial dis-
tribution predicted far more heavily infested hosts than were observed in Anopheles sp.
(L ANCIANI & B OYETT 1980). This indicates that heavily infected hosts disappear from
the population. Another possibility of a poor fit may be an indication of mixing among
groups with different levels of infection (B.P. S MITH & M C I VER 1984b).
How much e ne rg y does a larva gain through parasitism? Depending on the species, a larva
may increase its volume from 3-600 fold (7-5 a-c). For some Eylais spp. larval nutrition
provides up to 45 % of the adult body mass (L ANCIANI 1971). In water mite species para-
sitic on odonates, commonly more than 50 larval mites per host are encountered (B.P.
S MITH 1988). These larvae may undergo an 80-90 fold increase in volume (M ÜNCHBERG
1938a) or 27 times increase in dry weight (M ITCHELL 1969b). Dipterans have little space
for such a large number of parasites, a maximum of 10-20 is normally observed. Arrenurus
spp. parasitic on dipteran hosts increase 7-16 fold in volume (M ÜNCHBERG 1938a). Similar
data were found by U LLRICH (1978) for Sperchon sp. and by E LLIS -A DAMS & D AVIDS
(1970) for Piona alpicola (N EUMAN , 1880). In Hydrachna, Eylais and Hydryphantes spe-
cies large numbers of larvae on the same host reduce the average terminal size of each at-
tached parasite and is probably a result of intraspecific competition by diminution in the
266 General part
nutrient extracted from their host (D AVIDS 1973a, L ANCIANI 1984, C OOK et al. 1989,
R EILLY & M C C ARTHY 1991). These crowded mites are likely to have a reduced fitness
because of lower fecundity or delayed maturation. A phenomenon known from Hydrachna
and Eylais spp. is that they parasitize host species of different sizes. Specimens living on
larger hosts realize a greater percentage of their total growth in the parasitic stage than
those on smaller hosts (L ANCIANI 1971, 1984, D AVIDS & S CHOOTS 1975, D AVIDS et
al. 1977b, R EILLY & M C C ARTHY 1991). Arrenurid larvae attached to the anal vein of
a dragonfly wing are behind in growth compared with those attached to the subcostal
vein (M ÜNCHBERG 1963a). Occasionally, the coexistence of a few well-developed larvae
along with numerous stunted ones on the same host suggests that during an early phase of
larval development some interaction allows further growth only to a sustainable number of
parasites (G ERECKE unpubl.).
A water mite larva harms its host by reducing the amount of energy available for growth,
reproduction, and maintenance and lowers survivorship. Unparasitized female insects pro-
duce significantly more eggs than parasitized ones, and this effect is proportional to the
parasite load or to the weight of the engorged mites compared to the weight of the
host (D AVIDS & S CHOOTS 1975, B.P. S MITH 1988, 1989c; L ANCIANI 1975, 1979a,
1984, L ANCIANI & M AY 1982, S MITH & M C I VER 1984b, F ORBES & B AKER 1991, R OLFF
2000, B RAUNE & R OLFF 2001). The feeding intensity of parasitized Cymatia bonsdorfi
(S AHLBERG ) (Hemiptera) was significantly greater than that of unparasitized bugs, possibly
as an attempt to compensate for the effects of nutritional drain by larval Hydrachna con-
jecta (R EILLY & M C C ARTHY 1991). Water mite parasitism should reduce damselfly male
mating success (F ORBES 1991, A NDRÈS & C ORDERO 1998), but this is denied by R OLFF et
al. (2000). R EINHARDT (1996) found that parasitic mites have a negative influence on the
flight ability of their host. Heavy mite infestation causes feeding scars in close proximity
and, in heavily loaded hosts, considerable areas of epidermis become eroded. Local lack of
epidermis seems to enfeeble the host, presumably owing to desiccation (ÅBRO 1982). At-
tached to the wing veins of Odonata, the larvae bring about lysis of the tissues and the
tracheas become filled with blood (P FLUGFELDER 1970). An increased asymmetry of the
wings may be the result (B ONN et al. 1996). For succesfully completing the life cycle how-
ever, it is obligatory for the mite to retain any negative influences to the host within strict
limits. In fact, the probability of water mite larvae returning to water is a function of their
host activity. As the detachment of larvae parasitic on chironomid adults was found to be
independent of the proximity of water by U LLRICH (1978), in these mites a preference for
host species which do not spend their aerial life away from aquatic habitats should be ex-
pected. Mite larvae on odonates detach at a significantly higher ratio from hosts with sub-
merged ovipostion behaviour compared with hosts ovipositing on the water surface (R OLFF
1997, R OLFF & S CHRÖDER 1999).
Deutonymphs and adults

In contrast to the larval stage, where parasitism is the rule, it is found only exceptionally in
deutonymphs and adults. Often such interactions between postlarval water mites and re-
presentatives of other taxa are restricted to commensalism. Species of the genera Najadicola
(Pionidae), Unionicola (Unionicolidae) and Dockovdia (Hygrobatidae) use freshwater
sponges or molluscs either temporarily (=t r a ns ie n t) as a site for oviposition and for trans-
formation (7-10 a), or more permanently as residents (= st a t io na r y). The complex life cy-
cle (7-10 b) includes, depending on the species, a larva that parasitizes a chironomid, returns
to water and then colonizes a sponge or mussel with which it may establish a life long as-
sociation (R.A.B AKER 1977, B OOTH & L EARNER 1978, H EVERS 1978a, 1980a, P ROCTOR
1990, E DWARDS & D IMOCK 1991, G LEDHILL 2003). A world wide survey of unionicoline
mite associations with molluscs is given by G LEDHILL & V IDRINE (2002).
7. Hydrachnidia 267
The molluscan host originally served as a pro te c te d site for the mite during transforma-
tion stages. The next evolutionary step is that after transformation the active stages did not
leave the mussel and showed a modification to a parasitic way of life (M ITCHELL 1955).
Apart from host specificity there is also site specificity. Najadicola ingens (K OENIKE , 1895)
Fig. 7-10: Water mites related to mussels and sponges; a, Unionicola aculeata, protonymph resting
stages on the inhalant siphon and some on the gills of Anodonta anatina (L.); grey spots are empty skins
of hatched mites, on the background eggs in the mantle tissue (Photo VAN M ECHELEN ); b, life cycle of
Unionicola crassipes (after B ÖTTGER 1972b).
268 General part
occurs in the pericard/renal and suprabranchial cavities (R.A.B AKER 1982), Unionicola
spp. in the mantle cavities of their hosts. Unionicola spp. generally distribute themselves
within rather specific regions and as a consequense several species can cohabit in the same
host individual on a spatiotemporal basis (M ITCHELL 1965b). D OWNES (1990) showed
that the morphology of mites can be influenced by the host species. No Unionicola
spp. have been reported from margaritiferine bivalves. A conjecture may be made that
when an ancient unionicolid assemblage had begun to coevolve with an ancestral union-
acean genetic stock, the margaritiferine clade may already have been genetically discrete
enough to be unamenable to coevolution with ancestral unionicolids (V IDRINE & B EREZA
1977a). A few species have associations with gastropods and all these associations are with
prosobranch gastropods (G LEDHILL 1985a).
Eggs of sponge mites may also hatch outside the sponge. Obviously, females only need the
sponge to lodge their eggs and the presence of sponge tissue is not necessary for egg devel-
opment. In contrast, eggs of mussel mites such as Unionicola intermedia and U. ypsilo-
phora (B ONZ , 1783) do not develop outside mussel tissue. Developmental time for eggs
as well as for protonymphs can last up to several months. This type of dormancy needs
further research (H EVERS 1980). D IMOCK (1985) observed more than 300 eggs of U. for-
mosa (D ANA & W HELPLEY, 1836) per cm2 of gill tissue. Sponge tissue can contain more
than 5000 mite eggs per ml (D AVIDS 1979). It is supposed that eggs in this amount hamper
the sponge, because no tissue could be observed, only eggs were found between the sponge
spicules.
The number of adult mites per molluscan host varies widely (H EVERS 1980). The abun-
dance of mites can increase with host size as is found for Unionicola formosa females (D I-
MOCK 1985). In contrast, M ITCHELL (1965b) never found more than two females and one
male U. fossulata (K OENIKE , 1895) in the mussel Lampsilis siliquoidea (B ARNES ), while the
pionid Najadicola ingens commonly occured with only one male and one female per host.
Unionicola fossulata females are usually found only as single specimens in the left or right
gill. Hence, it appears that females of this species tend to exclude other females by terrri-
torial behaviour. Deutonymphs were never found between gills, these sites were occupied
by actively ovipositing females (M ITCHELL 1965b). The presence of one U. fossulata male
seems to block the establishment of a second. Unionicola ypsilophora males display ter-
ritorial behaviour, they fight with other males, and as a result, there is usually one male per
host mussel. This intrasexual aggression, resulting in a female-defence polygyny, or a h ar-
e m m a t i n g s ys t em (D AVIDS et al. 1988), is also known from the North American species
Unionicola formosa and its sibling species U. foili E DWARDS & V IDRINE , 1994 (G ORDON et
al. 1979, D IMOCK 1983, 1985, D OWNES 1986, E DWARDS & D IMOCK 1991). Unionicola
ypsilophora males show no aggression against e.g. U. intermedia, but an infestation to-
gether with this species is rarely found. In many invertebrate taxa females are more numer-
ous than males (H AMILTON 1967), but such kind of female-defence polygyny has been de-
scribed very rarely.
The da m ag e to t he h ost is obvious in infestations of Najadicola ingens, occurring in
North America and Southeast Asia (V IDRINE 1996). This species physically tears the sup-
port structures of the gills and hampers the use of the mussel gill as a brood chamber
(H UMES & J AMNBACK 1950a, H UMES & R USSELL 1951). Damage to the host was also
found by R.A.B AKER (1977) and F ISHER et al. (2000), who demonstrated that Unionicola
spp. actually feed on host tissue or haemolymph. These mites probably do not ingest par-
ticulate material acquired by its host’s suspension feeding. W ELSH (1930, 1931, 1932b)
demonstrated a remarkable behavioural response to light by a mussel mite attributed to
Unionicola ypsilophora by the author (in reality, he possibly worked with Unionicola for-
mosa). Mites free of any chemical influence of their molluscan host consistently exhibited
positive phototaxis. However, in the presence of water from the mantle cavity of a host
7. Hydrachnidia 269
mussel, an immediate reversal to a photonegative state was observed. Positive phototaxis is

established in the European U. ypsilophora, but this species gives no response to molluscan
products (D IMOCK & D AVIDS 1985). This supports the contention of M ITCHELL (1957)
and V IDRINE (1980d) that the European U. ypsilophora and the North American U. for-
mosa are separate species. The characteristics of the inducing substance from the mantle
tissue of the mussel are described by D EL P ORTILLO & D IMOCK (1982), and J ACK & D I-
MOCK (1987). There is a strong indication of the occurrence of at least two separate photo-
physiological systems, evidenced by the photobehaviour of U. intermedia. Spectral- and
intensity-sensitivity data are available for a few species (R OBERTS et al. 1978a, D IMOCK
& D AVIDS 1985). Additional behavioural data on the ability of various unionicolids to
discriminate among potential host mussels are given by D OWNES (1986, 1989) and all ex-
isting data are summarized by D IMOCK (1988a). These behavioural preferences can be in-
terpreted as adaptations for locating potential hosts in the field and it is suggested that host
specificity is an important mechanism in maintaining reproductive isolation (E DWARDS
1999; E DWARDS & D IMOCK 1995a). This became obvious when a high genetic differentia-
tion was found between populations of U. formosa and U. foili, sibling species reproduc-
tively isolated by using different host species (E DWARDS & D IMOCK 1997). Furthermore,
there is an indication that host specifity among unionicolid water mites can vary geogra-
phically, owing to differences in geographic distribution of available hosts and differences
in host use (E DWARDS et al. 1998).
Commensal or parasitic relationships of adult water mites with organisms other than mol-
luscs have been observed in exceptional cases only: Parasitalbia sumatrensis K.V IETS , 1929
attaches to the abdomen and gill platelets of larval Povilla corporaali (L ESTAGE), a xylo-
phagus ephemeropteran living in the littoral of lake Singkarak, Sumatra (Indonesia). The
idiosoma of mature females becomes „physogastric“, growing to a pear shaped sac up to
2.700 lm in length. Astacocroton mollis H ASWELL , 1922 lives in the gill chamber of the
Australian decapod crustacean Astacopsis serratus H ASWELL , 1882. Hygrobates salaman-
drarum G OLDSCHMIDT et al., 2002 was found attached to newts from China. Its particu-
larly strong chelicerae allow for penetrating the hosts skin.
Life-histories
Hydrachnidia evolved from terrestrial ancestors. Their characteristic life history pattern is
similar to that of Trombidiidae and Johnstonianidae (terrestrial Parasitengona, chapter 6 of
this volume, 6-2). Hypotheses on their origin usually presume a non-swimming hydryphan-
toid-like ancestor (M ITCHELL 1957a, C OOK 1974, I.M.S MITH & O LIVER 1986, I.M.S MITH
& C OOK 1999). The endurance ability of some representatives of the Hydryphantidae to
withstand prolonged drought should be regarded as a primitive condition reflecting their
primarily terrestrial ancestors (W IGGINS et al. 1980). According to B ADER (1974a, 1977),
water mites should have a polyphyletic origin, with various groups of their ancestors start-
ing life in fresh water during different periods of earth history. In contrast, in his study on
the male reproductive system, B ARR (1972) came to the conclusion that, with the exception
of Hydrovolzioidea, the basic structure of this system appears to correspond in all recog-
nized water mite superfamilies, and W ITTE & O LOMSKI (1999) found a homologous struc-
tural composition of the ejaculatory complex also in Trombidia. Thus, the male reproduc-
tive system is consistent with a monophyletic clade Hydrachnidia + Trombidia. S MITH &
C OOK (1999) are of the opinion that the ejaculatory complex of Hydrovolzioidea is sec-
ondarily modified and this clade probably diverged from either hydryphantoid or eylaioid
ancestral stock. The eight superfamilies of Hydrachnidia are all characterized by plesioty-
pically 16 pairs of glandularia arranged in a regular pattern on the idiosoma (W ILES
1997b). This character state is considered as a synapomorphy of a monophyletic clade Hy-
drachnidia by numerous authors (see also chapter 6 of this book, 6-1). One might argue
270 General part
that a successful invasion of the new habitat necessitated a radically different method of
sperm transfer, and that similarities are the result of convergence. There is evidence that
water mites could have functioned with the same reproductive morphology as their terres-
trial relatives (D AVIDS & B ELIER 1979b).
One of the basic distinctions within Hydrachnidia is whether the larvae have remained
aerial, the ancestral condition, or whether there have been changes in morphology to pro-
duce swimming larvae. Until recently it was thought there were only two evolutionary lines
with swimming larvae, the Hydrachnidae and a group of „higher“ families on the „phy-
logenetic ladder“ starting with the Sperchontidae. However, it is now obvious that swim-
ming larvae have evolved indepently several times (S MITH et al. 2001). Also certain mem-
bers of the Wandesiinae (Hydryphantidae) possess aquatic larvae. Similarly, in the Hydro-
volzioidea members of the Hydrovolziidae have terrestrial, but of the Acherontacaridae,
aquatic larvae (T UZOVSKIJ et al. 2001).
Life span
The duration of the separate life cycle stages is rather variable among species and highly
dependent on temperature and other abiotic conditions. In general, the life span is limited
to one year with long-lived females. Since the chance of surviving to the following repro-
ductive season is generally small, the cost of delaying reproduction increases strongly and
soon outweighs the benefits of increasing in size. Exceptions with bi- or trivoltine life cycles
are made by those species living at low temperatures or investing a lot of maintenance by
having a strong exoskeleton or by unpalatable substances produced by the glandularia
(D AVIDS 1997). Development times of the various stages of several species are reported
in detail by B ÖTTGER (1962, 1972a,b), E LLIS -A DAM & D AVIDS (1970), D AVIDS
(1973a), D AVIDS et al. (1985), S TECHMANN (1978a), H EVERS (1980), M EYER &
S CHWOERBEL (1981) and M EYER (1985, 1989). Voltinism of species depends on the av-
erage length of the reproductive season relative to generation or development time, and
on climatic conditions. In cold-stenothermous rivulets with temperatures between 4
and 10 C, females of several species were found to live two to three years (B ADER
1977a, 1980b). Hydrodroma despiciens females can live at 20 C for more than one
year and still produce some eggs in their second year (M EYER 1985). The subsequent de-
velopment stages of Limnochares aquatica pass facultatively through two to three free-liv-
ing nymphal stages, exceptional in water mites. The adults live several years and there are
additional moults (B ÖTTGER 1972a). Unionicola crassipes (or a related species, see C RO-
WELL & D AVIDS 1979) in a Canadian Lake has one generation (R IESSEN 1982), but, in The
Netherlands two generations (D AVIDS et al. 1985). In the former, ice covers the lake surface
for about four and a half months. The combination of a prolonged unfavourable period
with a complex and generally lengthy life history may be major factors in limiting the Ca-
nadian population to a single generation per year. Biological factors (seasonal occurrence of
hosts for the larvae, prey for deutonymphs and adults) may tend to reinforce this climatic
limitation and also profoundly affect the exact nature of the life history, including the iden-
tity of the overwintering stage (R IESSEN 1982a). In the majority of species, fertilized fe-
males hibernate, but water mites may also hibernate as eggs, as parasitic larvae on their
hemipteran or coleopteran host, or as nymphs. In this context, the role of the resting stages
(proto- and tritonymph) which are hidden in the sediment and difficult to detect, is almost
unknown. In many taxa, males emerge a few days earlier than conspecific females. The
males of certain sperchontids and lebertiids occur some weeks earlier than the females
of the same generation (B ADER 1977a, 1980b). But, females live longer than males, the
latter generally mate and die within a few days to weeks after maturing. This explains
to a great extent the heavy preponderance of females in many species. Following a hypo-
thesis of G ERECKE (2002), stream-dwelling mites in temperate areas could have a double
7. Hydrachnidia 271
Fig. 7-11: Life cycle of Hydrachna conjecta; a, The female depositing her eggs in an air cavity of an
aquatic plant; b, swimming larvae in search for a corixid host ( e.g. Sigara striata), c, adult feeding on
corixid eggs (after D AVIDS 1979).
track strategy with parts of the population overwintering at the adult stage, other parts as
eggs. The percentage of overwintering specimens may change with annual conditions, de-
pending on climatic conditions and thus become a limitating factor for the distribution of
selected species at lower latitudes.
E g gs: Under temperate and cold climates, the majority of water mite species breed in
spring and early summer, other species in autumn or year-round (B ADER 1977a). No
data are available for species from the tropics, where year round reproduction is probable
in areas with little seasonal climatic change. A week to a month after insemination, egg
production starts (B ÖTTGER 1962, E LLIS -A DAM & D AVIDS 1970, D AVIDS 1973a, S TECH-
MANN 1978a, R IESSEN 1982). Water mites produce their eggs in a series of separate events
(„it e ropa ri ty“). This strategy only gives a high reproductive value when adult mortality
is low. Total egg production varies greatly among species, but also within populations,
depending on life conditions and food supply. Total egg numbers per female are: Eylais
discreta up to 13.000 (with clutch sizes of 1.000-2.000 over three months); Hydrachna
conjecta 500-1.000 (D AVIDS 1973a), Piona spp. (E LLIS -A DAM & D AVIDS 1970, R IESSEN
1982), Arrenurus spp. (C ASSAGNE -M ÉJEAN 1966a, S TECHMANN 1978a), Hydrodroma
272 General part
despiciens (M EYER 1985), and Hygrobates spp. (VAN H EZEWIJK & D AVIDS 1985) about
100-500, in several batches from less than 10 to 50 or more deposited over a period of one
to three months, in Arrenurus also up to ten months. Egg production is also significantly
influenced by temperature: Hydrodroma despiciens produced 600 - 800 eggs at 25 C, but
200-600 at 15 C, over a period of two to three months (M EYER 1985). Acerbitas sp. failed
to deposit eggs below 13 C (M ULLEN 1977). Cold-stenotherm species produce fewer eggs
than eurythermous species (WALTER 1922c) and females of several taxa minute in size are
known to deposit one egg at a time (e.g. species of Feltria, Aturus or Stygomomonia, M AR-
TIN pers. comm.). Egg development time in Hydrodroma despiciens is influenced by the
daily photoperiod and temperature (eight days at 25 C, 41 days at 15 C: M EYER 1989).
Arrenurus eggs develop in 10-14 days at 18-22 C, four to eight times longer at 8-18 C
(S TECHMANN 1978a), several months at 4-5 C (B ÖTTGER 1962). Other studies give similar
results. As long as temperature does not sink below a limit of 13 C, Atractides nodipalpis
T HOR , 1899 larvae do not leave the covering of the egg mass and stay motionless for at least
six months. They must be activated by lower temperatures. After a few months at a tem-
perature of 6 C, they will emerge at 13 C (U LLRICH 1978).
L a rva e an d N ym p hs: Water mite larvae parasitic on short-lived hosts such as dipterans,
typically need two to seven days to complete engorgement (E LLIS -A DAM & D AVIDS 1970,
B ÖTTGER 1972b, U LLRICH 1978, S TECHMANN 1978a, M EYER 1985). Longer development
times are observed in larvae parasitizing surface-dwelling Hemiptera (six to eight days,
B ÖTTGER 1972a), on damselflies (ten days, S TECHMANN 1978a), but in particular in larvae
hibernating on submerged Hemiptera and Coleoptera (up to ten months, D AVIDS 1973a,
B.P. S MITH 1988). In most cases, the engorged larva detaches from its host, re-enters the
water and transforms on a suitable substratum to a protonymph, but species hibernating on
their host (Eylais spp., Hydrachna spp.) moult there and remain attached also at the pro-
tonymphal stage and only the fully formed deutonymph leaves the host. This extended
association presumably reflects the large size and mainly aquatic existence of these hosts.
In Piona- and Arrenurus species the transformation to deutonymph lasts about four to se-
ven days (B ÖTTGER 1962, E LLIS -A DAM & D AVIDS 1970, S TECHMANN 1978a). Literature
data indicate that the duration of the deutonymphal stage varies widely from a couple of
weeks to five months or more. Deutonymphs of Hydrodroma despiciens live about four
weeks at 20 C, but do not enter into the tritonymphal stage at 10 C (M EYER 1985). In the
species studied by E LLIS -A DAM & D AVIDS (1970), M EYER (1985) and S TECHMANN
(1978a) the tritonymphal stage lasts three to eight days, but long-term quiescence in other
species is hypothesized by G ERECKE (2002).
Adaptations and Strategies

A state of sup pre sse d m e ta bo li sm is observed in eggs, larvae and nymphs of several
species (M ITCHELL 1955, B ÖTTGER 1962, D AVIDS 1973a, U LLRICH 1978, W IGGINS et
al. 1980, I.M.S MITH 1982, M EYER 1985, R EILLY & M C C ARTHY 1991, M ARTIN 1998,
M ARTIN & D AVIDS 2002). Several species occur only in early summer and this may be an
effect of the overall timing of the whole life cycle. Dormancy (escape in time) represents an
important phase in many life cycles. It is not only in enhancing the chances of survival
during adverse periods of the year, but also in the regulation of seasonal phenologies
by the synchronization of life cycles and the determination of patterns of voltinism (T AU-
BER et al. 1986). Diapause induction can be photoperiodic or thermoperiodic. In hibernat-
ing larvae (Hydrachna and Eylais spp.) the reproductive diapause of the insect host may
also play a part (D AVIDS et al. 1977b).
Larvae belonging to the Hygrobatoidea, Lebertioidea and Arrenuroidea, and probably se-
lected representatives of other superfamilies as well, exhibit a phoretic association to ma-
7. Hydrachnidia 273
ture larvae or pupae of their hosts („pre pa ra sit ic a tt en da nc e“) and become parasitic
only at the moment of the imaginal moult. P hor esy is a symbiotic process by which an
organism hitchhikes on the exterior of a host organism, disembarking as the host traverses a
suitable environment. As the phoretic organism becomes more committed to one particular
host species, behaviour of that host becomes an important selective force in the evolution of
traits affecting symbiosis in the phoretic. The formation of a symbiotic relationship is a
logical and necessary step prior to parasitism. Once parasitism is accomplished in a taxon,
some aspects of the phoretic activity become maladaptive and would be removed from the
life cycle (H OUCK 1999). In this case parasitism may emerge as an evolutionary phenome-
non when the phoretic organism extends its dependence on the host from simple habitat
dispersal to nutrition (ATHIAS -B INCHE 1994, H OUCK 1994). Phoresy may indicate the ori-
gin of the evolutionary line along which parasitism in most water mites has been developed.
The advantage of a parasitic phase is that energy is gained in a short period of time. There-
fore, larval parasitism prevents a constraint in development time. A trade-off is the risk of
finding a proper host and the return to a suitable habitat for further development. A con-
straint for parasitic larvae is the life cycle of the insect host. As a consequence species with
non-parasitic larvae can start egg production early in spring and the benefit may be a longer
reproductive period (D AVIDS 1997).
It is generally accepted among parasitologists that the evolution of parasites in most in-
stances has not kept pace with the evolution of the host. The hypothesis may be formulated
that when insect hosts evolved towards an aquatic life, adult water mites adjusted sooner to
an aquatic life than their parasitic larvae. This explains why in water mites belonging to
early derivative clades, larvae often resemble the terrestrial forms more than do deuto-
nymphs or adults (D AVIDS & B ELIER 1979b). Animals with a small progeny are assumed
to suffer little loss in the juvenile stages. In contrast, it is accepted that parasites produce
many offspring as compensation for losses during their complicated life cycle. The huge
amount of eggs produced by Eylais species gives an indication of the enormous loss of lar-
vae in their pre-parasitic period (D AVIDS 1997). It is of basic importance for these species
that the parasitic growth is so substantial, that this loss may be balanced by a high egg
production. In contrast, adult Arrenurus spp. live for two to three years, but often produce
not more than 100 eggs per female (B ÖTTGER 1962, C ASSAGNE- M ÉJEAN 1966a, S TECH-
MANN 1978a). Due to their remarkably strong exoskeleton much of their energy may be
allocated into maintenance and this may decrease vulnerability to predation (D AVIDS
1997). Species known to be larval parasites of odonates are often characterized by having
large egg clutches from which small larvae hatch. In contrast, species known to be parasites
of dipterans produce larger larvae hatching from smaller clutches. The probability of host
discovery and competition between parasites is directly correlated to the two tactics de-
scribed above (W.J. C OOK et al. 1989).
A variation from the basic life-cycle is the lo ss of the pa ras iti c phase, in which larvae
forego parasitic feeding. Larvae may emerge from eggs with sufficient nutrition to allow
them to transform into protonymphs, or, they do not leave the egg, further development
takes place and the deutonymph hatches as the first active stage. A reproductive trade-off
between the size of the eggs and number of eggs would be expected in such cases (D AVIDS
1997). The lack of this association has been ascertained only in a few exceptional cases and
is presumed to be apotypic (B.P. S MITH 1998), obviously representing an evolutionary blind
alley. In fact, examples are reported only from species closely related to species with para-
sitic larvae and no example is known of a taxonomic radiation in a clade after loss of para-
sitism.
274 General part
Deformities, diseases and epibionts

Misshaped specimens are regularly found in populations of Hydrachnidia. They usually
occur in the form of bifurcations, rarely trifurcations, of legs (with the branches bearing
claws directed against each other - K OENIKE 1888b, T HOR 1926c, L ÁSKA 1955), the sup-
pression of sclerites and appendages, and the reduction, or multiplication, of setae and
acetabula. Intersexual specimens may combine a female external genital field with the pre-
sence of an ejaculatory complex, or they present a male genital field without having devel-
oped the ejaculatory complex. Usually, such intersexes do not exhibit secondary sexual
characters found in males of the species to which they belong. An account of such phenom-
ena found in the genus Atractides is given by G ERECKE (2003).
Little is known about infections and parasites of water mites. Parasitic species of Micro-
sporidia (Cnidosporidia, Protozoa) have been recorded in species of the genera Sperchon
(D AVIES 1960, L IPA 1962) and Limnochares (L ARSSON 1990) but no records about fungi or
bacteria attacking water mites are available. YAKIMENKO et al. (1997) report the presence
of arboviruses in deutonymphs and adults of species representing several water mite genera
in Western Siberia.
Frequently, especially in species living in muddy habitats rich in organic material, water
mites are found covered by epibionts such as suctorians, vorticellid ciliates or diatoms.
Nothing is known about a possible specifity of protozoans as epibionts of these mites. In-
stead, the cyanobacterium Sokolovia neumaniae E LENKIN (1926) described from Buchara,
Uzbekistan as growing on all legs of Neumania triangularis P IERSIG , 1894, a species
incerta, probably synonym of N. limosa (K OCH , 1836) or N. deltoides P IERSIG , 1894,
has not been found again since the original description (K OMÁREK & A NAGNOSTIDIS
2005). Epibionts of water mites could be an interesting subject for future research.
Habitat
Hydrachnidia occur throughout the world, except Antarctica, in almost every type of fresh-
water habitat. They inhabit running as well as standing waters, temporary, permanent,
hyporheic interstitial waters and springs, usually in close association with the bottom sedi-
ments or aquatic macrophytes (D I S ABATINO et al. 2002). Species belonging to the genera
Pontarachna and Litarachna have evolved to a marine life (WALTER 1925). Water mites are
not known from ephemeral water bodies, the deep ground water of porose aquifers or
strongly polluted waters; in caves and karst aquifers water mites have been recorded
only occasionally, often as single individuals and mostly representatives of surface dwelling
species.
Standing waters
Most limnophilous taxa tolerate a broad range of temperatures which they encounter in
weed beds and silty substrata in small water bodies and in the littoral and sublittoral zones
of lakes. A few cold-adapted stenophiles are restricted to either arctic or alpine glacial
lakes. The most extensive studies on the distribution of water mites in lakes are those
of C. A NGELIER (1953), B AGGE (1989), B IESIADKA (1972b, 1980a), B ÖTTGER & M IER-
WALD (1990), D AVIDS et al. (1994), D I S ABATINO et al. (2004), K OWALIK (1977, 1978),
M EYER & S CHWOERBEL (1981), M OTHES (1964a), P EŠIĆ (2001), P IECZY N~ SKI (1959, 1960,
1976), K. V IETS (1924h, 1930f, 1931d), R AMAZZOTTI (1947), K.V IETS & K.O.V IETS
(1954d), K.O.V IETS (1979b) and WALTER (1922c). Most of the lakes studied are in central
Europe.
Generally, with increasing depth, a decrease in density and species number is observed.
B AGGE (1979a, 1983) found 79 % of the total number of species of a lake in Finland
7. Hydrachnidia 275
in the upper 1.5 m, 40 % from 5 - 10 m depth and only 15 % from 20 - 30 m; below 40

meters mites were absent. The mite fauna at various depths is composed differently and
depends mainly on habitat differentiation. European species considered as characteristic
for the profundal zone are Arrenurus coronator T HOR , 1900, A. nobilis N EUMAN ,
1880 and Huitfeldtia rectipes T HOR , 1898. In boreal lakes the stenothermous species Neo-
brachypoda ekmani (WALTER , 1911) and Acalyptonotus violaceus WALTER , 1911 are
found (L UNDBLAD 1927c; B AGGE 1983,1989). D I S ABATINO et al. (2004) report various
cold stenothermous species from Alpine lakes which at lower altitude are restricted to
springs and spring streams.
K ARL V IETS (1924h, 1930f) made extensive faunistic studies on the vertical distribution of
water mites in 50 lakes, mainly situated in northern Germany. He collected mites in quite
large numbers down to a depth of 30 m. The trophic level of these lakes must have been low
at that time, because water mites do not occur in anoxic waters. He observed that during
wintertime the mites inhabit deeper layers than in summertime. In studies on ditches, canals
and ponds in The Netherlands (S MIT & VAN DER H AMMEN 2000, VAN DER H AMMEN
1992) a high diversity of water mite communities was documented, their composition
being influenced by salt content, degree of eutrophication and dimensions. B IESIADKA
& C ICHOCKA (1997) studied so-called „Lobelia-type“ lakes, which are slightly calcareous
and favour a specific faunistic character compared to other lakes. Water mite communities
of various types of small water bodies were studied and compared furthermore by B IESIAD-
KA (1972b), B IESIADKA & K OWALIK (1980) and C ICHOCKA (1996b). Data on water mites
from brackish waters are given by K.V IETS (1925f) and G ERECKE (1991), from bogs by
K.V IETS (1938e) and C ICHOCKA (1996b, 1998), and from sodium rich waters by P ONYI
& P ONYI (1958).
In the pelagic zone of European lakes, water mites occur only sporadically, mostly they are
representatives of the genera Unionicola and Piona (S CHWOERBEL 1972). In contrast, true
planktonic mites were found in large numbers in several American lakes: In Lake Madden,
Panama, Piona limnetica B IESIADKA , 1975 attained a maximum density of 2.000 indivi-
duals per m2 (150 individuals per m3) (G LIWICZ & B IESIADKA 1975). Similar high densities
were found for Piona constricta (W OLCOTT, 1902) in the plankton community of Heney
Lake, Canada (R IESSEN 1980, 1982) and for Centrolimnesia boopis G ERECKE & F ISHER -
H ARTIG , 1997 in a lake in Amazonian Ecuador (G ERECKE et al. 1997). In all these lakes,
diel vertical migration of pelagic water mites was observed.
The highest numbers of water mites occur in the community of the littoral fauna, especially
in the vegetation zone. Differences are found between communities of emergent (e.g.
Phragmites) and submergent (e.g. Potamogeton or Chara) vegetations (P IECZY N ~ SKI
1959, 1960, 1964, K OWALIK 1977, K.O.V IETS 1979b, Z AWAL 1992). P IECZY N ~ SKI
(1976) proposed that individual numbers in the littoral zone should be of the order of
1.000 individuals per m2. The highest density, 39.400 individuals per m2, is given by T U-
ZOVSKIJ (1972a). M OTHES (1964a) noted 230-280 individuals per m2 in the oligotrophic
Lake Stechlin. T EN W INKEL et al. (1989) found about 1.000 Hygrobates individuals per m2
on a shallow sandy littoral. These data indicate the considerable significance of water mites
as elements of benthic communities.
Running waters
Many representatives of water mites invading streams evolved from the plesiotypically
globular body shape to a wedge-shaped and flattened body designed to produce a down-
ward pressure in response to water flow; locomotion necessitated evolution of a generally
crawling gait. In natural streams, species numbers are generally rather high and may reach,
or occasionally exceed, 50 species at a single collecting site. The most extensive studies on
276 General part
the distribution of water mites in streams were in mountain areas of Central and Northern
Europe: England (G LEDHILL 1973), Sweden (L UNDBLAD 1930c, 1962), Poland (K OWALIK
& B IESIADKA 1981; C ICHOCKA 1996a), Germany (G ERECKE 2002; M ARTIN 1997; M AR-
TIN & S PETH 1996; M EYER 1986; S CHWOERBEL 1959, 1964a; K.V IETS 1959d; K.O.V IETS
1955a, 1957c), the Alps (WALTER 1922c, 1944; M OTAŞ 1928e, B ADER 1963, 1975). Com-
parable data from Southern Europe come from the Pyrenees (C. A NGELIER 1951b, E. A N-
GELIER et al. 1985, V INÇON 1987), Central and Southern Italy (C ICOLANI & D I SABATINO
1985, C ICOLANI et al. 1991) and Sicily (G ERECKE 1991a, F ERRITO 1994). The water mites
from lowland brooks in The Netherlands were surveyed by VAN DER H AMMEN & S MIT
(1996). Larger streams and rivers are insufficiently investigated in Europe. Data available
from the upper Rhine between Basel and Koblenz (B ADER 1988a, L ANGE 1990) and from
the middle Neman in Belarus (B IESIADKA et al. 2004) indicate the presence of rhithrobionts
known also from lotic parts of lower order streams, and a highly diversified standing water
fauna in pools and oxbow ponds when naturally conserved reaches are present.
Most rheophilic species are strictly bound to characteristic stream sectors or microhabitats.
In the middle, and especially in the lower course of a river, species common in standing
water, may also occur. Temperature and substratum in combination with water quality
have an authoritative influence on density and species composition (D I S ABATINO et al.
2000). In the Black Forest (SW Germany), S CHWOERBEL (1959) distinguished four rhithral
sectors, and a fifth one in the epipotamon, each with characteristic water mite commu-
nities. B IESIADKA (1974) found a great variability of habitats in a Carpathian river and
divided it on the base of the occurrence of water mites into two zones. S MITH et al.
(2001) observed up to 5000 individuals per m2 of substratum in running water riffles
of streams in North America. M EYER (1994) reports a medium annual density of about
600-1000 individuals per m2 in a small stream in southern Germany with a contribution
by abundance of 5.5 % to the total invertebrate community. Regional differences in species
composition can often be explained by environmental variables with a particular influence
of geological factors (S CHWOERBEL 1959, M EYER 1986, G ERECKE 1991). At a lower scale,
different substratum types are characterized by communities of species which obviously
display distinct habitat preferences (G ERECKE 2002). Also many stream dwelling water
mites are colourful, possibly as a deterrent against predators in correlation with the repel-
lent production of glandularia, but others have a mimetic coloration. Probably, thanks to
their defence system, and in contrast to all other invertebrates in streams, lotic water mites
have their maximum activity during daytime. This is evidenced by the opposite abundance
pattern in drift experiments, where water mites exhibit peak occurrences during daylight
(S CHMIDT 1969, M ARTIN 1999).
Compared with species from lakes and ponds, investigations on lotic mites started about a
century later and important details were documented only during the past fifty years. Con-
sequently, little is known about faunal changes under anthropogenic pressure. Compari-
sons between more or less polluted stream sectors regularly testify for a strong sensitivity of
water mites against organic load and heavy metals (e.g. B IESIADKA & K ASPRZAK 1977;
B OLLE et al. 1977; C ICOLANI & D I S ABATINO 1985, 1988, 1991, 1992; I LLIES 1954;
K OWALIK & B IESIADKA 1981; L EARNER et al. 1971; M EYER 1986; P ETROVA 1985;
P UNĆOCHÁŘ 1972; S CHWOERBEL 1964; Y OUNG 1969). In a 25-year monitoring in the
upper Danube area, G ERECKE & S CHWOERBEL (1991) document how temporal changes
of domestic waste-water pollution are reflected by water mite species number and diversity.
C ICOLANI & D I S ABATINO (1992) show that the impact on water mite populations caused
by stream regulation is qualitatively and quantitatively similar to the influences of in-
creased organic pollution. The extinction of species from catchments of organically pol-
luted and/or regulated rivers could be caused by direct impact on the mite population, or be
a secondary consequence of the disappearing of suitable insect host species. Observations
7. Hydrachnidia 277
of a particular sensitivity of water mites to heavy metals suggest that in this case a direct
influence is acting on the mite population (G ERECKE & S CHWOERBEL 1991). The potential
of water mites as indicators of heavy metal pollution is an interesting, but little studied
subject. The water mite fauna of streams appeared to be deteriorated in The Netherlands
due to canalization and eutrophication (VAN DER H AMMEN & S MIT 1996).
Springs
The water mite fauna of springs has attracted attention in various parts of Europe. A par-
ticularly rich set of data is available for the following areas: Sweden (L UNDBLAD 1926a,
1968), Central European secondary chain of mountains (B IESIADKA & K OWALIK 1978b,
B IESIADKA et al. 1990, G ERECKE et al. 2005, S CHWOERBEL 1959, 1959a, K.V IETS 1925g,
K.O.V IETS 1955a), the Alps (B ADER 1975, 1977a, 1994, G ERECKE et al. 1998, G ERECKE &
M ARTIN 2006, WALTER 1922c), the Appennines (C ICOLANI & D I S ABATINO 1990, C ICO-
LANI et al. 1995), the Tyrrhenian islands (G ERECKE 1991, G ERECKE & D I S ABATINO 1996)
and the Pyrenees (R OCA & G IL 1992). From all these studies, as well as from research in
other continents (Northern America: S MITH 1991, Latin America: G OLDSCHMIDT 2001)
it is clear that the Hydrachnidia is a group of animals with a particularly high diversifica-
tion in springs. With regard to the percentage of crenobionts (species strictly bound to
spring habitats and not found in other habitat types), no other taxon has produced
such a high number of specialists (e.g., D I S ABATINO et al. 2003). Crenobionts found in
high numbers in spring sources, occur only as scattered specimens in the springbrook
and are completely replaced by rhithrobiontic assemblages further downstream (e.g.,
B IESIADKA 1974, G ERECKE et al. 1998).
Traditionally, most of these species are considered to be stenobiontic, in particular bound to
the year round stable temperature of springs. However, some of them may be found in small
pools bordering the spring area which are much warmer during summer, and are easily
reared in the laboratory at room temperature. Furthermore, in the tropics where the gra-
dient of temperature stability between spring and stream is negligible (without noticeable
seasonal change far distant from the spring), crenobiont water mite taxa are also found
(e.g. G OLDSCHMIDT 2001). Therefore, still unclear additional factors obviously contribute
to the attractiveness of these habitats to water mites. G ERECKE & D I S ABATINO (1996) and
D I SABATINO et al. (2000) postulate that the presence of a diverse dipteran fauna, as well as
the importance of springs as egg deposition sites for insect species not strictly bound to
springs, provide particularly favorable conditions for these egg predators and parasites.
In any case, water mites merit particular attention in the documentation and monitoring
of spring faunas, and their habitat requirements merit further research (C ANTONATI &
O RTLER 1999). Species of the subgenus Partnuniella (Wandesia, Hydryphantidae) and
all known members of the monotypic family Thermacaridae are thermobionts and are ex-
clusively found in hot springs, at temperatures from 32-43 C (S OKOLOW 1927b, M IT-
CHELL 1960, S CHWOERBEL 1987, M ARTIN & S CHWOERBEL 2002). Monatractides alger-
iensis (L UNDBLAD , 1942) is a thermophilous species in North Africa found in various ha-
bitats, but in southern Europe bound to thermal springs (D I SABATINO et al. 1992). Species
only found at higher elevations in mountains, or having a boreoalpine distribution, can be
considered cold stenotherms. However, many of the species listed under this category by
S CHWOERBEL (1959), have in recent times been also detected in warmer climates, and other
factors, such as preference for certain host insects, are probably more important factors
triggering the presence / absence of localized species.
For the terminology of spring types and the ecology of their specific inhabitants, important
details are given, and the bibliography reviewed, by D I S ABATINO et al. (2003) and G ER-
ECKE & M ARTIN (2006). It is particularly interesting that, in contrast to the representatives
278 General part
of several other groups of invertebrates, no similarity is found between the water mite fau-
nas of springs on the one hand and the hyporheic interstitial on the other (G ERECKE et al.
2005). There are no obvious morphological trends which characterize spring dwelling
mites compared with rhithrobionts. Both in ancient groups (e.g. Protziinae, Tartarothya-
dinae, Euthyadinae of the family Hydryphantidae), and in more derivative clades of which
some members have secondarily invaded spring habitats (e.g. some Atractides spp., Hygro-
batidae), soft bodied species as well as species with more or less extensive dorsal and ventral
shields may be crenobionts. As a general rule, reduction of swimming setae has developed
in crenobionts to a major extent compared with stream dwelling species (S CHWOERBEL
1959). The phenomenon of crenobiosis of some species is differentiated at a regional scale,
in general changing with geographical latitude: some are found as strict crenobionts in
central European lowlands and secondary mountains, or in high mountains of southern
Europe, but occur in the profundal of Scandinavian lakes or in the benthos of lakes at
high elevation in the Alps (L UNDBLAD 1962, D I S ABATINO et al. 2004).
Interstitial waters
Mostly in the second half of the 20th century, studies on the hyporheic interstitial have
demonstrated the presence of a diverse water mite fauna with a high percentage of char-
acteristic species in different parts of Europe. The most important examples (including
references to a widely scattered bibliography) are: England and Scotland (G LEDHILL
1971, 1973a, 1982b); Central-East European mountain chains (B IESIADKA 1975, G ERECKE
et al. 2005, S CHWOERBEL 1959a,1961a, K.V IETS 1955a, 1959d), the Alps (R ENSBURG
1971, S CHWOERBEL 1962b, WALTER 1947), Pyrenees (E. A NGELIER 1949b); The Balcans
(from here came the very first record of a stygobiont mite, K.V IETS 1932b, WALTER 1947,
M OTAŞ et al. 1947d, M OTAŞ 1962b); Apennines (D I S ABATINO et al. 1996, D I S ABATINO &
C ICOLANI 1999), Tyrrhenian Islands (G ERECKE 1991, G ERECKE & D I S ABATINO 1996).
Results from a world-wide survey of the subterranean water mite fauna (S CHWOERBEL
1986 a-c) and the analysis by S MITH & C OOK (1999), indicate a high number of taxono-
mically isolated clades living in hyporheic waters. Many of them have uncertain taxonomic
relationships, are endemic to restricted geographical areas or have disjunct distributions. In
Europe, there is a clear tendency of a decrease in diversity from south to north, with no
records of hyporheobiont Hydrachnidia published so far from Scandinavia (G ERECKE
2002). However, assuming the period of glaciations did not permit the survival of inter-
stitial populations in Fennoscandia, life conditions surely have turned favourable and some
interstitial species may have succeeded in recolonizing the area. As in the case of spring
dwelling mites, interstitial species may also show regional differences regarding habitat
preference (G ERECKE & D I S ABATINO 1997).
Most hyporheobiont species live in superficial riverine gravels. In contrast to hyporheic
representatives of other taxa, they do not usually extend to deeper layers of groundwater
(G ERECKE et al. 2005). From deep wells or cave waters, water mites have been recorded
only very rarely. However, troglobiontic water mite species have been detected at several
occasions in New Zealand (I MAMURA 1978, 1979b, C OOK 1992). Life conditions in the
interstitial habitat and evolutionary trends of hyporheobiont mites were studied by
S CHWOERBEL (1961, 1967). Interstitial species are generally smaller and more slender
than their surface dwelling relatives and have lateral eyes with lenses reduced in size,
and pigmentation reduced or absent. They are highly thigmotactic, specialized for crawling
or running through interstices in sand or gravel deposits and tend to have relatively long
appendages with enlarged tactile setae. Possibly as an adaptation for anchorage in sediment
microspaces (not, as sometimes suggested, for peristaltic movement), some species are ver-
miformly elongated (in the European fauna Stygothrombium spp., Wandesia spp. and
Parawandesia chappuisi). In contrast to the general tendency towards dwarfism, such spe-
7. Hydrachnidia 279
cies may be rather large and are often found at the interface between gravel and large stones
or rocks. Compared with the surface sediments, the hyporheic interstitial habitat is more
stable regarding water chemistry and temperature. S CHWOERBEL (1959, 1967) proposed its
function as a refugium in periods of drought or extreme temperatures. However, this hy-
pothesis was not confirmed by the experimental studies of B OULTON et al. (2003, 2004). In
contrast to older suggestions (S CHWOERBEL 1961a) and just like their surface-dwelling
relatives, hyporheic mites are also bound to insect hosts for their parasitic larval stage
(G ERECKE 1996). Consequently they are somewhat restricted for the colonization of deeper
sediments. However, their parasitic-phoretic phase does provide better means for coloniza-
tion than other members of that biocenosis have.
Temporary pools and semi-aquatic habitats

