WATER MITES of the GENUS TIPHYS KOCH, 1836 (ACARIFORMES: PIONIDAE) in RUSSIA P.V. Tuzovsky

Acarina 19 (2): 113–212 © Acarina 2011

WATER MITES OF THE GENUS TIPHYS KOCH, 1836 (ACARIFORMES: PIONIDAE) IN RUSSIA P.V. Tuzovsky Institute of Biology of Inland Waters, Russian Academy of Sciences, Borok, Yaroslavl Province, 152742 Russia

ABSTRACT: This study presents a detailed taxonomic review of water mites of the genus Tiphys Koch, 1836 (Hygrobatoidea: Pionidae) found in Russia during a long-term survey period between 1970 and 2008. The review includes (re)descriptions and illustrations of all known active stages of 18 Tiphys species found in this country: Tiphys bullatus (Thor, 1899), T. ensifer (Koenike, 1895), T. uchidai Viets, 1956; T. lapponicus (Neuman, 1880), T. latipes (Müller, 1776), T. pistillifer (Koenike, 1908), T. gladiator Tuzovskij, 2005; T. ornatus Koch, 1836; T. scaurus (Koenike, 1892), T. scaurellus Tuzovskij, 1983; T. torris Koenike, 1914; T. anadyrensis sp. n., T. chaunensis sp. n., T. kamchatkaensis sp. n., T. kolymaensis sp. n., T. minutiporus sp. n, T. samaraensis sp. n. and T. yaroslavlensis sp. n. Separate keys to all species based on females, males, deutonymphs and larvae are presented.

KEY WORDS: water mites, Pionidae, Tiphys, morphology, systematics, identification keys, female, male, deutonymph, larva, Russia.

CONTENTS differentiated deutonymph, and eight-legged sexually differentiated adult), and three inactive rest- Introduction ...... …. 113 ing stages (prelarva, protonymph and tritonymph). Methods and material ...... … 114 The resting stages are strongly reduced and repre- Morphology ...... … 115 sented only by integument; they do not have or- Systematics ...... 118 gans except for provisional ones. Larvae differ Genus Tiphys Koch, 1836 …...... 118 greatly from other active stages in their ecology Tiphys ornatus Koch, 1836 ...... 121 and morphology. Water mite larvae typically para- T. anadyrensis sp. n...... … 127 sitize aquatic and semi-aquatic insects, although T. scaurus (Koenike, 1892) ...... 130 in some cases larvae transform into deutonymphs T. scaurellus Tuzovskij, 1983 ...... 137 without feeding. Deutonymphs and adults are free- T. lapponicus (Neuman,1880) ...... 145 living predators feeding on various small inverte- T. latipes (Müller, 1776) ...... 152 brates. Adults and deutonymphs are well adapted T. kamchatkaensis sp. n...... 159 for a subaquatic life, but most larvae have adapta- T. kolymaensis sp. n...... 163 tions for aerial existence. The idiosoma of water mites is typically provided with a constant number T. torris Koenike, 1914 ...... 168 of setae, which are associated with glandularia in T. bullatus (Thor, 1899) ...... 175 deutonymphs and adults but not in larvae. T. samaraensis sp. n...... 175 The present study is a taxonomic revision of T. yaroslavlensis sp. n...... 177 water mites of the genus Tiphys Koch, 1836 (Hy- T. minutiporus sp. n...... 179 grobatoidea: Pionidae) occurring in the fauna of T. chaunensis sp. n...... 184 Russia. It includes descriptions of all known stag- T. ensifer (Koenike, 1895) ...... 188 es of species found in this country and provides T. uchidai Viets, 1956 ...... 196 keys to all of these species and stages. T. pistillifer (Koenike, 1908) ...... 196 The world fauna of the genus Tiphys currently T. gladiator Tuzovskij, 2003 ...... 204 includes about 50 species and subspecies (K.O. Keys to species of the genus Thyas ...... 208 Viets 1987). In the monograph “Limnofauna Eu- References ...... 211 ropaea”, K.O. Viets (1978) reported 9 species in Europe: Tiphys bullatus (Thor, 1899), T. convexi� INTRODUCTION palpis Ponyi, 1956; T. ensifer (Koenike, 1895), T. Water mites, inhabiting various types of fresh- lapponicus (Neuman, 1880), T. latipes (Müller, water biotopes, comprise a vast taxonomic group 1776), T. ornatus Koch, 1831; T. pistillifer (Koe- within the cohort Parasitengona (Acariformes) nike, 1908), T. scaurus (Koenike, 1892) and T. named in the acarological literature as Hydracari- torris Koenike, 1914. From the territory of the for- na, Hydrachnellae or Hydrachnidia. Like other mer USSR, the following species were recorded members of the Parasitengona, the life cycle of by Sokolow (1940): Tiphys bullatus, T. ensifer, T. water mites typically includes an egg, three active latipes, T. ornatus, T. scaurus and T. torris. Inves- stages (six-legged larva, eight-legged sexually un- tigations of water mites from different regions of

113 P. V. Tuzovsky

Russia over the past thirty years allowed me to de- Russia between 1970 and 2008. The review in- scribe two further species, T. scaurellus Tuzovskij cludes (re)descriptions and illustrations of all known (Tuzovsky 1983) from the Magadan Province and active stages of 18 species found so far in this coun- T. gladiator Tuzovskij from the Yaroslavl Prov- try. Separate keys to all species based on females, ince (Tuzovsky 2005). In addition, two more spe- males, deutonymphs and larvae are presented. cies have been found in the territory of Russia: T. METHODS AND MATERIAL lapponicus (Neuman, 1880) from Ladoga Lake (Sokolow and Yankovskaya 1962; Yankovskaya Water mites of the genus Tiphys live in tem- 1965) and T. pistillifer (Koenike, 1908) from the porary ponds, bogs, springs and streams. A dip net Yaroslavl Province (Tuzovsky 2008). with 0.2 mm mesh size was used to collect mites In water mites, including representatives of the in habitats with water plants (pools, bogs, littoral genus Tiphys, species descriptions and ecological zone of lakes and water bodies). For standing-wa- research deal mainly with the adult stage (Piersig ter sites without vegetation, I used a scraper to cut 1897–1900; Koenike 1909; Soar and Williamson off the upper layer of substrate. In flowing streams 1929; Motaş 1928; Viets 1936; Sokolow 1940; the scraper and the dip net loaded by a few stones Lundblad 1956, 1962; Besseling 1964; Szalay were used. The contents of sample was washed 1964, etc). For most water mites, larvae and deuto and placed into a large bucket with water, and then nymphs either are unknown or too poorly studied to small portions of that sample were examined part be included. Generalized data on pre-imaginal stag- by part in a white dish. Mites (adults and deuto es are given in a few publications for larvae (Spar- nymphs) were taken by a pipette and placed into ing 1959; Prasad and Cook 1972; Smith 1976; cupping-glasses or test-tubes. After that, mites Wainstein 1980; Smith and Cook 1991) and for were either mounted on permanent slides or fixed deutonymphs (Viets 1936; Tuzovsky 1990). for long-term storage. Morphology of Tiphys larvae has been de- Permanent slides were made with fresh speci- scribed for the following taxa: T. americanus (Mar- mens not fixed in Koenike’s-fluid using Hoyer’s shall), T. diversus (Marshall) (Prasad and Cook medium (Baker and Wharton 1952). Mouthparts 1972; Smith 1976); T. ornatus (Piersig 1897–1900; were separated first; then the body was covered Uchida 1932; Sparing 1959; Prasad and Cook with a cover slip and the contents was gently 1972); T. bullatus (Viets 1919); T. daisetsuensis squeezed out by means of slow pressing from the Imamura (Imamura 1954); T. eremitus Prasad and posterior to the anterior end. When the body was Cook (Prasad and Cook 1972); T. vernalis Habeeb, clean, it was washed and mounted on a glass slide. 1954 (Smith 1976), T. brevipes (Habeeb, 1954) The mouthparts were arranged so that pedipalps, (Smith 1976); T. scaurus, T. cooki Smith (Smith chelicerae and capitulum can be easily seen in lat- 1976); T. haliki Conroy (Conroy 1982), T. scaurel� eral view. In the case of working with specimens lus Tuzovskij (Tuzovsky 1983) and T. pistillifer that had been in fixative for a long time, mites (Koenike, 1908) (Tuzovsky 2008). were dissected along lateral line, cleared and A key of water mite larvae proposed by Wain- washed out in the water. Mouthparts, dorsal and stein (1980) contains very short descriptions of ventral sides were mounted on a slide. The mite larvae of four Tiphys-species: T. ornatus, T. lati� body and its mouth parts were always mounted on pes, T. ensifer and T. lutescens (Hermann, 1804). the same slide. The latter species is usually assigned to the genus To rear larvae, water mites were maintained Pionopsis (K.Viets 1956; Prasad and Cook 1972; in laboratory conditions. Eggs and larvae were ob- Cook 1974, Smith 1976; K.O. Viets 1987). tained from females kept individually in glass or Information on the morphology of deuto transparent plastic vessels (diameter 10–15 mm, nymphs has been given for a few species: Tiphys height 15 mm). Fragments of tree leaves, sedge bullatus (Viets 1919; Soar and Williamson 1929); stalks or other substrates from the habitat where T. latipes (Piersig 1897–1900); T. ornatus (Piersig mites had been collected were added to the ves- 1897–1900, 1901; Koenike 1909; Sokolow 1940); sels. Collected deutonymphs were held until their T. scaurus (Besseling 1974). A key to water mite transformation into adult mites. When the deuto deutonymphs (Tuzovsky 1990) included descrip- nymphs turn to the adult mites, an additional feed- tions of T. lapponicus, T. ornatus and T. scaurellus. ing is usually necessary for them. Young chirono- The present work gives a detailed taxonomic mid larvae and small crustaceans collected in the review of water mites of the genus Tiphys found in same water body as the deutonymphs were usually

114 Water mites of the genus Tiphys offered to freshly moulted adults. It was seldom Sce — scapulares externae (VI) possible to bring young deutonymphs (with coxal Li — lumbales internae (VIII) plates occupying more than half ventral body sur- Le — lumbales externae (VIII) face) to the size required for transformation to Si — sacrales internae (IX) adult mites even with additional feeding. Mature Se — sacrales externae (IX) deutonymphs (with relatively shorter coxal plates) Ci — caudales internae (X) could transform to adult mites even without the Pi — praeanales internae (XI) additional feeding. Vessels with deutonymphs, Pe — praeanales externae (XI) adult mites or eggs were always covered with a Ai — anales internae (XII) gaze or cover glass as mites often attempted to Ae — anales externae (XII) leave them, especially when no additional sub- Other structures of idiosoma and appendages: strate was provided. Other details for collecting i1–i5 — slit organs. water mites, their maintenance in laboratory con- P-1–5 — pedipalp segments (trochanter, femur, ditions, and mounting of slides are given in Prasad genu, tibia, tarsus) and Cook (1972), Cook (1974), Sokolow (1940), I–Leg 1–6 — leg number and segments 1–6 (tro- Tuzovsky (1987, 1990, 2007). chanter, basifemur, telofemur, genu, tibia, tarsus) Unless stated otherwise, all material used in tr –trochanter, the present study was collected by the author in f –femur different regions of Russia in the period 1970– g –genu 2008. Moreover, the collections of water mites de- ti –tibia posited in the Institute for Biology of Inland Wa- ta –tarsus ters of Russian Academy of Science (Borok, bf — basifemur Yaroslavl Province) were investigated. Material tf — telofemur for comparison was loaned from the private col- s1 — proximal solenidion on tibia of leg I–II in lection of Dr. Reinhard Gerecke (Zoological Insti- larva tute of the University of Tübingen, Germany) and s2 — distal solenidion on tibia of leg I–II in larva Dr. Henk van der Hammen (The Netherlands). as — acanthoid seta (terminal seta on tarsi I–III) Descriptions and redescriptions are given ac- dp.1–dp.3 — dorsal plates cording to the standard schemes proposed by Pras- vl.1–vl.4 — ventrolateral plates ad and Cook (1972) (for larvae) and Tuzovsky ey –eye (1990, 2007) (for adults and deutonymph). The eyp — eye plate terminology for idiosomal setae and slit organs ags — anterior genital sclerite folows Tuzovsky (1987). All measurements are in pgs — posterior genital sclerite micrometers. The number of measured specimens gp — genital plate is indicated for each stage. Only little-known syn- ac1–3 — genital acetabula (first, second, third) onymy is provided; exhaustive synonymies for bsch — basal segment of chelicera most species considered in the present paper were st — cheliceral stylet provided by Viets (1956), Lundblad (1962) and C1–C4 — coxal setae K.O. Viets (1987). ap1–ap3 — coxal apodemes Abbreviations and designations used in the aasc — anterior anal sclerite text and figures pasc — posterior anal sclerite Idiosomal setae referred to respective segments: pregen — pregenital plate Fch — frontales chelicerarum (I) gen — genital plate Fp — frontales pedipalporum (II) cx I–cx IV — coxal plates Vi — verticales internae (III) epp — excretory pore plate Ve — verticales externae (III) tmas — transverse muscle attachment scar Oi — occipitales internae (IV) ur — urstigma Oe — occipitales externae (IV) Specimen depositories and reference to ac- Hi — humerales internae (V) cession numbers are given using the following ab- He — humerales externae (V) breviation: IBIW — the Institute of Biology of Hv — humerales ventralia (V) Inland Waters of the Russian Academy of Science Sci — scapulares internae (VI) (Borok, Yaroslavl Province, Russia).

115 P. V. Tuzovsky

1 2

Figs 1–2. General view of idiosoma and terminology of adult mite of the genus Tiphys (scheme): 1 — dorsal view, 2 — ventral view. See abbreviations in the text.

7 3 8

Figs 3–9. The scheme of measurements of details of a structure of adult mite of the genus Tiphys: 3 — external genital organ of female, 4 — external genital organ of male, 5 — chelicera, 6 — pedipalp, 7 — leg, 8 — leg III claws of female, 9 — leg III claws of male. 116 Water mites of the genus Tiphys

Morphology Adult mites Although water mites lack distinct traces of segmentation or subdivision into tagmata (Fig. 1), arrangement of dorsal plates and glandularia likely reflect the ancestral segmentation. Tuzovsky (1987) suggested that the body of water mites of all active stages consists of 10 recognizeable segments and anal lobe (the numbering takes in ac- count segment VII which is reduced in all acari- form mites (Sitnikova 1978): I — segment of chelicerae, II — segment of pedipalp, III — verti- cal (legs I), IV — occipital (legs II), V — humeral (legs III), VI — scapular (legs IV), (VII — reduced), VIII — lumbal, IX — sacral, X — caudal, XI — praeanal and XII — anal lobe. The segments VIII–X are genital ones. The four anterior segments of the body constitute the anterior tagma (proterosoma), and the remaining ones represent the posterior tagma (hysterosoma), Fig. 10. Ventral view of deutonymphs of the genus Tiphys although in water mites these tagmata are not de- (scheme). limited. The idiosoma bears five pairs of lyriform organs (i1–i5). The anterior pair (i1) is always situ- trum is short, located ventrally and not reaching ated in immediate proximity to lateral eyes and re- the anterior edge of capitulum. ferred to proterosoma; others of four pairs (i2–i5) The chelicerae (Fig. 5) consist of two seg- correspond to anterior segments of hysterosoma, ments, the large basal segment with convex dorsal i.e. humeral, scapular, lumbar and sacral segments. edge and a small, crescent-shaped movable chela For the systematics of the genus Tiphys, shape (or stylet). Pedipalps (Fig. 6) consist of five seg- and size of idiosomal plates, legs III and IV, coxal ments: trochanter, femur, genu, tibia and tarsus. plates of all legs, pedipalps and genital field in The pedipalpal tibia has a dorsoapical or distolat- adult mites are the most taxonomically valuable eral peg-like seta. characters. The following three pairs of plates are Legs (Fig. 7) consist of six segments: tro- recognized on the dorsal surface of idiosoma (Fig. chanter, basifemur, telofemur, genu, tibia and tar- 1): one pair of anteromedial plates, one pair of an- sus. Coxae of all legs from plates on the ventral terolateral plates, and one pair of posteromedial surface of the body. On each side, coxae I–II and plates. The ventral surface of the body bears the coxae III–IV are fused forming two large coxal following plates (Fig. 2): four pairs of lateral plates plates in females (Fig. 2) and in deutonymphs (ventrolateralia 1–4), and two unpaired genital sc- (Fig. 10). Plates of the posterior coxal group are lerites (anterior and posterior). The medial mar- well separated in females and deutonymphs and gins of coxae IV are well developed. close to each other in males. Legs are provided The genital field (Fig. 3) is provided with a with swimming setae, the segments from trochant- pair of plates and typically three pairs of acetabula er to tibia bear numerous thin and thick setae; tarsi (as an exception, in T. mitchelli Cook, four to six bear mainly thin setae. The two tarsal claws on all pairs are present). Genital plates are separated in pairs of legs in adult and deutonymphs have two females, but fused to each other in males (Fig. 4). symmetric denticles each (Fig. 8), claws of leg III In general, the anterior genital sclerite of the male in males are asymmetric (Fig. 9). lies separate, but occasionally it is fused to the genital plate. Genital setae are variously arranged Deutonymphs along the lateral and posterior edges of genital Deutonymphs are rather similar to females plates, in females more numerous than in males. and differ mainly in smaller size and external As typical for all parasitengone mites, the ca- structure of the genital area. The genital opening is pitulum bears two pairs of setae (h1–h2). The ros- absent. External structures of the genital field (Fig.

117 P. V. Tuzovsky

11 12

Fig. 11–12. General view of idiosoma and abbreviations of larva of the genus Tiphys (scheme): 11 — dorsal view, 12 — ventral view. See abbreviations in the text.

10) are represented by the anterior and posterior moderately developed, and setae Ai and Ae are pairs of acetabula (ac. 1–2), two genital plates (gp), rather long. The anterior four pairs of lyriform or- a genital sclerite (gs), and a pregenital sclerite (pre� gans (i1–i4) are situated dorsally; the fifth pair (i5) gen). Three pairs of genital setae are located along is situated usually ventrally between the excretory the anterior edge of the genital plates. The number pore plate and setae Ci or Se. and arrangement of idiosomal setae and slit organs The capitulum (Fig. 13) has a well developed are similar to adults. All idiosomal plates are devel- rostrum; the anterodorsal setae (h1) of the capitu- oped to a lesser extent or some parts of plates are lum are longer and thinner than the ventral ones reduced, and the number of setae on pedipalpal and (h2). The basal segments of chelicerae (Fig. 14) leg segments is lower than in adult mites. are rather large and medially fused to each other. The cheliceral stylet is crescent-shaped, without Larvae subapical denticles. Pedipalps (Fig. 15) are short The morphology of larvae greatly differs from and stocky, have five free segments with the seta- that of deutonymphs and adult mites. Larvae have tion formula (P-1–5): 0, 1, 2, 4, 8 (s). Pedipalpal three pairs of legs and a well sclerotized idiosoma femur and genu are relatively elongated; the tibia (Figs 11–12). The formula of idiosomal setation is is short with a thick, curved dorsodistal claw; the 2–2–2–4–4–4–4–4–2–4–4. Idiosomal segment III tarsus is very small. Legs consist of five segments lacks setae Ve; segment V lacks setae Hv, segment (Fig. 16). Claws have a large central empodium XII (anal lobe) bears two pairs of setae. There are and a pair of thin ambulacra. Number of setae on no glandularia. The dorsal surface of idiosoma segments of legs is constant, with the following (Fig. 11) has a large shield bearing four pairs of setation formula, except eupathidia: I–Leg. 1–5: 1, setae (Fch, Fp, Vi, Oi). Lateral eyes are separated 7, 5 (s), 10–11 (2s), 14 (s, as); II–Leg. 1–5: 1, 7, 5 from each other and situated on small platelets on (s), 11 (2s), 13 (s, as); III–Leg. 1–5: 1, 6, 5(s), 10 each side. Coxae of legs II and III are fused to each (s), 11 (as). Sensillae (chemosensory setae) are other but are separated from coxae I; coxae I have situated only on the three distal segments of legs. two setae; coxae II and III each bear one seta (Fig. 12). Larval organs or urstigmata (ur) are small and SYSTEMATICS more or less circular, occupy a lateral position on Genus Tiphys Koch, 1836 the body border between coxae I and II and are Type species: Tiphys sagulatus Koch, 1836. provided with a cup. The excretory pore plate is By original designation.

118 Water mites of the genus Tiphys

13 14

Figs 13–16. The scheme of measurements of details of a structure of larvae of the genus Tiphys: 13 — capitulum, 14 — chelicera, 15 — pedipalp, 16 — leg I.

Diagnosis. Adult. Characters of the family Pi- plate, posterior eyes in soft integument. Suture onidae; body soft, with small dorsal and ventral line between coxal plates II–III incomplete, oblit- platelets. Coxal plates in females in four groups, in erated medially; four pairs of setae on coxal plates; males usually posterior coxal groups (in some spe- two pairs of setae on excretory pore plate. Bases cies also anterior coxal groups) fused medially; of setae C4 close to corresponding suture lines be- posteromedial setae of coxae IV hair-like; anterior tween coxal plates II and III (separated from them coxal group with short posterior apodemes; medial by 1–2 diameters). Setae Ci is very long and whip margins of coxae IV well developed; typically with like. Number of setae on legs I–III, except eu- three pairs of genital acetabula but in one species, T. pathidia, as follows: I–Leg 1–5: 1, 7, 5 (s), 10–11 mitchelli Cook, four to six pairs present; typically, (2s), 13 (s, as); II–Leg 1–5: 1, 7, 5 (s), 11 (2s), 13 male pregenital sclerite separated from genital field (s, as); III–Leg 1–5: 1, 6, 5(s), 10 (s), 11 (as). (occasionally fused with the remainder of the geni- Geographic Distribution. Holarctic (Cook tal field); male without petiole; leg IV of male, es- 1974). pecially genu (IV–Leg-4), exhibiting sexual dimor- Remarks. Karl Viets (1936, 1956) divided phism; tarsus (III–Leg-6) modified for the genus into three subgenera, Tiphys s. str., Pi� spermatophore transfer and exhibiting slight sexual onides Thor, 1911 and Acercopsis Viets, 1926, dimorphism; peg-like seta at tip of P–IV usually based on male features expressing the sexual di- well developed and without numerous heavy setae. morphism. In the subgenus Tiphys, the genital bay Deutonymph. Similar to females, differing of male is relatively shallow, the genital field bears mainly by smaller size and external structures of scattered setae; basifemur of leg II is without a the genital field. Gonopore is absent. External ventrodistal hump, genu of leg IV does not termi- structures of the genital field are represented by nate with a single heavy and curved seta, tibia of two acetabula (ac. 1–2) and three setae, which are leg II is without heavily thickened (spatulate or located on a plate on each side, and genital (gen) sword-like) setae. In Pionides, the genital bay is and pregenital (pregen) sclerites; genital plates very shallow, the genital field bears scattered set- separate. All dorsal plates are developed to a lesser ae; genu of leg IV bears terminally a single, long extent or absent. and curved seta, basifemur of leg II is without a Larvae. Dorsal plate large, covering nearly ventrodistal hump and tibia of leg II is without entire idiosoma, with four pairs of proterosomal heavily thickened setae. In Acercopsis, the genital setae. Anterior eyes on each side located on a bay is relatively deep, the genital field bears nu-

119 P. V. Tuzovsky

Table Known active stages of Tiphys species recorded in Russia Stages Species Male Female Deutonymph Larva Tiphys ornatus + + + + T. anadyrensis + – – – T. scaurus + + + + T. scaurellus + + + + T. lapponicus + + + + T. latipes + + + + T. kamchatkaensis + + – – T. kolymaensis – + – + T. torris + + + + T. samaraensis – + – – T. yaroslavlensis + – – + T. minutiporus – + – + T. ensifer + + + + T. chaunensis – + – + T. bullatus + + – – T. uchidai + – – – T. pistillifer + + – + T. gladiator + + – – merous setae arranged in four clusters; genu of leg eggs usually remained in their body. Larvae parasit- IV does not terminate with a single, heavy and ize imagines of all subfamilies of the family Chi- curved seta as in the subgenus Pionides, basife- ronomidae (Smith and Oliver 1976, 1986). mur of leg II has a ventrodistal hump, tibia of leg Adults and deutonymphs are active predators, II with one or two broadened (spatulate or sword- feeding on small crustaceans or juvenile insect lar- like) setae. This system was followed by subse- vae (Smith 1976). According to observations in quent acarologists (Sokolow 1940; Szalay 1974; the middle latitude zone of the European part of Besseling 1974; Cook 1974; K.O. Viets 1987, etc. Russia, adults and deutonymphs of Tiphys species and also in the present work); although Smith occur throughout the whole period in temporary (1976) considered Pionides as a distinct genus. waters (April–July), but are most abundant in the Classification of the genusTiphys and remain- first part of this period (Sokolow 1940; Tuzovsky ing Tiphysinae is based exclusively on male fea- 1974, 1996, 2007). Sokolow (1940) characterized tures expressing sexual dimorphism. Females, some species as representatives of so-called deutonymphs and larvae exhibit species-specific “springtime fauna” although he did not give de- characters, but cannot be referred to a particular tails concerning their phenology. genus or subgenus. Males have a short life, usually no longer than Biology. Larvae of almost all Tiphys species one week. Spermatophore transfer is direct, involv- from Russia (Table) obtained from females in the ing copulation (Viets 1914; Mitchell 1957; Smith laboratory failed to transform to deutonymphs in and Oliver 1986). Males die soon after sperm trans- the lab. This means that they require a parasitic fer, while females die after egg laying. Most fe- phase to continue the development — only larvae males finish egg-laying before temporary basins of T. scaurellus had transformed into the deutonymph disappear; larvae hatch and find a host. Subsequent without feeding (personal observations). Females destiny of larvae depends on these hosts, whether lay eggs on diverse substrata (usually leaves or the host will lay eggs in the same water basin or will stalks of plants). The greatest number of eggs (30– find another one; larvae quit their hosts when these 60) was laid by females of T. ornatus, females of return to the water and moult to protonymphs. If a other species produced 10–40 eggs. In the labora- water body dries out rapidly, larvae can find a shel- tory, females commonly laid not all eggs, some ter among plant roots and dead parts of plants. A 120 Water mites of the genus Tiphys

17 18

19 18–23

21 23 20 20 25 24 17, 24, 25 Figs 17–25. Tiphys ornatus, female: 17 — seta Fch, 18 — anteromedial and anterolateral dorsal plates, 19 — posteromedial dorsal plates, 20 — ventral view, 21–23 — genital plates, 24–25 — excretory pore. Scale bars = 100. complete drying out of inhabited water basins com- and 3 deutonymphs: Russia, Samara Province, monly does not take place; the bottom of habitats National natural Park “Samara Luka”, village colonized by Tiphys species retains enough mois- Koltsovo, sedge bogs, May–June 1992–1994, P.V. ture for survival of the mites, especially in forest Tuzovsky; 12 females, 5 males, 12 deutonymphs biotopes. If rains partially refill such basins, further and 42 larvae reared in laboratory (IBIW): Russia, development can take place; deutonymphs and Yaroslavl Province, Nekouz Distr., Borok, tempo- adults appear. All postlarval stages (and possibly rary ponds, sedge bogs, sedge-sphagnum bogs, also eggs) can hibernate in temporary waters, but April–July 2000–2004, P.V. Tuzovsky. there are no exact data on their physiological state Diagnosis. Female. Medial margin of coxal in that period. In the course of my investigations in plate IV approximately twice as long as medial the Yaroslavl province (Europe: Russia), males and margin of coxae III. Posteromedial and posterolat- females of T. torris were recorded in a brook in July, eral margins of coxae IV forming a large and acute August and September. Deutonymphs of T. ornatus angle. Genital plates elongate, acetabula arranged were found in water bodies that had been thawed by in an arc. Gonopore and genital plates approxi- rain. In those waters, at a water temperature of +2 mately subequal in length. Pedipalpal genu with 3 C°, mites were found immobile, but quickly began setae, external lateral seta longer than the two oth- moving when brought to laboratory conditions. er setae; base of external lateral seta situated near distal end of segment; pedipalpal tibia slender, Tiphys (Tiphys) ornatus Koch, 1836 both ventral setae inserted on very small tubercles Figs 17–53 near middle of segment; distolateral spine of tibia Material. 11 females, 1 male: Russia, Maga- 2.5–3.0 times shorter than tarsus. Claws of legs dan Province, Anadyr Distr., Markovo: sedge- I–III larger than claws of legs IV. sphagnum bog on the left bank of the Anadyr river, Male. Coxal plates IV greatly expanded, 5 km upstream of the settlement, July 1978, P.V. touching each other posteromedially; medial mar- Tuzovsky; 5 females, 1 male: Russia, Magadan gin of coxal plate IV 3–4 times as long as medial Province, Tenka Distr., village Agrobasa, June– margin of coxal plate III; posteromedial portions July 1979, P.V. Tuzovsky; 1 female: Russia, Kam- of coxal plates IV forming a deep rectangular bay chatka Province, Ust’-Kamchatsk Distr., sedge- of the genital field; genu of leg IV expanded dors- sphagnum bog on right bank of the Kamchatka oventrally with slightly convex dorsal and ventral river, 40 km upstream from Ust’-Kamchatsk city, margins, tibia IV slightly curved with single fan- 18 July 1983, P.V. Tuzovsky; 14 females, 19 males shaped dorsodistal seta and 6–8 distal long swim-

121 P. V. Tuzovsky

26, 27 26 28

29 27 29 28

Figs 26–29. Tiphys ornatus, female: 26 — chelicera, 27 — pedipalp, 28 — genu, tibia and tarsus of leg IV, 29 — claw. Scale bars: 26–27 = 100, 28 = 200, 29 = 50. ming setae; tarsus IV slender, slightly tapering regular transverse row, or setae Ai located slightly distally with numerous short, fine setae and 5–6 anterior to setae Ae; setae Ai shorter than setae Ae; relatively long, thick ones. Claws of legs III asym- excretory pore located near posterior margin of ex- metric, large claw with two nearly subequal den- cretory pore plate. Solenidion on genu of leg I lon- ticles, small claw with relatively long external ger than solenidia on tibia I; solenidion on genu II denticle and short internal one. longer than distal solenidion on tibia II, proximal Deutonymph. Dorsum usually without true solenidion on tibia II shorter than distal one. dorsal plates; medial margin of coxae IV slightly Description. Female. Dorsum with relatively longer than medial margin of coxae III. Postero- large anteromedial, small anterolateral and pos- medial angle of coxae IV moderately developed. teromedial platelets (Figs 18–19). Anteromedial Pedipalpal genu with 2 setae, external lateral seta plates and posteromedial platelets elongate, ante- longer than dorsodistal one, base of external lat- rolateral platelets transverse. Suture line between eral seta located distally to middle of segment; coxal plates III and IV slightly oblique, directed both ventral setae on tibia slightly separated and posteromedially. Medial margin of coxal plate IV situated distally to middle of segment on minute approximately twice as long as medial margin of tubercles, length of distolateral spine of tibia ap- coxae III. Posteromedial margin and posterolateral proximately in 1.5–2.0 times shorter than tarsus. margin of coxae IV forming large acute angle. Sc- Larva. Dorsal shield elongate (ratio length/ lerites bearing setae Hv, situated on soft integu- width 1.5–1.6). Setae Fch and Vi slightly longer ment between coxal plates II and III, but closely to than Fp and Oi. Dorsal shield with cell-shaped re- posterior margin coxal plate II. Three pairs of ticulation. Coxal setae C1 shorter than setae C2–C4, small unequal platelets present along lateral mar- which are approximately equal in length. Base of gins of coxal plates II–IV. setae C4 on coxal plate III located near medial end Genital plates elongate (Figs 21–23) with of suture line between coxal plates II and III. Pos- three acetabula and 10–17 thin short setae each, teromedial apodemes of coxal plates I distinctly de- acetabula arranged in an arc; all acetabula approx- veloped, apodemes of coxal plates II and III slightly imately equal in size; distance between anterior developed. Transverse muscle attachment scar pres- and medial acetabula, and distance between me- ent. Excretory pore plate wider than long, ratio dial and posterior acetabula less than diameter of length/width = 0.5–0.7; anal setae or in an almost one acetabulum. Gonopore and genital plates ap-

122 Water mites of the genus Tiphys

30, 32 31

31 32 30

Figs 30–33. Tiphys ornatus, male: 30 — ventral view, 31 — genu, tibia and tarsus of leg III, 32 — telofemur, genu, tibia and tarsus of leg IV, 33 — claws of leg III. Scale bars = 100. proximately subequal in length. Excretory pore Measurements, n=10. Idiosoma length 1200– with small anterior sclerite (Figs 24–25). 2050, length of medial margin coxa III 60–90, Basal segment of chelicera (Fig. 26) with length of medial margin coxa IV 125–165; length/ large dorsal hump, cheliceral stylet relatively width of anteromedial dorsal plates 65–70/20–25; large, pointed. length/width of genital plate 175–230/65–100; Pedipalps (Fig. 27) slender. Ventral margin of length/width of acetabula (ac. 1–3): 48–67/30–45, femur straight or slightly convex, ventral margin 48–60/36–45, 48–72/36–45; width of anterior of genu concave. Pedipalpal genu with three setae, genital sclerite 60–85, length of chelicera basal external lateral seta longer than two other setae; segment/claw 175–220/65–100; length of pedipal- base of external lateral seta situated near distal end pal segments (P-1–5) 45–65, 130–205, 90–140, of segment. Pedipalpal tibia long, slender with 195–255, 55–75; length of distolateral peg-like short distolateral peg-like seta, two ventral and seta on tibia 12–18; length of leg segments (1–6) several dorsal setae and single dorsoproximal I–Leg: 85–125, 170–245, 200–325, 285–435, spine. Both ventral setae inserted near middle tibia 245–480, 355–415; II–Leg: 100–140, 190–285, on very small tubercles, distolateral peg-like seta 200–360, 275–455, 315–475, 375–425; III–Leg: of tibia with rounded tip, 2.5–3.0 times shorter 120–150, 185–280, 135–345, 185–455, 250–500, than tarsus. Pedipalpal tarsus with single long 290–410; IV–Leg: 185–245, 170–320, 135–430, proximal solenidion, with five thin setae and four 245–615, 340–650, 405–490. thick unequal distal spines. Male. Dorsal surface and coxae of legs I–III All legs with swimming setae; two anterior similar to female. Coxae plates IV greatly expand- pairs of legs with short swimming setae, two pos- ed, fused with each other posteromedially but sep- terior pairs of legs with long swimming ones (Fig. arated anteriorly, but interspace between posterior 28). One or two swimming setae present on genu coxal groups sclerotized. Medial margin of coxal of legs I, 5–7 on tibia I, 5–9 on genu II, 7–12 on plate IV 3–4 times as long as medial margin of tibia II, 6–13 on genu III, 10–12 on tibia III, 4–12 coxal plate III. Posteromedial portions of coxal on genu IV, 8–14 on tibia IV. Claws of legs I–III plate IV forming a deep more or less rectangular are larger than claws of legs IV. Internal denticle bay nearly completely embracing genital field. of claw of legs I–III rather wide with rounded tip, Genital plates triangular; all acetabula relatively external denticle thin and pointed (Fig. 29). large, approximately subequal in size, and ar-

123 P. V. Tuzovsky

39 34

34, 36, 37 37 36 38 Figs 34–39. Tiphys ornatus, deutonymph: 34 — seta Fch, 35 — ventral view, 36 — excretory pore, 37 — pedipalp, 38 — genu, tibia and tarsus of leg IV, 39 — claw. Scale bars: 34, 36–37 = 50; 35, 38 = 100, 39 = 25. ranged in a triangle. Anterior genital sclerite small, of medial margin of coxa IV 270–345; length of with two pairs of short and fine setae and -some anteromedial dorsal plates 60–70, width 15–18; times fused with genital plates. All sclerites bearing length of genital plates 120–130, their general width ventral idiosomal setae, lying in soft integument. 215–240; length/width of genital acetabula (ac. Excretory pore with small anterior sclerite. Capitu- 1–3): 48–55/30–42, 48–60/36–45, 42–60/30–45; lum, chelicera and pedipalps as in female, but pedi- length of basal segment of the chelicera 135–170; palpal tibia with more numerous dorsal setae. length of cheliceral stylet 65–75; length of pedipal- Genu III (Fig. 31) slightly shorter than tibia pal segment (P-1–5): 40–50, 120–145, 80–100, III, tarsus III longer than tibia III; genu III with 145–185, 40–57; length of distolateral spine on 4–5 and tibia III with 8–10 long swimming setae. tibia 13–17; length of leg segments: I–Leg. 1–6: Telofemur IV (Fig. 32) considerably shorter than 85–105, 170–180, 200–230, 285–300, 245–280, genu IV. Genu IV expanded dorsoventrally with 390–400; II–Leg. 1–6: 100–125, 190–195, 200– slightly convex dorsal and ventral margins, bear- 220, 275–295, 315–335, 395–415; III–Leg. 1–6: ing numerous thin dorsal and ventral setae and 120–125, 135–180, 185–205, 250–270, 290–335; several relatively short distal sword-like setae; IV–Leg 1–6: 185–205, 170–205, 135–180, 245– dorsal setae shorter and thicker than ventral ones. 310, 340–410, 400–490. Tibia IV slightly curved, with concave dorsal and Deutonymph. Dorsum usually without true convex ventral margins, with numerous unequal dorsal plates, sometimes fragments of anteromedial setae, a single fan-shaped dorsodistal seta and 6–8 plates present. Suture line between coxal plates III long distal swimming setae. Tarsus IV slender, and IV oblique, directed posteromedially. Medial slightly tapering distally, with numerous short, margin of coxae IV slightly longer than medial mar- fine setae and 5–6 relatively long, unequal thick gin of coxae III. Posteromedial angle of coxae IV ones. Claw of legs I and II as in female. Claws of moderately developed. Genital plates separated, legs III asymmetric, large claw with nearly sub- oblique, directed anteromedially, with more or less equal clawlets, small claw with relatively long parallel anterior and posterior margins. Both aceta- dorsal clawlet and short ventral clawlet (Fig. 33). bula approximately subequal in size, distance be- Measurements, n=10. Body length 730–960, tween them less than diameter of one acetabulum. length of medial margin of coxa III 60–115, length Postgenital sclerite larger than pregenital sclerite.

