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87Mise au pointParasite, 2007, 14, 87-100

HISTORICAL REVIEW OF THE GENUS DERMANYSSUS DUGÈS, 1834(ACARI: MESOSTIGMATA: DERMANYSSIDAE)

ROY L.* & CHAUVE C.M.*

Summary:

A synthetic review of the historical systematics of DermanyssusDugès, 1834 (Acari: Mesostigmata: Dermanyssidae) is provided.The classification at the specific level in this early genus has notreally been clarified during more than a century despite itseconomic impact, and the history of the genus is complex andincludes various stages. Moreover, Dermanyssus currently includes23 species, whereas the last review took only 18 species intoaccount. Changes in the species status and position in the genusDermanyssus from 1834 until today are presented. The evolutionof the generic definition is explored and compared with othergenera of the group. How the discrimination between the differentspecies evolved in the genus is also examined. Some difficulties inthe specific definitions are discussed. A current diagnosis of thegenus Dermanyssus is given. A table of the species included inthis genus since its first description along with their respectivecurrent positions, a list of the currently included species inDermanyssus with their hosts, and a world map presenting theirgeographic distribution are provided.

Résumé : REVUE HISTORIQUE DU GENRE DERMANYSSUS (ACARI :MESOSTIGMATA : DERMANYSSIDAE)

La systématique historique de Dermanyssus Dugès, 1834 (Acari :Mesostigmata : Dermanyssidae) est révisée de manièresynthétique. La classification au niveau spécifique de ce vieuxgenre n’a pas été véritablement clarifiée durant plus d’un sièclemalgré son impact économique, si bien que son histoire estquelque peu complexe et présente diverses étapes. En outre, legenre Dermanyssus englobe à l’heure actuelle 23 espèces, tandisque la dernière révision prenait seulement 18 espèces en compte.Les changements de statut et de position des espèces dans legenre Dermanyssus depuis 1834 jusqu’à présent sont présentés.Le processus d’évolution de la définition du genre est exploré parcomparaison avec d’autres genres du groupe. La manière dont ladiscrimination entre les différentes espèces a évolué est aussiconsidérée. Certaines difficultés dans la définition spécifique sontdiscutées. Une description diagnostique actualisée du genreDermanyssus est fournie. Un tableau des espèces qui ont étéincluses dans le genre depuis sa création avec leur positionrespective, une liste des espèces actuelles du genre Dermanyssusavec leurs hôtes, ainsi qu’une carte de leur distribution mondialesont fournis.

KEY WORDS : Acari, Mesostigmata, Gamasida, Dermanyssus, historicalreview, systematics.

MOTS CLÉS : Acari, Mesostigmata, Gamasida, Dermanyssus, revue historique,systématique.

* Laboratoire de Parasitologie et Maladies Parasitaires, École Natio-nale Vétérinaire de Lyon, 1, avenue Bourgelat, 69280 Marcy-L’Étoile,France.Correspondence: Lise Roy.Tel.: 00 33 4 78 87 25 74 – Fax: 00 33 4 78 87 25 77.E-mail : [email protected]

group name, including only obligatory ectoparasites.Radovsky (1966, 1967) separated this group into the twofollowing families, depending on some morphologicaland biological characters: Macronyssidae Oudemans,1936 and Dermanyssidae. Really Dermanyssidae appearphylogenetically closer to free-living Laelapids than toMacronyssidae. Consequently, Moss (1968, 1978) consi-dered only the two following genera to be included inDermanyssidae: Dermanyssus and Liponyssoides. In thispaper, we follow this last classification.Along with Liponyssoides, Dermanyssus possesses somemorphological mainly located in the mouthparts adap-tations to hematophagic habits. Adult females, proto-nymphs and deutonymphs possess conspicuously thinand elongated chelicerae, with a second segment adap-ted to hematophagy. Faces of opposed second seg-ments are medially concave, so that they may form atube by joining together through which blood is with-drawn. Chelae are conspicuously reduced, even ifdigits can be seen with a scanning electron microscope(cf. Fig. 2; Radovsky, 1969; Phillis, 2006). Male cheli-

INTRODUCTION

The genus Dermanyssus Dugès, 1834 (Acari: Meso-stigmata: Dermanyssidae) includes hematopha-gous mite species which are ectoparasites of

birds. Dermanyssus is the type genus of a family whosename has represented various groups all along 19th cen-tury, with more or less internal splitting. DermanyssidaeKolenati, 1859 included first mites with diverse habits,some of them being obligatory ectoparasites and othersfree-living or facultative ectoparasites. Berlese (1892)separated the former and the last group in two diffe-rent families: Dermanyssidae and Laelapidae respecti-vely. Then numerous steps occurred, Dermanyssidaestatus alternating from subfamily-group name to family-

ROY L. & CHAUVE C.M.

88 Mise au pointParasite, 2007, 14, 87-100

Fig. 1. – Distribution map of non-gallinae species of Derma-nyssus. A: Europe. B: Asia. C: Africa. D: America. Note: a symbolcorresponds to the presence of one species in a country (admi-nistrative).