In a strict sense, no drought-resisting stages are known in Hydrachnidia. For inhabitants of
te m po ra ry p ool s two strategies can be distinguished for tolerating or avoiding drought
(W IGGINS et al. 1980). A number of species belonging to the Hydryphantidae, Pionidae and
Arrenuridae reproduce in springtime before surface water disappears. These taxa aestivate
in the dry pool mainly as nymphs or adults and can tolerate drought for considerable per-
iods provided a minimum humidity is maintained. Species of Hydrachnidae and Eylaidae
are also found in temporary pools in springtime. Eylais spp. retain an essentially terrestrial
existence as larvae while all other instars are fully adapted aquatic organisms unable to
withstand drought. Eylais and Hydrachna larvae leave the pool attached to their hosts
and survive as parasites of their hosts for a period of up to ten months. Temporary waters
can differ widely from each other: for instance, coastal temporary dune-waters are very rich
in water mite species, but are inhabited by characteristic species assemblages depending on
hydrology. Dune areas with a high-lying hinterland can be distinguished from dune areas
with a low-lying hinterland on the basis of their water mite communities (S MIT & VAN DER
H AMMEN 1992). A classification of the water mite fauna in various types of temporary
running waters is given by G ERECKE (1991) who distinguishes three basic types of su m -
m e r-d ry str ea m s: (1) completely dry in summer and lacking a lotic water mite fauna also
in winter and spring time, but with a few pioneer colonizers of the genera Eylais and Hy-
drachna in pool areas; (2) with remnant pools in summer, the mite community dominated
by stagnant water species which in springtime may be found also in lotic areas, but under
unfavorable conditions for these species; (3) with a subterranean flow persisting during
summer, the lotic mite fauna more or less rich, dominated by species particularly adapted
to survival of summer drought at the nymphal stages such as Sperchon algeriensis (L UND-
BLAD , 1942) and Atractides inflatus (WALTER , 1925). A remarkable assemblage of water
mites was found in qu ag fe ns an d ca rr, a semi-aquatic habitat (S MIT & VAN DER H AM-
MEN 1996). The number of species in carr is much lower than in quagfens. No typical spe-
cies of quagfens can be distinguished, although their fauna is characterized by a number of
species almost confined to terrestrializing water bodies which elsewhere are found in tem-
porary waters (e.g. B ÖTTGER & V ÖLKL 1987).
Rare and threatened species

During preparative work for this book, intense revisional work on several families allowed
the reassessment of numerous species described a long time ago. Due to bad documentation
and/or loss of type material, many of them had to be designated as species incertae. Their
names should not be used in future faunistic work and they should not be included in taxo-
nomic lists from now on. Numerous other names proved to be junior synonyms of species
known under a different name at the time of their description. In a few cases species new to
science were detected or important additions to our knowledge on geographical distribu-
280 Special glossary
tion were made. The presently known number of about 700 species recorded from the area
treated in this book appears to be realistic. The number of species not yet recorded from this
area will probably be equalled or exceeded by the number of (poorly documented) species
placed into synonymy during future taxonomic studies.
Based on knowledge accrued up to 2004, G ERECKE & L EHMANN (2005) attempted to high-
light possible threatened species in the fauna of central Europe. At present it would not
make sense to present a ’Red List’, but species of the following categories merit particular
attention during future research (for details concerning the Limnofauna Europaea, see
K.O.V IETS 1978: (A) found only once at one site or in a very limited area; (B) restricted
to one region of the Limnofauna; (C) found in two or three Limnofauna regions, but as
records of single specimens or disjunct populations only. Among species falling into these
categories, those not recorded for 50 years or more are of particular interest. An important
result of this analysis is that more than one third (38 %) of the 85 possibly threatened spe-
cies are typical of standing waters, mostly ponds and ditches, while crenobionts (23 %),
surface dwelling rhithrobionts (20 %) and interstitial species (18 %) are less represented.
However, this result should be checked by investigations on a local scale. In The Nether-
lands, one of the few countries with a documentation sufficient for long term faunistic
monitoring of water mites, a comparison of historical and present day data show a
very high loss of rhithrobiont species (S MIT & VAN DER H AMMEN 1996). Pollution
and/or regulation of streams and stagnant water bodies is obviously the main reason
for the disappearance of many water mite species from extensive areas of the central Eu-
ropean landscape (see above). Each local extinction is correlated with a loss in genetic di-
versity, but due to their generally extended distribution, survival of populations in local
refuges is probable also for rare species of water mites. Future faunistic work should focus
on the importance of such refuge areas. In the systematic part of this work, the particular
risk for the species indicated as potentially threatened by G ERECKE & L EHMANN (2005)
will be discussed.
Special glossary
(!) arrows lead to the general glossary of the introductory chapter to acari, page 33-37.
acetabular plate see „genital field“
anchoral process proximal extension of the (!) gnathosoma serving as an attachment site of mus-
cles which move mouth parts
antagonistic bristle pointed dorsal projection of P-4 flanking P-5; (!) palp.
anterior keel see „ejaculatory complex“
anterior ramus see „ejaculatory complex“
apical setae see „ejaculatory complex“
body pores fine pores in sclerotized plates of Halacaridae and Hydrachnidia, at least in some
Hydrachnidia species facilitating gas exchange processes.
cauda extension of the posterior part of the idiosoma, more or less distinctly set off
from the remainder of the body; frequently found only in males of species
with sexual dimorphism in idiosoma shape; posteriolateral extensions of the
cauda (found mainly in male representatives of Arrenuridae) are called „pygal
lobes“
chelate morphological state of (!) chelicera or (!) palp with two segments acting to-
gether like the branches of scissors.
coxoglandularia see „glandularia“
distal arm see „ejaculatory complex“
dorsal furrow see „dorsal shield“
7. Hydrachnidia 281
dorsal shield large sclerotized shield covering the whole dorsal idiosoma, originating from the
fusion of the (!) „dorsalia“, in general not including (!) „lateral eyes“ and (!)
„lateroglandularia“; in most cases, this shield separated from the (!) ventral
shield by a ring-shaped membranous area called „dorsal furrow“
dorsalia dorsal muscle attachment sclerites; plesiotypically paired, in a total number of
eleven on each side, in a regular pattern following the original segmentation: pre-
and postfrontalia (prefr, postfr) flanking the area of the (!) median eye , some-
times fused with the (!) frontale; five dorsocentralia (dc) paralleling the median
line; four dorsolateralia (dl) arranged more laterally; sclerites may be fused in
various ways; fusion of all sclerites produces the (!) dorsal shield .
dorsocentralia see „dorsalia“
dorsolateralia see „dorsalia“
dorsoglandularia see „glandularia“
ejaculatory complex distal part of the male internal genital system, responsible for the composition
and deposition of (!) „spermatophores“; plesiotypically a complex organ made
up of unpaired keels („anterior, posterior keel“; lat. „carina anterior, posteri-
or“), paired sclerite arms („distal, proximal arm“, lat. brachium distale, prox-
imale; „proximal processus“, lat. processus proximalis), a proximal vesicular
structure („proximal chamber“, lat. cella proximalis), and an apical sclerite
bridge („anterior ramus“, lat. ramus anterior) bearing paired setae („apical se-
tae“, lat. setae apicales), in many clades modified for indirect or direct sperma-
tophore transmission. (= „genital skeleton“; [German] „Penisgerüst“)
genital field area surrounding the opening of the reproductive system („gonopore“), consist-
ing of the following elements: paired sclerotized plates called „acetabular plates“
or „genital plates“ ([German] „Genitalplatten“) when (!) acetabula are located
on their surface, but „genital flaps“ or „genital valves“ ([German] „Genitalk-
lappen“) when acetabula are placed beneath them and can be covered by the
door-like closing flaps; small unpaired anterior („pregenital“) and posterior
(„postgenital“) sclerites delimiting the slit-shaped gonopore; occasionally, a de-
pressed area surrounding the male gonopore is called „genital pit“
genital flap, pit,
plate, valve see „genital field“
genital skeleton see „ejaculatory complex“
glandulae limnesiae see „glandularium“
glandularium defence organ typical for, and a synapomorphy of, Hydrachnidia; consisting of a
round plate bearing a sensory seta and a glandular opening; plesiotypically, glan-
dularia are distributed over the idiosoma in a characteristic ground pattern re-
flecting the original segmentation of the arthropod body: six pairs of „dorso-
glandularia“ flanking the median line, four pairs of „lateroglandularia“ in a
row along the lateral line, four pairs of „coxoglandularia“ placed on or near
the coxae, and four pairs of „ventroglandularia“ surrounding the anal area;
a particular pair of glandularia found in species of the family Limnesiidae on
coxae 4, occasionally called „glandulae limnesiae“; for further terminological
details, see 7-1, 7-2.
hypostomal setae pair of setae on the ventral surface of the gnathosomal rostrum in water mite
larvae
lateroglandularia see „glandularium“
petiole small projecting sclerite at the posterior end of the idiosoma, mainly found in
males of Arrenurus-species, often combined with the formation of a cauda (see
there)
posterior keel see „ejaculatory complex“
postfrontalia see „dorsalia“
282 Key to superfamilies
postgenital sclerite see „genital field“

postocularia paired, knob-shaped, seta-bearing sclerites in the ground plan of Hydrachnidia
located posteriorly from the (!) lateral eyes
prefrontalia see „dorsalia“
pregenital sclerite see „genital field“
preocularia paired seta-bearing sclerites in the ground plan of Hydrachnidia located ante-
riorly from the (!) lateral eyes
proximal arm see „ejaculatory complex“
proximal chamber see „ejaculatory complex“
proximal processus see „ejaculatory complex“
pygal lobes see „cauda“
subchelate morphological state of (!) palp with the terminal segment P-5 acting against P-4
like a clasp-knife (occasionally called „uncate“)
swimming setae elongated setae arranged mainly on III-L-4/5 and IV-L-4/5, more rarely also on
the anterior legs and/or on more proximal segments; in general in rows or as
single elements distally on the anterior segment surface, reaching in length at
least 50 % of the following segment. (occasionally called „swimming hairs“)
uncate see „subchelate“
ventralia paired ventral muscle attachment sclerites; plesiotypically found in water mites
in the circumanal area in a total number of four on each side („ventrale 1-4“);
sclerites may be fused in various ways; fusion of all sclerites and the coxae pro-
duces the (!) ventral shield.
Special part
Area treated
At genus level the keys for Hydrachnidia include Europe as a whole. At species level, the
area treated includes Central and Northern Europe extending in the West to the Southwes-
tern margin of the Pyrenees, in the South to the Southern margin of the Alps and in the East
to the Eastern state borders of Austria, Slovakia, Poland and the Baltic countries.
Key to superfamilies
Apart from Stygothrombioidea, Hydrachnoidea and Hydrovolzioidea, the superfamilies of Hydrach-
nidia are obviously para- or polyphyletic (C OOK 1974, T UZOVSKIJ 1987, PANESAR 2004). Notwith-
standing this fact, for two reasons the system proposed by C OOK (1974) is retained in this work: (1)
This partitioning allows us to keep a survey of the highly diversified families; (2) despite several new
approaches, we are still far from understanding the phylogenetic systematics of the major clades of
Hydrachnidia and newly detected taxa tend to complicate, rather than facilitate, the classificatory
system.
Indications for the identification of preadult stages are given in the paragraphs dealing with families,
genera and species. Surveys on preadult taxonomy have been published for larvae by S PARING (1959:
Palaearctic) and C OOK & S MITH (2000: Nearctic), for deutonymphs by T UZOVSKIJ (1987). A special
volume of this series will be dedicated to the larval stage of Hydrachnidia.
1 Legs with three claw-like, sickle-shaped appendages (7-16 c): paired, ventrally pectinate, true
claws, and an empodium similar in shape, but without pectination; idiosoma vermiform (7-16
a), without muscle attachment sclerites; rod-shaped frontal sclerite (7-13 a) with one unpaired,
and three paired setae; eyes strongly reduced or completely absent; legs covered by rather uniform
hairs over the segment surface („hypertrichous“, 7-16 b-c); palp compact (7-14 a), segments 1-3
fused, segments 4+5 forming a functional unit; P-5 without claws, bearing one stiff terminal seta;
pale reddish species of interstitial waters. . . . . . . . . . . . . . . . Stygothrombioidea (page 289)
7. Hydrachnidia 283
– Legs with two (or occasionally less) claw-like appendages (7-13 h-k): empodium not claw like;
claws rarely with ventral pectination; idiosoma rarely vermiform (7-12 a), with or without muscle
attachment sclerites; if a rod-shaped frontal sclerite is present (e.g., 7-13 b), never bearing an un-
paired anterior seta; eyes various, in most cases with paired lateral eyes, sometimes also with an
unpaired frontal eye (7-13 e, g); legs rarely hypertrichous, in general with distal margin setae stron-
ger than setae on remaining segment surface (7-13 h-k); palp various (7-14 b, e-f, h-i, l-o), P-5
generally with claws, rarely bearing only one stiff terminal seta. . . . . . . . . . . . . . . . . . . . 2
2 Palp chelate (P-4 forming a chela with P-5) and P-1 longer than P-2, P-3 longer than P-4, P-4
reduced in size, maximum height of palps in P-1, the following segments gradually tapering
(7-14 b); gnathosoma in ventral view characteristically narrowed at the base of the slender, pointed
rostrum (7-14 b, d), chelicerae needle-like (basal segment + claw fused to a single unit, 7-14 c);
genital field (7-15 c) with maximum width in the anterior part, bearing numerous acetabula lo-
cated anteriorly from, or on the level of, the anterior margin of the gonopore; rather large, globular
red species of standing waters. . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachnoidea (page 333)
– If palp chelate, neither P-1 longer than P-2, nor P-4 shorter than P-3, maximum height of palps in
general in segments P-2-4 (7-14 e-f, h-i, l-o); gnathosomal rostrum, if present, generally not
pointed (7-14 i), chelicerae two-segmented, not needle-like (7-14 k, p); genital field various in
shape, if numerous acetabula present, they are arranged in a different manner or the genital field
has not its maximum width anteriorly from the gonopore (7-15 a-b, e-o); larger or smaller species,
various in colour and shape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Lateral eyes, if present, incorporated into a frontal sclerite (7-13 b-d, f); palps not chelate (7-14 e-f,
h), in some groups transformed by fusion of segments (e.g., 7-14 f), claws of P-5 straight (7-14 h) or
reduced to fine tips (7-14 f); mouth opening surrounded by a circular, membranous fringe (7-14 g)
or (in Apheviderulicidae, so far not recorded from the study area, 7-14 f) reduced to a fine slit in the
centre of a conspiciously enlarged gnathosoma; acetabula numerous, or on roundish sclerites
flanking the gonopore (7-15 a), or in the membranous perigenital area (7-15 b), or extremely re-
duced in size scattered all over the membranous idiosoma surface (7-15 d); glandular openings
occasionally with semilunar, fan-shaped openings (7-13 f, 7-15 a); large, red species of temporary
and permanent standing waters (outside the study area also pale inhabitants of springs and inter-
stitial waters). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylaoidea (page 301)
– Lateral eyes, if present, not incorporated into a frontal sclerite (but occasionally very close to such
a sclerite, 7-13 g); palps various in shape, P-5 in general with hook-like distal claw setae (7-14 i, l-
o); mouth opening various, but neither surrounded by a circular fringe, nor in the centre of an
extremely enlarged idiosoma; acetabula in three pairs or numerous (7-15 e-o), in rare cases in-
visible by light microscope; glandular openings never fan-shaped (7-13 e, g). . . . . . . . . . . . 4
4 I/II-L directed anteriorly, III/IV-L directed posteriorly, the two leg groups separated by a large inter-
space (7-12 b); genital field minute, in the centre of the ventral idiosoma, devoid of acetabula
(these numerous, mostly invisible in light microscope and inserted as minute cups on the medial
coxal surface); in the postgenital area two large unpaired plates, the anterior one with the anal
pore, or all perigenital plates and platelets fused to a shield embracing the genital field; dorsum
with two shields in „T“ position (7-12 c); glandular openings and setae on paired paletelets (7-12
b, arrow); flattened inhabitants of springs, cascades and interstital waters. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrovolzioidea (page 294)
– If legs arranged in two groups, they are not orientated into two opposite directions, and the inter-
space is less extended (2-4 f-g, 7-12 a, d); genital field various, (with the exception of the marine
Pontarachnidae) always with groups of acetabula arranged in various ways (7-15 e-o); if dorsal
shields present, not arranged in a „T“ position; glandularia and their associated setae together on
one single platelet (7-13 g, arrow) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Palps chelate (7-3 b, d, 7-14 i, arrow: P-4 with a dorsal extension or a heavy dorsal seta dorsally
flanking P-5); idiosoma smooth or with regularly arranged muscle insertion sclerites (7-12 d), only
in exceptional cases with extended dorsal and ventral shields; acetabula in various arrangements
in, or around, the genital field (7-15 e, g-h, l), but never embedded in a sclerotized ventral shield;
diversified morphological adaptations, but mostly larger species, all kinds of inland waters. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydryphantoidea (vol. 7/2-2)
– Palps not chelate (7-14 l-o); idiosoma various; if a sclerotized ventral shield is present, it may
include also some, or all acetabula (7-12 f-h, 7-15 m-o).. . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Palps uncate: P-4 with a ventrodistal extension opposing P-5 (7-14 o) or at least P-4 ventrally
bearing a projecting setal tubercle (7-14 m); idiosoma in general heavily sclerotized (7-12 f); geni-
tal field without movable genital flaps; acetabula numerous, or on fixed plates flanking the go-
nopore (often fused to the ventral shield as given in 7-15 n), or lined up in the membranous area
flanking the gonopore, or both (7-15 m); various in colour and dimensions, mostly in standing
waters, but also in spring and interstitial habitats. . . . . . . . . . . . . . Arrenuroidea (vol. 7/2-3)
– Palps not uncate (7-14 l, n: P-4 without extensions flanking P-5); movable flaps may be present in
the genital field (7-15 f), acetabula in three pairs or numerous - or located between the flaps (7-15
f), or on immovable genital plates (7-15 i-k) - if these plates are shifted to the posterior margin and
variously fused with surrounding sclerites, in ventral view acetabula may be visible tangentially (2-
2 b, 7-12 g, 7-15 o); various in colour and dimensions, all kinds of habitats. . . . . . . . . . . . 7
7 Acetabula located in two rows between the gonopore and movable genital flaps (7-15 f, 2-4 f) (if,
in rare exceptions of species so far not recorded in the study area, they are fused to the flaps, a
suture indicates the former line of separation between acetabula and flaps); if a ventral shield is
present, genital flaps remain always separate and movable. . . . . . . . Lebertioidea (vol. 7-2/2)
– Acetabula located on immovable plates flanking the gonopore, they are fully integrated, no suture
lines are recognizable between acetabula and plate surface (7-12 g-h; 7-15 i, k); acetabula-bearing
plates may be fully integrated into the ventral shield; in this case, acetabula are in three to four
pairs and directed ventrally (7-12 h), or multiplied and directed caudally (7-12 g, 7-15 o).. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hygrobatoidea (vol. 7-2/3)
7. Hydrachnidia 285
Fig. 7-12: Hydrachnidia, habitus; a, Wandesia female, ventral view (from D I S ABATINO et al. 2002); b,
Hydrovolzia male, ventral view (detail: glandulare); c, Hydrovolzia male, dorsal view (arrow: dorsal
furrow), d, Panisopsis male, dorsal view (from D I S ABATINO et al. 2002), e, Sperchon male, ventral
view (detail: glandulare); f, Arrenurus male, ventral view; g, Aturus female, ventral view, h, Ljania
male, ventral view.
Fig. 7-13: Hydrachnidia; a-g, frontal sclerites, h-k, legs; a, Stygothrombium; b, Limnochares; c, d,
Eylais; e, Euthyas; f, Piersigia; g, Hydryphantes; h, Thyas IV-L; i, Hydryphantes (Polyhydryphantes)
IV-L; k, Trichothyas I-L.
7. Hydrachnidia 287
Fig. 7-14: Hydrachnidia, mouthparts; a, Stygothrombium palp; b, Hydrachna gnathosoma and palp
laterally; c, Hydrachna chelicera; d, Hydrachna gnathosoma dorsal view; e, Limnochares palp; f,
Apheviderulix gnathosoma and palps ventral view; g, Eylais gnathosoma ventral view; h, Eylais
gnathosoma and palp lateral view; i, Hydryphantes gnathosoma and palp lateral view; k, Hydry-
phantes chelicera; l, Lebertia palp; m, Mideopsis palp; n, Atractides palp; o, Arrenurus palp; p, Arre-
nurus chelicera.
Fig. 7-15: Hydrachnidia, genital fields; a, Piersigia female; b, Limnochares female; c, Hydrachna
male; d, Eylais male (acetabula in the integument); e, Protzia female; f, Sperchon female; g, Hydry-
phantes s.str. male; h, Hydryphantes (Polyhydryphantes) female; i, Hygrobates male; k, Hygrobates
female; l, Wandesia (Pseudowandesia) female; m, Balcanohydracarus female; n, Arrenurus female;
o, Aturus male.
7. Hydrachnidia 289
Superfamily Stygothrombioidea
1980 Stygothrombioidea M ULLEN & G RANDJEAN , Folia Parasitol. 27: 154.
Di ag no sis: Idiosoma (7-16 a) vermiform. Integument papillate, without sclerotized mus-
cle attachments, often folded. Glandularia openings not on distinct sclerite platelets, but
surrounded by unpapillate integument, each flanked by a pair of setae, one of them long,
the other one more or less reduced (under light microscopy often visible only as a fine pit).
Frontal shield („crista metopica“, 7-13 a) bearing one unpaired anterior seta + three pairs
of setae, no frontal eye, lateral eyes reduced or completely absent. Coxae in four groups, on
each side Cx-I+II and Cx-III+IV fused to two coxal plates, these plates separated by a wide
interspace. Genital field without distinct sclerites, gonopore anteriorly flanked by a row of
fine setae; three pairs of stalked, cupuliform acetabula in perigenital integument; excretory
pore as a longish slit similar in dimensions to genital opening, located near posterior idio-
soma. Gnathosoma (7-16 a, d) large and pyriform, at the end of a protrusible membranous
tube, with orifice directed dorsally and flanked by two pairs of setae on a well developed
rostrum, a further pair of particularly modified setae inserted anteriorly from the palp in-
sertions. Chelicerae with finely curved piercing claws. Palp (7-17) characteristic due to the
fusion of trochanter + femur + genu (P-1-3), segments tibia + tarsus (P-4-5) unfused, but
forming a morphological unit orientated at 90 to the basal segment, a characteristic, nee-
dle-like terminal seta on P-5. Legs (7-16 b-c) with large numbers of rather uniform fine
setae covering the whole surface of all segments, not concentrated near the distal segment
margins; particular whip-like setae in the dorsal centre of segments 4-6, a particular area of
densely arranged fine hairs on the ventrodistal sector of I-L-6 (7-16 b); distal leg segments
with three appendages: a pair of falciform, ventrally pectinate claws and an unpaired em-
podium, not pectinate, but similar in shape to the claws.
Di sc ussi on : In early investigations, members of this group were treated as derivative (or
plesiomorphic) representatives of the terrestrial Trombidiidae (T HOR 1935; A NDRÉ 1949).
As a consequence, information concerning Stygothrombioidea is lacking in important text
books on Hydrachnidia (K.V IETS 1956b, K. O. V IETS 1987) and the significance of the
particular morphology of the members of this group for understanding the phylogeny
of the Hydrachnidia was overlooked for a long time (V ERCAMMEN -G RANDJEAN 1980, T U-
ZOVSKIJ 1987). All known members of Stygothrombioidea are characterized by the pre-
sence of the following diagnostic character combination in both larval, and free-living post-
larval stages (V ERCAMMEN -G RANDJEAN 1980): (1) Setation of the frontal shield (larval
„scutum“, adult „crista metopica“) with seven setae (unpaired anteromedian seta, paired
trichobothria, paired mediolateral setae, paired posterolateral setae); (2) leg segments 3-5
(tibia, genu, femur) each bearing a „stomatoid“ (opening in the sclerotized integument
probably homologue of a slit organ); (3) empodium of legs claw-like; (4) acetabula cupuli-
form; (5) palp with fused segments 1-3 (trochanter – genu); (6) legs with high numbers of
uniform setae („neotrichous“). The rather poor character differentiation in larval and post-
larval stages is one of the reasons to consider the members of Stygothrombidioidea as re-
presentatives of an early derivative clade, possibly the sister group of all remaining Hy-
drachnidia. The presence of some of these characters in other taxa of Hydrachnidia (Lim-
nocharidae, Hydryphantidae, Wandesiinae), but there always restricted to larval or post-
larval stages, is best explained as a consequence of convergent or divergent evolution (V ER-
CAMMEN -G RANDJEAN 1980). Further research should reveal if stomatoids are really homo-
logues of slit organs; presence of slit organs on legs would be a singular feature never ob-
served in other representatives of the Acari.
A still doubtful case is Charonothrombium racovitzai M OTAŞ & TANASACHI , 1946, with a
reduced number of frontal setae. As the type series of this species is missing and the figures of
290 Superfamily Stygothrombioidea
Fig. 7-16: Stygothrombium chappuisi; a, ventral habitus; b, I-L posteriorly; c, IV-L anteriorly; d, an-
terior part of gnathosoma (rostrum and left palp ventrally); e, palp.
7. Hydrachnidia 291
the original description are rather sketchy, this taxon must be considered a species dubia,
possibly incorrectly described or based on a misshapen specimen. A true trombidiid mite
dwelling in interstitial waters is Hygrothrombidium tibiotarsale R OBAUX , 1969, a represen-
tative of the monotypic family Hygrothrombidiinae R OBAUX , 1969 only known from India.
Family Stygothrombiidae T HOR , 1935

1935 Stygothrombiinae T HOR , Zool. Anz. 109: 107-112.
Di ag no sis: Only family of Stygothrombioidea, with the characters of the superfamily.
Genus Stygothrombium K.V IETS , 1932

1932e Stygothrombium K.V IETS , Zool. Anz. 100: 173
1934c Cerberothrombium K.V IETS, Zool. Anz. 106 (5/6). 121
1946 Charonothrombium M OTAŞ & TANASACHI, Notat. biolog., Bucarest 4 (1-3): 7
1969 Hydrothrombium R OBAUX , Ann. Spéléol. 24 (1): 196
1974 Victathrombium H ABEEB, Leafl. Acad. Biol. 61: 1
Typus generis: Stygothrombium karamani K. V IETS , 1932, by original designation.
Di ag no sis: Only genus of the family, with the characters of the superfamily.
Di sc ussi on : A revisional study of larvae and adults of about 16 (still unpublished) species
from North America revealed no distinct character patterns suitable for separation of sys-
tematic units within the members of the family (M ULLEN & V ERCAMMEN -G RANDJEAN
1980, V ERCAMMEN -G RANDJEAN 1980). As a consequence, all formerly described genera
and one subgenus were proposed as synonyms, but with a contradiction within the paper of
V ERCAMMEN -G RANDJEAN (1980): On page 163, his proposal is to retain three subgenera,
Stygothrombium s.str., Cerberothrombium and Baicalacarus S OKOLOW, 1944, while on
page 154 he argues convincingly in favour of a synonymization of Cerberothrombium
with Stygothrombium. The latter conclusion is retained here, with Stygothrombium as
the unique subgenus present in the study area. Due to their vermiform idiosoma, among
the species in the study area Stygothrombium species are most similar to members of the
genera Wandesia and Parawandesia. They are most easily distinguished by the presence of
three leg appendages functioning as claws, the neotrichy on legs, the particular pyriform
gnathosoma bearing a particular seta near the palp insertion, and the number of palp seg-
ments reduced by the fusion of P-1-3.
Pr eadu lt st ages: L arvae: M OTAŞ (1928f), V ERCAMMEN -G RANDJEAN (1980) and
M ULLEN & V ERCAMMEN -G RANDJEAN (1980), all without identification to species level.
Host range: Plecoptera larvae (for details of parasitism see YASICK et al. 2003).
Key to species
Ex clude d a s s pe cie s i nce r tae: Stygothrombium gallicum A NDRÉ , 1949; S. gallicum delphinense
A NDRÉ , 1949 - see discussion of S. chappuisi.
1 Basal palp segment (P-1-3) elongated (length/height 4 2.5); apical seta of the terminal palp seg-
ment long (4 1/3 length P-5) and rod-shaped (7-17 b, e); gnathosomal rostrum elongated (length
4 50 lm) and more slender. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Basal palp segment shorter (length/height 5 2.0); apical seta of the terminal palp segment shorter
(5 1/3 length P-5) and leaf-shaped (7-17 c-d, f); gnathosomal rostrum shorter (length 5 50 lm)
and more robust. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Palp segments rather robust (length/height P-1-3 2.7, P-4+5 5 3.0), ventral setae on P-1-3 re-
stricted to the distal sector, ventral margin of P-4+5 straight (7-17 b). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stygothrombium angelieri (page 293)
292 Superfamily Stygothrombioidea
Fig. 7-17: Stygothrombium, palps; a, indet. species, deutonymph, general scheme of segmentation and
setation (after V ERCAMMEN -G RANDJEAN 1980); b, S. angelieri (after A NDRÉ 1949); c, S. minor (after
A NDRÉ 1949); d, S. gallicum delphinense (after A NDRÉ 1949); e, S. bispinosum (combined from ho-
lotype and S CHWOERBEL 1962, Fig. 1a); f, S. gallicum (species incerta, after A NDRÉ 1949).
7. Hydrachnidia 293
– Palp segments more slender (length/height P-1-3 4 3.5, P-4+5 4 3.0), some of the ventral setae on
P-1-3 located in the proximal sector, ventral margin of P-4+5 concave (7-17 e) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stygothrombium bispinosum (page 293)
3 P-1-3 with two strong ventrodistal setae; medial setae on P-4/5 particularly robust and thickened,
terminal seta pear-shaped (7-17 c) . . . . . . . . . . . . . . . . . Stygothrombium minor (page 294)
– P-1-3 with one strong ventrodistal seta; setae on P-4/5 less robust, terminal seta more slender
(7-17 f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stygothrombium chappuisi (page 293)
1. Stygothrombium angelieri A NDRÉ , 1949 (7-17 b)

1949 Stygothrombium angelieri A NDRÉ , Bull. Mus. Paris 21: 685.
De sc ri pti on : Measurements (original description): Idiosoma length 2900, frontal shield
length 260 lm. Gnathosoma length 420 lm, width 150 lm, rostrum length 80, distal width
60 lm, with a truncate anterior margin laterally extending in two short tips. Palp (setation:
7-17 b) basal segment P-1-3 length 130, P-4+5 length 90, distal seta length 30 lm.
S im i la r sp e c ie s : Stygothrombium armatum K.V IETS , 1934 c, a species known from Ma-
cedonia, agrees in palp proportions, but differs in more slender palp segments, the ventral
setae of P-1-3 located both in the proximal and distal sectors of the segment, a longer distal
seta on P-5 and P-1-3 and P-4+5 having nearly straight dorsal margins.
H a bit at a nd B io lo gy : Hyporheobiont. Life cycle unknown.
Di stri bu ti on: France, eastern Pyrenees, only known from the type locality (Ariège,
Ariège River near the village l’Hospitalet, altitude 1420 m asl).
2. Stygothrombium bispinosum S CHWOERBEL , 1962 (7-17 e)

1962b Stygothrombium bispinosum S CHWOERBEL , Zool. Anz. 168: 294.
De sc ri pti on : Adult unknown; weakly defined species, based on one single deutonymph
and consequently measurement values of little use for comparison with other species. Also
palp setation obviously reduced as compared with adults and not suitable as a diagnostic
character. In the original description, some features of the palp were misinterpreted due to
overlooked segment limits: mistakes pertaining to the position of setae and the presence of a
virtual sclerotized extension (in reality absent). Figure 7-17 e combines the information
from S CHWOERBEL’s Fig. 1 a and new information from the somewhat compressed holo-
type.
S im i la r s pe c ie s: The differences to S. angelieri (see there) and a similar species known
from Macedonia, S. armatum K.V IETS , 1934 c (with a higher number of palp setae, a longer
distal seta on P-5 and nearly straight dorsal margins of palp segments P-1-3 and P-4+5
respectively) need further discussion when the adult of S. bispinosum is detected. All diag-
nostic characters might be subject to changes from deutonymph to adult.
Di stri bu ti on: Austria, Vorarlberg, only known from the type locality (Bizauer Bach near
Bizau).
3. Stygothrombium chappuisi WALTER , 1947 (7-16)