124 Water mites of the genus Tiphys

40 41 Figs 40–41. Tiphys ornatus, larva: 40 — dorsal view, 41 — ventral view. Scale bar: 40–41 = 100.

46, 47 48

47 46

44 43

42–45 45 42 Figs 42–48. Tiphys ornatus, larva: 42–45 — excretory pore plate, 46 — capitulum (ventral view), 47 — chelicerae, 48 — pedipalp. Scale bars: 42–47 = 50, 48 = 100. Ventral margin of femur straight or slightly convex, shorter than tarsus. Legs II–IV with swimming set- ventral margin of genu straight or slightly concave. ae; legs II with short swimming setae, two posterior Pedipalpal tibia longer than femur, with rather long pairs of legs with long swimming ones (Fig. 38). distolateral peg-like seta, two ventral setae and 1–2 Two swimming setae present on genu II and genu dorsodistal ones. Both ventral setae separated and III, 2–4 on tibia II, 4–6 on tibia III, 1–2 on genu IV situated in distal half of tibia on very small tubercles, and 5–7 on tibia IV. Claws of legs I–III larger than distolateral peg-like seta rather large, 1.5–2.0 times claws of legs IV, in shape as in females (Fig. 39).

125 P. V. Tuzovsky

49–51

52, 53 49

Figs 49–53. Tiphys ornatus, larva: 49 — leg I, 50 — leg II, 51— leg III, 52 — claws of leg I, 53 — claws of leg III. Scale bars: 49–51 = 50, 52–53 = 25. Measurements, n=5. Body length 570–855, Larva.Dorsal shield elongate (ratio length/ length of medial margin of coxa III 42–50, length width 1.5–1.6). Setae Fch and Vi slightly longer of medial margin of coxa IV 60–65; length of gen- and thicker than Fp and Oi. Setae Oe longest and ital plates 60–65, width 37–42; diameter of genital heaviest of all dorsal setae, and setae Li shorter acetabula (ac. 1–2): 18–22, 21–25; length of basal than other dorsal setae. Dorsal shield with punc- segment of chelicera 90–120; length of cheliceral tate scale-like patterns. stylet 40–55; length of pedipalpal segments Setae C1 thin and shorter than setae C2–C4, (P-1–5): 24–30, 70–90, 40–55, 85–100, 32–40; which are approximately equal in length; C4 much length of distolateral peg-like on tibia 20–27; heavier than C1, base of setae C4 on coxal plate III length of leg segments: I–Leg. 1–6: 49–55, 55–70, located anteriorly, very close to medial end of su- 80–90, 105–130, 125–165, 145–180; II–Leg. 1–6: ture line between coxal plates II and III. Postero- 55–65, 65–75, 75–100, 110–130, 130–165, 155– medial apodemes of coxal plates I developed dis- 180; III–Leg. 1–6: 55–65, 65–80, 75–100, 95–140, tinctly, apodemes of coxal plates II and III 145–170, 145–180; IV–Leg. 1–6: 80–100, 90– relatively slightly developed. Transverse muscle 100, 95–115, 135–170, 170–205, 180–210. attachment scar well developed and located in the

126 Water mites of the genus Tiphys posteromedial portions of coxal plates III. Setae Remarks. European larvae of T. ornatus dif- Ci very long and heaviest of all ventral idiosomal fer from North American specimens attributed to setae and situated on small bases, setae Si shortest this species (characters states from Prasad and of all ventral idiosomal setae. Coxal plates I–III Cook, 1972 in parentheses) in the well elongated with a pore like structure and slightly developed dorsal shield, length/width 240–310/150–190, ra- reticulations on it. tio 1.5–1.6 (slightly elongated, length/width 216– Excretory pore plate (Figs 42–45) wider than 249/157–172, ratio 1.3–1.4), and the scale-like long, ratio length/width = 0.5–0.7. Setae Ai and Ae patterns occupying nearly the whole ventral sur- located on anterior half of plate and forming al- face of capitular base (present as a circle in the most regular transverse row, or setae Ai located middle of capitular base). slightly in front of setae Ae; Ai shorter than Ae. Ecology. Temporary ponds, sedge-sphagnum Excretory pore located near posterior margin of bogs and lakes with macrophytes; permanent wa- excretory pore plate. Capitulum (Fig. 46) small, ters (Smit and Van der Hammen 2000). rounded posteriorly, scale like patterns occupy Geographical distribution. Holarctic: Eu- nearly all ventral surface of capitular base; anteri- rope, Asia, North Africa (Algeria), North America or hypostomal setae longer and thinner than poste- (USA) (Sokolow 1940; Viets 1956; Cook 1956; rior ones. Dorsodistal setae of femur and genu of Lundblad 1968; K.O. Viets 1978). pedipalps short, thin and approximately equal in Tiphys (Tiphys) anadyrensis sp.n. length; proximal lateral genual seta very long and very heavy; tibia with 3 approximately subequal Figs 54–61 setae and large dorsodistal pointed claw; tarsus Material. Holotype male (IBIW 2303): Asia, very small with1 short solenidion, 2 long setae and Russia, Magadan Province, Anadyr Distr., Marko- 5 short unequal simple ones. vo: small lake on right bank of the Anadyr river, Number of setae on legs I–III (Figs 49–51), 13 km downstream of the settlement, 1 July 1978, except eupathidia: I–Leg. 1–5: 1, 7, 5 (s), 11 (2s), P.V. Tuzovsky. Paratype: 1 male, Russia: Magadan 14 (s, ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, Province, Anadyr Distr., Markovo: collected in ac); III–Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). For- some lake as holotype, 11 June 1981, P.V. Tu- mula of large setae on leg segments: I–Leg 1–5: 0, zovsky. 2, 1, 1, 0; II–Leg 1–5: 0, 2, 2, 3, 0; III–Leg 1–5: 0, Diagnosis. Male. Anterior coxal groups fused 2, 2, 3, 0. Solenidion on genu I longer than so- to each other posteromedially; medial margin of lenidia on tibia I, both solenidia on tibia I approxi- coxal plate IV 3.8–4.8 times as long as medial mately subequal in length; proximal solenidion on margin of coxal plate III; posteromedial portions tibia II shorter than distal one, solenidion on genu of coxal plates IV forming a deep bay almost com- II longer than distal solenidion on tibia II; solen- pletely enclosing genital field. Genu IV expanded idion on genu III shorter than solenidion on tibia dorsoventrally with distinctly convex dorsal and III. On tarsi of legs I and II, empodia bent at a right ventral margins. Tibia IV slightly curved with angle and ambulacra only slightly bent (Fig. 52). concave dorsal and convex ventral sides bearing Distal part of empodium of leg III turned onto numerous setae and a single short, thick peg-like ventral side at obtuse angle (Fig. 53). seta; long distal swimming setae absent. Claws of Measurements, n=10. Length of the dorsal legs III asymmetric; large claw with two long shield 240–310, width 150–190; length of the anal curved nearly subequal in length denticles, exter- plate 12–18, width 23–35; length of the capitulum nal denticle pointed with short dorsal spur, inter- 95–100; length of basal segment of the chelicera nal denticle with rounded tip; small claw with two 67–95, length of the chelicera stylet 16–20; length short pointed denticles, external denticle larger of medial margin of the coxae I 70–80, length of than internal one. medial margin of the coxae II+III 110–140; length Differential diagnosis. The male of T. anady� of excretory pore plate 12–18, width 23–32; length rensis sp. n. is similar to the male of T. ornatus, of the pedipalpal segments (P-1–5): 6–8, 30–32, from which it differs by the structure of the ap- 19–24, 8–10, 6–7; length of the legs segments: I– pendages. The anterior coxal group of T. anady� Leg. 1–5: 32–44, 32–36, 35–38, 40–52, 52–60; II– rensis is fused to each other posteromedially (Fig. Leg. 1–5: 38–44, 28–40, 32–42, 52–58, 72–80; 56), genu IV (Fig. 59) has distinctly convex dorsal III–Leg. 1–5:36–46, 35–42, 38–45, 54–65, 68– and ventral margins and tibia IV is without long 85. distal swimming setae. In contrast, in T. ornatus,

127 P. V. Tuzovsky

55 54 55, 56

57 56

Figs 54–57. Tiphys anadyrensis sp. n., male: 54 — seta Fch, 55 — anteromedial and anterolateral plates, 56 — ventral view, 57 — pedipalp. Scale bars = 100. the anterior coxal group is separated (Fig. 30), coxal plate III. Posteromedial portions of coxal genu IV has slightly convex dorsal and ventral plates IV forming a deep bay almost completely en- margins while tibia IV is with long distal swim- closing genital field; lateral margins of these coxal ming setae (Fig. 32). plates convex. Genital plates triangular and fused to Description. Male. Body oval, setae Fch long, each other posteromedially; all acetabula relatively whip-like (Fig. 54), other idiosomal setae filiform. large, approximately subequal in size, and arranged Integument soft, with very fine strips. Dorsum with in triangular shape. Anterior genital sclerite very relatively large anteromedial and small anterolater- small, with two pairs of short, fine setae. Genital al plates (Fig. 55), posteromedial platelets not de- plates with 2 short, fine setae each. Sclerites, bear- veloped; anteromedial plates larger than anterolat- ing setae Hv and Pe, situated on soft integument. eral ones. Coxal plates occupying most of the Excretory pore with small anterior sclerite. Capitu- ventral surface (Fig.56), coxae IV considerably lon- lum with short anchoral projection, both pairs of ger than wide, touching posteriorly but separated setae equal in length. The basal segment of chelic- anteriorly, interspace between posterior coxal era with large dorsal hump. Cheliceral stylet rela- groups sclerotized. Anterior coxal groups fused to tively large, pointed. each other posteromedially; anterior apodemes Pedipalps (Fig. 57) rather slender. Pedipalpal rather large, extending beyond medial edge of coxal trochanter with a single short dorsodistal seta, fe- plate III, posterior apodemes slightly developed. mur with 3–4 proximal and 2 dorsodistal short se- Coxal plate I with 4 lateral thin setae and 1 postero- tae. Ventral margin of femur straight, ventral mar- medial thin seta, coxal plate II with 3 lateral thin gin of genu slightly concave. Pedipalpal genu with setae, coxal plate III with 3 lateral setae and 1 me- 3 setae, external lateral seta considerably longer dial seta, coxal plate IV with 2 posteromedial setae than two other setae; base of external lateral seta and 3–4 short lateral ones. Medial margin of coxal situated distally to middle of segment. Pedipalpal plate IV 3.8–4.8 times longer than medial margin of tibia long, slender, with short distolateral spine, 2

128 Water mites of the genus Tiphys

Figs 58–61. Tiphys anadyrensis sp. n., male: 58 — genu, tibia and tarsus of leg III, 59 — leg IV, 60 — claw of leg II, 61 — claws of leg III. Scale bars: 58 = 100, 59 = 200; 60–61 = 50. ventral and numerous dorsal setae and 2 proximal genu IV in distal half without setae. Tibia IV spines. Both ventral setae inserted near the middle slightly curved with concave dorsal and convex of tibia. Distolateral spine of tibia 3.0–4.0 times ventral sides and with numerous thin setae and longer than length of tarsus. Pedipalpal tarsus with single short peg-like dorsodistal seta; long distal single long proximal solenidion, with 5 thin setae swimming setae absent. Tarsus IV slender, slightly and 4 thick unequal distal spines. tapering distally, with numerous short and fine se- Genu III (Fig. 58) shorter than tibia III, tarsus tae and with 10–11 relatively long unequal ones. III slightly shorter than tibia III; genu III with 4–5 Claws of legs I–II larger, than claws of legs IV. and tibia III with 8–10 long swimming setae. Te- Internal denticle of claws of legs I–II with more or lofemur of leg IV (Fig. 59) considerably shorter less rounded tip, external denticle pointed (Fig. than genu IV. Genu IV dorsoventrally expanded 60). Claws of legs III asymmetric (Fig. 61), large near the middle of segment comparing to telofe- claw with two long, curved and nearly subequal in mur IV, slightly tapering distally, bearing numer- length denticles; internal denticle with rounded ous dorsal and ventral thin setae, and several rela- tip; external denticle pointed, with short spur; tively short and thick setae; dorsal setae shorter small claw with two short pointed unequal denti- than ventral ones. Part of dorsodistal surface of cles, external denticle larger than internal one.

129 P. V. Tuzovsky

63 62

64 63–70

65 68 69

70 66 67 Figs 62–70. Tiphys scaurus, female: 62 — seta Fch, 63 — anteromedial and anterolateral dorsal plates, 64 — posteromedial dorsal plates, 65 — ventral view, 66–69 — genital plates, 70 — excretory pore. Scale bars = 100.

Measurements (n=2). Body length 810–850, 2000–2004, P.V. Tuzovsky; 1 male and 1 female: length of medial margin of coxa III 60–78, length The Netherlands, H. van der Hammen. of medial margin of coxa IV 285–300; length of Diagnosis. Female. Medial margin of coxal anteromedial dorsal plates 35–42, theirs width plate IV slightly longer than medial margin of 18–21; length of genital plates 95–110, theirs coxal plate III. Posteromedial margin and postero- width 95–100; length/width of genital acetabula lateral margin of coxae IV forming large obtuse (ac. 1–3): 36–42/47–50, 60–62/42–44, 48–55/30– angle. Genital plates longer than wide, more or 36; length of basal segment of the chelicera 163– less triangular with 3 acetabula and 9–16 short and 180; length of cheliceral stylet 57–65; length of fine setae each, genital opening longer than genital pedipalpal segment (P-1–5): 40–50, 114–116, 80– plates, anterior genital sclerite 2–3 times wider 82, 163–172, 40–41; length of distolateral spine than posterior one. Pedipalpal genu usually with on tibia 12–18; length of leg segments: I–Leg. 3(2) setae, external lateral seta long and located 1–6: 80–90, 160–165, 180–195, 250–255, 260– near distal end of segment; pedipalpal tibia with 295, 365–375; II–Leg. 1–6: 95–105, 145–170, short distolateral spine (3 times shorter than tar- 195–205, 260–270, 305–310, 370–380; III–Leg. sus), 2 ventral and several dorsal setae and a single 1–6: 114–122, 145–165, 105–115, 120–140, 210– dorsoproximal spine; both ventral setae located 220, 245–255; IV–Leg.1–6: 180–190, 160–165, near the middle of tibia on small tubercles. 130–135, 325–345, 375–410, 445–480. Male. Coxal plates IV greatly expanded, Ecology. Lake with macrophytes. touching each other posteromedially, medial mar- Geographical distribution. Asia, Russia: gin of coxal plate IV 2.5–2.8 times longer than Magadan Province. medial margin of coxal plate III, posteromedial margins of coxal plate IV oblique, directed poste- Tiphys (Tiphys) scaurus (Koenike, 1892) rolaterally, genital bay shallow, most of genital Figs 62–98 field lying outside genital bay. Genu IV expanded Material. 2 females: Russia, Magadan Prov- dorsoventrally, with distinctly convex dorsal mar- ince, Tenka Distr., village Agrobasa, 22 June 1979, gin and weakly convex ventral one, bearing nu- 19 females, 7 males, 3 deutonymphs and 34 larvae merous dorsal and ventral setae, and several rela- reared in laboratory (IBIW): Russia, Yaroslavl tively short sword-like setae; dorsal setae arranged Province, Nekouz Distr., Borok, temporary ponds, in two groups: proximal group with several rela- sedge bogs, sedge-sphagnum bogs, April–July tively short setae, distal group with several long

130 Water mites of the genus Tiphys

Figs 71–74. Tiphys scaurus, female: 71 — chelicera, 72 — pedipalp, 73 — genu, tibia and tarsus of leg IV, 74 — claw. Scale bars: 71–73 = 100, 74 = 50. whip-like setae and with rather numerous flat and of suture line between coxal plates II and III. wide setae, the latter ones shorter than the whip- Transverse muscle attachment scar absent. Setae like setae. Claws of legs III asymmetric, but more Ci long and thicker than all ventral idiosomal set- or less subequal in size with massive external and ae, situated on small bases; setae Si shortest. Dor- internal denticles and thin spur between them sal shield and coxal plates I–III with cell-shaped each. reticulations. Excretory pore plate more or less Deutonymph. Dorsum with well developed circular, ratio length/width = 0.8–1.1, setae Ai anteromedial plates only; medial margin of coxae shorter than Ae, excretory pore located in centre of IV slightly longer than medial margin of coxae III, excretory pore plate. Solenidion on genu I slightly posteromedial angle of coxae IV moderately de- longer than solenidia on tibia I, latter ones sub- veloped, with short apodeme. Genital plates with equal in length; proximal solenidion on tibia II more or less parallel anterior and posterior mar- twice shorter than distal one, solenidion on genu II gins, postgenital sclerite considerably larger than shorter than distal solenidion on tibia II. pregenital sclerite, excretory pore unsclerotized. Description. Female. Body oval; setae Fch Pedipalpal femur with 3 dorsodistal setae; genu long, whip-like (Fig. 62), other idiosomal setae with 2 setae, external lateral seta 3–4 times longer filiform. Integument soft with very fine strips. than dorsodistal seta, base of external lateral seta Dorsum with relatively large anteromedial plates, situated distally to middle of segment; tibia with small anterolateral (Fig. 63) and posteromedial rather large distolateral spine (0.45–0.70 of tarsus platelets (Fig. 64). Coxal plate I usually with 4 lat- length), ventral setae situated close to each other, eral thin setae and 1 posteromedial thin seta, coxal on small tubercles distally to middle of segment. plate II with 2–3 lateral thin setae (Fig. 65). Apo- Larva. Dorsal shield elongate (ratio length/ deme of coxal plate I moderately developed and width 1.4–1.6). Setae Fch and Vi slightly longer usually not extending beyond medial margin of than Fp and Oi. Seta C1 shorter than setae C2–C4, coxal plate III. Suture line between coxal plates III which approximately equal in length; bases of se- and IV slightly oblique, directed posteromedially. tae C4 on coxal plate III located near medial end Medial margin of coxal plate IV longer than me-

131 P. V. Tuzovsky dial margin of coxal plate III. Posteromedial mar- length of medial margin of coxa IV 70–85; length gin and posterolateral margin of coxae IV forming of anteromedial dorsal plates 48–60, width 15–24; large obtuse angle with distinct apodeme. Scler- length of genital plates 130–140, width 50–80; ites, bearing setae Hv, Pe and other idiosomal ven- length/width of genital acetabula (ac. 1–3): 36– tral setae, lying free in soft integument. Four pairs 42/24–30, 36–42/24–30, 36–45/ 25–30; width of of small unequal platelets present along lateral anterior genital sclerite 95–125; length of basal margins of coxal plates II–IV. segment of the chelicera 130–140, length of cheli- Genital plates more or less triangular (Figs ceral stylet 55–60; length of pedipalpal segment 66–69) with 3 acetabula and 9–16 short, fine setae (P-1–5): 30–40, 105–125, 70–82, 120–140, 55– each. All acetabula approximately subequal in 58; length of distolateral spine on tibia 18; length size, second and third acetabula closely together of leg segments: I–Leg. 1–6: 65–82, 95–115, 130– and form more or less right transverse row in pos- 145, 175–195, 195–205, 200–220; II–Leg. 1–6: terior portion of genital plate; distance between 70–82, 105–115, 135–150, 175–195, 195–215, first acetabulum and third acetabulum larger than 200–230; III–Leg. 1–6: 80–83, 110–125, 130–140, distance between second and third acetabula. An- 170–190, 210–220, 200–230; IV–Leg. 1–6: 120– terior ends of genital plates usually more or less 130, 130–150, 170–190, 235–255, 260–280, 250– pointed, but sometimes anterior end rounded (Fig. 260. 68). Genital opening longer than genital plates, Male. Dorsal surface, coxae I–III similar to anterior genital sclerite 2–3 times wider than pos- those in female. Anterior apodemes rather large, terior one. Excretory pore with small anterior sc- extending posteriorly to medial edge of coxal plate lerite (Fig. 70). Capitulum with short anchoral III; posterior apodemes little developed (Fig. 75). projection, both pairs of setae equal in length. The Coxae plates IV greatly expanded, touching each basal segment of chelicera (Fig. 71) with large other posteromedially, with 2 short and fine setae dorsal hump, cheliceral stylet relatively large, on each side of this area, situated rather close to pointed. each other; interspace between posterior coxal Pedipalp rather stout (Fig. 72). Pedipalpal tro- groups sclerotized. Medial margin of coxal plate chanter with single short dorsodistal seta, femur IV 2.5–3.0 times longer than medial margin of with 3–4 short proximal setae and 2 dorsodistal coxal plate III. Suture line between coxal plates III ones. Ventral margin of femur straight, ventral and IV nearly transverse. Posteromedial margins margin of genu concave. Pedipalpal genu usually of coxal plate IV oblique, directed posterolateral- with 3(2) setae (proximal dorsal seta sometimes ly, and forming a shallow bay partially surround- absent), external lateral seta longer than two other ing external genital organ; lateral margins of these setae; base of external lateral seta situated near dis- coxal plates convex. Genital plates triangular and tal end of segment. Pedipalpal tibia slender with fused to each other medially, all acetabula rela- short distolateral peg-like seta, 2 ventral setae and tively large, approximately subequal in size, and several dorsal ones; both ventral setae inserted on arranged in triangle. Anterior genital sclerite small, small tubercles distally, distolateral peg-like seta curved, with 2 pairs of short and fine setae, some- approximately 3 times longer than tarsus. Pedipal- times it fused with genital plates. Genital plates pal tarsus with long proximal solenidion, 5 thin se- with 2–3 short and fine setae each. Sclerites bear- tae and 4 short and thick unequal distal setae. ing setae Hv close to posterior margin of coxae II; All legs with swimming setae; two anterior other sclerites bearing ventral idiosomal setae ly- pairs of legs with short swimming setae, two pos- ing free in soft integument. Excretory pore with terior pairs of legs with long swimming ones (Fig. small anterior sclerite. Capitulum, chelicera and 73). One swimming seta present on genu I, 2 on pedipalps similar to female, but pedipalpal tibia tibia I and genu II, 2–3 on genu III and tibia II, 3–4 with more numerous dorsal setae. on genu III and IV, 5–8 on tibia III and IV. Claws Genu III (Fig. 76) shorter than tibia III, and of legs I–III larger than claws of legs IV; both den- tarsus III considerably longer than tibia III; genu ticles pointed and approximately subequal in III with 5–6 rather long distal setae, tibia III with length, but internal denticle slightly wider than ex- 6–9 long swimming setae, and tarsus III with nu- ternal one (Fig. 74). Claw lamella moderately de- merous short thin setae. Telofemur of leg IV (Fig. veloped with convex ventral margin. 77) short, expanded distally; genu IV large, nar- Measurements (n=10). Length of body 800– rowed proximally, expanded dorsoventrally with 940, length of medial margin of coxa III 54–60, convex dorsal and ventral margin, bearing numer-

132 Water mites of the genus Tiphys

75–77

75 78 77 Figs 75–78. Tiphys scaurus, male: 75 — ventral view, 76 — genu, tibia and tarsus of leg III, 77 — telofemur, genu, tibia and tarsus of leg IV, 78 — claws of leg III. Scale bars: 75–77 = 100, 78 = 50. ous dorsal and ventral setae and several relatively 1–6: 55–65, 90–100, 105–115, 140–155, 150–165, short distal ones; dorsal setae arranged in two 210–220; II–Leg. 1–6: 65–75, 95–105, 105–125, groups: proximal group with several relatively 135–155, 150–165, 200–230; III–Leg. 1–6: 73– short setae, distal group with several long whip- 90, 90–100, 70–90, 85–100, 115–130, 170–190; like setae and with a few sword-like setae; latter IV–Leg.1–6: 120–130, 95–115, 85–95, 160–180, setae shorter than whip-like ones. Tibia IV slightly 220–245, 220–230. curved with concave dorsal and convex ventral Deutonymph. Similar to female, but smaller. sides with numerous unequal setae and a single Dorsum with well developed anteromedial plates short, thick dorsodistal peg-like seta. Tarsus of leg only (Fig. 79). Setae Fch long, thin (Fig. 80). Cox- IV slender, slightly tapering distally with numer- al plate I usually with 3 lateral thin setae and 1 ous short and fine setae, and 3–6 unequal long posteromedial thin seta, coxal plate II with 1 lat- swimming setae. Claw of legs I and II similar to eral thin setae, coxal plate III with 1 medial seta those in female. Claws of legs III (Fig. 78) asym- and 1–2 lateral thin setae, coxal plate IV with 2 metric; large claw with thick external denticle and medial seta and 1 lateral seta (Fig. 81). Apodeme thin internal one; small claw with thin pointed un- of coxal plate I short and not extending beyond equal denticles. medial margin of coxal plate III. Suture line be- Measurements (n=7). Body length 570–615, tween coxal plates III and IV oblique, directed length of medial margin of coxa III 70–75, length posteromedially. Medial margin of coxae IV of medial margin of coxa IV 180–200; length of slightly longer than medial margin of coxae III. anteromedial dorsal plates 36–42, theirs width Posteromedial angle of coxae IV moderately de- 15–24, length of genital plates 65–72, theirs gen- veloped with short apodeme. Sclerite bearing seta eral width 160–180; length/width of genital aceta- Hv located near posterior margin of coxal plate II, bula (ac. 1–3): 36–42/24–30, 42–45/30–36, 30– other ventral idiosomal setae situated on soft in- 36/24–36; length of basal segment of the chelicera tegument. Genital plates separated, oblique, di- 120–125; length of cheliceral stylet 48–57; length rected anteromedially, with more or less parallel of pedipalpal segment (P-1–5): 30–33, 90–105, anterior and posterior margins, bearing 2 acetabula 45–65, 45–50; length of distolateral peg-like spine and 3 short and fine setae each. Both acetabula ap- on tibia 15–18; length of leg segments: I–Leg. proximately subequal in size, distance between

133 P. V. Tuzovsky

80 79 79–81, 83

81 82 83

Figs 79–83. Tiphys scaurus, deutonymph: 79 — anterior dorsal plates, 80 — seta Fch, 81 — ventral view, 82— pedipalp, 83 — genu, tibia and tarsus of leg IV. Scale bars: 79–81, 83 = 100, 82 = 50. them usually less than diameter of one acetabulum of medial margin of coxa IV 30–35; length of an- on each plate. Postgenital sclerite considerably teromedial dorsal plates 30–35, width 12–14, larger than pregenital sclerite. Excretory pore non- length of genital plates 57–65, width 32–35; diam- sclerotized. eter of genital acetabula (ac. 1–2): 16–19, 16–18; Pedipalpal trochanter (Fig. 82) without setae, length of basal segment of chelicera 97–100, femur with 3 dorsodistal setae. Ventral margin of length of cheliceral stylet 30–35; length of pedi- femur straight or slightly convex, ventral margin palpal segment (P-1–5): 23–25, 55–65, 30–35, of genu concave. Pedipalpal genu with 2 setae, ex- 55–60, 25–30; length of distolateral spine on tibia ternal lateral seta 3–4 times longer than dorsodis- 15–18; length of leg segments: I–Leg. 1–6: 40–45, tal seta; base of external lateral seta inserted dis- 55–60, 58–65, 90–95, 100–105, 120–125; II–Leg. tally to middle of segment. Pedipalpal tibia with 1–6: 45–50, 55–60, 60–65, 95–105, 110–115, rather long distolateral peg-like spine, 2 ventral 130–135; III–Leg. 1–6: 50–55, 60–65, 60–65, setae and 1–2 dorsodistal ones. Both ventral setae 100–105, 115–125, 130–135; IV–Leg. 1–6: 60– situated closely to each other and inserted on very 65, 65–70, 75–80, 105–115, 125–135, 145–150. small tubercles in distal half of tibia. Length of Larva. Body small and oval; dorsal shield distolateral spine of tibia 0.45–0.70 of tarsus truncated anteriorly, rounded posteriorly, slightly length. Pedipalpal tarsus with single long proxi- indented anterolaterally near anterior eyes, with mal solenidion, 3 thin setae and 4 thick distal convex lateral margins (Fig. 84). Dorsal shield spines. elongate (ratio length/width 1.4–1.6), covering al- All legs with swimming setae; two anterior most all dorsum in unengorged larva and bearing pairs of legs with short swimming setae, two pos- 4 pairs of setae (Fch, Fp, Vi, Oi). Setae Fch and Vi terior pairs of legs with long swimming setae (Fig. slightly longer than Fp and Oi. Setae Oe, Hi, He, 83). One swimming seta present on tibia I, 2 on Sci, Sce and Li located on the soft and wrinkled tibia II and genu II–IV, 3–4 on tibia III, 4–5 on membrane. Setae Oe longest and thickest of all tibia IV. Claws of legs I–III larger, than claws of dorsal setae, setae Li shorter than other dorsal set- legs IV. Internal denticle of claw of legs I–III rath- ae. Lateral sides of coxae II+III expanding onto er wide with rounded tip, external denticle thin dorsal side and lying beside setae Oe and other and pointed. hysterosomal setae located on interscutal mem- Measurements (n=3). Body length 80–540, brane. Dorsal shield with cell-shaped reticula- length of medial margin of coxa III 24–30, length tions.

134 Water mites of the genus Tiphys

84 85 Figs 84–85. Tiphys scaurus, larva: 84 — dorsal view, 85 — ventral view. Scale bar: 84–85 = 100. 91 92

91, 92 93

89 86 87 90 88 85–90 Figs 86–93. Tiphys scaurus, larva: 86–90 — excretory pore plate, 91 — capitulum (ventral view), 92 — chelicerae, 93 — pedipalp. Scale bars: 86–90 = 50; 91, 93 = 30, 92 = 35.

Coxal plates I–II relatively small; anterior which are fused, suture line between them incom- coxal plates separated from coxal plates II and III, plete and developed only in their lateral parts (Fig.

