Taxonomic Notes on Larvae of Willmannella (Acari

20 Systematic & Applied Acarology Special Publications (1997) 1 Systematic & Applied Acarology Special Publications (1997) 1, 5-18 5

Key to species of Molothrognathus (adult female)

1 Setae sc2 much longer than other dorsal setae ...... 2 - Setae sc subequal to or slightly longer than other dorsal setae ...... 9 2 Taxonomic notes on larvae of Willmannella 2 Setae sc2 and c2 of about equal length ...... 3 - Setae sc2 longer than setae c2 ...... 5 (Acari: Microtrombidiidae) and description 3 Setae f (69-71) about as long as h1 (67-74) ...... colei Swift of a new species from Iran - Setae f shorter than h1 ...... 4 4 Setae sc2, c2, f and h1 relatively long, their lengths being 100, 100, 45-50, ZHI-QIANG ZHANG1 & ALIREZA SABOORI2 and 70, respectively ...... dilucidus Kunzetnov 1 International Institute of Entomology, CAB INTERNATIONAL, 56 QueenÕs Gate, - Setae sc , c , f and h shorter, their length being 72-85, 63-72, 32-38, and 2 2 1 London SW7 5JR, UK 50-60, respectively ...... terrulantus Meyer & Ueckermann 2 5 Setae c shorter than sc and v ; striae simple, not discernible mid-dorsally Department of Entomology, College of Agriculture, Tarbiat Modarres University, 2 1 2 P.O. Box 14155-4838, Tehran, Iran between setae v1 and d ...... crusis Summers & Schlinger - Setae c2 longer than sc1 ...... 6 6 Subequal c2 and v2 longer than sc1 ...... minutus Soliman - Setae c2 longer than setae v2; v2 and sc1 of about equal length ...... 7 Abstract 7 Medial prodorsum with normal striae, not shield-like ...... fulgidus Summer & Schlinger The genus Willmannella Feider (Acari: Trombidioidea: Microtrombidiidae), - Medial prodorsum with weaker striae than surrounding area, shield-like .... 8 previously known only from New Guinea, Java and Europe, is recorded from Middle

8 Dorsal setae sc1 (28), sc2 (84-92), f (30-34), h1 (50-60) relatively long, other Asia (Iran) for the first time. Willmannella kazerunica sp. nov. is described from a dorsal setae >= 20; sc2 ≈ 2.8 times as long as f ...... mehrnejadi sp. nov. larva parasitic on an undetermined dipteran in Kazerun, Iran. A revised definition of larval Willmannella is proposed and the systematic position of this genus within the - Dorsal setae sc1 (13-19), sc2 (57-76), f (13-19), h1 (25-32) relatively long, Microtrombidiidae is discussed. other dorsal setae no longer than 20; sc2 ≈ 4 times as long as f ...... phytocolus Meyer & Ueckermann Key words: Willmannella, systematics, parasite, W. kazerunica sp. nov. 9 Palptibia swollen; palptarsus about 1/3 length of palptibial claw ...... tumpipalpus Meyer & Ueckermann - Palptibia normal ...... 10

10 Setae sc2 longer than c2; setae f longer than h1; palptibial claw about 3/4 Introduction length of palptarsus ...... flatichelus Meyer & Ueckermann - Setae sc2 and c2 about equal length; setae f longer or shorter than h1 ...... 11 The genus Willmannella was erected by Feider (1952) with Ottonia phyllophora 11 Medial prodorsum with weaker striae than surrounding area, looking like a Canestrini, 1897 as the type species, which was described from New Guinea and shield ...... 12 is also known from Java (Thor & Willmann 1947). Feider (1952) included two - Medial prodorsum with normal striae, not looking like a shield ...... 13 other European species in this genus: Willmannella racovitzai (Feider, 1948), 12 Tibia I with a single baculiform solenidion ...... conantae Swift originally placed in Microtrombidium (Enemothrombium) Berlese, and - Tibia I with 2 solenidia sharing an alveolus ...... Willmannella franzi (Willmann, 1950), originally placed in Campylothrombium ...... parmatus Meyer & Ueckermann 6 Systematic & Applied Acarology Special Publications (1997) 1 Systematic & Applied Acarology Special Publications (1997) 1, 19-24 19