1947 Stygothrombium chappuisi WALTER , Verh. Naturforsch. Ges. Basel 58: 147.
1955 Stygothrombium hercyniense K.V IETS , Arch. Hydrobiol. 50: 58.
De sc ri pti on : (after female specimens from Berchtesgaden and Luxembourg): Idiosoma
(7-16 a) length/width 2000-2300/500-600, Cx-I+II length/width 145-150/100-115;
Cx-III+IV length/width 180-200/83-90. Ac-3 length/width 37-70/23-35. Gnathosoma
294 Superfamily Hydrovolzioidea
length 220-255, basal width 90-95, width at rostrum base 23-24, at rostrum tip 30-31. Palp
(7-16 e) P-1-3 length/height 42-45/27-28; P-4-5 length/height 45-50/23-28, distal seta
length 45-60 lm. Legs (7-16 b-c) with dense setae cover, distal segments of I/II-L rather
variable in proportions, obviously easily inflatable like balloons, length/height I-L-3 85-
100/63-100; I-L-4 98-110/55-125; I-L-5 108-130/55-60; I-L-6 140-165/73-125; II-L-3
65-80/38-45; II-L-4 93-100/35-45; II-L-5 108-120/35-40; II-L-6 130-155/38-40; III-L-3
70-80/33-40; III-L-4 98-110/33-45; III-L-5 115-125/33-40; III-L-6 130-135/33-38; IV-
L-3 103-110/30-40; IV-L-4 125-135/30-38; IV-L-5 140-155/30-35; IV-L-6 145-170/33-
38; IV-L-6 claw length 55-60, empodium length 55-63 lm.
S im i la r s p e ci e s : Stygothrombium karamani K.V IETS , 1932, described from Macedonia
differs in the spherical, not elongate shape of all acetabula. Stygothrombium gallicum A N-
DRÉ , 1949, a species incerta based on a single deutonymph from the Eastern Pyrenees (river
Tech between Prats-de-Mollo and Tech-sur Tech), is rather similar to S. chappuisi in the
shape of the gnathosoma and palp (7-17 f). A series of differential characters in palp seta-
tion and location of frontal shield setae are most probably stage-dependent. Stygothrom-
bium gallicum delphinense A NDRÉ , 1949 (published as „S. g. delphiniensis“) from the
French Alps (Drôme, Ruisseau de Vernaison près des Barraques-en-Vercors, altitude
660 m asl) differs from its stem species only in stage dependent measurements; for the shape
and setation of the palp see 7-17 d. Both taxa are in need of revision, possibly synonyms of
S. chappuisi.
H a bit at a nd B i ol og y: Hyporheobiont, in low and middle order streams. Larvae para-
sitic on stonefly larvae (G ERECKE & M ARTIN 2006).
Di stri bu ti on: Most frequent species of the genus in the study area (G ERECKE et. al.
2005): Romanian Carpathians, Alps, Central European middle elevation mountains, (?)
Pyrenees.
4. Stygothrombium minor A NDRÉ , 1949, nov. stat. (7-17 c)

1949 Stygothrombium angelieri var. minor A NDRÉ , Bull. Mus. Paris 21: 688.
De sc ri pti on : Idiosoma length 2450, frontal shield length 170 lm. Gnathosoma length
270, width 90, rostrum length 50, distal width 50 lm. Palp (setation: 7-17 c) basal segment
P-1-3 length 45, P-4+5 length 50, distal seta length 15 lm, inflated and pear-shaped.
S im i la r spe c ie s: The most similar, S. chappuisi, is distinguished by the lower number of
strong distoventral setae on P-1-3. The difference in shape of the terminal seta of P-4+5 may
depend on the observer’s angle of view - in several specimens of S. chappuisi it was found
flattened and slightly willow-leaf shaped.
Di stri bu ti on: France, Basses-Pyrénées, only known from the type locality (Gave d’Os-
seau at Pont du Germe, one km north from Arudy, altitude 400 m asl).
Superfamily Hydrovolzioidea
1905 Hydrovolziidae T HOR , Zool. Anz. 28: 509.
Di ag no sis: Idiosoma flattened, with two unpaired plates in a „T“ arrangement (anterior
plate wider than long, posterior plate distinctly longer than wide, 7-19), each bearing four
pairs of setae; 2-11 pairs of small platelets in the integument surrounding the central plates.
Median eye absent, lateral eyes generally present, in capsules. Slit organs plesiotypically
present in five pairs. Glandularia with gland openings and associated setae on separate
7. Hydrachnidia 295
platelets; antenniformia, postocularia, and dorsoglandularia 3-6 (trichobothria Fp, Oi;

setae Oe, Hi, Sci, Li sensu T UZOVSKIJ 1987) lacking glandularia, secondary reduction
of further glandularia and idiosoma setae possible. Sclerotization of the ventral idiosoma
surface variable, with four to eight plates (7-20 a) to the formation of a continuous ventral
shield including also the coxae (7-18 a). Genital field and excretory pore sclerite arranged
close to each other in the interspace between the posterior margin of Cx-I+II and the medial
margin of Cx-III+IV. Gonopore flanked by movable flaps, genital field roundish in males
(Fig. 7-18 a, 7-20 a, b), elongate in females (7-20 c); no acetabula in the genital field, but
analogue organs arranged on lateral and posterior surface of all coxae (under light micro-
scopy visible only in slightly sclerotized juveniles). Lateral margins of all coxae, but espe-
cially of Cx-IV, extending over the level of the lateral idiosoma margin. Gnathosoma
(7-20 a) with well-developed rostrum. Chelicerae not fused, consisting of a large basal seg-
ment and a short, crescent-shaped claw. Palps (7-20 d, e) five-segmented, not chelate. Legs
in two distinct groups, with I/II-L directed anteriorly, and III/IV-L directed posteriorly
(7-20 a), without swimming setae, sometimes IV-L with sexual distinctions.
Di sc ussi on : Several former statements about the morphology of hydovolzioid mites were
corrected by recent research (for details see T UZOVSKIJ et al. 2001). The two families
included in Hydrovolzioidea represent two extremely divergent evolutionary lines with
regard to larval behaviour and morphology, but are held together by indisputable synapo-
morphies such as „bipartite“ glandularia and a transfer of acetabula to the coxal surface
in adults. Their taxonomic relations to other clades of water mites are still enigmatic.
Obviously, their similarities to members of the Halacaroidea stressed by several authors
(M OTAŞ 1962b, S CHWOERBEL 1986b) result from convergent evolution in some aspects
of idiosoma morphology (T UZOVSKIJ et al. 2001).
Key to families
1 Coxae close to each other, and forming, together with lateral plates, genital sclerites, excretory,
and postexcretory plates, a continuous shield covering the whole ventral idiosoma surface, suture
line separating Cx-I+II visible only in its lateral part (7-18 a); platelets flanking the mediodorsal
plates often numerous, forming one single line and rather uniform, with little difference in shape
between gland-bearing, and gland-free platelets (7-18 b); no slit organs visible; margins of dor-
socentral shields often with poreform pits; male IV-L six-segmented and with secondary sexual
differences . . . . Acherontacaridae - only genus in the study area: Acherontacarus (page 295)
– Sclerotization of the ventral surface incomplete, with well developed strips of soft integument
surrounding genital field and other plates (7-20 a); suture line separating Cx-I+II usually well de-
veloped; platelets flanking the mediodorsal plates (7-19) often in lower numbers, more hetero-
morphic, occasionally arranged in two lines; slit organs present; no pits on the lateral margins
of dorsal shields; male IV-L without secondary sexual differences or five-segmented. . . . . . . . .
. . . . . . . . . . . . . . . . Hydrovolziidae - only genus in the study area: Hydrovolzia (page 297)
Family Acherontacaridae C OOK , 1967

1967 Acherontacarinae C OOK , Mem. Amer. Ent. Inst. 9: 15.
2001 Acherontacaridae C OOK : T UZOVSKIJ , B ENFATTI & G ERECKE , Acarologia 41: 451.
Di ag no sis: Dorsum with two central plates surrounded 10 pairs of platelets bearing
alternately setae and accompanying glandularia (7-18 b); Posteroventral setae caudales in-
ternae (Vgl-3) and praeanales internae (Vgl-1) always without glandularia. No slit organs
visible. Dorsocentral plates frequently with poreform pits arranged at their margins in spe-
cies-specific numbers (not in the species depicted in 7-18 b). Venter with coxae in four
groups, closely approached to each other, and forming together with lateral plates, genital
sclerites, anal, and postanal plates a continuous shield covering the whole ventral idiosoma
Fig. 7-18: Acherontacarus nicoleiana; a, male ventral view; b, male dorsal view; c, female palp medial
view; d, male IV-L terminal segment; e, female circumgenital field.
surface (7-18 a). Suture line separating coxae I+II visible only in its lateral part. Distal seg-
ments of male IV-L with secondary sexual differences (7-18 d).
Di sc ussi on : The family is monotypic, but the taxonomic placement of Stygovolzia
I MAMURA , 1957 (Japan) and Hydrovolziella WALTER , 1935 (West Africa) is still uncertain
(see discussion of Hydrovolziidae).
Pr eadu lt stages: T UZOVSKIJ et al. (2001), G ERECKE & B ENFATTI (2005).
Genus Acherontacarus K.V IETS , 1932

1962 Acherontacarellus L UNDBLAD, Ark. Zool. (2) 15 (16): 285.
1967 Acherontacaropsis C OOK , Mem. Amer. Ent. Inst. 9: 16.
1989 Neoacherontacarus B ADER , Bijdr. Dierk. 59 (1): 33.
Typus generis: Acherontacarus halacaroides K.Viets, 1932; monobasic.
Di ag no sis: As given for the family.
Di sc ussi on : A total number of 13 species of the genus is known, all recorded exclu-
sively from the Mediterranean basin and Macaronesia (G ERECKE & B ENFATTI 2005,
VALDECASAS , et. al. 2005). Only one species is known from the study area.
Pr ea du lt sta ge s: Larvae and Deutonymphs: T UZOVSKIJ et al. (2001), (G ERECKE &
B ENFATTI 2005). Larvae highly derivative in morphology, attaching under water to the
body surface of water beetles. Host range: Coleoptera (Dytiscidae: Deronectes, Agabus;
Hydraenidae: Hydraena. Like Hydrovolzia larvae, they are regularly found penetrating
7. Hydrachnidia 297
strongly sclerotized parts of their hosts body surface, but without preference of particular
areas: They were found on the elytral surface, on sternal sclerites, at leg segments and also
on the head capsule. Deutonymphs similar to adults in the presence of acetabula on the
coxae, three or more pairs of dorsolateral platelets (in Hydrovolziidae only two pairs),
genital field completely lacking sclerotized structures. Association of deutonymphs to
adults is possible on the base of characters of the dorsal idiosoma and palp.
1. Acherontacarus nicoleiana VALDECASAS , A RTHEAU & B OUTIN , 2005 (7-18)

2005 Acherontacarus nicoleiana VALDECASAS , A RTHEAU & B OUTIN , J. Nat. Hist. 39 (34): 3138.
De sc ri pti on : Idiosoma slender (total length/width males 1294/900, females 1352/843,
postdorsal plate males 890/480, females 965/525 lm). Dorsal plates (7-18 b) without la-
teral pits, setae on postdorsal plate distanced from lateral margin. Anterior coxal plates
medially separated by a distinct suture line. In both sexes, circumgenital field maximum
width posteriorly, on the level of the convexely extending lateral margins of genitalia 3,
anteriorly strongly narrowed, in particular in males, and genitalia 2 formed as minute, tri-
angular platelets (7-18 a). IV-L with sexual dimorphism, in females simple and without
particular setae, in males stout, the terminal segment bearing three rows of strong, var-
iously modified setae (7-18 d). Gnathosoma with ventrodistal setae strong and stout,
but with only one tip, dorsodistal setae more slender. Palp (7-18 c) slender, P-2 particularly
elongated, P-4 with a pair of strong mediodistal setae - the proximal one slightly longer
than the distal one.
S im i la r sp e ci e s : Acherontacarus nicoleiana is most similar to A. dividuus B ADER , 1989,
described from Algeria and only known in the male sex (see G ERECKE & B ENFATTI 2005)
in the absence of lateral pits on the dorsal plates, the predorsal plate relatively narrow,
postdorsal plate caudally narrowed, and in males the two platelets anterior from the genital
field strongly different in size (genitale 2, placed laterally, by far smaller than the medial
genitale 1). Acherontacarus dividuus differs from A. nicoleiana in minor dimensions, idio-
soma and palps stouter in shape and the ventrodistal gnathosoma mitten-shaped, double-
pointed. Acherontacarus ruffoi G ERECKE & B ENFATTI 2005, known from a single female
collected in Andalusia, differs in a more stout idiosoma shape, minor dimensions (e.g. post-
dorsal shield length/width 590/423) and the genitalia 2 rectangular in shape.
Pr ea du lt sta g es: Larva: unknown. Deu to ny m ph: VALDECASAS et al. (2005).
H a bit at an d B i ol og y: Found in a 8.5 m deep well in an agriculturally used Quaternary
flood plain, together with a community characterized by stygobiont crustacean species.
Life cycle unknown.
Di stri bu ti on: France, only known from the type locality (Eastern Pyrenees, river Têt ba-
sin, Corbère, well near Comelade river).
Family Hydrovolziidae T HOR , 1905

1905 Hydrovolziidae T HOR , Zool. Anz. 28: 509.
Di ag no sis: Dorsum, in addition to glandularia, bearing at most three pairs of narrow,
longish, lateral platelets. Posteroventral setae caudales internae (Vgl-3) and praeanales in-
ternae (Vgl-1) associated with glandularia. Five pairs of slit organs present, with i3 and i4
placed on the pair of (sometimes fused) narrow plates flanking the posterodorsal shield; no
pits on the lateral margins of dorsal shields. Venter with well developed strips of soft in-
tegument between coxal plates, and surrounding genital field and other plates and shields.
Coxae arranged in four groups, suture line Cx-I/II usually well developed. Male IV-L with-
out secondary sexual differences or five-segmented.
Fig. 7-19: Hydrovolzia males, dorsal idiosoma; a, H. placophora; b, H. cancellata.
Di sc ussi on : In addition to Hydrovolzia, two further monotypic genera erected for species
from Japan (Stygovolzia I MAMURA , 1957) and West Africa (Hydrovolziella WALTER , 1935)
have been attributed to this family, both insufficiently documented and uncertain in taxo-
nomic placement (T UZOVSKIJ et al. 2001).
Pr eadu lt stages: T UZOVSKIJ et al. (2001) and bibliography cited there.
Genus Hydrovolzia T HOR , 1905

1905 Polyxo M ONTI , Rendiconti Istituto Lombardo di Scienze e Lettere, Milano 38: 168. (preocc.)
1905 Hydrovolzia T HOR , Zool. Anz. 28: 506.
Typus generis: Hydrovolzia halacaroides = H. placophora (M ONTI , 1905); monobasic.
Di ag no sis: Genital flaps in both sexes with rounded, not angular margins (7-20 b-c), in
their anterior half flanked by a pair of glandularia. Male IV-L six-segmented, without sec-
ondary sexual characters.
Pr ea du lt st ag e s: Larvae and Deutonymphs: T UZOVSKIJ et al. (2001) and references given
there. Larvae leave the water after hatching and attach to their hosts in semiterrestrial en-
vironments Host range: Diptera (Empididae); Coleoptera (Staphylinidae, Scirtidae); Hete-
roptera (Mesoveliidae), terrestrial mites (Trombidiidae). For attachment sites, they do not
7. Hydrachnidia 299
Fig. 7-20: a-d Hydrovolzia placophora; a, male ventral view; b, male genital field; c, female genital
field; d, male P-4/5; e, H. cancellata, male palp.
prefer membranous intersegmental surfaces as most Hydrachnidia larvae, but are regularly
found penetrating strongly sclerotized parts of their hosts body surface, mainly leg femora
and tibiae.
Key to species
Excluded as species incertae: H. vietsi C. A NGELIER , 1949a and H. baderi G IL , 1990 (see H. placo-
phora).
1 Mediodorsal sclerites with minute, round pores (diameter 5 10 lm, 7-19 a); postdorsal plate wide
(maximum width 4 300 lm, length/width ratio 5 1.9); postexcretory plate transverse (width 4
190 lm, length/width 5 1.0, 7-20 a); P-4 more stout, with maximum height in the distal part of
the segment (7-20 d). . . . . . . . . . . . . . . . . . . . . . . . . . Hydrovolzia placophora (page 300)
– Structure of the mediodorsal sclerites in the central part lattice-like, with large, often irregularly
shaped pores (diameter 4 10 lm, 7-19 b); postdorsal plate narrow (maximum width 5 300 lm,
length/width ratio 4 2.0); postexcretory plate roundish or slightly elongated (width 5 190 lm,
length/width 4 1.0); P-4 more slender, with maximum height in the centre of the segment (7-20 e)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrovolzia cancellata (page 300)
1. Hydrovolzia cancellata WALTER , 1906 (7-19 b, 7-20 e)

1905 Hydrovolzia cancellata WALTER , Zool. Anz. 30: 570.
1930 Hydrovolzia oscensis K.V IETS, Arch. Hydrobiol. 21 (2): 192.
De sc ri pti on : Dimensions of idiosoma sclerites and appendages considerably variable
(specimens from Tyrrhenian islands and in parentheses holotype after G ERECKE 1991):
length/width predorsal plate 170-230/290-360 (230/380), postdorsal plate 480-640/
190-290 (710/300), excretory plate 100-170/80-160 (158/158); postexcretory plate
140-210/90-175 (246/164) lm; length/height palp segments P-1 8-12/50-52, P-2 85-93/
52-60 (83/50), P-3 62-70/50-61 (62/54), P-4 108-118/38-44 (96/37), P-5 43-46/18 lm.
S im i la r spe c ie s: The synonymization of H. oscensis from Spain and Corsica (G ERECKE
1991) needs verification since on Corsica Hydrovolzia larvae have been found which differ
from continental populations (T UZOVSKIJ et al. 2001).
Pr ea du lt sta g es: Larva and deutonymph: T UZOVSKIJ et al. (2001).
B io lo gy : Larvae parasitic on Diptera (family Empididae, Clinocera, Wiedemannia -
T UZOVSKIJ et al. 2001).
H a bit at : Rhithrobiont, in running waters with well developed hygropetric areas (rheo-
crenes, cascades of low and middle order streams), rare.
Di stri bu ti on: Central and Eastern Mediterranean (Maghreb - Balkan), a single record
from the southern margin of the study area (WALTER 1906b: locus typicus in the Southern
Alps).
2. Hydrovolzia placophora (M ONTI , 1905) (7-19 a, 7-20 a-d)

1905 Polyxo placophora M ONTI , Rendiconti Istituto Lombardo di Scienze e Lettere, Milano 38:
168.
1905 Hydrovolzia halacaroides T HOR , Zool. Anz. 28: 507.
De sc ri pti on : Measurements (specimens from South Italy, G ERECKE 1991): length/width
predorsal plate 180-230/380-440, postdorsal plate 500-600/300-350, excretory plate 130-
170/140-210; postexcretory plate 170-250/200-300 lm; length/height palp segments P-1
8-12/41-45, P-2 73-81/48-52, P-3 70-73/51-58, P-4 111-116/38-41, P-5 41-44/15-17 lm.
S im i la r sp ec i es : The identity of two further species recorded from the Pyrenees, H. vietsi
C. A NGELIER , 1949a and H. baderi G IL , 1990, needs clarification - probably, both are sy-
nonyms of H. placophora.
Pr ea du lt st ag es : Larva: WALTER (1922a), L UNDBLAD (1930a), K.V IETS (1936d), S PA-
RING (1959). D eu ton ym p h: B ADER (1975).
B io lo gy : Larvae parasitic on Diptera of the family Empididae (Clinocera, Roederioides),
Hemiptera (Hebrus ruficeps (T HOMSON )) and Coleoptera of the families Scirtidae (Elodes),
and Staphylinidae (Lesteva) (T UZOVSKIJ et al. 2001).
H a bit at : Crenobiont. In the southern part of the distribution area mostly in seepage wa-
ters with high amount of sand and gravel substrata („rheopsammocrenes“, G ERECKE & D I
S ABATINO 1997); in Central and Northern Europe also in rheocrenes and weakly flowing
rheohelocrenes rich in moss vegetation.
Di stri bu ti on: Europe; frequent in mountains of the central and northern part, more lo-
calized at lower elevations; in the south restricted to the upper parts of higher mountains
(4 1000 m asl.).
7. Hydrachnidia 301
Superfamily Eylaoidea
1815 Eylaidae L EACH , Trans. Linn. Soc., London 11: 399.
Di ag no sis: Red mites with papillose membranous integument. Frontal eye absent, but a
clear area in the central plate may be interpreted as a remnant; a frontal sclerite always
present, generally well developed and including the lateral eye capsules (7-13 b-d, f),
but sometimes (in eyeless species) reduced to a minute sclerotized bar. Coxae in four groups
and these usually widely separated. Acetabula always minute and numerous, or in two
groups on plates flanking the gonopore, or stalked and scattered in the circumgenital
area, or distributed all over the membranous idiosoma integument. Mouth opening sur-
rounded by a frilled ring of very fine hairs („villi“, e.g., 7-14 g-h, 7-25 b - a structure re-
duced in Apheviderulicidae, 7-14 f). Palp leg-like (neither chelate nor uncate); chelicera
two-segmented, claw plesiotypically very stout and dark in colour (7-14 g-h, arrows);
Legs with strong claws not retractable due to lack of a claw furrow. Idiosoma and appen-
dages without obvious secondary sex characters.
Di sc ussi on : Synapomorphic character states of the four highly different clades included
in this superfamily are the stout and strong cheliceral claws darkened in colour, obviously
adapted for penetrating hard sclerotized surfaces, and the fusion of the lateral eye capsules
to the margin of the frontal sclerite. The absence of the first character (Stygolimnocharinae:
lateral eyes reduced) or the second character (Apheviderulicidae: extremely modified
mouthparts) is best explained as an apotypical modification. Members of both groups
display clear sister group relationships with taxa plesiomorphic in the conditions in ques-
tion.
Key to families
1 Gnathosoma unique in shape among water mites (due to partial fusion of basal palp segments with
the gnathosomal base, 7-14 f) being dorsoventrally flattened and broadened distally bearing a
minute, slit-shaped mouth opening, surface with numerous fine, feathered setae; movable parts
of palp appearing unisegmental, but are large, digitiform and work against each other in a ho-
rizontal plane; number of glandularia exceeding the ground pattern of Hydrachnidia. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apheviderulicidae
(only known from larval and deutonymphal stages; monotypic family; not recorded from
the study area; three creno- or hyporheobiont species found in California, Morocco and
Italy - G ERECKE et al. 1999)
– Gnathosoma not flattened, its distal surface formed by the generally large, round mouth opening
surrounded by a fringe of fine villi (e.g., 7-14 g-h, 7-22 a, 7-25 a, b), apart from the two paired
buccal setae without setation on the gnathosomal surface; palps typically five segmented, but oc-
casionally extremely reduced in size and with only 2-4 segments (7-22 b), working parallel to each
other in a vertical plane; idiosoma smooth or with sclerotized muscle attachments as shown in 7-
21 a, 7-23 b, number of glandularia at maximum 16 pairs. . . . . . . . . . . . . . . . . . . . . . . . 2
2 Frontal shield large, including lateral eye capsules and two pairs of glandularia (7-13 f, 7-23 c-d)
(or in eye-less species from outside the study area, minute, rhombic or Y-shaped); dorsal muscle
attachments may be sclerotized large plates or unsclerotized; gonopore flanked by paired, setae-
bearing plates; acetabula may be located on these plates, surrounded by these plates, or lined up
in the membranous integument parallel to the gonopore; chelicera very stout, basal segment
only slightly, or not extending beyond the posterior margin of gnathosoma. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Piersigiidae (page 306)
– Frontal shield never including glandularia, but lateral eyes always present (e.g. 7-13 b-d, 7-21 b);
dorsal muscle attachments may be in the form of fine rods or (generally) be unsclerotized; setae
flanking the gonopore not on plates, in groups on common sockets or free in the membranous
integument; acetabula lying free in membranous integument between gonopore and Cx-IV (7-
15 b, 7-21 c), or distributed all over the idiosoma surface (7-15 d, 7-25 a); chelicera more slender,
basal segment extending far beyond the posterior margin of the gnathosoma (7-14 h). . . . . 3
302 Superfamily Eylaoidea
3 Frontal sclerite rhombic (7-13 b, 7-21 b); Cx-III/IV longer than wide, fused over the length of their
facing borders (7-21 c); stalked acetabula in the perigenital area (7-21 c); European species with-
out swimming setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnocharidae (page 302)
– Frontal sclerite „spectecle-shaped“ (7-13 c-d), consisiting of a transverse bar between the lateral
eyes sometimes extremely reduced so that the eye capsules appear fused side by side (7-13 c), never
extending posteriorly beyond the level of the eye capsules; Cx-III/IV wider than long, fused only
medially, laterally separated by a triangular membranous indentation (7-25 a); acetabula not con-
centrated in the perigenital area, but distributed all over the idiosoma surface as minute, barely
visible cups (7-15 d); legs with swimming seteae. . . . . . . . . . . . . . . . . . Eylaidae (page 310)
Family Limnocharidae G RUBE , 1859

1859 Limnocharidae G RUBE , Arch. Naturk. Liv-, Ehst- Kurlands 1: 434.
Di ag no sis: Colour red, integument papillate; idiosoma inflated and easily deformed by
folding, also in the living animal (7-7 c), often distally tapering. Frontal sclerite (7-13 b,
7-21 b) with a short anterior and a long posterior extension, lateral eyes well developed,
medially placed and fused with the frontal sclerite. Anterior and posterior coxal plates
separated by an extended membranous area; Cx-I+II quadratic or parallelogram-shaped,
Cx-III+IV fused all over their facing borders, with an equally concave medial margin
(7-21 c). Acetabula (20 to more than 200 pairs) small, stalked in the circumgenital field;
genital setae more numerous in males than in females, arranged on elongate platelets or in
the membranous perigenital integument (scattered or bundled in groups). Excretory pore
with or without a sclerotized ring. Gnathosoma (7-22 a) with a circular mouth opening
surrounded by a frilled ring. Basal segments of chelicerae long, medially fused. Palps mi-
nute, not chelate (7-22 b). No sexual dimorphism in legs, claws simple.
Di sc ussi on : This family is well defined by the characteristic rhombic frontal sclerite and
the slender shape of the posterior coxal groups, while all other characters appear to be
plesiomorphic conditions. Members of Limnocharidae are unique among water mites
due to the possibility of supernumerary moults at the nymphal and adult stages (see
B ÖTTGER 1972a; for a discussion of taxonomic consequences see under Limnochares
aquatica). Two subfamilies have been described: Limnocharinae with two genera, dis-
tributed worldwide, and the monotypic Rhyncholimnocharinae recorded from America
and Australia. In the study area, only species of the genus Limnochares (subfamily Lim-
nocharinae) are present.
Subfamily Limnocharinae
1880 Limnocharinae C LAUS, Grundzüge der Zoologie. Zum wissenschaftlichen Gebrauche, etc. 4,
Marburg 1: 653.
Di ag no sis: Gnathosoma not attached to a long, protrusible tube; palp four- or five-seg-
mented.
R em a rk: Two genera are recognized, based on the number of palp segments. Only species
of Limnochares are recorded from the area covered.
Genus Limnochares L ATREILLE , 1796

1796 Limnochares L ATREILLE, Précis de charactères génériques des insectes: 181.
Type species: Acarus aquaticus L INNAEUS , 1758 by original designation.
Di ag no sis: Gnathosoma not attached to a protrusable membranous tube; palp five-seg-
mented.
7. Hydrachnidia 303
R em a r ks : Limnochares includes the subgenera Limnochares s.str. and Cyclothrix

W OLCOTT, 1905. In the study area, only species of the subgenus Limnochares, character-
ized by the absence of swimming setae, are present. They are characterized by the presence
of high numbers of minute, cup-shaped structures distributed over the idiosoma surface.
These elements may represent supernumerary slit organs or glandularia with reduced
setae. In older specimens they can be more sclerotized, but are easily distinguished
from true muscle attachment sclerites in their irregular arrangement and high numbers.
Key to species.
1 Dorsum without sclerotized muscle attachments; anterolateral edges of Cx-I forming an obtuse
angle (7-22 c - but internal apodemes present, without setae, distinctly projecting and bent in-
wards); a high number of acetabula (60 to more than 200 pairs) scattered along several lines
in the area between the medial margin of Cx-IV and the gonopore; idiosoma length up to
5000 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnochares aquatica (page 303)
– Dorsum with sclerotized muscle attachments (7-21 a: three or four paired rod-shaped sclerites, one
pair of minute roundish sclerites, one elliptical posterodorsal sclerite generally unpaired); ante-
rolateral edges of Cx-I forming a digitiform projection with dense setae cover directed anterolat-
erally (7-21 c); acetabula in lower numbers (15-50, exceptionally up to 80 pairs); idiosoma length
5 2000 lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnochares azubi (page 304)
1. Limnochares aquatica (L INNAEUS , 1758) (7-7 c, 7-22 c)

1758 Acarus aquaticus L INNAEUS, Systema naturae ed. 10 reformat., 1: 617.
1778 Acarus holosericus D E G EER , Mémoires pour servir à l’histoire des Insectes: 149.
De sc ri pti on : Idiosoma length 1000-5000 lm; very variable in shape, somewhat rectan-
gular in a resting state, without sclerotized muscle attachments. Frontal shield rather slen-
der (length/width 4 500/250 lm, ratio 4 2.3). Genital field in males with numerous setae
bundled in four distinct groups, in females with fewer hairs scattered over the area flanking
the gonopore; a spherical internal structure at the posterior end of the female called „ovi-
positor“ by K.V IETS (1936d), but more probably involved in spermatophore uptake. Ex-
cretory pore not surrounded by sclerotization. Gnathosoma with apex of rostrum slightly
curved ventrally, P-5 retracted within P-4 and terminating distally in a seta. Legs without
swimming setae and incapable of swimming. Measurements of males: length/width Cx-I+II
(excluding the anterior apodeme) 315-430/145-220, Cx-III+IV 340-520/250-430; palp
segments length P-1 14-16, P-2 55-80, P-3 40-50, P-4 50-60, P-5 20-30, total length
185 lm. Measurements of a female: length/width Cx-I+II (excluding the anterior apodeme)
315-610/150-270, Cx-III+IV 410-740/135-270; palp segments length P-1 15, P-2 92, P-3
58, P-4 70, P-5 38, total length 270 lm.
Pr eadult stages: Larva: WAINSTEIN (1980). D eu ton ym p h: T UZOVSKIJ (1990).
S im i la r sp e ci e s: Not confusable with any species recorded from the study area, but taxo-
nomic state of the worldwide populations attributed to L. aquatica in need of revision (see
„Biology“).
B io lo gy : B ÖTTGER (1972a) gives an extensive description of the life-history. Adults and
nymphs appear year-round, feeding on chironomid larvae, in particular on representatives
of tube-dwelling species which are attacked by piercing their heads with the help of the
strong cheliceral claws (7-7 c). Oviposition in Central Europe, April-August, in one to three
clutches per female (including 15-500, generally 4 50 red eggs, egg diameter 170-230 lm)
on plants or other substrates. Neighbouring eggs in clutches not fusing, remaining isolated
from each other by distinct envelopes. Larvae hatch after four weeks at 20 C, then swim
to the water surface and run about in search of a host; characteristic shadow reflex (in a
shadow they freeze and then lift themselves up and wave their first pair of legs in the air).
Hosts are reached by climbing, not jumping; emergence of parasitic larvae May-September.
Host range: Heteroptera adults and larvae (Gerridae, Hydrometridae, Mesoveliidae, at-
tachment sites hardly specific, but thorax preferred), Odonata adults (P ETZOLD & M ARTIN
2004 and bibliography cited there, but rejected as hosts in experiments of B ÖTTGER
1972a). Duration of the parasitic stage only 5-10 days, parasite load/host specimens gen-
erally low (one to five), rarely up to 20; on growing Heteropteran larvae, L. aquatica lar-
vae, unique in water mites, are capable to detach before the host’s moult and fix again on
the newly moulted host; moult to the protonymphal stage after detachment from the host,
hidden in crevices of the sediment; duration of the protonymphal stage one to four weeks.
Before reaching sexual maturity, depending on the nutritional state of the individuals one or
two eight-legged nymphal stages may be passed, but no tritonymphal resting stage has been
observed. Instead, again unique among water mites, adults can become inactive and enter
into resting stages with supernumerary moults, explaining the observation of egg-bearing
resting stages by L UNDBLAD (1927c). Limnochares aquatica can reach an age of several
years. Details of the morphology of L. aquatica and their change occurring during the
supernumerary nymphal and adult moults are not yet well documented. Consequently, di-
agnostic features are far less defined than we should expect for one of the most conspicuous
water mite species of our study area. Also, the presence of undetected sister species hidden
behind the name of L. aquatica is possible.
H a bit at : Riparian sediments of permanent stagnant waters or pools of streams, often in
decaying wood, roots and leaf litter; also in eutrophic or acidic water bodies; less common
on sandy bottoms.
Di stri bu ti on: Holarctic.
2. Limnochares azubi G ERECKE , 2005 (7-21, 7-22 a-b)

2005 Limnochares azubi G ERECKE , Zootaxa 106: 52.
De sc ri pti on : Idiosoma length 900-1600 lm; generally one unpaired and four paired dor-
sal sclerites with tuberculate surface, but occasionally additional muscle attachments can
be found. Frontal shield short and robust (length/width in males 200-300/100-150, in
females 230-350/130-200 lm, ratio 1.6-2.4). Anterolateral edge of Cx-I with a digitiform
extension densely covered by fine setae. Cx-I maximum length/anterior margin in males
265-315/139-180, in females width 275-460/139-234 lm. Genital field with 15-80 (gen-
erally 5 50) pairs of acetabula, mostly arranged in a line parallel to the medial margin of
Cx-IV; genital setae in both sexes not arranged in groups; excretory pore generally unscler-
otized, but in one particularly large specimen (possibly after a supernumerary adult moult)
with a strong sclerite ring. Palp five-segmented, P-4 relatively minute.
Pr ea du lt sta g es: L a rv a: not described. De uto ny m ph : D AVIDS et al. (2005).
S im i la r sp e c ie s : Limnochares danielei G ERECKE & S MIT, 2005, known only from Cy-
prus, differs in having enlarged anterior coxae, a sclerotized ring surrounding the excretory
pore of all specimens, and a relatively longer and more inflated P-4 (D AVIDS et al. 2005).
Habitat & Biology: Helocrenes and low order streams, from 200 to 1660 m asl. Life cycle
unknown.
Di stri bu ti on: West Mediterranean (Italy, Corsica, Spain, Morocco), one record from
the southernmost edge of the study area: Italy, Trentino-Alto Adige („L. aquatica“, err.,
C REMA et al. 1996).
7. Hydrachnidia 305
Fig. 7-21: Limnochares azubi female (after D AVIDS et al. 2005); a, dorsal view; b, frontal sclerite; c,
coxal field and genital field.
Fig. 7-22: (a-b) Limnochares azubi male (after D AVIDS et al. 2005); a, gnathosoma lateral view; b,
palp; c Limnochares aquatica male, anterior coxae.
Family Piersigiidae O UDEMANS, 1902

1902 Piersigiidae O UDEMANS, Tijdschr. Entomol. 45: 61.
Di ag no sis: Frontal sclerite large, including two pairs of glandularia (7-23 c-d) or (in eye-
less species) reduced to a single rhombic or Y-shaped platelet bearing the postocularia; dor-
sal muscle attachments unsclerotized or forming well sclerotized symmetrically arranged
plates (7-23 a). Posterior margin of Cx-IV with short, pointed apodemes. Acetabula from
five to numerous pairs, lying on two paired genital plates (genera Stygolimnochares and
Piersigia) or lined up in the membranous integument flanking the gonopore (Parawande-
sia). Chelicera short and stout, basal segment only slightly extending beyond posterior mar-
gin of gnathosoma. Legs without swimming setae.
Fig. 7-23: Piersigia; a, P. intermedia female, ventral view (after L UNDBLAD 1962); c-d, frontal shield
(after K.V IETS 1936d); b, P. koenikei, dorsal view (after K.V IETS 1936d); c, P. limophila female;
d, P. intermedia female.
7. Hydrachnidia 307
Key to genera
1 Idiosoma vermiformly elongated and, apart from the minute frontal plate, without sclerotization,
lateral eyes absent; gonopore flanked by a pair of small setae-bearing plates and not more than 10
pairs of acetabula lying free in the integument; P-4 with a strong peg-like mediodistal seta. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parawandesia E. A NGELIER , 1951
Monotypic genus in Stygolimnocharinae C OOK , 1967; not recorded from the study area: Para-
wandesia chappuisi E. A NGELIER , 1951, hyporheobiont, Corsica, Sardinia (G ERECKE & C OOK
1995).
– Idiosoma oval in shape, with a large frontal plate including the well developed lateral eyes and two
pairs of glandularia and lines of symmetrically arranged dorsal muscle insertion sclerites (7-23 b);
gonopore flanked by two pairs of large plates each bearing numerous acetabula (7-24 a-d); P-4
without a peg-like seta (subfamily Piersigiinae). . . . . . . . . Piersigia P ROTZ , 1896 (page 307)
Subfamily Piersigiinae
1902 Piersigiinae O UDEMANS, Tijdschr. Entomol. 45: 61.
Di ag no sis: Frontal plate large, including lateral eyes and two pairs of glandularia. Dor-
sum with numerous symmetrically arranged muscle insertion sclerites. Gonopore flanked
by two pairs of large plates each bearing numerous acetabula. Palp segment P-4 without a
peg-like seta.
R em a rk: Apart from the genus Piersigia, including species recorded from Europe and
North America, the subfamily includes only one further genus, Austrapiersigia S MIT,
1998, described from the Australian region.
Genus Piersigia P ROTZ, 1896

1896 Piersigia P ROTZ , Zool. Anz. 19: 410.
Typus generis: Piersigia limophila P ROTZ , monobasic.
D iagnos i s : Frontal shield large, formed by the fusion of several plates, in some species still visible as
elements connected by sclerotized bridges, including two pairs of glandularia and the capsules of well
developed lateral eyes. Muscle attachments sclerotized as symmetrically arranged plates; a clear area in
the frontal shield may represent a vestigial median eye (C OOK 1974). Glandularia sclerites only par-
tially surrounding the glandularia; lateroglandularia platelets much larger than dorsoglandularia pla-
telets. Acetabula on minute stalks, lying in four groups in the soft integument, surrounded by ring-
shaped plates. Excretory pore on a large sclerite. Palp segment P-3 with several pectinate distal setae.
Pr ea du lt st ag e s: L ar va : WAINSTEIN (1980), I MAMURA & M ITCHELL (1967a). Deu to -
nymph : T UZOVSKIJ (1990); I MAMURA & M ITCHELL (1967a), with reduced number of
acetabula and a more trapezoid sclerite between anterior and posterior genital field.
Key to species
1 Posteromedial extension of frontal shield triangular, usually tapering posteriorly; this part of
shield in direct contact (continuous porosity) with the two pore plates located anteriorly (7-23 c).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Piersigia limophila (page 308)
– Posteromedial extension of frontal shield rectangular or tongue-shaped, not tapering posteriorly,
this part of shield not in direct contact (interrupted porosity) with the two pore plates located
anteriorly (7-23 b, d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Genital plates touching, anterior plates with more than 50, posterior plates with more than 60
acetabula (7-24 a, c); palp slender (7-24 f) . . . . . . . . . . . . . Piersigia intermedia (page 308)
– Genital plates separated, anterior plates with up to 20, posterior plates with up to 30 acetabula
(7-24 d); palp stocky (7-24 e). . . . . . . . . . . . . . . . . . . . . . . . . Piersigia koenikei (page 308)
1. Piersigia intermedia W ILLIAMSON , 1912 (7-23 a, d, 7-24 a, c, f)

1912 Piersigia intermedia W ILLIAMSON, Proc. Phys. Soc. 18: 251.
1954 Piersigia americana H ABEEB, Leaflets Acad. Biol. 2: 1.
De sc ri pti on : Ma l e: length 1450-1630, width 1050-1060 lm. Area between genital
plates with many long setae. Fe m a le : length 1800-2400, width 1140-1920 lm. Medial
plate of lateral eyes consisting of one large plate, with concave margins in the middle, and
two anterolaterally smaller plates. Anterior and posterior genital plates touching. Anterior
genital plate with more than 50, posterior plate with more than 60 acetabula. Area between
genital plates with few, relatively short setae. Palp slender; P-2 with two or three setae in
ventrodistal corner, P-3 with five to seven setae at distal margin. Unpaired medial plate of
eye consisting of one large plate, with two smaller ones laterally.
Pr ea du lt sta g es: L a rv a: not described. De uto ny m ph : L UNDBLAD (1962).
H a bit at an d B io lo gy : Reported from a wide range of permanent as well as temporary
habitats (VAN M AANEN et al. 1997), in eutrophic and dystrophic waters, with a pH around
6.5; preferably in temporary and semi-aquatic biotopes (terrestrializing waters, quagfens
and reedlands), often reported from Sphagnum-pools. Life cycle unknown.
Distribution. Holarctic. Widespread in Europe, but not known from southern Europe.
More recent records: Sweden (L UNDBLAD 1962), Russia (T UZOVSKIJ 1986a), The Nether-
lands (VAN M AANEN et al. 1997).
2. Piersigia koenikei K.V IETS , 1909 (7-23 b, 7-24 d-e)

