Systematic Review of Feather Mites of the Genus Sturnotrogus MIRONOV

BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE ENTOMOLOGIE. 76:55-81, 2006 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUU RWETE NSCHAPPEN ENTOMOLOGIE, 76: 55-81, 2006

Systematic review of feather mites of the genus Sturnotrogus MIRONOV, 1989 (Astigmata: Pteronyssidae) from starlings (Passeriformes: Sturnidae) in Africa and Europe by Sergey V. MIRONOV, Georges WAUTHY

Abstract general appearance, representatives of the genus Sturno trogus are medium-sized pteronyssids (adults about 300- We prese nt an improved diagnosis of the genu s Sturnotrogus 500 f..im in length) with relatively well-developed dorsal MIRONOV, 1989 and a key and diagnoses to species detected or poten tially expected on starlings (Sturnidae) in Africa and Europe. Two shields. Mites of this genus are commonly located on species groups, truncatus and subtruncatus, are recog ni zed within ventral surface the vanes of the primary flight feathers. the genus. Five new species of the genus Sturnotrogus are described The genus Sturnotrogus was originally established from starlings in Africa: Sturnotrogus acridorheres sp. n. from (MIRONOV, 1989) for six species removed from the Acridoteres tristis (LINNAEUS , 1766), S. /amprotornis sp. n. from polyphyletic genus Pteron.yssoides HULL , 1931 , which Lamprotornis splendidus (VIEILLOT, 1822) (type host), L. chalcums NORDMANN, 1835, L. cha/ybaeus (HEMPRICH & EHRENBERG , 1828), even after the generic revision of Pteronyssidae (FACCINI L. iris (OUSTALET, 1879), L. nitens (LINNAEUS , 1766), L. purpurop & ATYEO, 1981) continued to incorporate almost all pter tents RUPPELL, 1845, L. corruscus corruscus NoRDMANN , 1835, L. onyssids with free epimerites I in both sexes. At present, chloropterus elisabeth (STRESEMANN, 1924), S. onychognathi sp. n. among pteronyssid genera restricted to passeriforms, from Onychognathus moria (LI NNAUES, 1766), S. poeopterae sp. n. from Po eoptera lugubris BONAPARTE, 1854, and S. superbus sp. n. the five genera, Mouchetia GAUD, 1961, Pteronyssoides, from Lamprotornis superbus RuPPELL, 1845. Scutulanyssus MrRONOV, 1985, Sturnotrogus, and Tima Host associations of the genus Sturnotrogus are summari zed and lin.yssus MIRONOV, 2001, constitute the Pteronyssoides briefly di scussed. Representatives of th e truncatus spec ies group are generic group, which is characterized by free epimerites associated with starling species distributed in Europe, Africa, and I. Within this group, Sturnotrogus is differentiated by the Central and Oriental Asia ; species of the subtruncatus group occur on starlings and fan-tailed fl ycatchers of th e genus Rhipidura VIGORS following combination of features: in males, the opistho & HoRSFI ELD, 1827 (Rhipiduridae) in the Indo-Malayan region. It somal lobes are short and wide and are truncated api is hypothesized that Sturnidae are the original hosts of the ge nu s cally, the terminal cleft is not expressed, and the opistho Sturnotrogus, while Rhipiduridae are the secondary hosts. somal membrane is a narrow entire band on the posterior Key words: Pteronyssidae, Stumotrogus, systematics, host associa margin of opisthosoma (between bases of setae ps2); in tions, Sturnidae. females, the lateral sclerites (fragments of hysteronotal shield) are always clearly outlined and have a lanceolate form, the opisthosomal sclerite is a large unpaired plate Introduction or (if this sclerite is not split from the hysteronotal shield) the opisthosomal area of the hysteronotal shield is not Feather mites of the family Pteronyssidae (Astigmata: dissected medially. Regarding the status of the genus, Analgoidea), which currently includes over 150 species in the review of supraspecific feather mite taxa of the in 23 genera (FACCINI & ATYEO, 1981; GAUD & ATYEO, world by GAUD & ATYEO (1996), Sturnotrogus was 1996; MIRONOV, 2001, 2005; MIRONOV & WAUTHY, decreased in rank and treated as a subgenus of the genus 2005a), are mainly distributed on birds from the orders Scutulanyssus. We suggest that the decreasing of rank Passeriformes and Piciformes, with only a few species and inclusion into Scutulanyssus are groundless, because recorded from the Coraciiformes. As highly specialized the authors of that review did not take into consideration permanent ectoparasites inhabiting plumage, pteronys the general structure of the female hysteronotal shield sids are typical representatives of the feather mite mor for determination of pteronyssid genera. In contrast to photype adapted to inhabit vanes of large feathers. Within most feather mite families, the structure of the female this family, the genus Sturnotrogus MIRONOV, 1989 is hysteronotal shield varies greatly among different group one of ten pteronyssid genera restricted to Passeriformes ings of Pteronyssidae, and the general arrangement and (FACCINI & ATYEO, 1981; MIRONOV, 2001). Up to now, form of hysteronotal shield fragments may be used for this genus included eight species known from passerines quite clear diagnoses of pteronyssid taxa at the generic from two avian families, six from starlings (Sturnidae) rank (MIRONOV, 2001; MIRONOV & WAUTHY, 2005a; and two from fan-tailed flycatchers (Rhipiduridae). In MIRONOV & GALLOWAY, 2006). 56 Sergey V. MIRONOV, Georges WAUTHY

The two initially discovered species of Sturnotrogus (1996). Regarding the terms used for hysteronotal shield were described by TRouESSART (1885) in the context of fragments in females, we generally follow the scheme the genus Pteronyssus RoBIN, 1877: Pteronyssus trunca proposed by MIRONOV (1992) and MIRONOV and WAU tus TROUESSART, 1885 was described from the common THY (2005a). starling, Stumus vulgaris (LINNAEUS, 1758) in Europe, All measurements in the descriptions are given in and Pteronyssus subtruncatus TRouESSART, 1885 from micrometres (pm). A full set of measurements is given the hill myna, GraCLtla religiosa (L!NNAEUS , 1766), in only for the holotype (male) and one paratype (female); Indonesia. For over 80 years, experts recognized only the range of idiosomal size (length, width) is displayed these two pteronyssid species on starlings. HuLL (1931; for the rest of the paratype specimens. 1934) placed these species in the genus Pteronyssoides, Measuring technique for particular structures: which was established for pteronyssids with free epimer (i) distance between different pairs of setae is the short ites I. GAUD and co-authors (GAUD & Mouc HET, 1959; est distance between the transverse levels formed by GAUD & TILL , 1961; GAUD, 1977) reported "Ptero setae of respective pairs; nyssoides subtruncatus" from two starling species, Acri (ii) prodorsal shield length is measured along midline, dotheres tristis (LINNAEUS, 1766) and Poeoptera lugu and width is greatest width at posterior margin; bris BONAPARTE, 1854 in Africa; however, these authors (iii) hysterosoma is measured from the level of sejugal actually dealt with new species (see remarks for new furrow to bases of setae h3 ; species from each respective host). Later, GAUD (1968) (iv) hysteronotal shield length in males is the greatest described two species from Rennell Island, one from length from the anterior margin to bases of setae h3; the Rennell starling, Aplonis insularis MAYR, 1931, and width is measured at anterior margin; one from the Rennell fan-tailed flycatcher, Rhipidura (v) distance between prodorsal and hysteronotal shield rennelliana MAYR , 1931 (Rhipiduridae). The latter case in both sexes and the length of transventral sclerite represented the first record of Sturnotrogus species from in males are measured along midline; rhipidurids. Finally, three more species were described (vi) central sclerite length in females is the greatest from starlings, two from Asia (MIRONOV, 1987, 1989) length; width is measured at anterior margin. and one from Africa (MIRONOV & KoPIJ, 2000), and Specimen depositories and reference accession one more species was found on rhipidurids in Vietnam numbers are given using the following abbreviations: (M!RONOV, 1993). AMNH - American Museum of Natural History, New In the course of our investigations offeather mites asso York, USA; AMU - A. Mickiewicz University, Poznan, ciated with African passerines (MIRONOV & WAUTHY, Poland; GAUD- uncatalogued collection of Prof. J. GAUD 2005a-d, 2006a-c), we detected five new Stumotrogus in Zoological Institute of the Russian Academy of Sci species on star! i ngs and re-examined most material of ences, Saint Petersburg, Russia; IRSNB - uncatalogued this genus used by previous experts. In the present paper, collection of Prof. A. FAIN, Institut royal des Sciences we give an emended diagnosis of the genus Stumotro naturelles de Belgique, Brussels, Belgium; MRAC - gus and a systematic review of all species associated Musee royal de !'Afrique centrale, Tervuren, Belgium; with starlings in Africa and Europe, including descrip NU - Nebraska University, Lincoln, USA; TRT - Col tions of new species. We also include in the review those lection of E . TROUESSART in Museum national d' Histoire Stumotrogus species that have not yet been formally naturelle, Paris, France; UGA - University of Georgia, recorded from these continents but whose presence is Athens, USA; ZISP- Zoological Institute of the Russian quite expected, because their common hosts do occur in Academy of Sciences, Saint Petersburg, Russia. Where these areas. the collection number consists of two sections, the first section refers to the collection number of the mite speci men and its depository (if another depository is not spe Material and methods cifically mentioned); the second is a collection number of the respective host specimen. Location data are given The material used in the present study was received from in their original form, as on slide specimens; where the three main sources: Musee royal de !'Afrique central (Ter country name or important location in the original label vuren, Belgium), Institut royal des Sciences naturelles de was missed, it is added to collection data text in rectan Belgique (Brussels, Belgium), and Zoological Institute gular brackets. Systematics and scientific names of birds of the Russian Academy of Sciences (St. Petersburg, follow DI CK INSON (2003). Russia). Some type specimens and comparative materi als were loaned from the Museum national d'Histoire naturelle (Pari s, France) and A. Mickiewicz University Taxonomy (Poznan, Poland). An emended diagnosis of the genus and descriptions of new species are given in the standard form used for Pteronyssidae Oudemans, 1941 pteronyssid taxa (FACCIN I & ATY EO, J 98J; M I RONOV, Sturnotrogus MIRONOV, 1989 1992, 2001). The general morphological terms, and leg and idiosoma l chaetotaxy follow GAUD & ATYEO TYPE SP EC IES: Pteronyssus truncatus TROU ES SART, 1885. II Review of Sturnotrogus species from starlings in Afri ca and Europe 57

