BULLETIN DE L'!NSTITUT ROYAL DES SCIENCESNATURELLES DE BELGIQUE ENTO!v!QLOGIE, 75: 155-214, 2005 BULLETIN VAN HET KON INK.LIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN ENTOMOLOGIE 75: 155-214, 2005 A review of the feather mite genus Pteronyssoides HULL, 1931 (Astigtnata: Pteronyssidae) from African and European passerines (Aves: Passeriformes) with analysis of tnite phylogeny and host associations by Sergey V. MIRONOV & Georges WAUTHY Abstract representatives of the feather mite morphotype adapted to inhabit vanes of large feathers, the flight feathers and A systematic review of the feather mite genus Pteronyssoides R OBIN, the large upper covert feathers of wings (MtRONOV, 1877 is g iven, including a n improved diagnosis of the genus and ). subgenera, description of 14 new species from African passerines, 1999 cladistic analysis of the genus and a brief discussion of associations Among s ix pteronyssid genera restricted to passerines with passeriform hosts. New species are described as follows: Pter­ (FACCINI & ATYEO, 1981; MtRONOV, 2001 ), the genus onyssoides emberizae sp.n. from Emberizajlaviventris (Emberizidae), Pteronyssoides HULL, 1931 (Astigmata: Pteronyssidae) P. eupasseris sp. n. from Passer griseus (Ploceidae), P. euplecti sp. n. from Euplectes hordaceus (Pioceidae), P. faini sp. n. from Lonclwra is a relatively species-rich taxon that includes 14 species bicolor poensis (Estrildidae), P. f oudiae sp. n. from Foudia madagas­ associated with various families of higher passerines. cariensis (Pioceidae), P. microscutatus from Euplectes axillaris Representatives of the genus are medium- and small­ (Pioceidae), P. ovoscutatus sp. n. from Cyanomitra verticalis (Nectar­ iniidae), P. oxylabis from Oxylabes madagascariensis (Timaliidae), sized pteronyssids (260-400 f.tm in length), most com­ P. plocei sp. n; from Ploceus superciliaris (Pioceidae), P. pytiliae sp. n. monly located on the venh·al surfaces of large covert from Pytilia melba belli (Estrildidae), P. que/eae sp. n. from Quelea feathers. The most noticeable morphological peculiarity quelea (Pioceidae), P. serini sp. n. from Serinus sulphurallls (Fringil­ lidae), P. triangularis sp. n. from Pants caeruleus (Paridae), and of this genus among other pteronyssids is a great varia­ P. viduinus sp. n. from Vidua macroura (Viduidae). bility of the hysteronotal sclerite complex in females. Due Maximum parsimony analysis based on 45 morphological characters to this feature, identification of species, in conh·ast to genus into two sub­ confirmed current taxonomic s ubdivision of the most feather mite taxa, is much easier for the females than genera, Pteronyssoides s. str. and Holonyssoides MIRONOV, 1993, and revealed three species groups w ithin the nominal subgenus (nectari­ for the males. niae, ovoscutatus, and parinus). Two contrasting morphological The genus Pteronyssoides was originally established tendencies are displayed by the s ubgenera. The main tendency in based on their free epimerites I, a feature that is relatively 1-lo/onyssoides is s trengthening of dorsal shields in both sexes, while three different lineages of Pteronyssoides s. str. show a trend to reduced rare in Pteronyssidae where the most common state is the hysteronotal shield in females. fusion of epimerites (HULL, 1931; GAUD & MOUCHET, Based on the phylogenetic relationships between mite species and 1959, GAUD & TILL, 1961). In the generic revision of their cun·ently known distribution among passerines, and taking into consideration phylogenetic relationships between higher passerine pteronyssids (F ACC INI & ATY EO, 1981 ), its diagnosis was taxa, it is hypothesized that the genus Pteronyssoides originated on enlarged and specified and one species was removed the a ncestors the parvorder Passerida. Owing to subsequent cospecia­ to the genus Metapteronyssus GAUD, 1981; however, tion, its representatives are widely dispersed on two major passeridan lineages (Passeroidea and Sylvoidea); due to host-switchi ng events in Pteronyssoides continued to encompass mites with very the course of evolution, these mites also colonized tits (Paridae) within diverse appearances. Further, two species groups, Passerida and two phylogenetically distant lineages (Dicruridae and obscurus and truncatus, were arranged in separate genera, to the core Corvoidea (parvorder Corvida). Paradisaeidae) belonging Scutulanyssus MI RONOV, 1985 and Sturnotrogus Key words: Pteronyssidae, Pteronyssoides, systematics, phylogeny, MIRONOV, 1989, respectively (MIRONOV, 1985, 1989). host associations, cospeciation. Finally, based on general structure of the dorsal shields, two subgenera, Pteronyssoides s. str. and Holonyssoides MIRONOV, 1993, were established within the genus Introduction (MIRONOV, 1993). Representatives