1909 Piersigia koenikei K.V IETS, Abh. naturw. Ver. Bremen 19: 453.
De sc ri pti on : Male reported only once (H ALBERT 1944, without measurements): genital
plates narrower and more elongate than in female, the porose area between the two acet-
abula-bearing areas more oblique, 18 acetabula on each of the anterior, 24 on each of the
posterior plates. F e m al e: length 1360-1670, width 940-1060 lm; body somewhat taper-
ing anteriorly. Medial plate of lateral eyes consisting of one large plate, in the middle with
concave margins, and two anterolaterally smaller plates. Two pairs of genital plates; an-
terior and posterior plates not touching. Anterior plate with up to 20 acetabula, posterior
plate with up to 30 acetabula. Palp relatively short.
H a bit at an d B io lo gy : Reported from a forest pool, a spring brook, a pond and a semi-
permanent ditch; given its occurrence with other temporary water species, very likely with a
preference for habitats of that type. Life cycle unknown.
Di stri bu ti on: Germany, Ireland and Ukraine (Bukowina). A record from The Nether-
lands (VAN M AANEN et al. 1997) refers to an erroneous identification of P. intermedia
(S MIT et al. 2003); rare.
3. Piersigia limophila P ROTZ , 1896 (7-23 c, 7-24 b, g)

1896 Piersigia limophila P ROTZ, Zool. Anz. 19: 410.
De sc ri pti on : Mal e : length 1670, width 1330 lm. Unpaired median plate of lateral eyes
slightly narrower in male than in female, of variable shape, somewhat triangular and
slightly tapering posteriorly, occasionally reduced in size and not tapering. Anterior and
posterior pair of genital plates touching. Anterior pair of plates elongate, posterior pair
angular to circular. Genital plates with about 60-65 acetabula (in N-American specimens
posterior pair with about 18-57 acetabula, anterior pair with 30-55 acetabula). Anterior
genital plates with a straight or convex medial margin. Medial sclerotized space of genital
field with numerous long setae. Palp elongate, with a dorsal row of four to five pectinate
7. Hydrachnidia 309
setae on P-2 and and a meso-ventral row of five to eight pectinate setae on P-3 (data from
N-American specimens, I MAMURA & M ITCHELL 1967a). Fe m al e : length 2000-2270,
width 1400-1570 lm. Both pairs of genital plates with about 80 acetabula (in N-American
specimens with 38-67 in posterior plate and 38-64 in anterior plate). Anterior genital plates
with a somewhat geniculate medial margin. Medial space between genital plates with few,
short setae.
Fig. 7-24: Piersigia; a-d genital field; a, P. intermedia female (after L UNDBLAD 1962); b, P. limophila
female (after K.V IETS 1936d) ; c, P. intermedia male (after L UNDBLAD 1962); d, P. koenikei female
(after K.V IETS 1936d); e-g palp; e, P. koenikei female (after K.V IETS 1909a); f, P. intermedia male (after
K.V IETS 1936d); g, P. limophila female (after I MAMURA & M ITCHELL 1967).
Pr ea du lt st ag e s: L ar va : WAINSTEIN (1980), I MAMURA & M ITCHELL (1967a). Deu to -

nymph : T UZOVSKIJ (1990); I MAMURA & M ITCHELL (1967a).
B io lo gy : Egg-development: one to two months. Larvae move fairly slowly on the surface
of the water, and do not jump as do thyasid larvae, parasitic on hydrophilid water beetles
(genus Enochrus): attached to the dorso-lateral membranes of the abdomen where they
remain attached also during the protonymphal stage, leaving the host as a fully developed
deutonymph (data from North America: I MAMURA & M ITCHELL 1967a). These records
suggest that P. limophila is relatively long-lived and capable to survive unfavourable per-
iods during the phoretic phase of its life cycle. Adults are active when the water level is
adequate, but probably suspend their activity when water levels are low.
H a bit at : Semi-aquatic habitats in the mud of shallow marshes and swamps with a fluc-
tuating water level (I MAMURA & M ITCHELL 1967a). The species is able to creep in the
narrow spaces among masses of sodden rotting vegetation. It can crawl in open water
or burrow itself in the mud, and is never found far away from the water margin. S MIT
& VAN DER H AMMEN (2000) reported the species from small puddles in reedlands.
Di stri bu ti on: Holarctic. Widespread, but rarely reported. In Europe so far known from
Germany, France, Poland, Russia and The Netherlands (S MIT & VAN DER H AMMEN 2000).
Family Eylaidae L EACH , 1815

1815a Eylaidae L EACH, Trans. Linn. Soc., London 11: 399.
Di ag no sis: Colour red; integument soft, subtly lineated. Dorsum without dorsalia. La-
teral eyes in two groups of two, each group enclosed in a capsule, these close together and
usually joined medially by a sclerotized intercapsular bridge forming together a „spectacle-
like“ median eye plate. The eye bridge bears a pair of setae. Coxae in four groups, third and
fourth coxae separated except near the medial margins. Genital field well forward in ven-
tral surface with numerous, small, scattered acetabula. Gonopore in males simple, only
surrounded by a pair of small setae-bearing sclerites, except in males belonging to the sub-
genus Syneylais which have a well developed sclerite surrounding the gonopore. Excretory
pore usually surrounded by a narrow, sclerotized ring. Gnathosoma with a large circular
mouth opening. Mouth in a wheel-like membrane fringe with setae forming a suctorial disk
on the ventral surface of gnathosoma, well developed air sacs associated with capitulum.
Pharyngeal plate usually conspicuous. Blunt cheliceral claws project through the mouth
opening. Palp five-segmented, not chelate and bearing numerous, often pectinate setae.
P-2 and P-3 usually with an extensive area of articular membrane on the ventral side. First
three pairs of legs with numerous swimming setae. Leg claws simple. Fourth pair of legs
without swimming setae and trailed behind as the animal swims.
R em a r ks : Eylais is the only genus occurring in the area covered. The only other genus in
the family, the monotypic Rhyncheylais L UNDBLAD , 1938, has only been recorded from
South America (Paraguay).
Genus Eylais L ATREILLE, 1796

1796 Eylais L ATREILLE , Précis de charactères génériques des insectes: 182.
Diagnosis: Characters of the family Eylaidae. Idiosoma usually oval to egg shaped. Coxae
for the most part narrow, sclerotization often forming a latticework framed by a distinct
ridge. Gnathosoma characterized by two paired proximal apodemes in dorsal and ventral
position (7-14 h). Chelicerae with blunt tips projecting into the mouth. P-2, P-3, as well as II-
L and III-L ventrally with extensive areas of poorly developed sclerotization. P-3 usually
with a distal prominence and thereon a group of setae, P-5 usually shorter than P-4.
7. Hydrachnidia 311
Fig. 7-25: Eylais sp.; a, male, ventral view; b, male gnathosoma, lateral view.
Pr ea du lt sta ge s: L ar va : A general description is given by WAINSTEIN (1963b, 1980).

See also P RASAD & C OOK (1972), L ANCIANI (1969), N IELSEN & D AVIDS (1975). De uto -
ny m ph : Similar to adult but lacking gonopore, eye bridge less sclerotized. Integument
dorsally with transverse striation, those in the adult running longitudinally (B ESSELING
1964)
B io lo gy : Larvae are aerial and after hatching crawl through the surface film to seek their
aquatic hemipteran or coleopteran host. The larvae remain attached from a couple of
weeks to eleven months, depending on the season and their life cycles. Often a bivoltine
life cycle pattern is found with a regular pattern of a summer generation with short-lived
larvae and a winter generation with long-lived, hibernating larvae (L ANCIANI 1970). The
protonymphal phase is passed on the host and the mites emerge as deutonymphs. Adults
and deutonymphs feed on ostracoda or (more frequently) cladocerans (rare reports on
the uptake of both groups of microcrustacea by the same species) (K UDRINSKAJA 1950,
W OHLTMANN 1991). A detailed analysis of the functional morphology of the mouth
apparatus is given by Z HAVORONKOVA (1992).
H a bit at : In shallow standing waters, temporary and permanent ponds, between the ve-
getation, occasionally in lakes and in slowly running waters or pools of fast flowing
streams.
Di stri bu ti on: Cosmopolitan.
R em a rks : The genus Eylais is very distinct, but the species are highly variable and there-
fore often difficult to separate on adult morphology. The problem is that most descriptions
are, besides pharyngeal plate and palps, mainly based on the shape of the eye bridge, which
varies considerably within a species (S OKOLOW 1930a; L UNDBLAD 1962; K.O.V IETS
1949b, 1950a,b). The same holds for the palps. They may be subject to sexual dimorphism
and even left and right palps of one individual may differ considerably (S OAR & W ILLIAM-
SON 1925-1929; K.O.V IETS 1949b). Furthermore, one has to keep in mind that females are
larger, not only in the lengths of pharyngeal plate, palps and legs, but also in the number of
setae on the individual palp and leg segments. In the first decades of the 20th century, va-
riation of these characteristics was hardly known. As a consequence, many species were
described of which the majority later fell into synonymy. Larvae are less variable and show
more reliable characters for separating species. In view of the particularly confused taxo-
nomic situation, and contrary to other genera, a more extended synonymy is given here for
Eylais. However, for a complete synonymy the reader is also here referred to K.V IETS
(1956b) and K.O.V IETS (1987).
The subgeneric classification is based on larval characteristics (WAINSTEIN 1968) and the
structure of the pharyngeal plate (C OOK 1974). In accordance with K.O.V IETS (1987), no
reference will be made to this taxonomic level, but the subgenera occurring in the region
covered are briefly described:
Subgenus E yl ai s s. str. L ATREILLE , 1796: Posterior part of pharyngeal plate with a pha-
ryngeal ridge (e.g., 7-30 a, e, h-i). Male gonopore surrounded by two sclerotized plates (As
given in 7-32 k for a species of Meteylais, see below). Known larvae with one pair of setae
and six pairs of setal bases on the dorsal plate. (The subgenus Pareylais S ZALAY, 1934 was
synonymized with Eylais s. str. by WAINSTEIN 1963b, based on larval characteristics). Spe-
cies: Eylais bisinuosa, E. extendens, E. mutila, E. planipons, E. rimosa, E. setosa, E. tan-
tilla, E. tullgreni.
Subgenus S yn e yl ai s L UNDBLAD , 1936: Characteristics of pharyngeal plate as in Eylais s.
str.; male gonopore surrounded by a single sclerotized plate (7-32 h, l-m). Species: Eylais
discreta and E. infundibulifera.
7. Hydrachnidia 313
Subgenus Prote y la is S ZALAY, 1933: Posterior end of pharyngeal plate lacking a ring-like
area, pharyngeal plate only indistinctly set off from the posterolateral projections of the
gnathosoma (7-30 c). Species: Eylais degenerata.
Subgenus M et ey l ai s S ZALAY, 1934: Posterior end of pharyngeal plate generally lacking a
ring-like area (7-30 b, f). Male gonopore as in subgenus Eylais (7-32 k). Seven pairs of setae
on the larval dorsal shield. Species: Eylais curvipons, E. koenikei, E. hamata.
Subgenus S pi ne yl ai s WAINSTEIN , 1968: Based on larval characteristics only, cannot be
fitted into an adult system. The larval gnathosoma is greatly enlarged and possesses a well
developed, sharply pointed anterior projection which extends well anterior to distal end of
palps. Species: Eylais glubokensis (species incerta, see discussion of E. extendens).
Table 7-1: Genus Eylais; a list of species incertae recorded from the study area and bordering terri-
tories (following K. V IETS , 1956b).
The original descriptions of the following species are insufficient (important details missing or in-
complete, location of type material unknown, type material damaged or lacking). For each taxon,
the nation named first includes the type locality.
Abbreviations:B = Belgium; CS = Czech Republic and Slovakia; D = Germany; E = Spain; GB = Great
Britain; HU = Hungary; I = Italy; N = Norway; NL = The Netherlands; F = France; IRL = Ireland;
PL = Poland; RO = Romania; RUS = Russia; S = Sweden; YU = former Yugoslavia.
E. affinis D ADAY, 1901 RUS
E. akmolinski gracilipalpis WALTER, 1927 F
E. angustipons T HOR, 1899 N, CS, RUS
E. babori T HON, 1899 CS
E. bicornuta H ALBERT, 1904 IRL, GB, E (= infundibulifera?)
E. cavipontia G EORGEVITSCH, 1906 YU
E. celtica H ALBERT, 1903 IRL
E. clitellata G EORGEVITSCH, 1906 YU, I
E. cornuta T HOR, 1901 N
E. dividua S OAR, 1901 GB
E. dubia D ADAY, 1901 HU
E. duplex T HOR, 1899 N, YU ( = mutila?)
E. elliptica P IERSIG, 1904 RUS (= extendens?)
E. emarginata P IERSIG, 1899 RUS, Europe, Asia (= extendens?)
E. eugeni T HON, 1903 HU, NL, D
E. gigas P IERSIG, 1904 RUS, D, GB, S (= discreta?)
E. hungarica D ADAY, 1901 HU
E. infundibulifera acuta D ADAY, 1901 HU, D
E. insularis T HOR, 1899 N, CS, IRL, S
E. krendowskii P IERSIG, 1904 RUS (= extendens?)
E. latipons T HON, 1899 CS, N, RUS, HU
E. media M USSELIUS, 1912 PL
E. muelleri bifissa D ADAY, 1901 RUS (= extendens?)
E. muelleri mrázeki S ZALAY, 1934 (nec T HON , 1903) HU (= extendens?)
E. neglecta T HOR, 1899 N, Europe, Asia
E. occulta T HOR, 1899 S, N, D (monstr.?)
E. prelipceani H USIATINSCHI, 1937 RO
E. producta D ADAY, 1901 HU (= infundibulifera?)
E. saratovi P IERSIG, 1904 RUS, B, POL
E. semipons T HOR, 1926 D
E. skorikowi P IERSIG, 1904 RUS
E. symmetrica H ALBERT, 1903 IRL, RUS (= extendens?)
E. voeltzkowi D ADAY, 1901 (non K OENIKE, 1897) RUS, Asia (= extendens?)
E. wilsoni S OAR, 1917 GB, D, E, USA (monstr.?)
Key to species
The synonymy in this genus is very confusing, because of great variation in shape and measurements of
the characters. Many species are incompletely described, sometimes being based only on the shape of
the eye plate. Many authors have described deutonymphs instead of adults, which causes confusion
e.g. regarding the shape of the eye bridge and the measurements of the palp segments. In certain species
separating females from deutonymphs is difficult. Be aware that males are smaller than females, a
difference most obvious in eye plates and palps. The smaller the species, the less are differences in
measurements between males and females. Only males or ovigerous females provide reliable determi-
nations! Table 7-1 gives a list of species incertae excluded from the key.
In reference to the species listed for the study area by K.O.V IETS (1978), the following changes are
important (for discussion of details see D AVIDS et al. 2005):
Excluded: Eylais bergstroemi WALTER, 1911 and E. incurvata K.V IETS, 1911 (both syn. to E. koenikei:
L UNDBLAD , 1962); E. glubokensis U DALZOV, 1907 (species dubia - lack of an adequate description of
the adult). Now included: (considered by former authors as synonyms or species incertae, but recently
differently interpreted - or recent new records) Eylais bisinuosa, E. planipons, E. rimosa, E. telma-
tobia, E. triarcuata and E. tullgreni.
1 Eye bridge wide (4 1.5 times the width of an eye capsule - in critical cases check E. telmatobia)
and thin (7-28 b, g) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Eye bridge more narrow (5 1.5 times the width of an eye capsule), often thickened (e.g. 7-28 a, c)
.................................................................. 3
2 Eye bridge straight (7-28 g), pharyngeal plate enlarged and posteriorly strongly rounded, with a
ring-like thickened area bearing two knob-shaped extensions (7-30 f) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais hamata (page 326)
– Eye bridge V shaped (7-28 b), pharyngeal plate over the whole length about equal in width, poster-
iorly without extensions (7-30 b) . . . . . . . . . . . . . . . . . . . . . . . Eylais curvipons (page 323)
3 Distance between the eye capsules about 1.5 times greater than width of one eye capsule (7-29 g);
gnathosoma (7-31 e) with a very large mouth disk; pharyngeal plate large, at the level of the pos-
terior gnathosoma apodemes, with a narrowing followed by lateral extensions . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais telmatobia (page 331)
– Distance between the eye capsules equal to, or less than, the width of one eye capsule (e.g., 7-28 a,
c); mouth disk minor, pharyngeal plate various in shape . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Gnathosoma (7-30 c) posterior to the mouth disk forming only a narrow bridge of porous scler-
otization (not larger than the posterior apodemes of the gnathosoma); pharyngeal plate only
indistinctly separated from the posterior apodemes, strongly dilated as in E. hamata, but lacking
a ring-like thickened area and posterior projections . . . . . . . . . Eylais degenerata (page 323)
– Gnathosoma with an extensive area of porous sclerotization posterior to the mouth opening (cov-
ering an area larger than the radius of the mouth disk, e.g., 7-30 d, g, h); pharyngeal plate distinctly
separated from the posterior apodemes, various in shape . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Reticulation on coxae forming rounded meshes (7-32 b, c) . . . . . . . . . . . . . . . . . . . . . . . . 6
– Reticulation on coxae forming elongated meshes mostly extending between the anterior and pos-
terior margins (7-32 a, d, e). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 Anterior margin of eye bridge convex, on the level of, or slightly projecting beyond, the anterior
edge of the capsules (7-28 k ); length of eye capsule 4 200 lm; female genital field with paired
plates bearing long setae (7-32 f) . . . . . . . . . . . . . . . . . . . . . . . . . Eylais mutila (page 328)
– Anterior margin of eye bridge straight or concave, posterior to the anterior edge of the capsules
(7-28 i); length of eye capsule 5 200 lm; female genital field without paired plates, a few setae
scattered in the circumgenital field (7-32 g) . . . . . . . . . . . . . . . . Eylais koenikei (page 327)
7 Males (ejaculatory complex showing through the integument in the genital area): gonopore sur-
rounded by a sclerotized plate, which is bulging out ventrally (7-32 h, l-m). Females (no ejacu-
latory complex visible, in ovigerous specimens eggs showing through the integument): one pair of
rather large plates flanking the gonopore, with 15 or more long setae (7-32 n) . . . . . . . . . . 8
– Males: genital field various, but gonopore not surrounded by a single genital plate. Females:
genital plates absent or hardly visible, with at most 15 short setae. . . . . . . . . . . . . . . . . . . 9
7. Hydrachnidia 315
8 Eye capsules (7-28 d) anteriorly enlarged and including the neighbouring setae in their porose area;
eye bridge generally projecting anteriorly, but sometimes not extending beyond the level of the eye
capsules. Male genital plate with maximum width anteriorly and three posterior extensions, setae
in two groups (7-32 l-m). Female genital plates parallel or only slightly diverging posteriorly (7-
32 n) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais discreta (page 324)
– Eye capsules (7-28 h) anteriorly not enlarged, no setae included in their porous area; eye bridge
with a pointed medial extension projecting anteriorly to the level of the eye capsules. Male genital
field with maximum width in the centre, many setae extending all over the medial area surround-
ing the gonopore (7-32 h). Female genital plates strongly diverging posteriorly (7-32 i) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais infundibulifera (page 327)
9 Pharyngeal plate with maximum width at an extension in the posterior quarter (occasionally ma-
ximum width reached in the middle and from here to the end lateral margins subparallel (7-30 e,
7-31 b, f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Pharyngeal plate with maximum width in the middle (7-30 a, i, 7-31 a, c-d, g) . . . . . . . . 12
10 Pharyngeal plate (7-31 b) stout, posterior width 4 250 lm, setal papillae on anterior margin of eye
bridge bulging out and flanking a slit-like medial indentation (7-29 c) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais rimosa (page 330)
– Pharyngeal plate more slender, posterior width 5 250 lm; anteriomedial indentation of eye bridge
absent, if present not slit-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Gnathosoma slender, porosity in the area posterior to mouth disk rather homogenous (7-30 e),
transverse ridge on the pharyngeal plate strongly developed; P-4 medially with about 5-6 pectinate
setae restricted to the distal part of the segment (7-26 c, arrow) . Eylais extendens (page 325)
– Gnathosoma more robust, porosity in the area posterior to mouth disk differentiated in an anterior
part with large pores, distinctly separated from a posterior part with smaller pores (7-31 f), trans-
verse ridge on the pharyngeal plate often less pronounced; P-4 medially with more than 15 pecti-
nate setae, some in proximal half of segment (7-27 e, arrows). . . Eylais triarcuata (page 332)
12 Eye bridge with a more or less pointed extension reaching beyond the level of the anterior edge of
the eye capsules; seta papillae about mid length of eye bridge (7-29 b). . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais relicta (page 329)
– Eye bridge with anterior margin not extending beyond level of anterior edge of eye capsules; seta
papillae near its anterior margin (e.g., 7-29 f, i) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Eye bridge not deeply indented anteriorly (7-29 d); on the ventral gnathosoma an anterior area
with large pores, separated from a posterior area with small pores by a distinct separation line
halfway between mouth disk and posterior margin (7-31 c); palp (7-27 b): P-3 with no clear pro-
jection, P-4 medially with many setae, P-5 longer than 1/2 P-4. . . . . Eylais setosa (page 330)
– Eye bridge with a more or less distinct anterior indentation (e.g. 7-29 a, i) ; on the ventral gnatho-
soma no distinct separation line between an anterior area with large pores and a posterior area
with minute ones (e.g. 7-30 i, 7-31 d); palp various in shape and setation . . . . . . . . . . . . 14
14 Diameter of mouth disk less than distance between posterior margin of mouth disk and posterior
margin of gnathosoma (measured along the median line, 7-30 a); setae on projection of P-3 needle-
like (7-26a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais bisinuosa (page 323)
– Diameter of mouth disk greater than distance between posterior margin of mouth disk and pos-
terior margin of gnathosoma (measured along the median line, 7-30 i, 7-31 d, g); setae on projec-
tion of P-3 short and broad (7-26 i, 7-27 d, f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Eye bridge anteriorly hardly indented (7-29 a) porous area of eye capsules anteriorly enlarged,
occasionally enclosing the neigbouring setae . . . . . . . . . . . . . . . Eylais planipons (page 329)
– Eye bridge with a deep anterior slit (7-29 f, i), porous area of eye capsules anteriorly not enlarged,
not enclosing the neigbouring setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Length of pharyngeal plate 4 _ 300 lm, length of P-4 about 300 lm (palp: 7-27 d) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais tantilla (page 331)
– Length of pharyngeal plate 5 300 lm, length of P-4 5 250 lm, eye bridge very small with a deep
anterior slit (palp: 7-27 f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eylais tullgreni (page 332)
Fig. 7-26: Eylais, palp; a, E. bisinuosa female; b, E. discreta female; c, E. extendens female; d, E.
hamata male; e, E. hamata female; f E. infundibulifera female; g, E. koenikei male (after L UNDBLAD
1962); h, E. mutila male; i, E. planipons female; k, E. relicta male.
7. Hydrachnidia 317
Fig. 7-27: Eylais, palp; a, E. rimosa female; b, E. setosa female; c, E. extendens male; d, E. tantilla
female; e, E. triarcuata female; f, E. tullgreni female.
Fig. 7-28: Eylais, frontal sclerite; a, E. bisinuosa male; b, E. curvipons male; c, E. degenerata female; d,
E. discreta male; e, E. extendens male; f, E. glubokensis female (after U DALZOV 1907); g, E. hamata
female; h, E. infundibulifera male; i, E. koenikei female; k, E. mutila female.
7. Hydrachnidia 319
Fig. 7-29: Eylais, frontal sclerite; a, E. planipons female; b, E. relicta male; c, E. rimosa female; d, E.
setosa male; e, E. extendens male; f, E. tantilla female; g, E. telmatobia male (after S ZALAY 1956b); h,
E. triarcuata female; i, E. tullgreni male.
Fig. 7-30: Eylais, gnathosoma in ventral view; a, E. bisinuosa female; b, E. curvipons female (after
S OKOLOW 1925a); c, E. degenerata female; d, E. discreta male; e, E. extendens male; f, E. hamata
female; g, E. infundibulifera male; h, E. mutila male; i, E. planipons male.
7. Hydrachnidia 321
Fig. 7-31: Eylais, gnathosoma in ventral view; a, E. relicta male; b, E. rimosa female; c, E. setosa male;
d, E. tantilla female; e, E. telmatobia male (after S ZALAY 1956b); f, E. triarcuata female; g, E. tullgreni
male.
Fig. 7-32: Eylais; a-e, sculpturing of coxae; a, E. extendens female; b, E. koenikei male; c, E. mutila
male; d, E. tantilla female; e, E. setosa female; f-n, genital field; f, E. mutila female; g, E. koenikei
female; h, E. infundibulifera male; i, E. infundibulifera female; k, E. hamata male; l, E. discreta
male, ventral view; m, E. discreta male, lateral view; n, E. discreta female.
7. Hydrachnidia 323
1. Eylais bisinuosa P IERSIG , 1899 (7-26 a, 7-28 a, 7-30 a)

1899a Eylais bisinuosa P IERSIG , Zool. Anz. 22: 62.
1919d Eylais bisinuosa nodipons V IETS, Arch. Naturg. 83: 160.
De sc ri pti on : Frontal sclerite width 240-280 lm, anterior margin with broad medial in-
dentation, muscle attachments generally bulging out anteriorly, setae close to anterior mar-
gin. Gnathosoma slender and long, most of the upper portion with large pores, pharyngeal
plate elliptical in outline. Mouth disk diameter 120-150 lm, P-3 prominence with 6-12
needle-like setae, some of them pectinate, P-4 length 200-230 lm, medially with three
smooth and four to six pectinate setae, four or five of them near the distal margin.
Ma le : Idiosoma length 1500-3000. Fe m al e : Idiosoma length 2500-3600 lm.
P re a d ul t st a g e s : L a r v a: WAINSTEIN (1963b, 1980). D e u t on y m ph : T UZOVSKIJ (1990).
S im i la r spe c ie s: Various authors synonymize E. bisinuosa with E. tantilla and others
consider E. bisinuosa as a species incerta. However, the characters presented here in
the key allow for a clear distinction of the species.
H a bit at an d B io lo gy : Small eutrophic water bodies. Larvae parasitic on the water bee-
tle Hydraena palustris E RICHSON (Hydraenidae) (Z AWAL 2004).
Di stri bu ti on: Palaearctic.
2. Eylais curvipons S OKOLOW, 1922 (7-28 b, 7-30 b)

1922 Eylais curvipons S OKOLOW, Fauna Petropolitanae Cat. 5 (2): 21.
1922 Eylais longipons I SOSSIMOFF (non D ADAY, 1901), Arb. Student.-Ver. Naturfr. Univ. Kasanj 2:
45.
De sc ri pti on : Pharyngeal plate elliptical in outline, posteriorly without hooks. Fe m al e :
length 3700, width 3000; eye capsule length/width 210/130, frontal plate width 590 lm.
Male unknown.
Pr eadult stages: Larva: WAINSTEIN (1963b, 1980). Deutonymph not described.
H a bit at an d B i olo gy : Ponds (Sweden: L UNDBLAD 1962). Oviposition in June, larvae
hatch in December.
Di stri bu ti on: NE Europe (recorded from Russia, Lithuania, Poland and Sweden).
3. Eylais degenerata K OENIKE , 1897 (7-28 c, 7-30 c)

1897a Eylais degenerata K OENIKE , Abh. naturw. Ver. Bremen 14: 292.
1906 Eylais pseudorimosa P IERSIG, Zool. Jb. Syst. 23: 380.
1902c Eylais variabilis T HOR, Ann. South African Mus. 2: 450.
1903a Eylais mrazeki T HON (non muelleri mrazeki S ZALAY, 1934), S. B. böhm. Ges. Wiss. II. Cl. 19: 3.
1911a Eylais angulata K.V IETS, Zool. Anz. 37: 155.
1911a Eylais galeata K.V IETS, Zool. Anz. 37: 156.
1912 Eylais consors S ZALAY, Állat. Közlemén 11: 70.
1912 Eylais eregliensis S ZALAY, Állat. Közlemén 11: 73.
1912 Eylais stagnalis S ZALAY (non H ALBERT, 1903), Állat. Közlemén 11: 77.
1914g Eylais taurica K.V IETS, Arch. Hydrobiol. 9: 560.
1939 Eylais variabilis H USIANTINSCHI (non T HOR, 1902), Zool. Anz. 127: 104
De sc ri pti on : Frontal sclerite anterior margin variable, often with a slight indentation;
posterior margin sometimes slightly bulging. Eye capsule length 160-250 lm. Sculpturing
of coxae as in E. extendens (compare 7-32 a). P-3 bulging, with long pectinate setae, P-4
medially with about eight setae (five, located in the distal part, pectinate), P-5 more taper-
ing than in other species. M al e: Idiosoma length 2100 – 4000 lm. F em a l e: Idiosoma
length 2500 – 4000 lm.
R em a r k: K URT O.V IETS (1950a, see there) synonymized all subspecies described in the
first half of the 20th century.
H a bit at a nd B iol og y : In small ponds and pools of lowland streams, often in temporary
waters. Life cycle unknown.
Di stri bu ti on: Reported from Southern Europe, South Africa, Asia Minor, India and Chi-
na. In Central Europe one record from Slovakia (L ÁSKA 1964b).
4. Eylais discreta K OENIKE , 1897 (7-26 b, 7-28 d, 7-30 d, 7-32 l-n)

1897a Eylais discreta K OENIKE , Abh. naturw. Ver. Bremen 14: 292.
1897c Eylais spinipons T HOR , Arch. Math. Naturv. 20: 8. (= deutonymph)
1897c Eylais tenuipons T HOR , Arch. Math. Naturv. 20: 7.
1899g Eylais discissa T HON , Zool. Anz. 22: 441.
1899g Eylais meridionalis T HON , Zool. Anz. 22: 445
1908a Eylais aurita K OENIKE , Abh. naturw. Ver. Bremen 19: 254.
1911 Eylais discreta stagnalis H ALBERT, Proc. Irish Acad. Dublin 31: 5.
1919d Eylais infundibulifera meridionalis K.V IETS , Arch. Naturg. 83: 158.
1935c Eylais porosipons K.V IETS , Zool. Anz. 110: 273.
1962 Eylais infundibulifera meridionalis B ÖTTGER , Zool. Jahrb. Syst. 89: 501.
De sc ri pti on : Pharyngeal plate close to equal in width over the total length or ellipsoid.
Gnathosoma with large pores except for a small strip at the posterior border. P-3 with a
weak protuberance, bearing 12-14 long pectinate setae, P-4 with numerous setae (7-26 b).
Ma le : Idiosoma length 2700-3400 lm; F e m al e: Idiosoma length 2600-5200 lm.
Pr eadu lt stages : Larva: B ÖTTGER (1962a); N IELSEN & D AVIDS (1975). D eu to-
ny m ph : not described.
B io lo gy : Bivoltine. Adults are found from spring to late summer (in The Netherlands from
mid-April until September, with a peak in late summer, S MIT & VAN DER H AMMEN 2000).
Direct sperm transfer by alignment of the genital openings (B ÖTTGER 1962, see there also
for a detailed description of developmental times of all stages). Females can produce up to
13.000 eggs over a period of two to three months (D AVIDS 1973a). Larvae of the first an-
nual generation hatch in early spring, as an exception also in winter. After reaching their
host, they stay for two to four days between head and pronotum before crawling to the
abdominal tergites (B ÖTTGER 1962). They parasitize corixid species of the genera Corixa,
Callicorixa, Hesperocorixa, Sigara and Cymatia (N IELSEN & D AVIDS 1975, B IESIADKA &
C ICHOCKA 1994). Larval growth is slow in winter and early spring, but considerable from
mid-March onwards. L ANCIANI (1971) showed that about 15 % of the total individual
growth takes place at the larval stage. Larvae of the autumn generation remain attached
to their hosts for a long period and hibernate on their host. In April they leave their hosts as
deutonymphs (D AVIDS et al. 1977b). W OHLTMANN (1991) reports on both Cladocera and
Ostracoda as typical food of Eylais discreta, while in experiments of S CHEIDEGGER (1999)
ostracods were not accepted. In laboratory experiments, the latter author found very high
daily consumption rates on individuals of Daphnia magna (13 mg per diem in deuto-
nymphs and 35-68 mg in adults, with an uptake of up to 305 individuals per diem
when prey of the first instar was offered, and up to 18 individuals per diem when large,
last instar prey was available).
7. Hydrachnidia 325
H a bit at : Shallow lakes, ponds and ditches rich in macrophytes (B ÖTTGER 1962a). Ac-
cording to S MIT & VAN DER H AMMEN (2000) in fresh, relatively nutrient-poor waters.
Di stri bu ti on: Palaearctic, widely distributed especially in northern Europe and Siberia
(L UNDBLAD 1968).
5. Eylais extendens (M ÜLLER , 1776) (7-26 c, 7-27 c, 7-28 e, 7-29 e, 7-30 e, 7-32 a)
1776a Hydrachna extendens M ÜLLER , Zool. Dan. prod.: 190.
1897a Eylais undulosa K OENIKE , Abh. naturw. Ver. Bremen 14: 283.
1897a Eylais muelleri K OENIKE , Abh. naturw. Ver. Bremen: 282.
1899a Eylais limnophila P IERSIG , Zool. Anz. 22: 63.
1899a Eylais soari P IERSIG , Zool. Anz. 22: 67.
1899g Eylais similis T HON, Zool. Anz. 22: 446.
1899g Eylais tenera T HON , Zool. Anz. 22: 444.
1899 Eylais mosquensis C RONEBERG, Bull. Soc. Natural. 13: 97.
1902 Eylais unisinuata C RONEBERG , Bull. Soc. Natural. 16: 98.
1903b Eylais instabilis H ALBERT, Ann. Mag. Natur. Hist. 12: 510.
1910b Eylais similis gibberipons K.V IETS , Abh. naturw. Ver. Bremen 20: 171.
1907b ? Eylais glubokensis U DALZOV, Arb. Hydrobiol. Stat. See Glubokoje 2: 232.
1911 Eylais sugambra K OEP, Verh. naturh. Ver. preuß. Rheinl. Westfal., Bonn: 276
1911a Eylais thienemanni K OENIKE , Zool. Anz. 37: 321.
1919a Eylais soari valentiana K.V IETS , Abh. natur. Ver. Bremen 24: 23.
1934a Eylais extendens monofissa S ZALAY, Ann. Mus. Nation. Hungar. 28: 277.
1939 Eylais szalayi H USIATINSCHI, Zool. Anz. 127: 102.
De sc ri pti on : Frontal sclerite with eye bridge not reaching beyond the anterior margin of
eye capsules, considerably varying in shape (7-28 e, 7-29 e), sometimes enlarged as in E.
triarcuata (7-29 h); setae insertions may bulge out the anterior margin. Sculpturing of coxae
as in 7-32 a. Gnathosoma entirely covered by large pores except for a narrow strip at the
posterior edge. P-3 strongly bulging out, with 10-22 smooth or pectinate setae; P-4 distally
with at least three pectinate setae, medially with at least five smooth setae; P-5 hardly half
the length of P-4. M al e: Idiosoma length 2200-3500. Fe m al e : Idiosoma length 1500-
5000. Pharyngeal plate maximum width 250 lm. In the circumgenital field between
the first coxae 10-15 setae of two different size groups. The long ones surrounding the
anterior part of the genital opening sometimes seem to be inserted on a small plate, which
is hard to see.
Pr eadu lt stages: L arva: WAINSTEIN (1963b, 1980), N IELSEN & D AVIDS (1975) - see
also under „Similar species“. De ut ony m ph : not described.
S im i la r spe c ie s: A detailed analysis of the variability and synonymy of E. extendens was
published by K.O.V IETS (1949b). The larvae described by WAINSTEIN (1980) under the
names of E. muelleri and E. mosquensis, two species synonymized with E. extendens
by K.V IETS (1956b), are rather similar to larvae of the latter species. As M ARTIN
(2003) stated, morphological features such as length and width of the larval dorsal plate
may be subject to considerable variability within the progeny of one adult. Another prob-
lem is that WAINSTEIN ’s figures are depicted in various styles, so that comparison becomes
difficult. Currently both E. mosquensis and E. muelleri should be considered synonyms of
E. extendens. Eylais glubokensis U DALZOV, 1907 was considered a species incerta by
K.V IETS (1956b), but described at the larval stage by WAINSTEIN (1968, 1980) and later
recorded from Poland by C ICHOCKA (1983) and Z AWAL (2004) from larvae which were
found parasitic on the coleopteran Limnebius truncatellus T HUNBERG (Hydraenidae). As

the description of the adult of this species does not provide details useful for separating it
from E. extendens, the reasons for attributing the very characteristic larvae to that species
remain unclear. Rearing experiments are necessary in order to determine if E. extendens
represents a complex of species which can only be separated at the larval stage (D AVIDS et
al. 2005).
B io lo gy : Eylais extendens is possibly bivoltine (Z AWAL 2004) and hibernates as egg or as
parasitic larva (N IELSEN & D AVIDS 1975). Larvae were first recorded parasitic on water
beetles of the family Haliplidae (Haliplus, N IELSEN & D AVIDS 1975), later on 26 different
water beetle species of the families Dytiscidae (Acilius, Bidessus, Colymbetes, Graptodytes,
Hydroporus, Hyphydrus, Scarodytes), Haliplidae (Haliplus, Peltodytes), Noteridae (No-
terus), Laccophilidae (Laccophilus) and Hydrophilidae (Anacena, Laccobius, Helochares,
Enochrus) (Z AWAL 2004). S PARING (1959) recorded E. extendens larvae parasitic on
corixids. However, there are doubts with regard to their correct taxonomic assignment
(D AVIDS et al. 2005).
H a bit at : In all types of standing and slowly flowing waters, permanent and temporary,
oligotrophic and eutrophic, dystrophic and calcareous (L UNDBLAD 1968), but sensitive to
high electrolyte concentrations and therefore absent from very brackish waters.
Di stri bu ti on: Western Palaearctic; one of the most common species of the genus in
Europe and Asia Minor.
6. Eylais hamata K OENIKE , 1897 (7-26 d, 7-28 g, 7-30 f, 7-32 k)

1897a Eylais hamata K OENIKE , Abh. naturw. Ver. Bremen 14: 282.
1922a Eylais hamata alpina WALTER, Rev. Suisse Zool. 29: 247.
1901d Eylais georgei S OAR, Sci.-Goss. (n.s.) 8: 68.
1901a Eylais longipons D ADAY, Math. Természtr. Értes. 19: 94.
1905b Eylais marenzelleri T HON, Ann. Naturh. Hofmus. 20: 158.
De sc ri pti on : Frontal sclerite dumb-bell-shaped, eye bridge about twice the width of an
eye capsule, narrow and straight or slightly bent posteriorly, anterior margin slightly un-
dulating. Gnathosoma short, length about twice the width. Pharyngeal plate extending
considerably beyond posterior margin of gnathosoma, with two small (not always distinct)
extensions. P-3 hardly bulging. Great variety in eyebridge and length of palp segments be-
tween males and females. M al e: Idiosoma length 2400-5500 lm. Genital field with two
rod shaped setae-bearing plates. Palp rather stout (7-26 d). F em a l e: Idiosoma length
3300-7000 lm. Palp more slender and with higher numbers of setae (7-26 e).
Pr eadult stages: Larva: WAINSTEIN (1963b, 1980). De uto ny m ph: Differing remark-
ably from adult in the shape of P-5, T UZOVSKIJ (1990), G ERECKE (1991).
S im i la r s p e ci e s : S EZEK & ÖZKAN (2000) give measurements of a male, a female and a
deutonymph of E. hamata from Turkey and argue for a separation of the sister species E.
rahmi ÖZKAN , 1982 which was first described from a deutonymph. Eylais rahmi should
differ in generally major dimensions of adults, but minor dimensions of the deutonymph,
and a more slender eye bridge in all active postlarval instars. However, as the dimensions of
the holotype of E. rahmi described by ÖZKAN (1982) fit the variability range given by S E-
ZEK & ÖZKAN for the deutonymph of E. hamata, it is most probably a synonym of the
latter. In view of the generally high variability of dimensions in Eylais adults, there are
good reasons to attribute both populations described from Turkey to E. hamata. Other-
wise, the taxon published by S EZEK & ÖZKAN as E. rahmi would merit a new name.
7. Hydrachnidia 327
B io lo gy : Adults found in springtime, disappearing after the beginning of June (S MIT &
VAN DER H AMMEN 2000). Larvae hatching in August (WAINSTEIN 1963b), parasitic on
water beetles of the family Dytiscidae (species of Dytiscus and Cybister: P IATAKOV
1915a, b) and Helophoridae (Helophorus flavipes FABRICIUS : Z AWAL 2004).
H a bit at : All types of small standing waters.
Di stri bu ti on: Palaearctic, widely distributed in Europe, but never abundant. Also known
from northern Africa, Siberia, China and Asia Minor (L UNDBLAD 1968).
7. Eylais infundibulifera K OENIKE , 1897 (7-26 f, 7-28 h, 7-30 g, 7-32 h-i)