135 P. V. Tuzovsky

97, 98 95

94–96

Figs 94–98. Tiphys scaurus, larva: 94 — leg I, 95 — leg II, 96 — leg III, 97 — claws of leg I, 98 — claws of leg III. Scale bars: 94–96 = 50, 97–98 = 25. 85). Anterior margin of coxal plates I concave and Ai located in front of setae Ae, latter ones longer considerably shorter than posterior margin. Setae than setae Ai. Excretory pore open near midlength C1 shorter than setae C2–C4, which approximate- of excretory pore plate. ly equal in length. Bases of setae C4 on coxal plate Capitulum elongate (Fig. 91), capitular base III located near medial end of suture line between relatively narrow with rounded or straight poste- coxal plates II and III. Posteromedial apodemes of rior edge, anterior hypostomal setae longer and coxal plates I well developed, apodemes of coxal thinner than posterior ones. Basal segments of plates II absent, and apodemes of coxal plates III chelicerae (Fig. 92) fused by their medial parts poorly developed. Transverse muscle attachment and tapering proximally; suture line well devel- scar not developed in the posteromedial angles of oped along their whole length. Cheliceral stylet coxal plates III. Setae Ci, Si, Se, Le, Pi, Pe situated small and crescent in shape. on soft interscutal membrane in posterior part of Pedipalp trochanter (Fig. 93) short and without body. Setae Ci longer and thicker than all ventral setae; dorsodistal setae of femur and genu of pedi- idiosomal setae, and situated on small bases; setae palps short, approximately equal in size, proximal Si shortest and thin. Coxal plates I–III with devel- lateral genual seta thick and very long; tibia with 3 oped cell-shaped reticulations. Urstigma occupy- short approximately subequal setae, and large dor- ing lateral position on border between coxae I and sodistal claw. Pedipalpal tarsus with 1 short solen- II and provided cup. idion, 2 long setae and 5 unequal simple ones. Shape of excretory pore plate highly variable The general number of setae on legs I–III (Figs 86–90), ratio length/width = 0.8–1.1. Setae (Figs 94–96), except eupathidia, is as follows: I–

136 Water mites of the genus Tiphys

Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); II–Leg. Geographical distribution. Europe, Asia, 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); III–Leg. 1–5: 1, USA(?) (Sokolow 1940; Viets 1956; Cook 1956; 6, 5(s), 10 (s), 11 (ac). Formula of large setae on Lundblad 1968; K.O. Viets 1978). leg segments as follows: I–Leg 1–5: 0, 3, 1, 1, 0; Tiphys (Tiphys) scaurellus Tuzovskij, 1983 II–Leg 1–5: 0, 4, 2, 3, 0; III–Leg 1–6: 0, 3, 2, 3, 0. Solenidia on genu, tibia and tarsus of leg I approx- Figs 99–135 imately subequal in length; proximal solenidion Material. Holotype male (IBIW 2567): Asia, on tibia II twice shorter than distal one, solenidion Russia, Magadan Province, Tenka Distr., village on genu II shorter than distal solenidion on tibia II; Agrobasa, sedge-sphagnum bog near village, 8 solenidion on genu of leg III shorter than solenid- July 1979, P.V. Tuzovsky. Paratypes: 5 females, 3 ion on tibia III. Central claw and lateral claws of males, 1 deutonymph and 5 larvae reared in labo- all legs almost subequal in size; on tarsi of legs I ratory (IBIW): Russia, Magadan Province, Tenka and II central claw bending at right angle and am- Distr., village Agrobasa, sedge-sphagnum bogs bulacra only slightly bent (Fig. 97); central claw June–July 1979, P.V. Tuzovsky. Additional mate- of tarsus III turned onto ventral side at obtuse an- rial: 13 females, 12 males, 4 deutonymphs and 29 gle (Fig. 98). larvae reared in laboratory (IBIW): Asia, Russia: Measurements (n=10). Length of dorsal shield Magadan Province, Anadyr Distr., Markovo: 260–280, its width 170–180; length of excretory sedge-sphagnum bog on the left bank of the An- pore plate 30–36, width 30–36; length of capitu- adyr river, 5 km upstream of the settlement, July lum 70–75; length of basal segment of chelicera 1978, P.V. Tuzovsky; 3 females, 4 males and 4 60–68, length of chelicera stylet 12–14; length of deutonymphs: Kamchatka Province, Ust’-Kam- medial margin of coxae I 70–80, length of medial chatsk Distr., sedge-sphagnum bog on right bank margin of the coxae II+III 120–130; length of pe- of the Kamchatka river, 40 km upstream from dipalpal segments (P-1–5): 6–8, 26–30, 20–22, Ust’-Kamchatsk city, 18 July 1983, P.V. Tuzovsky; 7–8, 6–7; length of legs segments: I–Leg. 1–5: 1 female and 1 male, Europe, Russia: Arkhangelsk 32–36, 32–34, 30–32, 42–44, 60–64; II–Leg. 1–5: Province, Vaigach island, peat-bog, 13 July 2004, 34–36, 32–36, 30–32, 50–52, 72–76; III–Leg. 1–5: O.L. Makarova. 35–40, 36–40, 38–40, 54–58, 73–78. Diagnosis. Female. Medial margin of coxal Remarks. The morphology of the European plate IV 1.3–1.6 times as long as medial margin of larva of T. scaurus differs from larva of this spe- coxal plate III, posteromedial margin and postero- cies described from Canada in the following char- lateral margin of coxae IV forming obtuse angle. acters (characters states of the Canadian larva T. Sclerites bearing setae Hv fused with posterior scaurus are indicated in parentheses, data after margin of coxal plate II; sclerites bearing other id- Smith 1976): transverse muscle attachment scar is iosomal setae lying free in soft integument. Geni- absent (present); base of seta C4 on coxal plate III tal plates more or less triangular with 3 acetabula and the medial end of suture line between coxal and 8–10 short and fine setae each; all acetabula plates II and III are separated (close to each other); small, approximately subequal in size, and usually length of dorsal plate 260–280 (227–243), width arranged in an arc. Pedipalp slender, genu with 2 of dorsal plate 170–180 (147–156), length of ex- setae, external lateral seta considerably longer cretory pore plate 30–36 (27–30), width of anal than dorsodistal seta, base of external lateral seta plate 30–36 (27–32). situated near middle of segment. Pedipalpal tibia Unfortunately, Smith (1976) did not give with short distolateral spine (3–4 times shorter characteristics of the T. scaurus female from North than tarsus), both ventral setae situated on small Americ and cited the data of Cook (1956); the lat- tubercles distally to middle of segment. ter author gave only brief characteristics of T. Male. Coxae plates IV greatly expanded, scaurus female received from The Netherlands. It touching each other posteromedially; medial mar- is possible, that North American specimens should gin of coxal plate IV 3–4 times longer than medial be treated as a subspecies of T. scaurus or even a margin of coxal plate III; suture line between cox- distinct species. al plates III and IV transverse. Posteromedial mar- Ecology. Temporary ponds, sedge-sphagnum gins of coxal plate IV oblique, directed posterolat- bogs, streams with macrophytes which are subject erally, and forming a deep bay almost completely to periodic drying, but occurs also in permanent enclosing genital field. Sclerites bearing setae Hv waters. fused with posterior margin of coxae II; other sc-

137 P. V. Tuzovsky

99 99–101 100

101

104 105

103–105

102 103 106 107 105–107 Figs 99–107. Tiphys scaurellus, female: 99 — seta Fch, 100 — anteromedial and anterolateral dorsal plates, 101 — posteromedial dorsal plates, 102 — ventral view,103–105 — genital plates, 106–107 — excretory pore. Scale bars: 99–101 =50, 102–107 = 100. lerites bearing ventral idiosomal setae lying free in tibia with rather a long distolateral seta (0.45–0.55 soft integument. Genu IV expanded dorsoventral- of tarsal length), both ventral setae situated in dis- ly with almost straight dorsal margin and convex tal half tibia on very small tubercles. ventral one, dorsal setae arranged in two groups: Larva. Body small and nearly circular in proximal group with several relatively short setae, shape; dorsal shield convex and slightly elongate and distal group with numerous long, thick setae. (ratio length/width 1.15–1.20), setae Fch and Vi Claws of legs III asymmetrical, large claw with slightly longer and thicker than Fp and Oi. Setae two long approximately subequal denticles; small Oe longest and thickest of all dorsal setae, setae Li claw with long external denticle and short internal shorter than other dorsal setae. Setae C1 and C4 one, external denticle with short, thin ventral considerably shorter than setae C2–C3, which ap- spur. proximately equal in length. Base of setae C4 on Deutonymph. Dorsum with well developed coxal plate III and medial end of suture line be- anteromedial plates only, setae Fch relatively short tween coxal plates II and III well separated. Dor- and thin. Medial margin of coxae IV 1.0–1.2 times sal shield and coxal plates I–III with reticulations. longer than medial margin of coxae III. Postero- Excretory pore plate oval or nearly circular (ratio medial angle of coxae IV moderately developed. length/width = 0.9–1.1), excretory pore placed be- Sclerites bearing setae Hv fused with posterior tween rows of setae Ai and Ae in anterior half of margins of coxae II, other ventral idiosomal setae anal plate. Solenidia on genua and tibiae I–III rela- lying free in soft integument. Genital plates sepa- tively short; solenidion on genu I not longer than rated, slightly oblique, directed anteromedially, distal solenidion on tibia I; proximal solenidion on both acetabula approximately subequal in size, tibia I and II slightly shorter than distal one, solen- distance between them usually equal to diameter idion on genu II slightly shorter than distal solen- of acetabulum on each plate. Postgenital sclerite idion on tibia II. considerably larger than pregenital sclerite. Pedi- Description. Female. Body oval; setae Fch palpal femur with 3 dorsodistal setae; genu with 2 long, whip-like (Fig. 99), other idiosomal setae setae, external lateral seta approximately twice as filiform. Dorsum with relatively large anterome- long as dorsodistal seta; base of external lateral dial, small anterolateral, and small posteromedial seta located near middle of segment. Pedipalpal platelets (Figs 100–101). Anteromedial plates rel-

138 Water mites of the genus Tiphys

111 108

110

109

Figs 108–111. Tiphys scaurellus, female: 108 — chelicera, 109 — pedipalp, 110 — genu, tibia and tarsus of leg IV, 111 — claw. Scale bars: 108–110 = 100, 111= 50. atively large and elongate, anteromedial and pos- proximately subequal in size, and usually arranged teromedial platelets oval or almost circular; some- in an arc or obtuse triangle; acetabula relatively times anterolateral platelets not developed. Coxal small and occupy less than half surface of genital plate I usually with 4 lateral thin setae and 1 pos- plate. Genital opening longer than genital plates, teromedial thin seta, coxal plate II with 2–3 lateral anterior genital sclerite considerably wider than thin setae (Fig. 102). Apodeme beneath postero- posterior one. Excretory pore with very small an- medial edge of coxal plate I moderately devel- terior sclerite (Figs 106–107). oped, not reaching or slightly extending beyond Capitulum with short anchoral projection, anterior edge of coxal plate III. Suture line be- both pairs of setae equal in length. Basal segment tween coxal plates III and IV complete and oblique, of chelicera (Fig. 108) large, with strong convex directed posteromedially. Medial margin of coxal dorsal margin and nearly straight ventral one. Che- plate IV 1.3–1.6 times longer than medial margin liceral stylet moderately developed, pointed. of coxal plate III. Posteromedial margin and pos- Pedipalps (Fig. 109) rather slender. Pedipal- terolateral margin of coxae IV forming right or pal trochanter with single short dorsodistal seta, obtuse angle. Sclerites bearing setae Hv close to or femur with 3–4 proximal and 2 short dorsodistal fused with posterior margin of coxal plate II; sc- setae. Ventral margin of femur straight, ventral lerites, bearing other idiosomal setae, lying free in margin of genu concave. Pedipalpal genu with 2 soft integument. setae, external lateral seta considerably longer Genital plates elongate and more or less trian- than dorsodistal seta; base of external lateral seta gular (Figs 103–105), with 3 acetabula and 8–10 situated distally to middle of segment. Pedipalpal short and fine setae each. All acetabula small, ap- tibia relatively thin, slender, with short distolateral

139 P. V. Tuzovsky

115, 116

112

115

117 114

113 116

Figs 112–117. Tiphys scaurellus, male: 112 — ventral view, 113 — genital field, 114 — genu, tibia and tarsus of leg III, 115 — genu of leg IV, 116 — tibia and tarsus of leg IV, 117 — claws of leg III. Scale bars: 112–114 = 100, 115–116 = 250, 117 =50. peg-like spine, 2 ventral and several fine dorsal se- length of cheliceral stylet 65–75; length of pedi- tae and 1 spine proximally to middle of segment. palpal segment (P-1–5): 40–50, 105–140, 80–100, Both ventral setae inserted distally to middle of 120–170, 40–50; length of distolateral spine on tibia on small tubercles. Distolateral spine of tibia tibia 12–16; length of leg segments: I–Leg. 1–6: 3–4 times shorter than tarsus. Pedipalpal tarsus 70–100, 120–130, 145–195, 200–270, 220–295, with long proximal solenidion, 5 thin setae and 4 220–305; II–Leg. 1–6: 80–105, 110–165, 150– thick unequal distal spines. 190, 200–260, 220–280, 225–280; III–Leg. 1–6: All legs with swimming setae; two anterior 80–115, 115–160, 130–175, 175–230, 220–295, pairs of legs with short swimming setae, two pos- 220–285; IV–Leg. 1–6: 120–165, 145–190, 160– terior pairs of legs with long swimming ones (Fig. 215, 240–310, 275–350, 250–325. 110). One swimming seta present on genu I, 2–3 Male. Dorsal surface, coxae of legs I–III sim- on tibia I and genu II, 2–4 on genu III, 3–5 on ilar to those in female. Anterior apodemes rather tibia II, 5–7 on genu IV and tibia III and 5–8 on large, extending beyond anteromedial edge of tibia IV. Claws of legs I–III larger than claws of coxal plate III; posterior apodemes slightly devel- legs IV; both denticles subequal in length (Fig. oped (Fig. 112). Coxal plates IV greatly expanded, 111), internal denticle wide with rounded tip, ex- touching each other along their whole medial mar- ternal denticle thin and pointed. Claw lamella gins, with 2 posteromedial fine setae close togeth- moderately developed, with convex ventral mar- er on each side. Medial margin of coxal plate IV gin. 3–4 times longer than medial margin of coxal plate Measurements (n=10). Body length 700– III. Suture line between coxal plates III and IV 1305, length of medial margin of coxa III 55–75, transverse. Posteromedial margins of coxal plate length of medial margin of coxa IV 80–110; length IV concave and forming deep bay almost com- of anteromedial dorsal plates 45–60, width 20–27, pletely encompassing genital organ; lateral mar- length of genital plates 95–140, width 55–75; di- gins of these coxal plates convex. Genital plates ameter of genital acetabula (ac. 1–3): 24–27, 24– triangular, all acetabula approximately subequal 27, 25–30; width of anterior genital sclerite 115– in size, and occupy half or lesser surface of plates 155; length of basal segment of chelicera 140–170, (Fig. 113). Anterior genital sclerite very small and

140 Water mites of the genus Tiphys

119 120 123 118 119–122

122

121

121, 123

Figs 118–123. Tiphys scaurellus, deutonymph: 118 — anterior dorsal plates, 119 — seta Fch, 120 — ventral view, 121— pedipalp, 122 — genu, tibia and tarsus of leg IV, 123 — claw. Scale bars: 118–120, 122 = 100; 121, 123 = 50. curved, with two pairs of short and fine setae; gen- length of medial margin of coxa IV 250–305; ital plates with 2 short and fine setae each. Scler- length of anteromedial dorsal plates 40–45, theirs ites bearing setae Pe not fused with posteromedial width 12–18, length of genital plates 90–100, their margin of coxae IV; sclerites bearing setae Hv general width 210–220; length/width of genital close to posterior margin of coxae II; other scler- acetabula (ac. 1–3): 48–55/30–40, 42–48/30–35, ites bearing ventral idiosomal setae lying free in 48–60/35–40; length of basal segment of the che- soft integument. Excretory pore with small ante- licera 130–145; length of cheliceral stylet 55–58; rior sclerite. Capitulum, chelicerae and pedipalps length of pedipalpal segment (P-1–5): 35–50, similar as in female, but pedipalpal tibia with more 100–122, 55–80, 120–140, 40–42; length of disto- numerous dorsal setae. lateral spine on tibia 12–16; length of leg seg- Genu III (Fig. 114) shorter than tibia III, tar- ments: I–Leg. 1–6: 70–90, 120–140, 135–165, sus III slightly tapering distally, tibia and tarsus 185–230, 220–255, 240–285; II–Leg. 1–6: 80– nearly subequal in length; genu III with 4–5 rela- 100, 110–145, 130–155, 180–205, 180–230, 220– tively short swimming setae, tibia III with 10–12 245; III–Leg. 1–6: 95–115, 120–155, 105–125, long swimming ones. Genu IV (Fig. 115) expand- 150–180, 185–205; IV–Leg. 1–6: 135–165, 120– ed distally, with widely rounded distal end, with 155, 90–115, 230–270, 260–310, 285–310. numerous dorsodistal and ventral setae and a few Deutonymph. Similar to female, but smaller. short setae; dorsodistal setae longer than ventral Dorsum with anteromedial plates only (Fig. 118), ones. Tibia IV slightly curved, with concave dor- anteromedial and posteromedial platelets rudi- sal margin and convex ventral one, with numerous mentary or absent. Setae Fch relatively short (Fig. unequal dorsal and ventral setae and a single peg- 119). Coxal plate I usually with 3 lateral thin setae like dorsodistal seta, and 5–7 long swimming setae and 1 posteromedial thin seta, coxal plate II with 1 (Fig. 116). Tarsus IV slender, slightly tapering dis- lateral thin setae, coxal plate III with 1 medial seta tally, with numerous short and fine setae and 7–9 and 1–2 lateral thin setae, coxal plate IV with 2 long unequal swimming setae. medial seta and 1 lateral seta (Fig. 120). Apodeme Claw of legs I and II as in female. Claws of of coxal plate I moderately developed, not reach- legs III (Fig. 117) asymmetrical, large claw with ing or slightly extending beyond anteromedial two approximately subequal denticles, small claw margin of coxal plate III. Suture line between cox- with long external denticle and short internal one. al plates III and IV slightly oblique, directed pos- Measurements (n=10). Length of body 650– teromedially. Medial margin of coxae IV 1.0–1.2 815, length of medial margin of coxa III 65–75, times longer than medial margin of coxae III. Pos-

141 P. V. Tuzovsky

124 125

Figs 124–125. Tiphys scaurellus, larva: 124 — dorsal view, 125 — ventral view. Scale bar: 124–125 = 100. teromedial angle of coxae IV moderately devel- larger than claws of legs IV. Internal denticle of oped, obtuse, with short apodeme. Sclerites bear- claw of legs I–III rather wide with rounded tip, ing setae Hv close together or fused with posterior external denticle thin and pointed (Fig. 123). margins of coxae II, other ventral idiosomal setae Measurements, n=9. Body length 400–530, lying free in soft integument. Genital plates sepa- length of medial margin of coxa III 30–35, length rated, oblique, directed anteromedially, bearing 2 of medial margin of coxa IV 35–45; length of an- acetabula and 3 short and fine setae each. Aceta- teromedial dorsal plates 30–35, width 10–12, bula approximately subequal in size, distance be- length of genital plates 60–65, width 35–38; diam- tween them usually equal to one diameter of ac- eter of genital acetabula (ac. 1–2): 21–24, 18–22; etabulum on each plate. Postgenital sclerite length of basal segment of the chelicera 92–105; considerably larger than pregenital one. Excretory length of cheliceral stylet 340–45; length of pedi- pore with small anterior sclerite. palpal segment (P-1–5): 28–30, 65–70, 45–50, Pedipalpal trochanter (Fig. 121) without seta, 75–80, 28–32, length of distolateral spine on tibia femur with 3 dorsodistal setae. Ventral margin of 15–17; length of leg segments: I–Leg. 1–6: 48– femur straight or slightly convex, ventral margin 50,65–70, 65–70, 100–105, 110–115, 128–135; of genu slightly concave. Pedipalpal genu with 2 II–Leg. 1–6: 48–52, 68–72, 68–73, 100–105, 115– setae, external lateral seta approximately twice as 120, 140–145; III–Leg. 1–6: 55–58, 70–75, 68–73, long as dorsodistal seta; base of external lateral 100–105, 120–130, 130–140; IV–Leg.1–6: 80–84, seta inserted near middle of segment. Pedipalpal 80–85, 88–92, 130–135, 145–155, 155–162. tibia slightly longer than femur, with rather long Larva. Body small and nearly circular in distolateral spine, 2 ventral and 1–2 dorsodistal shape. Dorsal shield with straight or convex ante- setae. Both ventral setae situated in distal half of rior margin, rounded posteriorly, very convex lat- tibia on very small tubercles. Length of distolateral margins, it anterolateral margins are weakly eral spine of tibia 0.40–0.55 of length of tarsus. concave (Fig. 124). Dorsal shield convex and Pedipalpal tarsus with long proximal solenidion, 3 slightly elongate (ratio length/width 1.15–1.20), thin setae and 4 thick distal spines. covering almost all dorsum in unengorged larva All legs with swimming setae; two anterior and bearing 4 pairs of setae (Fch, Fp, Vi, Oi). Se- pairs of legs with short swimming setae, two pos- tae Fch and Vi slightly longer and thicker than Fp terior pairs of legs with long swimming ones (Fig. and Oi. Setae Oe, Hi, He, Sci, Sce and Li located 122). One swimming seta present on genu leg II on the soft wrinkled membrane. Setae Oe longest and tibia of leg I, 2 on genu III–IV and tibia II, 3–4 and thickest of all dorsal setae, setae Li shortest. on tibia III, 4–6 on tibia IV. Claws of legs I–III Lateral sides of coxae II+III passing onto dorsal

142 Water mites of the genus Tiphys

129 128

130

126 127 Figs 126–130. Tiphys scaurellus, larva: 126–127 — excretory pore plate, 128 — capitulum (ventral view), 129 — chelicerae, 130 — pedipalp. Scale bars: 126–130 = 50. side and almost extending to setae Oi and other Excretory pore plate (Figs 126–127) oval or hysterosomal setae located on interscutal mem- nearly circular, ratio length/width = 0.9–1.1. Ex- brane. Dorsal shield with reticulations. cretory pore in anterior half of excretory pore Coxal plates I–II relatively small; anterior plate, setae Ai shorter than Ae and placed close to coxal plates separated from coxal plates II and III anterior margin of this plate; setae Ae located near wich are fused to one another, suture line between middle of excretory pore plate. them incomplete and developed only in their lat- Capitulum (Fig. 128) short, with wide base, eral parts (Fig. 125). Anterior margin of coxal anterior hypostomal setae long and thin, posterior plates I concave and shorter than posterior one. setae short, spine-like. Basal segments of chelic- Setae C1 and C4 considerably shorter than C2 and erae (Fig. 129) fused by their medial parts and ta- C3, the latter are approximately equal in length. pering proximally; suture line well developed Base of seta C4 on coxal plate III and medial end along their whole length. Cheliceral stylet small of suture line between coxal plates II and III well and crescent in shape. separated. Posteromedial apodemes of coxal plates Pedipalp trochanter (Fig. 130) short and with- I small, sometimes they are absent, apodemes of out setae; dorsodistal seta of femur and genu short coxal plates II not developed, and apodemes of and approximately equal in size, proximal lateral coxal plates III poorly developed. Transverse genual seta very long and thick; tibia with 3 ap- muscle attachment scar not developed in postero- proximately subequal short and thin setae and with medial angles of coxal plates III. Setae Ci, Si, Se, large dorsodistal claw. Pedipalpal tarsus with short Le, Pi, Pe situated on soft interscutal membrane in solenidion, 2 long setae and 5 short unequal ones. the posterior part of the body. Setae Ci longer and The general number of setae on legs I–III thicker than all ventral idiosomal setae, and situ- (Figs 131–133), except eupathidia, as follows: I– ated on small bases; setae Si shortest and thin. Leg. 1–5: 1, 7, 5 (s), 11(2s), 13 (s, ac); II–Leg. Coxal plates I–III with reticulations. Urstigma 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); III–Leg. 1–5: 1, small, occupying lateral position on the border be- 6, 5(s), 10 (s), 11 (ac). Formula of large setae on tween coxae I and II, and provided with a cup. leg segments as follows: I–Leg 1–5: 0, 2, 1, 1, 0;

143 P. V. Tuzovsky

134, 135

131 134

132

133 135

131–133

Figs 131–135. Tiphys scaurellus, larva: 131 — leg I, 132 — leg II, 133 — leg III, 134 — claws of leg I, 135 — claws of leg III. Scale bars: 131–133 = 100, 134–135 = 25.

II–Leg 1–5: 0, 2, 2, 3, 0; III–Leg 1–6: 0, 2, 2, 3, 0. lum 85–90; length of basal segment of chelicera Solenidia on genua and tibiae of legs I–III rela- 70–76, length of chelicera stylet 18–20; length of tively short. Solenidion on genu I slightly shorter medial margin of coxa I 85–90, length of medial than distal solenidion on tibia I; proximal solen- margin of coxae II+III 135–140; length of pedi- idion on tibia I slightly shorter than distal one, so- palpal segments (P-1–5): 9–10, 38–40, 23–25, lenidion on genu II and both solenidia on tibia II 10–11, 8–9; length of legs segments: I–Leg. 1–5: nearly equal in length; solenidion on genu III and 40–45, 39–43, 37–40, 58–60— 80–92; II–Leg. solenidion on tibia III subequal. Central claw and 1–5: 40–45, 41–45, 43–46, 70–73, 88–97; III–Leg. lateral claws of legs I nearly subequal in size; on 1–5: 40–45, 45–49, 43–47, 72–76, 102–107. tarsi of legs I and II, empodium bent at right angle Ecology. Temporary ponds, sedge-sphagnum and ambulacra only slightly bent (Fig. 134). Distal bogs, peat-bogs. part of empodium of leg III turned to ventral side Geographical distribution. Europa, Russia: at obtuse angle (Fig. 135). Arkhangelsk Province; Asia, Russia: Magadan Measurements, n=10. Length of dorsal shield Province, Kamchatka Territory, Primorsk Territo- 330–340, its width 280–290; length of excretory ry, Khabarovsk Territory (Tuzovsky 1983, 1990; pore plate 46–55, width 48–52; length of capitu- Semenchenko et al 2010).

144 Water mites of the genus Tiphys

Tiphys (Tiphys) lapponicus (Neuman, 1880) Deutonymph. Dorsum with well developed Figs 136–171 anteromedial plates only. Medial margin of coxae Material. 6 females, 1 male, 3 deutonymphs: IV longer than medial margin of coxae III. Pos- Russia, Magadan Province, Anadyr Distr., Marko- teromedial angle of coxa IV moderately developed vo: lake on the left bank of the Anadyr river, 5 km with long apodeme. Genital plates with 2 aceta- upstream of settlement, June–July 1978 and 1981, bula and 3 short and fine setae each, both aceta- P.V. Tuzovsky; 6 females, 28 larvae reared in labo- bula relatively large and approximately subequal ratory (IBIW): Magadan Province, Tenka Distr., in size, distance between acetabula usually less settlement Sibik-Tyellakh, July–August 1987, P.V. than one diameter of acetabulum on each plate. Tuzovsky; 11 female, 5 males, 6 deutonymphs: Postgenital sclerite considerably larger than pre- Magadan Province, Chaun Distr., temporary ponds genital sclerite. Excretory pore non-sclerotized. on right bank of the Chaun river, 18 km upstream Pedipalpal genu with 2 setae, external lateral seta from mouth of the river, July–August 1982, P.V. longer than dorsodistal seta; base of external lat- Tuzovsky. eral seta located near distal end of segment; bases Diagnosis. Female. Medial margin of coxal of ventral setae on tibia separated and located in plate IV twice as long as medial margin of coxal middle of segment, distolateral spine of tibia 5–6 plate III. Posteromedial margin and posterolateral times as short as tarsus. margin of coxae IV forming obtuse angle with Larva. Dorsal shield elongate (ratio length/ short apodeme. Genital plates more or less trian- width 1.5–1.6), covering almost all dorsum in un- gular, with 3 acetabula and 10–13 short, fine setae engorged larva; setae Fch longer than Fp, Vi and each; all acetabula small, approximately subequal Oi. Setae C1 considerably shorter than setae C2, in size, and usually arranged in obtuse triangle or setae C3 and C4 longer than setae C2; base of se- in an arc; genital opening longer than length of tae C4 on and medial end of suture line between genital plates, anterior genital sclerite 2–3 times as coxal plates II and III well separated. Transverse width as posterior one. Pedipalps moderately de- muscle attachment scar absent. Dorsal shield and veloped; genu with 4 setae, distal external lateral coxal plates I–III with reticulations. Excretory seta considerably longer than other dorsal setae; pore plate oval, ratio length/width 1.1–1.3. Setae base of distal external lateral seta attached near Ai located in front of setae Ae, last longer than Ai. distal end of segment; tibia with short distolateral Excretory pore open between rows setae Ai and Ae spine (4–5 times shorter than tarsus), both ventral in anterior half of excretory pore plate Solenidion setae long and located in middle of segment, ven- on genu I and both solenidia on tibia I subequal in tral tubercles not developed. Claws of legs I–III length; solenidion on genu II and proximal solen- are larger than claws of legs IV; both denticles ap- idion on tibia II subequal in length and slightly proximately subequal in length, external denticle shorter than distal solenidion on tibia II. thin-pointed, and internal denticle expanded dis- Description. Female. Body oval; setae Fch tally with rounded tip. Claw lamella moderately long, whip-like (Fig. 136), other idiosomal setae developed with nearly straight ventral margin. filiform. Integument soft with very fine strips. Male. Coxae plates IV greatly expanded, Dorsum with relatively large anteromedial plate- touching each other posteromedially, medial mar- lets (Fig. 137), anterolateral and posteromedial gin of coxal plate IV 3.1–4.5 times longer than platelets rudimentary or absent. Coxal plate I usu- medial margin of coxal plate III. Suture line be- ally with 4 lateral thin setae and 1 posteromedial tween coxal plates III and IV nearly transverse. thin seta, coxal plate II with 2–3 lateral thin setae Posteromedial margins of coxal plate IV slightly (Fig. 138). Apodeme of coxal plate I moderately bowed or straight, oblique, directed posterolater- developed and usually not extending beyond me- ally; genital bay shallow, most of the genital field dial margin of coxal plate III. Suture line between lying outside the genital bay. Genital plates trian- coxal plates III and IV oblique, directed postero- gular, fused medially. Anterior genital sclerite medially. Medial margin of coxal plate IV twice as small, transverse, with two pairs of short and fine long as medial margin of coxal plate III. Postero- setae. Genital plates with 4 short and fine setae medial margin and posterolateral margins of coxae each. Sclerites bearing setae Hv closely together IV forming right or obtuse angle with short apo- with posterior margin of coxae II; other sclerites deme beneath this angle. Sclerites, bearing setae bearing ventral idiosomal setae lying free in soft Hv other idiosomal setae, lying free in soft integu- integument. ment.

145 P. V. Tuzovsky

136 136, 137

139 140

140–142

144 143

138 142 141 Figs 136–144. Tiphys lapponicus, female: 136 — seta Fch, 137 — anteromedial dorsal plates, 138 — ventral view, 139–142 — genital plates, 143–144 — excretory pore. Scale bars: 136–142 =100, 143–144 = 50.

145 145–147, 149–150

148 147

146

149

148

150

Figs 145–150. Tiphys lapponicus, female: 145 — chelicera, 146 — pedipalp, 147 — pedipalpal tibia, 148 — genu, tibia and tarsus of leg IV, 149–150 — claw. Scale bars= 100.

146 Water mites of the genus Tiphys

151 152 154

153

Figs 151–154. Tiphys lapponicus, male: 151 — ventral view, 152 — genu, tibia and tarsus of leg III, 153 — telofemur, genu, tibia and tarsus of leg IV, 154 — claws of leg III. Scale bars: 151–153 = 100, 154 =50.

Genital plates more or less triangular (Figs ones, both ventral setae inserted near middle of 139–142), with 3 acetabula and 10–13 short, fine tibia; distolateral peg-like spine of tibia short with setae each. All acetabula small, subequal in size obtuse tip. Pedipalpal tarsus with one long proxi- and usually arranged in an obtuse triangle or some- mal solenidion, 5 thin setae and 4 thick distal un- times in an arc; acetabula occupying approximate- equal. ly half or less of area of genital plate; distance be- Legs II–IV with swimming setae; second pair tween first and second acetabula and second and of legs with short swimming setae, two posterior third acetabula usually less than diameter of one pairs of legs with long swimming ones (Fig. 148). acetabulum. Genital opening longer than genital Three swimming setae present on genu of legs II, plates, pregenital sclerite 2–3 times as wide as 4–5 on tibia II and genu III, 5–6 on genu IV, 6–8 postgenital sclerite. Excretory pore non-sclero- on tibia III and 8–9 on tibia IV. Claws of legs I–III tized (Fig. 143) or with very small anterior sclerite are larger, than claws of legs IV; both denticles (Fig. 144). subequal in length, external denticle thin pointed, Capitulum with short anchoral projection, and internal denticle expanded distally with round- both pairs of setae equal in length. Basal segment ed tip (Fig. 149–150). Claw lamella moderately of chelicera (Fig. 145) large, with strong convex developed with nearly straight ventral margin. dorsal margin and slightly concave ventral margin. Measurements (n=10). Body length 1000– Cheliceral stylet moderately developed, pointed. 1315, length of medial margin of coxa III 65–80, Pedipalps (Fig. 146) rather slender. Pedipal- length of medial margin of coxa IV 110–150; pal trochanter with a single short dorsodistal seta, length of anteromedial dorsal plates 55–70, theirs femur with 3–5 proximal and 2 dorsodistal setae. width 15–20, length of genital plates 160–195, Ventral margin of femur straight or slightly con- theirs width 80–125; length/width of genital ac- cave, ventral margin of genu concave. Pedipalpal etabula (ac. 1–3): 42–48/30–36, 40–45/30–35, genu with 4 setae, distal external lateral seta con- 36–48/30–36; width of anterior genital sclerite siderably longer than other dorsal setae; base of 65–80; length of basal segment of chelicera 170– distal external lateral seta attached near distal end 205, length of cheliceral stylet 73–80; length of of segment. Pedipalpal tibia slightly longer than pedipalpal segment (P-1–5): 45–50, 155–172, 95– femur with short distolateral spine (4–5 times as 125, 160–180, 73–82; length of distolateral spine short as tarsus), usually 2 (sometimes 3, Fig. 147) of tibia 12–18; length of leg segments: I–Leg. 1–6: rather long ventral setae and several fine dorsal 90–100, 160–190, 180–220, 250–285, 270–295,

147 P. V. Tuzovsky

155 155, 156, 158 156

159

158 157

Figs 155–159. Tiphys lapponicus, deutonymph: 155 — anterior dorsal plates, 156 — seta Fch, 157 — ventral view, 158— pedipalp, 159 — genu, tibia and tarsus of leg IV. Scale bars = 100.