A

D. alaudae

D. trochilinis

D. Triscutatus

D. transvaalensis

D. rwandae

D. quintusD. prognephilus

D. passerinusD. nipponensis

D. longipesD. hirundinis

D. hirsutus

D. grochovskae

D. gallinoidesD. faralloni

D. chelidonis

D. carpathicus

D. brevis

D. brevirivulus

D. antillarum

D. americanus

D. wutaiensis

B C

D

cerae are broader and male chelae are enlarged, witha long spermadactyl.The poultry red mite D. gallinae (De Geer, 1778) isvery common in layer houses in Europe. The economicimpact of this parasite is quite important and may takemany forms, including the following: downgradedeggs, decreased egg production, anaemia, possibledeath from exsanguination. The poultry red mite canalso transmit diseases such as avian spirocheatosis, fowlcholera, salmonellosis, etc. Despite such economicimportance, the classification of this genus at specieslevel has been in a state of confusion for many years.About 40-50 years ago, some authors began workingprecisely on the genus Dermanyssus. According tosome of them, D. gallinae may not be the only Der-manyssus parasitising laying hens. Consequently, someother closely related species might often have beenconfused with this species. A rather low host-specifi-city and a rather wide geographic distribution of Der-manyssus species contribute to obscure the issue.Most species of this genus are not very host specific:for instance, more than 30 bird species are potentialhosts for D. gallinae (Zemskaya, 1971) and 40 bird spe-cies (belonging to eight different orders) for D. hirun-dinis (Hermann, 1804) (Moss et al., 1970; Moss, 1978;Fend’a & Schniererová, 2004; Fend’a, unpublished data).Most of the Dermanyssus species are nidicolous.Although some of them can be found frequently on thehost and can deposit their eggs on its feathers (D. gro-chovskae Zemskaya, 1961, D. quintus Vitzthum, 1921and D. americanus Ewing, 1922), most of them climbonto their host only to get a meal and then go backto their hiding-place in the host nest or roost. Moreover,

many species of Dermanyssus are distributed on morethan one continent (Fig. 1). The history of Dermanyssusis very complicated and has never been extensively exa-mined. Moreover, this early genus currently includes23 species, whereas the last review of the genus tookonly 18 species into account. For both these reasonsand in order to get a clear view of the genus beforereviewing it, it seemed necessary to examine it curso-rily from its description until the present and to checkthe current species included in it. In order to get a viewof the generic history, the text will be broken down asfollows: changes in the species status and position inthe genus Dermanyssus from 1834 until today are pre-sented first, then the generic definition and its evolu-tion are explored. Afterwards, the species definitionsand their difficulties will be examined. Finally, concer-ning the genus as is currently defined, a list of the cur-rently included species in Dermanyssus is provided.Abbreviations: Setal terminology follows Lindquist &Evans, 1965 for the dorsum and Evans, 1963 for the legs.

CHANGES IN THE STATUS ANDTHE POSITION OF SPECIES IN THE GENUSDERMANYSSUS FROM 1834 UNTIL TODAY

Dugès described the genus Dermanyssus in 1834,in which he included five new species: D. aviumDugès, 1834, D. vespertilionis Dugès, 1834,

D. convolvuli Dugès, 1834, D. oribatis Dugès, 1834, andD. hominis Dugès, 1834. The type-species D. gallinae(De Geer, 1778) was described by De Geer in the

HISTORICAL REVIEW OF THE GENUS DERMANYSSUS


Fig. 2. – Left: characte-ristically reduced chelain D. gallinae. Right: se-cond segments of cheli-cerae medially concavein D. gallinae (electronscanning microscope).

genus Acarus. In 1834, Dugès named it D. avium, andconsidered A. gallinae, although senior, synonymouswith D. avium. D. gallinae was later reinstated as thesenior synonym (Koch, 1836). D. vespertilionis has beensuppressed by the International Commission of Zoo-logical Nomenclature (ICZN) under the plenary powersfor the principle of priority, but not for homonymy(Melville & Smith, 1987). About D. oribatis and D. convo-

lvuli, Dugès only noted for each: species name, fol-lowed by a comma and the personal latin pronounnobis. Without any description and as species namessuggest host associations far from the common ones,birds, in Dermanyssus, D. oribatis and D. convolvulimight be deemed nomina dubia. In any case, theycannot be included in Dermanyssidae sensu Radovsky(1966). D. hominis seems to have been omitted by all



Species included or previouslyincluded in Dermanyssus Current position Comments

1778 * D. gallinae (De Geer, 1778) Dermanyssidae Dermanyssus* D. alaudae (Schrank, 1781) Dermanyssidae Dermanyssus In 1781, Schrank named alaudae the

seventh species he described in 1776.* D. hirundinis (Hermann, 1804) Dermanyssidae DermanyssusD. truncatus (Olfers, 1816) synonymy ➣ D. alaudaeD. hominis (Dugès, 1834) synonymy ➣ D. gallinae Bory de Saint Vincent described this

species in 1823 and more completely in1828 without any name nor systematicposition within the Acari group. Dugès placed it in genus Dermanyssus and named it in 1834.

D. avium Dugès, 1834 synonymy ➣ D. gallinaeD. vespertilionis Dugès, 1834 suppressed ICZN direction 66: suppressed under

the plenary powers for the principle of priority, but not for homonymy.

D. convolvuli Dugès, 1834 ?D. oribatis Dugès, 1834 ?D. musculi Koch, 1836 Macronyssidae Steatonyssus This species has complicated history.

Oudemans (1936) considered it junior synonym of A. musculi Shrank, which he placed in genus Steatonyssus (homo-nymy and synonymy in the same time).Evans and Till (1966:278-279) suggestedthat S. musculi Schrank could be a junior synonym of Ornithonyssus bacotiHirst, 1913.

D. arcuatus Koch, 1839 Hirstionyssidae EchinonyssusD. carnifex Koch, 1839 Hirstionyssidae Echinonyssus Tenorio (1984) treats this species as

nomen dubium.D. coriaceus Koch, 1839 synonymy ➣ D. arcuatusD. lanius Koch, 1839 synonymy ➣ D. carnifexD. noctulae Koch, 1839 synonymy ➣ D. arcuatusD. murinus Lucas, 1840 Macronyssidae SteatonyssusD. avium Wagner, 1841 synonymy ➣ D. murinusD. pipistrellae Koch, 1841 synonymy ➣ D. arcuatusD. lacertarum (Contarini, 1843) ?D. natricis Gervais, 1844 Macronyssidae OphionyssusD. musculi Johnston, 1849 Hirstionyssidae Echinonyssus A complicated history. Seems to be

conspecific to D. musculi Koch which is conspecific to A. musculi Shrank.