1897a Eylais infundibulifera K OENIKE , Abh. naturw. Ver. Bremen 14: 284.
1899a Eylais bifurca P IERSIG, Zool. Anz. 11: 64.
1899g Eylais meridionalis T HON, Zool. Anz. 22: 445.
1901b Eylais dadayi P IERSIG, in: P IERSIG & L OHMANN , Acarina, Hydrachnidae und Halacaridae:
308.
1901d Eylais projectus S OAR, Sci.-Goss. (n.s.) 8: 69.
1901a Eylais infundibulifera acuta D ADAY, Math. Természtr. Értes. 19: 84.
1901a Eylais incisa D ADAY, Math. Természtr. Értes. 19: 87.
1930d Eylais infundibulifera lundbladi K.V IETS, Arch. Hydrobiol. 21: 200.
De sc ri pti on : Frontal sclerite with eye bridge (7-28 h) small but variable in width, with a
pointed anterior extension and posterior margin with a deep bay; eye capsules longer than
wide. Palp (7-26 f) stout, P-3 thick, slightly bulging with numerous short, smooth and pec-
tinate setae, P-4 with numerous medial setae, most of them pectinate, P-5 stout and apically
blunt, with characteristic nail-like distal setae. Gnathosoma with large pores in the anterior
half, separated distinctly from a posterior area of smaller pores; oral disk large, pharyngeal
plate in the middle as wide as it is posteriorly. Gonopore on a cone surrounded by many
stout, long setae. Ma le : Idiosoma length 3000-4000. F em a le : Idiosoma length 3500-
5000 lm. Genital plates each bearing 10-28 setae. Palp much larger and with many
more setae than in male.
Pr eadu lt s tages: L arva: WAINSTEIN (1962, 1980), N IELSEN & D AVIDS (1975). De u -
to ny m ph: not described.
B io lo gy : Larvae parasitic on Corixidae of the genera Corixa, Sigara and Cymatia (N IEL-
SEN & D AVIDS 1975; B IESIADKA & C ICHOCKA 1994), attached to the abdominal tergites.
Hibernating possibly as egg or attached to the host as parasitic larva or protonymph. The
statement by P IATAKOV (1915 a, b) concerning parasitism on beetles is surely erroneous.
H a bit at : Standing eutrophic waters, especially lowland ponds, lakes and ditches, but also
in alpine waters. Not very common. Reported from calcareous waters by L UNDBLAD
(1968), from relatively nutrient-poor waters by S MIT & VAN DER H AMMEN (2000).
Di stri bu ti on: Holarctic. Widely distributed in Europe, especially northern and central
Europe. Also reported from Turkey, Asia and Canada (L UNDBLAD 1968).
8. Eylais koenikei H ALBERT, 1903 (7-26 g, 7-28 i, 7-32 b, 7-32 g)

1903b Eylais koenikei H ALBERT, Ann. Mag. natur. Hist., London 12: 507.
1911b Eylais incurvata K.V IETS, Abh. Naturw. Ver. Bremen 20: 339.
1911a Eylais bergstroemi WALTER , Naturwissenschaftliche Untersuchungen des Sarek-Gebirges in
Schwedisch-Lappland, geleitet von Dr. Axel Hamberg 4: 588.
De sc ri pti on : Frontal sclerite with seta insertions away from anterior margin of eye
bridge. Eye capsules length/width about 200/120 lm, with medial margin various in shape
(convex as given in 7-28 i or concave). Coxae reticulate. Gnathosoma with area posterior to
mouth disk equal in length to diameter of mouth disk; pharyngeal plate broad, somewhat
distended in its anterior portion, lateral margins parallel, posteriorly with strong ring-like
area. Palps rather long and slender; P-3 slightly bulging with seven or eight setae, a few of
them pectinate; P-4 medially with 7-12 long setae. Mal e : Idiosoma length 2200. Fe m al e :
Idiosoma length 1600-4000. Eye sclerite width 372, eyebridge length/width 127/105 lm. A
few setae in the anterior part of the genital field.
Pr eadult stages: Larva: WAINSTEIN (1980). D eu ton ym p h: T UZOVSKIJ (1990).
B io lo gy : Larvae parasitic on dytiscid beetle species of the genera Hydroporus, Grapto-
dytes and Hyphydrus (Z AWAL 2004). In The Netherlands, adults found mainly in spring-
time (S MIT & VAN DER H AMMEN 2000)
H a bit at : Eutrophic ponds and pools of slowly running waters, peat bogs, coastal dune
waters. In springtime colonizing astatic waters (L UNDBLAD 1968, Z AWAL 2004).
Di stri bu ti on: Palaearctic. Rare; not recorded from Southern Europe.
9. Eylais mutila K OENIKE , 1897 (7-26 h, 7-28 k, 7-30 h, 7-32 c, f)

1897a Eylais mutila K OENIKE , Abh. naturw. Ver. Bremen 14: 285.
1897c Eylais foraminipons T HOR , Arch. Math. Naturv. 20: 8.
1907b Eulais longipalpis U DALZOW, Arb. Hydrobiol. Sat. See Glubokoje 2: 230.
1911 Eylais setipalpis K.V IETS , Abh. Naturw. Ver. Bremen 20: 341.
1919a Eylais cordifera K.V IETS , Abh. Naturw. Ver. Bremen 24: 7.
1930d Eylais rimosa proceripalpis K.V IETS, Arch. Hydrobiol. 21: 204. (= deutonymph)
1931a Eylais rimskykorssakowi S OKOLOW, Zool. Jahrb. Syst. 61: 467.
De sc ri pti on : Frontal sclerite with eye bridge situated anteriorly between the eye capsules,
longer than one capsule in width, anterior margin convex. Width of eyebridge half of the
length. Ventral surface of gnathosoma posterior to the mouth disk 2/3 covered by large
pores, in the posterior third with fine porosity or smooth. Pharyngeal plate ellipsoid. Coxae
reticulate as given in 7-32 c. Palp long and slender, with numerous setae; P-3 not bulging
distoventrally, medially with long setae not grouped together, P-4 with about 40 setae. P-5
as long as P-3. One of the largest species of the genus. Ma le : Idiosoma length 4000-4500
lm; Fe m al e : Idiosoma length 3100-6000 lm.
Pr eadu lt st ages : L arva: WAINSTEIN (1980). De u ton ym p h: K.V IETS (1930d, under
the name Eylais rimosa proceripalpis, see D AVIDS et al. 2005).
S im i la r spe c ie s: As no diagnostic differences can be found which would allow for a se-
paration of adult E. longipalpis from Eylais mutila, these taxa were considered conspecific
by most authors (see K.O.V IETS 1987). However, WAINSTEIN (1980) described larvae
reared from eggs of a species which he called E. longipalpis and at this stage he found
morphological differences from E. mutila. It remains unclear how he separated females
of Eylais mutila from E. longipalpis, but we must keep in mind the possibility that E. mutila
is a complex of species which are recognizeable at the larval stage only (D AVIDS et al. 2005).
B io lo gy : Oviposition in June, the majority of the eggs hatch in August, the rest in Sep-
tember. Larvae parasitic on water beetles of the families Dytiscidae (Dytiscus: P IATAKOV
1915b, Ilybius and Rhantus: L UNDBLAD 1927c, Acilius, Agabus, Colymbetes, Dytiscus,
Hydaticus, Hyphydrus, Ilybius, Platambus, Rhantus: Z AWAL 2004), Noteridae (Noterus),
Haliplidae (Haliplus) and Hydrophilidae (Cercyon, Anacena and Laccobius: Z AWAL
2004).
7. Hydrachnidia 329
H a bit at : In lakes, ponds, flowing waters, mostly in oligotrophic waters. Also found in
eutrophic, calcareous and brackish waters (L UNDBLAD 1968).
Di stri bu ti on: Palaearctic. Most frequently recorded from Northern and Central Europe.
10. Eylais planipons WALTER , 1924 (7-26 i, 7-29 a, 7-30 i)

1924d Eylais planipons WALTER , Bull. Soc. Sci., Cluj 2: 93.
1930d Eylais puripons K.V IETS, Arch. Hydrobiol. 21: 212.
1936e Eylais bisinuosa oligotricha L UNDBLAD, Ark. Zool. 29A: 11.
1942a Eylais planipons novata K.V IETS, Zool. Anz. 138: 211.
1962a Eylais canariensis L UNDBLAD, Ark. Zool. (2) 15: 288.
1977a Eylais tantilla D AVIDS (non K OENIKE , 1897), Biol. Gallo-Hellenica 6: 241.
De sc ri pti on : Frontal sclerite rather broad, eye capsule anteriorly wider than posteriorly,
with barely distinct medial margin; neighbouring setae included into the area of very fine
porosity of the capsules. Length of eyebridge about the width of one capsule or somewhat
longer, anterior margin straight or slightly indented. Mouth disk large, diameter larger than
the distance between the disk and the posterior margin of the gnathosoma. Palps small, P-3
distoventrally slightly bulging, distally with 6-10 short and thick setae, some of them pec-
tinate. P-4 medially with three to four medial setae and three to four pectinate distal setae.
One of the smallest species. Ma l e: Idiosoma length 1200-1500. Cx-III transversally mea-
sured 400-500. Frontal sclerite width 200-280, length of eye capsule about 120. Length of
P-4 less than 200 lm.
S im i la r s p e ci e s : Eylais gibbosa K.V IETS, 1930 from Spain is closely related to Eylais
planipons, but differs in having longer eye capsules, with the setal insertions not included
in the capsules. Furthermore, palps are longer and with higher numbers of setae, especially
P-4 (K.V IETS 1930d).
H a bit at & B io lo gy : Small ponds and ditches, life cycle unknown.
Di stri bu ti on: Mediterranean (Greek Islands, Spain, French Pyrenees); Iran (A SADI et al.
2002).
11. Eylais relicta H ALBERT, 1911 (7-26 k, 7-29 b, 7-31 a)

1911 Eylais relicta H ALBERT, Proc. Irish Acad., Dublin 31: 6.
1930a Eylais discreta stagnalis S OKOLOW (non H ALBERT, 1911), Zool. Jahrb. Syst. 59: 153.
1937 Eylais stagnaliformis S OKOLOW, Trav. Stst. limnol. Lac Baikal 7: 71.
1938 Eylais laminifera B ESSELING, Ent. Ber., Amst. 10: 76.
1950b Eylais infundibulifera relicta K.V IETS, Neue Ergebn. Probleme Zoologie, Leipzig: 1016.
De sc ri pti on : Frontal sclerite anteriorly with triangular extension (pointed or flattened),
eye bridge not extending over the level of the anterior margin of the eye capsules, setae
located close to the middle of the bridge, posterior margin with a deep bay. Eye capsules
broad posteriorly but contracted anteriorly. Pharyngeal plate slender. P-2 with nine to ten
strong setae on distal medial margin, proximomedial surface of the segment completely
devoid of setae. P-3 not bulging distally, with about 16-20 long setae. P-4 medially rich
in setae, P-5 apically with a group of slender setae. Ma le : Idiosoma length 2500-3200
lm. Palp (7-26 k) rather compact. F em a le : Idiosoma length 5000 lm. Palp more slender.
Pr eadu lt st ages: L arva: B IESIADKA & C ICHOCKA (1997). De u ton ym p h: not de-
scribed.
Habitat & Biology: Lakes and pools. Larvae parasitic on water beetles of the family Gyr-
inidae (Gyrinus distinctus A UBÉ: B IESIADKA & C ICHOCKA 1997).
Di stri bu ti on: Western Palaearctic, rare, known from scattered sites in Sweden, Den-
mark, Germany, England, Ireland, The Netherlands, Poland and Russia.
12. Eylais rimosa P IERSIG , 1899 (7-27 a, 7-29 c, 7-31 b)

1899a Eylais rimosa P IERSIG , Zool. Anz. 22: 65.
1899g Eylais tenera T HON , Zool. Anz. 22: 444.
1899g Eylais similis T HON , Zool. Anz. 22: 446.
De sc ri pti on : Frontal sclerite anterior margin generally with a slit or V-shaped depressi-
on, and setal insertions strongly bulging, but highly variable in shape (S OKOLOW 1930a).
Ventral surface of gnathosoma with large pores restricted to the anterior half, posteriorly
gradually diminishing in size. Pharyngeal plate maximum width 250 lm or more, with a
strongly developed ring-like area. P-3 with 12-14 setae on the projecting distal portion,
three to four of them pectinate. P-4 medially with 10-12 setae, five to seven of them pec-
tinate, four inserted at distal margin. Mal e : Idiosoma length 1500. Fe m al e : Idiosoma
length 2600-3800 lm.
Pr eadult stages: Larva: WAINSTEIN (1980). D eu ton ym p h: not described.
Habitat & Biology: Small standing water bodies, life cycle unknown.
Di stri bu ti on: Palaearctic. Recorded from Finland, Germany, England, Ireland, Italy,
Poland, Russia, Slovakia, Switzerland and Japan.
13. Eylais setosa K OENIKE , 1897 (7-27 b, 7-29 d, 7-31 c, 7-32 e)

1897a Eylais setosa K OENIKE , Abh. Naturw. Ver. Bremen 14: 281.
1899g Eylais latipons T HON , Zool. Anz. 22:440.
1907b Eylais okaënsis U DALZOV, Arb. hydrobiol. Stat. See Glubokoje 2: 233.
1910b Eylais amplipons K.V IETS , Abh. Naturw. Ver. Bremen 20: 166.
1910b Eylais trapezipons K.V IETS , Abh. Naturw. Ver. Bremen 20: 169.
1919d Eylais kuehnei K.V IETS , Arch. Naturg. 83: 161.
1929a Eylais extendens setosa L UNDBLAD , Sjön Tåkerns och Flora 5: 14.
1931a Eylais ussuriensis S OKOLOW, Zool. Jahrb. Syst. 61: 466.
1940a Eylais setosa goldfeldi S OKOLOW, Faune de l’URSS 5: 44.
De sc ri pti on : Frontal sclerite rather variable, but eye bridge essentially trapezoid (axis of
the eye capsules faintly converging posteriorly). Eye bridge with anterior margin generally
straight, with muscle attachments and setae insertions bulging or not. Surface of the
gnathosoma posterior to the mouth disk with a clear separation between an anterior
part bearing large pores and a posterior part with smaller pores. Pharyngeal plate with
maximum width distally, but only slightly diverging lateral margins in the distal two thirds.
P-3 only slightly bulging mediodistally, with about 10 nail-like robust setae. P-4 with about
20 setae, 10 of them (more frequently in the distal part) pectinate. Mal e : Idiosoma length
1400-3700. Fe m a le : length 1500-4000. Genital field with two small genital plates with
10-15 short setae.
Pr eadu lt s tages: L arva: WAINSTEIN (1963b, 1980); N IELSEN & D AVIDS (1975). De u -
to ny m ph: not described.
B io lo gy : Oviposition in May-June. Larvae parasitic on water beetles of the families
Haliplidae (Haliplus), Dytiscidae (Hygrotus, Hyphydrus) (N IELSEN & D AVIDS 1975)
7. Hydrachnidia 331
and Hydrophilidae (Coelostoma) (Z AWAL 2004). They attach to the abdominal tergites and
hibernate on their hosts. In food selection experiments, deutonymphs and adults showed a
clear preference for Daphnia longispina M ÜLLER against other prey species of the genera
Bosmina, Ceriodaphnia, Sida and Simocephalus (PAVELJEVA & Z ANKAI 1971).
H a bit at : Littoral of lakes and pools rich in macrophytes, in eutrophic, oligotrophic, dys-
trophic and slightly brackish waters (L UNDBLAD 1968).
Di stri bu ti on: Palaearctic. Common and widely distributed in Europe. Also reported
from Siberia and Japan (L UNDBLAD 1968).
14. Eylais tantilla K OENIKE , 1897 (7-27 d, 7-29 f, 7-31 d, 7-32 d)

1897a Eylais tantilla K OENIKE , Abh. naturw. Ver. Bremen 14: 287.
1903c Eylais nullipons T HOR, Nyt Mag. Naturv. 41: 73.
1906 Eylais bisinuosa T HON (non P IERSIG , 1899), Arch. naturw. Landesdurchforsch. Böhmen 12: 60.
1934a Eylais rimosa S ZALAY (non P IERSIG , 1899), Ann. Mus. nation. Hungar. 28: 275.
1919d Eylais bisinuosa nodipons K.V IETS, Arch. Naturg. 83: 160.
De sc ri pti on : Eye bridge very variable. The type specimen has a U-shaped anterior inden-
tation, in other specimens it may be narrower and slit-shaped. Mouth disk less in diameter
than the distance between the mouth disk and the posterior margin of the gnathosoma.
Proximal apodemes of gnathosoma rather long, pharyngeal plate ellipsoid, maximum
width about twice the width at the level of the posterior ring. Ventral surface of the gnatho-
soma posteriorly from mouth disk with large pores in one to two rows restricted to a nar-
row area bordering the mouth disc. P-3 bulging with short and thick setae, a few of them
pectinate. P-4 medially with four smooth setae, distally with four to five pectinate setae.
Ma le : Idiosoma length 1400-1800. Fe m al e : Idiosoma length 2200 lm. Genital field with
two plates bearing 10-15 short setae.
Pr eadult stages: Larva: WAINSTEIN (1963b, 1980). De uto ny m ph: not described.
B io lo gy : A typical springtime species (L UNDBLAD 1968). In The Netherlands not found
after the beginning of June (S MIT & VAN DER H AMMEN 2000). Oviposition early in sum-
mer, larvae hatch in July-August (WAINSTEIN 1963b). Parasitism not known.
H a bit at : Temporary and permanent standing waters rich in vegetation, also eutrophic or
polluted.
Di stri bu ti on: Palaearctic, rare, but found all over Europe. Also reported from eastern
Siberia, China and Algeria.
15. Eylais telmatobia S ZALAY, 1933 (7-29 g, 7-31e)

1933d Eylais megalostoma telmatobia S ZALAY, Zool. Anz. 104: 324.
De sc ri pti on : Frontal sclerite variable in shape, anterior margin weakly undulated, pos-
terior margin on the level of the middle of the eye capsules. Setal pores at the anterior mar-
gin of the eye bridge, close to the eye capsules. Eye bridge shorter and wider than in E.
hamata and E. curvipons, but longer than in the other European species. Anterior part
of ventral surface of gnathosoma behind mouth disk with large pores, the posterior
part with small pores. Ma l e: Idiosoma length 2300. Length of P-4 ca. 420. F e ma l e: Idio-
soma length 3200. Mouth disk diameter 300-360. Pharyngeal plate length 4 500, maxi-
mum width 280-310. P-3 without prominence, length P-4 ca. 470 lm.
H a bit at a nd B io lo gy : Small temporary waters and littoral of shallow lakes. Life history
unknown.
Di stri bu ti on: Hungary, Poland, Slovakia (L ÁSKA 1971).
16. Eylais triarcuata P IERSIG , 1899 (7-27 e, 7-29 h, 7-31 f)

1899a Eylais triarcuata P IERSIG , Zool. Anz. 22: 66.
1910a Eylais muelleri triarcuata WALTER , Arch. Hydrobiol. 5: 174.
1942 Eylais rimosa triarcuata ŞOAREC , Ann. Scient. Univ. Jassy 29: 63.
De sc ri pti on : Frontal sclerite (7-29 h) similar to E. extendens (7-28 e, 7-29 e), width
about 300 lm. Eye bridge anteriorly with three arches. Pharyngeal plate, gnathosoma
and palp larger than in E. extendens. Gnathosoma ventral surface behind the mouth
disk with an area of large pores, distinctly set off from the posterior part with a rough
skin. Sculpturing of coxae as in E. extendens (7-32 a). P-3 with ventrodistal edge bulging
less than in E. extendens with about 15 faintly pectinate setae. P-4 medially with more than
15 pectinate setae distributed over the whole medial side, distally 5 pectinate setae. M al e :
2500-4000. Fe m a le : length 3000-5000 lm.
Praeadult stages, Habitat and Biology: No data published.
Di stri bu ti on: Western Palaearctic. Reported from Russia, Bulgaria, Poland, Germany,
Italy and Scandinavia.
R em a r k: Eylais triarcuata was considered a synonym of E. extendens by K.V IETS (1956b)
and L UNDBLAD (1962). However, the setation on the palp allows a clear distinction. Prob-
ably, the two species have often been confused in faunistic surveys and their distribution
and habitat preference need reinvestigation.
17. Eylais tullgreni T HOR , 1899 (7-27 f, 7-29 i, 7-31 g)

1899b Eylais tullgreni T HOR , Foreløbig meddelelse, Kristiania: 7.
De sc ri pti on : Frontal sclerite with very small eye bridge, anteriorly and posteriorly with a
deep small slit, which can be wider than indicated in 7-29 i. Muscle attachment at the pos-
terior side of the bridge. Ventral area of the gnathosoma behind the mouth disk short, pha-
ryngeal plate narrow, extending far beyond the posterior margin of the gnathosoma. P-3
ventrodistal prominence with only three to five short broad setae. P-4 length about 250 lm,
medially with 8-11 setae (three smooth and five to eight pectinate), laterally with four
smooth setae. F e m al e: length 2000-2400 lm. Male not described.
Pr eadult stages: Larva: See WAINSTEIN (1963b, 1980). De u ton ym p h: not described.
B io lo gy : Z AWAL (2004) found small numbers of larvae on water beetles of the genera
Helophorus and Laccobius. Larvae can be found in August-September (WAINSTEIN
1963b).
H a bit at : Z AWAL (2004) found the larvae in small eutrophic water bodies and in the lit-
toral of lakes.
Di stri bu ti on: Poland, Russia, Sweden.
R em a r k: K.V IETS (1956b) considered this species as a species incerta, L UNDBLAD (1962)
proposed its synonymy with E. tantilla, while WAINSTEIN (1963a, 1980) found morpho-
logical differences at the larval stage which indicate that this is a species on its own. How-
ever, the diagnostic features presented here and their variability should be re-examined with
more material from a wider geographic area.
7. Hydrachnidia 333
Superfamily Hydrachnoidea
1815a Hydrachnides L EACH, Trans. Linn. Soc., London 11: 399.
Di ag no sis: Idiosoma spherical (7-6). Integument papillate, usually red; sclerotization of
dorsum variable, from almost none to a nearly complete armour leaving only minute mem-
branous slits in the posterodorsal area. Sclerotization generally with indistinct margins (pa-
pillate membranous areas grading to sclerotized fields, with the pattern of papillosity re-
flected in sclerotized parts as a pattern of more dense porosity), porosity of sclerites very
fine and dense. Lateral eyes in capsules, never included into, but often embraced by, dorsal
sclerites (7-36 a-b, d); median eye absent or present. Coxal plates typically in four groups
(7-33 a), occasionally joined by secondary sclerotization, number of setae on coxae very
low, no setae associated with leg insertions or plate margins; Cx-III+IV with concave mar-
gins embracing the genital field; Cx-IV larger than Cx I-III, with the leg insertions distinctly
further from the median line than Cx-I-III. Genital field (e.g. 7-38) consisting of an un-
paired, often anteriorly indented sclerite bearing numerous minute acetabula and a few
scattered setae in the anterior, and the posteriorly directed gonopore flanked by groups
of setae in the posterior part; excretory pore surrounded by a sclerotized ring. Gnathosoma
(7-33 c-d) typically with a long, pointed rostrum. Chelicerae with the two segments fused
forming a stylet (7-33 b). Palp (e.g. 7-34) chelate, with P-1 usually being the largest seg-
ment, P-3 elongate and taking over the function of P-4 of other water mites, P-4 and P-5
reduced in size, acting together as a chela. Legs laterally compressed, all segments bearing
numerous, rather uniform, finely pinnate setae distributed all over the segment surface, but
no particularly differentiated setae at distal segment margins; numerous long swimming
setae on the posterior surface of II-IV-L-4/5, additional long ventral setae arranged on
III-L-4/5 and IV-L-3-5; distal segments of all legs shortened, with simple, minute claws.
Ma le s: Genital field with a high number of densely arranged setae in the posterior part;
gonopore completely surrounded by sclerotized lips, longish or enlarged.
Fe m a le s: Genital field with lower numbers of setae, in the posterior part in two groups:
one group of setae surrounding a pair of enlarged pores (modified acetabula?) lying be-
tween acetabular field and gonopore, a further group of longer setae at posterior plate
margin; gonopore sclerotized only anteriorly, posteriorly membranous, with a tubular ovi-
positor, which is usually inverted inside the body; the whole genital field movable with a
„flipflap mechanism“: in normal position posterior part deeply embedded in a membra-
nous fold, of the posterior group of setae at most the very tips externally visible, extrusible
by a hinged joint at the anterior margin, in extended position the gonopore directed ven-
trally, the acetabula directed anteriorly, the setal bases visible.
Family Hydrachnidae L EACH , 1815

1815a Hydrachnides L EACH , Trans. Linn. Soc. London 11: 399.
Di ag no sis: Only family of the superfamily, with the characters of the superfamily.
Genus Hydrachna M ÜLLER , 1776

1776 Hydrachna M ÜLLER, Hallager: 188.
1916 Anohydrachna, Diplohydrachna T HOR, Rev. Russe Entomol. 16: 46.
1916 Limnohydrachna, Monohydrachna, Odontohydrachna, Schizohydrachna T HOR, Rev. Russe
Entomol. 16: 47.
1931b Rhabdohydrachna K.V IETS, Zool. Anz. 93: 184.
1933b Scutohydrachna K.V IETS, Zool. Anz. 103: 162.
334 Superfamily Hydrachnoidea
1934b Tetrahydrachna L UNDBLAD, Ark. Zool. 28 A 3: 30.

1966 Chitohydrachna H ABEEB, Leafl. Acadian Biol. 41: 1.
1966 Hymenohydrachna WAINSTEIN, Inst. Biol. Vnutrenn. Vod, Trudy 12 (15): 183.
1976c Piersigihydrachna, Thorihydrachna WAINSTEIN, Inst. Biol. Vnutrenn. Vod, Trudy 31 (34): 134.
Fig. 7-33: Hydrachna; a-b, H. processifera male; a, coxal and genital area; b, chelicera; c, H. skor-
ikowi female gnathosoma lateral view; d, H. processifera male gnathosoma, lateral view; e, H. incisa
female gnathosoma lateral view.
7. Hydrachnidia 335
1976c Microhydrachna WAINSTEIN, Inst. Biol. Vnutrenn. Vod, Trudy 31 (34): 140.
Typus generis: Hydrachna cruenta M ÜLLER , 1776 by subsequent designation of L ATREILLE
(1796).
Di ag no sis: Only genus of the family, with the characters of the superfamily.
R em a rks : As pointed out by L UNDBLAD (1969) and C OOK (1974), the system of subge-
neric classification used by former authors (e.g. K.V IETS 1931b) is unsatisfacory. With the
exception of Bargena K OENIKE , 1893 (lacking a median eye and not occuring in the area
covered), the definition of subgenera is based on the presence and shape of the dorsal scler-
ites. The fact that males and females of the same species occasionally fit in different sub-
genera emphasizes the artificial character of this system. Furthermore, there is much va-
riation in the shape of dorsal sclerites and plates within species. For these reasons, the for-
mer subgeneric classification is abandoned here. In agreement with K.O.V IETS (1987) we
also treat the genera and subgenera erected by WAINSTEIN (1966, 1976c) as synonyms of
Hydrachna. In an early phase of systematics, a high number of species were described based
on doubtful characters often subject to individual variability (integument structure, shape
and arrangement of frontal plates, outline of coxal plates). Many taxonomic questions re-
mained open up to the present day and several species are frequently named which are not
well defined in their diagnostic characters. Here, we follow in many details L UNDBLAD
(1962, 1968) who lumped numerous older taxa. However, future investigations on life cy-
cles and larval morphology, following the example of WAINSTEIN (1976c), could demon-
strate that some of the taxa treated here as species, in reality represent species complexes. In
such cases, revival of species names here treated as synonyms, will make sense only if it is
accompanied by laboratory rearings, redefinition of species at larval and both sexes of the
adult stage and neotype designation.
B io lo gy : Especially in species inhabiting temporary ponds and swamps, females produce
high numbers of eggs which are deposited in the aerenchyma or, more rarely, the paren-
chyma of submerged macrophytes with the help of a pointed ovipositor (7-11). As oviposi-
tion is prepared for by the drilling of a hole into the tissues with the help of the gnathosomal
rostrum, incorrect observations lead earlier authors to the wrong conclusion that Hydrach-
na species were herbivores. Larvae are good swimmers and find their insect hosts under
water. The known host range includes water bugs (Nepomorpha) and water beetles (Dy-
tiscidae, Hydrophilidae). With their characteristically enlarged gnathosoma, Hydrachna
larvae attach generally to external surfaces, only rarely in the subelytral space. As their
strongly developed mouth parts allow for perforation of sclerotized parts of the host’s in-
tegument, in contrast to most other water mite larvae, they are not restricted to interseg-
mental membranes. They are found often in large numbers on single host individuals, but
only one, or a few specimens grow up and are capable of developing further (7-5 a-c). Also,
the postlarval moult takes place on the host and the protonymph remains attached, as a
large, drop-shaped appendix. As these first developmental stages may last up to ten
months, a large part of the individual life of some species proceeds in this parasitic-phoretic
phase. At this stage, larvae are capable of surviving drought periods in terrestrial habitats,
profitting from the behavioral and physiological adaptations of their hosts. The deutonym-
phal moult takes place when a mature protonymph comes in contact with water. Again,
Hydrachna species profit from their hosts, in this case from their hosts habitat selection
capacity. In contrast to most other water mites, Hydrachna deutonymphs hatch with a con-
siderably large body size and are capable of quickly reaching the tritonymphal and adult
stages. As both deutonymphs and adults often feed on the eggs of their potential hosts, there
are particularly strong ties between the mite and its host during the whole life cycle of the
mite. Most species (including all species so far recorded from Europe) have only a few re-
duced idiosoma sclerites allowing for expansion of the membranous parts. Depending on
the availability of food resources, dimensions in the same species may vary extremely and
single specimens can reach an idiosoma size of up to 9.000 lm (D AVIDS 1973a, S TEVENS &
G REVEN 1994, 1995, 1999).
Pr eadu lt stages: L arvae: WAINSTEIN (1980). Deutonymphs: usually with two separate
provisional genital plates bearing numerous acetabula (T UZOVSKIJ 1990).
Key to species
The following species listed for the study area by K.O.V IETS (1978) are excluded:
Hydrachna aspratilis K OENIKE , 1897: Juist, Germany, described from a deutonymph; reported from
several East European countries (species incerta, D AVIDS et al. 2005); H. bivirgulata P IERSIG , 1897d:
Sachsen, Germany (species incerta, D AVIDS et al. 2005); H. denudata P IERSIG , 1896: Sachsen, Ger-
many (species incerta, D AVIDS et al. 2005); H. extorris K OENIKE : type locality of the only record un-
clear (possibly outside Europe - for this reason not treated by K.V IETS 1936); for diagnostic characters
see D AVIDS et al. 2005 and discussion of H. incisa; H. inermis P IERSIG , 1895: Sachsen, Germany (spe-
cies incerta, D AVIDS et al. 2005); H. marita WAINSTEIN , 1966: Russia, Poland (incompletely described,
see „similar species“ of H. geographica); H. perpera K OENIKE , 1908: Sachsen, Germany (synonym to
H. processifera, D AVIDS et al. 2005); Hydrachna papilligera K.V IETS , 1919: Braunschweig, Germany;
described from a deutonymph, possibly synonym to H. skorikowi (species incerta, D AVIDS et al. 2005);
H. piersigi K OENIKE , 1897: Sachsen, Germany (species incerta, D AVIDS et al. 2005); H. regulifera
K OENIKE , 1908: Bremen, Germany (described from a deutonymph erroneously assumed a female,
possibly synonym to H. geographica (species incerta, D AVIDS et al. 2005); H. williamsoni S OAR ,
1908: British Isles (species incerta, G LEDHILL & K.O.V IETS 1976, D AVIDS et al. 2005).
1 Dorsum with one large frontal shield (7-36 d); if this shield posteriorly indented, the indentation
never deeper than the median length of the shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– If (in rare cases) a single frontal shield is present, it is apparently formed by the medial fusion of the
anterior edges of paired plates, leaving a posteromedial indentation much deeper than the medial
length of the shield (7-36 a); otherwise frontal idiosoma with a pair of plates (anteriorly enlarged,
rod-shaped or oval) or without sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Palp stocky (P-2 length/height 5 _ 1.0, P-3 length/height 5 4.0), P-3 basally distinctly thicker than
in the middle (7-34 f); gnathosomal rostrum in dorsoventral view basally not narrowed; posterior
margin of frontal shield may be indented. . . . . . . . . . . . . . Hydrachna skorikowi (page 355)
– Palp slender (P-2 length/height 1.0, P-3 length/height 4 4.0), P-3 ventral margin in its basal part
concave, maximum height in the middle (7-35 a, c); gnathosomal rostrum in dorsoventral view
basally narrowed; frontal shield usually with straight posterior margin. . . . . . . . . . . . . . . . 3
3 Medial margins of Cx-III pointed, distanced from each other (distance excretory plate diame-
ter); posteromedial angle of Cx-IV not prominent, rounded (7-38 a); P-3 distinctly longer than P-2
(7-35 a); median eye on the level of, or behind the posterior margin of lateral eye capsules; male
genital field similar to that of H. skorikowi (7-39 e), longer than wide, collar surrounding the male
gonopore poorly developed or absent.. . . . . . . . . . . . . . . . . . Hydrachna cruenta (page 346)
– Medial margins of Cx-III with an extended subparallel sector, approaching each other (distance =ˆ
excretory plate diameter); posteromedial angle of Cx-IV strongly projecting and pointed (7-39 b);
P-3 only a little longer than, equal to, or shorter than, P-2 (7-35 c); median eye at the level of centre
of lateral eyes; male genital field wider than long, gonopore surrounded by a protruding collar,
particularly projecting at the anterior margin (7-39 b) (female undescribed).. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna inversa (page 353)
4 Frontal area with a pair of longish sclerites extending between (and often including) pre- and post-
ocularia; these sclerites occasionally fusing anteromedially (7-36 a-b, f).. . . . . . . . . . . . . . . 5
– Frontal area with several minute sclerites (maximum diameter not exceeding 50 % of the distance
between pre- and postocularia capsules, occasionally completely lacking sclerites - 7-36 c, e,
7-37 d-f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
5 Frontal plates anteriorly enlarged (P OPP 1959: „South America shaped“), occasionally anterome-
dially fused (7-36 a, b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Frontal plates narrow, rod-shaped (7-36 f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6 Palp stout (length/height P-2 1.0-1.2, P-3 1.8-2.5, P-4 1.5-1.9 - 7-35 d); posteromedial edges of
Cx-IV broadly rounded, not extended (7-38 c); frontal plates never fused medially. . . . . . . . 7
7. Hydrachnidia 337
– Palp more slender (length/height P-2 1.7-2.0, P-3 3.0, P-4 2.0 - 7-34 e, g-h); posteromedial edges
of Cx-IV forming more or less narrowed extensions (7-37 a, 7-39 d, g); frontal plates may fuse
medially. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Female genital field transverse (7-38 c), distinctly wider than long; medial margin of frontal plates
with an angular indentation . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna conjecta (page 344)
– Female genital an inverted heart-shape, not wider than long (7-38 b); medial margin of frontal
plates weakly concave; (little defined species, male unknown). Hydrachna distincta (page 347)
8 Palp slender (length/height P-3 4 3.8 strongly narrowed basally) (7-34 g) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna juncta (page 353)
– Palp less slender (length/height 5 3.8, not strongly narrowed basally) (7-34 e, h, 7-35 b) . . 9
9 Posteromedial extension of Cx-IV narrow („finger-shaped“); male genital field longish, gonopore
pointed anteriorly (7-39 g). . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna globosa (page 349)
– Posteromedial extension of Cx-IV larger („tongue-shaped“, 7-39 d); male genital field more en-
larged, gonopore rounded anteriorly (7-39 c). . . . . . . . . . . . Hydrachna goldfeldi (page 350)
10 Gnathosoma and palp very stocky (length/height P-2 5 1.5, P-3 5 2.5, P-4 length/height 5 2.0,
7-34 b), rostrum shorter than base. . . . . . . . . . . . . . . . . Hydrachna crassipalpis (page 345)
– Gnathosoma and palp slender (length/height P-2 4 1.8, P-3 4 3.5, P-4 length/height 4 2.2, 7-35
f), rostrum longer than base. . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna comosa (page 344)
11 Gnathosoma slender, rostrum distinctly longer than base (rostrum/base length 4 1.4, 7-33 e);
Cx-IV with numerous (two to eight) setae; palp slender (P-2 length/height 4 1.5, P-3 length/height
4 3.5), P-1 without setae; P-3 with straight, anteriorly slightly converging dorsal and ventral mar-
gins (7-34 c, d) (for an exceptional species with a long gnathosomal rostrum but differing in other
characters, not with certainty recorded from the study area, see discussion of H. incisa) . . 12
– Gnathosoma more compact, rostrum slightly longer or shorter than base (rostrum/base length 5
1.4, 7-33 c or similar to 7-33 d); Cx-IV bearing one seta only; palp various in shape, P-1 bearing
one or two dorsodistal setae, P-3 ventral margin basally concave, distally convex (e.g., 7-34 i) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 Frontalia associated with postocularia crescent-shaped (7-36 c), maximum length 350; Cx-IV
bearing two to five setae; smaller in dimensions (idiosoma length 2000-4000, coxal field length
1400-2000); male gonopore transverse, with straight or slightly indented anterior margin (7-39 f)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna incisa (page 351)
– Frontalia associated with postocularia rod-shaped (7-37 d), length 400-700; Cx-IV bearing six
to nine setae; a very large species (idiosoma length males 3500-7000, females 5000-9000, coxal
field length 2000-2600); male gonopore elongated, anteriorly pointed or parabola-shaped (7-38 d)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna geographica (page 348)
13 Ventral margin of P-2 straight or weakly concave (7-34 i); paired sclerites in the frontal area round,
not fused to each other or to preocularia (7-37 e); medial margins of Cx-III and Cx-IV meeting in
an obtuse angle or together forming a concave genital bay . . . . . . . . . . . . . . . . . . . . . . . 14
– Ventral margin of P-2 distinctly concave (7-34 a); some of the paired sclerites in the frontal area
and preocularia may be variously fused forming rod shaped platelets (7-36 e, 7-37 f); medial mar-
gins of Cx-III and Cx-IV subrectangular (7-38 f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14 Palp longer (male: length P-2 4 200, P-3 4 300, P-4 4 120) and more slender (length/height P-3
4 2.5, 7-34 i); male genital field length/width 640/600, acetabula small, more numerous, medially
extending over the anterior half of the plate (7-33 a).. . . . Hydrachna processifera (page 355)
– Palp shorter (male: length P-2 5 150, P-3 5 200, P-4 5 110) and stouter (length/height P-3 5 2.2,
7-35 e), male genital field length/width 365/360, acetabula larger, less numerous, medially re-
stricted to the anterior third of the genital plate; female unknown. . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna danubialis (page 346)
15 Posterior to preocularia a pair of elongated, rod-shaped platelets; these sclerites often fused to
preocularia (7-37 f); P-3 more slender (length/height 4 3.3, 7-35 g); male gonopore (7-38 f) sur-
rounded by rather robust, less densely arranged setae . . . . . . . . Hydrachna leegei (page 354)
– Without elongated rod-shaped platelets posterior to preocularia; preocularia with a small poste-
rior extension or round (7-36 e); P-3 stout (length/height 5 3.0, 7-34 a); male gonopore as given in
7-38 f for H. leegei, but surrounded by finer, more densely arranged setae . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrachna incognita (page 352)
Fig. 7-34: Hydrachna, palps; a, H. incognita female; b, H. crassipalpis female; c, H. incisa female; d,
H. incisa male (lateral setae on P-3 lacking); e, H. goldfeldi female; f, H. skorikowi male; g, H. juncta
female; h, H. globosa male; i, H. processifera male.
7. Hydrachnidia 339
Fig. 7-35: Hydrachna, palps; a, H. cruenta female; b, H. goldfeldi lectotype female; c, H. inversa male,
combined after WALTER & M OTAŞ (1927) and L UNDBLAD (1956a); d, H. conjecta female; e, H. da-
nubialis male (after L ÁSKA 1964); f, H. comosa male; g, H. leegei female.
Fig. 7-36: Hydrachna, frontal area; a, H. juncta female with fused frontal sclerites; b, H. juncta female
with separated frontal sclerites; c, H. incisa female; d, H. skorikowi female; e, H. incognita female; f,
H. crassipalpis female.
7. Hydrachnidia 341
Fig. 7-37: Hydrachna, a-c, female genital fields; a, H. juncta, with medial margin of Cx-III+IV; b, H.
incisa; c, H. incognita, with medial margin of Cx-III+IV; d-f, frontal areas, d, H. geographica male;
e, H. processifera male; f, H. leegei female.
Fig. 7-38: Hydrachna, genital fields; a, H. cruenta female, with medial margin of Cx-III+IV; b, H.
distincta female, holotype; c, H. conjecta female, with medial margin of Cx-III+IV; d, H. geographica
male; e, H. crassipalpis male, after L ÁSKA (1958); f, H. leegei male.
7. Hydrachnidia 343
Fig. 7-39: Hydrachna, male genital field and posterior coxae; a, H. conjecta; b, H. inversa, combined
after WALTER & M OTAŞ (1927) and L UNDBLAD (1956a); c, H. goldfeldi, genital field; d, H. goldfeldi,
medial margin of Cx-III+IV; e, H. skorikowi; f, H. incisa; g, H. globosa.
1. Hydrachna comosa K OENIKE , 1896 (7-35 f)