285–325; II–Leg. 1–6: 90–105, 170–205, 185– genu III with 4–5 and tibia III with 8–9 long swim- 220, 230–285, 285–320, 285–325; III–Leg. 1–6: ming setae. Telofemur IV (Fig. 153) very short, 95–125, 160–205, 180–235, 260–305, 300–360, genu IV slightly expanded dorsoventrally in distal 285–325; IV–Leg.1–6: 160–190, 195–230, 220– half, bearing a few dorsal and ventral setae, and 270, 325–390, 375–425, 340–395. several relatively short lateral setae; dorsal setae Male. Dorsal surface, coxae of legs I–III as in arranged in two groups: proximal group with sev- female. Anterior apodemes rather large, extending eral relatively short and thin setae; distal group beyond medial margin of coxal plate III; posterior with a few long and thick setae. Tibia IV slightly apodemes slightly developed (Fig. 151). Coxae curved, with concave dorsal margin and convex plates IV greatly expanded, touching each other ventral one, with numerous unequal setae but posteromedially, with 2 short and fine setae close without long swimming setae, and a single short together in this region on each side. Medial mar- fan-shaped seta. Tarsus of leg IV considerably gin of coxal plate IV 3.1–4.5 times longer than longer than tibia, slightly tapering distally with medial margin of coxal plate III. Suture line be- numerous short setae and 6–8 long unequal swim- tween coxal plates III and IV nearly transverse. ming setae. Posteromedial margins of coxal plate IV slightly Claw of legs I and II similar to female. Claws bowed or straight, oblique, directed posterolater- of legs III (Fig. 154) asymmetrical, large claw ally; lateral margins of these coxal plates convex. with 2 long subequal denticles, clawlet with rela- Genital plates triangular, fused medially; all aceta- tively long external and short internal pointed den- bula subequal in size. Anterior genital sclerite ticles. small, transverse with two pairs of short and fine Measurements (n=6). Length of body 650– setae; genital plates with 2–4 short, fine setae each. 860, length of medial margin of coxa III 60–80, Sclerites, bearing all ventral idiosomal lying free length of medial margin of coxa IV 240–270; in soft integument. Excretory pore unsclerotized. length of anteromedial dorsal plates 40–55, width Capitulum, chelicerae and pedipalps as in fe- 12–15; length of genital plates 105–130, general male, but pedipalpal tibia with more numerous width 220–245; length/width of genital acetabula dorsal setae. (ac. 1–3): 42–60/36–42, 36–60/35–42, 42–60/36– Genu of leg III (Fig. 152) shorter than tibia 48; length of basal segment of chelicera 145–165; III, and tarsus III slightly longer than tibia III; length of cheliceral stylet 55–65; length of pedi-

148 Water mites of the genus Tiphys

160 161

Figs 160–161. Tiphys lapponicus, larva: 160 — dorsal view, 161 — ventral view. Scale bar: 160–161 = 100. palpal segment (P-1–5): 30–40, 115–130, 80–90, acetabulum. Postgenital sclerite larger than prege- 115–140, 48–57; length of distolateral spine of nital sclerite. Excretory pore unsclerotized. tibia 9–12; length of leg segments: I–Leg. 1–6: Pedipalpal trochanter without seta, femur 75–80, 145–155, 145–165, 195–235, 210–290, with 3 dorsodistal setae (Fig. 158). Ventral margin 325–350; II–Leg. 1–6: 75–100, 130–155, 145– of femur straight or slightly curved, ventral mar- 165, 195–235, 200–255, 310–345; III–Leg. 1–6: gin of genu concave. Pedipalpal genu with 2 setae, 90–100, 120–155, 95–105, 120–130, 160–190, external lateral seta longer than dorsodistal seta 200–240; IV–Leg.1–6: 145–180, 150–170, 105– and inserted near distal end of segment. Pedipal- 115, 195–230, 265–295, 445–490. pal tibia with short distolateral peg-like spine, 2 Deutonymph. Dorsum with well developed ventral and 2 dorsodistal setae; bases of ventral anteromedial plates only (Fig. 155). Setae Fch setae separated and inserted near middle of seg- long and thin (Fig. 156). Coxal plate I usually with ment. Distolateral seta of tibia short, with obtuse 3 lateral thin setae and 1 posteromedial thin seta, tip, 5–6 times shorter than tarsus. Pedipalpal tar- coxal plate II with 1 lateral thin setae, coxal plate sus with single long proximal solenidion, 3 thin III with 1 medial seta and 1–2 lateral thin setae, setae and 4 distal unequal setae. coxal plate IV with 2 medial seta and 1 lateral seta Legs II–IV with swimming setae; legs II with (Fig. 157). Apodeme of coxal plate I moderately short swimming setae, two posterior pairs of legs developed and not reaching medial margin of cox- with long swimming ones (Fig. 159). One or 2 al plate III. Suture line between coxal plates III swimming setae present on genu legs II–IV and and IV oblique, directed posteromedially. Medial tibia II, 2–3 on tibia III, 4–5 on tibia IV. Claws of margin of coxae IV longer than medial margin of legs I–III larger, than claws of legs IV; both den- coxae III. Posteromedial angle of coxae IV moder- ticles approximately subequal in length, external ately developed, with short apodeme. Sclerites denticle thin and pointed, internal denticle ex- bearing setae Hv and other ventral idiosomal setae panded distally with rounded tip. Claw lamella lying free in soft integument. Genital plates sepa- slightly developed, with straight ventral margin. rated, oblique, directed anteromedially, bearing 2 Measurements, n=9. Body length 530–735, acetabula and 3 short and fine setae each. Both ac- length of medial margin of coxa III 24–36, length etabula approximately subequal in size, distance of medial margin of coxa IV 45–55; length of an- between them usually lesser than one diameter of teromedial dorsal plates 25–48, theirs width 9–15,

149 P. V. Tuzovsky

164 165

166

162 163

Figs 162–166. Tiphys lapponicus, larva: 162–163 — excretory pore plate, 164 — capitulum (ventral view), 165 — chelicerae, 166 — pedipalp. Scale bars: 162–163 = 50, 164–165 = 100, 166 =20. length of genital plates 60–70, theirs width 35–40; sides of coxae II+III passing onto dorsal side and diameter of genital acetabula (ac. 1–2): 22–30, almost extending to setae Oi and other hysterosom- 24–32; length of basal segment of the chelicera al setae located on interscutal membrane. Dorsal 120–125; length of the cheliceral stylet 55–57; shield with a reticulations. length of pedipalpal segment (P-1–5): 24–32, 84– Coxal plates I–II relatively small; anterior 92, 55–65, 87–92, 44–48; length of distolateral coxal plates separated from coxal plates II and III spine on tibia 7–11; length of leg segments: I–Leg. which fused, suture line between them incomplete 1–6: 48–52, 80–85, 100–115, 130–140, 150–165, and developed only in their lateral parts (Fig. 161). 170–185; II–Leg. 1–6: 52–57, 80–90, 98–103, Anterior margin of coxal plates I concave and con- 148–155, 165–175, 185–195; III–Leg. 1–6: 57– siderably shorter than posterior margin. Setae C1 63, 80–90, 103–110, 140–150, 170–180, 180–195; considerably shorter than setae C2–C4, base of se- IV–Leg.1–6: 88–95, 88–95, 120–125, 160–168, tae C4 and medial end of suture line between cox- 200–210, 225–235. al plates II and III well separated. Posteromedial Larva. Body small and oval; dorsal shield apodemes of coxal plates I–III poorly developed, truncated anteriorly, rounded posteriorly, with sometimes apodemes 2 absent. Transverse muscle convex lateral margins and weakly concave ante- attachment scar not developed in the posterome- rolateral margins near eyes plates (Fig. 160). Dor- dial angles of coxal plates III. Setae Ci, Si, Se, Le, sal shield elongate (ratio length/width 1.5–1.6), Pi, Pe situated on the soft interscutal membrane in covering almost all dorsum in unengorged larva posterior part of body. Setae Ci very long, thicker and bearing 4 pairs of setae (Fch, Fp, Vi, Oi); setae than all ventral idiosomal setae and situated on Fch longer than Fp, Vi and Oi. Setae Oe, Hi, He, small bases; setae Si shortest and thin. Coxal plates Sci, Sce and Li located on the soft wrinkled mem- I–III with reticulations on it. Urstigma occupyinga brane. Setae Oe longest and thickest of all dorsal lateral position on the border between coxae I and setae, setae Si slightly shorter than Sce. Lateral II and provided with cups.

150 Water mites of the genus Tiphys

167

170

168

171

167–169

170, 171 169

Figs 167–171. Tiphys lapponicus, larva: 167 — leg I, 168 — leg II, 169 — leg III, 170 — claws of leg I, 171 — claws of leg III. Scale bars: 131–133 = 100, 134–135 = 25.

Excretory pore plate (Figs 162–163) oval, ra- Pedipalp trochanter (Fig. 166) short and with- tio length/width = 1.1–1.3. Setae Ai located near out setae; femur with 1 short and thin seta; genu anterior margin of excretory pore plate, setae Ae with 1 very long proximal seta and short and thin longer than setae Ai and placed near middle of this distal one; tibia with 3 approximately subequal se- plate. Excretory pore located in anterior half of ex- tae, and large dorsodistal claw. Pedipalpal tarsus cretory pore plate between rows setae Ai and Ae with 1 short solenidion, 2 long and relatively thick Capitulum (Fig. 164) with a wide base, ante- setae and 5 unequal thin setae. rior hypostomal setae longer and thinner than pos- All legs rather long. General number of setae terior ones. Basal segments of chelicerae (Fig. on legs I–III (Figs 167–169), except eupathidia, is 165), fused at their medial parts and tapering prox- as follows: I–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, imally; suture line between them well developed ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); III– along their whole length. Cheliceral stylet small Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). Formula of and crescent in shape. large setae on leg segments as follows: I–Leg 1–5:

151 P. V. Tuzovsky

0, 2, 1, 1, 0; II–Leg 1–5: 0, 2, 2, 3, 0; III–Leg 1–6: sclerite considerably wider than posterior one. Pe- 0, 1, 2, 3, 0. Solenidion on genu of leg I and both dipalpal genu with 3 setae, external lateral seta solenidia on tibia I approximately equal in length; longer than other dorsal setae; base of external lat- solenidion on genu II and proximal solenidion on eral seta situated proximally to the middle of the tibia II slightly shorter than distal solenidion on segment. Pedipalpal tibia with short distolateral tibia II; solenidion on genu of leg III shorter than seta (2.5–3.0 times shorter than tarsus), both ven- solenidion on tibia III. Central claw and lateral tral setae situated on small tubercles distally to the claws on tarsi of legs I–II almost subequal in size; middle of the segment. on tarsi of legs I and II, empodium bent at right Male. Coxal plates IV greatly expanded, angle and ambulacra only slightly bent (Fig. 170). touching each other along whole medial margin, Distal part of empodium of leg III turned onto posterior margin of coxal plate IV nearly trans- ventral side at obtuse angle (Fig. 171). verse, genital bay shallow. Medial margin of coxal Measurements (n=10). Length of dorsal shield plate IV 3.2–4.0 times longer than medial margin 350–390, its width 225–245; length of anal plate of coxal plate III. Genital plates triangular, fused 38–55, its width 38–45; length of the capitulum medially. Genital plates with 4–5 short, fine setae 105–112; length of basal segment of the chelicera each. Sclerites bearing setae Pe fused with pos- 95–102, length of cheliceral stylet 20–26; length teromedial margin of coxae IV; sclerites bearing of medial margin of coxae I 80–96, length of me- setae Hv fused with posterior margin of coxae II; dial margin of the coxae II+III 155–180; length of other sclerites, bearing ventral idiosomal setae, ly- anal plate 44–55, width 38–44; length of pedipal- ing free in soft integument. Genu IV greatly ex- pal segments (P-1–5): 8–10, 40–43, 26–29, 9–12; panded dorsoventrally in proximal half of seg- length of legs segments: I–Leg. 1–5: 35–40, 38– ment. Tibia IV slightly curved with a single thick 44, 44–48, 62–64, 84–86; II–Leg. 1–5: 35–44, dorsodistal seta and long ventrodistal swimming 38–54, 48–52, 72–75, 100–105; III–Leg. 1–5: 44– setae. Tarsus and tibia IV nearly subequal in 48, 45–52, 48–52, 76–80, 110–115. length. Claws of legs III asymmetrical, large claw Ecology. Temporary ponds, sedge bogs, lakes with subequal external and internal denticles, ex- with macrophytes, streams. ternal denticle provided by thin spur; small claw Geographical distribution. Europe (Viets with short external and short internal denticles. 1936, 1956; Lundblad 1968, K.O. Viets 1978), Deutonymph. Dorsum without true dorsal Asia; Russia: Leningrad Province, Komi Repub- plates. Medial margin of coxae III slightly longer lik, and Magadan Province (Sokolow and Yank- than medial margin of coxae III. Genital plates ovskaya 1962; Tuzovsky 1990). separated, oblique, directed anteromedially, both acetabula subequal in size, distance between them Tiphys (Tiphys) latipes (Müller, 1776) usually more than one diameter of acetabulum. Figs 172–204 Postgenital and pregenital sclerites subequal in Material. 3 females, 1 male: Asia, Russia: size, situated closely to each other between ante- Kemerovo Province, Prokopjevsk Distr., village rior ends of genital plates. Pedipalpal genu with 2 Sharap, spring, 15 July 1973, 5 females: Magadan setae, external lateral seta considerably longer Province, Tenka Distr., village Agrobasa, June– than dorsodistal one; base of external lateral seta July 1979, 26 females, 11 males, 3 deutonymph situated distally to middle of segment. Pedipalpal and 30 larvae reared in laboratory (IBIW); Europe, tibia with very long distolateral seta (0.65–0.70 of Russia: Yaroslavl Province, Nekouz Distr., near length of tarsus), base of ventral setae separated settlement Borok, temporary pond, sedge bogs, and situated on small tubercles in the middle of the sedge-sphagnum bogs, April–July 2002–2006, segment. P.V. Tuzovsky. Larva. Dorsal shield elongate (ratio length/ Diagnosis. Female. Medial margin of coxal width 1.5–1.6), covering almost complete dorsum plate IV longer than medial margin of coxal plate in unengorged larva and bearing only 2 pairs of III. Posteromedial and posterolateral margins of trichobothria (Fp, Oi), setae Fch and Vi situated on coxae IV forming obtuse angle with a short apo- subcutaneous extensions of dorsal shield. Setae deme. Genital plates lying in an arc or more or less Fch and Vi slightly longer than Fp and Oi. All cox- triangular, with 3 acetabula and 8–11 short and al setae (C1–C4) long and approximately subequal fine setae each. Genital opening and genital plates in length. Base of setae C4 on coxal plate III situ- approximately subequal in length, anterior genital ated closely to medial end of suture line between

152 Water mites of the genus Tiphys

172 173

174 173, 174

176 177

175

176–180

180 179 178 Figs 172–180. Tiphys latipes, female: 172 — seta Fch, 173 — anteromedial and anterolateral dorsal plates, 174 — posteromedial plates, 175 — ventral view, 176–179 — genital plate, 180 — excretory pore. Scale bars: 172 =50, 173–180 = 100. coxal plates II and III. Transverse muscle attach- III. Suture line between coxal plates III and IV ment scar on coxal plate III present. Dorsal shield oblique, directed posteromedially. Medial margin and coxal plates I–III porous. Excretory pore plate of coxal plate IV longer than medial margin of wider than long or nearly as long as wide, ratio coxal plate III. Posteromedial and posterolateral length/width = 0.8–1.0, setae Ae longer than setae margins of coxae IV forming right or obtuse angle Ai; excretory pore open in posterior half of anal with short apodeme. Sclerites, bearing setae Hv, plate. Solenidion on genu of leg I slightly longer fused with posteromedial margin of coxa II, other than distal solenidia on tibia I which subequal in idiosomal setae lying free in soft integument. length; solenidion on genu II shorter than proxi- Genital plates slightly elongate and lying in mal solenidion on tibia II, latter slightly shorter an arc or more or less triangular (Figs 176–179) than distal solenidion on tibia II. with 3 acetabula and 8–11 short and fine setae Description. Female. Body oval; setae Fch each. All acetabula small, subequal in size and ar- long, whip-like (Fig. 172), other idiosomal setae ranged in an arc or in a triangular shape; acetabula filiform. Dorsum with relatively large anterome- occupying approximately half of area of the geni- dial platelets, small anterolateral and posterome- tal plate; distance between first and second aceta- dial platelets (Figs 173–174). Anteromedial and bula, and second and third acetabula lesser than posteromedial plates elongate, anteromedial plate- diameter one acetabulum. Genital opening and lets oval or nearly circular. Coxal plate I usually genital plates subequal in length, anterior genital with 4 lateral thin setae and 1 posteromedial thin sclerite wider than posterior one. Excretory pore seta, coxal plate II with 3 lateral thin setae, coxal with a very small anterior sclerite (Fig. 180). plate III with 1 medial seta and 3 lateral setae, Capitulum with short anchoral projection, coxal plate IV with 1 medial seta and 4–5 lateral both pairs of setae equal in length. Basal segment setae (Fig. 175). Apodeme of coxal plate I moder- of chelicera (Fig. 181) large, with obtuse dorsal ately developed and usually in mature specimens hump and slightly concave ventral margin. Cheli- extending beyond medial margin of coxal plate ceral stylet relatively small, pointed.

153 P. V. Tuzovsky

181

181, 182

183 184

182

Figs 181–184. Tiphys latipes, female: 181 — chelicera, 182 — pedipalp, 183 — genu, tibia and tarsus of leg IV, 184 — claw. Scale bars: 181–183= 100, 184 = 50. Pedipalps (Fig. 182) moderately developed. or less rounded tip (Fig. 184). Claw lamella mod- Pedipalpal trochanter with a short dorsodistal seta, erately developed with slightly convex or straight femur with 3–4 dorsoproximal and 2 dorsodistal ventral margin. setae. Ventral margin of femur straight or slightly Measurements (n=10). Body length 650– convex, ventral margin of genu slightly concave. 1200, length of medial margin of coxa III 48–60, Pedipalpal genu with 3 setae, external lateral seta length of medial margin of coxa IV 65–90; length considerably longer than other dorsal setae; base of anteromedial dorsal plates 50–65, width 18–25, of external lateral seta inserted proximally to midlength of genital plates 120–165, width 60–95; dle of segment. Pedipalpal tibia thin, slender, with length/width of genital acetabula (ac. 1–3): 48– a short distolateral seta, 2 ventral setae and 2–3 60/35–42, 42–55/35–42, 48–65/36–45; width of short and fine dorsal setae. Both ventral setae in- anterior genital sclerite 78–90; length of basal seg- serted in distal portion of tibia on small tubercles. ment of chelicera 140–155, length of cheliceral Distolateral peg-like seta of tibia 2.5–3.0 times stylet 55–60; length of pedipalpal segment (P-1–5): shorter than tarsus. Pedipalpal tarsus tapering dis- 40–48, 110–125, 60–80, 120–140, 50–60; length tally with long proximal solenidion, 5 thin setae of distolateral spine on tibia 20–24; length of leg and 4 thick unequal distal setae. segments: I–Leg. 1–6: 55–75, 105–125, 115–130, Legs II–IV with swimming setae; second pair 145–165, 160–180, 200–210; II–Leg. 1–6: 75–80, of legs with short swimming setae, two posterior 110–130, 120–140, 155–180, 170–195, 220–230; pairs of legs with long swimming setae (Fig. 183). III–Leg. 1–6: 80–90, 105–130, 120–140, 155–175, Three or four swimming setae present on genu and 185–220, 195–220; IV–Leg. 1–6: 135–155, 115– tibia of legs II, 2–3 on genu III, 4–5 on genu IV 145, 155–175, 210–230, 235–255, 235–245. and tibia III, and 4–7 on tibia IV. Claws of legs I– Male. Dorsal surface, coxae of legs I–III as in III larger than claws of legs IV; both denticles ap- female. Anterior apodemes rather large, extending proximately subequal in length, external denticle beyond medial margin of coxal plate III; posterior thin pointed, and internal denticle wide with more apodemes slightly developed (Fig. 185). Coxae

154 Water mites of the genus Tiphys

185 186 187

185–187 188 Figs 185–188. Tiphys latipes, male: 185— ventral view, 186 — genu, tibia and tarsus of leg III, 187 — telofemur, genu, tibia and tarsus of leg IV, 188 — claws of leg III. Scale bars: 185–187 = 100, 188 =50. plates IV greatly expanded, touching each other and thinner than ventrodistal setae. Tibia IV nearly along whole medial margins, abruptly depressed straight with a few rather long and thick dorsal se- posteromedially, with 2 short and fine setae ante- tae, several short and thick ventral setae, a single rior to depressed region on each plate. Posterior thick dorsodistal peg-like seta and long distal margin of coxal plate IV nearly transverse, genital swimming setae. Tarsus IV with numerous short bay shallow. Medial margin of coxal plate IV 3.0– and fine setae and 5–7 relatively long unequal 4.0 times longer than medial margin of coxal plate thick dorsal setae. III. Suture line between coxal plates III and IV Claw of legs I and II as in female. Claws of oblique, directed posteromedially. Genital plates legs III (Fig. 188) asymmetrical, large claw with triangular, fused medially; all acetabula approxi- approximately subequal external and internal den- mately equal in size and occupying about one half ticles, external denticle provided with thin spur; of area of plates. Anterior genital sclerite small small claw with subequal external and internal and curved, with two pairs of short and fine setae. denticles. Genital plates with 2–5 short, fine setae each. Sc- Measurements (n=10). Body length 530–735, lerites bearing setae Pe fused with posteromedial length of medial margin of coxa III 65–80, length margin of coxae IV; sclerites bearing setae Hv of medial margin of coxa IV 250–280; length of fused with posterior margin of coxae II; other sc- anteromedial dorsal plates 55–60, theirs width lerites bearing ventral idiosomal setae lying free in 20–25, length of genital plates 72–90, theirs gen- soft integument. Excretory pore with small ante- eral width 180–250; length/width of genital aceta- rior sclerite. bula (ac. 1–3): 36–48/30–36, 42–48/36–42, 37– Capitulum, chelicerae and pedipalps as in fe- 55/30–42; length of basal segment of chelicerae male, but pedipalpal tibia with 3–7 dorsal setae. 105–155; length of cheliceral stylet 40–57; length Genu III (Fig. 186) shorter than tibia III, tar- of pedipalpal segment (P-1–5): 32–40, 90–120, sus III and tibia III approximately equal in length; 50–65, 95–125, 40–55; length of distolateral spine genu III with 3–4 relatively short, and tibia III on tibia 18–21; length of leg segments: I–Leg. with 6–8 long swimming setae. Telofemur IV (Fig. 1–6: 55–75, 80–125, 90–130, 115–170, 130–190, 187) short, genu IV greatly expanded dorsoven- 185–255; II–Leg. 1–6: 65–80, 85–130, 90–140, trally in proximal half with almost straight dorsal 120–175, 130–195, 185–245; III–Leg. 1–6: 75– margin and very large ventral hump, bearing nu- 100, 80–125, 55–90, 80–125, 120–180, 145–205; merous dorsal and ventrodistal setae and several IV–Leg. 1–6: 120–155, 90–145, 60–100, 145–195, relatively short lateral setae; dorsal setae longer 180–235, 180–235.

155 P. V. Tuzovsky

189, 191

191 190 189

Figs 189–191. Tiphys latipes, deutonymph: 189 — ventral view, 190— pedipalp, 191 — genu, tibia and tarsus of leg IV. Scale bars = 100. Deutonymph. Dorsum without true dorsal with 2 setae, external lateral seta considerably lon- plates. Setae Fch long, thin. Coxal plate I usually ger than dorsodistal seta; base of external lateral with 3 lateral thin setae and 1 posteromedial thin seta inserted proximally to middle of segment. Pe- seta, coxal plate II with 1 lateral thin setae, coxal dipalpal tibia with very long distolateral seta (0.6– plate III with 1 medial seta and 1–2 lateral thin 0.7 times as long as tarsus), 2 ventral and 1–2 dor- setae, coxal plate IV with 1 medial seta and 1 lat- sodistal setae; base of ventral setae inserted on eral seta (Fig. 189). Apodeme of coxal plate I small tubercles near distal end of segment. Pedi- moderately developed and not reaching medial palpal tarsus with long proximal solenidion, 3 thin margin of coxal plate III. Suture line between cox- setae and 4 thick and short distal spines. al plates III and IV oblique, directed posteromedi- Legs II–IV with swimming setae; second pair ally. Medial margin of coxae III slightly longer of legs with short swimming setae, two posterior than medial margin of coxae III. Posteromedial pairs of legs with long swimming setae. One or angle of coxae IV obtuse, with rather long apo- two swimming seta present on genu legs II–IV and deme. Sclerites bearing setae Hv fused with poste- tibia II–III, 4–5 on tibia IV (Fig. 191). Claws of rior margin of coxae II, setae Pe and other ventral legs I–III larger than claws of legs IV; both denti- idiosomal setae lying free in soft integument. cles subequal in length, external denticle thin and Genital plates separated, oblique, directed antero- pointed, internal denticle wide with rounded tip. medially, bearing 2 acetabula and 3 short and fine Claw lamella slightly developed, with nearly setae each. Both acetabula approximately equal in straight ventral margin. size, distance between them usually more than one Measurements, n=3. Body length 420–570, diameter of acetabulum. Genital and pregenital sc- length of medial margin of coxa III 30–35, length lerites subequal, situated closely together and of medial margin of coxa IV 33–38; length of gen- placed between anterior ends of genital plates. Ex- ital plates 60–66, width 24–30; diameter of genital cretory pore with very small anterior sclerite. acetabula (ac. 1–2): 18–21, 18–22; length of basal Pedipalpal trochanter without seta, femur segment of chelicera 100–105; length of cheliceral with 1 dorsoproximal seta and 2 dorsodistal short stylet 33–36; length of pedipalpal segment (P-1–5): setae (Fig. 190). Ventral margin of femur straight, 24–327, 72–75, 42–48, 69–78; length of distolat- ventral margin of genu concave. Pedipalpal genu eral spine on tibia 28–32; length of leg segments:

156 Water mites of the genus Tiphys

192 193

Figs 192–193. Tiphys latipes, larva: 192 — dorsal view, 193 — ventral view. Scale bar: 192–193 = 50.

199 198

197

194 195 196 Figs 194–199. Tiphys latipes, larva: 194–196 — excretory pore plate, 197 — capitulum (ventral view), 198 — chelicerae, 199 — pedipalp. Scale bar: 194–199 = 50.

I–Leg. 1–6: 42–48, 42–48, 57–60, 78–82, 90–96, Larva. Body small and oval; dorsal shield 108–115; II–Leg. 1–6: 42–48, 48–54, 60–66, 81– truncated anteriorly, rounded posteriorly, with con- 87, 97–102, 115–123; III–Leg. 1–6: 50–54, 48–57, vex undulating lateral margins, anterolateral mar- 54–60, 84–90, 112–115, 115–122, 180–195; IV– gins weakly concave near eye platelets (Fig. 192). Leg. 1–6: 70–75, 60–66, 78–82, 105–110, 125– Dorsal shield elongate (ratio length/width 1.5–1.6), 138, 130–145. covering almost complete dorsum in unengorged

157 P. V. Tuzovsky

203, 204

200

203

200–202

201

204

202

Figs 200–204. Tiphys latipes, larva: 200 — leg I, 201 — leg II, 202 — leg III, 203 — claws of leg I, 204 — claws of leg III. Scale bars = 25. larva and bearing only 2 pairs of trichobothria (Fp, and developed only in their lateral parts (Fig. 193). Oi), setae Fch and Vi situated on subcutaneous ex- Anterior margin of coxal plates I concave and tention of dorsal plate. Setae Fch and Vi subequal nearly twice shorter than posterior margin. All and longer than Fp and Oi. Setae Oe, Hi, He, Sci, coxal setae (C1–C4) long and subequal in length. Sce and Li located on soft wrinkled membrane. Se- Base of setae C4 on coxal plate III situated closely tae Oe longest of all dorsal setae, setae Hi longer to medial end of suture line between coxal plates than setae He, setae Sci and Sce equal in length and II and III. Posteromedial apodemes of coxal plates slightly longer than Li. Lateral sides of coxae II+III I relatively large, posteromedial apodemes of cox- coming onto dorsal side and lying beside setae Oi al plates II and III slightly developed. Transverse and other hysterosomal setae located on interscutal muscle attachment scar and posteromedial margin membrane. Dorsal shield porous. of coxal plate III separated. Setae Ci, Si, Se, Le, Pi, Coxal plates I–II relatively small; anterior Pe situated on the soft interscutal membrane in the coxal plates separated from coxal plates II and III posterior part of the body. Setae Ci very long and which fused, suture line between them incomplete thicker than all ventral idiosomal setae and situat-

158 Water mites of the genus Tiphys ed on small bases; setae Si considerably shorter Ecology. Temporary ponds, sedge bogs, than setae Se. Coxal plates I–III porous. Urstigma sedge-sphagnum bogs, springs, lakes. Also in per- have a lateral position on border between coxae I manent waters. and II and provided with cups. Geographical distribution. Europe (Viets Excretory pore plate (Figs 194–196) slightly 1936 1956; Sokolow 1940; Lundblad 1968; K.O. wider than long or nearly as long as wide, ratio Viets 1978), Asia, Russia: Magadan Province. length/width = 0.8–1.0, both pairs of setae situated Tiphys (Tiphys) kamchatkaensis sp.n. in anterior half of plate, setae Ai shorter than Ae. Excretory pore open in posterior half of excretory Figs 205–213 pore plate. Material. Holotype male (IBIW 4601): Asia, Capitulum (Fig. 197) with wide base, anterior Russia, Kamchatka Province, Ust’-Kamchtsk Dis- hypostomal setae longer and thinner than posterior tr., Ust’-Kamchatsk settlement: sedge-sphagnum setae. Basal segments of chelicerae (Fig. 198) bog on right bank of the Kamchatka river, 20 km fused at their medial parts and tapering proximal- upstream of the settlement, 26 June 1983, P.V. Tu- ly; suture line well developed along their whole zovsky. Paratypes: 2 females collected in some length. Cheliceral stylet small, crescent-shaped. bog as holotype, 23 June 1983, P.V. Tuzovsky. Pedipalp trochanter (Fig. 199) short and with- Diagnosis. Female. Medial margin of coxal out setae; dorsodistal seta of femur and genu of plate IV longer than medial margin of coxal plate pedipalp short, thin and approximately equal in III. Posteromedial and posterolateral margins of size, proximal lateral genual seta long and very coxae IV forming obtuse angle with short apo- thick; tibia with 3 approximately subequal short, deme. Genital plates elongate and more or less tri- thin setae and large dorsodistal claw. Pedipalpal angular, with 3 acetabula and 7–9 short, fine setae tarsus with short solenidion, 2 long and 5 unequal each. Genital opening and genital plates approxi- simple setae. mately subequal in length, anterior and posterior All legs short and stout. General number of genital sclerites slightly developed and approxi- setae on legs I–III (Figs 200–202), except eu- mately equal in width. Pedipalpal genu with 3 se- pathidia, as follows: I–Leg. 1–5: 1, 7, 5 (s), 11 tae, external lateral seta shorter than dorsal margin (2s), 13 (s, ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 of segment, and only slightly longer than other (s, ac); III–Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). For- dorsal setae; base of external lateral seta situated mula of large setae on leg segments as follows: I– proximally to middle of segment; tibia with mod- Leg 1–5: 0, 3, 1, 2, 0; II–Leg 1–5: 0, 2, 2, 4, 0; erately long distolateral spine (2.5 times as short III–Leg 1–6: 0, 2, 2, 4, 0. Solenidion on genu of as tarsus), both ventral setae located on small tu- leg I slightly longer than distal solenidia on tibia I, bercles distally to middle of segment. Claws of which is approximately subequal in length; solen- legs I–III larger than claws of legs IV; both denti- idion on genu II shorter than proximal solenidion cles approximately subequal in length, external on tibia II, the latter slightly shorter than distal so- denticle thinly-pointed, internal denticle wide and lenidion on tibia II; solenidion on genu of leg III with rounded tip; claw lamella moderately devel- shorter than solenidion on tibia III. Central claw oped with convex ventral margin. on tarsi of all legs shorter than lateral claws (Figs Male. Coxae plates IV greatly expanded, 203–204); on tarsi of legs I and II empodia hook- fused with one another along whole medial mar- like, on tarsus III empodium only slightly bent. gins, abruptly depressed posteromedially. Posteri- Measurements (n = 10). Length of dorsal or margin of coxal plate IV nearly transverse. Me- shield 214–220, width 130–140; length of excre- dial margin of coxal plate IV 4.8 times longer than tory pore plate 16–19, width 19–23; length of ca- medial margin of coxal plate III. Suture line be- pitulum 67–70; length of basal segment of chelic- tween coxal plates III and IV complete, slightly era 67–70, length of cheliceral stylet 14–16; length oblique, directed posteromedially. Genital plates of medial margin of coxae I 64–70, length of me- triangular, fused medially. Anterior genital sclerite dial margin of coxae II+III 83–95; length of pedi- small transverse with two pairs of short, fine setae palpal segments (P-1–5): 5–6, 25–29, 18–21, 6–7, and fused with genital plates. Genu III with 3, tib- 4–5; length of legs segments: I–Leg. 1–5: 25–29, ia III with 4 long swimming setae. Genu IV great- 18–22, 22–26, 30–33, 40–43; II–Leg. 1–5: 31–33, ly expanded dorsoventrally with maximum height 26–29, 25–29, 38–41, 49–53; III–Leg. 1–5: 32–35, in the middle of segment, tibia IV nearly straight 26–29, 28–33, 41–46, 70–75. with lance-shaped and pointed dorsodistal seta

159 P. V. Tuzovsky

206–208 206

207

208 209 205

Figs 205–209. Tiphys kamchatkaensis, female: 205 — seta Fch, 206 — anteromedial and anterolateral dorsal plates, 207 — posteromedial plates, 208 — ventral view, 209 — genitoanal area. Scale bars: 205 =50, 206–208 = 100, 209 = 75. and with 10–12 long distal swimming setae. Tar- length of the segment, Fig. 211 (longer, Fig. 182), sus and tibia of leg IV nearly subequal in length. the posterior angle of coxal plates IV is obtuse, Claws of legs III asymmetrical: large claw with Fig. 208 (right or acute, Fig. 175). two denticles, external denticle larger than inter- Description. Female. Body oval; setae Fch nal one and provided with thin dorsal spur; small long, whip-like (Fig. 205), other idiosomal setae claw with two short stout denticles, external den- filiform. Integument soft with very fine strips. ticle larger than internal one and bearing very short Dorsum with relatively large anteromedial plates and thin spur. (Fig. 206), anterolateral and posteromedial plate- Differential diagnosis. The new species is lets (Fig. 207) little developed. Coxal plate I usu- closely related to T. latipes. The male of T. kam� ally with 4 lateral thin setae and 1 posteromedial chatkaensis sp. n. differs from the male of T. lati� thin seta, coxal plate II with 3 lateral thin setae, pes by the following characters (character states of coxal plate III with 1 medial seta and 3 lateral se- the male of T. latipes are given in parentheses): tae, coxal plate IV with 1 posteromedial seta and setae Hv and Pe are lying free in soft integument, 3–4 lateral setae (Fig. 208). Apodeme of coxal Fig. 214, (seta Hv fused with posterior margin of plate I moderately developed. Suture line between coxa II, seta Pe fused with posteromedial margin coxal plates III and IV complete and slightly of coxa IV, Fig. 185); genu IV is expanded dors- oblique, directed posteromedially. Medial margin oventrally with maximum height near middle of of coxal plate IV longer than medial margin of segment, Fig. 216 (in proximal portion, Fig. 187); coxal plate III. Posteromedial and posterolateral tibia IV with a relatively long lance-shaped dorso- margins of coxae IV forming obtuse angles with distal seta (Fig. 217) (with short peg-like seta, Fig. short apodemes. Sclerites bearing seta Hv close to 187), small claw of leg III has two unequal denti- posteromedial margin of coxa II, other idiosomal cles, external denticle is considerably larger than setae lying free in soft integument. internal one and with very short spur (Fig. 218) Genital plates elongate (Fig. 208), lying in an (with two subequal denticles, external denticle arc or nearly triangular (Fig. 209) with 3 acetabula without spur, Fig. 188). The female of T. kamchat� and 7–9 short and fine setae each. All acetabula kaensis sp. n. differ from the female of T. latipes small and approximately of equal size, and usually by the following characters (character states of the arranged in triangular shape; acetabula occupy ap- female of T. latipes are given in parentheses): the proximately about half area of genital plate; dis- genital plates is crescent, Fig. 208 (triangular, Figs tance between first and second acetabula usually 176–179), the lateral setae on P-3 shorter than more than distance between second and third ac-

160 Water mites of the genus Tiphys

210–211

211 210

212

213 Figs 210–213. Tiphys kamchatkaensis, female: 210 — chelicera, 211 — pedipalp, 212 — genu, tibia and tarsus of leg IV, 213 — claw. Scale bars: 210 = 100, 213 = 50. etabula. Genital opening and genital plates ap- rather long distolateral seta, 2 ventral setae and sev- proximately equal in length, anterior and posterior eral short and fine dorsal setae; both ventral setae genital sclerites slightly developed and approxi- located distally to middle of segment on small tu- mately equal in width. Excretory pore with very bercles. Distolateral seta of tibia approximately 2.5 small anterior sclerite. times shorter than tarsus. Pedipalpal tarsus tapering Capitulum with short anchoral projection, distally with 1 long proximal solenidion, 5 thin se- both pairs of setae equal in length. Basal segment tae and 4 thick distal unequal spines. of chelicera narrow and curved proximally, straight Legs II–IV with swimming setae; second pair distally (Fig. 210). Cheliceral stylet relatively of legs with short swimming setae, two posterior large, pointed. pairs of legs with long swimming ones (Fig. 212). Pedipalpal trochanter with a single short dor- Two swimming setae present on genu and tibia of sodistal seta, femur with 3 proximal and 2 dorsodis- legs II, 2–3 on genu III, 3–4 on genu IV and tibia tal setae (Fig. 211); ventral margin of femur and III, 5–6 on tibia IV. Claws of legs I–III larger, than genu straight. Pedipalpal genu with 3 short setae, claws of legs IV; both denticles approximately external lateral seta shorter than dorsal margin of subequal in length, external denticle thin and segment and only slightly longer than other dorsal pointed, internal denticle wide with rounded tip setae; base of external lateral seta situated proxi- (Fig. 213). Claw lamella moderately developed mally to middle of segment. Pedipalpal tibia with with slightly convex ventral margin.