D. flavus Kolenati, 1857 Macronyssidae MacronyssusD. glutinosus Kolenati, 1857 synonymy ➣ M. granulosusD. granulosus Kolenati, 1857 Macronyssidae MacronyssusD. ambulans Thorell, 1872 Haemogamasidae HaemogamasusD. richiardii Canestrini & Fanzago, 1877 ?D. sylviarum Canestrini & Fanzago, 1877 Macronyssidae OrnithonyssusD. hirundinis Berlese, 1889 homonymy ➣ nomen novum:

D. chelidonis* D. longipes Berlese &Trouessart, 1889 Dermanyssidae Dermanyssus nomen dubium.

1889 * D. passerinus Berlese & Trouessart, 1889 Dermanyssidae Dermanyssus species inquirenda.

Table I (to be continued).

reviewers of Dermanyssus until today (cf. § Specieswhose nomenclatural status is not clear). To sum up,only one of the five initially included species has beenconsidered in the subsequent studies.After these descriptions, many species were created inDermanyssus by other authors. We list 57 species whichare included or have been included in the genus(Table I). 32 species are changed in status or position:ten species have been synonymized with other Der-manyssus species. Two species receive nomina novabecause they are deemed junior homonyms. One wassuppressed under the plenary powers (cf. above). 16 spe-cies are now included in some other groups: five inthe other genus of Dermanyssidae Liponyssoides, sevenin several genera of the family Macronyssidae, threein the family Hirstionyssidae, one in the family Hae-mogamasidae. Four are incertae sedis or species inqui-renda (cf. § Species whose nomenclatural status is notclear). Finally, one species is suggested here beingsynonymized (D. hominis). As a result, 23 species areincluded in Dermanyssus.Strandtmann & Wharton (1958) listed ten species inDermanyssus: D. gallinae, D. hirundinis, D. quintus,D. americanus, D. oti Ewing, 1925, D. evotomydisEwing, 1933, D. prognephilus Ewing, 1933, D. brevis

Ewing, 1936, D. scutatus Ewing, 1936, D. chelidonisOudemans, 1939.Evans & Till (1962) recognized 14 species, two ofwhich, overlooked by Strandtmann & Wharton (1958),were considered doubtful but not to be invalidated(D. passerinus Berlese & Trouessart, 1889 and D. lon-gipes Berlese & Trouessart, 1889). Two others had beendescribed after 1958 (D. triscutatus Krantz, 1959 andD. grochovskae), one was a new species (D. trans-vaalensis Evans & Till, 1962). Another one, which hadbeen considered synonymous with D. gallinae byOudemans, was restored (D. alaudae (Schrank, 1781)).From the ten listed species in Strandtmann and Whar-ton 1958, two were synonymized (D. oti with D. ame-ricanus and D. evotomydis with D. gallinae).Moss (1968) also included 14 species in Dermanyssus,but not exactly the same as Evans & Till (1962). Thedifferences were: the newly named D. hirsutus Moss& Radovsky, 1967 (= D. scutatus, praeocc.), the recentlydescribed D. gallinoides Moss, 1966 and D. faralloniNelson & Furman, 1967 (in a footnote, because thisspecies had been described as Moss’s paper went topress), and the omission of D. passerinus and D. lon-gipes. In 1978, Moss added four species: D. antillarumDusbabek & Cerny, 1971, D. trochilinis Moss, 1978,



Species included or previouslyincluded in Dermanyssus Current position Comments

1902 D. albatus Oudemans, 1902 synonymy ➣ D. arcuatusD. aegyptius Hirst, 1913 Dermanyssidae LiponyssoidesD. muris Hirst, 1913 Dermanyssidae LiponyssoidesD. sanguineus Hirst, 1914 Dermanyssidae Liponyssoides* D. quintus Vitzthum, 1921 Dermanyssidae Dermanyssus* D. americanus Ewing, 1922 Dermanyssidae DermanyssusD. oti Ewing, 1925 synonymy ➣ D. americanusD. evotomydis Ewing, 1933 synonymy ➣ D. gallinae* D. prognephilus Ewing, 1933 Dermanyssidae DermanyssusD. brasiliensis Fonseca, 1935 Dermanyssidae Liponyssoides* D. brevis Ewing, 1936 Dermanyssidae DermanyssusD. scutatus Ewing, 1936 homonymy ➣ nomen novum:

D. hirsutus* D. chelidonis Oudemans, 1939 Dermanyssidae Dermanyssus This species has been described in 1889

by Berlese as D. hirundinis. Because ofhomonymy, Oudemans renamed it in 1939.

* D. triscutatus Krantz, 1959 Dermanyssidae Dermanyssus* D. grochovskae Zemskaya, 1961 Dermanyssidae Dermanyssus* D. transvaalensis Evans & Till, 1962 Dermanyssidae DermanyssusD. intermedius Evans & Till, 1964 Dermanyssidae Liponyssoides* D. gallinoides Moss, 1966 Dermanyssidae Dermanyssus* D. faralloni Nelson & Furman, 1967 Dermanyssidae Dermanyssus* D. hirsutus Moss & Radovsky, 1967 Dermanyssidae Dermanyssus* D. antillarum Dusbabek & Cerny, 1971 Dermanyssidae Dermanyssus* D. trochilinis Moss, 1978 Dermanyssidae Dermanyssus* D. carpathicus Zeman, 1979 Dermanyssidae Dermanyssus* D. nipponensis Uchikawa & Kitaoka, 1981 Dermanyssidae Dermanyssus* D. brevirivulus Gu & Ting, 1992 Dermanyssidae Dermanyssus* D. wutaiensis Gu & Ting, 1992 Dermanyssidae Dermanyssus

1993 * D. rwandae Fain, 1993 Dermanyssidae Dermanyssus

Table I. – Species included or previously included in Dermanyssus listed in chronological order with their present position/status. Spe-cies names preceded by * are here included in Dermanyssus.

D. passerinus and D. longipes, the last two being consi-dered incertae sedis and not included in the key foridentification. Thus, in the last review of genus Der-manyssus, only 18 species were included.The five species which have been described after Moss’slast review are: D. brevirivulus Gu & Ting, 1992, D. car-pathicus Zeman, 1979, D. nipponensis Uchikawa &Kitaoka, 1981, D. rwandae Fain, 1993, D. wutaiensis Gu& Ting, 1992.