1896b Hydrachna comosa K OENIKE , Zool. Anz. 19: 359.
De sc ri pti on : Paired frontal plates extending from pre- to postocularia very slender,
slightly S-shaped, surface finely granular. Coxae 4 with one seta, posteromedial corner
broadly rounded, not extended. Gnathosoma rostrum slightly longer than base. Palp slen-
der, approximate setae numbers; P-1 one dorsodistal, P-2 numerous (about 10) dorsal and
two lateral, P-3 three dorsal and five lateral. M al e: Idiosoma length 2700 lm. Genital field
as wide as long, heart-shaped, anteriorly and posteriorly indented. Fe m al e : Idiosoma
length up to 4200, coxal field 1300. Genital field length/width 560/730 lm, posteriorly
concave, with a deep anterior indentation extending over more than three quarters of
the plate. Gnathosoma length 1680 (rostrum 930), chelicera 2400; palp total length
1660, length/height P-1 390/500 (0.8). P-2 400/210 (1.9), P-3 600/130 (4.6), P-4 190/
80 (2.4), P-5 80/43 (1.9) lm.
Pr ea du lt sta g es: L a rv a unknown; Deu to ny m ph: T UZOVSKIJ (1990).
S im i la r spe c ie s: Specimens with misshaped frontalia (sclerotization occasionally inter-
rupted ) might be confused with H. leegei, a species characterized by distinctly minor gen-
eral dimensions and a stouter P-3 (length/height 5 4.0).
B io lo gy : Little known; adults most frequently recorded during springtime.
H a bit at : Temporary and permanent standing waters.
Di stri bu ti on: Western Palaearctic. Reported from Northern, Western and Central Eur-
ope; apparently rare everywhere, e.g. in Sweden only known from a few records around
Uppsala (L UNDBLAD 1962).
2. Hydrachna conjecta K OENIKE , 1895 (7-5 e-f, 7-35 d, 7-38 c, 7-39 a)

1895a Hydrachna conjecta K OENIKE, Rev. biol. Nord France 7: 145.
De sc ri pti on : Of the European fauna, together with H. leegei, the species most reduced in
size. Frontal plates slender, never fusing anteromedially. Coxae 4 bearing one seta. Palp
rather robust; approximate setae numbers: P-1 without dorsodistal seta; P-2 five to seven
dorsal, one lateral; P-3 two to three dorsal, one lateral. M al e : Idiosoma length 2300-3300;
frontalia 580-700; coxal field 920; genital field length/width 400-430/340-390 lm, ante-
riorly to the anterior edge of the gonopore with a weakly produced, medially concave ex-
tension. Gnathosoma length 550-590, rostrum 220-230, chelicera 750-800; palp total
length 585-630, length/height P-1 125-130/195220 (0.6), P-2 160-170/150-155 (1.1),
P-3 175-190/80 (2.2-2.4), P-4 80-90/45-50 (1.8), P-5 45-50/25-27 (0.9) lm. F em a l e: Idio-
soma length 2100-2800; frontalia 800; coxal field 1200; genital field length/width 270/480
lm, strongly transverse, with weakly convex posteromedial margin. Gnathosoma length
770, rostrum 370; palp total length 770, length/height P-1 170/290 (0.6), P-2 200/200
(1.0), P-3 250/105 (2.4), P-4 100/55 (1.8), P-5 50/30 (1.7) lm.
S im i la r s p e ci e s : Hydrachna georgei S OAR , 1908, described from the British Isles and
later recorded from France (C. A NGELIER 1950a), is possibly a synonym of H. conjecta
(D AVIDS 1973a). In view of poor documentation it is considered a species dubia by G LED-
HILL & K.O.V IETS (1976).
Pr eadu lt st ages: L arva: D AVIDS (1973a), WAINSTEIN (1980). D eu ton ym p h: unlike
many other Hydrachna-deutonymphs, with fused provisional genital plates (D AVIDS
1973a, T UZOVSKIJ 1990).
B io lo gy : D AVIDS (1973a) studied this species in detail, and unless stated otherwise, most
of the following data refer to this paper. Larvae parasitize water bugs of the family
Corixidae (Hemiptera). The following species are known as host: Cymatia coleoptrata
7. Hydrachnidia 345
(FABRICIUS ), Hesperocorixa linnei (F IEBER ), Sigara striata (L INNAEUS ), Corixa punctata

(I LLIGER ), Cymatia bonsdorffi (S AHLBERG .), Callicorixa praeusta (F IEBER ), Sigara semi-
striata (F IEBER ) and S. stagnalis (L EACH ). Sigara falleni (F IEBER ) seems to be resistant.
The food of H. conjecta consists of eggs of Corixidae, e.g. Sigara striata. Longer stalked
eggs, e.g those of Cymatia coleoptrata, are rarely taken. Oviposition takes place from May
to June; females lay a maximum of 1500 eggs. The parasitic stage takes about 17 days, the
free living nymphal stage five to nine days and adults live for about one month. There are
two or three generations per year. The species hibernates as egg or at a nymphal stage.
H a bit at : All kind of standing waters, also in canals and dune lakes or slow flowing
streams; tolerant to eutrophication and higher salinity.
Di stri bu ti on: Palaearctic - widespread in Europe, occurring north up to central Sweden
(L UNDBLAD 1968); populations formerly recorded from North-America now considered as
a separate species, Hydrachna davidsi B.P. S MITH , 1987.
3. Hydrachna crassipalpis P IERSIG , 1897 (7-34 b, 7-36 f, 7-38 e)

1897d Hydrachna crassipalpis P IERSIG, Zool. Anz. 20: 352.
De sc ri pti on : Coxae 3 with a nose-like projecting anteromedial edge, medial margin
slightly convex; Cx-IV bearing one seta, medially slightly concave, with distinct, but
not very projecting posteromedial extension bearing a border of secondary sclerotization
forming a distinct, hook like posterolateral edge. Gnathosoma stout, with a short, ventrally
bent rostrum. Palp very stout, P-2 ventrally strongly concave or with a kink, P-3 inflated,
ventral margin with a concave base, but over most part of the segment strongly convex,
dorsal margin straight, distally projecting in a right angle, P-4 with maximum height dis-
tally, dorsodistal projection of P-4 and P-5 very strong; approximate setae numbers; P-1
two or three dorsodistal, P-2 8-10 dorsal and three or four lateral; P-3 three or four dorsal
(one of them at the distal edge) and four or five lateral. Mal e : Idiosoma length 2800; fron-
tale length/width 880-1240/120; coxal field length 1150-1200, genital field length/width
600-700/700 lm, deeply indented anteriorly and laterally slightly narrowed at the level of
anterior edge of gonopore. Gnathosoma length 700-800, rostrum 300-400; chelicera 1200-
1400; palp total length 800, length/height P-1 200/260 (0.8), P-2 190/160 (1.2), P-3 250/
110 (2.3), P-4 100/55 (1.9), P-5 58/30 (1.9) lm. F em a le : Idiosoma length 2500; frontale
length/width 800-1000/140; coxal field length 1150-1250; genital field length/width 440-
450/600 lm. Gnathosoma length 950-1000, rostrum 350-500; palp total length 880-890,
length/height P-1170-200/300-320 (0.5-0.7), P-2 215-230/190-210 (1.0-1.2), P-3 280-305/
130-135 (2.2-2.3), P-4 110-125/65-70 (1.6-1.9), P-5 60-75/35 (1.7-2.1) lm.
Pr eadult stages: Larva: WAINSTEIN (1976c, 1980). De ut ony m ph : T UZOVSKIJ
(1990).
S im i la r sp e c ie s : Hydrachna crassipalpis is not documented by type material, but the
description of the characteristic palp (P IERSIG 1897d, Fig. 6) is sufficient for recognizing
this species, and a neotype could be designated in future (locus typicus: Leipzig, Großzscho-
cher, forest pools). L UNDBLAD (1969) pointed out that H. crassipalpis is possibly a synonym
of H. bivirgulata, a species described in the same year from the same type locality. As there
is no information on important morphological details and the type material of H. bivir-
gulata is lost, D AVIDS et al. (2005) considered it a species dubia.
H a bit at a nd B io lo gy : Small standing water bodies. Life cycle unknown.
Di stri bu ti on: Central and Northern Europe, rare. Probably often recorded under the
name H. bivirgulata.
4. Hydrachna cruenta M ÜLLER , 1776 (7-35 a, 7-38 a)

1776 Hydrachna cruenta M ÜLLER, Hallager: 190.
De sc ri pti on : Anterior margin of frontal shield with a large, anteriorly rounded extension
projecting far beyond the lateral eye capsules, posterior margin convex, straight or with a
minute indentation, similar to H. skorikowi (7-36 d), but medial eye located more poster-
iorly, on the level of the posterior margin of the lateral eye capsules. Medial margins of
Cx-III separated from each other and pointed; Cx-IV with one seta, posteromedial edges
truncated and broad. Gnathosoma rostrum longer than base. Palp slender, P-2 distinctly
longer than high, P-3 ventral margin basally concave, distally convex, basal part of P-4
(excluding the dorsodistal extension) distinctly longer than P-5; approximate setae num-
bers: P-1 two dorsodistal, P-2 six to seven dorsal, two or three lateral, P-3 two to three
dorsal, two medial. Mal e : Idiosoma length 2000-3000; coxal field 1150; genital field
length/width 580/580 lm, maximum width anteriorly, only slightly narrowed posteriorly,
posterior margin on the level of posterior edges of Cx-IV, gonopore distinctly longer than
wide; in general shape similar to H. skorikowi (7-39 e), but anteromedially with a slight
indentation and bearing a higher number of setae (about 20 pairs) in the acetabular region.
Gnathosoma length 790, rostrum 410; palp total length 820, length/height P-1 170/260
(0.7), P-2 200/155 (1.3), P-3 290/70 (4.1), P-4 115/50 (2.3), P-5 45/25 (1.8) lm. Fe m al e :
Idiosoma length 2000-3000; coxal field 1070; genital field length/width 320/500 lm,
transverse oval, with large acetabula which in the anterior part extend over the whole plate
surface (not divided into two groups by a medial, acetabula-free strip). Gnathosoma length
1050, rostrum 650; palp total length 1090, length/height P-1 220/290 (0.8), P-2 250/160
(1.6), P-3 440/90 (4.9), P-4 130/60 (2.2), P-5 50/30 (1.7) lm.
S im i la r s pe c ie s: The undescribed female of H. inversa should differ from H. cruenta in
the presence of pointed extensions at the posteromedial margins of Cx-IV and the relatively
short P-3; possibly, the position of the frontal eye (more anterior in H. inversa) is subject to
individual variability.
P r ea d u lt s t a ge s : La r v a: WAINSTEIN (1980), D AVIDS (1973a). De ut ony m ph : with
paired frontal plates (S OKOLOV 1940a).
B io lo gy : Larvae parasitic on Corixidae (Hemiptera): Callicorixa praeusta (F IEBER ), Co-
rixa dentipes T HOMSON , Corixa punctata (I LLIGER ), Sigara distincta (F IEBER ), Sigara
falleni (F IEBER ), S. fossarum (L EACH ), S. lateralis (L EACH ), S. nigrolineata (F IEBER ), S.
semistriata (F IEBER ), S. striata (Li NNAEUS ), Cymatia coleoptrata (FABRICIUS ), C. bonsdorffi
(S AHLBAUER ), Hesperocorixa linnei (F IEBER ) and H. sahlbergi (F IEBER ) (D AVIDS 1973a,
B IESIADKA & C ICHOCKA 1994).
H a bit at : Standing waters, occasionally in slow flowing streams; eurytopic, occurring in
oligotrophic and eutrophic waters, tolerant to high salinity.
Di stri bu ti on: Holarctic; widespread in Europe, in Sweden occurring far north (L UND-
BLAD 1968).
5. Hydrachna danubialis L ÁSKA , 1964 (7-35 e)

1964 Hydrachna (Rhabdohydrachna) danubialis L ÁSKA, Věstn. Česk. Spol. Zool. 28: 217.
De sc ri pti on : Female unknown. Frontal area with a pair of minute, knob-shaped sclerites
halfway between pre- and postdorsal and a pair of larger, irregularly shaped platelets fused
to postdorsalia, no sclerites associated with predorsalia. Medial margin of Cx-IV concave,
posteromedial corner extended with hook-shaped laterally-directed secondary sclerotiza-
tion. Palp very stout, numbers of setae low (setation similar to H. processifera). Ma le :
Idiosoma length/width 1760/1580, eye capsules maximum diameter 140, distance 300;
coxal field length/width 785/1465; genital field length/width 365/360 lm, acetabula scat-
7. Hydrachnidia 347
tered over the anterior third of the plate, leaving an extended strip of free sclerite surface
between the posterior acetabula and the anterior margin of the posterior setation. Gnatho-
soma length 560, rostrum length 220, chelicera length 770; palp total length 543, length/
height P-1 122/164 (0.74), P-2 118/122 (0.97), P-3 157/79 (1.99), P-4 96/44 (2.18), P-5
50/- lm.
S im i la r sp ec i e s : L ÁSKA compared this species to H. silvestrii R IBAGA , 1902 from South
America and H. tenuissima K.V IETS , 1935 from Sumatra, species distincly different in the
shape of the genital field. Notwithstanding the artificial character of its collection sites, H.
danubialis is most probably a representative of the European fauna, rather similar to H.
leegei. The taxonomic significance of the diagnostic features should be investigated with
variability studies.
H a bit at an d B i olo gy : Artificial rice fields temporarily planted for agricultural experi-
ments. Life cycle unknown.
Di stri bu ti on: Only known from man-made habitats in Slovakia.
6. Hydrachna distincta K OENIKE , 1897 (7-38 b)

1897b Hydrachna distincta K OENIKE, Zool. Anz. 20: 394.
De sc ri pti on : Male unknown (females and males ascribed to H. distincta by H ALBERT
1911 probably representing H. conjecta, see below), data given here based exclusively
on the (damaged) female holotype. In the morphology of frontal area, coxae and mouth-
parts very similar to H. cruenta. Coxae 4 with one seta. Uncommonly in Hydrachna, the
female genital field an inverted heart-shape and anteriorly bluntly pointed. Approximate
palp setae numbers: P-1 without seta (artifact?), P-2 four dorsal, three lateral, P-3 three
dorsal, no lateral setae. Fe m a le : Idiosoma length 3000; frontale 820; coxal field 1250
lm; Cx-IV posterior margin as in H. cruenta; genital field length/width 400/450 lm, ante-
riorly bluntly pointed, posteriorly slightly concave, acetabula extending also onto the dor-
solateral part of the sclerite (in unextended position directed to the centre of the body and
facing the folded membrane flanking the genital field). Gnathosoma length 750, rostrum
350, palp total length 840, length/height P-1 200/320 (0.6, P-2 240/230 (1.0), P-3 240/120
(2.0), P-4 110/70 (1.6), P-5 50/30 (1.7) lm.
S im i la r spe c ie s: The populations described by British and Irish authors under the name
of H. distincta, including the only published male specimen (H ALBERT 1911), agree with H.
conjecta and are best ascribed to that species (D AVIDS et al. 2005). They were attributed to
H. distincta due to minor differences in shape of the frontal sclerites (gradually tapering
posteriorly, medial margin equally concave, not angular) lying obviously within the varia-
bility range of H. conjecta. It is reasonable to assume that the holotype of H. distincta is a
misshaped specimen of H. conjecta. However, since WAINSTEIN (1980) found minor mor-
phological differences between larvae attributed by him to H. conjecta respectively H. dis-
tincta, the question remains open until the variability of the related species is investigated
on the basis of more adults of both sexes.
Pr eadult stages: Larva: WAINSTEIN (1980). De u ton ym p h: not described.
B io lo gy : B IESIADKA & C ICHOCKA (1994) found larvae attributed to this species almost
exlusively parasitic on Hesperocorixa sahlbergi (F IEBER ).
H a bit at : Described from a lake, further uncertain records from (eutrophic) ponds.
Di stri bu ti on: Recorded from Drecksee, Schleswig-Holstein, Germany. Further uncertain
records from Eastern and Central Europe.
7. Hydrachna geographica M ÜLLER , 1776 (7-37 d, 7-38 d)

1776 Hydrachna geographica M ÜLLER, Hallager: 190.
De sc ri pti on : Dorsal idiosoma of living specimens marked by a variously ramified black
cross on a red background; no sclerites associated with preocularia; a pair of rod-shaped
sclerites flanking the postocularia, between these and the lateral eye sclerites up to five pairs
of minute sclerotized dots, occasionally some of them fused. Medial margin Cx-III equally
convex, Cx-IV nearly straight; posteromedial angle of Cx-IV extending, with a cleaver-
shaped extension of secondary sclerotization; five to nine setae on Cx-IV. Gnathosoma
with a very long, weakly curved rostrum (length rostrum/gnathosomal base 4 1.5).
Palp very slender, P-1 with a narrow base, without setae; P-2 dorsal margin in the basal
half strongly convex, in the distal half straight and here nearly parallel to the ventral mar-
gin, with 12-20 dorsal and up to six lateral setae; P-3 slender, slightly inflated in the centre,
with three to eight dorsal setae and five to eight medial setae; P-4 with a straight or convex
dorsal margin, maximum height in the middle or distally. M al e : Idiosoma length 5000-
6000, frontal sclerite 400; coxal field 2000; genital field length/width 880/840 lm, sub-
triangular, anteriorly indented, some of the anterior acetabular setae grouped closely to-
gether; gonopore forming a deep indentation at the posterior margin of the genital field.
Gnathosoma length 1660, rostrum 1000, chelicera 2200; palp total length 1730; length/
height P-1 300/460 (0.7), P-2 500/280 (1.8), P-3 620/140 (4.4), P-4 220/90 (2.4), P-5 90/40
(2.3) lm. Fe m a le : Idiosoma length 6000-9000, frontal sclerite 560-620, coxal field 2300-
2600; genital field length/width 630-700/840-1000 lm, anteriorly more or less indented,
with equally convex or bluntly pointed posterior margin. Gnathosoma length 2400, ros-
trum 1600, chelicera 2800; palp total length 2280-2410, length/height P-1 400-420/590-
640 (0.7), P-2 670-710/370-380 (1.8-1.9), P-3 850-930/190-195 (4.5-4.8), P-4 250/110
(2.27, P-5 100-110/50-60 (1.7-2.2) lm.
Pr eadu lt stages: L arva: WAINSTEIN (1980). D eu ton ym p h: K.V IETS (1936d), B ESSE-
LING (1964). Deutonymphs of H. geographica and H. incisa differ from all other European
species in the presence of more than one seta on Cx-IV. Distinction between these two spe-
cies at the deutonymphal stage probably possible based on Cx-IV setae number and dimen-
sions (both should be greater in H. geographica).
S im i la r spe c ie s: Notwithstanding numerous locality records from many parts of Europe,
this species has only rarely been discussed with regard to diagnostic morphological fea-
tures. For a long time defined only from idiosoma coloration and its large dimensions,
it is possible that it has frequently been confused with the closely related H. incisa. On
the other hand, several synonyms or related species dubiae have been published during
the past hundred years (see D AVIDS et al. 2005).
B io lo gy : Larvae parasitize in the subelytral space of larger dytiscid beetle species, mainly
Dytiscus marginalis LINNAEUS , less frequently on Acilius sulcatus (LINNAEUS ), Dytiscus
lapponicus G YLLENHAL and D. dimidiatus B ERGSTRÖM , rarely on some other species (Z A-
WAL 2003); on Cybister sp. (P IATAKOV 1915a). In Poland H. geographica is the species most
frequently encountered parasitic on dytiscid beetles, while it was found rare in Sweden
(L UNDBLAD 1968) and The Netherlands (S MIT & VAN DER H AMMEN 2000): during a large
faunistic inventory in the province of North-Holland, no parasitised dytiscid beetles were
found.
H a bit at : Pools, ponds and the macrophyte belt of lakes; in Poland, numerous larvae were
found in a peat bog with Sphagnum (Z AWAL 2003).
Di stri bu ti on: Holarctic; widespread in Europe, in Sweden found only in the southern
part of the country (L UNDBLAD 1968).
7. Hydrachnidia 349
8. Hydrachna globosa (D E G EER , 1778) (7-34 h, 7-39 g)

1778 Acarus aquaticus globosus D E G EER, Mémoires pour servir à l’histoire des Insectes: 146.
1897b Hydrachna uniscutata T HOR, Arch. Math. Naturv. 19: 19.
1916 Hydrachna gallica T HOR, Rev. Russe Entom. 16: 56.
1962 Hydrachna globosa neumani L UNDBLAD , Ark. Zool. (2) 14 (1): 8.
1964b Hydrachna globosa rotundata L ÁSKA, Biológia, Cas. Slov. Akad. vied 19: 922.
De sc ri pti on : Paired frontal plates broad anteriorly, tapering posteriorly, anteriorly sepa-
rated from each other by membranous integument, or fused to each other, occasionally
including the frontal eye, anterolateral margins near eyes concave, with a more or less de-
veloped lateral extension embracing the eye plate, medial margins with step-like indenta-
tion anteriorly to the postoculare. Medial margin of Cx-III nose-shaped, projecting, medial
margin of Cx-III+IV strongly concave; posteromedial corner of Cx-IV generally narrow
and extended. Palp slender, ventral margin P-2 concave, P-3 slightly concave basally,
slightly convex distally; approximate palp setae numbers: P-1 one dorsodistal, P-2 four
or five dorsal and one lateral, P-3 four to six (two dorsal, one to three lateral, one longer
ventrodistal). Ma le : Idiosoma length 1800-2900; frontal plate length/width 600-700/
270-300, coxal field length 800-1000; genital field length/width 360-470/370-480 lm,
rounded, anteriorly with a small indentation, posteriorly truncate, surface divided into
three sectors nearly equal in size (anterior right and left bearing acetabula, posterior bear-
ing densely arranged setae and the rather large gonopore), acetabula various in size (in
particular, some rather large ones in the posterolateral parts of the acetabula fields).
Gnathosoma length 700-830, rostrum 370-450, chelicera 850-1000; palp total length
850, length/height P-1 180-250/225-270 (0.8-1.1), P-2 220/225 (1.0), P-3 195-275/
65-80 (2.9-3.6), P-4 90-110/45-55 (2.0-2.2), P-5 44-50/20-25 (1.8-2.3) lm. F em a le :
Idiosoma length 1800-3600, frontal plate length/width 650-830/270-320; coxal field
length 800-1000, genital field length/width 290-300/360-500 lm, anteriorly slightly in-
dented, posteriorly rounded. Gnathosoma length 800-1000, rostrum 450-570, chelicera
1030-1250; palp total length 885, length/height P-1 200-325/280-315 (0.7-1.0), P-2
210-250/125-150 (1.6-2.4), P-3 250-300/75-85 (3.3-3.8), P-4 110-125/45-60 (1.9-2.4),
P-5 40-55/25-28 (1.6-2.0) lm.
S im i la r sp ec i es: The variable shape of the dorsal plates has led to the description of nu-
merous taxa now regarded as junior synonyms of H. globosa (see above for synonyms still
in use in recent bibliography, and K.O.V IETS 1987 for a complete overview). For the (sub)-
species most commonly cited, H. (globosa) uniscutata, with the paired frontal plates fused
anteriorly and including the frontal eye, and H. (globosa) separata, with the frontal eye
excluded from the fused frontal plates, L UNDBLAD (1962) demonstrated that all transitio-
nal modifications of this character state can be found coexisting in natural populations. In
contrast, S PARING (1959) reported on morphological and behavioral differences between
larvae of H. globosa s. str.-like adults, parasitizing Nepa rubra L INNAEUS and Ranatra li-
nearis (L INNAEUS ), and larvae of H. uniscutata-like adults, parasitizing Notonecta glauca
L INNAEUS . The same applies to H. gallica, a species similar to H. globosa in the shape of
dorsal plates, but supposed to differ in broader posteromedial corners of Cx-IV. According
to L UNDBLAD (1962), this character is subject to individual variability and D AVIDS et al.
(2005) could not detect diagnostic morphometric differences in populations attributed to
H. gallica by C ICHOCKA . However, C ICHOCKA (pers. comm.) observed a different position
of the median eye (at the posterior margin of the lateral eyes, behind the postocularia in H.
globosa - near the centre or at the anterior margin of lateral eyes, between the postocularia,
in H. gallica). As B IESIADKA & C ICHOCKA (1994) observed larvae of H. globosa s. str.-like
adults parasitizing on Nepa cinerea, but larvae of adults attributed to H. gallica restricted
to Ranatra linearis, there is some evidence that the question concerning the synonymy of
these two species merits additional investigation over a wider geographical range. For the
time being we follow L UNDBLAD (1962) in considering H. uniscutata and its subspecies (see
K.O.V IETS 1987) as well as H. gallica, as junior synonyms of H. globosa, keeping in mind
that this taxon could represent a species complex. Further synonyms of the stem species are
Hydrachna globosa rotundata L ÁSKA , 1964, and probably also H. rubicunda K.V IETS ,
1930d described from Spain and later recorded by L UNDBLAD (1956a) from Portugal (D A-
VIDS et al. 2005).
Pr eadult stages: Larva: WAINSTEIN (1980), D AVIDS (1973a).

Deut onymph : T UZOVSKIJ (1990).
B io lo gy : Eggs are placed by the ovipositor in the aerenchyma of higher aquatic plants,
larvae hatch after about four weeks at 20 C. Swimming under the surface of the water
larvae attack the ventral side of juveniles at the fifth larval stage or adults of aquatic het-
eropterans (see „similar species“ for discussion of differential host preference). When the
juvenile host moults, those larvae that have just started feeding are detached with the exu-
viae to which they are attached, while other larvae on juveniles pass through a phoretic
phase and are able to move to the imago at the moment of moulting. Larvae which do
not find a host die after about ten days. Each host can be infested by several dozen parasites.
Further development takes place only at low temperatures so that parasitic larvae of H.
globosa can come into hibernation. The life cycle is completed in the course of one
year. In spring, deutonymphs leave their host and are found mainly in April-June, adults
are able to survive for a few weeks only, from June to August (B ÖTTGER 1972a).
H a bit at : All kinds of standing waters, occasionally in slow running streams; in oligo-
trophic and eutrophic waters (L UNDBLAD 1968); also in brackish waters, but highly saline
waters are avoided (S MIT & VAN DER H AMMEN 2000); in the Mazurian lakelands in peat
bogs C ICHOCKA (1998, partly under the name of H. gallica).
Di stri bu ti on: Palaearctic; widespread in Europe, in Sweden only in the middle and
southern part of the country (L UNDBLAD 1968).
9. Hydrachna goldfeldi T HOR , 1916 (7-34 e, 7-35 b, 7-39 c-d)

1916 Hydrachna goldfeldi T HOR, Rev. Russe Entomol. 16: 59.
De sc ri pti on : Paired frontal plates with hardly developed or absent anterolateral exten-
sions. Coxae 3 with medial margin slightly convex, anteriorly forming an acute angle; Cx-
IV with medial margin equally concave, posteromedial edge broadly rounded (diameter of
this area twice that of excretory pore sclerite), with a large border of secondary sclerite, or
cleavage-shaped, or divided into two parts (one of them directed medially, the other one
posteriorly). Gnathosoma stout, with short rostrum bent posteriorly with a kink; P-2 ven-
tral margin concave, P-3 dorsal and ventral margins slightly undulating, P-4 maximum
height basally, P-5 rather slender; approximate setation: P-1 without setae; P-2 six dorsal
at regular distances, one medial, P-3 three dorsal and three lateral. Ma le : Idiosoma length
2200-2800; frontale length/width 700/270; coxal field length 1170; genital field length/
width 400/480 lm, with straight anterior and equally rounded lateral margins, gonopore
large, an unpaired minute extension at its anterior edge. Gnathosoma length 900, rostrum
450, palp total length 885, length/height P-1 180/250 (0.7), P-2 250/170 (1.5), P-3 280/100
(2.8), P-4 130/55 (2.4), P-5 45/25 (1.8) lm. Fe m a le : Idiosoma length 3000-4500; frontale
length/width 820/290, coxal field length 1120-1250; genital field length/width 300-400/
520-600 lm; similar to 7-37 a, but anterior indentation less distinct, the anterior third of
the acetabular field devoid of setae, and the „modified acetabulum“ not anterior from, but
between posterior setae fields. Gnathosoma length 1000, rostrum 500; chelicera 1240-
1300; palp total length 1030, length/height P-1 180-200/330-340 (0.6), P-2 250-290/
7. Hydrachnidia 351
170-195 (1.5-1.6), P-3 320-350/85-100 (3.5-3.8), P-4 135-140 /55-65 (2.2-2.5), P-5 50/
25-28 (1.8-2.0) lm.
Pr ea du lt sta g es: L a rv a: not described. De uto ny m ph: S OKOLOV (1928b).
B io lo gy : Larvae parasitic on Notonecta glauca L INNAEUS (B IESIADKA & C ICHOCKA
1994); in The Netherlands and Sweden adults only in springtime, from May at latest
to the end of June (S MIT & VAN DER H AMMEN 2000; L UNDBLAD 1962, 1968).
H a bit at : Temporary and permanent standing waters.
Di stri bu ti on: Western Palaearctic; reported from a limited number of countries in
Northern, Western and Eastern Europe, apparently everywhere rare.
10. Hydrachna incisa H ALBERT, 1903 (7-33 e, 7-34 c-d, 7-36 c, 7-37 b, 7-39)
1903a Hydrachna incisa H ALBERT, Zool. Anz. 26: 265.
De sc ri pti on : Frontal area with only one pair of crescent-shaped sclerites near postocu-
laria and a few sclerotized dots halfway between lateral eyes and postocularia. Coxae 4
with pointed posteromedial extensions, bearing two to five setae. Gnathosomal rostrum
extremely elongate; palps slender, approximate setation: P-1 without setae, P-2 17-21 dor-
sal and four lateral, P-3 five to six dorsal and six to seven lateral. Ma le : Idiosoma length
3500, width 2700; frontale length 210-300; coxal field 1400-1500, genital field length/
width 690-750/660-700 lm, heart-shaped, anterior margin with a narrow indentation
posteriorly continuing in a tongue-shaped furrow, posterolateral margins slightly concave,
gonopore large, directed posteriorly, with anterior margin equally concave or slightly in-
dented in the centre, 10-15 pairs of setae scattered in the acetabular field, a dense cover of
rather long setae in the tapering posterior area, in the immediate surroundings of the
gonopore a line of short, spine-like setae, a pair of minute humps near the anterolateral
edge of gonopore. Gnathosoma length 1550-1800, rostrum 1000-1200; chelicera
2000-2200; palp total length 1650-1710, length/height P-1 240-260/390-440 (0.6), P-2
520-560/250-260 (2.1-2.2), P-3 520-580/120-150 (3.8-3.9), P-4 220-250/80-90 (2.8),
P-5 60-70/50 (1.2-1.4) lm. Fe m a le : Idiosoma length 4000-5000; frontal sclerite
280-320; coxal field 1400-2000; genital field length/width 460-600/640-830. Gnathosoma
length 1800, rostrum 1100-1300; chelicera 2150-2680; palp total length 1785-2050,
length/height P-1 220-270/370-470 (0.57-0.59), P-2 580-660/250-310 (2.13-2.32), P-3
680-800/140-150 (4.86-5.33), P-4 240-250/90-100 (2.50-2.67), P-5 65-70/35-45
(1.44-2.0) lm.
S im i la r sp ec i es: Hydrachna bulgarensis (K.V IETS , 1940), described from a lake in the
Rila mountains, differs from H. incisa in having the genital field with anterior margin not
indented, presence of an elevated median ridge between the acetabula-bearing areas and a
slightly more stout palp (D AVIDS et al. 2005). Further studies are needed in order to under-
stand if these differences are of taxonomic importance. Hydrachna marita WAINSTEIN ,
1966, described from larvae, deutonymphs and adults in both sexes collected in Russia,
and later recorded also from Poland (C ICHOCKA 1983: larvae parasitic on dytiscid water
beetles, e.g. Rhantus latitans S HARP ) agrees well with H incisa (idiosoma maximum length
male 3000, female 4000 lm; frontal sclerites crescent-shaped, elevated number of setae on
Cx-IV, gonopore slit transverse, anterior margin not pointed, palp similar in setation). Dif-
ferences are found in the male genital field (very slight tapering posteriorly) and palp (sur-
prisingly stout - such as to induce considerable doubt if both sexes in the original descrip-
tion represent the same species). As no information is given on the shape of the gnatho-
somal rostrum, the species is not included in the determination key. WAINSTEIN (1966)
stated that this species reproduces by direct spermatophore transfer and hypothesized a
similar reproductive mechanism in all species with a slit-shaped, transverse male gonopore.
He introduced for these the subgenus Hymenohydrachna, a taxon not accepted by

K.O.V IETS (1987). This possible sister species of H. incisa, as well as particular reproduc-
tive mechanisms within this species group, merit attention in future research. Hydrachna
extorris K OENIKE , 1897, a species described from a single specimen of uncertain geogra-
phical origin, agrees with H. incisa in the combination of a long gnathosomal rostrum and
idiosoma and gnathosoma rather large in dimensions, but smaller than in H. geographica
(e.g., coxal field length 1600, genital field length/width 550/780, chelicera length 1960
lm). Differences are found in the presence of only one seta on Cx-IV, and the more stout
palp (length/height P-2 1.6, P-3 3.9, P-4 2.2) with lower setae numbers. As no specimens
with this character combination have been recorded from the study area, D AVIDS et al.
(2005) propose that H. extorris is not a member of the Central European fauna.
H a bit at a nd Bi ol og y: Stagnant waters. Life cycle unknown (but, see under „similar
species“).
Di stri bu ti on: British Isles, The Netherlands, South Germany, little documented; a doubt-
ful record of a deutonymph from Mecklenburg (T HOR 1928b).
11. Hydrachna incognita (WAINSTEIN , 1976) (7-34 a, 7-36 e, 7-37 c)

1976c Microhydrachna incognita WAINSTEIN, Inst. Biol. Vnutrenn. Vod; Trudy 31: 140.
De sc ri pti on : Frontal area with muscle attachments mostly as separate sclerotized dots,
not fused to longish platelets; only the preocularia may be drop-shaped due to a minute
posterior extension, and the postocularia are attached to a crescent-shaped sclerite. Exten-
sion of sclerotization rather variable, often different also on the right and left side of one
specimen. Coxae 4 bearing one seta. Palp robust, approximate setation: P-1 two dorsodis-
tal, P-3 five to seven dorsal and two or three lateral, P-3 two dorsal and three or four lateral.
Ma le : Sclerite near postoculare length 160, coxal field 700, genital field length/width 420/
480 lm. Gnathosoma length 600, rostrum 310; chelicera 960; palp total length 600, length/
height P-1 150/180 (0.8), P-2 145/95 (1.5), P-3 185/70 (2.6), P-4 85/40 (2.1), P-5 35/20
(1.8) lm. Fe m a le : Sclerite near postoculare length 110-180, coxal field 640-950 lm; gen-
ital field deeply indented anteriorly, acetabula posteriorly extending to the level of the tip of
the indentation, in the posterior part groups of hairs, genital field (7-37 c) length/width
310-410/430-550. Gnathosoma length 790-920, rostrum 410-470, chelicera 1150-
1360; palp total length 760-935, length/height P-1 170-200/200-260 (0.8-0.9), P-2
200-220/120-145 (1.5-1.7), P-3 230-280/80-95 (2.9-3.0), P-4 110-115/50-53 (2.2), P-5
45-50/25-27 (1.7-2.0) lm.
S im i la r sp ec i e s: This species was mostly defined from larvae, with scanty information
about a female adult, the male is described for the first time by D AVIDS et al. (2005). Hy-
drachna incognita, H. leegei and a further species described from Russia, H. vulpes (W AIN-
STEIN , 1976c), form a group of taxa which at the adult stage are characterized by generally
small dimensions grading into each other with regard to shape and arrangement of frontal
sclerites. In both sexes, H. incognita differs from H. leegei mostly in the absence of elongate
sclerites placed near, or fused to, the preocularia, and a stouter P-3 (length/height ratio 5
3.0, in H. leegei 4 3.3). Males differ furthermore in the genital field being more parallel-
sided and with setae surrounding the gonopore present in higher number and density, but
minor in size. Hydrachna vulpes, only known in the female, represents the extreme point of
reduction of the anterior sclerites (completely absent, preocularia on circular sclerites or
with a minute posterior appendix). Adult H. vulpes further differ from both other species in
having acetabula extending all over the surface of the genital plate and the absence of par-
ticular groups of fine setae surrounding the gonopore. Along with H. lupus (WAINSTEIN ,
1976c), at present only known at the larval stage, these taxa need a better definition based
on male morphology.
7. Hydrachnidia 353
Pr eadult stages: Larva: WAINSTEIN (1976c, 1980). De ut ony m ph : not described.