Krausse. All three species of this genus were known only from adults then. Larval Willmannella were described by Feider (1956) who successfully reared W. racovitzai larvae in culture from adults. This is the only species of this genus known at both larval and post-larval stages. The host associations of this genus Key to species of the genus Molothrognathus are unknown. In this paper, we describe a new species of Willmannella from a (Prostigmata: Caligonellidae) with larval mite ectoparasitic on an undetermined dipteran in Kazerun, Iran. This is description of a new species from Iran the first record of this genus in Middle Asia and the first record of the host association of larval Willmannella. We also compare this larva with that of W. racovitzai and propose a revised definition for larval Willmannella. We also LAIRONG LIANG1 & ZHI-QIANG ZHANG2 discuss the systematic position of Willmannella within the Microtrombidiidae. 1Department of Environmental & Resource Biology, Fudan University, Shanghai 200433, China 2 The terminology and abbreviations used in this paper are found in Robaux CAB International Institute of Entomology, 56 QueenÕs Gate, London SW7 5JR, UK (1967) and Kethley (1990). Measurements in the description are given in micrometres (µm). Abstract

Molothrognathus mehrnejadi Liang & Zhang sp. nov. is described from females Willmannella kazerunica sp. nov. (Figs. 1-10) collected in pistachio collar soil and bark in Rafsanjan, Iran. A key to world species of Molothrognathus is provided. Description Larva: Holotype partially fed and swollen. Colour in life unknown. Length Key words: Molothrognathus mehrnejadi, new species, Caligonellidae of idiosoma (from anterior end of scutum) 400, maximum width near level of setae c3 and d3 270. A medium-sized larva as measured by total length of legs (IP 972). Introduction Prodorsum covered predominantly by a pentagonal scutum (Fig. 1). Antero-lateral parts of scutum deflexed, with 2 antero-lateral angles directed The genus Molothrognathus was erected by Summers and Schlinger (1955) with ventrally in normal position. Scutum striate longitudinally with punctations M. leptostylus Summers & Schlinger, 1955 as type species. Members of this between striations; striations fade near 2 corners postero-lateral of setae PL and genus have peritremes originating dorsally immediately behind the stylet area anterior to setae AM; striations much denser in central area than deflexed condyle, descending on the laterobasal margins of the stylophore. Summers and antero-lateral corners, striations on latter becoming slightly reticulate in Schlinger (1995) included three species in the genus. Further contributions to posterior part. Scutum slightly convex posteriorly, covering anterior margin of the taxonomy of Molothrognathus were made by Smiley and Morse (1968) and scutellum, but strongly concave laterally at level of trichobothria (S) to Swift (1996) in the US, Soliman (1971) and Meyer and Ueckermann (1989) in accommodate eyes and associated plates. Setae AM and AL nude, their Africa, and Kuznetzov (1978) in the former USSR. Up to now, 13 species of combined length shorter than distance between their insertions (AM + AL < Molothrognathus have been described worldwide. The present paper describes a MA). Setae PL with very fine barbs, located very near postero-lateral angles of new species from Iran. A key to known species of Molothrognathus is also scutum; PL obviously longer than AM and AL, its length subequal to distance provided. ≈ between insertions of AL and PL (PL AP). Trichobothria S thin and nude, The terminology and notation used in this paper are found in Swift (1996). situated antero-mediad of PL; S longer than PL and reaching beyond tip of PL. All measurements are in micrometres. In the description, averages of measurements are given with ranges in parenthesis. 18 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 7

Robaux, P. (1967) ƒtude des larves de Thrombidiidae. I. - La larve de Thrombidium mediterraneum Berlese 1910. Acarologia, 9, 395-410. Robaux, P. (1974) Recherches sur le dŽveloppement et la biologie des acariens ÇThrombidiidaeÈ. MŽmoires du MusŽum National dÕHistoire Naturelle SŽrie A, AM 85, 1-186. Southcott, R.V. (1994) Revision of the larvae of the Microtrombidiinae (Acarina: Microtrombidiidae), with notes on life history. Zoologica, 144, 1-155. Thor, S. & Willmann, C. (1947) Trombidiidae. Tiereich, 71b, 187-541. Vercammen-Grandjean, P.H. (1973) Sur les statuts de la famille des Trombidiidae Leach, 1815 (Acarina: Prostigmata). Acarologia, 15, 102-114. Welbourn, W.C. (1984) Phylogenetic studies on Trombidioidea. In: Griffiths, AL D.A. & Bowman, C.E. (eds.) Acarology VI. Chichester: Ellis Horwood Ltd. pp. S 135-142. Welbourn, W.C. (1991) Phylogenetic studies of the terrestrial Parasitengona. In: PL Dusb‡bek, F. & Bukva, V. (eds.) Modern Acarology 2. Prague, Academia and The Hague, SPB Academic Publishing bv. pp. 163-170. c3 c1 Willmann, C. (1950) Auffallige neue Formen unter den Trombidiiden (Acari). c2