EVOLUTION OF THE GENUS DEFINITIONCOMPARED WITH THE OTHER GENERAOF THE GROUP

This early genus definition follows a somewhatcomplex evolution, and it is necessary toexplore its history throughout the literature in

order to understand it. Dugès (1834) described thegenus Dermanyssus as follows: “Palporum articulus5us minimus; labium acutum; mandibulae maribuschelatae, ungue longissimo, feminis ensiformes; corpusmolle; pedes antici longiore; coxae contiguae. Larvaehexapodae, adultis vix dissimiles”. Such a morpholo-gical description appears today extremely general andfits most current mesostigmates. About a century later,in 1923 and in 1936, Ewing provided short surveys ofthis genus in North America. The first one, includedin a review of North American dermanyssids, listedonly two species, and the second survey, being a com-pact summation, included several recently describedspecies.In 1958, Strandtmann and Wharton, in a large opusreviewing the classification of the mesostigmates para-sitic on vertebrates, pointed out the serious need ofrevision of the genus Dermanyssus: “The genus is inneed of revision. It is doubtful that all the species listedbelow really are specific entities” (Strandtmann &Wharton, 1958, p. 122).From then on, three steps can be distinguished, on thewhole, which lead to the current and stabilized des-cription. Many other genera have been created, whichare more or less closely related to Dermanyssus.First, two of these genera are very closely related toDermanyssus: Allodermanyssus Ewing, 1923 and Lipo-nyssoides Hirst, 1913. The exploration of both thesegenera compared with Dermanyssus helped the defi-nition to become more precise. Krantz (1959) and Sheals(1962) took part in the evolution of the genus defini-tion, discussing the relationships among the three clo-sely related genera. According to both the authors,Allodermanyssus was not valid anymore. But Krantz consi-dered Allodermanyssus synonymous with Dermanyssus,whereas Sheals considered Allodermanyssus synony-mous with Liponyssoides. This discordance induced a

deeper investigation of the description of Derma-nyssus. Krantz described the first Dermanyssus specieshaving a divided dorsal shield in the adult stage: D. tris-cutatus (dorsal shield short, several metanotal scutellapresent). He also pointed out the fact that the discri-mination between Allodermanyssus and Dermanyssusbased on the character incomplete/complete dorsalshield in the adult stage is not correct anymore. Buthe neglected to consider the genus Liponyssoides. Shealsexamined the three genera together. In order to explainthe new synonymy he established between Alloder-manyssus and Liponyssoides, he provided some argu-ments concerning the ontogeny (one seta less on thefemur and one less on the palp genu in the adult stagein Dermanyssus).Another apparently important argument concerned thechaetotaxy of the dorsal shield and applied, accordingto Sheals, not only to Dermanyssinae but also to Macro-nyssinae (today Macronyssidae; cf. infra): he consideredthe presence/absence of seta j3 a character related tothe host group in both taxa (present in all parasites ofmammalian and absent in all parasites of birds). How-ever, it should be noted that the more recently des-cribed species D. trochilinis is an exception to such ahypothesis: it is parasitic on birds and doesn’t lack j3.Moreover, genus Ornithonyssus (Macronyssidae) lacksj3 and includes species which are parasitic on mam-mals (e.g. O. bacoti on rodents). In short, Sheals waswrong in this last hypothesis.Secondly, Evans & Till (1962) stabilized Dermanyssusdescription: they wrote the first worldwide monographon the genus Dermanyssus. Many generic characterswere based on the ontogeny and the chaetotaxy ofshields and legs.Finally, Radovsky (1966, 1967) established Macro-nyssinae (Mesostigmata: Dermanyssidae) as themacronyssid family. This status change is very impor-tant. From then on, dermanyssids species i.e. Der-manyssus spp. and Liponyssoides spp., are to be dis-tinguished from macronyssids in having the 2nd

segment of the chelicerae elongate and very slender(the 1st one is elongate, and differently conformed,in Macronyssids), the chelae reduced (edentate, buteach digit visible with an optic miscroscope in Macro-nyssids) and a deutonymphal stage which needs ablood meal in order to accomplish its moulting (deu-tonymphs moult without feeding, as do larvae, inmacronyssids).

CURRENT DIAGNOSIS OF DERMANYSSIDAE

Dermanyssids are characterized among Dermanyssoi-dea by the following characters:. Adult femalesGnathosoma-chelicerae: distal segment (= 2nd) of thefemale chelicerae conspicuously elongated and slender,





Dermanyssus species Host species

D. alaudae (Schrank, 1781) Alauda arvensis Alauda: Alaudidae: PasseriformesLululla arborea Alauda: Alaudidae: Passeriformes

D. americanus Ewing, 1922 Carpodacus lexicanus Carpodacus: Fringilloidea: PasseriformesEmberiza cioides Emberiza: Fringillidae: PasseriformesOtus asio Otus: Strigidae: StrigiformesPasser domesticus Passer: Passeridae: PasseriformesP. montanus Passer: Passeridae: PasseriformesSerinus canaries Serinus: Fringillidae: PasseriformesSitta sp. Sitta: Sittidae: Passeriformes

D. antillarum Dusbabek & Cerny, 1971 Mimus polyglottos orpheus Mimus: Sturnidae: PasseriformesPasser domesticus Passer: Passeridae: PasseriformesAccipiter striatus fringilloides Accipiter: Accipitridae: CiconiiformesTachornis phoenicobia Tachornis: Apodidae: Apodiformes

D. brevirivulus Gu & Ting, 1992 Galerida cristata leautungensis Galerida: Alaudidae: Passeriformes

D. brevis Ewing, 1936 Alauda arvensis Alauda: Alaudidae: PasseriformesEremophila alpestris Eremophila: Alaudidae: Passeriformes

D. carpathicus Zeman, 1979 Phoenicurus phoenicurus Phoenicurus: Muscicapidae: PasseriformesParus major Parus: Paridae: Passeriformes