DI AGNOS IS: Both sexes: Epimerites I free. Unpaired seta SP EC IES INCLUDED: The genu s includes 13 species vi present. Prodorsal shield as narrow trapezoid, extend arranged in two species groups based on the structure of ing beyond row of scapular setae, not encompassing setae the hysteronotal shields in males and females (see Table cl; scapular setae se in lateral incisions of the shield l and di agnoses of species groups) . (Figs. lA, 3A). Setae c2 hair-like, short, length less than half the distance between setae se. Setae c3 lanceolate, R EMARK: The genu s Sturn.otrogus is most similar to the spiculiform, or seti fo rm (Figs. 2C, SD, 6D). Setae dp2 of genus Scutulan.yssus, and differs from it by the following palpae hair-like, simple. Setae ba of tarsi I-II hair-like, combination features: in males, the opisthosomal lobes short. Genua! solenidion al longer than tarsal sol enidion are truncated apically, terminal cleft is not expressed, wl on legs I; genu a! solenidion a2 absent. Tarsus III with and the opisthosomal membrane is a narrow entire band 5 setae. Ventral membrane of tarsus I about 1/2- 2/3 of on the posteri or margin of opisthosoma between bases of the length of the segment. Surface of coxal fields I, II setae ps2; in females, the lateral opisthosomal sclerites covered with striated tegument. are always clearl y outlined and have a lanceolate fo rm, Male: Opisthosomal lobes short and wide, truncate the opisthosomal sclerite is a large unpaired plate, or (if or bluntly rounded. Posteri or margin of opisthosomal this sclerite is not split from the hysteronotal shield) the lobes between bases of setae ps2 with narrow enti re opi sthosomal area of hysteronotal shield is not dissected opisthosomal membrane; free margin of the membrane medi all y. In the males of Scutulanyssus, opi sthosomal straight or shallowly concave or with small median lobes have a cl early convex posterior margin, terminal incision, but membrane is never interrupted (Figs. 2A, cleft is small but clearly expressed, opisthosomal mem B, 4C, 12C). Terminal cleft not expressed. Supranal brane is present only on terminal margin of lobes and concavity short, open posteriorly. Setae c2 in antero interrupted in terminal cleft; in the females, opisthoso median angle of humeral shields or mesal to it (Figs. mal scl erites are paired or (if this fragment is not split lA, 4A). Setae psi anterior to or at the level of setae from the hysteronotal shield) the opisthosomal part of h3. Hysteronotal shi eld without scl erotized ridges; ante hysteronotal shield has a deep median incision, lateral rior angles acute or blunt-angul ar. Coxal fi elds III open. sclerites are of irregul ar fo rm, separated or fused with Transventral scl erite and epiandrium present. Epi opisthosomal sclerites. andrium usuall y horseshoe-shaped and fused with posterior margin of transventral sclerite; branches of epiandrium encompass genital apparatus. Anal discs Key to African and European species of Stumotrogus circula r, large, with finely striated membrane. Adana! shield represented by pair of poorl y scleroti zed plates Females or absent. Adana! membranes present. Setae h3 long seti fo rm comparable in length to setae h2. Tarsus III l. Opisthosomal shield not separated from the main body elongated, slightly curved, commonly with cl aw-li ke of hysteronotal shield (s ubtruncatus group) (Fig. apex. Setae r longer than tarsus III, seta w spiculiform, 13D) ...... S. subtruncatus (TROUESSA RT, 1885 ) seta s narrowly lanceolate or spiculifo rm, seta f thick Hysteronotal shield separated by transverse band ened (Figs. 2G, H). Tarsus IV without dorsobasal spine, of soft stri ated tegument into central sclerite and longer than tibia IV; setae d, e modified into barrel opi sthosomal sclerite (truncatus group) (Fig. 3A) 2 shaped suckers (Fig. 2I). 2. Central sclerite noticeably elongated, 2-2.5 times Female : Idiosoma moderately elongated, without longer than wide at anterior margin (Fig. 3A, 13B , opi sthosomal lobes. Arrangement of hysteronotal shield C) ...... 3 fragments in one of two patterns: (I) large approximately Central sclerite 1.5-1.8 times longer than wide at rectangul ar central sclerite, pair of lateral opisthoso anterior margin (Figs. SA-C) ...... 5 mal sclerites and large unpaired opisthosomal sclerite 3. Central sclerite approximately 2.2-2.5 times longer (truncatus group) (Fig. 3A); (2) main body of hyster than wide (Fi gs. 3A, 13C); length of idiosoma longer, onotal shield entire, i.e. not split into central sclerite 415-450...... 4. and opi sthosomal sclerite, lateral sclerites completely Central sclerite approxi mately 2 times longer than or incompletely split from hysteronotal shield (subtrun wide (Fig. 13B); length of idiosoma shorter, 360- catus group) (Fig. 13D). Lateral sclerites always Ian 405 ...... S. superbus sp.n. ceolate in form, in most known species (except for S. 4. Posteri or margin of opisthosoma between setae h3 subtruncatus) clearl y separated from the main body of with scleroti zed median extension; setae c3 lanceo- hysteronotal shield . Hysteronotal gland openings gl on late, wide, 22-30 x 6. 5-7 (Figs. 3A, B) ...... anterior end of lateral sclerites. Epigynium bow-shaped ...... S. truncatus (TROUESSART, 1885) or semicircul ar, with pair of short lateral extensions, tips Posteri or margin of opi sthosoma without median of epigynium not touchi ng epimerites lila; scl erotized sclerotized extension; setae c3 na rrowly lanceolate, fo lds of oviporus short (Fig. 3B). External copulatory with thin acute apex, 35-45 x 3-3.3 (Figs. 13C) . . . tube absent. S. creatophorae MIRONOY & Koru, 2001 5. A nterior angles of central sclerite not extended lat- HosTs: Passerifo rmes: Stu rn idae and Rhipiduridae. erally (Fig. 8B) ...... S. lamprotornis sp. n. I I 58 Sergey Y. MIRONOV, Georges WAUTHY