Zoologischer Anzeiger, 145, 1100-1113. d3

d1 d2

Published 31 December 1997

 Systematic and Applied Acarology Society e1 e2

f1 f2 e3

m h1 µ 100

FIGURE 1. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). Dorsal view. 8 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 17

(1991) revised his classification of this group into two families and four subfamilies: Eutrombidiidae (Eutrombidiinae) and Microtrombidiidae s.s. (Microtrombidiinae s.s., Feideriinae and Manriquiinae). In a recent review of larvae of Microtrombidiinae s.l., Southcott (1994) or1 grouped Microtrombidiinae s.l. and Eutrombidiinae in Microtrombidiidae s.l. and included Willmannella in his key to larval genera of Microtrombidiinae s.l., which is the Microtrombidiidae s.s. of Welbourn (1991). 100 µm s.c The concept of Microtrombidiidae sensu Welbourn (1991) is used in this paper. Accordingly, Willmannella is considered here a member of the restricted Microtrombidiinae, based on larval morphology. This genus seems to be most closely related to Campylothrombium Krausse and Fissitrombium Southcott.

1a 1b Acknowledgements

Space and facilities used by the senior author during this study were kindly 2b provided by the Department of Entomology, The Natural History Museum, London. A critical review of this manuscript by Dr. Anne Baker of The Natural History Museum, London is greatly appreciated. 3a

3b References

Canestrini, G. (1897) Acaroidei della N. Guinea. Termeszetrajzi FŸzetek, 20, 461-474. Feider, Z. (1948) Sur quelques acariens de Roumanie appartenant a la famille des Trombidiidae. Bulletin de la Section Scientific de lÕAcademie de la Republique Populaire Roumanie, 30(9), 578-587. Feider, Z. (1952) Impartirea genului Microtrombidium Haller 1882 in mai multe genuri. Buletin Stiintific Sectiunea de Stiinte Biologice, Agronomice, ps 1-3 Geologice si Geografice, 4(3), 587-629. Feider, Z. (1956) O noua larva de acarian din subfamilia Microtrombidiinae si importanta sa in tipizarea larvelor si in explicarea factorilor morfogenetici. Academia Republique Populaire Roumanie Filiala Iasi Studii si Cercetari Stiintifice Biologie si Stiintifice Agricole, 7(1), 10-33. Kethley, J. (1990) Acarina: Prostigmata (Actinedida). In Dindal, D.L (ed.) Soil Biology Guide. New York, Chichester, Brisbane, Toronto & Singapore: FIGURE 2. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). Ventral view. John Wiley & Sons. pp. 667-677. 16 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 9

