Sergey Mironov1 & Georges Wauthy2
1 Zoological Institute RAS, Saint Petersburg, Russia 2 Institut royal des Sciences naturelles de Belgique, Bruxelles, Belgique
SYSTEMATIC REVIEW AND PHYLOGENY OF THE FEATHER MITE GENUS METAPTERONYSSUS GAUD, 1981 (ASTIGMATA: PTERONYSSIDAE) ASSOCIATED WITH AFRICAN PASSERINES
Mironov, S.V. & G. Wauthy, 2006. Systematic review and phylogeny of the feather mite genus Metapteronyssus Gaud, 1981 (Astigmata: Pteronyssidae) associated with African pas- serines. – Tijdschrift voor Entomologie 149: 21-53, figs. 1-20, tables 1-3. [issn 0040-7496]. Published 1 June 2006. A systematic review of the feather mite genus Metapteronyssus Gaud, 1981 is given, including an improved diagnosis of the genus, revised key to species, description of four new species from African passerines, cladistic analysis of the genus, and a brief discussion of associations with passerine hosts. The type species Metapteronyssus glossifer (Gaud, 1953) is redescribed based on the type material from Euplectes franciscanus (Isert, 1789) (Ploceidae); new species found in the course of study are described as follows: Metapteronyssus amadinae sp. n. from Amadina fasciata alexanderi Neumann, 1908 (Estrildidae), M. anaplecti sp. n. from Anaplectes rubriceps (Sundevall, 1850) (Ploceidae), M. hordacei sp. n. from Euplectes hordaceus Linnaeus, 1758 (Ploceidae), and M. pseudonigritae sp. n. from Pseudonigrita cabanisi (Fischer et Reichenov, 1884) (Passeridae). Brief diagnoses are provided for all formerly described species. Maximum parsimony based on 34 morphological characters revealed three phylogenetic line- ages treated as species groups: angolensis, glossifer, and plocepasseri. Two clear morphological tendencies realized independently in these lineages are traced. One tendency is a progressive reduction of the central hysteronotal sclerite in females; in most derived species of each lineage it is reduced to a thin rod-shaped sclerite or is completely lost. The other tendency is a narrow- ing of the opisthosomal terminus and development of terminal membranous lobules in males that is realized in each lineage in a different way. Based on the phylogenetic relationships among mite species, their currently known distribution among passerines, and taking into consideration phylogenetic relationships between higher passerine host families, we hypothesize that the genus Metapteronyssus originated on the ances- tors the superfamily Passeroidea (parvorder Passerida). We suggest that owing to subsequent cospeciation, its representatives have been dispersed on estrildid finches (Estrildidae), weavers (Ploceidae), and sparrows (Passeridae), and apparently have gone extinct on other families of Passeroidea. Correspondence: S.V. Mironov, Zoological Institute, Russian Academy of Sciences, Saint Petersburg, 199034, Russia. E-mail:����������������� astigmata@�zin.ru������ G. Wauthy, Institut royal des Sciences naturelles de Belgique, rue Vautier 29, B - 1000, Bruxelles, Belgique. E-mail: [email protected] Key-words. – Feather mites; Analgoidea; Pteronyssidae; Metapteronyssus; systematics; host- parasite associations; Passeriformes, Africa.
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Table 1. Characters used in the phylogenetic analysis.
No Characters and coding
Males and females 1 Prodorsal shield posterior to setae se in both: developed (0), not developed (1). 2 Epimerites I: free (0), fused as a narrow U (1). 3 vertical seta vi: present (0), absent or rudimentary (1). 4 Size of setae ba of tarsi I, II: long, setiform (0), reduced to small spine or absent (1). 5 Spine–like seta ba of tarsi I, II: present (0), absent (1).