Central sclerite with anterior angles extended later Seta s of tarsus III situated on inner margin of the ally (Figs. 8A, C, D) ...... 6 segment (Figs. 7A, E, 11A, E) ...... 5 6. Anterior angles of central sclerite acute. Setae el Anterior angles of hysteronotal shield acute but not approximately at the transverse level of openings gl elongated; anterior margin of this shield straight or (Fig. 80) ...... S. pastoris (MIRONOV, 1987) slightly concave. Seta s of tarsus III situated on ven Anterior angles bluntly rounded, often with irregu tral surface of the segment, noticeably distant from lar margin. Setae el distinctly posterior to the level inner margin (Figs. SA, E, F) ...... 6 of openings gl (Figs. 8A, l3A) ...... 7 5. Lateral margins of hysteronotal shield with small 7. Setae c3 distinctly lanceolate, wide, 25-30 x 6-6.5. irregular incisions at level of setae cp; setae ps3 Openings gl closer to level of setae d2 than to level commonly on small sclerites (Figs. 7A, C) ...... of setae el (Fig. 8A) ...... S. acridotheres sp. n...... S. pastoris (MIRONOV, 1987) Setae c3 spiculiform or narrowly lanceolate with very Lateral margins of hysteronotal shield without inci thin apex, 38-55 x 3-3.5. Openings gl closer to level of sions; setae ps3 commonly on soft tegument (Figs. setae el than to level of setae d2 (Fig. 8C) ...... 8 11A, C) ...... 8. Lateral sclerites narrow, 3-3.5 times longer than ...... S. creatophorae M IRONOV & KOPIJ, 2000 wide. Anterior margin of opisthosomal sclerite 6. Length of idiosoma 340-370; width of opisthosoma straight (Fig. 8C). ldiosoma longer, 425-475 ...... (distance between setae ps2 ) 100-110 ...... 7 ...... S. onychognathi sp. n. Length of idiosoma 275-320; width of opisthosoma Lateral sclerites noticeably thickened in anterior 65-75 ...... 8 part because of strongly convex inner margin, 2-2.5 7. Posterior margin of opisthosomal membrane straight. times longer than greatest wide. Anterior margin of Opisthosoma with almost rectangular posterolateral opisthosomal sclerite concave (Fig. l3A). ldiosoma angles. Setae c3 shorter, 45-50 in length (Figs. SA shorter, 355-375 ...... S. poeopterae sp n. F) ...... S. lamprotornis sp. n. Posterior margin of opisthosomal membrane shal Males lowly concave. Opisthosoma with rounded postero lateral angles. Setae c3 longer, 55-60 in length (Figs. I. Idiosoma wide and short, approximately 1.2-1.3 6A-F) ...... S. onychognathi sp. n. times longer than wide. Anterior angles of hyster 8. Tips of epiandrium commonly extend to level of onotal shield blunt-angular. Gland furrows clearly setae 4a (Fig. 9B). Adana! membranes without scle- demark antero-lateral fragments from the main body rotized rib (Fig. 9C) ...... S. poeopterae sp. n. of hysteronotal shield. Opisthosomal membrane with Tips of epiandrium not extending to level of setae 4a small median incision (Figs. 12A-C). (subtruncatus (Fig. lOB). Adana! membranes with sclerotized rib group) . . . . . S. subtruncatus (TROUESSART, 1885) (Fig. IOC) ...... S. superbus sp. n. Idiosoma relatively elongated, 1.4-2 times longer than wide. Anterior angles of hysteronotal shield Group truncatus acute. No pieces separated or demarked from antero lateral parts of hysteronotal shield. Opisthosomal DIAGNOSIS: In male, hysteronotal shield with acute ante membrane with straight or slightly concave margin rior angles (Figs. lA, 7A, 11 A); in female, central and (Figs. IA, 4A, SA). (truncatus group) ...... 2 opisthosomal sclerites are clearly separated, lateral scler 2. Setae c3 distinctly lanceolate and relatively wide ites lanceolate in form and always clearly separated from and short, 25-30 x 5-6.5 (Figs. 2C, 40) ...... 3 the main body of hysteronotal shield (Figs. 3A, 8A-D, Setae c3 spiculiform or narrowly lanceolate, 35-55 x 13A-C). 2.5-3.5 (Figs. 50, 60) ...... 4 Included species: Stumotrogus acridotheres sp. n., S. 3. Bases of setae ps3 not surrounded by small trian creatophorae MIRONOV & KoPIJ, 2000, S. lamprotornis gular sclerites. Setae cJ approximately equidistant sp. n., S. minimus MIRONOV, 1989, S. onychognathi sp. from prodorsal and hysteronotal shields or close to n., S. pastoris (MIRONOV, 1987), S. poeopterae sp. 11., S. posterior margin of prodorsal shield. Seta s of tar superbus sp. 11. , and S. truncatus (ThoUESSART, 1885), sus III clearly extends beyond apex of the segment. ldiosoma noticeably elongated, 1.6-1.85 longer than l. Sturnotrogus truncatus (TROUESSART, 1885) (Figs. lA, wide (Figs. lA, B, 2B, G, H) ...... B, 2A-L, 3A, B) ...... S. truncatus (TROUESSART, 1885) Bases of setae ps3 on small triangular sclerites. Pteronyssus truncatus ThouESSART, 1885: 49; CAN Setae cl closer to anterior margin of hysteronotal ESTRIN!, 1886: 272; CANESTRIN I & KRAMER, 1899: 80; shield than to prodorsal shield. Seta s of tarsus III VITZTHUM, 1929: 97; RADFORD, 1958: 157. scarcely extending beyond apex of the segment. Idi Pteronyssus quadratus var. trunca.tus: BERLESE, 1886: osoma 1.4-1.5 times longer than wide (Figs. 4A-E) fasc. 26, No. 3; BONNET & TI MON-DAVID, 1934: 258 ...... S. acridotheres sp. n. Pteronyssoides truncatus: HULL, 1934: 202, RADFORD, 4. Anterior angles of hysteronotal shield strongly elon 1953: 205; GAUD & TILL, 1961: 276; FACCINI & ATYEO, gated; anterior margin of this shield deeply concave. 1981: 54. I I Review of Sturnotrogus species from starlings in Africa and Europe 59

Pteronyssoides (Scutulanyssus) truncatus: MIRONOV, of Europe and Asia (ThouESSART, 1SS5 ; CANEST RINI 19S5: 202, Fig. 16g, 17g, 1Sg. & KRAMER, 1S99 ; VITZTHUM , 1929; HULL, 1934; Sturnotrogus truncatus: MIRONOV, 19S9: 120, Fig. 12, DUBININ, 1950, DUBININ & SOSNINA, 1952; SHUMILO, 1-3. 1960; GAUD & TILL, 1961 ; CERNY, 1964; FACCINI & ATYEO, 19S1, RAZOWSKI , 1997; MIRONOV, 19S5, 1989, MATERIAL EXAMINED: 2 male, 2 female syntypes (TRT 1996, 1997), and also from individuals introduced to 41Fl) ex Sturnus vulgaris LINNAEUS, 175S, [France], Canada (H.C. PROCTOR, personal communication). Maine et Loire, no other data; 3 males, 3 females GAUD and PETITOT (1948) reported S. truncatus from (GAUD), same host, [France], Rennes, XII.l969, coli. the common myna, Acridotheres tristis, in Indochina, unknown; 3 males, 4 females (GAUD), same host, The but after the examination of materials collected from Netherlands, X.l953, coli. unknown; 3 males (AMU that host in Saint Helene Island, GAUD (1977) referred 032, slides A, B), same host, Poland, Slonsk, 1.VII.19S3, mites associated with A. tristis to S. subtru.ncatus. Re coli. J. Dabert; 2 males, 2 females (AMU, no number), examination of materials from Saint Helene Island and same host, Poland, near Poznan, Jeziory, 22.V.2002, coli. Indochina shows that these authors actually dealt with a unknown; 15 males, 1S females, (ZISP 1466), same host, quite different Sturnotrogus species, which we describe Russia, Kaliningrad Province, Rybachy, 16.X.l979, S.V. below as a new species, S. acridotheres. GAUD & TILL Mironov; 4 males, 4 females (ZISP 1465), same host, (1961) also reported S. truncatus from Ptilostomus afer Russia, Leningrad Province, Gumbaritsy, 2.X.19S1, S.V. (LINNAEUS, 1766) (Corvidae) in Africa; however, this Mironov; 10 males, S females (ZISP 1467), same host, material actually also represented another new species Russia, [Astrakhan Province], Volga delta, Damchik, (seeS. lamprotornis). We suggest that Stu.1nu.s vulgaris 13.III.l940, V. Dubinin; 4 males, 3 females (ZISP 1463), and its close relative, the spotless starling St. u.nicolor (a same host, Russia, Novosibirsk Province, Chany Lakes, newly recorded host), are original and principal hosts of no date, B. Bykhovsky; 10 males, 12 females (ZISP S. truncatus. 1S30), same host, Russia, [Primorye, Khankqjskii Dis trict], Gornokhankajsk, 25.V.l954, L. Mulyarskaya; 1 2. Sturnotrogus acridotheres sp. n. (Figs. 4A-D, SA) male, 2 females (ZISP 1459), same host, Kazakhstan, North Kazakhstan Province, Konyukhovskii District, Pteronyssoides truncatus: sensu GAUD & PETITOT, 1948: 15.IX.1951, Bezukladnikov; 12 males, S females (ZISP 339, Fig. I, 2 (misidentification). 1S32), same host, Tadzhikistan, Tigrovaya Balka reserve, Pteronyssoides subtruncatu.s: sensu GAUD, 1977: 265 22.XII.1940, E. Sosnina; 3 males, 2 females (MRAC 1SO (misidentification). 397), same host, Morocco, Rabat, XII.l966, R. Dupuy; 6 males, 5 females (MRAC 1SO 401 and 1SO 402), same TYPE MAT ERIAL: Male holotype, 2 male and 7 female host, Morocco, Christian, 1.1949, coli. unknown; 5 males, paratypes (MRAC 129 OSO, 10 slides) ex Acridotheres · 6 females (MRAC 1SO 39S and 1SO 399) ex St. unicolor tristis (LINNAEUS, 1766): ST. HELENE ISLAND: TEMMINCK , 1S20, Morocco, Hazagan, VII.l950, coli. XII.l965, Belgian Zoological Mission, P. BASILEVSKY, P. unknown; 3 males, 3 females (MRAC 1SO 400), same BENOIT, and N. LEL EU P. host, Morocco, Fes, 1.1949, coil. unknown. OTHER MATERIAL EXAMINED: 5 males, 1 female (GAUD), DIAGNOSIS: Both sexes: setae c3 distinctly lanceolate ex A. tristis, [Cambodia], Siem Reap Province, VII.l946, 22-30 x 6.5-7 (Fig. 2C). Male: Hysteronotal shield with J. Gaud; Smales, 3 females (GAUD), same host, [Vietnam, out incision on lateral margin, anterior angles acute but Lam Dong Province, Di Linh District], Djiring, V.1946, not elongated, opisthosomal lobes truncate, margin of J. Gaud. opisthosomal membrane straight, without incisions, setae ps3 on soft tegument, adana! membranes with sclerotized DIAGNOSIS : Both sexes: setae c3 clearly lanceolate, short, rib (Figs. 1A, B, 2A, B), idiosoma 330-3SO long, 1.7- 25-30 x 6-6.5 (Fig. 4D). Male: Hysteronotal shield with l.S5 longer than wide, width of opisthosoma (distance out incision on lateral margins, anterior angles acute but between setae ps2) S0-90. Female: Hysteronotal shield not extended, posterior margin of opisthosomal lobes split into central and opisthosomal sclerites, central scle truncate, opisthosomal membrane with small blunt-angu rite 2.2-2.5 longer than wide and its anterior angles not lar incision, setae ps3 on small triangular sclerites, rib of extended (Fig. 3A), anterior margin of opisthosomal sc le adana! membranes poorly developed (Fig. 4C), idiosoma rite slightly concave, lateral sclerites narrow (3 or more 340-350 long, 1.4-1.5 times longer than wide, width of times longer than wide), openings gl much closer to the opisthosoma 100-110. Female: Hysteronotal shield split level of setae el than to level of setae d2, idiosoma 435- into central and opisthosomal sclerites, central sclerite 4SO long, 1.9-2.1 longer than wide. with anterior angles sli ghtly extended and rounded, 1.1- 1.2 times longer than wide, anterior margin of opisthoso REMARK: Sturnotrogus truncatus, the type species of mal sclerite straight or slightly sinuous, lateral sclerites the genus, was originally described from the common narrow, openings gl approximately equidistant from lev starling St. vulgaris in France (ThouESSART, 1SS5) ; now els of setae e2 and el (Fig. SA) , idiosoma 415-450 long, it is widely known from thi s host in numerous locations 1.6-1.7 longer than wide. 60 Sergey V. MIRONOV, Georges WAUTHY I I