161 P. V. Tuzovsky

215 216

214

214–217 217 218 Figs 214–218. Tiphys kamchatkaensis, male: 214— ventral view, 215 — genu, tibia and tarsus of leg III, 216 — telofemur and genu of leg IV, 217 — tibia and tarsus of leg IV, 218 — claws of leg III. Scale bars: 214–217 = 100, 218 =25. Measurements (n=2). Body length 600–650; plete, oblique, directed posteromedially. Genital length of medial margin of coxa III 50–55, length plates triangular, all acetabula approximately equal of medial margin of coxa IV 75–80; length of an- in size and occupying less than half the area of the teromedial dorsal plates 54–60, width 15–20; plates. Anterior genital sclerite small, with two length of genital plates 77–90, width 45–50; pairs of short, fine setae and fused with genital length/width of genital acetabula (ac. 1–3): 36– plates. Genital plates with 4–5 short, fine setae 43/23–30, 36–42/25–30, 35–42/30–36; width of each. Sclerites, bearing setae Hv, Pe and other idio- anterior genital sclerite 35–42; length of basal seg- somal setae lying free in soft integument. ment of chelicera 90–95, length of cheliceral sty- Genu III (Fig. 215) considerably shorter than let 30–33; length of pedipalpal segment (P-1–5): tibia III, tarsus III slightly longer than tibia III; 28–30, 72–78, 44–56, 77–85, 30–35; length of genu III with 3 and tibia III with 4 long swimming distolateral spine on tibia 20–21; length of leg seg- setae. ments: I–Leg. 1–6: 45–48, 65–72, 70–73, 83–90, Telofemur IV (Fig. 216) considerably shorter 97–102, 110–115; II–Leg. 1–6: 47–50, 65–72, than genu IV. Genu IV greatly expanded dors- 75–78, 90–95, 105–112, 115–125; III–Leg. 1–6: oventrally, with almost straight dorsal margin and 50–60, 70–75, 72–78, 88–95, 115–130, 115–120; large ventral hump with maximum height in mid- IV–Leg.1–6: 75–85, 80–85, 90–100, 130–140, dle of segment, bearing numerous dorsal and ven- 150–165, 140–150. trodistal setae, and several relatively short lateral Male. Dorsal surface, capitulum, chelicerae setae; dorsal setae very numerous and longer than and pedipalps as in female. Anterior apodemes ventrodistal setae. Tibia IV (Fig. 217) nearly rather large, extending beyond medial margin of straight, with numerous thin dorsal setae, lance- coxal plate III; posterior apodemes little developed shaped pointed dorsodistal seta, several rather (Fig. 214). Coxae plates IV greatly expanded, fused thick ventral setae, and 10–12 long distal swim- with one another along whole medial margins, ming setae. Tarsus and tibia IV nearly equal in abruptly depressed posteromedially, with 2 short length, with numerous short, fine setae and 4 rela- and fine setae anterior to depressed region on each tively long unequal thick, dorsal setae. plate. Posterior margin of coxal plate IV nearly Claw of legs I and II as in female. Claws of transverse. Medial margin of coxal plate IV 4.8 legs III (Fig. 218) asymmetrical, large claw with times longer than medial margin of coxal plate III. two rather long denticles, external denticle with a Suture line between coxal plates III and IV com- thin dorsal spur and longer than internal denticle;

162 Water mites of the genus Tiphys small claw with two short and stout denticles, ex- Larva. Dorsal shield elongate (ratio length/ ternal denticle with short and thin spur and larger width 1.5–1.6), covering almost complete dorsum than internal denticle. in unengorged larva. Setae Fch and Vi slightly lon- Measurements (n=1). Body length 510, length ger than Fp and Oi. Setae C4 slightly longer than of medial margin of coxa III 60, length of medial setae C1–C3 which approximately subequal in margin of coxa IV 29; length of anteromedial dor- length. Base of setae C4 on coxal plate III situated sal plates 440–45, width 10–15, length of genital slightly beneath medial end of suture line between plates 55, theirs general width 135; length/width coxal plates II and III. Transverse muscle attach- of genital acetabula (ac. 1–3): 30–36/27–34, 30– ment scar and posteromedial margins of coxal 36/25–30, 36–42/30–33; length of basal segment plates III fused to each other or slightly separated. of chelicera 145, length of cheliceral stylet 42; Dorsal shield and coxal plates I–III porous. Excre- length of pedipalpal segments (P-1–5): 35, 99, 60, tory pore plate nearly circular, ratio length/width = 108, 48; length of distolateral spine on tibia 18; 0.8–1.0, setae Ae longer than setae Ai; excretory length of leg segments: I–Leg. 1–6: 60, 78, 102, pore situated in posterior half of excretory pore 132, 155, 205; II–Leg. 1–6: 70, 78, 108, 145, 160, plate. Capitulum with wide base. Solenidion on 215; III–Leg. 1–6: 70, 60, 72, 90, 102, 170; IV– genu of leg I and solenidia on tibia I approximate- Leg. 1–6: 115, 95, 102, 170, 205, 210. ly subequal in length; solenidion on genu II short- Deutonymph. Unknown. er than solenidia on tibia II, both these solenidia Larva. Unknown. subequal, its bases well separated. Ecology. Sedge-sphagnum bog. Differential diagnosis. The new species is Geographical distribution. Asia, Russia: similar to T. scaurus. The female and larva of T. Kamchatka Province. kolymaensis sp. n. differ from the female and larva of T. scaurus by the following characters (charac- Tiphys (Tiphys) kolymaensis sp. n. ter states of the female and larva of T. scaurus are Figs 219–245 given in parentheses): female — the suture line Material. Holotype female (IBIW 2594): between coxal plates III and IV incomplete, oblit- Russia, Magadan Province, Tenka Distr., Agroba- erated medially, Fig. 222 (complete, Fig. 65), the sa: sedge-sphagnum bog on left bank of the Koly- pedipalpal genu with short lateral seta, Fig. 229 ma river, 1 km upstream of settlement, 26 June (with long lateral seta, Fig. 72), the lateral seta on 1979, P.V. Tuzovsky. Paratypes: 5 females and 5 P-3 located in the proximal half of segment (in larvae reared in laboratory (IBIW): 2 females, 21 distal half of segment); larva — the dorsal plate June 1979 and 3 females, 22 June 1979 collected and coxal plates punctate, Figs 233–234 (with re- in the same bog as holotype, P.V. Tuzovsky. ticulation, Figs 84–85), the base of seta C-4 close Diagnosis. Female. Dorsum with relatively to the suture line between coxal plates III and IV large anteromedial plates, anterolateral and pos- on each side (separated), seta C-4 is long, its tip teromedial platelets little developed. Suture line reaching posterior margin of Cx–III (short, its tip between coxal plates III and IV incomplete (oblit- not reaching posterior margin of Cx–III). erated medially), posteromedial and posterolateral Description. Female. Body oval; setae Fch margins of coxae IV forming obtuse angle with long, thin (Fig. 219), other idiosomal setae fili- short apodeme. Sclerites bearing setae Hv fused form. Integument soft with very fine strips. Dor- with posteromedial margin of coxa II, other idio- sum with relatively large anteromedial plates, somal setae lying free in soft integument. small anterolateral and posteromedial platelets Genital plates elongate and more or less trian- (Figs 220–221). Coxal plate I with 4 lateral thin gular with 3 acetabula and 12–13 short, fine setae setae and 1 posteromedial thin seta, coxal plate II each. Pedipalpal genu with 3(2) setae (sometimes with 3 lateral thin setae, coxal plate III with 1 me- dorsoproximal seta absent), external lateral seta dial seta and 2–3 lateral setae, coxal plate IV with shorter than dorsal margin of segment, and only 1–2 posteromedial setae and 4–5 lateral ones (Fig. slightly longer than other dorsal setae; base of ex- 222). Apodeme of coxal plate I moderately devel- ternal lateral seta situated proximally to middle of oped and usually in mature specimens not reach- segment. Pedipalpal tibia slender, both ventral se- ing to medial margin of coxal plate III. Suture line tae situated on small tubercles distally to middle between coxal plates III and IV incomplete (oblit- of segment; distolateral seta of tibia 3 times short- erated medially) oblique, directed posteromedial- er than tarsus. ly. Posteromedial and posterolateral margins of

163 P. V. Tuzovsky

219 220

221 220–220

227 226

222 224

223–227

223 225 Figs 219–227. Tiphys kolymaensis, female: 219 — seta Fch, 220 — anteromedial and anterolateral dorsal plates, 221 — posteromedial plates, 222 — ventral view, 223–225 — genital plate, 226–227 — excretory pore. Scale bars = 100. coxae IV forming obtuse angle, with short apo- base of external lateral seta located proximally to deme beneath this angle. Sclerites, bearing setae middle of segment. Pedipalpal tibia slender with Hv, fused with posteromedial margin of coxae II, relatively short distolateral peg-like spine (3 times other idiosomal setae situated on soft integument shorter than tarsus), 2 ventral setae, 2–3 thin dor- freely. sal ones and 1 dorsal seta proximally to middle of Genital plates more or less triangular, with 3 segment. Both ventral setae located on small tu- acetabula and 12–13 short, fine setae each. All ac- bercles distally to middle of tibia. Pedipalpal tar- etabula small, oval and approximately equal in sus tapering distally with single long proximal size, and usually arranged in an obtuse triangle; solenidion, five thin setae and four short and thick acetabula occupying about half the area of the distal unequal of different shape and length. genital plate; distance between first and second ac- Leg I (Fig. 230) without swimming setae; legs etabula and distance between second and third ac- II–IV with swimming setae, second pair of legs etabula usually less than length of one acetabulum with short swimming setae, two posterior pairs of (Figs. 223–225). Genital opening longer than gen- legs with long swimming setae (Fig. 231). Two or ital plates, anterior genital sclerites larger than three swimming setae present on genu and tibia II, posterior sclerite. Excretory pore with small ante- 3–4 on genu III and IV, 4–5 on tibia III, 5–7 on rior sclerite (Figs 226–227). tibia IV. Claws of legs I–III larger, than claws of Capitulum with rather long anchoral process- legs IV; both denticles approximately subequal in es, both pairs of setae equal in length. Basal seg- length, external denticle thin and pointed, internal ment of chelicera (Fig. 228) with large dorsal denticle wide with rounded tip (Fig. 232). Claw hump. Cheliceral stylet relatively large, pointed. lamella moderately developed with slightly con- Pedipalpal trochanter with a single short dor- vex ventral margin. sodistal seta, femur with 4 proximal and 2 unequal Measurements (n=6). Body length 670–1190, dorsodistal setae (Fig. 229). Ventral margin of fe- length of medial margin of coxa III+IV 125–140; mur straight, ventral margin of genu straight or length of anteromedial dorsal plates 60–65, width slightly concave. Pedipalpal genu with 3(2) setae 15–18, length of genital plates 115–130, width (occasionally dorsoproximal seta absent), external 55–66; length/width of genital acetabula (ac. 1–3): lateral seta shorter than dorsal margin of segment, 39–42/18–24, 36–42/27–30, 45–48/30–36; width and only slightly longer than other dorsal setae; of anterior genital sclerite 60–84; length of basal

164 Water mites of the genus Tiphys

228, 229, 232 232

229

228

230

231 230, 231

Figs 228–232. Tiphys kolymaensis, female: 228 — chelicera, 229 — pedipalp, 230 — genu, tibia and tarsus of leg I, 231 — genu, tibia and tarsus of leg IV, 232 — claw. Scale bars: 228, 229, 232 = 50, 230–231= 100. segment of chelicera 120–140, length of cheliceral twice as long as Li. Lateral sides of coxae II+III stylet 48–51; length of pedipalpal segment (P-1–5): passing onto dorsal side and almost extending to 36–42, 102–108, 60–66, 108–115, 50–54; length setae Oi and other hysterosomal setae located on of distolateral spine on tibia 17–18; length of leg interscutal membrane. Dorsal shield porous. segments: I–Leg. 1–6: 60–65, 70–78, 95–105, Coxal plates I–II relatively small; anterior 120–126, 135–145, 155–165; II–Leg. 1–6: 65–72, coxal plates separated from coxal plates II and III, 78–85, 105–110, 130–140, 150–157, 160–165; the latter fused to each other, suture line of coxal III–Leg. 1–6: 75–80, 75–85, 100–110, 125–135, plates II and III incomplete and developed only in 160–165, 160–165; IV–Leg. 1–6: 105–110, 95– lateral parts (Fig. 234). Anterior margin of coxal 105, 130–140, 185–195, 215–225, 200–210. plates I concave and considerably shorter than Larva. Body small and oval; dorsal shield posterior one. Setae C4 longer than setae C1–C3, truncated anteriorly, rounded posteriorly, with the latter approximately equal in length. Base of convex undulating lateral margins (Fig. 233). Dor- setae C4 on coxal plate III inserted closely to me- sal shield elongate (ratio length/width 1.5–1.6), dial portion of suture line between coxal plates II covering almost complete dorsum in unengorged and III. Posteromedial apodemes of coxal plates I larva and bearing 2 pairs of trichobothria (Fp, Oi) well developed, posteromedial apodemes of coxal and 2 pairs of simple setae (Fch, Vi). Setae Fch plates II rudimentary, and posteromedial apo- and Vi slightly longer and thicker than Fp and Oi. demes of coxal plates III small. Transverse muscle Setae Oe, Hi, He, Sci, Sce and Li located on the attachment scar and posteromedial margins of soft wrinkled membrane. Setae Oe longest of dor- coxal plates III fused or slightly separated. Setae sal setae, setae Hi longer than setae He, setae Sce Ci, Si, Se, Le, Pi, Pe situated on the soft interscutal

165 P. V. Tuzovsky

233 234 Figs 233–234. Tiphys kolymaensis, larva: 233 — dorsal view, 234 — ventral view. Scale bar: 233–234 = 50.

239 238, 239

238

236 240 235 237

235–237 Figs 235–240. Tiphys kolymaensis, larva: 235–237 — excretory pore plate, 238 — capitulum (ventral view), 239 — chelicerae, 240 — pedipalp. Scale bars: 235–237, 240 = 25, 238–239=50. membrane in posterior part of the body. Setae Ci 0.8–1.0. Setae Ai situated near anterior margin of very long and heaviest of all ventral idiosomal se- excretory pore plate, and shorter than Ae. Excre- tae, and situated on small bases; setae Si shortest tory pore situated posteriorly to middle of excre- than Se. Coxal plates I–III porous. Urstigma oc- tory pore plate. cupying lateral position on the border between Capitulum (Fig. 238) with wide base, anterior coxae I and II and provided with cups. hypostomal setae long and thin, posterior hypos- Excretory pore plate (Figs 235–237) wider tomal seta short spine-like. Basal segments of che- than long or nearly circular, ratio length/width = licerae (Fig. 239) fused by their medial parts and

166 Water mites of the genus Tiphys

241 244

241–243

244, 245

242

245

243

Figs 241–245. Tiphys kolymaensis, larva: 241 — leg I, 242 — leg II, 243 — leg III, 244 — claws of leg I, 245 — claws of leg III. Scale bars: 241–243 = 50, 244–245 = 25. well tapering proximally, with concave posterior mula of large setae on leg segments as follows: I– margins, lateral margins of these segments con- Leg 1–5: 0, 3, 1, 2, 0; II–Leg 1–5: 0, 3, 2, 4, 0; vex; suture line between segments distinctly de- III–Leg 1–6: 0, 2, 2, 4, 0. Solenidion on genu of veloped along their whole length. Cheliceral stylet leg I and solenidia on tibia I approximately sub- crescent-shaped, moderately in size. equal in length; solenidion on genu II shorter than Pedipalp trochanter (Fig. 240) short and with- solenidia on tibia II, both solenidia on tibia II equal out setae; dorsodistal setae of femur and genu of in length; solenidion on genu of leg III short, so- pedipalps thin, short and approximately equal in lenidion on tibia III relatively long. Central claw length, proximal lateral genual seta very long and on tarsi of all legs shorter than lateral claws (Fig. heavy; tibia with 3 approximately subequal thin 244–245); on tarsi of legs I and II empodia hook- setae and large dorsodistal claw. Pedipalpal tarsus like, empodium on tarsus III only slightly bent. very small, with 1 rather long solenidion, 2 long Measurements (n=5). Length of dorsal shield and 5 unequal short setae. 208–216, width 128–140; length of anal plate 20– Legs short and stout. The general number of 22, width 22–24; length of capitulum 80–87; setae on legs I–III (Figs 241–243), except eu- length of basal segment of chelicera 68–70, length pathidia, is as follows: I–Leg. 1–5: 1, 7, 5 (s), 11 of cheliceral stylet 15–16; length of medial margin (2s), 13 (s, ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 of the coxae I 54–61, length of medial margin of (s, ac); III–Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). For- the coxae II+III 89–97; length of pedipalpal seg-

167 P. V. Tuzovsky ments (P-1–5): 7–8, 26–28, 19–21, 7–8, 5–6; angular ventroproximal corner. Tibia IV hardly length of legs segments: I–Leg. 1–5: 27–29, 19– curved with a large distal projection bearing 8–12 20, 22–24, 28–32, 36–41; II–Leg. 1–5: 30–32, swimming setae and a single short peg-like seta. 22–25, 26–28, 34–38, 48–52; III–Leg. 1–5: 34–36, Tarsus of leg IV slightly longer than tibia, with 24–30, 28–30, 40–45, 54–60. numerous short and fine setae and 6–9 relatively Male. Unknown. long unequal thick dorsal setae. Both claws of Deutonymph. Unknown. legs III subequal, with three unequal denticles Ecology. Sedge-sphagnum bog. each. Geographical distribution. Asia, Russia: Deutonymph. Medial margin of coxae III Magadan Province. and medial margin of coxae III nearly subequal in length. Posteromedial angle of coxae IV moder- Tiphys (Tiphys) torris (Müller, 1776) ately developed with rather long apodeme. Scler- Figs 246–282 ites bearing setae Hv fused with posterior margin Material. 2 females, 1 male and 1 deutonymph: of coxae II. Genital plates with 2 acetabula and 3 Europe, Russia: Krasnodar territory, Seversk Dis- short, fine setae each. Posterior acetabulum larger tr., settlement Ubinskaya, brook, 9 May 1976, 18 than anterior one, distance between these aceta- females, 5 males, and 32 larvae reared in labora- bula usually equal or more than diameter of ante- tory (IBIW), Russia, Yaroslavl Province, Nekouz rior acetabulum on each plate. Postgenital and Distr., village Postyltsevo, temporary ponds, sedge pregenital sclerites well separated, genital sclerite bogs and brook near the settlement, June–Septem- considerably large than pregenital sclerite. Excre- ber 2000, 2006–2007, P.V. Tuzovsky; 1 female, 1 tory pore with very small anterior sclerite. Pedi- male: Spain, Aragon, Horta de San Juan, Nas de palps stocky, genu with 2 setae, external lateral Clinto, 17 April 1998, R. Gerecke. seta considerably longer than dorsodistal seta; Diagnosis. Female. Medial margin of coxal base of external lateral seta situated distally to plate III and medial margin of coxal plate IV sub- middle of segment; tibia short, with very long dis- equal in length or last slightly longer than medial tolateral seta (0.75–0.80 times as long as tarsus), margin of coxal plate III. Sclerites, bearing setae both ventral setae situated on small tubercles dis- Hv, fused with posteromedial margin of coxa II. tally to middle of segment. Genital plates more or less triangular with 3 ac- Larva. Dorsal shield elongate (ratio length/ etabula and 6–10 short and fine setae each; in ad- width 1.70–1.85), covering almost all dorsum in dition usually 2–3 genital setae situated on soft unengorged larva and bearing 4 pairs subequal se- integument between anterior genital sclerite and tae. Setae Oe longer of all dorsal setae, setae Sce anterior ends of genital plates on each side. Pedi- slightly longer than setae Li. Setae C1 shorter than palpal genu usually with 3(4) setae, external lat- coxal setae C2–C4, base of setae C4 on coxal plate eral seta 2–3 times longer than other dorsal setae; III situated closely to medial end of suture line be- base of external lateral seta situated distally to tween coxal plates II and III. Posteromedial apo- middle of segment; tibia with rather long distolat- demes of coxal plates I well developed, postero- eral seta (2.0–2.5 times shorter than tarsus), both medial apodemes of coxal plates II and III little ventral setae situated distally to middle of seg- developed. Transverse muscle attachment scar ment, external seta situated on a relatively large present. Setae Si considerably shorter than setae tubercle, internal seta on a small tubercle. Se. Dorsal shield and coxal plates I–III with retic- Male. Coxae plates IV greatly expanded, ulations. Excretory pore plate wider than long, touching each other along whole medial margins, anal setae situated near anterior margin of excre- abruptly depressed posteromedially. Medial mar- tory pore plate, setae Ai located slightly in front of gin of coxal plate IV approximately twice as long setae Ae, last longer than Ai; excretory pore open as medial margin of coxal plate III. Suture line in posterior half of excretory pore plate. Tibia of between coxal plates III and IV oblique, directed leg I with 10 setae, solenidion on genu of leg I and posteromedially. Genital plates triangular, fused distal solenidion on tibia I subequal in length, medially, with 4–5 short,fine setae each. Sclerites proximal solenidion on tibia I shorter than distal bearing setae Pe fused with posteromedial margin one; solenidion on genu of leg II and distal solen- of coxae IV; sclerites bearing setae Hv fused with idion on tibia II approximately subequal in length, posterior margin of coxae II. Genu of leg IV proximal solenidion on tibia II slightly shorter greatly expanded dorsoventrally with large rect- than distal solenidion.

168 Water mites of the genus Tiphys

246 246, 249 247

248 247–252

251 250

249 253 254

253, 254 252 Figs 246–254. Tiphys torris, female: 246 — seta Fch, 247 — anteromedial and anterolateral dorsal plates, 248 — posteromedial plates, 249 — ventral view, 250–252 — genital plate, 253–254 — excretory pore. Scale bars: 246–252 = 100, 253–254 = 50. Description. Female. Body oval; setae Fch each side. All acetabula approximately equal in long, whip-like (Fig. 246), other idiosomal setae size, and arranged in a triangle; acetabula occupy- filiform. Integument soft and striated. Dorsum ing about half or more than half of area of genital with relatively large anteromedial plates, small plate. Genital opening longer than genital plates, anterolateral and posteromedial platelets (Fig. anterior genital sclerite moderately developed and 247–248). Anteromedial plates elongate, postero- larger than posterior one. Excretory pore usually medial plates and anteromedial platelets oval. with anterior and posterior sclerites, last smaller Coxal plate I usually with 4(3) lateral thin setae than anterior one (Fig. 253), but sometimes poste- and 1 posteromedial thin seta, coxal plate II with rior sclerite absent (Fig. 254). 2–3 lateral thin setae, coxal plate III with 1 medial Capitulum with short anchoral projection, seta and 2–3 lateral setae, coxal plate IV with 1 both pairs of setae equal in length. Chelicera (Fig. medial seta and 3–5 lateral setae (Fig. 249). Apo- 255) with large basal segment, cheliceral stylet deme of coxal plate I moderately developed and moderately developed, pointed. usually not reaching medial margin of coxal plate Pedipalpal trochanter with single short dorso- III. Suture line between coxal plates III and IV distal seta, femur with 3–5 proximal and 2 dorso- complete and oblique, directed posteromedially. distal setae; ventral margin of femur straight or Medial margin of coxal plate III and medial mar- slightly convex, ventral margin of genu slightly gin of coxal plate IV subequal to or slightly longer concave (Fig. 256). Pedipalpal genu usually with than medial margin of coxal plate III. Posterome- 3 (sometimes 4) setae, external lateral seta 2–3 dial and posterolateral margins of coxae IV form- times longer than other dorsal setae; base of exter- ing right angle, with short apodeme. Sclerites nal lateral seta inserted distally to middle of seg- bearing setae Hv fused with posteromedial margin ment. Pedipalpal tibia slightly longer than femur, of coxa II, other idiosomal setae lying free in soft with rather long distolateral spine (2.0–2.5 times integument. shorter than tarsus), 2 ventral setae and several Genital plates (Figs 250–252) short, more or short and fine dorsal setae; both ventral setae less triangular, with 3 acetabula and 6–10 short, placed distally to middle of segment, external seta fine setae each. In addition, usually 2–3 genital se- situated on relatively large tubercle, internal seta tae situated on soft cuticle between anterior geni- on a small tubercle. Pedipalpal tarsus tapering distal sclerite and anterior ends of genital plates on tally, with single long proximal solenidion, with 5

169 P. V. Tuzovsky

255 257

256

258

Figs 255–258. Tiphys torris, female: 255 — chelicera, 256 — pedipalp, 257 — genu, tibia and tarsus of leg IV, 258 — claw. Scale bars: 255 = 50, 256–257 = 100, 258 = 30. thin setae and 4 thick distal setae of different shape 110, 95–125, 130–155, 145–180, 155–190; II– and length. Leg. 1–6: 55–75, 70–115, 95–130, 130–165, 150– Legs II–IV with swimming setae; legs II with 180, 160–205; III–Leg. 1–6: 55–85, 85–115, short swimming setae, two posterior pairs of legs 95–125, 145–165, 170–195, 170–195; IV–Leg. with long swimming ones (Fig. 257). Two swim- 1–6: 90–125, 95–140, 125–150, 180–205, 190– ming seta present on genu of legs II, 2–3 on tibia 230, 190–220. II, 3–4 on genu III and IV, 3–6 on tibia III and IV. Male. Dorsal surface, coxae of legs I–III, ca- Claws of legs I–III are larger than claws of legs pitulum, chelicera and pedipalps as in female. An- IV; external denticle longer than internal denticle terior apodemes rather large, extending beyond (Fig. 258). Claw lamella moderately developed medial margin of coxal plate III; posterior apo- with nearly straight ventral margin. demes small (Fig. 259). Coxae plates IV greatly Measurements (n=10). Body length 630–900, expanded, fused with one another along whole length of medial margin of coxa III 60–72, length their medial margins, abruptly depressed postero- of medial margin of coxa IV 70–90; length of an- medially, with 2 short and fine setae close together teromedial dorsal plates 36–45, width 18–25, anterior to depressed region on each plate. Pos- length of genital plates 90–105, width 65–100; teromedial margin of coxal plate IV concave, pos- length/width of genital acetabula (ac. 1–3): 24– terolateral margin convex. Medial margin of coxal 36/24–30, 29–36/24–30, 30–38/24–33; width of plate IV twice as long as medial margin of coxal anterior genital sclerite 75–105; length of basal plate III. Suture line between coxal plates III and segment of the chelicera 120–140, length of the IV oblique, directed posteromedially. Genital cheliceral stylet 48–60; length of pedipalpal seg- plates triangular, fused medially; all acetabula ap- ment (P-1–5): 30–40, 105–115, 65–75, 95–125, proximately subequal in sizes, and occupying 40–60; length of distolateral spine on tibia 21–24; about half of area of plates. Anterior genital scler- length of leg segments: I–Leg. 1–6: 55–65, 75– ite small, with two pairs of short and fine setae.

170 Water mites of the genus Tiphys

262 260, 261

260 259 261 Figs 259–262. Tiphys torris, male: 259— ventral view, 260 — telofemur, genu, tibia and tarsus of leg III, 261 — telofemur, genu, tibia and tarsus of leg IV, 262` — claws of leg III. Scale bars: 259–261 = 100, 262 =25.

Genital plates with 4–5 short and fine setae each. 38/24–32; length of basal segment of chelicera Sclerites, bearing setae Pe, fused with posterome- 105–140; length of cheliceral stylet 35–48; length dial margin of coxae IV; sclerites bearing setae Hv of pedipalpal segment (P-1–5): 25–40, 80–90, 40– fused with posterior margin of coxae II; other sc- 50, 80–90, 40–50; length of distolateral spine on lerites bearing ventral idiosomal setae situated on tibia 15–25; length of leg segments: I–Leg. 1–6: soft cuticle freely. 45–60, 70–95, 90–100, 120–125, 135–145, 190– Telofemur III (Fig. 260) short, genu III short- 215; II–Leg. 1–6: 45–65, 75–95, 95–105, 130– er than tibia III, and tarsus III slightly longer than 140, 150–165, 200–215; III–Leg. 1–6: 65–75, tibia III; telofemur III with 2, genu III with 3, and 85–95, 65–70, 75–85, 120–140, 160–170; IV– tibia III with 5–7 long swimming setae. Leg. 1–6: 95–120, 90–110, 65–85, 145–155, 195– Genu IV (Fig. 261) strongly expanded dors- 230, 230–245. oventrally, with distinct rectangular ventroproxi- Deutonymph. Dorsum with I pair anterome- mal corner, bearing rather numerous dorsal and dial dorsal plates only (Fig. 263), anterolateral and ventral unequal setae and several relatively short posteromedial platelets not developed. Setae Fch thick lateral pointed setae; dorsal setae thicker long, thin (Fig. 264). Coxal plate I usually with 3 than ventral ones. Tibia IV almost straight, with lateral thin setae and 1 posteromedial thin seta, large distal projection bearing 8–12 swimming se- coxal plate II with 1 lateral thin setae, coxal plate tae and single short peg-like seta, with a few dor- III with 1 medial seta and 1–2 lateral thin setae, sal and ventral setae. Tarsus IV slightly longer coxal plate IV with 2 medial setae and 1 lateral than tibia, with numerous short and fine setae and one (Fig. 265). Apodeme of coxal plate I moder- 6–9 unequal thick dorsal setae. Claw of legs I and ately developed, not extending beyond medial II similar to female with two denticles. Both claws margin of coxal plate III. Suture line between cox- of legs III (Fig. 262) subequal in shape and size, al plates III and IV slightly oblique, directed pos- with three unequal denticles each. teromedially. Medial margin of coxae III and me- Measurements (n=6). Body length 485–510, dial margin of coxae III nearly subequal in length. length of medial margin of coxa III 70–90, length Posteromedial angle of coxae IV moderately de- of medial margin of coxa IV 135–180; length of veloped, with rather long apodeme. Sclerites bear- anteromedial dorsal plates 30–28, theirs width ing setae Hv fused with posterior margin of coxae 18–24, length of genital plates 60–70, theirs gen- II; sclerites bearing setae Pe and other ventral id- eral width 140–200; length/width of genital aceta- iosomal setae lying free in soft integument. Geni- bula (ac. 1–3): 24–36/24–30, 30–36/24–30, 30– tal plates separated, oblique, directed anteromedi-

171 P. V. Tuzovsky

263 264

266

264, 267

265 263, 265, 266 267

Figs 263–267. Tiphys torris, deutonymph: 263 — anteromedial dorsal plates, 264 — seta Fch, 265 — ventral view, 266 — pedipalp, 267 — genu, tibia and tarsus of leg IV. Scale bars: 263, 265–266 = 50; 264, 267 = 100. ally, bearing 2 acetabula and 3 short and fine setae thin and pointed, and internal denticle wide with each. Posterior acetabulum larger than anterior rounded tip. Claw lamella well developed with one, distance between them usually equal or more convex ventral margin. than one diameter of anterior acetabulum on each Measurements, n=1. Body length 445, length plate. Postgenital and pregenital sclerites well sep- of medial margin of coxa III 36–45, length of me- arated, postgenital sclerite considerably large than dial margin of coxa IV 30–38; length of anterome- pregenital sclerite. Excretory pore with a very dial dorsal plates 12, width 9; length of genital small anterior sclerite. plates 60–66, theirs width 30; diameter of genital Pedipalpal trochanter without seta, femur acetabula (ac. 1–2): 15–17, 21–23; length of basal with 1 dorsoproximal seta and 2 dorsodistal setae segment of the chelicera 55, length of cheliceral (Fig. 266). Ventral margin of femur straight. Pedi- stylet 30; length of pedipalpal segment (P-1–5): palpal genu with 2 setae, external lateral seta con- 24, 60, 35, 57, 30; length of distolateral spine on siderably longer than dorsodistal seta; base of ex- tibia 24; length of leg segments: I–Leg. 1–6: 35, ternal lateral seta situated distally to middle of 42, 60, 72, 90, 115; II–Leg. 1–6: 39, 54, 63, 84, segment. Pedipalpal tibia with very long distolat- 100, 120; III–Leg. 1–6: 42, 54, 63, 84, 108, 120; eral spine, 2 ventral and 1–2 dorsodistal setae; IV–Leg. 1–6: 60, 54, 72, 93, 108, 125. both ventral setae situated on small tubercles dis- Larva. Body small and oval; dorsal shield tally to middle of segment; distolateral spine of truncated anteriorly, rounded posteriorly, lateral tibia 0.75–0.80 of length of tarsus. Pedipalpal tar- margins undulating convex, anterolateral margins sus with single long proximal solenidion, 3 thin weakly concave (Fig. 268). Dorsal shield elongate setae and 4 thick and short distal spines. (ratio length/width 1.70–1.85), covering almost Legs II–IV with swimming setae; second pair complete dorsum in unengorged larva and bearing of legs with short swimming setae, legs III–IV 4 pairs setae (Fch, Fp, Vi, Oi), all setae approxi- with long swimming setae (Fig. 267). Two swim- mately equal in length but trichobothria Fp and Oi ming setae present on genu of legs II–IV and tibia thinner than Fch and Vi. Setae Oe, Hi, He, Sci, Sce II, 3 on tibia III, 4 on tibia IV. Claws of legs I–III and Li located on the soft wrinkled interscutal larger, than claws of legs IV; both denticles ap- membrane. Setae Oe longer of all dorsal setae, se- proximately subequal in length, external denticle tae Sce and Li nearly equal in length. Lateral sides

172 Water mites of the genus Tiphys

268 268, 269 269 270

Figs 268–270. Tiphys torris, larva: 268 — dorsal view, 269 — ventral view, 270 — Leg I . Scale bars: 268–269 = 50, 270 =25. of coxae II+III passing onto dorsal side and almost Capitulum (Fig. 276) elongate, base of capit- extending to setae Oe and other hysterosomal set- ulum longer than wide; rostrum with long and thin ae located on interscutal membrane. Dorsal shield anterior hypostomal setae and short spine-like with reticulations. posterior ones. Basal segments of chelicerae (Fig. Coxal plates I–II relatively small; anterior 277) fused at their medial parts and tapering prox- coxal plates separated from coxal plates II and III imally; suture line well developed along their which fused each other, suture line between them whole length. Cheliceral stylet small and crescent incomplete and developed only in their lateral in shape. parts (Fig. 269). Anterior margin of coxal plates I Pedipalp trochanter (Fig. 277) short and with- concave and considerably shorter than posterior out setae; dorsodistal setae of femur and genu of one. Setae C1 shorter than other coxal setae (C2– pedipalps thin, short and approximately equal in C4), base of setae C4 on coxal plate III inserted size, proximal lateral genual seta very long and closely to medial end of suture line between coxal thick; tibia with 3 short subequal setae and large plates II and III. Posteromedial apodemes of coxal dorsodistal claw. Pedipalpal tarsus with 1 solen- plates I well developed, posteromedial apodemes idion, 2 long setae and 5 unequal short setae. of coxal plates II and III slightly developed. Trans- All legs relatively long (Figs 270, 279–280). verse muscle attachment scar situated near poste- General number of setae on legs I–III (Figs 258– rior margin of coxal plate III. Setae Ci, Si, Se, Le, 260), except eupathidia, is as follows: I–Leg. 1–5: Pi, Pe situated on soft interscutal membrane in the 1, 7, 5 (s), 10 (2s), 13 (s, ac); II–Leg. 1–5: 1, 7, 5 posterior part of body. Setae Ci very long and (s), 11 (2s), 13 (s, ac); III–Leg. 1–5: 1, 6, 5 (s), 10 thicker than all ventral idiosomal setae and situat- (s), 11 (ac). Formula of large setae on leg segments ed on small bases; setae Si shorter than setae Se. as follows: I–Leg 1–5: 0, 3, 1, 1, 0; II–Leg 1–5: 0, Coxal plates I–III with reticulations. Urstigma oc- 3, 2, 3, 0; III–Leg 1–6: 0, 2, 2, 3, 0. Proximal so- cupying lateral position on border between coxae lenidion on tibia I shorter than distal one, solenid- I and II and provided with cups. ion on genu I and distal solenidion on tibia I ap- Excretory pore plate wider than long (Figs proximately subequal in length; solenidion on 271–275). Both pairs of anal setae situated near genu II and distal solenidion on tibia II approxi- anterior margin of excretory pore plate, setae Ae mately subequal in length, proximal solenidion on longer than Ai. Excretory pore open in posterior tibia II slightly shorter than distal one; solenidion half of excretory pore plate. on genu III nearly twice as short as solenidion on

173 P. V. Tuzovsky

276–278

278 276 277

271

274

272

275 273 271–275 Figs 271–278. Tiphys torris, larva: 271–275 — excretory pore plate, 276 — capitulum (ventral view), 277 — chelicerae, 278 — pedipalp. Scale bars: 272–275 = 30, 276–278 =25.