Males 6 Lateral hysteronotal ridges in male: absent (0), present (1). 7 Median hysteronotal ridge: absent (0), present (1). 8 Posterolateral margins of opisthosoma: gradually attenuate to posterior end (0), strongly convex (1). 9 Position of openings gl: on lateral margins of hysteronotal shield (0), in postero-lateral incisions of hysteronotal shield (1). 10 Margin of opisthosoma between setae ps2: almost straight or moderately convex (0), with median extension of approximately trapezoidal form (1) 11 Entire opisthosomal membrane between setae ps2: absent (0), present (1). 12 terminal extensions of entire opisthosomal membrane: absent (0), present (1). 13 Membranous terminal lobules between setae h3: absent (0), present, short (1), present, finger-like (2). 14 Length of membrane or membranous extensions from bases of setae h3 to distal margin: less than 10 μm (0), 15-25 μm (1). 15 Subterminal membranes between setae h2 and h3: absent (0), present (1). 16 transventral sclerite along midline: equal or longer than 10 μm (0), less than 10 (1). 17 Position of setae 3a: on transventral sclerite (0), posterior to transventral sclerite (1). 18 * adanal shield: absent (0), present (1). 19 Median part of adanal shield: present (0), absent (1). 20 Lateral parts of adanal shield: not separated from median part (0), separated (1), absent (2). 21 tarsus III: with curved apex (0), with straight apex (1). 22 apex of tarsus III: claw-like (0), bidentate (1).
Females 23 Distance between setae e2: further apart from each other than distance between setae se (0), closer to each other than setae se (1). 24 Central sclerite: not differentiated from main body of hysteronotal shield (0), present (1), absent (2). 25 General form of central sclerite: longitudinal bar-like plate extending to trochanters III (0), ovate plate (1). 26 Width of central sclerite: greatest width over 1/3 of idiosoma width (0), width about 30-50 μm (1), less than 30 μm (2). 27 Splitting of central sclerite: not split (0), split into anterior and posterior fragments (1). 28 * Posterior end of central sclerite: free (0), fused with pygidial sclerites (1). 29 * Position of setae d1: on central sclerite (0), off central sclerite (1), variable in a species, on or off central sclerite (2). 30 Lateral sclerites: not separated from the main body of hysteronotal shield (0), absent (1), present (2). 31 General form of lateral sclerites: elongated plate of irregular form (0), narrow longitudinal band (1), ovate (2). 32 Posterior part of lateral sclerites: relatively wide (0), narrowed (1). 33 External copulatory tube: absent (0), present (1). 34 Length of external copulatory tube: short, less than 10 μm (0), long, 15-45μm (1)
(*) Parsimony uninformative characters.
Feather mites of the family Pteronyssidae Oude- secondary flight feather of their hosts. mans, 1941 (Astigmata: Analgoidea) currently in- This genus initially included a single species, clude over 130 species in 22 genera (Faccini & Atyeo Metapteronyssus glossifer (Gaud, 1953), described 1981, Gaud & Atyeo 1996, Mironov 2001, Mironov from Euplectes franciscanus (Isert, 1789) (Ploceidae) & Wauthy 2005a). Within this family, the genus and also reported from several other ploceids and Metapteronyssus Gaud, 1981 represents small-sized estrildid finches in Africa (Gaud 1953, Faccini & pteronyssid mites known from three families of pas- Atyeo 1981). Later, Mironov and Kopij (2000) de- serine birds, Estrildidae, Passeridae, and Ploceidae. As scribed the second species, M. angolensis Mironov for most pteronyssids, representatives of this genus et Kopij, 2000, from the blue waxbill Uraeginthus inhabit the ventral surfaces of vanes of primary and angolensis (Linnaeus, 1758) (Estrildidae). Finally,
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nine more new species were described from various small lobe-like extensions with membranous margins African sparrows, weavers, and estrildid finches and developed on the posterior end of the opisthosoma a key to species was proposed for the first time by between setae h3 (figs. 1a, b, 2a, b) are referred to Mironov (2002). as ‘terminal lobules’. In some species the border be- In the course of our systematic study of feather tween the membranous marginal part and scarcely mites associated with African passerines (Mironov & sclerotized basal part of the lobules may be not clearly Wauthy 2005a-c) we have found four new species of expressed. In any case, only the membranous part of Metapteronyssus and also re-examined type material these lobules is referred to as the ‘terminal membrane’. of all formerly described species. Our examination of Two small semicircular membranous extensions situ- the type series of M. glossifer and additional material ated