DESCRIPTION : Male holotype: ldiosoma length x width 3. Sturnotrogus lamprotornis sp. n. (Figs. SA-F, 8B) 350 x 245 (340-345 X 215-225 in 2 paratypes). Length of hysterosoma 230. Prodorsal shield: posterior margin TYPE MATERIAL: Male holotype and 3 female paratypes slightly convex, length along median line 94, width at (MRAC 180 380) ex Lamprotornis splendidus (VIEIL posterior margin 112, setae se separated by 89 (Fig. 4A). LOT, 1822): CAMEROON, 1959, collector unknown; 8 Setae c2 thin hair-like, situated mesal to anteromedian male and 8 female paratypes (MRAC 180 381 , 180 382), angle of humeral shields; setae c31anceolate, 30 in length, same data. 6.5 in width. Hysteronotal shield: anterior margin almost straight, length along midline 220, width 154. Distance OTHER MATERIAL EXAMINED: 7 males, 5 females (MRAC between prodorsal and hysteronotal shields along midline 180 374, 180 375) ex L. chalcurus, NORDMANN , 1835, about 30. Opisthosomal lobes truncate, terminal mem Cameroon, Adomaoua, IY.l951 , collector unknown; 1 brane with shallow median incision. Length of supranal male, 1 female (MRAC 180388) ex L. chalybaeus (HEMP concavity 32. Dorsal measurements: c2:d2 80, d2:e2 92, RICH & EHRENBERG, 1828), Kenya, Njoro, VII.1964, d2:gl24, gl:el 16-17, e2:h3 35, e2:e2 76, h2:h2 81, h3:h3 collector unknown; 1 male, 3 females (MRAC 180392) 46. Transventral sclerite 11 in length along median line. ex L. nitens (LINNAEUS, 1766), South Africa, Cape Prov Tips of epiandrium extending to base of genital appara ince, East London, X.l966, collector unknown;2 males, 1 tus; genital arch 28 x 16. Adana] shields as pair of poorly female (MRAC 147367, 3 slides) ex L. splendidus, Cam sclerotized plates. Setae ps3 on small triangular sclerites. eroon, Yaounde 22.Y.1959, collector unknown; 4 males, Adana] membranes with longitudinal sclerotized rib. 2 females (MRAC 148138, 6 slides), same host, Cam Diameter of anal discs 22. Ventral measurements: ps2: eroon, no other data; 7 males (IRSNB) ex L. purpurop ps2 102, ps3:ps3 38, ps3:h3 40. Tarsus III 60 long, setae tents RuPPELL, 1845, Rwanda, Akanyaru, 18.11.1955, w 33 long; setae s 20 long, not extending beyond apex of A. Fain; 2 males (IRSNB), same data except 9.1.1955; 2 the segment. Tarsus IV subequal in length to tibia IV. females (IRSNB), same data except 15.X.I955; 2 males, Female paratype: Idiosoma length x width 440 x 260 (in 2 females (NU 8267, deposited in ZISP) ex L. iris (Ous other 7 series 415-450 x 245-275). Length of hysterosoma TALET, 1879), West Africa, Sierra Leone, 12.IX.l961 , Fr. 305. Prodorsal shield as in the male, 103 x 127, setae se Allison; 1 male (UGA 3338 AMNH 210 863, deposited separated by 102. Setae c2 short hair-like, situated on in ZISP) ex L. corruscus corruscus NoRDMANN, 1835, striated tegument. Setae c3 lanceolate 27 x 6.5. Arrange Kenya, Saukuri, 10.III.l923, K. Caldwell; 1 female (UGA ment of hysteronotal shield fragments: central sclerite, 3348 AMNH 668 723, deposited in ZISP) ex L. chlorop pair of lateral opisthosomal sclerites, and opisthosomal tents elisabeth (STRESEMANN, 1924), Mozambique, N. sclerite. Central sclerite with slightly extending ante Mozambique, Lalana, Suli river, 28.V.l925, H.C. Muller; rior angles, length along midline 172, width at anterior 9 males, 3 females (MRAC 180 383 - 180 386, 4 sides) margin 152. Lateral opisthosomal sclerites with rounded ex Ptilostomus afer (LINNAEUS, 1766) (Corvidae), Sen posterior end. Opisthosomal sclerite with almost straight egal, Vl.l955, collector unknown. anterior margin, posterior margin convex, poorly sclero tized, greatest length 86, greatest width 134. Posterior DIAGNOSIS: Both sexes: setae c3 spiculiform, 40-50 margin of opisthosoma between setae h3 straight, with x 3-3.5 (Fig. 50). Male: Hysteronotal shield without out terminal extension. Setae d2 on lateral margins of incision on lateral margins, anterior angles acute but central sclerite, setae el closer to the level of openings gl not elongated, opisthosomal lobes truncate, margin of than to level of setae d2. Setae psi approximately at the opisthosomal membrane straight, without incision, setae level of setae h2. Dorsal measurements: c2:d2 124, d2: ps3 on soft tegument, rib of adana] membranes poorly e2 97, d2:gl34, gl:el 24, e2:h3 60, h2:h2 118, h3:h3 83. developed (Figs. SA-C), idiosoma 340-370 long, 1.3- Epigynium bow-like, 32 x 108. 1.4 times longer than wide, width of opisthosoma 105- 110. Female: Hysteronotal shield split into central and REMARK : The new species described above was previ opisthosomal sclerites, central sclerite 1.4 times longer ously recorded by GAUD & PETITOT (1948) as Ptero than wide, its anterior angles rectangular, not extended, nyssoides truncatus from the common myna, Acridoth anterior margin of opisthosomal sclerite concave, open eres tristis, in Cambodia and Vietnam; further, GAUD ings gland setae el approximately at the same transverse (1977) detected this species from the same host on level , lateral sclerites with rounded posterior end (Fig. Saint Helene Island and re-identified all material from 8B), idiosoma 425-445 long, about 1.55-1.6 longer than the common myna as Pteronyssoides subtruncatus. Re wide. examination of the materials of Gaud from Saint Helene Island and Indochina has shown that A. tristis actually DES CR IPTlON: Male holotype: Idiosoma length x width bears a new species, which is quite different from trueS. 358 x 260 (340-370 x 245-280 in 8 paratypes). Length subtruncatus. of hysterosoma 230. Prodorsal shield: posterior margin straight, length along median line 96, width at posterior ETYMOLOGY: The specific epithet derives from the margin 110, setae se separated by 84. Setae c2 thin hair generic name of the type host and is a noun in a genitive like, situated mesal to anteromedian angle of humeral case. shields; setae c3 spiculiform, 48 in length, 3.5 in width. Review of Sturnotrogus species from starlings in Africa ami Europe 61

