© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.atmuseum KOENIG BONNER ZOOLOGISCHE MONOGRAPHIEN 52 2004 Herausgeber: Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn Arne R. Panesar EVOLUTION IN WATER MITES (HYDRACHNELUE, ACnNEDIDIA,ACARI) A revision of the Anisitsiellidae KOENIKE, 1910 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Schrifrleitung / Editor • . Karl-L. Schuchmann Zoologisches Forscliungsinstitut und Museum A. Koenig (ZFMK) Ornittiologie Adenauerallee 160 D-53 113 Bonn, Germany Email: [email protected] Die Reihe BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) erscheint seit 1971 in regelmäßiger Folge. Mit der vorliegenden Ausgabe werden nur noch Originalbeiträge in englischer Sprache publiziert. Die BZM nehmen vor allem solche Arbeiten auf, die zu umfangreich sind, um in der Zeitschrift BONNER ZOOLOGISCHE BEITRÄGE zu erscheinen. Schwerpunkte der BZM sind: Systematik, Taxonomie, Bio- geographie, Anatomie und Evolutionsbiologie. Alle eingereichten Manuskripte werden begutachtet. Autoren werden gebeten, sich vor Manuskriptvorlage mit dem Schriftleiter in Verbindung zu setzen. Erwerb der BZM: Bibliothek, Zoologisches Forschungsinstitut und Museum A. Koenig (ZFMK), Adenauer- allee 1 60, D-53 1 1 3 Bonn, Germany. The peer-reviewed series BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) has existed since 1971. The BZM consist of original zoology papers too long for inclusion in our institute's journal BONNER ZOOLOGISCHE BEITRAGE. Preferred manuscript topics are: systematics, taxonomy, biogeography, ana- tomy, and evolutionary biology. Manuscripts must be in English. Authors are requested to contact the editor prior to manuscript submittal. The BZM may be obtained from: The Library, A. Koenig - Zoological Research Institute and Museum of Zoology, Adenauerallee 160, D-531 13 Bonn, Germany. Author of this issue: Arne R. Panesar Published: April 25, 2004 Bonner zoologische Monographien Nr. 52, 144 pp. Editorial assistants of this issue: Angela Schmitz, Brian Hillcoat Price: EUR 42,- © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Arne R. Panesar EVOLUTION IN WATER MITES (HYDRACHNELLAE, ACTINEDIDIA, ACARI) A revision of the Anisitsiellidae KOENIKE, 1910 © Zoologisches Forschungsinstitut unci Museum Alexander Koenig, Bonn © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at PANESAR, EVOLUTION IN WATER MITES CONTENTS Abstract 6 Acknowledgments 7 Abbreviations 9 1 . Introduction 11 Hydrachnellae, systematic position and origin 11 Evolution of the water mite exoskeleton 13 Life history and behavior 14 Phylogenetic concepts referring to behavior 18 General morphology 19 2. Objectives of the study 24 The position of acetabula and their role in systematics of Hydrachnellae 24 The Rheolimnesia problem 26 Questions investigated 26 3. Material and Methods 28 Material 28 Material from field collections in India 28 Material from museums and private collections 30 Methods 30 Sampling methods, conservation, and preparation 30 Drawings, EM-photographs and, measurements 30 Parsimony calculation 30 4. What are Hygrobatoidea sensuTvLZows\d 1987"? 31 Discussion of segmental remnants 31 Suggested terms for metameric structures 36 Testing the superfamily concept of Tuzovski (1987) with a phylogenetic approach 36 Consequences for the concept of Hygrobatoidea 38 Diagnosis of the Hygrobatoidea (sT.) 39 5. Systematics in the Hygrobatoidea Koch, 1842 {sensiiTvaowski 1987) 40 Family group "^/^^-rc/^o/^-like" Hygrobatoidea 40 Bandakiopsidae fam. Nov. 42 Genus: Bandakiopsis Smith, 1979 [2 spp.] 43 Bandakiopsis phaluti sp. nov. 45 Genus: Cookacarus'Qdj:^, 1977 [1 sp.] 45 Genus: Oregonacarus Smith., 1989 [1 sp.] 47 Family group Limnesia-Xike' Hygrobatoidea 48 Family Limnesiidae 49 Subfamily Bandakiinae (subfam. nov.) 51 Genus: BandakiaThot, 1913 [20 spp.] 52 3 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 52/2004 Bandakia himachali sp. now. 55 Bandakia curvipalpis sp. nov. 57 Bandakia gangetica sp. nov. 59 Bandakia kulluensis sp. nov. 60 Genus: UtaxataxH^^itth, 1964 [2 sg. 9 spp.] 62 Utaxatax (s.s.) brahmeri sp. nov. 64 Utaxatax (s.s.) crassipalpis sp. nov. 66 Utaxatax (s.s.) gereckei sp. nov. 67 Utaxatax (s.s.) parvati sp. nov. 67 Subfamily Bharatoniinae (subfam. nov) 68 Genus: Bharatonia Cook, 1967 [1 sp.] 69 Genus: Shivatonia ^tn. nov. [1 sp.] 70 Shivatonia acetabidensis sp. nov. 72 Subfamily Anisitsiellinae (stat. nov.) 72 Platymamersopsis-^'conp 75 Genus: PlatymamersopsisVitts, 1914 [12 spp.] 76 Genus: Paddelia g^n. nov. [1 sp.] 78 Paddelia eichhorniae sp. nov. 79 Genus: Gilatonia K.O. Viets & Böttger, 1974 [1 sp.] 80 Mamersides-group 82 Genus: Hydrobäum ia Halik, 1930 [1 sp.] 82 Genus: Mamersides Viets, 1935 [2 spp.] 83 Genus: MamersopsidesVlets, 1916 [2 spp.] 85 Genus: Af/stmmö/JiH Nordenskiöld, 1905 [3 spp.] 86 Genus: Navamamersides Cook, 1967 [2 spp.] 87 Mahemamersides-groxip 88 Genus: Mahemamersides Ba.der, 1985 [1 sp.] 88 Genus: Nilgiriopsis Cook, 1967 [1 sp.] 90 Anisitsiellides-group 91 Genus: Aijisitsiellides Lundbhd, 1941 [11 spp.] 91 Genus: Zelandatonia Cook, 1992 [1 sp.] 93 Genus: Sigthoriella Besch, 1964 [1 sp.] 94 Nilotonia-gvoup 95 Genus: Anisitsidartia Cook, 1966 [1 sp.] 96 Genus: Anisitsiella Daday, 1905 [2 spp.] 97 Genus: MamersellaVieis, 1929 [2 SG, 4 spp.] 100 Genus: Rutacams h\xn(d)o\2id, 1937 [2 SG, 5 spp.] 103 Genus: M7öföw/Vz (Nordenskiöld, 1905) 105 Subgenus: Nilotonia (s.s.) Thor, 1905 108 Nilotonia indica Walter, 1928 110 Nilotonia cooki sp. nov. 110 S\})ogen\\s: Mamersonia (Vieis, 110 Snhgenxxs: Manotonia (yieis, \^'55) III Nilotonia {Manotonia) shivaisp. nov 112 Subgenus: Dartia Soa.r, 1917 114 4 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at PANESAR, EVOLUTION IN WATER MITES Subgenus: Tadjikodartia^didev, 1980 115 ' Subgenus: DartoniaVitts, 1929 116 Subgenus: DartiellaNitxs,, 1929 116 Nilotonia {Dartielld) fiindamentalis 118 Taxa incertae sedis 119 Genus: RanautoniaYleis, 1929 [1 sp.] 119 Genus: Sigthoria Ko^mkc, 1907 [1 sp.] '. 120 Genus: Stygomameropsis Qook, 1974 [2 spp.] 121 Genus: Psammotorrenticola Ang^eliei, 1951 [1 sp.] 123 Family group of Hygrobates-Yike and Mideopsis-like" Hygrobatoidea 124 6. Summary 125 7. Future research 132 8. Keys 133 Key 1: Adult key to the Superfamily Hygrobatoidea Koch, 1842 {sensu TmoYski) 133 Key 2: Families " Sperchon-iike' and Limnesia-Yike" Hygrobatoidea 133 9. References 135 10. Index 141 5 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 52/2004 ABSTRACT A cladogram for the Hydrachnellae (water mites) superfamilies resulting from a parsimony calculation is presented and discussed [(Hydryphantoidea (Hygrobatoidea (Wandesioidea (Hydrachnoidea (Hydrovolzioidea (Piersigioidea (Eylaoidea, Limnocharoidea) (Length = 13; Ci = 92; Ri = 87)]. The information for the data set was gained during the analysis of studies on body plans and segmental remnants in Hydrachnellae (Tuzovski 1987, Wiles 1997, Weigmann 2001). The concept of the superfamily Hygrobatoidea Koch, 1 824 {sensu Tuzovski 1 987) is confirmed in the present study by the above-mentioned analysis. The families Anisitsiellidae and Limnesiidae, in traditional systems (e.g.. Cook 1974) placed within two separate superfamilies, are both placed within the Hygrobatoidea in the system of Tuzovski (1987). This study proves that they are closely related to each other. Several years of investigations on the water mite fauna of the Himalayas have yielded new genera and species belonging to the Anisitsiellidae-Limnesiidae Complex, one of them intermediate between the families Anisitsiellidae and Limnesiidae. To resolve taxonomic contradictions in current systematics, a critical revision of the 132 species and 31 genera placed under the family Anisitsiellidae Koenike, 1910, is undertaken. The revision leads to the proposal of major changes in the taxonomic system, including the suggestion of a new family and two new subfamilies. The family Bandakiopsidae (fam. nov.) is proposed for 3 of the genera formerly placed under Anisitsiellidae {Bandakiopsis, Cookacarus, Oregonacariis). All the other Anisitsiellidae genera are included in one of the 3 following Limnesiidae subfamilies: Bandakiinae (subfam.nov), Bharatoniinae (subfam. nov.) and Anisitsiellinae (stat. nov.). From the material collected in India, 2 new genera {Shivatonia, Bharatoniinae, Limnesiidae, and /'^/(M'/z^, Anisitsiellinae, Limnesiidae) and 13 new species {Bandakiopsis phaliid, Bandakia curvipalpis, Bandakia gangetica, Bandakia himachali, Bandakia kulluensis, Utaxatax brahmeri, Utaxatax crassipalpis, Utaxatax gereckei, Utaxatax parvati, Shivatonia acetabulensis, Paddelia eichhorniae, Nilotonia cooki, and Nilotonia shivai) are described. New synonyms suggested include: Anisitsiellidae as junior synonym to Limnesiidae, Nilotoniinae as junior synonym to Anisitsiellinae, Mamersellides as junior synonym to Anisitsiella, Tadagatonia as junior synonym to Nilotonia,