279

281

282 280

Figs 279–282. Tiphys torris, larva: 279 — leg II, 280 — leg III, 281— claws of leg I, 282 — claws of leg III. Scale bar: 279–282 = 25.

174 Water mites of the genus Tiphys tibia III. Central claw and lateral claws on tarsi of ner (slightly expanded dorsoventrally and without all legs approximately subequal in size (Fig. 281– ventroproximal corner). 282); empodium on tarsi of legs I and II bent al- Ecology. Temporary ponds, sedge bogs, lakes, most at right angle, on tarsus III empodium only brooks. slightly bent. Geographical distribution. Europe (Viets Measurements (n=10). Length of dorsal shield 1936, 1956; Sokolow 1940; K.O. Viets 1978). 200–227, width 115–128; length of the anal plate 13–20, width 25–27; length of capitulum 60–65; Tiphys (Tiphys) samaraensis sp. n. length of basal segment of chelicera 57–60, length Figs 283–290 of cheliceral stylet 13–16; length of medial margin Material. Holotype female (IBIW 7796): Eu- of coxae I 60–67, length of medial margin of cox- rope, Russia, Samara Province, National natural ae II+III 90–96; length of pedipalpal segments Park “Samara Luka”, village Koltsovo, sedge bog, (P-1–5): 4–7, 22–26, 16–19, 6–7; length of the 29 June 1981, P.V. Tuzovsky. legs segments: I–Leg. 1–5: 25–29, 25–27, 28–30, Diagnosis. Female. Suture line between cox- 35–38, 47–50; II–Leg. 1–5: 28–32, 28–30, 28–32, al plates III and IV nearly transverse. Medial mar- 40–43, 57–64; III–Leg. 1–5: 28–32, 32–35, 41–45, gin of coxal plate IV approximately 3 times longer 57–67. than medial margin of coxae III. Posteromedial Remarks. Tibia of leg I in larva T. torris with margin and posterolateral margin of coxae IV 10 setae; in contrast, tibia I at larvae in majority of forming large right angle with distinct apodeme species of the genus Tiphys with 11setae. beneath this angle. Sclerites bearing setae Hv Ecology. Temporary ponds, sedge bogs, lakes, fused with posterior margins of coxal plates II. brooks. Also in permanent waters. Genital plates elongate, roughly crescent, with 3 Geographical distribution. Europe, Asia, acetabula and 8–9 thin short setae each, third ac- North America: Alaska (Sokolow 1940; Viets etabulum larger than two other acetabula. Genital 1956; Cook 1956; Lundblad 1968; K.O. Viets opening slightly longer than genital plates. Ante- 1978). rior genital sclerite lesser than posterior one. Ex- cretory pore non-sclerotized. Pedipalps slender, Tiphys (Tiphys) bullatus (Thor, 1899) genu with 3 setae, external lateral seta longer than I was not able to examine any specimens of T. two other setae, base of external seta situated in bullatus. The diagnosis of this species is based on middle of segment. Pedipalpal tibia long, slender reference data (Thor 1899; K. Viets 1936; Sokolow with rather large distolateral spine (2.5 times 1940). shorter than tarsus), both ventral setae situated on Diagnosis. Male. External lateral seta on pe- very small tubercles near distal end of segment. dipalpal genu situated in middle of segment; ven- Claws of legs I–IV subequal in sizes. tral tubercles on pedipalpal tibia poorly developed, Differential diagnosis. The female of the new distal spine on tibia less than 0.25 length of tarsus; species is similar to the female of T. ornatus, from genu IV without ventroproximal corner, tibia IV which it differs by the following characters (charac- with distal protrusion. ter states of the female of T. ornatus are given in Female. Idiosoma very similar to that in fe- parentheses): coxal plate IV has right posteromedial male of T. torris. angle, Fig. 284 (with acute angle, Fig. 20), external Remarks. This species is similar to T. torris. setae of pedipalpal genu are situated in the middle The adults mites of T. bullatus differ from T. torris of segment, Fig. 287 (near distal end, Fig. 27), both by the following characters (characters states of T. ventral seta are inserted near distal end of pedipal- bullatus indicated in parentheses are from Thor pal tibia (near middle of the segment), claws of legs 1899; K. Viets 1936; Sokolow 1940): in both sex- I–IV are subequal in sizes (claws of legs I–III con- es, the base of external lateral seta on the pedipal- siderably larger than claws of legs IV). Claw of legs pal genu situated distally to the middle of segment I–IV are subequal in size too in the female of T. (in the middle of this segment), external seta on ensifer, but in this species the medial margins of pedipalpal tibia situated on a large tubercle (small coxal plates III and IV are nearly equal in length tubercle), distolateral spine of tibia 2.0–2.5 times (Fig. 344), pedipalpal tibia is without ventral tuber- shorter than tarsus (approximately 4 times shorter cles (Fig. 348), genital plates are wide and with than tarsus); in male, genu IV is greatly expanded 13–17 setae each. In contrast, in the female T. sa� dorsoventrally, with distinct ventroproximal cor- maraensis sp. n., the medial margin of coxal plate

175 P. V. Tuzovsky

283 283, 286, 287

284

286

284, 285

285 287

Figs 283–287. Tiphys samaraensis, female: 283 — seta Fch, 284 — ventral view, 285 — excretory pore, 286 — chelicera, 287 — pedipalp. Scale bars = 100.

IV is 3 times longer than the medial margin of cox- Capitulum with short anchoral projection, al plate III, pedipalpal tibia bears ventral tubercles, both pairs of setae equal in length. Chelicera with and genital plates are narrow with 8–9 setae each. large basal segment and relatively small pointed Description. Female. Body oval; setae Fch stylet (Fig. 286). long, thin (Fig. 283), other idiosomal setae hair- Pedipalps slender (Fig. 287). Pedipalpal tro- like. Integument soft with very fine strips. Dorsal chanter with single short dorsodistal seta, femur plates not found. Coxal plate I with 1 posterome- with 4 proximal and 2 dorsodistal setae. Ventral dial thin seta, coxal plate II with 3 lateral thin set- margin of femur straight or slightly convex, ven- ae, coxal plate III with 1 medial seta and 1 lateral tral margin of genu concave. Pedipalpal genu with one, coxal plate IV with 2 medial setae and 5 lat- 3 setae, external lateral seta longer than two other eral ones (Fig. 284). Apodeme beneath postero- setae; base of external seta situated near middle of medial medial edge of coxal plate I moderately segment. Pedipalpal tibia long, slender with rather developed. Suture line between coxal plates III large distolateral spine (2.5 times shorter than tar- and IV nearly transverse, medial margin of coxal sus), 2 ventral and several thin dorsal setae and plate IV 3 times as long as medial margin of coxae single thick dorsal spine near middle of segment; III. Posteromedial margin and posterolateral mar- both ventral setae inserted on very small tubercles gin of coxae IV forming right angle with distinct near distal end of segment. Pedipalpal tarsus with apodeme beneath this angle. Sclerites bearing set- single long proximal solenidion, 5 thin setae and 4 ae Hv fused with posterior margins of coxal plates thick distal unequal spines. II; other idiosomal setae situated on soft cuticle All legs long, thin (Figs 288–289). Legs II–IV freely. Genital plates elongate, roughly crescent, with swimming setae; second pair of legs with with 3 acetabula and 8–9 thin short setae each. short swimming setae, two posterior pairs of legs Third acetabulum larger than two other acetabula; with long swimming ones. One swimming seta or distance between first and second acetabula, and two swimming setae present on genu of legs II, distance between second and third acetabula lesser 2–3 on genu II, 3–5 on tibia II and III, 5 on genu than diameter one acetabulum. Genital opening IV, 9–10 on tibia IV. Claws of legs I–III and claw slightly longer than genital plates. Anterior genital of legs IV subequal in sizes. Internal denticle of sclerite small, and not larger than posterior one. claw of legs I–IV rather wide with rounded tip, Excretory pore unsclerotized (Fig. 285). external denticle thin and pointed (Fig. 290).

176 Water mites of the genus Tiphys

288

289

290

Figs 288–290. Tiphys samaraensis, female: 288 — genu, tibia and tarsus of leg I, 289 — genu, tibia and tarsus of leg IV, 290 — claw. Scale bars: 288–289 = 100, 290 = 50.

Measurements (n=1). Body length 810, length coxal plate III. Posterior margins of coxal plates of medial margin of coxa III 42–48, length of me- IV slightly concave. Genital plates triangular and dial margin of coxa IV 120–125; length of genital fused with anterior genital sclerite; all acetabula plates 145–150, width 55; length/width of genital relatively large, approximately subequal in sizes. acetabula (ac. 1–3): 47–49/27–30, 36–42/24–30, Sclerites bearing setae Hv fused with posterior 48–50/41–43; width of anterior genital sclerite 30; margins of coxal plates II; sclerites bearing setae length of basal segment of chelicera 160; length of Pe fused with posterior margins of coxal plates IV. cheliceral stylet 65; length of pedipalpal segment Pedipalpal genu with 3 setae, external lateral seta (P-1–5): 50, 132, 72, 155, 65; length of distolat- considerably longer than two other setae; base of eral spine of tibia 27; length of leg segments: I– external lateral seta situated proximally to middle Leg. 1–6: 120, 155, 170, 260, 310, 365; II–Leg. of segment; tibia with short distolateral spine (3 1–6: 130, 170, 230, 285, 335, 390; III–Leg. 1–6: times as short as tarsus), both ventral setae situated 125, 165, 215, 270, 340, ?; IV–Leg. 1–6: 185, 220, on small tubercles distally to middle of segment. 250, 335, 400, 385. Genu IV slightly expanded distally, with slightly Male. Unknown. convex dorsal margin and concave ventral one, Deutonymph. Unknown. bearing 6 relatively short thick setae and rather Larva. Unknown. numerous long distal ones; tibia IV slightly curved, Ecology. Sedgy bog. with concave dorsal and convex ventral sides, with Geographical distribution. Europe, Russia: single short fan-like seta, 6 short sword-like vent- Yaroslavl Province. rolateral setae, and 7 long distal swimming setae. Claws of legs III asymmetric, large claw with 2 Tiphys (Tiphys) yaroslavlensis sp. n. thick curved denticles, external denticle with short Figs 291–299 and fine spur; small claw with 2 unequal straight Material. Holotype male (IBIW 9366): Rus- pointed denticles. sia, Yaroslavl Province, Nekouz Distr., village Differential diagnosis. The male of the new Pogorelka, sedge-sphagnum bog, 18 May 2006, species is similar to the male of T. torris, from P.V. Tuzovsky. which differs by the following characters (charac- Diagnosis. Male. Medial margin of coxal ter states of the female of T. torris are given in plate IV 3.3 times longer than medial margin of parentheses): base of external lateral seta on pedi-

177 P. V. Tuzovsky

292 292, 293 291

294

291, 294, 295

295 293

Figs 291–295. Tiphys yaroslavlensis sp. n., male: 291 — seta Fch, 292 — anteromedial and anterolateral dorsal plates, 293 — ventral view, 294 — chelicera, 295 — pedipalp. Scale bars: 291, 294–295 = 50, 292–293 = 100. palpal genu situated proximally to the middle of short lateral setae. Medial margin of coxal plate segment, Fig. 295 (distally to the middle of seg- IV 3.3 times longer than medial margin of coxal ment, Fig. 256); genu IV slightly expanded dis- plate III. Posterior margins of coxal plates IV tally, without ventroproximal corner, the height of slightly concave. anterior end of this segment is lesser than the heigh Genital plates triangular and fused with ante- of distal end of telofemur, Fig. 297 (genu of leg IV rior genital sclerite; all acetabula relatively larger, is greatly expanded dorsoventrally with well de- approximately subequal in sizes, and occupy more veloped ventroproximal corner, the height of ante- than one half of surface of plates. Sclerites bearing rior end of genu is larger than the high of distal setae Hv fused with posterior margins of coxal end of telofemur, Fig. 261); claws of legs III are plates II; sclerites bearing setae Pe fused with pos- asymmetrical, the large claw has 2 thick curved terior margins of coxal plates IV, other ventral id- denticles, the external denticle with short and fine iosomal setae situated on soft cuticle freely. spur; the small claw has 2 unequal straight pointed Capitulum with short anchoral projection, denticles, Fig. 299 (both claws of legs III are sub- both pairs of setae equal in length. Chelicera with equal with three unequal denticles each, Fig. large basal segment and relatively small stylet 262). (Fig. 294). Description. Male. Body oval; setae Fch Pedipalps (Fig. 295) stout. Pedipalpal tro- long, whip-like (Fig. 291), other idiosomal setae chanter with single short dorsodistal seta, femur filiform. Integument soft striated. Dorsum with with 3 proximal and 2 dorsodistal unequal setae. relatively large anteromedial and small anterolat- Ventral margin of femur straight, ventral margin eral platelets (Fig. 292), posteromedial platelets of genu slightly concave. Pedipalpal genu with 3 not developed; anteromedial plates elongate, ante- setae, external lateral seta considerably longer rolateral platelets oval. Anterior apodemes extend- than two other setae; base of external lateral seta ing beyond medial edge of coxal plate III (Fig. attached proximally to middle of segment. Pedi- 293). Coxae plates IV greatly expanded, fused palpal tibia with relatively short distolateral spine with one another along whole medial edges. Coxal (3 times shorter than tarsus), 2 ventral and several plate I with 4 lateral thin setae and 1 posterome- dorsal setae; both ventral setae located on small dial thin seta, coxal plate II with 3 lateral thin set- tubercles distally to middle of segment. Pedipalpal ae, coxal plate III with 3 lateral setae and 1 medial tarsus with single long proximal solenidion, with seta, coxal plate IV with 2 posteromedial and 4 5 thin setae and 4 thick distal unequal spines.

178 Water mites of the genus Tiphys

296, 297

297

299

296 298

Figs 296–299. Tiphys yaroslavlensis sp. n., male: 296 — genu, tibia and tarsus of leg III, 297 — leg IV, 298 — claws of leg II, 299 — claws of leg III. Scale bars: 296–297 = 100, 298 = 50, 299 = 25.

Genu III (Fig. 296) short, tarsus III longer Measurements, n=1. Body length 580, length than tibia III; genu III with 3, and tibia III with 6 of medial margin of coxa III 78, length of medial long swimming setae. Genu of leg IV (Fig. 297) margin of coxa IV 260; length of anteromedial nearly not expanded dorsoventrally relatively te- dorsal plates 55, theirs width 20; length of genital lofemur IV, with slightly convex dorsal margin plates 96, their general width 240; length/width of and straight ventral margin, bearing a few rela- genital acetabula (ac. 1–3): 36–42/42–48, 42/48, tively short thick dorsolateral setae and 7 rather 42/48; length of basal segment of the chelicera long distal ones. Tibia IV slightly curved with 125, length of the cheliceral stylet 51; length of concave dorsal and convex ventral sides, with a pedipalpal segment (P-1–5): 42, 110, 60, 120, 50; few long dorsal setae, single short fan-like seta, 6 length of distolateral spine on tibia 21; length of short sword-like ventrolateral setae, and 7 long leg segments: I–Leg. 1–6: 72, 95, 120, 155, 170, distal swimming setae. Tibia and tarsus IV sub- 240; II–Leg. 1–6: 72, 95, 120, 160, 185, 250; III– equal in length, last slender, slightly tapering dis- Leg. 1–6: 72, 90, 90, 115, 168, 198; IV–Leg. 1–6: tally with numerous short and fine setae and 7 138, 120, 120, 170, 235, 235. relatively long thick unequal setae. Ecology. Sedgy bog. Claws of legs I–II are larger, than claws of Geographical distribution. Europe, Russia: legs IV. Internal denticle of claw of legs I–II rather Yaroslavl Province. wide with rounded tip, external denticle thin and Tiphys (Tiphys?) minutiporus sp. n. pointed (Fig. 298). Claws of legs III asymmetric (Fig. 299); large claw with 2 thick curved denti- Figs 300–323 cles, external denticle with short and fine spur; Material. Holotype female (IBIW 4475): small claw with 2 straight pointed denticles, exter- Asia, Russia, Magadan Province, Tenka Distr., vil- nal denticle longer and thicker than internal den- lage Agrobasa: sedge-sphagnum bog on the left ticle. bank of the Kolyma river, 1 km upstream of the

179 P. V. Tuzovsky

301 300

302 301, 302, 305

303

305 304

Figs 300–305. Tiphys minutiporus, female: 300 — seta Fch, 301 — anteromedial and anterolateral dorsal plates, 302 — posteromedial plates, 303 — ventral view, 304 — genital field, 305 — excretory pore. Scale bars: 300 = 50, 301–305 = 100. settlement, 18 July 1983, P.V. Tuzovsky. Para- muscle attachment scar and posteromedial margins types: 4 larvae reared in laboratory. of coxal plates III slightly separated. Dorsal shield Diagnosis. Female. Medial margin of coxal and coxal plates I–III porous. Excretory pore plate plate IV slightly longer than medial margin of wider than long, ratio length/width = 0.81–0.85; coxal plate III. Posteromedial and posterolateral both pairs of anal setae subequal in length. Excre- margins of coxae IV forming right or blunt angle tory pore situated distally to middle of excretory with very short apodeme beneath this angle. Scler- pore plate. Solenidion on genu of leg I and solenidia ites bearing setae Hv fused with posteromedial on tibia I approximately subequal in length; solen- margin of coxa II. Genital plates triangular with 3 idion on genu II shorter than distal solenidion on acetabula and 10–12 short and fine setae each. All tibia II, bases of solenidia on tibia II separated, dis- acetabula very small and approximately subequal tal solenidion longer than proximal one. in size, arranged in triangular; distance between Differential diagnosis. The new species is first and third acetabula equal nearly 3 diameters similar to T. scaurellus. The female and larva of T. of one acetabulum, distance between second and minutiporus sp. n. differ from the female and larva third acetabula equal 2 diameters of one acetabu- of T. scaurellus by the following characters (char- lum Pedipalpal genu with 3 short approximately acter states of the female and larva of T. scaurellus subequal in length setae, external lateral seta twice are given in parentheses): female — the genital as short as dorsal margin of segment; base of ex- plates are broad, length/width ratio 1.4, Figs 303– ternal lateral seta situated in middle of segment. 304 (elongate, length/width ratio 1.7–2.0, Figs Tibia slender with short distolateral spine (3 times 102–107), the genital acetabula are minute, ac.1–3: as short as tarsus), both ventral setae situated near 16–18, 17–19, 21–23 (relatively large, ac. 1–3: middle of pedipalpal on small tubercles. 24–27, 24–25, 25–30), pedipalpal genu with 3 Larva. Dorsal shield elongate (ratio length/ short subequal setae, Fig. 306 (with 2 unequal se- width 1.55–1.60), covering almost all dorsum in un- tae, Fig. 109); larva — the dorsal plate is elongate, engorged larva. Setae Fch and Vi slightly longer and length/width ratio 1.5–1.6, Fig. 310 (broad, length/ thicker than Fp and Oi. All coxal setae (C1–C4) ap- width ratio, 1.15–1.20, Fig. 124), the tmas pres- proximately subequal in length. Base of setae C4 on ent, Fig. 311 (absent, Fig. 125), the bases of setae coxal plate III situated closely to medial end of su- C4 are placed near the suture line between the ture line between coxal plates II and III. Transverse coxal plates II and III (well separated), the excre-

180 Water mites of the genus Tiphys

306

307

309

308

Figs 306–309. Tiphys minutiporus, female: 306 — pedipalp, 307 — tibia and tarsus of leg II, 308 — genu, tibia and tarsus of leg IV, 309 — claw. Scale bars: 306–308 = 100, 309 = 50. tory pore open in posterior half of the excretory Genital plates triangular, with 3 acetabula and pore plate, Figs 312–315 (in anterior half of the 10–12 short and fine setae each (Fig. 304). All ac- excretory pore plate, Figs 126–127. etabula very small and subequal in size, arranged Description. Female. Body oval; setae Fch in triangular; acetabula occupy considerably lesser long, whip-like (Fig. 300), other idiosomal setae than one half of surface of genital plate; distance filiform. Integument soft with very fine strips. between first and third acetabula equal to nearly 3 Dorsum with relatively large anteromedial plates, diameters of one acetabulum, distance between small anterolateral and posteromedial platelets second and third acetabula equal to 2 diameters of slightly developed (Figs 301–302). Coxal plate I one acetabulum. Genital opening longer than gen- with 3 lateral thin setae and 1 posteromedial thin ital plates, anterior genital sclerites considerably seta, coxal plate II with 3 lateral thin setae, coxal wider than posterior one. Excretory pore with an- plate III with 1 medial seta and 2–3 lateral setae, terior sclerite (Fig. 305). coxal plate IV with 1 medial seta and 3–4 lateral Capitulum with anchoral processes, both pairs ones (Fig. 303). Apodeme beneath posteromedial of setae equal in length. Basal segment of chelic- medial edge of coxal plate I moderately developed era with large dorsal hump. Cheliceral stylet rela- and slightly extending beyond medial edge of cox- tively large, pointed. Pedipalpal trochanter with al plate III. Suture line between coxal plates III single short dorsodistal seta, femur with 3 proxi- and IV complete oblique, directed posteromedial- mal and 2 dorsodistal short setae, ventral margin ly. Medial margin of coxal plate IV slightly longer of femur and genu straight (Fig. 306). Pedipalpal than medial margin of coxal plate III. Posterome- genu with 3 approximately subequal in length se- dial and posterolateral margins of coxae IV form- tae, both proximal setae twice as short as dorsal ing right or obtuse angle with very short apodeme margin of segment; base of external lateral seta at- beneath this angle. Sclerites, bearing setae Hv, tached near middle of segment. Pedipalpal tibia fused with posteromedial margin of coxa II, other slender with short distolateral peg-like spine (3 idiosomal setae situated on soft cuticle freely. times as short as tarsus), 2 ventral setae and sev-

181 P. V. Tuzovsky

310 311 Figs 310–311. Tiphys minutiporus, larva: 310 — dorsal view, 311 — ventral view. Scale bar: 310–311 = 100. eral short and fine dorsal ones. Both ventral setae III–Leg. 1–6: 85, 90, 120, 150, 185, 185; IV–Leg. situated on small tubercles near middle of seg- 1–6: 120, 105, 150, 205, 235, 210. ment. Pedipalpal tarsus tapering distally with sin- Larva. Body small and oval; dorsal shield gle long proximal solenidion, with five thin setae truncate anteriorly, rounded posteriorly, with con- and four short, thick unequal distal spines. vex lateral edges, and weakly concave anterolater- Legs I without swimming setae, tibia and tar- ally (Fig. 310). Dorsal shield elongate (ratio sus cylindrical (Fig. 307). Legs II–IV with swim- length/width 1.55–1.60), covering almost all dor- ming setae; second pair of legs with short swim- sum in unengorged larva and bearing 4 pairs of ming setae, two posterior pairs of legs with long setae: Fch, Fp, Oi, Vi. Setae Fch and Vi slightly swimming ones (Fig. 308). Two swimming setae longer and thicker than Fp and Oi. Setae Oe, Hi, present on genu and tibia II, 3 on genu III and tib- He, Sci, Sce and Li located on the soft wrinkled ia II, 4–5 on tibia III, 5 on genu IV, 7 on tibia IV. membrane. Setae Oe and He nearly subequal in Claws of legs I–III larger, than claws of legs IV; length and slightly longer than all dorsal setae, se- both denticles approximately equal in length, in- tae Hi longer than setae He, setae Sce longer Li. ternal denticle wide with rounded tip, external Lateral sides of coxae II+III coming onto dorsal denticle thin and pointed; claw lamella moderately side and almost extending to setae Oi and other developed, with slightly convex ventral margin hysterosomal setae located on interscutal mem- (Fig. 309). brane. Dorsal shield porous. Measurements, µm. Body length 1005, length Coxal plates I–II relatively small; anterior of medial margin of coxa III 60, length of medial coxal plates separated from coxal plates II and III margin of coxa IV 75; length of anteromedial dor- which fused to each other, suture line between sal plates 72, width 30, length of genital plates them incomplete and developed only in their lat- 120, width 85; diameter of genital acetabula (ac. eral parts (Fig. 311). Anterior margin of coxal 1–3): 16–18, 17–19, 21–23; width of anterior gen- plates I slightly concave and considerably shorter ital sclerite 102; length of basal segment of the than posterior one. All coxal setae (C1–C4) sub- chelicera 205, length of cheliceral stylet 63; length equal in length, but C4 thicker than C1–C3. Base of pedipalpal segment (P-1–5): 42, 115, 72, 132, of setae C4 on coxal plate III attached closely to 57; length of distolateral spine of tibia 23; length medial end of suture line between coxal plates II of leg segments: I–Leg. 1–6: 78, 78, 125, 150, and III. Posteromedial apodemes of coxal plates I 175, 185; II–Leg. 1–6: 78, 85, 120, 155, 180, 200; well developed, posteromedial apodemes of coxal

182 Water mites of the genus Tiphys

316–318 318

317 316

312

313 314 315

312–315 Figs 312–318. Tiphys minutiporus, larva: 312–315 — excretory pore plate, 316 — capitulum (ventral view), 317 — chelicerae, 318 — pedipalp. Scale bars: 312–315 = 25, 316–318 =20. plates II and III small. Transverse muscle attach- Pedipalpal trochanter (Fig. 317) short and ment scar and posteromedial margins of coxal without setae; distolateral seta of femur and genu plates III slightly separated. Setae Ci, Si, Se, Le, of pedipalps thin, short and approximately equal Pi, Pe situated on the soft interscutal membrane in in size, proximal lateral genual seta very long and the posterior part of the body. Setae Ci longer and heavy; tibia with 3 approximately subequal short, thicker than all ventral idiosomal setae and situat- thin setae and large dorsodistal claw. Pedipalpal ed on small bases; setae Si considerably shortest tarsus with one solenidion, 2 long and 5 unequal than setae Se. Coxal plates I–III porous. Urstigma thin setae. occupying lateral position on the border between All legs short and stout. The general number of coxae I and II and provided with cups. setae on legs I–III (Figs 319–321), except eupathid- Excretory pore plate (Figs 312–315) wider ia, as follows: I–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, than long, ratio length/width = 0.80–0.85. Both ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); III– pairs of anal setae approximately equal in length. Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). Formula of large Setae Ai located near anterior margin of excretory setae on leg segments as follows: I–Leg 1–5: 0, 3, 1, pore plate, setae Ae located near middle of lateral 2, 0; II–Leg 1–5: 0, 3, 2, 4, 0; III–Leg 1–6: 0, 2, 2, margins of the plate. Excretory pore located dis- 4, 0. Solenidion on genu of leg I and solenidia on tally to middle of excretory pore plate. tibia I approximately equal in length; solenidion on Capitulum (Fig. 316) with wide base, anterior genu II and proximal solenidion on tibia subequal hypostomal setae long, thin, posterior hypostomal and shorter than distal solenidion on tibia II, bases setae short, spine-like. Basal segments of chelic- of solenidia on tibia II well separated; solenidion on erae (Fig. 317) fused with one another along whole genu of leg III twice shorter than solenidion on tibia medial edges and tapering proximally, lateral mar- III. Central claw on tarsi of all legs shorter than lat- gins of these segments convex, posterior margin eral claws; empodium on tarsi of legs I and II hook- with small median incision. Cheliceral stylet small like (Fig. 322), empodium on tarsus III only slightly and crescent. bent (Fig. 323).

183 P. V. Tuzovsky

319

322

319–321

323

320

321

Figs 319–323. Tiphys minutiporus, larva: 319 — leg I, 320 — leg II, 321 — leg III, 322— claws of leg I, 323 — claws of leg III. Scale bars: 319–321 = 50, 322–323 = 25.

Measurements, n=4. Length of dorsal shield Tiphys (Tiphys?) chaunensis sp. n. 215–220, its width 135–140; length of anal plate Figs 324–340 18–20, its width 22–24; length of capitulum 78– 82; length of basal segment of chelicera 68–72, Material. Holotype larva (IBIW 4509): Asia, length of cheliceral stylet 15–16; length of medial Russia, Magadan Province, Chaun Distr., thermo- margin of the coxae I 57–64, length of medial carstic lake near mountain Neytlin, 9 August 1982, margin of the coxae II+III 83–95; length of pedi- P.V. Tuzovsky. Paratypes: 14 larvae reared in labo- palpal segments (P-1–5): 6–8, 30–32, 22–24, ratory together with holotype, 1 female, 23 July 8–10, 6–8; length of legs segments: I–Leg. 1–5: 1982. 28–32, 20–26, 25–29, 30–34, 40–45; II–Leg. 1–5: Diagnosis. Larva. Dorsal shield elongate (ra- 30–32, 28–32, 24–28, 36–40, 50–52; III–Leg. 1–5: tio length/width 1.5–1.6), surrounded by narrow 32–34, 28–30, 24–28, 36–40, 50–52. subcutaneous strip, setae Fch and Vi situated on Male. Unknown. subcutaneous strip; setae C1 shorter than C4, setae Deutonymph. Unknown. C2–C4 approximately equal in length, but C4 Ecology. Sedge-sphagnum bog. thicker than C2 and C3, bases of setae C4 on coxal Geographical distribution. Asia, Russia: plate III attached closely to medial end of suture Magadan Province. line between coxal plates II and III; transverse

184 Water mites of the genus Tiphys

324 325 Figs 324–325. Tiphys chaunensis, larva: 324 — dorsal view, 325 — ventral view. Scale bar: 324–325 = 100. muscle attachment scar on coxal plate III not de- ensis sp. n. with 5 dorsal setae, genu with 2 setae, veloped; excretory pore plate wider than long, and tibia without short dorsal spine (Fig. 340). The both pairs of anal setae nearly subequal in length larva of T. chaunensis differs from the larva T. en� and situated near middle of excretory pore plate, sifer by the following characters (character states excretory pore located near posterior margin of of the female of T. ensifer are given in parenthe- excretory pore plate; solenidion on genu I slightly sis): the dorsal shield is surrounded by the narrow longer than both solenidia on tibia I, which ap- subcutaneous strip, setae Fch and Vi are situated proximately subequal in length; solenidion on on the subcutaneous strip, Fig. 324 (dorsal shield genu of leg II considerably shorter than both so- without narrow subcutaneous strip, setae Fch and lenidia on tibia II; dorsal shield and coxal plates Vi situated on the dorsal shield, Fig. 360); the dor- with reticulations. sal shield and all coxal plates are covered with re- Female. Suture line between coxal plates III ticulations (porous pattern), the transverse muscle and IV complete and slightly obliquely postero- attachment scar on coxal plate III is absent, Fig. medially directed. Medial margin of coxal plate III 325 (present, Fig. 361). slightly shorter than medial margin of coxal plate Description. Larva. Body small and oval; IV. Posteromedial and posterolateral margins of dorsal shield truncate anteriorly, rounded posteri- coxae IV forming obtuse angle with short apo- orly, with slightly convex lateral edges and weakly deme beneath this angle. Pedipalp femur and genu concave anterolateral margins (Fig. 324). Dorsal with nearly straight ventral margins; genu with 2 shield elongate (ratio length/width 1.5–1.6), sur- setae, base of external lateral seta located near rounded by narrow subcutaneous strip and cover- middle of segment; tibia slender, with straight ing almost all dorsum in unengorged larva, bear- ventral margin, both ventral setae attached near ing only 2 pairs of trichobothria (Fp, Oi), setae distal end of segment, ventral tubercles not devel- Fch and Vi situated on subcutaneous strip. Setae oped. Distolateral spine of tibia approximately 2.5 Fch and Vi slightly longer and thicker than Fp and times shorter than tarsus. Oi. Setae Oe, Hi, He, Sci, Sce and Li located on the Differential diagnosis. The coxal plates and soft wrinkled membrane. Setae Oe longer of all pedipalpal tibia in females of the new species are dorsal setae, setae Sce and Li nearly equal in similar to those in T. ensifer. The pedipalpal femur length. Lateral sides of coxae II+III coming onto in females of T. ensifer with 6 dorsal setae, genu dorsal side and almost extending to setae Oe and with 3 setae, and tibia with short dorsal spine (Fig. other hysterosomal setae located on interscutal 348); in contrast, the pedipalpal femur of T. chaun� membrane. Dorsal shield with reticulations.

185 P. V. Tuzovsky

331

330 330, 331

332

327 328 329

326–329 326 Figs 326–332. Tiphys chaunensis, larva: 326–329 — excretory pore plate, 330 — capitulum (ventral view), 331 — chelicerae, 332 — pedipalp. Scale bars: 326–329 = 20, 330–331 =50, 332 = 100.