W. racovitzai are congeneric. Southcott (1994) provided a generic definition of Standard measurements of scutum and setae: AM 30; AL 25; PL 46; S 63; AA larval Willmannella based on the description of W. racovitzai by Feider (1956). 72; AW 117; PW 135; SB 99; ASB 135; PSB 33; SD 168; MA 72; AP 45; W With the description of W. kazerunica, the generic limit of Willmannella can 141 (at level of PL). now be more clearly defined. A revised description of Willmannella larvae is A pair of suboval eye plates (Fig. 1) located postero-laterad of scutum, 35 proposed below: long, 27 wide. Eye plates smooth. Anterior eyes (diameter 9) about twice as large as posterior eyes (diameter 4). Prodorsal scutum punctate and with longitudinal striations; antero- Hysterosoma dorsally with 22 setae arranged in 4 rows (c1-3, d1-3, e1-3, f1-2) lateral corners deflexed and with sparser striations than median part of and terminally with a row of 4 setae (h1-2). All these setae with very fine barbs. scutum. Scutellum punctate but with a pair of longitudinal striate Setae c1-2 on scutellum and others on individual small plates or platelets. strips (of not more three lines) near insertions of setal pair c1. Other Scutellum wider than scutum, punctate but with a pair of longitudinal striate dorsal setae on hysterosoma each arise from a small plate but two plates strips (of not more than three lines) laterad of insertions of c1. Measurements of nearest to scutellum significantly larger than others. Terminal setae h2 scutellum: HS 54; LSS 168; SS (c1) 48; SL 69. Maximum diameters of plates: more than twice as long as setae h1. Coxae I each with 1 nude and 1 c2 24, d1 37, e1 and h2 15, others 8. Dorsal hysterosomal setae similar in barbed seta, coxae II and III each with 1 barbed seta. Leg I and II each length. Setae h2 (95) more than twice as long as h1. with a pair of normal claws and a claw-like empodium, leg III with 2 Opisthosoma ventrally with 3 pairs of setae (ps1-3) and an anus (Fig. 2). asymmetric, modified claws but normal claw-like empodium; claws Setae ps finely barbed, 30-36 long, each with a small basal plate of diameter 6. each with a pair of barbs subterminally. Subcapitular setae short and Anus 28 long, at level of ps2 (right one). One ps2 (left) slightly antero-mediad of thick, branched with several blunt digitations. its normal position. Podosoma striate ventrally except for punctate coxal plates (Fig. 2). The systematic position of Willmannella within the family Striations longitudinal between pairs of coxal fields I, II and III, but transverse Microtrombidiidae is uncertain, in part due to changes in the status and extent of between coxal fields II and III on each side. Coxae I 83 long; coxal setae 1a family-group taxa within this group. nude, 31 long, located near basal 1/3 of anterior margin; coxal setae 1b barbed, Feider (1952) placed Willmannella in Microtrombidiinae s.l. when he erected at least 28 long (tip broken), located near middle of distal margin. Coxae II 71 it. Feider (1956) compared larval W. racovitzai with larvae of other genera in long; coxal setae 2b sparsely barbed, 28 long, located near antero-lateral corner. Microtrombidiinae s.l. and divided the then known species of this subfamily into Coxae III 60 long; coxal setae 3b finely barbed, located slightly posterior to four categories; he did not create any group names such as tribes for these mid-anterior margin. Intercoxal setae 3a (15) with very fine barbs, each arise categories. Feider (1956) considered W. racovitzai to be the most derived species from a platelet (Fig. 2). Urstigmata 15 in diameter, between distal corners of in terms of its adaptation to parasitism. coxae I and II. Vercammen-Grandjean (1973) split Microtrombidiinae s.l. into three Leg I (348, excluding claws) about twice as long as scutum. Trochanter I subfamilies within Trombidiidae s.l.: Microtrombidiinae s.s., Feideriinae and 39 long, with 1 subdorsal seta, barbed (Fig. 2). Femur I 62 long, with 1 nude Georgiinae. He placed Willmannella in Georgiinae. Vercammen-Grandjean seta near basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle and (1973) noted that only the larval stage of this genus was known then; this is not 3 barbed setae (2 dorsal and 1 postero-lateral) between 2/3 to 3/4 of segment correct. Feider (1952) erected Willmannella based on adults and reared larvae length (Fig. 3); all setae not longer than 1/2 length of femur. Genu I 23 long, from adults that he identified as W. racovitzai (Feider 1956). slightly longer than wide, with 2 dorsal solenidia (subequal s1 and s2, 43) near Welbourn (1984) included four tribes (Eutrombidiini, Manriquiini, basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle and 2 barbed Microtrombidiini and Feideriini) in the revised Microtrombidiinae s.l., but setae (1 dorsal and 1 postero-lateral) at » 3/4 of segment length; all setae about Willmannella was not listed as a member of any of these tribes. Welbourn 1.5 as long as length of genu, except dorsal seta (subequal to length of genu); 10 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 15

Remarks Palptibia typically bears three setae in larval Trombidioidea and related mites. In this new species, we observed only one nude seta and a large s1 f1 membranous area on the palptibia, although two sclerotized seta-like structures s2 f2 were present at the proximal end underneath the membrane. Robaux (1974) 3 likewise observed two setae and one seta-like structure on the palptibia of k Campylothrombium barbarum (Lucas) and one seta and two seta-like structures on the palptibia of Robauxia thaumapilosum (Robaux). He considered these seta-like structures on the palptibia to be true setae. We agree. We also observed a dorsal pit on the palpgenu of W. kazerunica; this pit could be a reduced seta. Most larval microtrombidiids do not have setae on the palpgenu. Southcott (1994) recently described Buandikia anneae and Patagonella echeverryae and observed a minute seta on the palpgenu and another on the palpfemur of both species. The only other genus in the Trombidioidea that has s k a seta on the palpgenu is Podothrombium of the Trombidiidae. f1

4 f2 Discussion

Feider (1956) gave a rather detailed description of W. racovitzai. His description contains many useful data but also some ÒproblemsÓ. He noted in his description (Feider 1956: 11) that lateral to a row of four s setae (each arising from a rectangular sclerite) is a group of two short setae, which are shown in his Fig. 1, but on page 31, he wrote in his French description of the mite: Òˆ lÕextrŽmitŽ grands, se trouve un groupe de trois petits 5 poilsÓ. One of the statements must be wrong. It is rather unsusal that these setae are found only on one side. In his Table 2 on page 8, Feider showed five setae on femur I and six setae on femur II; but in his Figs. 10 and 11, he showed five setae on both. However, most microtrombidiids have 6 setae on femur I and 5 setae on femur 100 mm II, as observed in W. kazerunica. His illustrations seem out of proportion, probably drawn free-hand instead