D. chelidonis Oudemans, 1939 Carduelis carduelis Carduelis: Fringillidae: PasseriformesDelichon urbica Delichon: Hirundinidae: PasseriformesHirundo rustica Hirundo: Hirundinidae: PasseriformesParus coeruleus Parus: Paridae: PasseriformesRiparia riparia Riparia: Hirundinidae: PasseriformesPtyonoprogne rupestris Ptyonoprogne = Hirundo: Hirundinidae: Passeriformes

D. faralloni Nelson & Furman, 1967 Oceanodroma homochroa Oceanodroma: Hydrobatidae: CiconiiformesCepphus columba Cepphus: Alcidae: CiconiiformesPtychoramphus aleutica Ptychoramphus: Alcidae: Ciconiiformes

D. gallinae (De Geer, 1778) Acrocephalus arundinaceus Acrocephalus: Sylviidae: PasseriformesAegolius funereus Aegolius: Strigidae: StrigiformesCarduelis carduelis Carduelis: Fringillidae: PasseriformesCarduelis spinus Carduelis: Fringillidae: PasseriformesColumba livia Columba: Columbidae: ColumbiformesDelichon urbica Delichon: Hirundinidae: PasseriformesEmberiza citrinella Emberiza: Fringillidae: PasseriformesErithacus rubecula Erithacus: Muscicapidae: PasseriformesFicedula albicollis Ficedula: Muscicapidae: PasseriformesFicedula hypoleuca Ficedula: Muscicapidae: PasseriformesHirundo rustica Hirundo: Hirundinidae: PasseriformesJynx torquilla Jynx: Picidae: PiciformesMerops apiaster Merops: Meropidae: CoraciiformesParus major Parus: Paridae: PasseriformesP. ater Parus: Paridae: PasseriformesPasser domesticus Passer: Passeridae: PasseriformesP. montanus Passer: Passeridae: PasseriformesPhoenicurus phoenicurus Phoenicurus: Muscicapidae: PasseriformesRemiz pendulinus Remiz: Paridae: PasseriformesRiparia riparia Riparia: Hirundinidae: PasseriformesSerinus canarius Serinus: Fringillidae: PasseriformesSitta europaea Sitta: Sittidae: PasseriformesSturnus vulgaris Sturnus: Sturnidae: Passeriformes

Other wild birds and numerous Galliformes, Anseriformesspecies of domestic fowl, etc. Sometimes on mammalian species (insectivora, rodents, man)

D. gallinoides Moss, 1966 Asyndesmus lewis Asyndesmus = Melanerpes: Picidae: PiciformesColaptes cafer (= C. auratus) Colaptes: Picidae: PiciformesDendrocopos pubescens Dendrocopos: Picidae: PiciformesDryocopus pileatus Dryocopus: Picidae: PiciformesSphyrapicus varius Sphyrapicus: Picidae: Piciformes

Table II (to be continued).




D. grochovskae Zemskaya, 1961 Nucifraga caryocatactes Nucifraga: Corvidae: PasseriformesDendrocopos leucotos Dendrocopos: Picidae: PiciformesD. major Dendrocopos: Picidae: PiciformesPicus awokera awokera Picus: Picidae: Piciformes

D. hirsutus Moss & Radovsky, 1967 Colaptes cafer (= C. auratus) Colaptes: Picidae: Piciformes

D. hirundinis (Hermann, 1804) Acrocephalus arundinaceus Acrocephalus: Sylviidae: PasseriformesA. palustris Acrocephalus: Sylviidae: PasseriformesA. scirpaceus Acrocephalus: Sylviidae: PasseriformesAnthus arboreus Anthus: Motacillidae: PasseriformesApus affinis Apus: Apodidae: ApodiformesAquila pomarina Aquila: Accipitridae: PasseriformesAnser anser Anser: Anatidae: AnseriformesAythya fuligula Aythya: Anatidae: AnseriformesA. ferina Aythya: Anatidae: AnseriformesChaetura pelagica Chaetura: Apodidae: ApodiformesColumba livia Columba: Columbidae: ColumbiformesDelichon urbica Delichon: Hirundinidae: PasseriformesDendrocopos pubescens Dendrocopos: Picidae: PiciformesFicedula albicollis Ficedula: Muscicapidae: PasseriformesHirundo rustica Hirundo: Hirundinidae: PasseriformesH. urbica Hirundo: Hirundinidae: PasseriformesIridoprocne bicolor Iridoprocne = Tachynecita: Hirundinidae: PasseriformesLanius minor Lanius: Laniidae: PasseriformesL. collurio Lanius: Laniidae: PasseriformesLuscinia megarhynchos Luscinia : Muscicapidae: PasseriformesMerops apiaster Merops: Meropidae: CoraciiformesMicropus affinis Micropus = Apus: Apodidae: ApodiformesParus caeruleus Parus: Paridae: PasseriformesP. major Parus: Paridae: PasseriformesP. palustris Parus: Paridae: PasseriformesPasser montanus Passer: Passeridae: PasseriformesP. domesticus Passer: Passeridae: PasseriformesPetrochelidon pyrrhonota Petrochelidon: Hirundinidae: PasseriformesPhoenicurus ochruros Phoenicurus: Muscicapidae: PasseriformesRemiz pendulinus Remiz: Paridae: PasseriformesRiparia riparia Riparia: Hirundinidae: PasseriformesSitta europaea Sitta: Sittidae: PasseriformesStrix aluco Strix: Strigidae: StrigiformesSturnus vulgaris Sturnus: Sturnidae: SturniformesTaeniopygia guttata castanotis Taeniopygia: Passeridae: PasseriformesTroglodytes aedon Troglodytes: Certhiidae: PasseriformesT. troglodytes Troglodytes: Certhiidae: PasseriformesTurdus torquatus Turdus: Muscicapidae: PasseriformesVireo olivaceus Vireo: Vireonidae: Passeriformes

Sometimes on mammalian species (insectivora, rodents)

