Acarina 11 (1): 3–29 © ACARINA 2003 ON SOME PROBLEMS IN THE SYSTEMATICS OF FEATHER MITES Î ÍÅÊÎÒÎÐÛÕ ÏÐÎÁËÅÌÀÕ Â ÑÈÑÒÅÌÀÒÈÊÅ ÏÅÐÜÅÂÛÕ ÊËÅÙÅÉ S.V. Mironov Ñ.Â. Ìèðîíîâ Zoological Institute, Russian Academy of Sciences, Universitetskaya emb. 1, Saint-Petersburg, 199034 Russia. E-mail: [email protected] Çîîëîãè÷åñêèé èíñòèòóò Ðîññèéñêîé Àêàäåìèè Íàóê, Óíèâåðñèòåòñêàÿ íàá. 1, Ñàíêò-Ïåòåðáóðã, 199034 Ðîññèÿ. E-mail: [email protected] Key words: feather mites, taxonomic system, phylogeny, idiosomal chaetotaxy, homology of chaetome Êëþ÷åâûå ñëîâà: ïåðüåâûå êëåùè, òàêñîíîìè÷åñêàÿ ñèñòåìà, ôèëîãåíèÿ, õåòîòàêñèÿ èäèîñîìû, ãîìîëîãèÿ õåòîìà ABSTRACT Two problems in the systematics of feather ñëåäîâàòåëÿìè ýòîé ãðóïïû, òîãäà êàê ñèñòåìà mites are discussed. The first problem concerns the ÎÊîííîðà [OConnor, 1982a] áîëåå àäåêâàòíî taxonomic frames of feather mites, which are an îòðàæàåò ôèëîãåíåòè÷åñêèå îòíîøåíèÿ ìåæäó ecological group within parasitic astigmatid mites, âûñøèìè òàêñîíàìè ïåðüåâûõ êëåùåé. Âòîðàÿ arrangement of recently recognized feather mite ïðîáëåìà çàêëþ÷àåòñÿ â óñòàíîâëåíèè ãîìîëî- families into superfamilies, and relationships among ãèè âåíòðàëüíûõ ãèñòåðîñîìàëüíûõ ùåòèíîê their higher taxa. Two general concepts in regard to (êîêñàëüíûõ è ãåíèòàëüíûõ) êàê ìåæäó ðàçëè÷- the taxonomic frame and taxonomic system of íûìè òàêñîíîìè÷åñêèìè ãðóïïàìè ïåðüåâûõ feather mites recently exist. The concept of Gaud êëåùåé. Ïðåäëîæåíî íåñêîëüêî ãèïîòåç è and Atyeo [1996] is most detailed and widely used îáñóæäàåòñÿ èõ ïðèìåíèìîñòü â îòíîøåíèè in practice by taxonomists and other investigators òåõ èëè èíûõ òàêñîíîìè÷åñêèõ ãðóïïèðîâîê of the group in question, while the concept of ïåðüåâûõ êëåùåé. O’Connor [1982a] most adequately reflects the INTRODUCTION phylogenetic relationships between higher taxa of feather mites. The second important problem in the Feather mites are a vast group of astigmatid study of feather mite concerns the homology of the mites that are permanent parasites or symbiotes of ventral hysterosomal setae (coxogenital setae) birds, and live on their plumage or skin [Dubinin, among different taxa of feather mites, and, respec- 1951, Peterson, 1975; Gaud, Atyeo, 1982a, 1996; tively, the assignation of chaetotactic nomencla- O’Connor 1982a, 1982b; Mironov, 1999; Dabert, ture to them. Several hypotheses of homology of Mironov, 1999; Proctor, Owen, 2000; Proctor the ventral hysterosomal setae in different taxo- 2003]. This group currently includes over 2400 nomic groups of feather mites are proposed and species (in 450 genera, 33–36 families, 3 super- discussed. families) distributed throughout the World and occurring on almost all recent orders of Aves, ÐÅÇÞÌÅ with the exception of penguins (Sphenisciformes). Ðàáîòà ïîñâÿùåíà îáñóæäåíèþ äâóõ ïðî- As it is very popular to stress in general papers on áëåì â ñèñòåìàòèêå ïåðüåâûõ êëåùåé. Ïåðâàÿ feather mites, this number of feather species is ñâÿçàíà ñ îïðåäåëåíèåì òàêñîíîìè÷åñêèõ ðàìîê estimated by experts to be less that 20% of possi- äëÿ ýòîé ýêîëîãè÷åñêîé ãðóïïèðîâêè àñòèãìàòè- bly existent species. In general appearance, feath- ÷åñêèõ êëåùåé, îáúåäèíåíèåì âûäåëÿåìûõ â er mites are rather typical astigmatid mites with íàñòîÿùåå âðåìÿ ñåìåéñòâ â íàäñåìåéñòâà è body size ranging from 300 to700 mm, and with ðîäñòâåííûìè ñâÿçÿìè âûñøèõ òàêñîíîâ ïåðüå- relatively well sclerotized tegument in compari- âûõ êëåùåé.  íàñòîÿùåå âðåìÿ ñóùåñòâóåò äâå son to free-living Astigmata. They are highly îñíîâíûå òàêñîíîìè÷åñêèå ñèñòåìû ïåðüåâûõ specialized and occupy discrete microhabitats on êëåùåé. Ñèñòåìà Ãî è Ýòèî [Gaud, Atyeo, 1996] the bodies of birds. For instance, these mites íàèáîëåå äåòàëèçèðîâàíà è øèðîêî èñïîëüçó- inhabit feathers only of certain type, and only åòñÿ íà ïðàêòèêå ñèñòåìàòèêàìè è äðóãèìè èñ- certain microareas within a fan of such feathers. S.V. Mironov Experts still disagree whether feather mites should feather mites in the near future, to understand be referred to as parasites or commensals. Some recent conceptions in this field and avoid errors. feather mites that inhabit the skin or quills and feed DISCUSSION respectively on the skin cells and internal corneous parts of feathers are obvious parasites according to The fist problem may be expressed as “What all definitions of parasitism. Most feather mites are feather mites in systematic and phylogenetic feed on a secretion produced by the oil gland, terms?” Of particular interest are the relationships which birds disperse over their plumage, and usu- of their highest taxa to each other and to other ally do not cause any visible damage to the host; astigmatids. It is worthy to point out that the scien- therefore, investigators were lead to believe that tific definition of “feather mites” (Federmilben — most feather mites are