of from a camera lucida. For example, setae h1 is shown <4 times as long as h2 in his Fig. 1, but it is over 5 times as long as h according to his measurements FIGURE 3-5. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). 3, femur, 2 genu and tibia I; 4, femur, genu and tibia II; 5, femur, genu and tibia III. All in (page 11). postero-lateral view. Only a re-examination of FeiderÕs specimen will resolve these problems. Despite all these inconsistencies, there seems little doubt that W. kazerunica and ZHANG & SABOORI: A new species of Willmannella from Iran 11

e 6 z2

50 mm

w e 7

FIGURE 6-8. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). 3, tarsus, claws and empodium I; 4, tarsus, claws and empodium II; 5, tarsus, claws and empodium III. All in postero-lateral view. 12 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 13 distal microseta k minute, 4 long. Tibia I 53 long, with solenidia f1 (45) at » ventrals, 4 antero-laterals and 1 postero-lateral) in distal half of segment; 3/4 and f2 (26) » 9/10 of segment length; microseta k (5) distal to f1; 6 barbed postero-lateral seta at 1/3 of segment length much longer than other setae; setae: 1 ventral at » 1/4 and another ventral at » 3/4, 1 antero-lateral at » 2/3 of ventral seta at distal end thicker and shorter than other setae; distal antero-lateral segment length, 1 antero-lateral and 2 postero-lateral distal to level of f1. seta subdorsal in location and branched along distal 2/3. Anterior claws (20) Tarsus I 88 long, with dorsal solenidia w (28) at » 1/4 and eupathidium z1 (44 longer than posterior claw (10); both thicker but shorter than claw-like mm) at » 2/3 of segment length (Fig. 6); postero-lateral z2 (18) near distal end; empodium (48); claws each with a pair of very small barbs near distal ends. famulus e (6) proximal to w; 17 barbed setae: 2 proximal to and 2 at level of Gnathosoma 94 wide, with posterior margin concave (Fig. 9). Palpi w, 8 between levels of w1 and z1, and 5 distal to level of z1. 2 claws (18) tapering, 60 long. Palptrochanter reduced. Palpfemur without setae, 24 long. shorter but thicker than claw-like empodium (28); claws each with a pair of Palpgenu 18 long, without obvious setae but with a dorsal pit (Fig. 10). small barbs near distal ends. Palptibia 18 long, with a nude seta (34) postero-laterally and a large (16 long Leg II (305, excluding claws) shorter than leg I. Trochanter II 33 long, and 7 wide) membranous area enclosing base of tibial claw; 2 sclerotized seta- with 1 dorsal barbed seta (Fig. 1). Femur II 58 long, with 1 nude seta near like structures at proximal end underneath membrane; tibial claw 9 long, basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle, and 2 recurved in lateral view and deeply divided. Palptarsus small (8 long and 8 in barbed setae (1 dorsal and 1 postero-lateral) between 2/3Ð3/4 of segment length diameter), with a postero-lateral solenidion w (7 m), a distal eupathidium z (6 (Fig. 4); 2 setae near middle longer than 1/2 length of femur. Genu II 22 long, um) and 6 nude setae; 1 postero-lateral seta 27 long and 1 ventral seta 39 long. slightly longer than wide, with 1 dorsal solenidion s (47) at » 1/3 of segment Chelicerae robust; cheliceral base 90 long and 30 wide; cheliceral blade 18 long, length, 2 barbed setae (1 antero-lateral and 1 postero-lateral) near 3/5 of segment sickled-shaped with 2 small teeth subterminally. Sclerotized oral ÒringÓ not length; all setae about 1.5 as long as length of genu; microseta k almost closed dorsally, with membranes outside it; outer ring margin with » 30 ÒteethÓ, imperceptible. Tibia II 48 long, with solenidia f1 (39) at » 1/8 and f2 (18) » while inner ring margin finely dentate; maximum distance between outer margin 7/8 of segment length; 5 barbed setae: 1 ventral at » 1/3 and 1 postero-lateral at 36. Oral setae (or1) nude, 13 long. Subcapitular setae (s.c) short (15) and thick » 2/3 of segment length, 2 antero-lateral and 1 postero-lateral slightly proximal (4 in diameter at base), branched along distal 2/3 with 9 blunt digitations (Fig. to level of f2. Tarsus II 74 long, tapering distally, with dorsal solenidia w (21) 9). Ventral surface of subcapitulum tongue-shaped and punctate posterior to slightly proximal to middle of segment length (Fig. 7); famulus e (4) lateral to setae s.c. w; 14 barbed setae: 5 proximal and 9 distal to level of w. 2 claws (18) shorter but thicker than claw-like empodium (31); claws each with a pair of small barbs Etymology near distal ends. This species is named after the type locality Kazerun in Iran. Leg III (319, excluding claws) shorter than leg I but longer than leg II. Trochanter III 47 long, with 1 barbed dorsal seta (Fig. 2). Femur III 66 long, Type specimen with 1 nude seta near basal end, 1 ventral barbed seta at » 1/3, 1 antero-lateral This species is described from a single larva [slide identification ARS- barbed seta at » 2/3 and 1 dorsal barbed seta at » 3/4 of segment length (Fig. 19950406-2] collected by H. Ostovan from an undermined dipteran in Kazerun, 5). Genu III 24 long, slightly longer than wide, with 1 dorsal solenidion s (50) Iran. The holotype larva is deposited in The Zoological Museum, College of at » 1/4 of segment length, 2 barbed setae (1 antero-lateral and 1 postero-lateral) Agriculture, University of Tehran, Tehran, Iran. near mid-segment. Tibia III 60 long, with 5 barbed setae: 1 subventral at » 1/3 and 1 postero-lateral at » 2/3 of segment length, 2 antero-laterals and 1 postero- Diagnosis of larva lateral within distal 1/3 of segment. Tarsus III trapezoidal in lateral view (Fig. This species differs from W. racovitzai