Coxal plates I–II relatively small; anterior Excretory pore open near posterior margin of ex- coxal plates separated from coxal plates II and III cretory pore plate. which fused to each other, suture line between Capitulum (Fig. 330) with wide base, anterior them incomplete and developed only in their lat- hypostomal setae long and thin, posterior hypos- eral parts (Fig. 325). Anterior margin of coxal tomal setae short spine-like. Basal segments of plates I concave and considerably shorter than chelicerae (Fig. 331) fused at their medial parts posterior one. Coxal setae C2–C4 approximately and tapering proximally, with straight posterior subequal in length and longer than C1, bases of edges; suture line well developed along their setae C4 on coxal plate III inserted closely to be- whole length. Cheliceral stylet small, crescent. neath from medial end of suture line between cox- Pedipalp trochanter (Fig. 332) short and with- al plates II and III. Posteromedial apodemes of out setae; femur with short and fine dorsal seta coxal plates I relatively large, posteromedial apo- near middle of segment; genu with very long and demes of coxal plates II rudimentary or absent, thick proximal seta and short and fine dorsal one; and posteromedial apodemes of coxal plates III dorsal setae on femur and genu approximately slightly developed. Transverse muscle attachment equal in length; tibia with 3 unequal thin setae and scar not developed. Setae Ci, Si, Se, Le, Pi, Pe large dorsodistal claw. Tarsus with 1 solenidion, 2 situated on the soft interscutal membrane in poste- long and 5 unequal short setae. rior part of body. Setae Ci longer and thicker than All legs (Figs. 333–335) rather slender. Gen- all ventral idiosomal setae, and situated on small eral number of setae on legs I–III, except eupathid- bases; setae Li considerably longer than setae Le, ia, as follows: I–Leg. 1–5: 1, 7, 5 (s), 11(2s), 13 (s, setae Si considerably shorter than setae Se. Coxal ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 (s, ac); III– plates with reticulations. Urstigma occupying lat- Leg. 1–5: 1, 6, 5(s), 10 (s), 11(ac). Formula of eral position on the border between coxae I and II large setae on leg segments as follows: I–Leg 1–5: and provided cups. 0, 2, 1, 1, 0; II–Leg 1–5: 0, 3, 2, 3, 0; III–Leg 1–6: Excretory pore plate (Figs 326–329) wider 0, 1, 2, 3, 0. Solenidion on genu of leg I slightly than long, ratio length/width 0.6–07. Anal setae longer than both solenidia on tibia I, which ap- situated near middle of excretory pore plate, both proximately equal in length; solenidion on genu of pairs of anal setae (Ai, Ae) nearly equal in length. leg II considerably shorter than both solenidia on

186 Water mites of the genus Tiphys

333

336

334

337 335

333–335

Figs 333–337. Tiphys chaunensis, larva: 333 — leg I, 334 — leg II, 335 — leg III, 336— claws of leg I, 337 — claws of leg III. Scale bars: 333–335 = 100, 336–338 = 25. tibia II; solenidion on genu of leg III shorter than Female. (Only coxal plates, chelicera and pe- solenidion on tibia III. Central claws on tarsi of all dipalps in satisfactory condition for description). legs shorter than lateral claws (Fig. 336–337), em- Coxal plate I with 4 lateral thin setae, coxal plate podia on tarsi of legs I and II bent nearly at a right II with 3 lateral thin setae, coxal plate III with one angle and ambulacra only slightly bent, empodium medial seta and 3–4 lateral setae, coxal plate IV of legs III turned onto ventral side at obtuse an- with 2 medial setae and 4 lateral ones (Fig. 338). gle. Apodeme beneath posteromedial medial edge of Measurements, n=10. Length of dorsal shield coxal plate I moderately developed and not ex- 295–345, width 180–220; length of anal plate 23– tending beyond medial edge of coxal plate III. Su- 25, its width 32–36; length of capitulum 110–118; ture line between coxal plates III and IV complete length of basal segment of chelicera 84–100, and oblique, directed posteromedially. Medial length of chelicera stylet 12–20; length of medial margin of coxal plate III shorter than medial mar- margin of coxae I 100–110, length of medial margin of coxal plate IV. Posteromedial and postero- gin of coxae II+III 140–150; length of pedipalpal lateral margins of coxae IV forming obtuse angle segments (P-1–5): 8–12, 35–45, 24–30, 8–12, 4–6; with short apodeme beneath this angle. Sclerites length of legs segments: I–Leg. 1–5: 36–44, 32– bearing setae Hv and other idiosomal setae situat- 36, 40–45, 55–60, 68–76; II–Leg. 1–5: 40–43, ed on soft cuticle freely. 38–45, 40–46, 60–68, 84–92; III–Leg. 1–5: 40–45, Chelicera with large basal segment and rela- 40–48, 44–50, 60–70, 84–96. tively small stylet (Fig. 339). Pedipalps rather

187 P. V. Tuzovsky

338

339

340

339, 340

Figs 338–340. Tiphys chaunensis, female: 338— coxal plates, 339 — chelicera, 340 — pedipalp. Scale bars: 338 = 100, 339– 340 = 50. slender (Fig. 340). Trochanter with single short Ecology. Thermocarstic lake. dorsodistal seta, femur with 3 proximal and 2 dor- Geographical distribution. Asia, Russia: sodistal setae; ventral margin of femur straight, Magadan Province. ventral margin of genu slightly concave. Genu with 2 setae, base of external lateral seta located Tiphys (Pionides) ensifer (Müller, 1776) near middle of segment. Tibia rather slender, Figs 341–372 slightly tapering distally with straight ventral mar- Material. 3 females Russia, Samara Prov- gin, with relatively short distolateral spine (2.5 ince, National natural Park “ Samara Luca”, vil- times shorter than tarsus), 2 ventral setae, and sev- lage Koltsovo, sedge bog, May 1998; 10 females, eral short and fine dorsal ones; both ventral setae 5 males, 1 deutonymph and 25 larvae reared in inserted distally, ventral tubercles lacking. Pedi- laboratory (IBIW): Russia, Yaroslavl Province, palpal tarsus tapering distally, with single long Nekouz Distr., Borok, temporary water bodies, proximal solenidion, 5 thin setae and 4 thick distal sedge bogs, May–July, 2001–2002, 2004, P.V. Tu- unequal spines. zovsky. Measurements, n=1. Length of medial margin Diagnosis. Female. Medial margin of coxal of coxa III 54, length of medial margin of coxa IV plate III slightly shorter than medial margin of 72; length of basal segment of chelicera 205, coxal plate IV. Sclerites bearing setae Hv and other length of cheliceral stylet 60; length of pedipalpal ventral idiosomal setae situated on soft cuticle segments (P-1–5): 40, 95, 55, 108, 54; length of freely. Genital plates roughly triangular with 3 ac- distolateral spine on tibia 21. etabula and 13–17 short and fine setae; all aceta-

188 Water mites of the genus Tiphys

341 342

343

345 346

344

342–346

Figs 341–346. Tiphys ensifer, female: 341 — seta Fch, 342 — anteromedial and anterolateral dorsal plates, 343 — posteromedial plates, 344 — ventral view, 345–346 — excretory pore. Scale bars: 341 = 50, 342–346 = 100. bula approximately subequal in sizes, and arranged denticle with short and thin dorsal spur; small claw in triangular. Genital opening and genital plates with relatively long external denticles and short nearly subequal in length; anterior genital sclerite internal one. Claws of legs IV lesser than claws of wider than posterior one. Pedipalpal genu usually legs I and II. with 3 setae, external lateral seta 2–3 times longer Deutonymph. Dorsum without true plates. than other dorsal setae, base of external lateral seta Medial margin of coxae III and medial margin of located near middle of segment; tibia slender with coxae III subequal in length. Posteromedial angle short distolateral spine (2.5–3.0 times shorter than of coxae IV obtuse with very short apodeme be- tarsus), both ventral setae attached near distal end neath this angle. Sclerites bearing setae Hv fused of segment, ventral tubercles not developed. Claws with posterior margin of coxae II; sclerites bearing of legs I–IV subequal in size; both denticles point- other ventral idiosomal setae situated on soft cuti- ed and approximately subequal in length. cle freely. Both acetabula subequal in sizes, dis- Male. Coxae plates IV greatly expanded and tance between them usually equal or lesser than fused with one another posteromedially, posterior diameter of acetabulum on each plate; genital sc- edge of coxal plate IV transverse and sinuous, lerite considerably larger than pregenital one. Ex- forming convexity with lateral edge; medial mar- cretory pore with relatively large anterior sclerite gin of coxal plate IV 2.0–2.5 times longer than and small posterior one. Pedipalpal genu with 2 medial margin of coxal plate III, suture line be- setae, external lateral seta considerably longer tween coxal plates III and IV nearly transverse; than dorsodistal seta; base of external lateral seta genital plates triangular and confined to gonopore attached distally to middle of segment; tibia with region. Sclerites bearing setae Hv fused with pos- large distolateral spine (0.75–0.80 times as long as terior margin of coxae II; sclerites bearing other tarsus), ventral setae situated near distal end of ventral idiosomal setae situated on soft cuticle segment, ventral tubercles lacking. Claws of legs freely. Telofemur IV with 2 long swimming setae, I–IV subequal in size. genu IV slightly expanded and bearing long, Larva. Dorsal shield elongate (ratio length/ curved, spine-like seta dorsodistally, which shorter width 1.6–1.7), bearing 4 pairs of setae, setae Fch than of segment. Claw of legs I and II similar to and Vi slightly longer and thicker than Fp and Oi. female. Claws of legs III asymmetric; large claw All coxal setae (C1–C4) subequal in length but C4 with two more or less subequal denticles, external thicker than C1–C3, base of setae C4 inserted

189 P. V. Tuzovsky

350 347

349

348

Figs 347–350. Tiphys ensifer, female: 347 — chelicera, 348 — pedipalp, 349 — telofemur, genu, tibia and tarsus of leg IV, 350 — claw. Scale bars: 347–350 = 100. closely to beneath from medial end of suture line medial seta and 2–3 lateral setae, coxal plate IV between coxal plates II and III. Transverse muscle with 1 medial seta and 4–5 lateral setae (Fig. 344). attachment scar present. Dorsal shield and coxal Apodeme beneath posteromedial medial edge of plates porous. Excretory pore plate wider than coxal plate I moderately developed and usually in long, setae Ai and Ae located near anterior margin immature specimens extending beyond medial of excretory pore plate, setae Ae slightly longer edge of coxal plate III. Suture line between coxal than Ai. Excretory pore open in posterior half of plates III and IV complete and oblique, directed excretory pore plate. Solenidion on genu of leg I posteromedially. Medial margin of coxal plate III slightly longer than both solenidia on tibia I; so- slightly shorter than medial margin of coxal plate lenidion on genu of leg II and both solenidia on IV. Posteromedial and posterolateral margins of tibia II approximately subequal in length. coxae IV forming obtuse angle with short apo- Description. Female. Body oval; setae Fch deme beneath this angle. Sclerites bearing setae long, whip-like (Fig. 341), other idiosomal setae Hv close or fused to posterior margin of coxal filiform. Dorsum with relatively large anterome- plate II, other idiosomal setae situated on soft cu- dial plates, small anterolateral and posteromedial ticle freely. Genital plates roughly triangular with platelets (Figs 342–343); anteromedial and pos- 3 acetabula and 13–17 short and fine setae; aceta- teromedial plates elongate, anterolateral plates bula in triangular approximately equal in size; ac- transverse. Coxal plate I usually with 4(3) lateral etabula occupy about half-surface of genital plate. thin setae and 1 medial thin seta, coxal plate II Genital opening and genital plates nearly equal in with 2 lateral thin setae, coxal plate III with one length; anterior genital sclerite wider than poste-

190 Water mites of the genus Tiphys

351

354 352 353 Figs 351–354. Tiphys ensifer, male: 351— ventral view, 352 — telofemur, genu, tibia and tarsus of leg III, 353 — telofemur, genu, tibia and tarsus of leg IV, 354 — claws of leg III. Scale bars: 351–353 = 100, 354 =50. rior one. Excretory pore usually with small ante- 5–6 on tibia IV. Claws of legs I–IV equal in size; rior sclerite (Fig. 345–346). both denticles pointed and approximately sub- Capitulum with short anchoral projection, equal (Fig. 350). Claw lamella moderately devel- both pairs of setae equal in length. Chelicera (Fig. oped, with slightly convex ventral margin. 347) with large basal segment and relatively small Measurements (n=10). Body length 600–855, pointed stylet. length of medial margin of coxa III 54–60, length Pedipalps slender (Fig. 348). Trochanter with of medial margin of coxa IV 65–79; length of an- 1 short dorsodistal seta, femur with 3–5 proximal teromedial dorsal plates 60–72, width 15–25, and 2 dorsodistal setae; ventral margin of femur length of genital plates 120–140, width 65–75; straight or slightly convex, ventral margin of genu length/width of genital acetabula (ac. 1–3): 42– straight or slightly concave. Genu with 3 setae, ex- 48/24–30, 36–45/28–30, 42–50/35–42; width of ternal lateral seta 2–3 times longer than other dor- anterior genital sclerite 60–95; length of basal seg- sal setae; base of external lateral seta located near ment of chelicera 165–170, length of cheliceral middle of segment. Pedipalpal tibia slender, slight- stylet 55–60; length of pedipalpal segment (P-1–5): ly tapering distally, with short dorsodistal spine 40–50, 120–140, 70–85, 130–165, 55–60; length (2.5–3.0 times shorter than tarsus), 2 ventral setae, of distolateral spine on tibia 20–26; length of leg several short and fine dorsal setae and 1 thick spine segments: I–Leg. 1–6: 65–85, 120–130, 120–130, proximally to middle of segment; both ventral se- 135–155, 160–180, 225–270; II–Leg. 1–6: 72–85, tae attached near distal end of tibia, ventral tuber- 120–135, 115–140, 135–165, 170–195, 225–270; cles not developed. Pedipalpal tarsus tapering dis- III–Leg. 1–6: 70–90, 115–140, 105–130, 135–165, tally, with 1 long proximal solenidion, 5 thin setae 175–205, 220–260; IV–Leg. 1–6: 120–150, 140– and 4 thick distal unequal spines. 155, 145–175, 195–235, 185–200, 240–295. All legs with swimming setae; legs I and II Male. Dorsal surface, capitulum, chelicera with short swimming setae, two posterior pairs of and pedipalps similar to those in female. Anterior legs with long swimming ones (Fig. 349). One apodemes rather large, extending beyond medial swimming seta present on genu I, 1–2 on telofem- edge of coxal plate III; posterior apodemes small ora III–IV and genu III, 2 on tibia I and genu II, (Fig. 351). Coxae plates IV greatly expanded, 3–4 on genu IV and tibia II, 4–7 on tibia III, and fused with one another posteromedially, with 2

191 P. V. Tuzovsky

355 358

355, 357, 359

356 359 357

Figs 355–359. Tiphys ensifer, deutonymph: 355 — seta Fch, 356 — ventral view, 357— pedipalp, 358 — genu, tibia and tarsus of leg IV, 359 — claw. Scale bars: 355, 357, 359 = 50; 356, 358 = 100. short and fine setae close together anterior to de- with numerous short and thin setae and 5–8 thick pressed region on each plate. Posteromedial edge unequal ones. of coxal plate IV transverse and sinuate, forming Claw of legs I and II similar to those in female. convexity with lateral edge. Medial margin of Claws of legs III (Fig. 354) asymmetric; large claw coxal plate IV approximately twice as long as me- with subequal denticles, external denticle with short dial margin of coxal plate III. Suture line between and thin dorsal spur; small claw with long external coxal plates III and IV nearly transverse. Genital denticles and relatively short internal one. plates triangular and confined to gonopore region; Measurements, n=5. Body length 570–700, all acetabula relatively large, subequal and occupy length of medial margin of coxa III 70–90, length more than half of plate surface. Anterior genital of medial margin of coxa IV 150–180; length of sclerite small with two pairs of short and fine set- anteromedial dorsal plates 36–48, width 9–12; ae. Genital plates with 2–3 short and fine setae length of genital plates 80–105, theirs general each. Sclerites bearing setae Hv fused with poste- width 175–215; length/width of genital acetabula rior margin of coxae II; sclerites bearing other (ac. 1–3): 24–36/24–30, 30–36/24–30, 30–38/24– ventral idiosomal setae situated on soft cuticle 32; length of basal segment of the chelicera 105– freely. Excretory pore with small anterior sclerite. 130; length of cheliceral stylet 40–50; length of Genu of leg III (Fig. 352) shorter than tibia pedipalpal segment (P-1–5): 32–40, 95–115, 60– III, tarsus III considerably longer than tibia III; te- 65, 120–140, 40–50; length of distolateral spine lofemur III with 2, genu III with 2–3, and tibia III on tibia 15–21; length of leg segments: I–Leg. with 5–6 long swimming setae. Telofemur IV (Fig. 1–6: 60–75, 105–125, 115–130, 130–155, 155– 353) with 2 long ventrodistal swimming seta; genu 175, 250–270; II–Leg. 1–6: 70–85, 115–130, 115– IV slightly expanded dorsoventrally relatively te- 140, 145–165, 170–195, 250–270; III–Leg. 1–6: lofemur and bearing large curved seta dorsodis- 70–85, 80–90, 55–75, 80–90, 130–145, 195–205; tally, it shorter than of segment. Tibia IV with IV–Leg. 1–6: 105–130, 105–125, 95–100, 120– slightly concave dorsal margin, with a few short 130, 175–220, 270–285. and thick setae, and 6–8 ventrodistal long swim- Deutonymph. Dorsum without true plates. ming setae. Tarsus IV slightly tapering distally, Setae Fch long, thin (Fig. 355). Coxal plate I with

192 Water mites of the genus Tiphys

360 361 Figs 360–361. Tiphys ensifer, larva: 360 — dorsal view, 361 — ventral view. Scale bar: 360–361=100.

2 lateral thin setae and 1 posteromedial thin seta, 0.80 times as long as tarsus. Pedipalpal tarsus with coxal plate II with 1 lateral thin setae, coxal plate single long proximal solenidion, 3 thin setae and 4 III with 1 medial seta and 1 lateral thin setae, cox- short, thick distal spines. al plate IV with 1 medial seta and 1 lateral one Legs II–IV with swimming setae; second pair (Fig. 356). Apodeme beneath posteromedial me- of legs with short swimming setae, legs III–IV dial edge of coxal plate I moderately developed with long swimming ones (Fig. 358). Genu II and not extending posteriorly to beneath medial edge III with 1 swimming seta, tibia II–III and genu IV of coxal plate III. Suture line between coxal plates with 2, and tibia IV with 3 swimming setae. Claws III and IV oblique, directed posteromedially. Me- of legs I–IV subequal in sizes; both denticles ap- dial margin of coxae III and medial margin of proximately subequal in length, external denticle coxae III subequal in length. Posteromedial angle thin-pointed, and internal denticle relatively wide of coxae IV obtuse, with very short apodeme be- (Fig. 359). Claw lamella slightly developed, with neath this angle. Sclerites bearing setae Hv fused nearly straight ventral margin. with posterior margin of coxae II; scleritesbearing Measurements, n=1. Body length 510; length other ventral idiosomal setae situated on soft cuti- of medial margin of coxa III 35, length of medial cle freely. Genital plates separated, oblique, di- margin of coxa IV 35–37; length of genital plates rected anteromedially, bearing 2 acetabula and 3 60, width 30; diameter of genital acetabula (ac. short and fine setae each. Both acetabula subequal 1–2): 18–21, 18–22; length of basal segment of in size, distance between them usually equal to or chelicera 90; length of cheliceral stylet 40; length lesser than one diameter of acetabulum on each of pedipalpal segment (P-1–5): 24, 66, 42, 72, 36; plate. Genital sclerite larger than pregenital scler- length of distolateral spine on tibia 27; length of ite. Excretory pore with small anterior sclerite. leg segments: I–Leg. 1–6: 42, 55, 54, 72, 88, 120, Pedipalpal trochanter without seta, femur II–Leg. 1–6: 48, 51, 54, 78, 95, 125; III–Leg. 1–6: with 3 dorsodistal setae (Fig. 357). Ventral margin 48, 48, 54, 72, 102, 125; IV–Leg. 1–6: 72, 60, 75, of femur straight, ventral margin genu slightly 102, 132, 150. concave. Genu with 2 setae, external lateral seta Larva. Body small and oval; dorsal shield considerably longer than dorsodistal one; base of truncate anteriorly, rounded posteriorly, with con- external lateral seta attached distally to middle of vex wavy lateral edges, it anterolateral margins segment. Tibia with very long dorsodistal spine, 2 are weakly concave (Fig. 360). Dorsal shield elon- ventral and 1–2 dorsodistal setae; ventral setae gate (ratio length/width 1.6–1.7), covering almost and situated near distal end of segment, ventral tu- all dorsum in unengorged larva and bearing 4 pairs bercles lacking. Dorsodistal spine of tibia 0.75– of setae (Fp, Oi, Fch, Vi), all setae approximately

193 P. V. Tuzovsky

366

365

365–367

362 367 364 362–364 363 Figs 362–367. Tiphys ensifer, larva: 362–364 — excretory pore plate, 365 — capitulum (ventral view), 366 — chelicerae, 367 — pedipalp. Scale bars: 362–364 = 25, 365–367 = 20. subequal in length, but setae Fch and Vi thicker considerably shortest than setae Se. Coxal plates than Fp and Oi. Setae Oe, Hi, He, Sci, Sce and Li I–III porous. Urstigma occupying lateral position located on the soft wrinkled membrane. Setae Oe on body border between coxae I and II and pro- longest of all dorsal setae, setae Sce longer than vided with cups. setae Li. Lateral sides of coxae II+III coming onto Excretory pore wider than long (Figs 362– dorsal side and almost extending to setae Oi and 364). Both pairs of anal setae situated near anterior other hysterosomal setae located on interscutal margin of excretory pore plate, setae Ae longer membrane. Dorsal shield porous. than setae Ai. Excretory pore placed in posterior Coxal plates I–II relatively small; anterior half of excretory pore plate. coxal plates are separated from coxal plates II Capitulum (Fig. 365) elongate, with rather and III, which are fused, suture line between wide base, anterior hypostomal setae long, thin, them incomplete and developed only in their lat- posterior hypostomal setae short and thick. Basal eral parts (Fig. 361). Anterior margin of coxal segments of chelicerae (Fig. 366) fused at their plates I concave and considerably shorter than medial parts and tapering proximally, suture line posterior one. All coxal setae (C1–C4) approxi- well developed along their whole length. Chelic- mately equal in length, base of setae C4 on coxal eral stylet small, crescent. plate III inserted closely to medial end of suture Pedipalp trochanter (Fig. 367) short and with- line between coxal plates II and III. Posterome- out setae; lateral seta of femur and genu thin, short dial apodemes of coxal plates I well developed, and approximately of similar length, proximal lat- posteromedial apodemes of coxal plates II and eral genual seta long and very thick; tibia with 3 III small. Transverse muscle attachment scar sit- approximately subequal setae and large dorsodis- uated near posterior margin of coxal plate III. Se- tal claw; tarsus with one rather long solenidion, 2 tae Ci, Si, Se, Le, Pi, Pe situated on soft inters- long and 5 unequal simple setae. cutal membrane in posterior part of body. Setae All legs short, stout. The general number of Ci very long and heaviest than other ventral idio- setae on legs I–III (Figs 368–370), except eu- somal setae, and situated on small bases; setae Si pathidia, as follows: I–Leg. 1–5: 1, 7, 5 (s), 11

194 Water mites of the genus Tiphys

371

368

372 368–370

369

370

Figs 368–372. Tiphys ensifer, larva: 368 — leg II, 369 — leg III, 370— claws of leg I, 371 — claws of leg I, 372 — claws of leg III. Scale bars: 368–370 = 50, 371–372 = 25.

(2s), 13 (s, ac); II–Leg. 1–5: 1, 7, 5 (s), 11 (2s), 13 plate 16–18, width 20–25; length of capitulum (s, ac); III–Leg. 1–5: 1, 6, 5(s), 10 (s), 11 (ac). 70–75; length of basal segment of chelicera 64– Formula of large setae on leg segments as fol- 68, length of chelicera stylet 12–14; length of me- lows: I–Leg. 1–5: 0, 3, 1, 3, 0; II–Leg. 1–5: 0, 3, dial margin of coxae I 64–68, length of medial 2, 4, 0; III–Leg. 1–6: 0, 2, 2, 4, 0. Solenidion on margin of coxae II+III 83–93; length of pedipalpal genu of leg I slightly longer than both solenidia segments (P-1–5): 6–8, 24–26, 20–22, 7–8, 4–6; on tibia I; solenidion on genu II and both solenid- length of legs segments: I–Leg. 1–5: 23–25, 19– ia on tibia II approximately subequal in length; 21, 22–24, 28–30, 41–44; II–Leg. 1–5: 30–33, solenidion on genu of leg III shorter than solen- 25–28, 24–27, 33–36, 50–52; III–Leg. 1–5: 30–33, idion on tibia III. Central claw and lateral claws 30–33, 25–28, 40–43, 51–54. on tarsi of all legs nearly subequal in length (Fig. Ecology. Temporary ponds, sedge-sphagnum 371–372); tarsi of all legs with hook-like empo- bogs, lakes. Also in permanent waters. dium, and lateral claws slightly bent. Geographical distribution. Europe, Asia Measurements, n=10. Length of dorsal shield (Viets 1936, 1956; Sokolow 1940; Lundblad, 200–210, width 120–130; length of excretory pore 1968; K.O. Viets 1978).

195 P. V. Tuzovsky

Fig. 373. Tiphys uchidai, male: 373 — telofemur, genu, tibia and tarsus of leg IV, from Uchida (1936). Distal short setae in genu IV are not show.

Tiphys (Pionides) uchidai Viets, 1956 as genital plates. Pedipalpal genu with 3 setae, external lateral seta longer than two other setae; base Fig. 373 of external lateral seta situated near to middle of Uchida (1936) gave brief characteristics and segment; tibia rather slender with short distolateral illustrated the telofemur, genu, tibia and tarsus IV spine (2.5 times shorter than tarsus), internal ven- of T. ensifer (Fig. 373) from the island of Sakhalin, tral seta situated on relatively large conic tubercle, which he has assigned to T. ensifer var. Karl Viets external ventral seta is on very small tubercle. In- (1956) elevated this variety to the rank of subspe- ternal denticle of claw of legs I–III wide with cies (T. ensifer uchudai), while K.O. Viets (1987) roundish tip, external denticle thin and pointed. considered it a full species, T. uchidai Viets, Claws of legs IV with short internal and long ex- 1956. ternal denticles, latter supplied with short spur. Remarks. The legs IV of T. ensifer differ Male. Coxae IV greatly expanded, their length from the legs IV of T. uchidai by the following exceeding total length of coxae I–III. Medial mar- features (characters states of T. uchidai are indi- gin of coxae III in 2.0–2.7 times shorter than length cated in parentheses based on Uchida 1936): te- of medial margin of coxae IV. Genital plates fused lofemur with 2 long swimming setae, Fig. 353 by anterior and posterior ends, acetabula arranged (without swimming setae, Fig. 373), genu with in an arc; genital setae clustered in four groups, relatively short curved seta dorsodistally, it length two anterior groups have a few setae (6–11), two shorter than of segment (with long curved seta posterior groups with more numerous setae (11– dorsodistally, it length longer than of segment). 19). Sclerites, bearing setae Pe, fused with pos- Tiphys (Acercopsis) pistillifer (Koenike, 1908) teromedial margin of coxae IV; sclerites, bearing setae Hv, fused with posterior margin of coxae II. Figs 374–410 Tibia of legs II with single spatulate seta. Genu of Material. 12 females, 5 males and 17 larvae leg IV expanded proximally, tibia IV curved reared in laboratory (IBIW): Russia, Yaroslavl slightly thickened near middle, with distolateral Province, Nekouz Distr., Borok, forest sedge bog leaf-shaped seta. Claws of legs III asymmetric near the settlement, April–August 2000–2003, with 3 unequal denticle each, large claw with rath- P.V. Tuzovsky. er long internal and median denticles and short Diagnosis. Female. Dorsum with 3 pairs of external one; small claw with long median denti- plates, venter with two pairs of rather large elon- cle and short internal and external ones, all denti- gate platelets and two pairs of small sclerites is cles pointed. present along a lateral line of a body. Medial mar- Larva. Dorsal shield elongate (ratio length/ gin of coxae III and IV approximately equal in width 1.4–1.6), covering almost all dorsum in un- length. Sclerites bearing setae Hv fused with pos- engorged larvae and bearing two pairs of trichobo- terior margin of coxae II. Genital plates triangular, thria (Fp, Oi). Setae Fch situated in anterolateral genital opening approximately 1.5 times as long corners of dorsal shield, and setae Vi occupying

196 Water mites of the genus Tiphys

374 375 Figs 374–375. Tiphys pistillifer, female: 374 — dorsal view, 375 — ventral view. Scale bar: 374–375 = 100. lateral position; both these pairs of setae in imma- all coxae with pore-like structure on it. Sclerites ture specimens separated from dorsal shield. Setae bearing setae Hv fused with posterior margin of C1 shorter than setae C2–C4 which approximately coxae II; sclerites bearing other ventral idiosomal equal in length. Base of setae C4 on coxal plate III setae situated freely on soft cuticle. Two pairs of located near medial end of suture line between rather large elongate platelets and two pairs of coxal plates II and III. Transverse muscle attach- small sclerites present along lateral idiosoma mar- ment scar present. Dorsal shield and coxal plates gins. I–III with pore like structure and slightly devel- Genital plates triangular (Figs 376–377), with oped reticulations on it. Excretory pore plate wider 3 acetabula and 6–9 thin short setae each. Lateral than long, ratio length/width = 0.6–0.7; setae Ai margins of genital plates distinctly concave. All and Ae approximately equal in size. Excretory acetabula approximately equal in size; distance pore located in posterior half of excretory pore between first and second acetabula, and distance plate. Solenidion on genu of leg I longer than so- between first and third acetabula more than diam- lenidia on tibia I; solenidia on genu and tibia of eter one acetabulum; distance between second and legs II equal in size. third acetabula usually lesser than diameter one Description. Female. Body oval; setae Fch acetabulum. Anterior genital sclerite wider than long, whip-like, other idiosomal setae filiform posterior one. Genital opening approximately 1.5 (Fig. 374). Integument soft, with very fine strips. times longer than genital plates. Dorsum with rather large anteromedial, relatively Capitulum with rather large anchoral projec- small anterolateral and posteromedial plates. Me- tion, both pairs of setae equal in length (Fig. 378). dial and posterior plates elongate, lateral plates The basal segment of chelicera (Fig. 379) convex, transverse. The first four pairs of lyriform organs cheliceral stylet relatively large, pointed. (i1–i4) located on dorsum, i5 in the posterior part Pedipalps (Fig. 380) relatively short, stout. of the dorsum or the venter. Trochanter with single dorsodistal seta. Ventral Anterior groups of coxae with rather large margin of femur and genu of pedipalp straight. Fe- apodemes (Fig. 375). Medial margin of coxae III mur with 3–4 proximal and 2 dorsodistal short se- and IV approximately equal in length. Suture line tae. Pedipalpal genu with 3 setae, external lateral between coxal plates III and IV obliquely postero- seta longer than two other setae; base of external medially-directed. Posteromedial margin of coxal lateral seta situated near middle of segment. Pedi- plate IV concave, and forming roughly right pos- palpal tibia with rather large distolateral spine (2.5 teromedial angle with lateral margin. Surface of times shorter than tarsus), 2 ventral and 2–4 dorsal

197 P. V. Tuzovsky

380

381 378–380 381

376

378

377 379

Figs 376–381. Tiphys pistillifer, female: 376–377 — genital plate, 378 — capitulum, ventral view, 379 — chelicera, 380 — pedipalp, 381 —distal part of pedipalpal tibia and tarsus. Scale bars: 376–380 = 50, 381 = 20. setae. Internal ventral seta on tibia situated on rel- internal and long external denticles, latter supplied atively large conic tubercle, base of external ven- with short spur (Fig. 385). tral seta on very small tubercle. Pedipalpal tarsus Measurements, n=10. Body length 685–800; (Fig. 381) with 1 long proximal solenidion, 5 thin length of medial margin of coxa III 45–55, length setae and 4 thick unequal dorsodistal spines. of medial margin of coxa IV 40–50; length of gen- Two anterior pairs of legs without true swim- ital plates 85–105, width 65–80; length of genital ming setae (Fig. 382). Three relatively short swim- acetabula (ac. 1–3) — 25–30, 25–30, 25–30; ming setae present on genu III, 3–4 on genu IV, length of basal segment of chelicera 145–155; 4(3) long swimming setae on tibia III and 5–7 on length of cheliceral stylet 54–60; length of pedi- tibia IV (Fig. 383). Claws of legs I–III larger than palpal segment (P-1–5) — 25–30, 90–97, 54–60, claws of legs IV. Internal denticle of claw of legs 90–105, 42–48; length of leg segments: I–Leg. I–III wide with roundish tip, external denticle thin 1–6 — 50–60, 60–80, 75–90, 100–120, 115–140, and pointed (Fig. 384). Claws of legs IV with short 135–165; II–Leg. 1–6 — 60–65, 60–75, 85–90,

198 Water mites of the genus Tiphys

382

382, 383

383

384

385 384, 385 Figs 382–385. Tiphys pistillifer, female: 382 — genu, tibia and tarsus of leg I, 383 — genu, tibia and tarsus of leg IV, 384 — claw of leg II, 385 — claw of leg IV. Scale bars: 382–383 = 70, 384–385 =20.