commensals. However, there Gm., Acariens plumicoles — Fr.) always directly are enough records that prove that under certain depended on the dominating taxonomic concept in conditions such normally commensal species cause regard to this group. To illustrate this and also show depluming itch and other diseases of birds [Shaw, the historical development of the taxonomic sys- 1966, Alwar et al., 1958; Alwar, 1970, Oba et al., tem of feather mites, it is expedient to trace the 1978, Rosen et al., 1988]. Therefore, the feather history of the systematics of this group. Gaud and mites that cause no visible damage to their hosts Atyeo [1996] proposed the first attempt to recog- may be considered to be potential parasites. nize and determine periods in feather mite studies, Since the discovery of this group of mites, the but the present paper proposes slightly different studies of this group have been mainly focused on subdivisions and other titles for the periods. the investigation of their biodiversity and improv- Primary period. The first feather mite was ing their taxonomic system. Ecological, physiolog- mentioned in the “Systema Nature” [Linnaeus, ical and anatomy investigations are still extremely 1758]; it was “Acarus passerinus Linnaeus, 1758” rare, apparently because they require rather com- from the chaffinch Fringilla coelebs Linnaeus. plicated techniques for rearing in experiments or Interestingly enough as a curious, the second feath- observations and collecting in natural conditions er mites species known up that moment, “Pediculus [For references on ecological studies see: Gaede, pari Linnaeus, 1758” from the big tit Paris major Knülle, 1987; Kim, 1989; Gaud, Atyeo, 1996; Linnaeus, was placed in that issue among the lice Mironov, 2000]. Phylogenetic studies of this group Anoplura, i.e. among insects, not mites. Sixty years are also in the stage of naissance. Dubinin [1951] later, Nitzsch [1818] established the first feather proposed the first phylogenetic scheme of feather mite genus Analges Nitzsh 1818, which united all mites in the 1950’s, but only a few phylogenetic feather mites know in that time. The period since reconstructions of several feather mite taxa have the first discovery of feather mites until the been carried out since [Mironov, 1991a, 1991b; late1860’s may be referred to as the primary period Dabert, Mironov, 1999; Mironov, Dabert, 1999; of investigation or the period of primary accumula- Dabert, Ehrnsberger, 1995, 1998; Dabert et al., tion of biodiversity data. 2001; Ehrnsberger et al, 2001]. Classical period. Robin [1868a, 1868b] for In spite of the extensive study of feather mites the first time used the term Avicolar Sarcoptidae, begun in the mid-20th century and the significant or Sarcoptides avicoles, a suprageneric taxon unit- progress in the elaboration of their taxonomic sys- ing 5 feather mite genera. Several years later, tem achieved by the end of this century, a number Robin and Megnin [1877] substituted Robin’s term of problems concerning the systematics of this for the term Sarcoptides plumicoles, and proposed group still remain. The present discussion does not the first taxonomic system with detailed morpho- intend to list and discuss all essential problems in logical characteristics of 5 genera recognized. Fur- the systematics of feather mites, but focuses on two ther, Trouessart and Megnin [1884a–1884c] treat- of them, which seem to be most important at this ing Sarcoptides plumicoles as a subfamily within point. It also does not propose any final resolutions, the ectoparasitic mite family Sarcoptidae Murray, because solving these problems requires complex 1877 proposed an arrangement of feather mite and long-term investigations. The main goal is to genera into three sections. From 1884 to 1916, point out these problems, propose possible hypoth- Trouessart with coauthors and in monographic esis and stimulate any studies to solve them. In papers contributed significantly to the systematics addition, this discussion would be helpful to the of feather mites and to knowledge of their biodiver- researchers, who begins or will begin to investigate sity. The final version of his taxonomic system for 4 On some problems in the systematics of feather mites this group [Trouessart, 1916] included four sec- feather mites. Besides, all suprageneric taxa rec- tions in the subfamily: ognized by Dubinin [1953] were provided with Analgeseae — 12 genera detailed morphological characteristics. The res- Pterolicheae — 35 genera toration of feather mites as a single taxon and the Proctophyllodeae — 9 genera publication of a three-volume taxonomic mono- Epidermopteae — 6 genera graph in a series “Fauna of the USSR” [Dubinin, Period of dispersion. In the taxonomic sys- 1951, 1953,
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