in many characters and can readily 8), 62 long dorsally and 54 long ventrally, with 13 barbed setae: 4 (1 ventral, 2 be distinguished from the latter by the following: (1) scutum length subequal to, antero-laterals and 1 postero-lateral) in proximal half and 9 (2 dorsals, 2 not significantly greater than, scutellum width; (2) eye plate smooth, not striate; 12 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 13 distal microseta k minute, 4 long. Tibia I 53 long, with solenidia f1 (45) at » ventrals, 4 antero-laterals and 1 postero-lateral) in distal half of segment; 3/4 and f2 (26) » 9/10 of segment length; microseta k (5) distal to f1; 6 barbed postero-lateral seta at 1/3 of segment length much longer than other setae; setae: 1 ventral at » 1/4 and another ventral at » 3/4, 1 antero-lateral at » 2/3 of ventral seta at distal end thicker and shorter than other setae; distal antero-lateral segment length, 1 antero-lateral and 2 postero-lateral distal to level of f1. seta subdorsal in location and branched along distal 2/3. Anterior claws (20) Tarsus I 88 long, with dorsal solenidia w (28) at » 1/4 and eupathidium z1 (44 longer than posterior claw (10); both thicker but shorter than claw-like mm) at » 2/3 of segment length (Fig. 6); postero-lateral z2 (18) near distal end; empodium (48); claws each with a pair of very small barbs near distal ends. famulus e (6) proximal to w; 17 barbed setae: 2 proximal to and 2 at level of Gnathosoma 94 wide, with posterior margin concave (Fig. 9). Palpi w, 8 between levels of w1 and z1, and 5 distal to level of z1. 2 claws (18) tapering, 60 long. Palptrochanter reduced. Palpfemur without setae, 24 long. shorter but thicker than claw-like empodium (28); claws each with a pair of Palpgenu 18 long, without obvious setae but with a dorsal pit (Fig. 10). small barbs near distal ends. Palptibia 18 long, with a nude seta (34) postero-laterally and a large (16 long Leg II (305, excluding claws) shorter than leg I. Trochanter II 33 long, and 7 wide) membranous area enclosing base of tibial claw; 2 sclerotized seta- with 1 dorsal barbed seta (Fig. 1). Femur II 58 long, with 1 nude seta near like structures at proximal end underneath membrane; tibial claw 9 long, basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle, and 2 recurved in lateral view and deeply divided. Palptarsus small (8 long and 8 in barbed setae (1 dorsal and 1 postero-lateral) between 2/3Ð3/4 of segment length diameter), with a postero-lateral solenidion w (7 m), a distal eupathidium z (6 (Fig. 4); 2 setae near middle longer than 1/2 length of femur. Genu II 22 long, um) and 6 nude setae; 1 postero-lateral seta 27 long and 1 ventral seta 39 long. slightly longer than wide, with 1 dorsal solenidion s (47) at » 1/3 of segment Chelicerae robust; cheliceral base 90 long and 30 wide; cheliceral blade 18 long, length, 2 barbed setae (1 antero-lateral and 1 postero-lateral) near 3/5 of segment sickled-shaped with 2 small teeth subterminally. Sclerotized oral ÒringÓ not length; all setae about 1.5 as long as length of genu; microseta k almost closed dorsally, with membranes outside it; outer ring margin with » 30 ÒteethÓ, imperceptible. Tibia II 48 long, with solenidia f1 (39) at » 1/8 and f2 (18) » while inner ring margin finely dentate; maximum distance between outer margin 7/8 of segment length; 5 barbed setae: 1 ventral at » 1/3 and 1 postero-lateral at 36. Oral setae (or1) nude, 13 long. Subcapitular setae (s.c) short (15) and thick » 2/3 of segment length, 2 antero-lateral and 1 postero-lateral slightly proximal (4 in diameter at base), branched along distal 2/3 with 9 blunt digitations (Fig. to level of f2. Tarsus II 74 long, tapering distally, with dorsal solenidia w (21) 9). Ventral surface of subcapitulum tongue-shaped and punctate posterior to slightly proximal to middle of segment length (Fig. 7); famulus e (4) lateral to setae s.c. w; 14 barbed setae: 5 proximal and 9 distal to level of w. 2 claws (18) shorter but thicker than claw-like empodium (31); claws each with a pair of small barbs Etymology near distal ends. This species is named after the type locality Kazerun in Iran. Leg III (319, excluding claws) shorter than leg I but longer than leg II. Trochanter III 47 long, with 1 barbed dorsal seta (Fig. 2). Femur III 66 long, Type specimen with 1 nude seta near basal end, 1 ventral barbed seta at » 1/3, 1 antero-lateral This species is described from a single larva [slide identification ARS- barbed seta at » 2/3 and 1 dorsal barbed seta at » 3/4 of segment length (Fig. 19950406-2] collected by H. Ostovan from an undermined dipteran in Kazerun, 5). Genu III 24 long, slightly longer than wide, with 1 dorsal solenidion s (50) Iran. The holotype larva is deposited in The Zoological Museum, College of at » 1/4 of segment length, 2 barbed setae (1 antero-lateral and 1 postero-lateral) Agriculture, University of Tehran, Tehran, Iran. near mid-segment. Tibia III 60 long, with 5 barbed setae: 1 subventral at » 1/3 and 1 postero-lateral at » 2/3 of segment length, 2 antero-laterals and 1 postero- Diagnosis of larva lateral within distal 1/3 of segment. Tarsus III trapezoidal in lateral view (Fig. This species differs from W. racovitzai in many characters and can readily 8), 62 long dorsally and 54 long ventrally, with 13 barbed setae: 4 (1 ventral, 2 be distinguished from the latter by the following: (1) scutum length subequal to, antero-laterals and 1 postero-lateral) in proximal half and 9 (2 dorsals, 2 not significantly greater than, scutellum width; (2) eye plate smooth, not striate; 14 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 11