Hysteronotal shield: anterior margin slightly concave, OTHER MATERIAL EXAM.INED: 3 males and 3 females greatest length 202, width 146. Distance between pro (NU 3568) ex 0. mario, South Africa: Transvaal, Potch dorsal and hysteronotal shields along midline about 60. efstroom, 1-6.IY.l953, F. Zumpt (material in ZISP). Opisthosomal lobes truncate, terminal membrane with straight margin without any incision. Length of supranal DIAGNOSIS: Both sexes: setae c3 setiform, 50-58 x 2.5- concavity 35. Dorsal measurements: c2:d2 71 , d2:e2 92, 3.5 (Fig. 60). Male: Hysteronotal shield without incision d2:gl 35 , gl:el 16-18, e2:h3 46, e2:e2 80, h2:h2 87, h3: on lateral margins, anterior angles acute, slightly elon h3 56. Transventral sclerite 13 in length along median gated, opisthosomal lobes with slightly convex posterior line. Tips of epiandrium extending to base of genital margin, opisthosomalmembrane with shallowly concave apparatus; genital arch 27 x 14. Adana! shields as pair margin, setae ps3 on soft tegument, adana! membranes of poorly sclerotized plates. Setae ps3 on soft tegument. without rib, idiosoma 350-365 long, 1.3-1.4 times longer Adana! membranes with poorly developed sclerotized than wide, width of opisthosoma (distance between setae rib. Diameter of anal discs 23-24. Ventral measurements: ps2) 105-112. Female: Hysteronotal shield split into ps2:ps2 106, ps3:ps3 42, ps3:h3 53. Tarsus III 60 long; central and opisthosomal sclerites, central sclerite 1.2- setae w 24 long; setae s, 21 long, situated on ventral side 1.3 times longer than wide, its anterior angles slightly of the segment and not extending beyond its apex. Tarsus extended and rounded, anterior margin of opisthosomal IV subequal in length to tibia IV. sclerite straight, openings gl closer to the level of setae el Female paratype: Idiosoma length x width 442 x 285 (in than to the level of setae d2, lateral sclerites with rounded other 10 paratypes 425-440 x 265-275). Length of hys posterior end, idiosoma 425-472 long, about 1.55-1.6 terosoma 310. Prodorsal shield as in the male, 112 x 132, longer than wide. setae se separated by 104. Setae c2 short hair-like, situ ated on striated tegument. Setae c3 spiculiform, slightly DESCRIPTION: Male holotype: ldiosoma length x width thickened in medial part, 42 x 3.5. Arrangement of hys 360 x 265 (in 3 specimens of other series 350-365 x teronotal shield fragments: central sclerite, pair of lateral 260-275). Length of hysterosoma 230. Prodorsal shield: opisthosomal sclerites, and opisthosomal sclerite. Cen posterior margin almost straight, length along median tral sclerite roughly rectangular, anterior margin slightly line 96, width at posterior margin 103, setae se separated narrower than posterior, length along midline 160, width by 83. Setae c2 thin hair-like, situated slightly mesal to at anterior margin 114. Lateral opisthosomal sclerites anteromedian angle of humeral shields; setae c3 setiform, with rounded posterior end. Opisthosomal sclerite with 58 in length, 2.5 in width. Hysteronotal shield: anterior slightly concave anterior margin, posterior margin con margin slightly concave, sinuous; greatest length 208, vex, with pair of poorly sclerotized incisions, greatest width 154. Distance between prodorsal and hysterono length 88, greatest width 130 (Fig. 8B). Posterior margin tal shields along midline about 55. Opisthosomal lobes of opisthosoma between setae h3 slightly convex. Setae slightly convex, terminal membrane with straight margin d2 off lateral margins of central sclerite, setae el almost without any incision. Length of supranal concavity 32. at the level of openings gl. Setae psi approximately at the Dorsal measurements: c2:d2 81, d2: e2 98, d2:gl27, gl:el level of setae h2. Dorsal measurements: c2 :d2 104, d2:e2 14-15, e2:h3 38, e2:e2 80, h2:h2 94, h3:h3 58. Transven 121 , d2:gl60, gl:el 9-10, e2 :h3 56, h2:h2 120, h3:h3 90. tral sclerite 15 in length along median line. Tips of epi Epigynium bow-like, 28 x 102. andrium extending to base of genital apparatus; genital arch 28 x 16. Adana! shields as pair of poorly sclerotized REMARKS: 1. In females collected from Lamprotornis plates. Setae ps3 on soft tegument. Adana! membranes chloropterus and L. iris, the anterior margin of opistho without sci erotized rib. Diameter of anal discs 24. Ventral somal sclerite is almost straight rather than in the type measurements: ps2:ps2 110, ps3:ps3 50, ps3:h3 50. Tar series and in specimens from remaining host species. sus III 58 long; setae w 26 long; setae s 21 long, situated 2. The material of S. lamprotornis from the piapiac, on ventral side of the segment and not extending beyond Ptilostomus c~fe r (LINNAEUS, 1766) (Corvidae), in our its apex. Tarsus IV subequal in length to tibia IV. opinion represents a case of accidental contamination. Female paratype: Idiosoma length x width 472 x 290 (in Based on this material, GAUD and TILL (1961) reported 3 specimens of other series 425-430 x 260-275). Length this species as Pteronyssoides truncatus from Pt. afer in of hysterosoma 330. Prodorsal shield as in the male, 112 Senegal. x 132 , setae se separated by 104. Setae c2 short hair-like, situated on striated tegument. Setae c3 setiform, slightly ETYMOLOGY: The specific epithet derives from the generic thickened, 54 x 3.5. Arrangement of hysteronotal shield name of the type host and is a noun in apposition. fragments: central sclerite, pair of lateral opisthosomal sclerites, and opisthosomal sclerite. Central sclerite 4. Sturnotrogus onychognathi sp. n. (Figs. 6A-F, 8C) roughly rectangular, with slightly extending anterior angles, anterior margin straight, length along midline TYPE MAT ERIAL: Male holotype (MRAC 180 395) ex Ony 165, width at anterior margin 140. Lateral opisthoso chognathus mario (LlN NAEUS, 1766), SOUTH AFRICA: mal sclerites with rounded posterior end. Opisthosomal Transvaal, IY.I972, collector unknown; female paratype sclerite with straight anterior margin , posterior margin (MRAC 180 394), same data. with pair of poorly sclerotized areas, greatest length 100, 62 Sergey V. MIRONOV, Georges WAUTHY greatest width 144. Posterior margin of opisthosoma 6. Sturnotrogus poeopterae sp. n. (Figs. 9A-F, 13A) between setae h3 straight. Setae d2 off lateral margins of central sclerite, setae el closer to the level of openings Pteronyssoides subtruncatus: sensu GAUD & MoucHET, gl than to the level of setae e2. Setae psl approximately 1959: 513 (misidentification). at the level of setae h3. Dorsal measurements: c2 :d2 122, d2:e2 112, d2:gl 56, gl:el 14-16, e2:h3 64, h2:h2 150, TYPE MATERIAL: Male holotype, 1 male and 3 female h3:h3 100. Epigynium bow-like, 30 x 115. paratypes (MRAC 180 377) ex Poeoptera lugubris BoNAPARTE, 1854: CAMEROON: Yaounde, 11.1956, ETYMOLOGY: The specific epithet derives from the coli. unknown; 2 male, 2 female paratypes (MRAC 180 generic name of the type host and is a noun in a genitive 376), same data. case. DIAGNOSIS: Both sexes: setae c3 setiform, 40-45 x 2.5- Sturnotrogus pastoris (MIRONOV, 1987) (Figs. 7A-E, 8D) 3.5 (Fig. 9D). Male: Hysteronotal shield without incision on lateral margins, anterior angles acute, slightly elon Pteronyssoides (Scutulanyssus) pastoris MIRONOv, 1987: gated, opisthosomal lobes with slightly convex posterior 529, Fig. 1. margin, opisthosomal membrane with shallow incision, Sturnotrogus pastoris: MIRONOv, 1989: 120, Fig. 12, 4- idiosoma 275-285 long, 1.45-1.6 longer than wide, width 6. of opisthosoma 65-70, setae ps3 on soft tegument, adana! membranes without rib. Female: Hysteronotal shield split MATERIAL EXAMINED: Male holotype, 2 male, 3 female into central and opisthosomal sclerites, central sclerite paratypes (ZISP 926) ex Sturnus roseus (LINNAEUS, 1.4-1.5 times longer than wide, its anterior angles slightly 1758): Kazakhstan: Djambul Province, Chokpak, extended and rounded, anterior margin of opisthosomal 4.IX.l984, S.V. Mironov; 1 male, 1 female (ZISP 2612), sclerite concave, openings gl closer to the level of setae same host, Tajikistan, Gissar Ridge, 6.VI.l951, L. Mul el than to that of setae d2 , lateral sclerites with acute yarskaya; 1 male, 1 female (ZISP 2611), same host, Turk posterior end, idiosoma 355-375 long, about 1.65-1.75 menistan, near Ashkhabad, ll.V.1972, D. Kurbanova; 7 longer than wide. male, 4 female (ZISP 2613) ex St. cineraceus Temminck, 1835, Russia, Primorye, Sudzukhin Reserve,l6.IV.1945, DESCRIPTION: Male holotype: ldiosoma length x width 285 coli. unknown. x 195 (275-285 x 170-185 in 3 paratypes). Length of hys terosoma 178. Prodorsal shield: posterior margin straight, DIAGNOSIS: Both sexes: setae c3 narrowly lanceolate, length along median line 76, width at posterior margin 78, 30-38 x 3-4 (Fig. 7D). Male: Hysteronotal shield com setae se separated by 63. Setae c2 thin hair-like, situated monly with small irregular incision on lateral margins, mesal to anteromedian angle of humeral shields; setae c3 anterior angles acute and elongated, opisthosomal lobes thick setiform, 42 in length, 3.5 in width. Hysteronotal with slightly convex posterior margin, opisthosomal shield: anterior margin slightly concave, greatest length membrane with slightly concave posterior margin bases 160, width 115. Distance between prodorsal and hyster of setae ps3 on small sclerites, adana! membranes with onotal shields along midline about 40. Opisthosomallobes out rib, idiosoma 335-360 long, 1.5-1.6 longer than wide, with slightly convex posterior margin, terminal membrane width of opisthosoma (distance between setae ps2) 80- margin with shallow incision. Length of supranal concav 90. Female: Hysteronotal shield split into central and ity 27. Dorsal measurements: c2:d2 50, d2:e2 76, d2: opisthosomal sclerites, central sclerite about 1.2-1.3 gl 22, gl:el 14-15, e2:h3 35, e2:e2 58, h2:h2 56, h3:h3 times longer than wide, its anterior angles extended and 35. Transventral sclerite 9 in length along midline. Tips acute (angles may be split from the main part of central of epiandrium not extending to base of genital apparatus sclerite), lateral margins with small incision at level of (left and right tips of epiandrium are unequal in length in trochanters Ill; anterior margin of opisthosomal sclerite the holotype and some paratypes); genital arch 20 x 10. straight or slightly sinuous, openings gl and setae el Adana! shields as pair of poorly sclerotized plates. Setae almost at the same transverse level, lateral sclerites with ps3 on soft tegument. Adana! membranes without sclero acute posterior tip, idiosoma 430-445 long, about 1.7-1.8 tized rib. Diameter of anal discs 15-16. Ventral measure longer than wide. ments: ps2:ps2 65, ps3:ps3 23, ps3: h3 40. Tarsus III 46 long; setae w 23 long; setae s 16 long, situated on ventral REMA RK: This species is known only from two star side of the segment and slightly extended beyond its apex. ling species, the rosy starling St. roseus (type host) and Tarsus IV subequal in length to tibia IV. white-cheeked starling St. cineraceus TEMMINCK, 1835, Female paratype: ldiosoma length x width 365 x 215 (in in Central Asia and Promorye region (Russia), respec 4 other paratypes 355-375 x 200-220). Length of hystero tively (MIRO NOv, 1987) . Although this species has not soma 248. Prodorsal shield as in the male, 78 x 96, setae been formally recorded from the territory of Africa and se separated by 76. Setae c2 short hair-like, situated on E urope, it is possible to expect it on the rosy starling in striated tegument. Setae c3 setiform, slightly thickened the southern and eastern parts of Europe and in northern in medial part, 40 x 2.5. Arrangement of hysteronotal Africa. shield fragments: central sclerite, pair of lateral opistho- Review of Sturnotrogus species from starlings in Africa and Europe 63 somal sclerites, and opisthosomal sclerite. Central scler c2:d2 SO, d2:e2 76, d2:gl22, gl: e114-15, e2:h3 35, e2:e2 ite roughly rectangular, with slightly extending anterior 58, h2:h2 56, h3:h3 35. Transventral sclerite 11 in length angles, anterior margin straight, lateral margins with along midline. Tips of epiandrium extending to midlevel small incision at level of trochanters III, length along of genital apparatus; genital arch 24 x 12. Adana! shields midline 140, width at anterior margin 95. Lateral opistho as pair of poorly sclerotized plates. Setae ps3 on soft teg somal sclerites with acute posterior end. Opisthosomal ument. Adana! membranes with oblique sclerotized rib. sclerite with slightly concave anterior margin, posterior Diameter of anal discs 18. Ventral measurements: ps2: margin concave and poorly sclerotized, greatest length ps2 72, ps3:ps3 36, ps3:h3 41. Tarsus III 48 long; setae 75, greatest width 115. Posterior margin of opisthosoma w 26 long; setae s 20 long, situated on ventral side of the between setae h3 concave. Setae d2 on lateral margins of segment and slightly extended beyond its apex. Tarsus IV centrals sclerite, setae el closer to the level of openings subequal in length to tibia IV. gl, than to the level of setae e2. Setae psi approximately Female paratype: Idiosoma length x width 375 x 210 at the level of setae h2. Dorsal measurements: c2:d2 78, (in 4 other paratypes 360-405 x 200-245). Length of d2:e2 97, d2 :gl38, gl:el 19-21, e2:h3 51 , h2:h2 lOS , h3: hysterosoma 260. Prodorsal shield as in the male, 94 x h3 80. Epigynium bow-like, 26 x 86. 108, setae se separated by 84. Setae c2 short hair-like, situated on striated tegument. Setae c3 thin spiculiform, REMARK: This species was recorded by GAUD & Mou 40 x 2.5. Arrangement of hysteronotal shield fragments: CHET (1959) under the name Pteronyssoides subtrunca central sclerite, pair of lateral opisthosomal sclerites, tus from Poeoptera lugubris in Cameroon. and opisthosomal sclerite. Central sclerite rectangular, elongated; anterior margin straight, anterior angles not ETYMOLOGY: The specific epithet derives from the generic extended, lateral margins slightly convex, length along name of the type host and is a noun in apposition. midline 144, width at anterior margin 70. Lateral opistho somal sclerites with acute posterior end. Opisthosomal 7. Sturnotrogus superbus sp.n. (Fig. lOA-F, 13B) sclerite with concave anterior margin, posterior margin with pair of poorly sclerotized areas, greatest length TYPE MATERIAL: Male holotype, 2 male, 2 female para 67, greatest width 114. Posterior margin of opisthosoma types (MRAC 180 390) ex Lamprotornis superbus RuP between setae h3 slightly sinuous. Setae d2 off lateral PELL, 1845: KENYA: Njoro, VI.1964, coli. unknown; 3 margins of central sclerite, setae el approximately equi male, 3 female paratypes (MRAC 180 389), same data. distant from levels of opening gland setae e2. Setae psi approximately at the level of setae h2. Dorsal measure DIAGNOSIS: Both sexes: setae c3 setiform or spiculiform, ments: c2:d2 94, d2:e2 103, d2:gl SO , gl:el 25-28, e2:h3 40-45 x 2.5-3.5 (Fig. lOD). Male: Hysteronotal shield 43, h2:h2 100, h3:h3 70. Epigynium bow-like, 29 x 87. without incision on lateral margins, anterior angles acute, slightly elongated, opisthosomal lobes with slightly con ETYMOLOGY: The specific epithet derives from the spe vex posterior margin, opisthosomal membrane shallowly cies name of the type host and is a noun in apposition. concave, setae ps3 on soft tegument, adana] membranes with poorly developed rib, idiosoma 285-320 long, 1.35- 8. Sturnotrogus creatophorae MIRONOV & KoPJ.J , 2000 1.55 longer than wide, width of opisthosoma 70-75. (Figs. 11A-E, 13C) Female: Hysteronotal shield split into central and opistho somal sclerites, central sclerite about 2 times longer than Sturnotrogus creatophorae MIRONOV & KoPIJ, 2000: wide, its anterior angles slightly extended and rounded, 328, Fig. Sc , d, 6a, c. anterior margin of opisthosomal sclerite concave, open ings gl closer to the level of setae el than to that of setae MATERIAL EXAMINED: 1 male, 1 female paratypes (ZISP d2, lateral sclerites with acute posterior end, idiosoma 4202) ex Creatophora cinerea (MEUSCHEN, 1787), South 360-405 long, about 1.65-1.8 longer than wide. Africa, Free State, Pellisier, 20.V.1989, R.A. EARLE.