of the genus Pteronyssoides are The feather mite fami ly Pteronyssidae (Astigmata: An­ known only from higher passerines (Oscines) of the Old algoidea) currently includes about 150 species in 22 World: most species were described from Africa (GAUD, '· genera (F ACC INI & ATY EO, 198 1; GAUD & ATY EO, 1996; 1952, 1957; GAUD & MOUCH ET, 1959; GAUD & TILL, MIRONOV, 1989, 2000, 200 I, 2003). As permanent 1961; MIRONOV & KOPIJ , 2000; MI RONOV, 2001), and a ectoparasites of birds, pteronyssids are mainly associated few species are known from the Palaearctic (MtRONOV, with Passeriformes and Piciformes, and a few species are 1985, 1989) and South-East Asia (SUGIMOTO, 1941; GAU D known from Coraciiformes. These mites are typical & PETITOT, 1948; MIRONOV, 1993). Neve11heless, species '' 156 Sergey V. MlRONOV & Georges WAUTHY diversity of this genus remains quite poorly explored, (iii) hysterosoma is measured from the level of sejugal because many avian species of the vast host-group Pas­ furrow to bases of setae h3. seriformes have never been examined. (iv) hysteronotal shield length in males is the greatest In the course of our study of feather mites associated length from the anterior margin to bases of setae h3; with African passerines we have found 14 new species width is measured at anterior margin. among materials obtained from various sources, and have (v) distance between prodorsal and humeral shield and re-examined all Pteronyssoides species formerly de­ the length of transventral sclerite in males are mea­ scribed from this area. In the present paper, we give an sured along midline. improved diagnosis of the genus and subgenera, a key and systematic review of all known species, descriptions of As f ACCIN I & ATYEO (1981) and MIRONOV (1 989) pro­ new species from African passerines, and a phylogenetic vided exhaustive synonymies for all fom1erly known analysis of this genus based on morphological characters. Pteronyssoides species we do not give synonymies in We also briefly discuss known host associations of the the present review. Specimen depositories and reference genus and provide a preliminary hypothesis of its evolu­ accession numbers are given using the following abbre­ tion on passerines. viations: AMNH- American Museum ofNatural Histmy, New York, USA; AMU - A. Mickiewicz University, Poznan, Poland; BMOC or MZUM- Museum.of Zool­ Material and methods ogy, University of Michigan, Ann Arbour, USA; IRSNB - uncatalogued collection of Prof. A. FAIN, lnstitut royal Specimens des Sciences naturelles de Belgique, Brussels, Belgium; NMB -National Museum of Bloemfontein, Free State, The material used in the present study was received from South Africa; NU- Nebraska University, Lincoln, USA; four main sources: Musee royal de I' Afrique central SAIMR - South African Institute of Medical Research, (Tervuren, Belgium), Institut royal des Sciences natur­ Johannesburg, South Africa; TRT - Collection of E. elles de Belgique (Bntssels, Belgium), University of TROUESSART in Museum national d'Histoire naturelle, Georgia (Athens, USA), and Zoological Institute of the Paris, France, UGA - University of Georgia, Athens, Russian Academy of Sciences (St. Petersburg, Russia). USA; USNM - National Museum of Natural Histmy, Some type specimens and important comparative materi­ Washington DC, USA; YSU- Youngstown State Uni­ als were loaned from the Museum national d'Histoire versity, Youngstown, Ohio, USA; ZISP - Zoological naturelle (Paris, France) and A. Mickiewicz University Institute of the Russian Academy of Sciences, Saint (Poznan, Poland). Petersburg, Russia. Where the collection number consists An emended diagnosis of the genus and descriptions of of two sections, the first section refers to the collection new species are given in the standard formats used for number of the mite specimens and its depositmy, if pteronyssid taxa (FACCINI & ATYEO, 1981; MIRONOV, another depositoty is not specifically pointed out; the 1992, 200 I). The general morphological terms, and leg second is a collection number of the respective host and idiosomal chaetotaxy follow GAU D & ATY EO ( 1996). specimen. Location data are given in an original form, Regarding the terms used for hysteronotal shield frag­ as in slide specimens. Systematics and scientific names of ments in females, we generally follow the scheme pro­ birds follow DICKINSON (2003), and passerine phylogeny posed by MIRONOV ( 1992) that was elaborated for the used in the discussion of host associations follows recent genus Pteroherpus GAUD, 1981; however, because the conceptions based on molecular studies (ERICSON et al., pair of sclerites in the posterior part of the opisthosoma 2002; ERICSON & JOHANSON, 2003; BARKER et al., 2004; in most Pteronyssoides species is entire and extends to