D. longipes Berlese & Trouessart, 1889 Passer domesticus Passer: Passeridae: Passeriformes(incertae sedis)

D. nipponensis Uchikawa & Kitaoka, 1981 Picus awokera awokera Picus: Picidae: Piciformes

D. passerinus Berlese & Trouessart, 1889 Emberiza cirtus Emberiza: Fringillidae: Passeriformes(incertae sedis) Ficedula albicollis Ficedula: Muscicapidae: Passeriformes

Jynx torquilla Jynx: Picidae: PiciformesParus major Parus: Paridae: PasseriformesPasser italiae Passer: Passeridae: PasseriformesSturnus vulgaris Sturnus: Sturnidae: Sturniformes

D. prognephilus Ewing, 1933 Progne subis subis Progne: Hirundinidae: PasseriformesColaptes cafer (= C. auratus) Colaptes: Picidae: PiciformesDendrocopos pubescens Dendrocopos: Picidae: PiciformesMelanerpes erythrocephalus Melanerpes: Picidae: PiciformesMolothrus ater Molothrus: Fringillidae: PasseriformesSialia sialis Sialia: Muscicapidae: Passeriformes

Table II (to be continued).

with chelae strongly reduced (Fig. 2); cornicules mem-branous, flexible (not acute as in free-living mesostig-mate species) and convergent; Podosoma-legs: coxaewithout spurs.. Adult males differ from adult females mainly inhaving more extensive sclerotization both ventrally anddorsally (holoventral shield in most cases, larger dorsalshield, including more dorsal setae than in female) andmodified chelicerae (less elongated and much broaderthan in female, chelae with a long spermadactyl on themovable digit). The tarsi of legs III and IV bear a tooth-like protuberance. Moreover, the genital orifice is cons-picuous and presternally situated.

KEY FOR DERMANYSSID GENERA

Few characters remain currently available for dia-gnosis between Dermanyssus and Liponyssoides. Hirst(1913) described Liponyssoides as a subgenus of Der-manyssus mainly based on a weak difference in theproportion of capitulum. Moss (1967) stated otherclear differences between them based on sternal shieldshape and chaetotaxy of dorsal shield, sternal shieldand legs. But the three species L. intermedius, D. tro-chilinis and partially D. antillarum appear as yetintermediate between both genera with chaetotaxy. Asa result, only following elements can be used for dia-gnosis of the two genera.Sternal shield roughly crescent-shaped; usually para-sitic on birds……………………DermanyssusSternal shield roughly hexagonal; parasitic onrodents.........................................Liponyssoides

SPECIES SPECIFIC CHARACTERSWITHIN THE GENUS

TRADITIONAL SYSTEMATICS

Here will be dealt with characters which havebeen used as arguments for species description,not with diagnostic characters. For all the follo-

wing characters, only adult females are to be conside-red.Most of specific-level discriminant characters are basedon chaetotaxy of the legs and dorsal shield and on therelative length of peritremes against the position fromterminating over coxae IV to over coxae III-I. Few other morphological characters are used. A markeddifference of the dorsal setae length is very conspi-cuous between central setae on dorsal shield (j4-j6 +“J” series except J5) and the other setae, which aresituated all around (J5, “z-Z” series, “r-R” series, “s”series), in the seven following species: D. alaudae,D. brevis, D. brevirivulus, D. hirsutus, D. grochovskae,D. quintus, D. rwandae. In these species, the centralseta length is near one-quarter the length of the per-ipheral ones, whereas they are all subequal in the otherDermanyssus species.A character concerning dorsal shield development isfound only in five species: mesonotal scutella are pre-sent only in D. americanus, D. antillarum, D. trans-vaalensis, D. triscutatus and D. wutaiensis. The natureof these platelets seems to correspond to the primarydorsal plates, which don’t become coalesced as they




D. quintus Vitzthum, 1921 Dendrocopos major Dendrocopos: Picidae: PiciformesD. pubescens Dendrocopos: Picidae: PiciformesDryobates leucotes Dendrocopos: Picidae: PiciformesD. major Dendrocopos: Picidae: PiciformesPicoides pubescens Picoides: Picidae: PiciformesP. tridactylus Picoides: Picidae: PiciformesPicus viridis Picus: Picidae: Piciformes

D. rwandae Fain, 1993 Apus affinis Apus: Apodidae: Apodiformes

D. transvaalensis Evans & Till, 1962 Hirundo spilodera Hirundo: Hirundinidae: PasseriformesPetrochelidon spilodera Petrochelidon: Hirundinidae: Passeriformes

D. triscutatus Krantz, 1959 Hirundo sp. Hirundo: Hirundinidae: PasseriformesPetrochelidon pyrrhonota Petrochelidon: Hirundinidae: Passeriformes

D. trochilinis Moss 1978 Trochilidae Trochilidae: Apodiformes

D. wutaiensis Gu & Ting, 1992 Passer montanus Passer: Passeridae: Passeriformes

Table II. – List of species currently included in Dermanyssus and their known host species, established with the help of following refe-rences: Berlese & Trouessart (1889), Bory de Saint-Vincent (1828), Dusbabek & Cerny (1971), Evans & Till (1962), Evans & Till (1964),Ewing (1922, 1933), Fain (1993), Fend’a & Schniererová (2004), De Geer (1778), Gu & Ting (1992), Haitlinger (1987), Hermann (1804),Krantz (1959), Moss (1966), Moss (1978), Moss (1970), Nelson & Furman (1967), Nosek & Lichard (1962), Schrank (1776), Uchikawa &Kitaoka (1981), Uchikawa & Takahashi (1985), Vitzthum (1921), Zeman (1979), Zeman & Jurík (1981), Zemskaya (1971), Fend’a (2006,unpublished data) and collection data from Pr. A. Fain.Taxonomic bird data follow Peterson’website.