(3) plates c2 and d1 oval, not subrectangular; (4) setae c1 mediad of, not lateral to, striation on scutellum; (5) setae h2 between 2-3, not 4, times as long as setae h1; (6) coxal setae 1b near middle of distal margin, not near antero-distal corner, of coxa I; (7) genu I with 4, not 3, barbed setae; (8) palptibia with 1, not 2, nude setae; (9) cheliceral blade with 2 , not 1, subterminal teeth ventrally; (10) leg lengths I-II-III 348-305-310, not 301-251-265.

w 9 or1 10

s.c

50 mm FIGURE 1. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). 9, gnathosoma, ventral view; 10, palp, dorsal view. 10 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 15

4 f2 Discussion

of from a camera lucida. For example, setae h1 is shown <4 times as long as h2 in his Fig. 1, but it is over 5 times as long as h according to his measurements FIGURE 3-5. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). 3, femur, 2 genu and tibia I; 4, femur, genu and tibia II; 5, femur, genu and tibia III. All in (page 11). postero-lateral view. Only a re-examination of FeiderÕs specimen will resolve these problems. Despite all these inconsistencies, there seems little doubt that W. kazerunica and 16 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 9

strips (of not more three lines) near insertions of setal pair c1. Other Scutellum wider than scutum, punctate but with a pair of longitudinal striate dorsal setae on hysterosoma each arise from a small plate but two plates strips (of not more than three lines) laterad of insertions of c1. Measurements of