DESCRIPTION: Male holotype: Idiosoma length x width DIAGNOSIS: Male: setae c3 narrowly Ianceolate with fili 315 x 215 (285-320 x 185-235 in 5 paratypes). Length form apex 45-53 x 2.5-3 (Fig. liD), hysteronotal shield of hysterosoma 198. Prodorsal shield: posterior margin without incision on lateral margins, anterior angles acute straight, length along median line 86, width at posterior and greatly elongated, opisthosomal lobes with almost margin 82, setae se separated by 67. Setae c2 thin hair straight posterior margin, opisthosomal membrane with like, situated mesal to anteromedian angle of humeral slightly concave posterior margin, setae ps3 on soft tegu shields; setae c3 thick setiform, 45 in length, 2.5 in ment, adana] membrane without rib, idiosoma 315-340 width. Hysteronotal shield : anterior margin slightly con long, 1.5-1.6 longer than wide, width of opisthosoma 82- cave, greatest length 178 , width 122. Distance between 90. Female: setae c3 narrowly lanceolate 35-40 x 3-3.5, prodorsal and hysteronotal shields along midline about hysteronotal shield split into central and opisthosomal 60. Opisthosomal lobes with almost straight posterior sclerites, central sclerite about 2.2-2.3 times longer than margin, terminal membrane margin slightly concave. wide, its anterior angles rectangular, anterior margin Length of supranal concavity 28. Dorsal measurements: of opisthosomal sclerite strongly concave, openings gl 64 Sergey V. MIRONOV, Georges WAUTHY I I