do in other Dermanyssus species in the adult stage. Thedorsal shield is more strongly reduced in D. triscutatusand D. antillarum than in the three other species: fromthe “J” series, only J1 is on the shield in D. triscutatusand in D. antillarum, whereas at least J1 and J2 are onthe shield in D. americanus, D. transvaalensis andD. wutaiensis. Moreover, the dorsal scutella are pro-portionally quite smaller in these three latter speciesthan in the two former.Three very conspicuous characters are found each ina single species and concern particular opisthoventralsetae:- a ventral neotrichy in form of a cluster of elongate,simple setae laterad of the anal shield is present onlyin D. hirsutus;- a U-shaped row of very large and deeply rooted setaeon the opisthogaster is present only in D. quintus;- several distally inflated setae situated posteriorly onthe idiosoma are present only in D. antillarum.No other Dermanyssus species possess such species spe-cific, apomorphic characters.Additionally, in D. quintus, the wider than long analplate is another species specific character.Nevertheless, apart from these characters, it should benoted that the leg and dorsal shield chaetotaxy providesmost of the characters traditionally used for species dis-crimination in the genus Dermanyssus. The reliability ofsome of them seems to be doubtful. Evans & Till (1962)emphasized with several remarks many intraspecificvariations, concerning the chaetotaxy: “The chaetotaxyof the venter of the opisthosoma shows considerableintraspecific variation” (p. 277). “The chaetotaxy of thevarious segments [of the legs] is considerably morevariable, both inter- and intra-specifically, than in thefree living and facultative parasitic Laelaptidae” (p. 278).Moss (1968) also noticed that characters of the legchaetotaxy usually seemed to be the most variable.Other than the chaetotactic characters of the legs anddorsal shield, the other traditionally most used charac-ter is the relative length of the peritreme. However thestates of this character don’t provide any defined limit.The extension of the peritreme varies continuouslyfrom coxa IV to coxa III to coxa IV to coxa I withouta sharp gap from one species to another in the genusDermanyssus and with intraspecifical variations. Indeed,from Moss (1978) the following data can be extracted,in increasing order: the peritreme extends from thecoxa IV “to middle of coxa III” in D. transvaalensisand D. chelidonis, “not as far as anterior margin of coxaIII” in D. alaudae and D. brevis, “to or past anteriormargin of coxa III” in D. americanus, “past anteriormargin of coxa III” in D. triscutatus, “to middle ofcoxa II” in D. hirsutus, “to middle or anterior marginof coxa II” in D. gallinae and D. gallinoides, “to ante-rior margin of coxa II” in D. grochovskae and D. hirun-dinis, “to middle of coxa I” in D. prognephilus, “past

posterior margin or to middle of coxa I” in D. trochi-linis and extends “anterad of base of coxa I” in D. faral-loni. Although the character states seem to overlap eachother, it takes part in several new species arguments.For instance, the first of the two characters which arecited by Uchikawa as a distinctive property in D. nip-ponensis is the remarkable length of the peritreme. Itslength is also the first of the five characteristics usedby Nelson and Furman in order to distinguish D. faral-loni from the a priori most closely related speciesD. hirundinis. Moreover, Oudemans (1939) cited thischaracter as the main difference distinguishing D. che-lidonis from D. hirundinis and D. gallinae. It would beof interest to investigate the reliability of such a cha-racter as an argument for species description.In short, many of the main traditionally used speciesspecific characters are problematic. Chaetotactic cha-racters of legs and dorsal shield are variable intraspe-cifically and the peritreme relative length doesn’t seemto provide states of characters that are precise enough.

AN ATTEMPT WITH NUMERIC TOOLS

Moss tried to use more objective tools than the tradi-tional systematics in order to explore the relationshipsbetween the Dermanyssus species: in 1967, he publi-shed a work on some numeric taxonomy theories(phenetics), in which he used the genus Dermanyssusas a model. He complained about the difficulties of fin-ding characters in this genus and considered this tobe due to a reduction and loss of structures correlatedwith parasitism. As a result, he selected quantitative cha-racters, which are much easier to find than qualitativeones. His analysis resulted in two subdivisions of thegenus: two subgenera, Dermanyssus and Microder-manyssus, are to be distinguished from one anotherby the setation of genu II (two al setae, one av andone pv setae in Dermanyssus (Dermanyssus); one alsetae, no av and pv setae in Dermanyssus (Microder-manyssus)) and by the size of the unengorged female’sbody. Subgenus Dermanyssus included D. chelidonis,D. gallinae, D. gallinoides, D. grochovskae, D. hirsutus,D. hirundinis, D. prognephilus, D. quintus, D. trans-vaalensis and D. triscutatus. Subgenus Microdermanys-sus included D. alaudae, D. americanus and D. brevis.Moreover, two species-groups were separated in thesubgenus Dermanyssus: the hirsutus-group, with,among other discriminant characters, the seta al1 ofpalp genu spiniform, and the gallinae-group, with theseta al1 of palp genu spatulate. The hirsutus-groupincluded D. grochovskae, D. hirsutus and D. quintus,whereas the gallinae-group included D. chelidonis,D. gallinae, D. gallinoides, D. hirundinis, D. progne-philus, D. transvaalensis and D. triscutatus. Moss usedthese new subdivisions in his 1968 and 1978 keys. The1978 article was somewhat more developed than the



one from 1968 and took three new species into account(D. antillarum, D. trochilinis and D. faralloni). It pro-vided a variety of information on the genus Derma-nyssus (hosts, phylogenetic results from Moss (1967),remarks on the high variability of some characters,...).The status of the two subgenera and the two species-groups is re-examined with the changes induced by thethree new species. Actually, the introduction ofD. antillarum and D. trochilinis into Moss’ pheneticanalysis introduced some problems with the subdivi-sions. Nevertheless, Moss decided to continue to reco-gnize the two subgenera and the two species-groupstemporarily because he was expecting some additionalelements from a new study he was preparing at thattime. Unfortunately, this study was never published.As well as these doubts about the interrelationships ofspecies in the genus Dermanyssus, Moss warned in1978 that any person who wanted to attempt identifi-cation with his key to remain careful because of theconsiderable variation within species: “The most usefulsetae for identification are those in the “j” series of thedorsum. [...] Leg setae are also useful in species iden-tification, but tend to be more variable within speciesthan dorsal setae. [...] Variation within species is consi-derable. One is advised to consider several charactersfrom several individuals of a sample when attemptingan identification. A recent example of such variationis described below for D. gallinoides. Most key cou-plets list several alternative features in view of this pro-blem.” (Moss, 1978:633-634). Moss also invalidatedone of the discriminant characters for D. gallinoides:“Dorsal shield scaling has been a reliable feature untilrecently for the separation of D. gallinoides and D. gal-linae. Two specimens just provided by N. Wilson havepredominantly scaled teeth in one case and comple-tely scaled teeth in the other, but otherwise key toD. gallinoides” (Moss, 1978:634)