nearest to scutellum significantly larger than others. Terminal setae h2 scutellum: HS 54; LSS 168; SS (c1) 48; SL 69. Maximum diameters of plates: more than twice as long as setae h1. Coxae I each with 1 nude and 1 c2 24, d1 37, e1 and h2 15, others 8. Dorsal hysterosomal setae similar in barbed seta, coxae II and III each with 1 barbed seta. Leg I and II each length. Setae h2 (95) more than twice as long as h1. with a pair of normal claws and a claw-like empodium, leg III with 2 Opisthosoma ventrally with 3 pairs of setae (ps1-3) and an anus (Fig. 2). asymmetric, modified claws but normal claw-like empodium; claws Setae ps finely barbed, 30-36 long, each with a small basal plate of diameter 6. each with a pair of barbs subterminally. Subcapitular setae short and Anus 28 long, at level of ps2 (right one). One ps2 (left) slightly antero-mediad of thick, branched with several blunt digitations. its normal position. Podosoma striate ventrally except for punctate coxal plates (Fig. 2). The systematic position of Willmannella within the family Striations longitudinal between pairs of coxal fields I, II and III, but transverse Microtrombidiidae is uncertain, in part due to changes in the status and extent of between coxal fields II and III on each side. Coxae I 83 long; coxal setae 1a family-group taxa within this group. nude, 31 long, located near basal 1/3 of anterior margin; coxal setae 1b barbed, Feider (1952) placed Willmannella in Microtrombidiinae s.l. when he erected at least 28 long (tip broken), located near middle of distal margin. Coxae II 71 it. Feider (1956) compared larval W. racovitzai with larvae of other genera in long; coxal setae 2b sparsely barbed, 28 long, located near antero-lateral corner. Microtrombidiinae s.l. and divided the then known species of this subfamily into Coxae III 60 long; coxal setae 3b finely barbed, located slightly posterior to four categories; he did not create any group names such as tribes for these mid-anterior margin. Intercoxal setae 3a (15) with very fine barbs, each arise categories. Feider (1956) considered W. racovitzai to be the most derived species from a platelet (Fig. 2). Urstigmata 15 in diameter, between distal corners of in terms of its adaptation to parasitism. coxae I and II. Vercammen-Grandjean (1973) split Microtrombidiinae s.l. into three Leg I (348, excluding claws) about twice as long as scutum. Trochanter I subfamilies within Trombidiidae s.l.: Microtrombidiinae s.s., Feideriinae and 39 long, with 1 subdorsal seta, barbed (Fig. 2). Femur I 62 long, with 1 nude Georgiinae. He placed Willmannella in Georgiinae. Vercammen-Grandjean seta near basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle and (1973) noted that only the larval stage of this genus was known then; this is not 3 barbed setae (2 dorsal and 1 postero-lateral) between 2/3 to 3/4 of segment correct. Feider (1952) erected Willmannella based on adults and reared larvae length (Fig. 3); all setae not longer than 1/2 length of femur. Genu I 23 long, from adults that he identified as W. racovitzai (Feider 1956). slightly longer than wide, with 2 dorsal solenidia (subequal s1 and s2, 43) near Welbourn (1984) included four tribes (Eutrombidiini, Manriquiini, basal end, 2 barbed setae (1 antero-lateral and 1 ventral) near middle and 2 barbed Microtrombidiini and Feideriini) in the revised Microtrombidiinae s.l., but setae (1 dorsal and 1 postero-lateral) at » 3/4 of segment length; all setae about Willmannella was not listed as a member of any of these tribes. Welbourn 1.5 as long as length of genu, except dorsal seta (subequal to length of genu); 8 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 17

1a 1b Acknowledgements

3b References

Canestrini, G. (1897) Acaroidei della N. Guinea. Termeszetrajzi FŸzetek, 20, 461-474. Feider, Z. (1948) Sur quelques acariens de Roumanie appartenant a la famille des Trombidiidae. Bulletin de la Section Scientific de lÕAcademie de la Republique Populaire Roumanie, 30(9), 578-587. Feider, Z. (1952) Impartirea genului Microtrombidium Haller 1882 in mai multe genuri. Buletin Stiintific Sectiunea de Stiinte Biologice, Agronomice, ps 1-3 Geologice si Geografice, 4(3), 587-629. Feider, Z. (1956) O noua larva de acarian din subfamilia Microtrombidiinae si importanta sa in tipizarea larvelor si in explicarea factorilor morfogenetici. Academia Republique Populaire Roumanie Filiala Iasi Studii si Cercetari Stiintifice Biologie si Stiintifice Agricole, 7(1), 10-33. Kethley, J. (1990) Acarina: Prostigmata (Actinedida). In Dindal, D.L (ed.) Soil Biology Guide. New York, Chichester, Brisbane, Toronto & Singapore: FIGURE 2. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). Ventral view. John Wiley & Sons. pp. 667-677. 18 Systematic & Applied Acarology Special Publications (1997) 1 ZHANG & SABOORI: A new species of Willmannella from Iran 7

Zoologischer Anzeiger, 145, 1100-1113. d3

d1 d2

Published 31 December 1997

 Systematic and Applied Acarology Society e1 e2

f1 f2 e3

m h1 µ 100

FIGURE 1. Willmannella kazerunica Zhang & Saboori sp. nov. (larva). Dorsal view.