much closer to the level of setae el than to that of setae REMARK : This species Was originally described from the d2, lateral sclerites with rounded posterior end, idiosoma hill myna, GraCLtla religiosa, in Indonesia (ThouESSA RT, 410-425 long, about 1.8-1.9 longer than wide. 1885). Subsequently, it was reported in Africa (GAUD & TI LL, 1961 ; GAUD, 1977) from Acridotheres tristis REM ARK: This species is known only from the type and Poeoptera lugubris. Our re-examination of these host, Creatophora cinerea in South Africa (MIRONOY & materials from Africa has shown that each of these two KOPJJ, 2000). hosts actually bears another species, which we desc ribed above as new species, S. acridotheres and S. poeopterae, Group subtruncatus respectively. These misidentifications may be appar ently explained by the following reasons: Sturn.otrogus DIAGNOSIS: In males, hysteronotal shield with blunt subtruncatus has never been redescribed or even fi gured angular or rounded anterior angles (Fig. 12A); in females: based on the materials from G. religiosa, and Sturnotro central and opisthosomal sclerites are not separated, lat ghs specimens collected from other hosts were not com eral sclerites separated or not separated from the main pared with type specimens or adequate materials from body of hysteronotal shield (Fig. 13D). the type host. INCLUDED SPECIES: Sturnotrogus albicolfis MIRONOV, In Europe, we have formally detected S. subtruncatus 1993, S. ostraconotus (GAUD, 1968), S. subtruncatus in the sample collected from its type host, G. religiosa, (TROUESSART, 1885), S. stroggylus (GAUD, 1968). Only imported to Belgium. Nevertheless, we believe that this S. subtruncatus occurs in the geographic region of our species may be also expected on the hill myna in wild present interest, three other species are associated with condition in some locations of Africa, where it has been hosts restricted to the Indo-Malayan reg ion. recently introduced, particularly in Saint Helene Island.

9. Sturnotrogus subtruncatus (TROUESSART, 1885) (Figs. 12A-E, 13D) Discussion

Pteronyssus truncatus var. subtruncatus TROU ESSA RT, In total, the genus Sturnotrogus now includes 13 species 1885: 49; CAN ESTRIN! & KRAM ER, 1899: 81; RADFORD, (Table I) recorded from 23 host species; most of known 1958 : 157. Sturnotrogus species (11) are associated with starlings Pteronyssoides truncatus var. subtruncatus: RADFORD, (Sturnidae) from various areas of the Old World , mainly 1953: 205. from Eurasia and Africa, and two species are known Pteronyssoides subtruncatus: GAUD & MouCHET, 1959: from fa n-tailed flycatchers of the genu s Rhipidura VIG 513 (misidentification); GAUD & TILL , 1961: 276 (misi ORS & HORSFI ELD, 1827 (Rhipiduridae) in Southeast Asia dentification); GAU D, 1977: 265 (misidentification); FAc and Indonesia. In host specificity, most species are mon CINI & ATYEO, 1981: 54. oxenous or oligoxenous, i.e. associated with one or with Sturnotrogus subtruncatus: MIRON OY, 1989 : 118. a few species of one genus; only Sturnotrogus lampro tornis is recorded from a large number (eight) species, all MATERIAL EXA MIN ED : 6 male, 3 female syntypes (TRT of the genus Lamprotornis TEMMINCK, 1820. In relation 37H14) ex Grantla religiosa LINNA EUS, 1758, [Indone to this, it is quite interesting to point out th at representa sia], "Java, Sumatra, Malacca", no other data; 3 males, 2 tives of such a morphologically homogenous genus are females (JRSNB), same host, Belgium , Antwerp, Avice di spersed on avian hosts of two phylogenetically di stant ntra, XII.l965, A. FAIN. families. Sturnidae and Rhipiduridae belong to different parvorders of oscines, Passerida and Corvida, respec DIAGN OS IS: Both sexes: setae c3 thick setiform 28-38 x ti ve ly (SIBLEY et al. , 1988 ; ERICSON et al., 2002; ERICSO N 2.5-3 (Fig. 12D). Male: Hysteronotal shield with convex & JOHANSON, 2003; BARK ER et al., 2004). Although the anterior margin , angles blunt-angul ar, small antero-lat pattern of host associations of Sturnotrogus species in eral pieces of this shield demarked , lateral margin with the context of respective avian families is obviously the out inci sions, opisthosoma lobes truncate with rounded result of co-speciation with hosts, it is quite unlikely that posterolateral angles, margin of opisthosomal membrane di stribution on such distantly related taxa as starlings and straight, with small median incisions, setae ps3 on soft fan -tailed flycatchers could be the result of co-speciation tegument, adana! membranes without rib, idiosoma 310- with a common ancestral host. Analysis of host associa 330 long, 1.2-1.3 longer than wide, width of opisthosoma tions within species groups, recogni zed in the course of 92-98. Female: Hysteronotal shield entire (main body of our taxonomic study, allows us to hypothesize the origi thi s shield not split into central and opi sthosomal scler nal and secondary hosts of Sturnotrogus. ites, laterals sclerites outlined but not split off), anterior The truncatus group apparently represent a deri ved angles of hysteronotal shield rounded, posterior quarter of grouping within the genu s, because females are character thi s shi eld with pair of short longitudinal grooves, lateral ised by the hysteronotal shield split into fragments (Figs. margins with pair of narrow incisions anterior to setae.f2, 8A-C). In most fa milies of Analgoidea, females are char openings gl closer to level of setae e 1 than to level of setae acterized by the relatively large and entire hysteronota l d2 , idiosoma 360-380 long, 1.5-1.6 longer than wide. shi eld, whil e onl y in fe males of Pteronyss id ae does one Rev iew of Sturnotrogus species from starlings in Africa and ,Europe 65 see such reducti on and splitting of the fe male hys trono sturnids are the ori ginal hosts of the ge nus, while rhipi tal shields, which is a very ch aracteri sti c evolutionary durids are the secondary hosts, and Sturnotrogus species trend di spl ayed within many pteronyss id genera (GAUD , were probabl y transferred onto the ancestor of fa n-tailed 1990; MIRONOV, 2001 , 2003; MIRONOV & WAU THY, fl ycatchers from some sturnids that originated in the Indo 2005a, 2006b). Representati ves of the truncatus group Malaya n reg ion. Taking into account that Sturnidae cur are restricted exclusively to sturnids and are associated rently includes about 11 4 species arranged in 23 or more with the most deri ved genera of thi s fa mily: Acridoth eres genera (DI CK INSON , 2003) and all members of this family VI EIL LOT, 1816, Creatophora LESSO N, 1847, Lamprotomis, are potenti al hosts of the genu s Stu rn otrogus, it is possible Onychognathus HARTLAUB, 1849 and Sturnus LI NNAEUS, to expect a number of new Sturnotrogus species on star 1758, whi ch are di stributed mainly in Afrotropic and Pal lings in tropical areas of the Old World that would help to earctic regions (SI BLEY & AHLQU IST, 1984). Within the understand better the pattern of host parasite associations subtruncatus group, which appears to be more primitive and co-speci ation process with these hosts. because the hysteronotal shield in fe males is entire (Fig. 13d), two species are known from sturnids, and two spe cies from rhipidurids. Representatives of the subtruncatus Acknowledgements group li ve on the relatively primitive sturnid genera, Aplo nis GouLD , 1836 and Gran t/a LI NNAEUS , 1758, di stributed T he authors wish to thank Prof. Dr. A. Fain (lnstitut roya l des Sciences naturell es de Belgique, Bru ssels, Belgium), Dr. H. Andre mainly in the Indo-Malayan region, in the sa me range as (Musee roya l de !'Afrique centrale, Tervuren, Belg ium), Dr. M. Jud Rhipidura species. It is also necessary to stress that many son (Museum nationa l d' Hi stoire naturelle, Paris, France) and Dr. J. relatives of sturnids within Passerid a have pteronyssid gen Dabert (A . Mickiewicz Uni versity, Poznan, Poland) fo r loaning mate era related to Sturnotrogus (ge nera of the Pteronyssoides ria ls used in the present study, and Dr. H.C. Proctor (University of group), whereas no cl ose relatives of rhipidurids within A lberta, Edmonton, Canada) fo r critica ll y reviewing the manuscri pt. T his research was supported by a grant from the Belgian Scienti fic Corvida have mites related to Sturn.otrogus (MI RONOV & Poli cy to SYM. The present in vesti gati on was based in part on the WAUT HY, 2005a, 2006b; MI RO NOV & GALLOWAY, 2006) materi a l belong ing to the coll ecti on U FC Z lN No 2-2.20 (Coll ecti on Based on these data, it is most reasonabl e to conclude that of Zoological Instit ute RAS, St. Petersburg, Ru ssia). 66 Sergey V. MIRONOV, Georges WAUTHY