B io lo gy : Larvae parasitic in the subelytral space of hydrophilid water beetles, most fre-
quently on Anacaena limbata (FABRICIUS ) and A. lutescens (S TEPHENS ), to a much lesser
extent on Helophorus granularis (LINNAEUS ) and H. griseus H ERBST (Z AWAL 2003).
H a bit at : Stagnant waters (peat bogs and mesotrophic lakes).
Di stri bu ti on: Russia, Poland: C ICHOCKA & B IESADKA (1994), C ICHOCKA (1996b,
1996c), C ICHOCKA (1998), Z AWAL (2003).
12. Hydrachna inversa WALTER , 1927 (7-35 c, 7-39 b)

1927 Hydrachna inversa WALTER, in WALTER & M OTAŞ , Trav. Labor. Piscicult. Univ. Grenoble 11:
74.
De sc ri pti on : Female undescribed (see below). Frontal shield anteriorly pointed, poster-
iorly tapering, posterior margin straight or concave, median eye at the level of the centre of
lateral eyes. The facing medial margins of Cx-III close to each other, rounded; posterome-
dial angle of Cx-IV strongly projecting. Gnathosoma with long, curved rostrum. Palp not
stout, but P-3 only a little longer, often even shorter than P-2; approximate setation: P-1 one
dorsodistal, P-2 one dorsal, one lateral, P-3 two dorsal, two lateral. Ma le : Idiosoma
length/width 2300/2000, frontal shield 720/900; coxal field 1140/1890; genital field
360/510 lm, anterior margin concave, gonopore surrounded by a collar which is particu-
larly protruding in its anterior part. Gnathosoma length/width 1000/555 lm, with long,
curved rostrum; chelicera length 1300; palp total length 1032, length/height P-1 210/300,
P-2 315/165, P-3 300/90, P-4 155/60, P-5 52/30 lm.
S im i la r sp ec i e s: Due to the uncommon enlarged shape of the male genital field unique
among European Hydrachna species, WALTER (in WALTER & M OTAŞ 1927) believed he
described a female, but V IETS (1930d) and L UNDBLAD (1956a) pointed out that the holo-
type was obviously a male. The latter author records both sexes, but does not describe the
female. It should be easily recognizeable by the particular shape of the medial margin of Cx-
III and P-3 being little longer or shorter than P-2.
H a bit at a nd B io lo gy : Swamps, ditches and pools of streams. Life cycle unknown.
Di stri bu ti on: Southeastern France (Var), Spain (Aragon, Gerona); only four records pub-
lished.
13. Hydrachna juncta WALTER , 1926 (7-34 g, 7-36 a-b, 7-37 a)

1926c Hydrachna juncta WALTER, Boll. Soc. Sci. natur. Maroc 6: 135.
1956a Hydrachna globosa juncta L UNDBLAD , Ark. Zool. (2) 10 (1): 23
De sc ri pti on : In many details very similar to H. globosa, including the shape of the frontal
plates which may be fused or not anteromedially. Gnathosomal rostrum a little longer than
base. Palp very slender, P-3 slightly longer than P-2; approximate setation: P-1 one dorso-
distal, P-2 eight dorsal, two lateral, P-3 two dorsal, three lateral. M al e: Idiosoma length
2500, frontal plate 865; gnathosoma 1207, chelicera 1534; palp total length 1265, P-1 240,
P-2 390, P-3 413, P-4 158, P-5 64 lm - no information on the genital field in the publication
of the only record of a male (L UNDBLAD 1956a). F em a le : Idiosoma length 2700, frontal
plate length/width 600-975/ (one part:) 270-480; coxal field length 1225-1350, genital
field length/width 270-450/420-560; gnathosoma length 820-1300, rostrum 500-700, che-
licera 1030-1560; palp total length 700-1300, length/height: P-1 200-250/270-345 (0.7), P-
2 240-400/165-185 (1. 5-2.2), P-3 300-435/70-100 (3.9-4.4), P-4 120-160/50-75 (2.0-2.6),
P-5 55-60/28-40 (1.5-2.2) lm.
S im i la r spe c ie s: A poorly defined species, originally named for the fusion of the fronta-
lia. Since this character has been found to be variable, both in H. juncta and in the closely
related H. globosa, the distinction is possible only on the base of palp measurements. Di-
mensions of palp segments can be measured only from mature adults, juvenile H. globosa
may have narrowed palp segments due to weak sclerotization.
H a bit at a nd B io lo gy : Most records from pools of streams. In Sicily in a high mountain
lakelet, but at lower elevations H. globosa was found (D AVIDS et al. 2005). Life cycle un-
known.
Di stri bu ti on: Morocco, Spain, France, S Italy (Sicily).
14. Hydrachna leegei K OENIKE , 1895 (7-35 g, 7-37 f, 7-38 f)

1895d Hydrachna leegei K OENIKE, Abh. naturw. Ver. Bremen 13: 230.
1897d Hydrachna maculifera P IERSIG, Zool. Anz. 20: 354.
1908b Hydrachna fuscata S OAR, J. Quekett microsc. Cl. (s. 2) 10 (63): 276
De sc ri pti on : In the European fauna together with H. conjecta the species most reduced in
size. Frontal area with a pair of rod-shaped anterior platelets, often including the preocu-
laria, more rarely disintegrated into several minute sclerites, and a pair of more enlarged,
often drop-shaped posterior platelets including postocularia; between these two platelets,
two or three minute sclerotized dots in triangular arrangement. Coxae 3 medially forming a
pointed projection, Cx-III/IV medially meeting in a distinct, almost right angle; postero-
medial corner of Cx-IV forming a rounded right angle, hardly projecting, the superficial
extension often rather narrow, directed posteriorly. Gnathosoma rostrum relatively short,
strongly bent ventrally with a kink. Palp stout, P-2 ventrally strongly concave, P-3 basally
concave, distally slightly convex, P-4 maximum height distally; approximate setation: P-1
two dorsal setae slightly away from the distal margin, P-2 five to seven dorsal, two to four
lateral, P-3 two dorsal and four lateral. Mal e : Idiosoma length 1700-2400, frontal scerite
near postocularia 200; coxal field 770; genital field (7-38 f) length/width 400/470 lm,
heart-shaped with rather sharp anterolateral angles, anterior margin indented, gonopore
long (about 1/3 of genital field). Gnathosoma length 620, rostrum 320, chelicera 870; palp
total length 625, length/height P-1 120/185 (0.7), P-2 180/105 (1.7); P-3 205/60 (3.4), P-4
85/35 (2.4), P-5 35/18 (1.9) lm. Fe m a le : Idiosoma length 1800-2700; frontal sclerite near
postocularia 180; coxal field 900; genital field length/width 320/440 lm, as given in 7-37 c
for H. incognita, anterior margin with a deep indentation reaching almost to the posterior
plate margin, posterior margin concave. Gnathosoma length 880, rostrum 480; chelicera
1150; palp total length 797, length/height P-1 170/250 (0.7), P-2 200/120 (1.7), P-3 280/75
(3. 7); P-4 105/45 (2.3); P-5 42/25 (1.7) lm.
Pr eadu lt st ages: L arva: WAINSTEIN (1980). De u ton ym p h: B ESSELING (1964), T U-
ZOVSKIJ (1990).
S im i la r spe c ie s: For a long time, Hydrachna maculifera was considered a subspecies of
H. leegei. L UNDBLAD (1962) pointed out that the presumptive differences between the two
taxa (anterior sclerites fused with the preocularia in H. leegei, separate in H. maculifera;
medial margin of Cx-IV longer in H. maculifera) are variable characters not suitable to
separate the two taxa. For further discussion, see H. incognita. Hydrachna fuscata, a spe-
cies synonymized with H. leegei by K.V IETS (1956b), is kept as a separate taxon on the base
of deutonymphal characters by T UZOVSKIJ (1990).
B io lo gy : A typical springtime species, in The Netherlands with a peak occurrence around
mid-May, after May rarely encountered (S MIT & VAN DER H AMMEN 2000); larvae (in
small numbers) on the water beetles Helophorus flavipes FABRICIUS and Laccobius
minutus L INNAEUS (Z AWAL 2003).
7. Hydrachnidia 355
H a bit at : Permanent and temporary small standing waters; both in eutrophic and oligo-
trophic waters (L UNDBLAD 1968).
Di stri bu ti on: Palaearctic; in Europe widespread, in Sweden restricted to the southern
part of the country (L UNDBLAD 1968).
15. Hydrachna processifera K OENIKE, 1903 (7-33 a-b, d, 7-34 i, 7-37 e)

1903 Hydrachna processifera K OENIKE, Zool. Anz. 26: 534.
1895c ? Hydrachna inermis P IERSIG, Zool. Anz. 18: 302. (species incerta: D AVIDS et al. 2005)
De sc ri pti on : Frontal area with two pairs of minute, knob-shaped sclerites arranged near
preocularia resp. halfway between pre- and postocularia and a pair of larger, irregularly
shaped platelets laterally from, or fused to postocularia; postocularia particularly distant
from preocularia, together with the lateral eyes forming the edges of a longitudinally rec-
tangular (in related species mostly quadrangular) field. Medial margin of Cx-IV straight or
slightly concave, posteromedial corner extended with hook-shaped laterally-directed sec-
ondary sclerotisation. Palp stout, P-2 ventral margin straight; approximate setation: P-1
two mediodistal setae close to each other, P-2 eight dorsal and two to three lateral, P-3
two dorsal and three lateral, P-4 one medial and one ventrodistal. Ma l e: Idiosoma length
2320-3440; frontal sclerite 120; coxal field 1200-1300; genital field length/width 640-650/
600-610 lm, heart-shaped, posteriorly extending beyond posterior margin of fourth coxal
plates, anteriorly indented, gonopore long, extending over the posterior third of the plate.
Gnathosoma length 1120-1150, rostrum 550-620 lm, relatively long and slightly curved;
palp total length 1095, length/height P-1 200-210/290 (0.7); P-2 270/220-230 (1.2), P-3
400-410/150 (2.7), P-4 145/702 (2.0), P-5 70-75/35-45 (1.6-2.1) lm. Fe m al e : Idiosoma
length 3000-4130; coxal field 1200-1400; frontal sclerite 75-100; genital field length/width
380-440/550-680 lm, anteriorly indented. Gnathosoma length 1250-1400, rostrum 600;
chelicera 1700-1800; palp total length 1115-1210, length/height P-1 200-230/310-370
(0.6-0.7), P-2 240-270/200-250 (1.1-1.4), P-3 450-520/110-150 (3.3-4.3), P-4 140/60-
75 (1.9-2.3), P-560-70/35-40 (1.6-1.8) lm.
S im i la r sp e ci e s : Hydrachna processifera is similar to H. inermis, a little documented
species described from Germany. The question if it is a junior synonym of H. inermis cannot
be addressed because the type material of H. inermis is lost. Consequently, D AVIDS et al.
(2005) consider H. inermis a species dubia. Hydrachna danubialis, only known in the male
sex, is similar in the formation of the frontal area, but differs in its smaller dimensions and a
more stout palp (see there). Hydrachna vaillanti K.V IETS , 1951, known from a single male
collected in the Central Sahara is similar from most points of view, but differs in the genital
field having a more deeply indented anterior margin (D AVIDS et al. 2005).
P r ea d u lt s t a ge s : L arv a : S PARING (1959), WAINSTEIN (1980, under the name of H. in-
ermis). D eu ton ym p h: T UZOVSKIJ (1990).
H a bit at a nd B i ol og y: Standing waters, pools of slow flowing streams. Larvae parasitic
on beetles of the family Dytiscidae (S PARING 1959).
Di stri bu ti on: Palaearctic; widespread in Europe, in Northern Europe occurring as far
north as northern Sweden (L UNDBLAD 1962).
16. Hydrachna skorikowi (P IERSIG , 1900) (7-5 a-c, 7-33 c, 7-34 f, 7-36 d, 7-39 e)
1900 Hydrachna schneideri skorikowi P IERSIG , Ann. Mus. zool. Acad. Sci., St. Pétersbourg: 483
1930d Hydrachna valida integra K.V IETS, Arch. Hydrobiol. 21: 224.
1936d Hydrachna skorikowi K.V IETS in D AHL , Tierwelt Deutschlands 31: 73.
1956a Hydrachna schneideri integra L UNDBLAD, Ark. Zool. 10: 20.
1991a Hydrachna skorikowi integra G ERECKE, Lauterbornia 7: 40.

2005 Hydrachna skorikowi D AVIDS et al., Zootaxa 1061: 50.
De sc ri pti on : Dorsal shield with posterior margin indented, straight or slightly convex.
Posteromedial margin of Cx-IV broadly rounded, sometimes truncated. Gnathosoma
stout, rostrum shorter than base, bent ventrally with a distinct kink. Palp stout, P-3 nearly
as high as long, with weakly concave ventral margin; approximate setation: P-1 one dor-
sodistal, P-2 eight dorsal, two lateral, P-3 two dorsal, three lateral. Ma le : Idiosoma length
2000-3000; frontal shield length/width 530-800/700-1000; genital field 480-500/420-450
lm, heart-shaped, anterior margin weakly indented, lateral margins facing Cx-IV slightly
concave. Gnathosoma length 600-650, rostrum 340; chelicera 830-900; palp total length
680, length/height P-1 150/200 (0.8), P-2 160/155 (1.0), P-3 220/110 (2.0), P-4 100/50
(2.0), P-5 50/25 (2.0) lm. Fe m al e : Idiosoma length 2000-3000; frontal shield length/
width 800-900/1150-1200; genital field 200/320 lm, slightly wider than long, anteriorly
slightly indented, located in the anterior part of the bay formed by Cx-III+IV, this bay pos-
terior to the genital field strongly narrowed by the approaching medial margins of Cx-IV;
genital field enlarged, general morphology similar to 7-37 a (H. juncta), but with the inter-
space between the acatabula fields larger. Gnathosoma length 670, rostrum 360, palp total
length 690, length/height P-1 145/220 (0.7); P-2 165/160 (1.0), P-3 230/100 (2.3); P-4 100/
50 (2.0), P-5 50/30 (1.7) lm.
S im i la r spe c ie s: There has been extensive discussion about the correct name for this spe-
cies (for details see D AVIDS et al. 2005). Specimens with a straight or convex, not indented
posterior margin of the frontal shield are found more frequently in the southern part of the
distribution area and were attributed to the subspecies integra K.V IETS , 1930. As this taxon
and the stem species grade into each other and „integra“-like specimens may be found co-
existing with typical specimens, this distinction is not accepted by D AVIDS et al. (2005).
Pr eadult stages: Larva: WAINSTEIN (1980), D AVIDS (1973a). Deu to ny m ph: T UZOVS-
KIJ (1990).
B io lo gy : B IESIADKA & C ICHOCKA (1994) found larvae of the species to be exclusively
parasitic on Sigara nigrolineata (F IEBER ), S TEVENS & G REVEN (1994) mainly on Sigara
lateralis (L EACH ), and also on Callicorixa praeusta (F IEBER ), Corixa punctata (I LLIGER )
and Sigara falleni (F IEBER ). Females oviposit in the stem, not in the air cavities, of macro-
phytes, e.g. Chara. According to S TEVENS & G REVEN (1994), in Central Europe it is very
likely bivoltine and hibernates as a parasitic larva; deutonymphs appear until the end of
April, adults from May onwards; between mid-June and mid-August all stages were found,
in September only larvae were present; maximum numbers of adults were present in May
and August. In The Netherlands, adults occur from the end of April until September, with a
peak in late summer (S MIT & VAN DER H AMMEN 2000). Although many species of fish do
not feed on water mites, S TEVENS & G REVEN (1994) observed that the introduced seed-
pumpkin (Lepomis gibbosus L INNAEUS ) predated on nymphs of Hydrachna skorikowi.
H a bit at : All kinds of standing and slow running waters; in the Mediterranean area reg-
ularly in pools of temporary streams, tolerant against high salinity (conductivity values up
to 8.700 lS/cm: G ERECKE 1991). Often in man-made inland fresh water habitats (S TEVENS
& G REVEN 1994).
Di stri bu ti on: Palearctic; in Europe widespread, in the north up to Central Sweden, also
recorded from Northern Africa and Asia.
7. Hydrachnidia 357
Acknowledgements
Preparation of this chapter found financial support from the Schure-Beijerinck-Popping
Foundation (Amsterdam) and the Freshwater Biological Association (Ambleside) for meet-
ings of the authors and from Pro Acarologia Basiliensis for preparation of Figure tables, a
work done carefully by C. A LLGEIER (Tübingen). The following colleagues helped with
loan of collection material: J. B ECCALONI (The British Museum, London), M. C ICHOCKA
(Teacher Training College Olsztyn), J. D UNLOP (Museum für Naturkunde der Humboldt-
Universität Berlin), A. H ÄNGGI (Naturhistorisches Museum Basel), P. J ÄGER (Senckenberg-
Museum Frankfurt), A. K OLIBÁC (Moravian Museum Brno), T. K RONESTEDT (Museum of
Natural History Stockholm), J. P. O’C ONNOR (National Museum of Ireland Dublin) and P.
T UZOVSKIJ (Hydrobiological Institute, Borok).
M ICHAEL S TEVENS (Düsseldorf) made available light microscope and scanning electron
microscope photographs from his thesis. The publication of Fig. 7-9a was made possible
by courtesy of the Nederlandse Entomologische Vereniging (Amsterdam). The Editor
houses Blackwell (Oxford, R UNDLE et al. 2002) and E. Schweizerbart (Stuttgart, Archiv
für Hydrobiologie) generously allowed the reprint of figures published in books and special
issues from their production.
Furthermore, we are thankful to D.R. C OOK (Paradise Valley), O. M AKAROWA (Moscow)
and P. M ARTIN (Kiel) for important discussions and helpful suggestions.
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Taxonomic index Araneae 377
Taxonomic index Araneae

Agelenidae 4, 6 lutescens, Clubiona 2
agricola, Pardosa 3 Lycosidae 1f, 9
amphibia, Aranea 5 maculata, Arctosa 3
antiqua, Argyroneta 4 menardi, Meta 3, 11
aquatica aquatica, Argyroneta 5 merianae, Metellina 3, 11
aquatica japonica, Argyroneta 5 Mesothelae 1
aquatica, Argyroneta 1-4, 5, 6, 10, 13 Metinae (Tetragnathidae) 3, 11
aquaticus, Araneus 5 minutissima, Theonoe 4
Araneae -Arachnida- 1, 6f mirabilis, Pisaura 3
Araneidae 4, 7, 12 montana, Tetragnatha 3
Argyroneta 4 nigrita, Tetragnatha 3
Argyronetidae 4, 7, 10, 13 nitidula, Singa 4
Argyronetinae 4 palustris, Argyroneta 4
arundineti, Robertus 4 patagiatus, Heliophanus 2
atra, Erigone 4 phragmitis, Clubiona 2
bellicosus, Rugathodes 12 piraticus, Pirata 3
bituberculatum, Hypomma 3f Pisauridae 3, 7, 9
bonneti, Larinia 4 piscatorius, Pirata 3
bourneti, Meta 3 plantarius, Dolomedes 3
canestrinii, Mendoza 2 purbeckensis, Pardosa 2
caricis, Enoplognatha 4 pusilla, Microlinyphia 4
caricis, Sitticus 2 radiata, Marpissa 2, 9
cinerea, Arctosa 2 reclusa, Clubiona 2
clathrata, Neriene 11 reimoseri, Tetragnatha 3
clercki, Pachygnatha 3, 10 Salticidae 2, 9
Clubionidae 2, 9 saturatior, Pardosa 3
cornutum, Hypomma 11 sclopetarius, Larinioides 4, 12
cornutus, Larinioides 4 stagnatilis, Clubiona 2, 9
Cybaeidae 4, 7 sticta, Crustulina 4
Cybaeinae 4 stigmosa, Arctosa 3
dearmata, Tetragnatha 3 striata, Tetragnatha 3
dentipalpis, Erigone 4 tenuitarsis, Pirata 3
Desidae 1 Tetragnathidae 3, 10
Dictynidae 4 Theridiidae 4, 7, 12
distinguendus, Sitticus 2 Theridiosomatidae 3, 11
dorsalis, Kaestneria 4 Thomisidae 2, 9
elegans, Larinia 4 triangularis, Linyphia 4
extensa, Tetragnatha 3, 10 ulmi, Xysticus 2
fallax, Clubiona 5 urinatoria, Aranea 5
fimbriatus, Dolomedes 3, 9 wagleri, Pardosa 3
floricola, Sitticus 2 yulei, Vectaraneus 4
gemmosum, Theridiosoma 1, 3, 11
graminicola, Heriaeus 2, 9
hemerobium, Theridion 4
heri, Hypsosinga 4, 12
hygrophilus, Pirata 3, 9
impigra, Microlinyphia 4, 11
instabilis, Rugathodes 4, 12
insularis, Pirata 3
ixobolus, Larinioides 4, 12
knorri, Pirata 3
latitans, Pirata 3
leopardus, Arctosa 3
Linyphiidae 1, 3, 11
longipes, Argyroneta 4

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378 Taxonomic index Acari I
Taxonomic index Acari I

underlined: accepted taxa dealt with in this book.
abbreviations: -Acarid.- = Acaridida (chapter 3); -Hal.- = Halacaridae (chapter 5);
-Hydr.- = Hydrachnidia (chapter 7); -Oribat.- = Oribatida (chapter 4);
-terr.P.- = terrestrial Parasitengona (chapter 6).
Abrolophus -terr.P.- 169, 173, 177, 239 Anohydrachna -Hydr.- 333

Acaridae -Acarid.- 44-46, 56f Anopheles -Insecta: Diptera 265
Acaridida 24, 28f, 38 Anystidae -terr.P.- 161
Acerbitas -Hydr.- 161, 256, 264, 272 Apheviderulicidae -Hydr.- 242, 264, 283, 301
Acherontacaridae -Hydr.- 242, 264, 270, 295, Apheviderulix -Hydr.- 287
363 Aphidae -Insecta: Homoptera 178, 231
Acherontacarellus -Hydr.- 296 Aphithrombium -terr.P.- 221
Acherontacaropsis -Hydr.- 296 aquatica, Limnochares -Hydr.- 244, 258f, 262,
Acherontacarus -Hydr.- 260, 295-297 264, 270, 302-305
Acilius -Insecta: Coleoptera 326, 328, 348 aquaticus globosus, Acarus -Hydr.- 349
Actinotrichida -Acari 21, 45, 155, 247 aquatilis, Schwiebea -Acarid.- 53
aculeata, Unionicola -Hydr.- 263, 267 Arachnida 15, 36, 43, 57, 117, 155, 178,
acuta, Eylais infundibulifera -Hydr.- 313, 327 204-206, 209, 212, 238, 369, 372
adustum, Allothrombium -terr.P.- 225 Arbovirus -Viruses 274
affinis, Eylais -Hydr.- 313 Argiopidae -Chelicerata: Araneae 236
Agabus -Insecta: Coleoptera 296, 328 armatum, Stygothrombium -Hydr.- 293
akmolinski gracilipalpis, Eylais -Hydr.- 313 Arrenurinae -Hydr.- 248, 260
Aletes -Hal.- 152 Arrenuroidea -Hydr.- 241, 243, 264f, 272,
algeriensis, Monatractides -Hydr.- 277 284
algeriensis, Sperchon -Hydr.- 279 Arrenurus -Hydr.- 252, 254, 256-258, 261,
Allothrombium -terr.P.- 159, 164, 180, 220f, 263-265, 271-273, 275, 281, 285, 287f
223-226 aspratilis, Hydrachna -Hydr.- 336
alpicola, Piona -Hydr.- 265 assimilis, Platynus -Coleoptera: Carabidae 52
alpina, Eylais hamata -Hydr.- 326 Astigmata -Acarid.- 26, 38f
alpinum, Valgothrombium -terr.P.- 210, 214 Atractides -Hydr.- 264, 272, 274, 277-279, 287
alpinus alpinus, Porohalacarus -Hal.- 130f Atractothrombium -terr.P.- 179, 183, 196-198,
alpinus brachypeltatus, Porohalacarus -Hal.- 201-204, 210
117, 130, 132 aturid mites -Hydr.- 23
alpinus, Halacarus -Hal.- 130 Aturinae -Hydr.- 23, 243
alpinus, Porohalacarus -Hal.- 130, 132, 155 Aturus -Hydr.- 272, 285, 288
americana, Limnochares -Hydr.- 248 aurita, Eylais -Hydr.- 324
americana, Piersigia -Hydr.- 308 australis, Physolimnesia -Hydr.- 257
amnicus, Limnozetes -Oribat.- 106, 107, 108 autumnalis, Neotrombicula -terr.P.- 183
Amphibia 256, 262, 364 azubi, Limnochares -Hydr.- 303-305
amplipons, Eylais -Hydr.- 330 babori, Eylais -Hydr.- 313
ampulligera, Hirstiosoma -terr.P.- 174f baderi, Hydrovolzia -Hydr.- 299f
Anacena -Insecta: Coleoptera 326, 328 Baicalacarus -Hydr.- 291
Anactinotrichida -Acari 15 Balaustium -terr.P.- 158, 167, 172-174, 176
andinus, Thermacarus -Hydr.- 262 Balcanohydracarus -Hydr.- 288
andrei, Lohmannella -Hal.- 146f balticus, Halacarus-Hal.- 126
andrei, Porolohmannella -Hal.- 146 barbigera, Thyas -terr.P.- 161
Andrevella -terr.P.- 172 Bargena -Hydr.- 335
angelieri minor, Stygothrombium -Hydr.- 294 basiliense, Microtrombidium -terr.P.- 212
angelieri, Stygothrombium -Hydr.- 291, 293 basteri, Thalassarachna -Hal.- 116
angulata, Eylais -Hydr.- 323 Bdellidae -Acari 12, 29-31
angustiforme, Allothrombium -terr.P.- 221 bergstroemi, Eylais -Hydr.- 314, 327
angustipons, Eylais -Hydr.- 313 beroni, Leptus -terr.P.- 170
Anisitsiellidae -Hydr.- 242 bicolor, Podothrombium -terr.P.- 229
annemiae, Chelomideopsis -Hydr.- 257 bicornuta, Eylais -Hydr.- 313
Anoetidae -Acarid.- 55 Bidessus -Insecta: Coleoptera 326

https://doi.org/10.1007/978-3-662-55958-1
Taxonomic index Acari I 379
bifissa, Eylais muelleri -Hydr.- 313 Cerambycidae -Insecta: Coleoptera 55

bifoliosum, Enemothrombium -terr.P.- 179, Ceratopogonidae -Insecta: Diptera 158, 188,
183, 197, 200, 211 211, 216f
bifurca, Eylais -Hydr.- 327 Cerberothrombium -Hydr.- 291
bifurcatus, Tanaupodus -terr.P.- 219, 237 Cercothrombium -terr.P.- 195
birds 254 Cercyon -Insecta: Coleoptera 328
bisinuosa nodipons, Eylais -Hydr.- 312, Ceriodaphnia -Crustacea: Cladocera 331
314-316, 318, 320, 323, 329, 331 chappuisi, Parawandesia -Hydr.- 278, 307
bisinuosa oligotricha, Eylais -Hydr.- 329 chappuisi, Soldanellonyx -Hal.- 140-142
bisinuosa, Eylais -Hydr.- 312, 314-316, 318, chappuisi, Stygothrombium -Hydr.- 290, 293
320, 323, 331 Chara -algae 275, 356
bispinosum Stygothrombium -Hydr.- 293 Charletonia -terr.P.- 169, 171-174, 176f
bispinum, Microtrombidium -terr.P.- 212 Charonothrombium Coelostoma -Insecta:
bivalves 262, 268 Coleoptera 289, 291
bivirgulata, Hydrachna -Hydr.- 336, 345 Chelicerata 15, 32, 34-36
blackflies -Insecta: Diptera 257 Chironomidae -Insecta: Diptera 158
blackwellae, Centrotrombidium -terr.P.- 185 Chitohydrachna -Hydr.- 334
bonsdorffi, Cymatia -Insecta: Heteroptera 266, ciliatus, Limnozetes -Oribat.- 106f
345f cinerea, Nepa -Insecta: Heteroptera 349
boopis, Centrolimnesia -Hydr.- 275, 363 Cladocera -Crustacea 250, 256f, 312, 324, 360,
Bosmina Crustacea: Cladocera 331 367, 370
botschi, Microtrombidium -terr.P.- 212 Cladotanytarsus -Diptera: Chironomidae 259
Brachionus -Rotifera 257 clavatum, Campylothrombium -terr.P.- 180,
brachypeltatus, alpinus, Porohalacarus -Hal.- 197f, 201, 207
117, 130, 132 Clinocera -Insecta: Diptera 300
brama, Abramis -Pisces 259 clitellata, Eylais -Hydr.- 313
breei, Trombidium -terr.P.- 233 coiffaiti, Dromeothrombium -terr.P.- 199
brevimanum, Trombidium -terr.P.- 170f, 233, Coleoptera -Insecta 50, 52, 55, 195, 272, 296,
236 298, 300, 312, 326
brunneus, Lasius -Hymenoptera: Formicidae 52 coleoptrata, Cymatia -Insecta: Heteroptera
bulbisetum, Trombidium -terr.P.- 235f 344-346
Callicorixa -Insecta: Heteroptera 324, 345f, Collembola -Insecta 260
356 Colymbetes -Insecta: Coleoptera 326, 328
Callidosoma -terr.P.- 172f comosa, Hydrachna -Hydr.- 337, 339, 344
Calyptostoma -terr.P.- 159f, 164, 167-171 confervae, Hydrozetes -Oribat.- 102-105
Calyptostomatidae -terr.P.- 168 confusum, Valgothrombium -terr.P.- 210, 214
Calyptostomatoidea -terr.P.- 158, 161, 164, 168 conisetum, Trombidium -terr.P.- 235
Camerotrombidium -terr.P.- 180, 196, 198, conjecta, Hydrachna -Hydr.- 251, 255f, 259,
201f, 204-206, 210 262, 266, 271, 337, 344
Campylothrombium -terr.P.- 180, 196-198, consors, Eylais -Hydr.- 323
201f, 207f constricta, Piona -Hydr.- 275
canariensis, Eylais -Hydr.- 329 Copepoda -Crustacea 117, 257
cancellata, Hydrovolzia -Hydr.- 299f Copidognathinae -Hal.- 119f
cancellata, Thyopsis -Hydr.- 249, 251 Copidognathopsis -Hal.- 123
Carabus -Coleoptera: Carabidae 48, 52 Copidognathus -Hal.- 113-115, 118, 120-124
cardinalis, Charletonia -terr.P.- 169, 171, 174, cordifera, Eylais -Hydr.- 328
176 Corixa -Insecta: Heteroptera 324, 327, 345f,
carnea, Piona -Hydr.- 252, 259f 356
carpaticum, Diplothrombium -terr.P.- 183, 188, Corixidae -Insecta: Heteroptera 327, 344-346
189f cornuta, Eylais -Hydr.- 313
Caspihalacarus -Hal.- 117f, 120, 125f coronator, Arrenurus -Hydr.- 275
castanipes, Melanotus -Coleoptera: corporaali, Povilla -Insecta: Ephemeroptera
Elateridae 55 269
cavernicola, Schwiebea -Acarid.- 45f, 48, 53, crassipalpis, Hydrachna -Hydr.- 337, 345
55, 76-80 crassipes, Unionicola -Hydr.- 257-259, 267,
cavipontia, Eylais -Hydr.- 313 270
celtica, Eylais -Hydr.- 313 cruenta, Hydrachna -Hydr.- 265, 335f, 339,
Centrotrombidium -terr.P.- 164, 184-187 342, 346f
Crustacea 312, 363 discreta, Eylais -Hydr.- 271, 312f, 315f, 318,
crustaceans 116, 157, 256, 262, 204-206, 212, 320, 322, 324, 329
269, 297 distincta, Hydrachna -Hydr.- 337, 342, 347
Cryptops -Myriapoda: Cryptopidae 55f distincta, Sigara -Insecta: Heteroptera 256, 346
Cryptostigmata -Oribat.- 56 distinctus, Gyrinus -Insecta: Coleoptera 330
Culicidae -Insecta: Diptera 372 dividuus, Acherontacarus -Hydr.- 297
culicoides, Centrotrombidium -terr.P.- 185 dividuus, Eylais -Hydr.- 313
culicoidium, Enemothrombium -terr.P.- 211 Dixidae -Insecta: Diptera 190
Curteria -terr.P.- 173, 175 Dockovdia -Hydr.- 266
curtipalpe, Podothrombium -terr.P.- 228 dolgarae, Lobohalacarus -Hal.- 132f
curvimandibulata, Lohmannella -Hal.- dolgarae, Porohalacarus -Hal.- 132
146-149, 152 Dolichothrombium -terr.P.- 221
curvimandibulata, Porolohmannella -Hal.- 147 Dolopius -Coleoptera: Elateridae 55
curvipons, Eylais -Hydr.- 313f, 318, 320, 323, dominikae, Stirlitrombidium -terr.P.- 195
331 Dromeothrombium -terr.P.- 199
cuspidifer, Arrenurus -Hydr.- 263 dubia, Eylais -Hydr.- 291, 313f, 344f, 348, 355
cvetkovi, Lohmannella -Hal.- 146, 148f dubiosum, Podothrombium -terr.P.- 229
cvetkovi, Porolohmannella -Hal.- 148 dubiosum, Valgothrombium -terr.P.- 213
Cybister -Insecta: Coleoptera 327, 348 duplex, Eylais -Hydr.- 313
Cyclops -Crustacea: Copepoda 257 Dytiscidae -Insecta: Coleoptera 296, 326-328,
Cymatia -Insecta: Heteroptera 266, 324, 327, 330, 335, 355
344-346 Dytiscus -Insecta: Coleoptera 263, 327f, 348
dacicum, Trombidium -terr.P.- 232-234 Eatoniana -terr.P.- 172
dactyloporus, Copidognathus -Hal.- 212 Echinothrombium -terr.P.- 179f, 196, 198-200,
Dactylothrombium -terr.P.- 180, 196f, 199, 201 208-210
dadayi, Eylais -Hydr.- 327 egregia, Malaconothrus -Oribat.- 98
damselflies -Insecta: Odonata 266, 357 egregium, Paratrombium -terr.P.- 226f
ekmani, Neobrachypoda -Hydr.- 275
danielei, Limnochares -Hydr.- 304
Eleutherengona -Acari 25, 29-31
danielopoli, Schwiebea -Acarid.- 48
elliptica, Eylais -Hydr.- 313
danubialis, Hydrachna -Hydr.- 337, 346, 355
Elodes -Insecta: Coleoptera 300
Daphnia -Crustacea: Cladocera 258f, 324, 331
emarginata, Eylais -Hydr.- 313
dariae, Podothrombium -terr.P.- 229
Empididae -Insecta: Diptera 298, 300
Dartia -Hydr.- 260 Enemothrombium -terr.P.- 179, 183, 196f,
davidsi, Hydrachna -Hydr.- 345 199f, 203-209, 211, 213f
dearmatum, Podothrombium -terr.P.- 229 Enochrus -Insecta: Coleoptera 310, 326
degenerata, Eylais -Hydr.- 313f, 318, 320, 323 Ephemeroptera -Insecta 257, 259, 264, 269,
Delphacidae -Insecta: Homoptera 117 274, 331, 353
delphinense, Stygothrombium gallicum episcopalis, Curteria -terr.P.- 175
-Hydr.- 291f, 294 eregliensis, Eylais -Hydr.- 323
deltoides, Neumania -Hydr.- 274 errans, Johnstoniana -terr.P.- 161, 179, 187,
dentipes, Corixa -Insecta: Heteroptera 346 191-194
dentipes, Troglohalacarus -Hal.- 133f Erythraeinae -terr.P.- 169, 172f
denudata, Hydrachna -Hydr.- 336 Erythraeoidea -terr.P.- 158, 160-162, 164, 168f,
despiciens, Hydrodroma -Hydr.- 253, 256, 264, 174-176
270, 272 Erythraeus -terr.P.- 159, 169, 171, 173-175,
diatoms -algae 96, 117, 274 177f
dimidiatus, Dytiscus -Insecta: Coleoptera 348 Ettmuelleria -terr.P.- 195
Dimorphothrombium -terr.P.- 199f eugeni, Eylais -Hydr.- 313
Dinothrombium -terr.P.- 228 Eupodina -Acari 25, 29f
Diplodontus -Hydr.- 260 eurynympha, Schwiebea -Acarid.- 42, 45, 48,
Diplohydrachna -Hydr.- 333 63-69
Diplothrombium -terr.P.- 183-192, 194 eurynympha, Tyroglyphus -Acarid.- 48
Diptera -Insecta 88, 158, 169, 190, 194f, 204, Euthyas -Hydr.- 264, 286
206, 208, 211, 213, 217, 231, 265, 298, 300 Eutrichothrombium -terr.P.- 209, 211
discissa, Eylais -Hydr.- 324 Eutrombidiinae -terr.P.- 196, 199
discrepans, Trischidothrombium -terr.P.- 199 Eutrombidium -terr.P.- 159f, 196f, 199
discreta stagnalis, Eylais -Hydr.- 324, 329 excavata, Camisia -Oribat.- 97
exigua, Piona -Hydr.- 259 georgei, Hydrachna -Hydr.- 344