105–120, 120–150, 135–170; III–Leg. 1–6 — 65– rior groups with more numerous setae (11–19). 72, 60–80, 70–85, 100–120, 135–165, 135–165; Sclerites bearing setae Pe fused with posterome- IV–Leg. 1–6 — 100–125, 75–95, 95–120, 155– dial margin of coxae IV; sclerites bearing setae Hv 180, 185–210, 180–210. fused with posterior margin of coxae II; other sc- Male. Dorsal surface, capitulum, chelicerae lerites bearing ventral idiosomal setae situated on and coxae of legs I–III similar to female. Coxal soft cuticle. Excretory pore with small anterior sc- plates in four groups and occupying most of ven- lerite. tral surface (Fig. 386). Anterior apodemes rather Pedipalps similar to female, but tibia with large, extending beyond medial edge of coxal plate more numerous dorsal setae. III; posterior apodemes not developed. Coxae IV Basifemur II (Fig. 388) with relatively large greatly expanded, their length exceeding total ventrodistal hump, tibia II with single spatulate length of coxae I–III. Medial margin of coxae III seta. Tibia III (Fig. 389) with 6–7 long swimming 2.0–2.7 times shorter than medial margin of coxae setae. Genu IV (Fig. 390) expanded proximally, IV. Posteromedial margin of coxae IV oblique and with maximal height in the middle of segment, forming bay where external genital organ situated; bearing 10–12 short, bent dorsal setae with round lateral edge of these coxae convex. Genital plates tips; ventral surface of genu straight, with long fused by anterior and posterior ends, acetabula ar- proximal swimming setae and numerous short ranged in an arc. Anterior and posterior acetabula pointed setae; in addition, swimming setae present oval, median acetabula usually rounded. Lateral in distolateral part of genu. Tibia IV (Fig. 391) margins of genital plate weakly concave; posterior curved, slightly thickened near middle, with con- margins converge at blunt angle. Genital setae cave dorsal and convex ventral sides; with 9–12 clustered in four groups (Fig. 387); two anterior dorsal and 6–7 short ventrolateral setae, with 6–7 groups have few setae (6–11) and located in two- long swimming setae, and distolateral leaf-shaped three rows in the anterior part of plate, two poste- seta (Figs 392–393). Tibia and tarsus IV approxi-

199 P. V. Tuzovsky

388

386 387 Figs. 386–388. Tiphys pistillifer, male: 386 — ventral view, 387 — genital plate, 388 — leg II. Scale bars: 386 = 100, 387–388 =50. mately some length, latter with numerous thin and teromedial margins are weakly concave (Fig. 396). thick setae. Claw of legs I and II approximately Dorsal shield elongate (ratio length/width 1.4–1.6), equal in sizes, with wide internal and narrow ex- covering almost all dorsum in unengorged larvae ternal denticles. Claws of legs III (Fig. 394) asym- and bearing two pairs of trichobothria (Fp, Oi). Se- metric, with 3 unequal denticle each, large claw tae Fch situated in anterolateral corners of dorsal with rather long internal and median denticles and shield, and setae Vi occupying lateral position; both short external one; small claw with long median these pairs of setae in immature specimens sepa- denticle and short internal and external denticles; rated from dorsal shield. Setae Oe, Hi, He, Sci, Sce, all denticles pointed. Claws of legs IV with short Li and the first 3 pairs of lyriform organs i1–i3( ) are internal and long external pointed denticles, latter located on the soft wrinkled membrane. Lateral supplied with short spur (Fig. 395). sides of coxae II+III coming onto dorsal side and Measurements, n=5. Body length 560–590; almost extending to setae Oi and other hysterosom- length of genital opening 64–70; length of genital al setae located on interscutal membrane. Dorsal plate 120–135, width 150–170; length (or diame- shield with a pore like structure and slightly devel- ter) of genital acetabula (ac. 1–3): 30–35, 25–30, oped reticulations on it. 30–35; length of basal segment of chelicera 120– Coxal plates I–II small; anterior coxal plates 126; length of cheliceral stylet 54–60; length of separated from coxal plates II and III which fused, pedipalpal segment (P-1–5) — 24–30, 84–90, 48– suture line between them is developed only in their 55, 90–96, 35–42; length of leg segments: I–Leg. lateral parts (Fig. 397). Anterior margin of coxal 1–6 — 55–65, 60–66, 75–95, 120–130, 125–135, plates I is straight or weakly concave and almost 160–175; II–Leg. 1–6 — 65–75, 65–73, 90–95, twice as short as posterior margin. Setae C1 short- 120–130, 120–135, 170–185; III–Leg. 1–6 — 65– er than setae C2–C4 which approximately equal in 80, 60–66, 54–60, 70–80, 115–120, 130–140; IV– length. Base of setae C4 on coxal plate III located Leg. 1–6 — 125–140, 70–85, 54–75, 125–135, near medial end of suture line between coxal plates 195–205, 185–205; length of medial margin of II and III. Posteromedial apodemes of coxal plates coxae III 48–60, length of medial margin of coxae I developed distinctly, apodemes of coxal plates II IV 110–125, length of leaf-shaped seta on tibia of and III slightly developed or reduced. Transverse legs II 50–55, its width 18–23. muscle attachment scar located in the posterome- Larva. Body small and oval in shape; dorsal dial angles of coxal plates III. Setae Ci, Si, Se, Le, shield truncate anteriorly, rounded posteriorly, it an- Pi, Pe and two pairs of lyriform organs (i4–i5)

200 Water mites of the genus Tiphys

394 389

389–391

394, 395

390 395

391

392 393

Figs 389–395. Tiphys pistillifer, male: 389 — genu, tibia and tarsus of leg III, 390 — basifemur, telofemur and genu of leg IV, 391— tibia and tarsus of leg IV, 392–393 — leaf-shaped seta, 394 — claws of tarsus III, 395 —claw of tarsus IV. Scale bars: 389–393 = 50, 394–395 = 20. situated on the soft interscutal membrane in the slightly anteriot to setae Ae; excretory pore located posterior part of the body. Setae Ci longest and in posterior half excretory pore plate. thickest of all ventral idiosomal setae, and situated Capitulum with wide base, anterior hypos- on small bases; setae Se shorter than Si. All lyri- tomal setae long, thin and posterior hypostomal form organs are ring-shaped. Coxal plates I–III setae short, spine-like (Fig. 402). Basal segments with pore-like structure and slightly developed re- of chelicerae (Fig. 403) fused by their medial ticulations on it. Urstigma occupying lateral posi- parts; suture line well developed along their whole tion on border between coxae I and II and provided length. Cheliceral stylet rather small and crescent. with cups. Pedipalp trochanter (Fig. 404) short and without Excretory pore plate wider than long (Figs setae; dorsal setae of femur and genu of pedipalps 398–401), ratio length/width = 0.6–0.7. Setae Ai approximately equal in size, proximal lateral gen- and Ae approximately equal in sizes. Anal setae ual seta very long and very heavy; tibia with one forming regular transverse row, or setae Ai located long seta, two short setae, and large dorsodistal

201 P. V. Tuzovsky

396 397 Figs 396–397. Tiphys pistillifer, larva: 396 — dorsal view, 397 — ventral view. Scale bar: 396–397= 50.

404 403 402

400 401 398 399

Figs 398–404. Tiphys pistillifer, larva: 398–401 — excretory pore plate, 402 — capitulum (ventral view), 403 — chelicerae, 404 — pedipalp. Scale bar: 398–404 = 20. claw. Pedipalpal tarsus with 1 short solenidion, 2 11 (ac). Formula of large setae on leg segments as long and 5 short thin setae. follows: I–Leg 1–5 — 0, 1, 1, 2, 0; II–Leg 1–5 — All legs rather slender (Figs 405–407). The 0, 1, 2, 4, 0; III–Leg 1–6 — 0, 2, 2, 4, 0. Solenidi- general number of setae on legs I–III , except eu- on on genu of leg I longer than solenidia on tibia I; pathidia, is as follows: I–Leg. 1–5 — 1, 7, 5 (s), solenidia on genu and tibia of legs II equal in size; 11(2s), 13 (s, ac); II–Leg. 1–5 — 1, 7, 5 (s), 11 solenidion on genu of leg III shorter than solenid- (2s), 13 (s, ac); III–Leg. 1–5 — 1, 6, 5(s), 10 (s), ion on tibia III. On tarsi of legs I and II, empodium

202 Water mites of the genus Tiphys

405 408

405–407

408–410 409 406

410 407

Figs 405–410. Tiphys pistillifer, larva: 405 — leg I, 406 — leg II, 407— leg III, 408 — claws of leg I, 409 — claws of leg II, 410 — claws of leg III. Scale bars: 405–407 = 40, 408–410 = 20. bent at a right angle, and ambulacra only slightly II+III 105–115; length of pedipalpal segments bent (Figs 408–409). Distal part of empodium of (P-1–5): 5–10, 24–32, 18–22, 7–11, 5–7; length of leg III turned onto ventral side at obtuse angle legs segments: I–Leg. 1–5: 32–42, 28–32, 28–35, (Fig. 410). 40–48, 55–65; II–Leg. 1–5: 35–39, 32–36, 32–37, Measurements, n=10. Length of dorsal shield 44–55, 65–77; III–Leg. 1–5: 35–42, 35–42, 32–36, 225–275, its width 155–175; diameter of urstigma 50–58, 70–77. 10–12; length of the anal plate 17–22, its width Ecology. Temporary ponds, sedge bogs, lakes, 32–35; length of capitulum 64–70; length of basal brooks. segment of chelicera 65–70, length of the chelic- Geographical distribution. Europe (Viets era stylet 13–19; length of medial margin of the 1956; Lundblad 1968; K.O. Viets 1978), Russia: coxae I 70–80, length of medial margin of coxae Yaroslavl Province (Tuzovsky 1996, 2008).

203 P. V. Tuzovsky

411 412 Figs 411–412. Tiphys gladiator, female: 411— dorsal view, 412 — ventral view. Scale bar: 411–412 = 100.

Tiphys (Acercopsis) gladiator Tuzovskij, 2005 oventrally with maximal height in middle of seg- Figs 411–430 ment, bearing 10 short and bent setae with round Material. Holotype male (IBIW 9102): Rus- tips; ventral surface of genu nearly straight, with sia, Yaroslavl Province, Nekouz Distr., Borok; long proximal swimming setae and numerous sedgesphagnum bog near the settlement, 22 Au- short pointed setae. Tibia IV slightly thickened gust 1974, P.V. Tuzovsky. Paratype: 1 female, near middle, with distolateral fan-shaped seta and same location as the holotype, 28. August 1974, considerably shorter than tarsus IV. Pedipalps P.V. Tuzovsky. similar to female, but tibia with more numerous Diagnosis. Female. Dorsum with two pairs dorsal setae. of anterior sclerites. Genital plates triangular with Description. Female. Body oval; setae Fch 3 acetabula and 11 thin short setae. Acetabula long, whip-like, other idiosomal setae filiform rather large and equal in sizes, genital opening ap- (Fig. 411). Integument soft, with very fine strips. proximately 1.5 times as long as genital plates. Dorsum with rather large anteromedial and rela- Pedipalpal genu with 3 setae, base of external lat- tively small anterolateral plates. Medial plates eral seta situated proximally to middle of segment; elongate, lateral plates transverse. Anterior and tibia moderately in length with relatively short posterior groups of coxae with rather large apo- with distolateral spine (2.5 times as short as tar- demes (Fig. 412). Medial edge of coxal plates III sus), both ventral setae situated near middle of and IV approximately equal in length, lateral edg- segment. es of coxal plate IV twice as long as lateral edges Male. Medial margin of coxae III in 1.5–2.0 of coxal plates III. Suture line between coxal plates times shorter than medial margin of coxae IV. III and IV obliquely posteromedially-directed. Genital plates fused by anterior and posterior ends, Posteromedial margin of coxal plate IV oblique, acetabula arranged in an arc. Genital setae clus- weakly concave or straight. Surface of all coxae tered in four groups; two anterior groups have a with fine pores. Sclerites, bearing all ventral idio- few setae (8–10) and located as two rows in the somal setae, situated freely on soft cuticle. Genital anterior part of plate, two posterior groups with plate triangular with 3 acetabula and 11 thin short very numerous setae. All sclerites, bearing ventral setae. Lateral margins of genital plates distinctly idiosomal setae, situated on soft cuticle. Tibia II concave, their internal and posterior margins with one rather long ventrodistal sword-like seta weakly concave. Acetabula rather large, separated and small clavate one. Genu IV expanded dors- by wide space. Genital opening 1.5 times longer

204 Water mites of the genus Tiphys

416 414 413

415 Figs 413–416. Tiphys gladiator, female: 413 — capitulum, ventral view, 414 — chelicera, 415 — pedipalp, 416 — distal part of pedipalpal tibia and tarsus. Scale bars: 413–414 = 50, 415 = 45, 416 =20. than genital plates. Excretory pore with small an- external denticles, latter with short spur (Fig. terior sclerite. 420). Capitulum elongate, with large anchoral pro- Measurements (n=1). Body length 610; length jection, both pairs of hypostomal setae subequal of medial margin of coxa III 30–36, length of me- (Fig. 413). Basal segment of chelicera (Fig. 414) dial margin of coxa IV 36–42; length of genital with large dorsal obtuse hump. Cheliceral stylet plates 90, its width 77; length of genital acetabula moderate in size, without ventral teeth. (ac. 1–3): 25, 30, 30; length of capitulum with an- Pedipalp (Fig. 415) rather short. Trochanter choral projection 185; length of basal segment of of pedipalp with one dorsodistal seta. Ventral mar- chelicera 125; length of cheliceral stylet 55; length gin of femur and genu of pedipalp straight. Femur of pedipalpal segment (P-1–5): 33, 90, 57, 90, 45; with 3–4 proximal and 2 dorsodistal setae. Genu length of leg segments: I–Leg. 1–6: 55, 80, 80, with 3 setae, external lateral seta longer than two 105, 125, 140; II–Leg. 1–6: 55, 85, 80, 105, 130, other setae; base of external lateral seta situated 155; III–Leg. 1–6: 75, 80,75, 105, 145, 145; IV– proximally to middle of segment. Pedipalpal tibia Leg. 1–6: 105, 80, 105, 165, 185, 205. relatively short, with relatively large distolateral Male. Dorsal surface, coxa of legs I–II (Fig. spine (2.5 times shorter than tarsus), 2 ventral and 421) similar to female. Anterior apodemes rather 2–3 dorsal setae; internal ventral seta on tibia in- large, extending posteriorly to beneath medial serted on relatively large conic tubercle, base of edge of coxal plate III; posterior apodemes not de- external ventral seta on small tubercle. Tarsus veloped. Coxae IV greatly expanded, their length (Fig. 416) with proximal solenidion, with 5 thin exceeding total length of coxae I–III. Medial mar- setae and 4 thick unequal dorsodistalsetae. gin of coxae III in 1.6 times shorter than length of Two anterior pairs of legs without swimming medial margin of coxae IV; lateral edge of coxa IV setae (Fig. 417). Two relatively short swimming 3 times longer than length of lateral edge of coxa setae present on genu III, 3 on genu IV, 4 long III. Posteromedial margin of coxae IV oblique and swimming setae on tibia III and 5–6 on tibia IV forming bay where external genital field situated; (Fig. 418). Claws of legs I–III larger than claws lateral edge of these coxae convex. Genital plates IV. Internal denticle of claw of legs I–III wide with fused by anterior and posterior ends, acetabula ar- roundish tip, external denticle thin and pointed ranged in arc. Anterior and posterior acetabula (Fig. 419). Claws IV with short internal and long oval, median acetabula rounded. Distance between

205 P. V. Tuzovsky

417

418

419 420

Figs 417–420. Tiphys gladiator, female: 417 — genu, tibia and tarsus of leg I, 418 — genu, tibia and tarsus of leg IV, 419 — claw of leg II, 420 — claw of leg IV. Scale bars: 417–418 = 100, 419–420 =20.

423

421 422

Figs. 421–423. Tiphys gladiator, male: 421 — ventral view, 422 — leg II, 423 — sword-shaped seta. Scale bars: 421, 423 = 100, 422 =50. first and second acetabula twice longer than than rior margins converge under blunt angle and form distance between median and third acetabula. Lat- median incision. Genital setae clustered in four eral edges of genital plate weakly concave; poste- groups; two anterior groups have few setae (8–10)

206 Water mites of the genus Tiphys

424, 427 424 425

427 425, 426

426

Figs 424–427. Tiphys gladiator, male: 424 — genu, tibia and tarsus of leg III, 425 — claw of tarsus II, 426 — claw of tarsus III, 427 — pedipalp. Scale bars: 424, 427 = 50; 425–426 = 20.

430

429

428

Figs 428–430. Tiphys gladiator, male: 428 — basifemur, telofemur, genu and of leg IV, 429 — fan-shaped seta, 430 — tarsus of leg III. Scale bar: 428–430 =80. and located as two rows in anterior part of plate, clavate one. Lateral margins of sword-shaped seta two posterior groups with very numerous setae. (Fig. 423), parallel to each other and not expanded All sclerites, bearing ventral idiosomal setae, situ- distally. Genu III (Fig. 424) with 2, tibia III with ated on soft cuticle. Excretory pore with tiny ante- 6–7 long swimming setae. Genu IV (Fig. 428) ex- rior sclerite. panded dorsoventrally, with convex dorsal surface Basifemur II (Fig. 422), with ventral hump, bearing 10 short, bent setae with round tips. Ven- tibia II with ventrodistal sword-like seta and small tral surface of genu nearly straight, with long

207 P. V. Tuzovsky proximal swimming setae and numerous short 7 (8) Claws of all legs equal in size ...... pointed ventral setae. In addition, swimming setae ...... T. samaraensis sp. n. present in distolateral part of genu; distal edge of 8 (7) Claws of legs I–III considerably larger than genu with small notch and short clavate seta. Tibia claws of legs IV IV (Fig. 398) slightly thickened near middle, with 9 (10) Suture line between coxal plates III and IV concave dorsal and convex ventral sides; with 7 incomplete, obliterated medially (Fig. 222) ...... dorsal, 7 short ventrolateral setae, 6–7 long swim- ...... T. kolymaensis sp. n. ming setae, and distolateral fan-shaped seta (Fig. 10 (9) Suture line between coxal plates III and IV 429). Tarsus IV (Fig. 430) with numerous thin and complete thick setae. Claw of legs I and II approximately 11 (16) Genital plates crescent- or half-moon sha- equal in length, with wide internal denticle and ped narrow external one (Fig. 425). Claws of legs III 12 (13) Posteromedial angle of coxal plate IV well (Fig. 426) asymmetric with 3 unequal denticle developed, acute (Fig. 20) ...... each, large claw with rather long internal and me- ...... T. ornatus Koch, 1836 dian denticles with rounded tips and short pointed 13 (12) Posteromedial angle of coxal plate IV mo- external denticle; small claw with 3 unequal point- derately developed, right or obtuse ed denticles. 14 (15) Anterior genital sclerite large (Fig. 102), Pedipalps similar to those in female, but tibia pedipalps slender (Fig. 109), setae Fch relatively with more numerous dorsal setae (Fig. 427). short (Fig. 99) ...... T. scaurellus Tuzovskij, 1983 Measurements (n=1). Body length 660; length 15 (14) Anterior genital sclerite small (Fig. 208), of seta Fch 130; length of genital opening 70; pedipalps stocky (Fig. 211), setae Fch long (Fig. length of genital plate 145, width 195; length (or 205) ...... T. kamchatkaensis sp. n. diameter) of genital acetabula (ac. 1–3): 38, 32, 16 (11) Genital plates triangular 32; length of capitulum with anchoral projection 17 (20) Two (rarely three) genital setae located in 185; length of basal segment of chelicera 115; the soft integument between anterior genital scle- length of cheliceral stylet 50; length of pedipalpal rite and genital plate on each side segment (P-1–5): 33, 90, 57, 95, 45; length of leg 18 (19) Ventral tubercles on pedipalpal tibia sli- segments: I–Leg. 1–6: 55, 85, 90, 120, 145, 170; ghtly developed, peg-like seta on tibia lesser than II–Leg. 1–6: 65, 90, 90, 120, 135, 185; III–Leg. 0.25 of tarsus length ...... T. bullatus (Thor, 1899) 1–6: 75, 90, 65, 75, 120, 155; IV–Leg. 1–6: 130, 19 (18) Ventral tubercles on pedipalpal tibia well 105, 75, 155, 190, 240; length of medial margin of developed, peg-like seta on tibia 0.4–0.5 of tarsus coxae III 55, length of medial margin of coxae IV length (Fig. 256) ...... T. torris Koenike, 1914 90, length of sword-shaped seta on tibia of legs II 20 (17) All genital setae located on genital plates 120, width 24; length of fan-shaped seta on tibia 21 (22) Genital acetabula very small, distance be- of legs IV 32, width 52–55. tween second and third acetabula approximately Ecology. Sedge-sphagnum bog. equal to two diameters of any acetabulum (Fig. Geographical distribution. Europe, Russia: 303) ...... T. minutiporus sp. n. Yaroslavl Province (Tuzovsky 2005). 22 (21) Genital acetabula relatively large, distance between second and third acetabula smaller than Keys to species of the genus Tiphys diameter of any acetabulum Females 23 (26) Medial margin of coxal plates IV longer 1 (6) Pedipalpal tibia without ventral tubercles than medial margin of coxal plates III 2 (3) Ventral setae on pedipalpal tibia located near 24 (25) Lateral setae on pedipalpal genu located middle of segment (Fig. 146) ...... proximally to middle of segment (Fig. 182); leg ...... T. lapponicus (Neuman, 1880) claw blade moderately developed, with nearly 3 (2) Ventral setae on pedipalpal tibia located dis- straight ventral margin, internal clawlet considera- tally bly wider than external one (Fig. 184) ...... 4 (5) Pedipalpal genu with 3 setae (Fig. 348 ...... T. latipes (Müller, 1776) ...... T. ensifer (Koenike, 1895) 25 (24) Lateral setae on pedipalpal genu located 5 (4) Pedipalpal genu with 2 setae (Fig. 340) ...... distally to middle of segment (Fig. 72); leg claw ...... T. chaunensis sp. n. blade well developed, with convex ventral margin, 6 (1) Pedipalpal tibia with more or less developed internal and external clawlets approximately equal ventral tubercles in wide (Fig. 74) ...... T. scaurus (Koenike, 1892)

208 Water mites of the genus Tiphys

26 (23) Medial margin of coxal plates III and IV 15.(12) Genital plates only partially placed in ge- equal in length nital bay, claws of leg III with 3 clawlets 27 (28) Dorsum with 3 pairs of platelets (Fig. 374), 16 (17) Base of external lateral seta on pedipalpal venter with 4 pairs of lateral platelets (Fig. 375), genu situated distally to middle of segment, exter- genital plates with 7–8 setae each (Figs 376–377) nal seta on pedipalpal tibia placed on large tuber- ...... T. cle, distolateral spine of this tibia 2.0–2.5 times pistillifer (Koenike, 1908) shorter than tarsus (Fig. 256); genu IV greatly ex- 28 (27) Dorsum with 2 pairs of platelets (Fig. 411), panded dorsoventrally with distinct ventroproxi- venter without lateral platelets (Fig. 412), genital mal corner (Fig. 261) ...... T. torris Koenike, 1914 plates with 11 setae each ...... 17 (16) Base of external lateral seta on pedipalpal T. gladiator Tuzovskij, 2003 genu situated in middle of segment, external seta on pedipalpal tibia placed on small tubercle, leng- Males th of distolateral spine of tibia approximately 4 ti- 1 (4) Basifemur II with ventral hump, tibia II with mes shorter than tarsus; genu IV weakly expanded distal spatulate seta or sword-like seta (subgenus dorsoventrally without ventroproximal corner ...... Acercopsis) ...... T. bullatus (Thor, 1899) 2 (3) Tibia II with spatulate seta (Fig. 388), genu 18 (11) Posteromedial portions of coxal plates IV of leg IV (Fig. 390) expanded dorsoventrally with not forming deep genital bay, genital plates placed ventroproximal protrusion, tibia IV with distolate- posterior to coxae IV ral leaf-shaped seta ...... 19 (20) Genu IV not expanded dorsoventrally in ...... T. pistillifer (Koenike, 1908) relation to telofemur IV (Fig. 297) ...... 3 (2) Tibia of leg II with sword-like seta (Fig. 422), ...... T. yaroslavlensis sp. n. genu of leg IV (Fig. 428) expanded dorsoventrally 20 (19) Genu of leg IV well expanded dorsoven- with maximum height near middle of segment, ti- trally in relation to telofemur IV bia IV with a distal fan-shaped seta ...... 21 (22) Tarsus IV considerably longer than tibia ...... T. gladiator Tuzovskij, 2003 IV (Fig. 153) ...... T. lapponicus (Neuman,1880) 4 (1) Basifemur II without ventral hump, tibia of 22 (21) Tarsus and tibia IV approximately equal in leg II without spatulate seta or sword-like length 5 (8) Genu IV with large dorsodistal curved sword 23 (26) Genu IV greatly expanded dorsoventrally, like seta (subgenus Pionides) with large ventral hump near middle of segment 6 (7) Sword like seta reaching to the middle of 24 (25) Setae Hv and Pe fused with posterior mar- genu IV (Fig. 353) ...... T. ensifer (Koenike, 1895) gins of coxal plates II and IV, respectively (Fig. 7 (6) Sword like seta almost as long as genu IV 185); tibia IV with short peg-like seta (Fig. 187); (Fig. 373) ...... T. uchidai Viets, 1956 tarsus IV with 6 short and thick unequal setae; 8 (5) Genu IV without dorsodistal curved sword small leg claw III with 2 subequal denticles (Fig. like seta (subgenus Tiphys) 188) ...... T. latipes (Müller, 1776) 9 (10) Anterior coxal groups fused to each other 25 (24) Setae Hv and Pe free, located on soft inte- medially (Fig. 56) ...... T. anadyrensis sp. n. gument (Fig. 214); tibia IV with rather long lance- 10 (9) Anterior coxal groups separated shaped seta (Fig. 217); tarsus IV with 4 short and 11 (18) Posteromedial portions of coxal plates IV thick subequal setae; small leg claw III with 2 une- forming a rather deep genital bay embracing geni- qual denticles (Fig. 218) ...... tal plates completely or partially ...... T. kamchatkaensis sp. n. 12 (15) Genital plates nearly completely situated 26 (23) Genu IV slightly expanded dorsoventrally, in genital bay, claws of leg III with 2 clawlets without ventral hump, with weakly convex dorsal 13 (14) Posterior coxal group separated (Fig. 30), and ventral margins (Fig. 77) ...... genital bay wide anteriorly, genu IV elongate with ...... T. scaurus (Koenike, 1892) maximum thickness near middle of segment (Fig. 32), tibia of leg III not expanded distally (Fig. 31) Deutonymphs ...... T. ornatus Koch, 1836 1 (2) Claws of all legs equal in size ...... 14 (13) Posterior coxal groups close to each other ...... T. ensifer (Koenike, 1895) (Fig. 112), genital bay narrow anteriorly, genu IV 2 (1) Claws of leg I–III considerably larger than expanded distally (Fig. 115), tibia III expanded claws of legs IV distally (Fig. 114) ...... 3 (10) Dorsal plates present ...... T. scaurellus Tuzovskij, 1983 4 (9) Excretory pore with more or less developed

209 P. V. Tuzovsky anterior sclerite, ventral setae on pedipalpal tibia 8 (9) Dorsal shield well elongate (Fig. 268), ratio short, located distally to middle of segment length/width = 1.70–1.85, with nearly straight la- 5 (6) Pedipalpal tibia with long distal peg-like teral margins ...... T. torris Koenike, 1914 seta, about 0.65 of tarsus length (Fig. 266) ...... 9 (8) Dorsal shield relatively wide, ratio length/ ...... T. torris Koenike, 1914 width < 1.5, with convex lateral margins 6 (5) Pedipalpal tibia with short distal peg-like 10 (11) Setae C1 and C2 on coxal plate I approxi- seta, lesser than 0.50 of tarsus length mately equal in length (Fig. 311) ...... 7 (8) Lateral seta on pedipalpal genu located near ...... T. minutiporus sp. n. middle of segment, distal peg-like setae on tibia 11 (10) Seta C1 on coxal plate I considerably shor- short, about 0.3 of tarsus length (Fig. 121), tibia ter than C2 IV with 5 swimming setae (Fig. 122) ...... 12 (13) Excretory pore plate with setae Ai and Ae ...... T. scaurellus Tuzovskij, 1983 subequal in length (Fig. 400), proximal and distal 8 (7) Lateral seta on pedipalpal genu located near solenidia on tibia II equal in length (Fig. 406) ...... distal end of segment, distal peg-like setae on tibia ...... T. pistillifer (Koenike, 1908) of moderate length, about 0.5 of tarsus length (Fig. 13 (12) Excretory pore plate with short setae Ai 82), tibia IV with 6 swimming setae (Fig. 84) ...... and long setae Ae (Fig. 43), proximal solenidion ...... T. scaurus (Koenike, 1892) on tibia II shorter than distal one (Fig. 50) ...... 9 (4) Excretory pore without anterior sclerite, ven- ...... T. ornatus Koch, 1836 tral setae on pedipalpal tibia long, located near mi- 14 (1) Transverse muscle attachment scar on coxal ddle of segment (Fig. 158) ...... plate III absent ...... T. lapponicus (Neuman,1880) 15 (16) Excretory pore plate wider than long (Fig. 10 (3) True dorsal plates absent 320–323) ...... T. chaunensis sp. n. 11 (12) Lateral seta on pedipalpal genu located 16 (15) Excretory pore plate as long as wide or proximally to middle of segment (Fig. 190), both longer than wide ventral setae of pedipalpal tibia placed distally, 17 (18) Dorsal shield nearly circular (Fig. 124), distance between genital acetabula longer than ratio length/width = 1.15–1.20 ...... diameter of acetabulum (Fig. 189) ...... T. scaurellus Tuzovskij, 1983 ...... T. latipes (Müller, 1776 18 (17) Dorsal shield elongate, ratio length/width 12 (11) Lateral seta on pedipalpal genu located = 1.4–1.6 distally to middle of segment (Fig. 37), both ven- 19 (20) Capitular base wider than long (Fig. 164), tral setae of pedipalpal tibia placed near middle of base of seta C4 and medial portion of suture line segment, distance between genital acetabula shor- between coxal plates II–III well separated from ter than diameter acetabulum (Fig. 35) ...... each other on each side (Fig. 161) ...... T. ornatus Koch, 1836 ...... T. lapponicus (Neuman,1880) 20 (19) Capitular base longer than wide (Fig. 91), Larvae base of seta C4 and medial portion of suture line 1 (14) Transverse muscle attachment scar on coxal between coxal plates II–III close to each other plate III present on each side (Fig. 85) ...... T. scaurus (Koenike, 1892) 2 (7) Dorsal and ventral shields punctate 3 (4) Solenidion on genu II and both solenidia on ACKNOWLEDGEMENTS tibia II (Fig. 369) subequal in length ...... The author expresses sincere gratitude for ...... T. ensifer (Koenike, 1895) valuable materials sent for processing, granted and 4 (3) Solenidion on genu of leg II shorter than both loaned for the study and for useful assistance to solenidia on tibia II perform this investigation to the following col- 5 (6) Proximal solenidion on tibia II shorter than leagues: Dr. K.A. Semenchenko (the Institute of distal one (Fig. 201), solenidion on genu I slightly Biology and Soil Sciences, the Far Eastern Branch longer than both solenidia on tibia I (Fig. 200) ..... of the Russian Academy of Sciences, Vladivostok, ...... T. latipes (Müller, 1776) Russia), Dr. O.L. Makarova (the Institute of prob- 6 (5) Both solenidia on tibia of leg II equal in leng- lems of Ecology and Evolution of the Russian th (Fig. 242), solenidion on genu I and both sole- Academy of Sciences, Moscow, Russia), Dr. R. nidia on tibia I (Fig. 241) approximately equal in Gerecke (the Zoological Institute, the University length ...... T. kolymaensis sp. n. of Tübingen, Germany) and Dr. Henk van der 7 (2) Dorsal and ventral shields with reticulation Hammen ��(The Netherlands). The�� author is espe-

210 Water mites of the genus Tiphys cially thankful to Dr. S.V. Mironov (the Zoologi- Prasad, V. and Cook, D.R. 1972. The taxonomy of wa- cal Institute of the Russian Academy of Sciences, ter mite larvae. Memoirs of the American Entomo� St. Petersburg), Dr. R. Gerecke and H. Smit (Zoo- logical Institute, 18: 1–326. logical Museum of Amsterdam, The Netherlands) Semenchenko, K.A., Abe, H. and Boeskorov, G.G. for critical comments, valuable advices and cor- 2010. [New data on water mite fauna (Acari, Hy- drachnidia, Halacaroidea) of Sakha Republic (Ya- rections proposed to the manuscript. kutia)]. Zoologichesky Zhurnal, 89 (2): 167–177. REFERENCES [in Russian] Sitnikova, L.G.1978 [The main evolutionary trends in Baker, E.W. and Wharton, G.W. 1952. An introduction Acari and problems of their monophyly]. Ento� to acarology. Macmillan, New York. 465 p. mol. Obozrenie, 57 (2): 431–457. [in Russian] Besseling, A.J. 1964. De Nederlandse Watermijten Smit, H. and Van der Hammen, H. 2000. Atlas van de (Hydrachnellae Latreille 1802). Monographieën Nederlandse watermijten (Acari: Hydrachnidia). van de Nederlandsche EntomologischeVereeni� Nederlandse Faunistische Mededelingen, 13: gung, 1: 1–199. 1–272. Conroy, J.C. 1982. New or interesting water mites (Ac- Smith, I.M. 1976. A study of the systematics of the wa- ari: Hydrachnellae) from the ponds in the Fort ter mite family Pionidae (Prostigmata: Parasiten- Whyte nature reserves. International Journal of gona). Memoirs of the Entomological Society of Acarology, 8 (2): 63–79. Canada, 98: 1–249. Cook, D.R. 1956. Preliminary studies of the Tiphysinae Smith, I.M. and Oliver, D.R. 1976. The parasitic asso- of the United States (Acarina: Pionidae). Annals of ciations of larval water mites with imaginal aquat- the Entomological Society of America, 49: 264– ic insects, especially Chironomidae. The Canadian 272. Entomologist, 108: 1427–1442. Cook, D.R. 1974. Water mite genera and subgenera. Smith, I.M. and Oliver, D.R. 1986. Review of parasitic Memoirs of the American Entomological Institute, associations of larval water mites Acari: Parasiten- 21: 1–860. gona: Hydrachnida) with insect hosts. The Cana� Gerecke, R. 1996. Untersuchungen über Wassermilben dian Entomologist, 118: 407–472. der Familie Hydryphantidae (Acari, Actinedida) Smith, I.M. and Cook, D.R. 1991. Water Mites. In: J. in der Westpalaearktis, II. Die Wassermilben der Thorp and A. Covich (Eds.). Ecology and Classifi- Familie Hydryphantidae Piersig, 1896 in den Mit- cation of North American Freshwater Inverte- telmeerländern. Archiv für Hydrobiologie. ��Supple� brates, San Diego: Academic Press, 16: 523–592. ment, 77 (3/4): 337–513. Soar, C.D. and Williamson, W. 1929. The British Hyd� Imamura, T. 1954. Studies on water-mites from Hok- racarina. III. Ray Society, London, 115: 8 +184 + kaido. Journal of the Hokkaido Gakugei Univer� 40 p. sity, B. Supplement, 1: 1–148. Sokolow, I.I. 1940. Hydracarina — vodyanye kleshchi. Koenike, F. 1909. Acarina, Milben. In: A. Brauer (Ed.). Chast’ I. Hydrachnellae. Fauna SSSR (novaya Die Süβwasserfauna Deutschlands. Jena (G. Fi- seriya No. 20. Paukoobraznye, 5 (2) [Hydracarina scher), 12: 13–184. — the aquatic mites. Part I. Hydrachnellae. Fauna Lundblad, O. 1956. Zur Kenntnis süd- und mitteleuro- of the USSR. (nouv. ser., no 20), Arachnides, 5(2)]. päischer Hydrachnellen. Arkiv för Zoologi, 10: Publisher: Nauka, Moscow-Leningrad, 24 s + 511. 1–306. [In Russian] Lundblad, O. 1962. Die Hydracarinen Schwedens. II. Sokolov, I.I. and Yankowskaya, A.I. 1962. [A review of Arkiv för Zoologi, (2) 14 (1): 1–635. Hydracarin fauna of Karelia]. Trudy Zoolog� Lundblad, O. 1968. Die Hydracarinen Schwedens. III. icheskogo Instituta, Moscow, 31: 172–191. [In Arkiv för Zoologi, (2) 21 (1): 1–633. Russian] Mitchell, R.D. 1957. The mating behavior of pionid Sparing, I. 1959. Die Larven der Hydrachnellae, ihre water mites. Annals of the Midland Naturalist, 58: parasitische Entwicklung und ihre Systematik. 360–366. Parasitologische Schriftenreihe, Jena, 10: 1–165. Motaş, C. 1928. Contribution à la connaissance des Hy- Szalay, L. 1964. Víziatkák Hydracarina — Fauna Hun� dracariens français, particulièrement du Sud-Est gariae, 72. Akademiei Kiadó, Budapest: 1–380. de la France. Travaux du Laboratoire Tuzovsky, P.V. 1974. [Distribution of water mites in the d’Hydrobiologie et de Pisciculture de l’Université temporary flood zone of the Rybinskoye water res- de Grenoble, 20: 1–373. ervoir and adjacent water basins]. Trudy Instituta Piersig, G.R. 1897–1900. Deutschlands Hydrachniden. biologii vnutrennikh vod, 25 (28): 202–229. [In Zoologica. Stuttgart, 19 (22): 1–601. Russian] Piersig, G.R. 1901. Hydrachnidae. In: R. Piersig and H. Tuzovsky, P.V. 1983. [A new water mite species of the Lohmann (Eds.). Acarina, Hydrachnidae und Ha- genus Tiphys (Pionidae, Acariformes)]. Biolog� lacaridae. Das Tierreich, 13: 18–336. icheskie nauki, 10: 31–87. [in Russian]

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