SPECIES WHOSE NOMENCLATURAL STATUS IS NOT CLEAR

About D. hominis, Dugès wrote as follows: “D. homi-nis; sorte d’acaride, Bory St-Vincent”. Bory de Saint-Vin-cent described it in 1823, in a memoir which was readduring a regular meeting of the French “Académie desSciences”. He did not attribute any name and any sys-tematic position within the Acari group to this species.Latreille and Savigny were designated to judge such aposition, but did not do so (cf. Académie des Sciences,1823-1828). Mites of this species were found infestingthe body of a woman. The description of this speciesand its illustration by Bory de Saint-Vincent (1828)appear very similar to D. gallinae. This last species hasbeen reported from humans several times (Beck, 1999;Cremer & Morrien, 1962; Holz J., 1954; Pampiglione etal., 2001). Moreover, there is no type material availableand the drawing of Bory de Saint-Vincent is not suffi-

cient to morphologically identify this species. As a resultD. hominis should be synonymized with D. gallinae.Two species are considered incertae sedis: D. passe-rinus Berlese & Trouessart, 1889 and D. longipes Ber-lese & Trouessart, 1889. Evans & Till (1962) consideredthem incertae sedis because of the inadequate des-criptions and the damaged type material in the Ber-lese collection. In Moss (1978), a part of the discus-sion refers to both of these species, in order to beginto solve this problem. In short, Zemskaya suggestedthat D. passerinus should be conspecific with D. ame-ricanus (which should be then a junior synonym), butdid not demonstrate this. Moss adds an argument:both hosts of these species seem to be conspecific too,according to an ornithologist. Moreover, according toMoss, it is most likely that D. passerinus and D. lon-gipes are conspecific. The type of D. longipes is tooopaque to confirm such a hypothesis.However, as the type of D. longipes (No. 52-47) isalmost opaque and essentially unusable according toMoss and as this species has not been cited for a longtime, it could be more appropriate to establish it as anomen dubium instead of incertae sedis. Indeed, thesystematic position within Dermanyssus does notappear to be doubtful, compared to its precise iden-tity, which is doubtful.As for D. passerinus, the type specimens are partiallyopaque and some papers include it in some acaro-faunal lists (Nosek & Lichard, 1962; Zemskaya, 1971;Zeman & Jurík, 1981). So, the problem is more impor-tant with this species, which we suggest be consideredspecies inquirenda.D. lacertarum and D. richiardii are also problematicspecies. D. richiardii had been collected on two dif-ferent species of insect, the hymenopteran Xylocopaviolaceus and the lepidopteran Cossus ligniperda (Canes-trini & Fanzago, 1877), which are not common hostsfor the Dermanyssus. As for D. lacertarum, it was trans-ferred from genus Ricinus by Canestrini (1877) in thesame paper, with the only following sentence: “Duespecie vi sono citate come nuove, il Ricinus lacer-tarum, e l’Acarus penetrans; il primo sembra un Der-manyssus, il secondo è una forma larvale”. From thatdate, we did not find any more information on thesetwo species. Maybe they should be established asnomina dubia. In any case, given the informationcited above, they can not be included in the genusDermanyssus anymore.

CONCLUSION AND PERSPECTIVES

The systematics of Dermanyssus is not completelyclear as yet. Its history is complex. Dermanyssusseems to be well defined today, but species

within the genus remain less clearly defined. Moreover,



not only was the last work on Dermanyssus not com-pletely carried out, but also five new species have beendescribed since this last review (Moss, 1978). The relia-bility of numerous traditional characters needs to bere-examined.Today, 23 species are included in this genus, two ofwhich are really doubtful. The status of D. longipes andD. passerinus is to be re-examined. D. americanusmight be a junior synonym of D. passerinus. D. homi-nis is a synonym of likely D. gallinae. Some other earlyspecies might likely be synonymized too. The reliabi-lity of the five species described after Moss (1978),needs to be checked and they have to be integratedin a review of the entire genus.For these reasons, it appears necessary to review thegenus Dermanyssus at the specific level, which we planto do, with the help of cladistic tools. Two major ques-tions need to be answered. First, the correct definitionof genus, even if it seems to be right using traditionaltools, has to be checked by testing the monophyly ofthe group. Secondly, the a priori most problematicquestion of the species definitions within the genusshould be explored, and maybe some species shouldbe synonymized. Finally, as morphological charactersseem to be insufficient, it seems necessary to add mole-cular characters to the phylogenetic analysis.

ACKNOWLEDGEMENTS

We are grateful for the help and advice ofP. Fend’a (Faculty of Natural Sciences, Come-nius University, Bratislava, Slovak Republic),

R. Nannelli (Istituto Sperimentale per la Zoologia Agra-ria, Firenze, Italia), J. Hallan (Texas A & M University,College Station, Texas), O. Bain and I. Roy (MuséumNational d’Histoire Naturelle, Paris, France), H. Ferrière(Université Paris I-Sorbonne, Paris, France), V. Marengo(Muséum d’Histoire naturelle, Lyon, France) and to Pr.F.J. Radovsky and an anonymous reviewer for cons-tructive reviewing and valuable suggestions.

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Reçu le 15 novembre 2006Accepté le 12 mars 2007



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