Tabl e I -Host associations and distribution of Sturnotrogus species

Mi te species and spec ies gro up Host species Hos t fam il y Distribut ion Reference

truncatus group

S. acridorh eres *Acridorh eres rrisris Sturnid ae Cambodia. Vi etnam , St. prese nt study Helene Is.

S. crearophorae *Creatophora cinerea Sturn idae South Africa Mironov & Kopij , 2000

S. /amproromis Lamproromis chalcurus Sturnidae Cameroon present study

L. chalybaeus Sturnidae Kenya present study

L. chloroprems elisaberh Sturnidae Mozambique prese nt study

L. C0 /"1"/I SC // S CO ri"I.I SC II S Sturnidae Kenya present study

L. iris Sturn idae Sierra Leone present study

L. uireus Sturnidae South Africa prese nt study

L. purpuroprem s Sturnidae Rwanda prese nt study

'' L. spleudidus Sturnidae Cameroon present study

)Pri/osromus af"er Corvidae Senega l Gaud & Ti ll, 1961 ; present study

S. lllillillll!S *Srunrus sr umiuus Sturnidae Ru ss ia (Pri morye) Mi rono v, 1993

s. onychognarhi ''' Ouyclroguarlrus m orio Sturnidae South Africa present study

S. pas/o ris *Srurnus roseus Sturnidae Ka za kh stan , Tajikista n Mirono v, 1987

S r. ciuereus Sturnidae Ru ss ia (Primorye) Mironov, 1987

S. poeoprera e ''P oeoprera /ugubris Sturnidae Cameroon present study

S. superbus '' Lwnproromis superbus Sturnidae Kenya present stud y

S. rmucatus *Sturnus vulgaris Sturnidae Europe, Morocco, Ru ss ia Trouessa rt. 1885; (Europea n part, West Faccini & Atyeo. Siberia, Primorye) , 198 1; Mironov, Ka zak hstan , Tadzhi ki stan , 1987; present Ca nada (A lbert a) study

St. uuicolor Stu rnidae Morocco present stud y

subtnmcatus group

S. a /bico!/is *Rhipidura albico!lis Rh ipiduridae Vietnam Mironov, 1993

S. ostracouotus ''Rhipidum reuuelliaua Rhipiduridae Renn ell ls. Gaud, 1968

s. stroggy!us *Aplonis insularis Sturnidae Rennell Is. Gaud. 1968

S. subtruncaru s *Graet!la religiosa Sturnid ae Ind ones ia, Belgium ''"'' Trouessa rt , 1885; present study

(*)Type host;(**) mite record ed from the imported host specim en; (t) probably accidental contaminati on. Review of Sturnotrogus species from starlings in Africa and, purope 67

A 8

Figs. I A-B - Sturnotrogus truncal us, ma le. A. Dorsal view. B. Ventral view. 68 Sergey Y. MIRONOV, Georges WAUTHY ' '

A h2 h2 c

E F wa dpl ra dp2

\I \I \) ~

s d

Figs. 2A-L - Stumotrogus truncal us, deta il s. A-I. Ma le. J-L Female. A. Opisthosoma, dorsal view. B. Opisthosoma, ventral view. C. Humera l shi eld, ventral view. D. Tarsus !, dorsal view. E. Tarsus 1, ventral view. F. Tarsus 11 , ventra l view. G. Tarsus III, dorsa l view. H . Tarsus Ill, ve ntral view. I. Tarsus !V, dorsal view. J. G nathosoma, ventra l view. K. Tarsus !11 , dorsal view. L. Tarsus IV, dorsal view. a m - adana! membrane, as- adana! shi eld , om - opisthosomal membrane, rb - rib of adana! membrane, sc - suprana l concavit y, vm - ve ntral membrane of tarsus I. Review of Stumotrogus species from starlings in Africa and Europe 69

Figs. 3A-B - Sturnotrogus truncatus, female. A. Dorsal view. B. Ventral view. Fragments of hysteronota l shield: cs - centra l sclerite. Is - latera l opisthosomal sclerite, os - opisthosoma l sclerite. 70 Sergey Y. MIRONOV, Georges WAUTHY

E ::::L 0 lO

E ::::L E 0 0 N

Figs. 4A-E- Stum otrogus acridotheres, male. A. ldiosoma, dorsal view. B. Hysterosoma, ventral view. C . Opisthosoma, ventral view. D. Seta cJ. E. Tarsus III , dorsa l view. Review of Sturnotrogus species from starlings in Afri c~ . and Europe 71

E E ::1. ::::t. 0 0 0 l!) N

Figs. 5A-F - Sl/(mutmgus lwnproturnis, male. A. ldiosoma, dorsal view. B. Hysterosoma, ve ntra l view. C. Opisthosoma, ventral view. D. Seta cJ. E. Tarsus I I L dorsa l view. F. Tarsus Ill. ventral view. 72 Sergey V. MIRONOV, Georges WAUTHY

E :::t 0 lO

Figs. 6A-F- Sturnotrogus onychognathi, ma le. A. ldiosoma, dorsal view. B. Hysterosoma, ve ntra l view. C. O pi sthosoma, ve n tra l view. D. Seta d . E. Tarsus lll, dorsal view. F. Tarsus Ill, ve ntra l view. I I

Review of Sturnotrogus species from starlings in Africa· and Europe 73

E ::i. 0 l()

E :::t 0 0 N

Figs. 7A-E - Sturnotrogus pas/oris. male. A. ldiosoma, dorsal view. B. Hysterosoma, ventral view. C. Opisthosoma, ventra l view. D. Seta d . E. Tarsus Ill, dorsal view. '' 74 Sergey V. MIRONOV, Georges WAUTHY

E :i 0 0 N 8

Figs. 8A-D- Sturnotrogus females, dorsal view of icliosoma. A. Stumotrogus acridotheres. B.S. /omprotornis. C. S. onychog nathi. D. Stu.rnutrugus pas/oris. Review of S-t u--rnotrog~ u s· ~s p~ec ~ies~o·~o m~st~ru·~r' m"'~·os•.• 111 ~AfII .· Ca~~~ an~~~--7:d EUl . Ope ?S

E ::::!.. 0 l()

E ::::L 0 0 N A

FigdA-F - ·· Sturnotrog . . . plsthosoma, v e ntral view. D· Seu t:d c3 po ·eo Ep · tTera arseu. sma I I 1l,e d · oAr ·sa ldl 1.vo ise owm. aF, . dTarsorsaul.s vIll1·e w' ve. B.n trHay l sv.t ieeJ.wo s oma . ventr. al view. C 0 . 76 Sergey V. MIRONOV, Georges WAUTHY I I

F E ::i.. 0 0 N

Figs. lOA-F - Stumotrogus superbus, male. A. Idiosoma, dorsal view. B. Hysterosoma, ventral view. C. Opisthosoma, ventral view. D. Seta c3. E. Tarsus Ill, dorsal view. fF Tarsus III, ventral view. rb - rib of adanalmembrane. Review of St·ul . no trogus species fro m stat. lin . gs Ill. Africa andI IEur ope 77

E :i. 0 1.!)

E :i. 0 0 N

Figs. llA-E - Stur' I o I rogus creatophorae m I . tral view. D. Seta c3 . .E Tal." s usa II e.I. A.do ldrsawl svoiemwa., dorsal view. B . Hysterosoma. ventral view· C · 0 p1.s t I10 soma, ven- 78 Sergey V. MIRONOV, Georges WAUTHY I I

Fi gs. 12A-E -Sturnotrogus subtruncotus, male. A. ldiosoma. dorsal view. B. 1-l ysterosoma, ventral view. C. O pi sthosoma, ven tral view. D. Seta c3. E. Ta rsus Ill, dorsa l view. Review of Sturnotrogus species from starlings in Africa and Europe 79

E :::i. 0 0 N

Figs. 13A-D - S!U m olrogus fe ma les. d orsal view of idiosoma. A. Slumolrugus poeoplerae. B. S. superbus. C. S. crealophorae. D. S. sub!rt.IIICOius. I I 80 Sergey Y. MIRONOV, Georges WAUTHY

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