exiguum, Podothrombium -terr.P.- 229 Gerridae -Insecta: Heteroptera 304
eximia, Johnstoniana -terr.P.- 182, 186f, gibberipons, Eylais similis -Hydr.- 325
192-194 gibbosus, Lepomis -Vertebrata 356
extendens monofissa, Eylais -Hydr.- 325 gigas, Eylais -Hydr.- 313
extendens setosa, Eylais -Hydr.- 330 glauca, Notonecta -Insecta: Heteroptera 349,
extendens, Eylais -Hydr.- 312f, 315-320, 322f, 351
325f, 332 globator, Arrenurus -Hydr.- 261, 263
extendens, Hydrachna -Hydr.- 325 globosa juncta, Hydrachna -Hydr.- 353
extorris, Hydrachna -Hydr.- 336, 352 globosa neumani, Hydrachna -Hydr.- 349
Eylaidae -Hydr.- 242, 264, 279, 301f, 310 globosa rotundata, Hydrachna -Hydr.- 349f
Eylais -Hydr.- 254, 256, 260, 262, 264-266, globosa separata, Hydrachna -Hydr.- 349
271-273, 279, 286-288, 310-332 globosa, Hydrachna -Hydr.- 337f, 343, 349f,
Eylaoidea -Hydr.- 241, 244, 253, 260, 264, 283, 353f
301 globosus, Acarus aquaticus -Hydr.- 349
falcatus, Leptognathus -Hal.- 146 glubokensis, Eylais -Hydr.- 313f, 318, 325
falleni, Sigara -Insecta: Heteroptera 256, 345f, glyptoderma, Copidognathus -Hal.- 120
356 goldfeldi, Eylais setosa -Hydr.- 330
Feltria -Hydr.- 272 goldfeldi, Hydrachna -Hydr.- 337-339, 343,
ferociforme, Microtrombidium -terr.P.- 212 350
filipes, Podothrombium -terr.P.- 180, 183, 222f, gracile, Allothrombium -terr.P.- 224f
228-231 gracilipalpis, Eylais akmolinski -Hydr.- 313
flavipes, Helophorus -Insecta: Coleoptera 327, gracilis, Malaconothrus -Oribat.- 98
354 granularis, Helophorus -Insecta: Coleoptera
foili, Unionicola -Hydr.- 268f 353
foraminipons, Eylais -Hydr.- 328 Graptodytes -Insecta: Coleoptera 326, 328
Forania -terr.P.- 172 griseus, Helophorus -Insecta: Coleoptera 353
forficatus, Lithobius -Myriapoda: Gyrinidae -Insecta: Coleoptera 330
Lithobiidae 52 Halacarellus -Hal.- 118, 120, 126-128, 134
formosa, Unionicola -Hydr.- 268f Halacaridae -Acari 18, 24, 28, 32f, 363, 369,
fossarum, Sigara -Insecta: Heteroptera 346 374f
fossulata, Unionicola -Hydr.- 268 Halacarinae -Hal.- 119, 125
foveolatus, Trimalaconothrus -Oribat.- 99f Halacaroidea -Acari 29, 113
fturum, Trombidium -terr.P.- 232 halacaroides, Acherontacarus -Hydr.- 296
fuliginosum, Allothrombium -terr.P.- 180, 221, halacaroides, Hydrovolzia -Hydr.- 200, 298
223-226 Halacarus -Hal.- 126, 130f, 134, 136
fuornum, Trombidium -terr.P.- 232 Haliplidae -Insecta: Coleoptera 326, 328, 330
furcatus, Zetomimus -Oribat.- 110-112 Haliplus -Insecta: Coleoptera 326, 328, 330
furcigerum, Neotrombidium -terr.P.- 183 hamata alpina, Eylais -Hydr.- 326, 331
fuscata, Hydrachna -Hydr.- 354 hamata, Eylais -Hydr.- 254, 313f, 316, 318,
fusicomum, Atractothrombium -terr.P.- 203, 320, 322, 326, 331
210 Hauptmannia -terr.P.- 172, 177
fusiforme, Camerotrombidium -terr.P.- 205 Helochares -Insecta: Coleoptera 326
galeata, Eylais -Hydr.- 323 Helophoridae -Insecta: Coleoptera 327
gallica, Hydrachna -Hydr.- 349f Helophorus -Insecta: Coleoptera 327, 332, 353f
gallicus, Walterella -Hal.- 132 Hemiptera -Insecta 262, 266, 270, 272, 300,
gallicum delphinense, Stygothrombium 312, 344, 346
-Hydr.- 291f hephaestios, Copidognathus -Hal.- 121f
gallicum, Stygothrombium -Hydr.- 291f, 294 hercyniense, Stygothrombium -Hydr.- 293
gallicus, Lobohalacarus -Hal.- 132f Hesperocorixa -Insecta: Heteroptera 324,
gallicus, Porohalacarus -Hal.- 130, 132f 345-347
Gamasida -Acari 15, 26, 28, 30 Heteroptera -Insecta 298, 304, 350
gastropods 253, 262, 268 heterotrichum, Trombidium -terr.P.- 233
geniculatum, Trombidium -terr.P.- 332, 334f Heterozetes -Oribat.- 94, 109f
geographica, Hydrachna -Hydr.- 336f, 341f, hexagonus, Punctoribates -Oribat.- 109
348, 352 hilmari, Willmannella -terr.P.- 195
georgei, Eatoniana -terr.P.- 172 Hirstiosoma -terr.P.- 174f
georgei, Eylais -Hydr.- 326 Hirstiosomatinae -terr.P.- 172f
hispanicum, Podothrombium -terr.P.- 181, Hyphydrus -Insecta: Coleoptera 326, 328, 330
229-231 hyrcanus, Caspihalacarus -Hal.- 117, 125f
Histiostomatidae -Acarid.- 39, 44f, 55 Ignacarus -Hydr.- 253, 260
histiostomatids -Acarid.- 42, 55 ignotum, Allothrombium -terr.P.- 221
holosericeum, Trombidium -terr.P.- 180, 183, Ilybius -Insecta: Coleoptera 328
222f, 226, 232-234, 236 incarnatum, Allothrombium -terr.P.- 224f
holosericum, Trombidium -Hydr.- 256 incisa, Eylais -Hydr.- 327
Holothyrida -Acari 15 incisa, Hydrachna -Hydr.- 334, 336-338,
Homocaligidae -Acari 25, 29f 340-343, 348, 351
Homocaligus -Acari 25, 30f incognita, Hydrachna -Hydr.- 337f, 340f, 352,
homocomum, Platytrombidium -terr.P.- 213, 354
237 incognita, Microhydrachna -Hydr.- 352
hungarica, Eylais -Hydr.- 313 incurvata, Eylais -Hydr.- 314, 327
hungaricum, Trombidium -terr.P.- 232, 235 inermis, Hydrachna -Hydr.- 336, 355
Hydaticus -Insecta: Coleoptera 328 inflatus, Atractides -Hydr.- 279
Hydracarina -Hydr.- 241 infundibulifera acuta, Eylais -Hydr.- 312f, 315f,
Hydrachna -Hydr.- 251, 253, 255-257, 259f, 318, 320, 322, 324, 327, 329
262f, 265f, 271f, 279, 287f, 325, 333-356 infundibulifera lundbladi, Eylais -Hydr.- 327
Hydrachnellae -Acari 241, 357 infundibulifera meridionalis, Eylais -Hydr.-
Hydrachnida -Acari 32 324, 359
Hydrachnidae -Hydr.- 242, 264, 270, 279, 327, infundibulifera relicta, Eylais -Hydr.- 329
333 infundibulifera, Eylais -Hydr.- 312f, 315f, 318,
Hydrachnides -Hydr.- 333 320, 322, 324, 327, 375
Hydrachnidia -Hydr.- 26, 29, 241, 244-246, inopinatus, Limnohalacarus -Hal.- 135
249, 259, 269f, 274, 277-282, 285-289, 299, instabilis, Eylais -Hydr.- 325
301 insulare, Paratrombium -terr.P.- 180, 222f,
Hydrachnoidea -Hydr.- 241f, 244, 260, 264, 227f, 238
282f, 333 insularis, Eylais -Hydr.- 313
hydrachnoides, Halacarus-Hal.- 131 integra, Hydrachna skorikowi -Hydr.- 355
Hydraenidae -Insecta: Coleoptera 296, 323, integra, Hydrachna valida -Hydr.- 355
326 intermedia, Milandanielia -terr.P.- 199f
Hydrometridae -Insecta: Heteroptera 304 intermedia, Piersigia -Hydr.- 307f, 374
Hydrophilidae -Insecta: Coleoptera 326, 328, intermedia, Unionicola -Hydr.- 260, 268, 358
331, 335 inversa, Hydrachna -Hydr.- 336, 339, 343, 346,
Hydroporus -Insecta: Coleoptera 326, 328 353
Hydrothrombium -Hydr.- 291 ipidis, Schwiebea -Acarid.- 48
Hydrovolzia -Hydr.- 256, 265, 285, 295f, Ips -Coleoptera: Ipidae 50
298-300 Isobactrus -Hal.- 118f, 152f
Hydrovolziella -Hydr.- 296, 298 Ixodida -Acari 15, 24, 28, 30
Hydrovolziidae -Hydr.- 242, 264, 270, 295-297 jankovskii, Carabus -Coleoptera: Carabidae 48
Hydrovolzioidea -Hydr.- 241f, 252, 264, 269f, Johnstoniana -terr.P.- 161, 179, 182, 184, 186f,
282f, 294f 190-195, 237f, 240
Hydrozetes -Oribat.- 94, 101-105, 111f Johnstonianidae -terr.P.- 159, 161f, 179f, 181,
Hydrozetidae -Oribat.- 94, 101, 103 184, 186f, 237f, 240
Hydryphantes -Hydr.- 256, 265, 286-288 juncta, Hydrachna -Hydr.- 337f, 340f, 353f,
Hydryphantidae -Hydr.- 241f, 248, 256, 264f, 356
269f, 277-279, 289 Kamertonia -terr.P.- 173, 237
Hydryphantoidea -Hydr.- 241f, 244, 253, 260, Kamitrombidium -terr.P.- 195
264f, 283 karamani, Stygothrombium -Hydr.- 291, 294
Hygrobates -Hydr.- 252, 257, 259, 264, 269, karlovaicus, Podothrombium -terr.P.- 229
272, 275, 288 kasjani, Ettmuelleria -terr.P.- 195
Hygrobatidae -Hydr.- 241, 243, 266, 278 klugkisti, Paratrombium -terr.P.- 227
Hygrobatoidea -Hydr.- 241, 243, 260, 264f, kneissli, Trombidium -terr.P.- 233f, 236
272, 284 koenikei, Eylais -Hydr.- 313f, 316, 318, 322,
Hygrothrombidiinae -Hydr.- 291 327
Hygrotus -Insecta: Coleoptera 330 koenikei, Piersigia -Hydr.- 306-309, 374
Hymenohydrachna -Hydr.- 334, 352 kordulae, Podothrombium -terr.P.- 229
Hymenoptera -Insecta 52, 55, 228 krendowskii, Eylais -Hydr.- 313
kuehnei, Eylais -Hydr.- 330 Lohmannella -Hal.- 118f, 146-149

Laccobius -Insecta: Coleoptera 326, 328, 332, Lohmannellinae -Hal.- 119, 146
354 longicollis, Hauptmannia -terr.P.- 172
Laccophilidae -Insecta: Coleoptera 326 longipalpe, Diplothrombium -terr.P.- 186-190
Laccophilus -Insecta: Coleoptera 326 longipalpis, Eulais [!]-Hydr.- 328
lacustris, Hydrozetes -Oribat.- 102-105, 111 longipalpis, Eylais -Hydr.- 328
laevinodis, Myrmica -Hymenoptera: longipes, Valgothrombium -terr.P.- 214f, 237
Formicidae 55 longipons, Eylais -Hydr.- 323, 326
laminifera, Eylais -Hydr.- 329 longipora, Nilotonia -Hydr.- 264, 363
lapponicus, Dytiscus -Insecta: Coleoptera 348 longisetus, Trhypochthoniellus -Oribat.- 95-97
laricis, Ips -Coleoptera: Ipidae 50 longispina, Daphnia -Crustacea: Cladocera 331
Lasius -Hymenoptera: Formicidae 52 loricatum, Cercothrombium -terr.P.- 195
lasseni, Lassenia -terr.P.- 182, 217f ludwinae, Diplothrombium -terr.P.- 188, 190
Lassenia -terr.P.- 217, 219 lundbladi, infundibulifera Eylais -Hydr.- 327
lateralis, Sigara -Insecta: Heteroptera 356, 372 lupus, Hydrachna -Hydr.- 352
latipons, Eylais -Hydr.- 313, 330 lusitanica, Trombella -terr.P.- 183
latum, Trombidium -terr.P.- 232, 234f lutescens, Anacaena -Insecta: Coleoptera 353
Lebertia -Hydr.- 257, 287 macrocarpum, Podothrombium -terr.P.- 230f
Lebertioidea -Hydr.- 241f, 260, 264f, 272, 284, maculata, Limnesia -Hydr.- 256, 360, 376
372, 375 maculator, Arrenurus -Hydr.- 258
leegei, Hydrachna -Hydr.- 337, 339, 341f, 344, magna, Daphnia -Crustacea: Cladocera 331
347, 352, 354 maior, Trimalaconothrus -Oribat.- 99f
lemnae, Hydrozetes -Oribat.- 24-26, 102-105, major, Valgothrombium -terr.P.- 210, 214f, 217
111 Malaconothridae -Oribat.- 94, 97, 112
lepidopterans 264 Malaconothrus -Oribat.- 92, 94-98, 112
Leptognathus -Hal.- 146, 150f mamillatus, Limnohalacarus -Hal.- 135f
Leptus -terr.P.- 169f, 173f, 175, 178 mammals 44, 254
Lesteva -Insecta: Coleoptera 300 marenzelleri, Eylais -Hydr.- 326
limbata, Anacaena -Insecta: Coleoptera 353 Margaritiferinae -Mollusca 268
limicola, Rutripalpus -Hydr.- 262, 364 marginalis, Dytiscus -Insecta: Coleoptera 348
Limnesia -Hydr.- 256, 360, 363, 376 marginatus, Dolopius -Coleoptera: Elateridae
Limnesiidae -Hydr.- 242f, 281, 363f, 370, 376 55
limnetica, Piona -Hydr.- 275 marita, Hydrachna -Hydr.- 336, 351
Limnochares -Hydr.- 244, 248f, 258f, 262, 264, Mastothrombium -terr.P.- 209, 211
270, 274, 286-288, 302f, 304f, 359, 368f, media, Eylais -Hydr.- 313
371f, 376 mediterraneum, Trombidium -terr.P.- 232, 235f
Limnocharidae -Hydr.- 242, 289, 302, 361, medium, Microtrombidium -terr.P.- 212
369, 371 megalochirum, Paratrombium -terr.P.- 227
Limnocharinae -Hydr.- 242, 253, 302 megalostoma telmatobia, Eylais -Hydr.- 331
Limnocharoidea -Hydr.- 241 Megninietta -Acarid.- 46
Limnohalacarinae -Hal.- 113, 119, 134, 156 Melanotus -Coleoptera: Elateridae 55
Limnohalacarus -Hal.- 116, 118f, 134-137, mentonensis, Forania -terr.P.- 172
145, 156 menzeli, Schwiebea -Acarid.- 53
Limnohydrachna -Hydr.- 333 meridionale, Allothrombium -terr.P.- 224f
limnophila, Eylais -Hydr.- 325 meridionalis, Eylais -Hydr.- 324, 327
Limnozetes -Oribat.- 94, 106-108, 111 meridionalis, infundibulifera Eylais -Hydr.-
Limnozetidae -Oribat.- 94, 106, 111 324, 359
Limonia -Insecta: Diptera 194 Mesoveliidae -Insecta: Heteroptera 265, 298,
Limoniinae -Insecta: Diptera 169, 191 304, 366
limophila, Piersigia -Hydr.- 306-310, 365 Meteylais -Hydr.- 312f
limosa, Neumania -Hydr.- 274 microcrustaceans 15, 256, 312, 363
linearis, Ranatra -Insecta: Heteroptera 349 Microhydrachna -Hydr.- 335
linnei, Hesperocorixa -Insecta: Heteroptera Microsporidia -Protozoa 274, 361, 366
345f Microtrombidiidae -terr.P.- 159, 162, 166,
Litarachna -Hydr.- 274 179f, 181, 194-198, 200f, 210, 237, 240
Lithobius -Myriapoda: Lithobiidae 52 Microtrombidium -terr.P.- 169, 180, 182, 196f,
Ljania -Hydr.- 285 201-209, 211-215, 237
Lobohalacarus -Hal.- 118, 120, 128f Mideopsis -Hydr.- 257, 287, 372
mihelcici, Microtrombidium -terr.P.- 212 neumaniae, Sokolovia -Cyanobacteria 274

Milandanielia -terr.P.- 199f nicoleianus, Acherontacarus -Hydr.- 296f
minor, Microtrombidium -terr.P.- 203, 212 nigrolineata, Sigara -Insecta: Heteroptera 356
minor, Stygothrombium -Hydr.- 292, 294 Nilotonia -Hydr.- 260, 264, 363
minor, Stygothrombium angelieri -Hydr.- 294 Nippotrombium -terr.P.- 221
minutissimum, Allothrombium -terr.P.- 221f nobilis, Arrenurus -Hydr.- 275
minutus, Laccobius -Insecta: Coleoptera 354 nodipalpis, Atractides -Hydr.- 272
mirabilis, Mirabilithrombium -terr.P.- 199 nodipons, Eylais bisinuosa -Hydr.- 323, 331
Mirabilithrombium -terr.P.- 199f Noteridae -Insecta: Coleoptera 326, 328
molliculum, Allothrombium -terr.P.- 224f Noterus -Insecta: Coleoptera 326, 328
mollis, Astacocroton -Hydr.- 269 nova, Schwiebea -Acarid.- 45f, 55, 82-86
molluscs 262, 266-269, 358, 362f, 364, 368 novata, Eylais planipons -Hydr.- 329
monardi, Soldanellonyx -Hal.- 141-143 novus, Trimalaconothrus -Oribat.- 100
monodactylus, Malaconothrus -Oribat.- 95, 97f nullipons, Eylais -Hydr.- 331
monoeciportuense, Trombidium -terr.P.- 232 occulta, Eylais -Hydr.- 313
monofissa, Eylais extendens -Hydr.- 325 Odonata -Insecta 266, 304, 357, 363, 369f
Monohydrachna -Hydr.- 333 Odontohydrachna -Hydr.- 333
montanum, Podothrombium -terr.P.- 229 oedipodarum, Phanolophus -terr.P.- 173, 175
monticola, Trombidium -terr.P.- 234f okaënsi, Eylais -Hydr.- 330
moravicum, Valgothrombium -terr.P.- 213 Oligochaeta 259
Morelacarus -terr.P.- 183 oligochaetes 15, 117, 125
mosquensis, Eylais -Hydr.- 325 Opilioacarida -Acari 15
mosquitoes -Insecta: Diptera 256f, 264f, 366, orbiculata, Midea -Hydr.- 261f, 367
368, 371 Oribatida -Acari 15, 18, 24f, 29, 30, 33-37, 39,
mrazeki, Eylais -Hydr.- 323 89, 91-112
mrazeki, Eylais muelleri -Hydr.- 323 oscensis, Hydrovolzia -Hydr.- 300
Mucronothridae -Oribat.- 93f Ostracoda -Crustacea 312, 324, 363f
Mucronothrus -Oribat.- 92, 94-96, 112, palustris, Heterozetes -Oribat.- 109f
muelleri, Eylais -Hydr.- 325 palustris, Hydraena -Insecta: Coleoptera 323
muelleri bifissa, Eylais -Hydr.- 313 Panisopsis -Hydr.- 285, 358
muelleri mrazeki, Eylais -Hydr.- 313 pannonicum, Podothrombium -terr.P.- 229
muelleri triarcuata, Eylais -Hydr.- 332 papilligera, Hydrachna -Hydr.- 336
multispinosum, Podothrombium -terr.P.- 229 Parasitengona -Acari 14f, 17-20, 24-29, 31, 36,
murorum, Balaustium -terr.P.- 176 158-240, 241, 244, 253, 259, 269, 359, 363,
muscarum, Trichotrombidium -terr.P.- 195, 370, 372, 376
199f Parasoldanellonyx -Hal.- 120, 137-140
mutila, Eylais -Hydr.- 312, 314, 316, 318, 320, Paratrombium -terr.P.- 180, 220-223, 226f
322, 328 Parawandesia -Hydr.- 278, 291, 306f
mutilus, Sperchon -Hydr.- 26, 28 parisiensis, Hydrozetes -Oribat.- 102f, 111
Mycobatidae -Oribat.- 94, 108 parisii, Cryptops -Myriapoda: Cryptopidae 55
Myriapoda 55f parkeri, Andrevella -terr.P.- 172
Myrmica -Hymenoptera: Formicidae 55 Partnuniella -Hydr.- 277
Naiadacarus -Acarid.- 44f, 55f, 87 parva, Johnstoniana -terr.P.- 179f, 192-194, 239
Najadicola -Hydr.- 266, 374 parviscutatus, Parasoldanellonyx 138f
Najadicola ingens -Hydr.- 267f, 365 parviscutatus, Soldanellonyx -Hal.- 137f
nasalis, Mucronothrus -Oribat.- 94-96 parvulum, Microtrombidium -terr.P.- 212
neapolitanum, Allothrombium -terr.P.- 224 passerini, Abrolophus -terr.P.- 170
neglecta, Eylais -Hydr.- 313 passimpilosus, Tanaupodus -terr.P.- 183,
nematodes 15, 262 218-220
nemoricola, Rhinothrombium -terr.P.- 217f Peltodytes -Insecta: Coleoptera 326
Neotrombicula -terr.P.- 183 peragile, Podothrombium -terr.P.- 230f
Neotrombidiidae -terr.P.- 181 pergrande, Allothrombium -terr.P.- 224f
Neotrombidium -terr.P.- 182f, 238 perpera, Hydrachna -Hydr.- 336
Neotyrrellia -Hydr.- 252, 364 pexatum, Camerotrombidium -terr.P.- 198,
Nepa -Insecta: Heteroptera 262f, 349 201, 205f, 210, 240
Neoacherontacarus -Hydr.- 296 Phanolophus -terr.P.- 173, 175
Nepomorpha -Insecta: Heteroptera 335, 359f Phragmites -macrophytes 122, 131, 275
Neumania -Hydr.- 257 phreaticus, Halacarellus -Hal.- 127, 157
piersigi, Hydrachna -Hydr.- 336 punctulatus, Malaconothrus -Oribat.- 98

Piersigia -Hydr.- 286, 288, 306-309, 373f puripons, Eylais -Hydr.- 329
Piersigihydrachna -Hydr.- 334 purpureum, Paratrombium -terr.P.- 227
Piersigiidae -Hydr.- 242, 264, 301, 306, 363 pusillum, Microtrombidium -terr.P.- 162, 179f,
Piersigioidea -Hydr.- 241 182, 197, 201, 211f, 215, 237
Piona -Hydr.- 251f, 256f, 259-261, 265, 271, pygiacum, Trombidium -terr.P.- 232
275, 359, 362, 366, 370 quisquiliarus, Abrolophus -terr.P.- 24, 26, 28,
Pionidae -Hydr.- 243, 259, 266, 279 171f, 174, 176
piriforme, Podothrombium -terr.P.- 229 rackae, Diplothrombium -terr.P.- 188, 190
placophora, Hydrovolzia -Hydr.- 256, 265, racovitzai, Charonothrombium -Hydr.- 289
298-300, 357 racovitzai, Willmannella -terr.P.- 195
placophora, Polyxo -Hydr.- 300 raeticum, Microtrombidium -terr.P.- 212
planipons, Eylais -Hydr.- 312, 314-316, 319f, raeticum, Trombidium -terr.P.- 232
329 rafalskii, Atractothrombium -terr.P.- 203, 210
planipons novata, Eylais -Hydr.- 329 rahmi, Eylais -Hydr.- 326, 371
Platambus -Insecta: Coleoptera 328 rapax, Johnstoniana -terr.P.- 179, 186, 192f,
Platynus -Coleoptera: Carabidae 52 195, 239
plurispinum, Microtrombidium -terr.P.- 212 rasum, Camerotrombidium -terr.P.- 205f, 210,
Podothrombium -terr.P.- 180f, 183, 220-223, 240
228, 231, 238 rectipes, Huitfeldtia -Hydr.- 275
polonica, Kamertonia -terr.P.- 173, 175, 237 regulifera, Hydrachna -Hydr.- 336
Polydiscia -terr.P.- 217f, 240 relicta, Eylais -Hydr.- 315f, 319, 321, 329
Polyhydryphantes -Hydr.- 286, 288 relicta, Eylais infundibulifera -Hydr.- 329
Polyxo -Hydr.- 298, 300 Rhabdohydrachna -Hydr.- 333, 346
Pontarachna -Hydr.- 274, 375 Rhantus -Insecta: Coleoptera 328, 351
Pontarachnidae -Hydr.- 243, 249, 283 Rhinothrombium -terr.P.- 217f
Pontarachninae -Hydr.- 241 rhodinum, Echinothrombium -terr.P.- 179f,
Pontarachnoidea -Hydr.- 241
198, 200, 208-210, 237
Porolohmannella -Hal.- 114, 118f, 146-148,
Rhombognathinae -Hal.- 119, 152, 156
150f, 155, 157
Rhombognathopsis -Hal.- 123
Porolohmannellinae -Hal.- 119, 150
Rhombognathus -Hal.- 152, 155
porosipons, Eylais -Hydr.- 324
rhopalicus, Trombidium -terr.P.- 232
Potamogeton -macrophytes 275
Rhyncholimnocharinae -Hydr.- 242, 302
praeusta, Callicorixa -Insecta: Heteroptera
345f, 356 rimosa proceripalpis, Eylais -Hydr.- 328
prelipceani, Eylais -Hydr.- 313 rimosa triarcuata, Eylais -Hydr.- 332
proceripalpis, Eylais rimosa -Hydr.- 328 rimosa, Eylais -Hydr.- 312, 314f, 317, 319, 321,
processifera, Hydrachna -Hydr.- 334, 336-338, 323, 330f
341, 346, 355 rimosum, Trombidium -terr.P.- 235f
processus, Malaconothrus -Oribat.- 98 rimskykorssakowi, Eylais -Hydr.- 328
producta, Eylais -Hydr.- 313 roari, Podothrombium -terr.P.- 231
profundus, Copidognathus -Hal.- 121, 123 Roederioides -Insecta: Diptera 300
projectus, Eylais -Hydr.- 327 romaniense, Centrotrombidium -terr.P.- 185,
Prosobranchia -Mollusca 268, 364 239
Prostigmata -Hydr.- 250, 359f, 366, 371f Ropohalacarinae -Hal.- 119, 154
Proteylais -Hydr.- 313 Ropohalacarus 119, 154f
Protzia -Hydr.- 264, 288, 358 rostratus, Mucronothrus -Oribat.- 94, 96
Protziinae -Hydr.- 242, 278 Rotifera 262
pseudorimosa, Eylais -Hydr.- 323 rubicunda, Hydrachna -Hydr.- 350
Pseudoscorpiones -Chelicerata 260 rubipes, Abrolophus -terr.P.- 170, 239
Pseudowandesia -Hydr.- 288 rubra, Nepa -Insecta: Heteroptera 349
Ptychopteridae -Insecta: Diptera 190 Rudaemannia -terr.P.- 172, 177
pulcherrimum, Dactylothrombium -terr.P.- rudaensis, Rudaemannia -terr.P.- 172
180, 196f, 199, 201, 240 ruffoi, Acherontacarus -Hydr.- 297
pulvinum, Allothrombium -terr.P.- 223 ruficeps, Hebrus -Insecta Heteroptera 300
punctata, Corixa -Insecta: Heteroptera 345f, rugosus, Limnozetes -Oribat.- 106-108
356 ruienensis, Schwiebea -Acarid.- 50
Punctoribates -Oribat.- 94, 107-109 rupestris, Erythraeus -terr.P.- 169, 171, 174f
sahlbergi, Hesperocorixa -Insecta: Soldanellonyx -Hal.- 120, 137f, 140-145

Heteroptera 346f Sperchon -Hydr.- 257, 264f, 274, 279, 285,
salamandrarum, Hygrobates -Hydr.- 269, 364 288, 364, 366
salarius, Ignacarus -Hydr.- 253 Sperchontidae -Hydr.- 241, 242, 270, 358
Salmo -Osteichthyes: Salmonidae 46, 56 Sperchontoidea -Hydr.- 241
sanguineum, Camerotrombidium -terr.P.- 205f, sphagni, Oribata -Oribat.- 106
210 Sphagnum -moss 97, 100f, 105f, 108f, 116f,
saratovi, Eylais -Hydr.- 313 308, 348
scapularis, Homocaligus -Acari 25, 30f Spineylais -Hydr.- 313
Scarodytes -Insecta: Coleoptera 326 spiniferum, Microtrombidium -terr.P.- 212
Schizohydrachna -Hydr.- 333 spinipons, Eylais -Hydr.- 324
schmuttereri, Schwiebea -Acarid.- 45, 50, 70, 81 spinosum, Echinothrombium -terr.P.- 209f
schneideri, Centrotrombidium -terr.P.- 164, spinosum, Podothrombium -terr.P.- 229-231
184-187 spinulosus, Bufo -Amphibia 262
schneideri integra, Hydrachna -Hydr.- 356 sponges 253, 262, 266f
schneideri skorikowi, Hydrachna -Hydr.- 355 squamata, Polydiscia -terr.P.- 217f, 240
schweizeri, Atractothrombium -terr.P.- 203 stagnaliformis, Eylais -Hydr.- 329
Schwiebea -Acarid.- 24, 26, 42, 44-46, 48, 50, stagnalis, Eylais discreta -Hydr.- 329
53, 55-57, 59-62, 64-86 stagnalis, Eylais -Hydr.- 323f
Scirtidae -Insecta: Coleoptera 298, 300 stagnalis, Sigara -Insecta: Heteroptera 345
scolytids -Insecta: Coleoptera 50, 56 Staphylinidae -Insecta: Coleoptera 298, 300
Scorpionida -Chelicerata 260 stellae, Kamitrombidium -terr.P.- 195
sculptus, Trimalaconothrus -Oribat.- 99-101 steudelii, Tanaupodus -terr.P.- 219f
scupiensis, Stygohalacarus -Hal.- 144f Stictochironomus -Insecta: Diptera 263
Scutohydrachna -Hydr.- 333 Stirlitrombidium -terr.P.- 195
sellnicki, Punctoribates -Oribat.- 107-109 stoneflies 294, 376
semipons, Eylais -Hydr.- 313, 373 strandi, Podothrombium -terr.P.- 230f
semistriata, Sigara -Insecta: Heteroptera 345f striata, Sigara -Insecta: Heteroptera 257, 271,
Sericothrombium -terr.P.- 232, 236 345, 361
serratus, Astacopsis -Crustacea: Decapoda 269 stuarti, Valgothrombium -terr.P.- 213, 216
setiger, Sperchon -Hydr.- 264, 364 Stygohalacarus -Hal.- 119, 144f
setipalpis, Eylais -Hydr.- 328 Stygolimnochares -Hydr.- 306
setosa goldfeldi, Eylais -Hydr.- 330 Stygolimnocharinae -Hydr.- 242, 301, 307
setosa, Eylais extendens -Hydr.- 330 Stygomomonia -Hydr.- 272
setosa, Eylais -Hydr.- 312, 315, 317, 319, 321f, Stygothrombidia -Hydr.- 241
330, 375 Stygothrombidioidea -Hydr.- 241
setosulum, Paratrombium -terr.P.- 227 Stygothrombiidae -Hydr.- 242, 291, 371, 374,
setosus, Aletes -Hal.- 152 376
setosus, Trhypochthoniellus -Oribat.- 95-97 Stygothrombiinae -Hydr.- 291, 368, 374
Sida -Crustacea: Cladocera 331 Stygothrombioidea -Hydr.- 242, 248, 282, 289,
Sigara -Insecta: Heteroptera 256f, 271, 324, 291, 368
327, 345f, 356, 361, 372 Stygothrombium -Hydr.- 24f, 263, 278, 286f,
sigthori, Camerotrombidium -terr.P.- 205 290f, 296, 357
siliquoidea, Lampsilis -Mollusca 268 Stygovolzia -Hydr.- 298
silvestrii, Hydrachna -Hydr.- 347 subferociforme, Microtrombidium -terr.P.- 212
similis, Eylais -Hydr.- 325, 330 subnudum, Podothrombium -terr.P.- 229
similis gibberipons, Eylais -Hydr.- 325 subterranea, Schwiebea -Acarid.- 55
Simocephalus -Crustacea: Cladocera 331 subtile, Allothrombium -terr.P.- 223
Simuliidae -Insecta: Diptera 263f, 361, 364, Sucidothrombium -terr.P.- 196, 202
369f sucidum, Sucidothrombium -terr.P.- 196, 202
skorikowi, Eylais -Hydr.- 313 suctorians 117, 274, 310
skorikowi, Hydrachna -Hydr.- 255, 257, 334, sugambra, Eylais -Hydr.- 325
336, 338, 340, 343, 346, 355f, 372f sulcatus, Acilius -Insecta: Coleoptera 348
skorikowi, Hydrachna schneideri -Hydr.- 355 sumatrensis, Parasitalbia -Hydr.- 269
skorikowi integra, Hydrachna -Hydr.- 356 susteri, Trombidium -terr.P.- 232
Smaridinae -terr.P.- 172f sylvaticum, Atractothrombium -terr.P.- 179f,
soari, Eylais -Hydr.- 325 183, 197f, 201, 203f, 210, 237
soari valentiana, Eylais -Hydr.- 325 symmetrica, Eylais -Hydr.- 313
Symphyla -Myriapoda 260 Trischidothrombium -terr.P.- 199f

Syneylais -Hydr.- 310, 312 Troglohalacarus -Hal.- 118, 120, 133f
szalayi, Eylais -Hydr.- 325 Trombella -terr.P.- 183
talpa, Schwiebea -Acarid.- 42, 45-48, 50, 53, Trombellidae -terr.P.- 181
57-62 Trombiculidae -terr.P.- 158, 162, 180f
Tanaupodidae -terr.P.- 166, 180f, 217f, 240 Trombidia -terr.P.- 158-161
Tanaupodus -terr.P.- 183, 217-220, 237 Trombidiidae -terr.P.- 159, 162, 179f, 181, 220,
tantilla, Eylais -Hydr.- 312, 315, 317, 319, 321- 222f, 236
323, 329, 331f Trombidioidea -terr.P.- 158-161, 167-170, 178-
Tartarothyadinae -Hydr.- 242, 278 180, 182f, 204
taurica, Eylais -Hydr.- 323 Trombidium -terr.P.- 159, 180, 220f, 226, 231,
tectiporus profundus, Copidognathopsis 233
-Hal.- 123 Troupeauia -Acarid.- 46
tectiporus, Copidognathopsis -Hal.- 123 truncata, Euthyas -Hydr.- 170f
tectiporus, Copidognathus -Hal.- 121, 123f truncatellus, Limnebius -Insecta: Coleoptera
tectiporus, Rhombognathopsis -Hal.- 123 326
telmatobia, Eylais -Hydr.- 314, 319, 321, 331 trutta fario, Salmo -Osteichthyes: Salmonidae
telmatobia, Eylais megalostoma -Hydr.- 331 46
tenera, Eylais -Hydr.- 325 tuberculatum, Allothrombium -terr.P.- 223
tenuipalpe, Microtrombidium -terr.P.- 212 tullgreni, Eylais -Hydr.- 312, 314f, 317, 319,
tenuipons, Eylais -Hydr.- 324 321, 332
tenuissima, Hydrachna -Hydr.- 347 turbellarians 262
Teresothrombium -terr.P.- 232 tymoni, Podothrombium -terr.P.- 229
terrestris, Hydrozetes -Oribat.- 103 typhlops, Parasoldanellonyx -Hal.- 138-140
Tetrahydrachna -Hydr.- 334 Tyroglyphus -Acarid.- 48
Thalassarachna -Hal.- 116 undulosa, Eylais -Hydr.- 325
Thermacaridae -Hydr.- 242 Unionacea -Mollusca 268
thienemanni, Eylais -Hydr.- 325 Unionicola -Hydr.- 250, 254, 257-260, 262,
thienemanni, Hydrozetes -Oribat.- 102-105 266, 268, 275
Thorihydrachna -Hydr.- 334 Unionicolidae -Hydr.- 243, 266
Thyadinae -Hydr.- 242 uniscutata, Hydrachna -Hydr.- 349f
Thyas -Hydr.- 256, 286 uniscutatus, Isobactrus -Hal.- 152f
Thyasinae -Hydr.- 368 uniscutatus, Porohalacarus -Hal.- 152f
Thysanoptera -Insecta 177 uniscutatus, Rhombognathus -Hal.- 152
tibiotarsale, Hygrothrombidium -Hydr.- 291, uniscutatus, Ropohalacarus -Hal.- 154f
370 unisinuata, Eylais -Hydr.- 325
ticks -Acari 15, 30, 366f ussuriensis, Eylais -Hydr.- 330
Tipula -Insecta: Diptera 194 vaillanti, Hydrachna -Hydr.- 355
Tipulinae -Insecta: Diptera 169 valentiana, Eylais soari -Hydr.- 325
toldti, Trombidium -terr.P.- 232 Valgothrombium -terr.P.- 166, 179, 196, 199,
Torrenticola -Hydr.- 257 213
trapezipons, Eylais -Hydr.- 330 valgum, Valgothrombium -terr.P.- 179, 198,
Trhypochthoniellidae -Oribat.- 94, 96 201, 210, 213f, 216
Trhypochthoniellus -Oribat.- 92, 94, 96 valida integra, Hydrachna -Hydr.- 355
triangularis, Neumania -Hydr.- 274 variabilis, Eylais -Hydr.- 323
triarcuata, Eylais -Hydr.- 314f, 317, 319, 321, velutinus, Calyptostoma -terr.P.- 169-171
325 verae, Podothrombium -terr.P.- 229
triarcuata, Eylais muelleri -Hydr.- 332 verrucosa, Sperchonopsis -Hydr.- 257
triarcuata, Eylais rimosa-Hydr.- 332 vertebrates 180, 254, 262
Trichoptera -Insecta 32, 256f, 259 Victathrombium -Hydr.- 291
trichosus, Tripochthonius -Oribat.- 97 vietsi, Hydrovolzia -Hydr.- 299f
Trichothyas -Hydr.- 286 vietsi, Trimalaconothrus -Oribat.- 99-101
Trichotrombidium -terr.P.- 199 violacea, Leptognathus -Hal.- 150f
tricuspidum, Neotrombidium -terr.P.- 182 violacea, Porolohmannella -Hal.- 114, 151
trigonum, Eutrombidium -terr.P.- 160, 196f, violaceus, Acalyptonotus -Hydr.- 275
199 violaceus, Carabus -Coleoptera: Carabidae 52
trimaculatus, Leptus -terr.P.- 169f, 174f visurgis, Soldanellonyx -Hal.- 141, 143f
Trimalaconothrus -Oribat.- 94, 98-101 voeltzkowi, Eylais -Hydr.- 313
vorticellid ciliates 274 williamsoni, Hydrachna -Hydr.- 336

vulpes, Hydrachna -Hydr.- 352 Willmannella -terr.P.- 195, 199f
wackeri, Halacarus -Hal.- 134, 136 wilsoni, Eylais -Hydr.- 313
wackeri, Limnohalacarus -Hal.- 135-137 wittei, Diplothrombium -terr.P.- 187-189, 191
Walterella -Hal.- 128, 132 wolffi, Allothrombium -terr.P.- 223
Wandesia -Hydr.- 277f, 285, 288, 291
wolmari, Allothrombium -terr.P.- 224f
Wandesiinae -Hydr.- 241f, 270, 289
xymenae, Lassenia -terr.P.- 219
Wandesioidea -Hydr.- 241
weberi quadriporus, Lobohalacarus -Hal.- 128f ypsilophora, Unionicola -Hydr.- 268f
weberi weberi, Lobohalacarus -Hal.- 129 zbigniewi, Diplothrombium -terr.P.- 188, 191
weberi, Lobohalacarus -Hal.- 24, 26, 118, 128f Zetomimidae -Oribat.- 14, 109
weberi, Walterella -Hal.- 128 Zetomimus -Oribat.- 94, 109, 111
Wiedemannia -Insecta: Diptera 300 Zygoptera -Insecta: Odonata 255, 265f

Gerecke (Ed.), 2006 - Süßwasserfauna Von Mitteleuropa, Vol. 7, 2-1 Chelicerata - Araneae, Acari I

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Gerecke (Ed.), 2006 - Süßwasserfauna Von Mitteleuropa, Vol. 7, 2-1 Chelicerata - Araneae, Acari I

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Süßwasserfauna von Mitteleuropa

Reinhard Gerecke Editor

Chelicerata: Araneae, Acari I

Kees Davids Joanna Makol

Ralf Deichsel Harry Smit

Antonio Di Sabatino Henk van der Hammen

Grzegorz Gabryś Gerd Weigmann

Reinhard Gerecke Andreas Wohltmann

Terence Gledhill Eberhard Wurst

Süßwasserfauna von Mitteleuropa

Gedruckt auf säurefreiem und chlorfrei gebleichtem Papier

Springer Spektrum ist Teil von Springer Nature

2. Order Acari – General Introduction and Key to major groups

3. Acari: Limnic Acaridida (Astigmata)

4. Acari: Limnic Oribatida

6. Acari: Terrestrial Parasitengona inhabiting transient biotopes

Dr. Ilse Bartsch Dr. Peter Jäger

1. Order Araneae (Peter Jäger)

2. Order Acarri – General Introduction and key to major groups

3. Acari: Limnic Acaridida (Astigmata) (Eberhard Wurst)

4. Acari: Limnic Oribatida (Gerd Weigmann and Ralf Deichsel)

5. Acari: Halacaridae (Ilse Bartsch)

Subfamily Lohmannellinae V IETS , 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

6. Acari: Terrestrial Parasitengona inhabiting transient biotopes

© Springer-Verlag GmbH Deutschland 2017

Key to spider families occurring on fresh water sites

1777), A. leopardus (S UNDEVALL, 1833), A. maculata (H AHN , 1822), A. stigmosa (T HORELL ,

bituberculatum (W IDER , 1834), Kaestneria dorsalis (W IDER , 1834), Microlinyphia impigra

Family Argyronetidae T HORELL , 1870

Genus Argyroneta L ATREILLE , 1804

Argyroneta aquatica (C LERCK , 1757) (1-2 a–f, 1-5 a–l)

2. Order Acari - General introduction and key to the

© Springer-Verlag GmbH Deutschland 2017

Morphology and terminology

Sampling and sample processing

Classification of undissected material

The major groups of mites found in aquatic habitats

Key to nymphs and adults

General Glossary Acari

chelicera appendage of the first prosomal segment of Chelicerata, in mites at least

tracheae, tracheoles see „respiratory apparatus“

3. Acari: Limnic Acaridida (Astigmata)

© Springer-Verlag GmbH Deutschland 2017

Solenidia and famulus

Appearance of chaetae in ontogeny

Rules for modifications of leg chaetome

phoresy; its formation is induced by a mixture of environmental and ge-

Key to the adults of limnic Acaridida

Family Acaridae L ATREILLE , 1802

1. Schwiebea talpa O UDEMANS , 1916 (3-3 a-b, 3-4–3-9)

Ma le (h om e om or phi c ): Length of idiosoma approximately 270 lm; body form ovoid;

2. Schwiebea eurynympha (O UDEMANS , 1911) (3-3 c-d, 3-9–3-15)

3. Schwiebea schmuttereri T ÜRK & TÜRK , 1957 (3-16–3-21)

4. Schwiebea cavernicola V ITZTHUM , 1932 (3-22–3-27)

smooth. D orsu m : Posterior margin of propodosomal shield with an incision of variable

Genus Naiadacarus FASHING , 1974 (3-33)

Family Histiostomatidae B ERLESE , 1897 (3-3 e, f, 3-34–3-35)

Fig. 3-34: Idiosoma (venter) of a histiosto-

Fig. 3-35: Hormosianoetus sp.. a, male (light

4. Acari: Limnic Oribatida

© Springer-Verlag GmbH Deutschland 2017

Key to families and genera

Family Mucronothridae K UNST, 1971