between the bases of setae h2 and h3 are referred revealed that in the generic revision of pteronyssids to as ‘subterminal membranes’ (Mironov 2002) (Faccini & Atyeo 1981) this species was redescribed (fig. 2b). The anal area bearing anal discs and anal incorrectly. These authors actually redescribed anoth- opening is flanked from lateral sides by a pair of the er species and gave incorrect data on host associations adanal membranes. that has also led to inappropriate interpretation of All measurements in the descriptions are given M. glossifer in subsequent publications (Mironov & in micrometres (μm). A full set of measurements is Kopij 2000, Mironov 2002). given only for the holotype (male) and one paratype The main goal of our paper is a taxonomic review (female); the range of idiosomal size (length, width) is and phylogenetic analysis of the genus Metapteronyssus displayed for the rest of the paratype specimens. based on morphological characters. The taxonomic Specimen depositories and reference accession part contains an improved diagnosis of the genus numbers are given using the following abbrevia- Metapteronyssus, a revised key to species, descriptions tions: amnh – American Museum of Natural History, of four new species, redescription of the true type New York, USA; irsnb – uncatalogued collection of species, and brief diagnoses and taxonomic remarks Prof. A. Fain, I�����������������������������������������nstitut royal des Sciences naturelles de for all formerly known species. We also briefly discuss Belgique��������������������, Brussels, Belgium; mrac – Musée�������������� royal de known host associations of Metapteronyssus and, based l’Afrique central,�������������������� Tervuren, Belgium; nmb – National on the phylogeny of this genus, provide a preliminary Museum of Bloemfontein, Free State, South Africa; hypothesis for its evolution on passerines. uga – University of Georgia, Athens, USA; usnm – US National Museum of Natural History, Belts- ville, Maryland, USA; zisp – Zoological Institute of Material and methods the Russian Academy of Sciences, Saint Petersburg, Specimens Russia. Where the collection number consists of The main part of material used in the present two sections, the first section refers to the collection study was received from the Musée royal de l’Afrique number of the mite specimens and its depository, if central (Tervuren, Belgium), other material was ob- another depository is not specifically indicated; the tained from the Institut royal des Sciences naturelles second is a collection number of the respective host de Belgique (Brussels, Belgium), National Museum specimen. In those cases where the recent country of Bloemfontein (Free State, South Africa), and name differs from that given in the slide-mounted University of Georgia (Athens, USA). specimen, location data also point out the original Diagnoses for the genus and species and descrip- country name, as written on the slide label; the ab- tions of new species are given in the standard formats breviation ‘coll.’ means ‘collector’. Taxonomy and used for pteronyssid taxa (Faccini & Atyeo 1981, scientific names of birds follow Dickinson (2003), Mironov 1992, 2001). The general morphological passerine phylogeny used in the discussion of host terms and leg and idiosomal chaetotaxy follow Gaud associations follows recent conceptions based on mo- & Atyeo (1996). Regarding the terms used for hyster- lecular studies (Sorenson & Payne 2001, Ericson & onotal shield fragments in females, we generally fol- Johanson 2003, Barker et al. 2004). low the scheme proposed by Mironov (1992) that was initially elaborated for the genus Pteroherpus Gaud, Phylogenetic analysis 1981. Qualitative morphological characters such as the For membranous structures of the opisthosoma in presence/absence of a structure or form of morpho- males of Metapteronyssus species we propose the fol- logical structure were used in the parsimony-based lowing terms, based in part on terms used by previous cladistic analysis (table 1). Two autoapomorphic authors. The entire membrane along all the posterior characters (unique features of hysteronotal shield in margin of the opisthosoma between bases of setae females of M. gaudi Mironov, 2002 and M. puylaerti ps2 (figs. 18a, b) is referred to as the ‘opisthosomal Mironov, 2002) were included to stress peculiarity of membrane’ following Faccini & Atyeo (1981). Two these species and also taking into consideration that
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these features may potentially represent characteris- absent (M. amadinae, M. pseudonigritae). Solenidion tics for species groupings, which could be recognised σ1 of genu I longer than ω1 on tarsus I; solenidion in future investigations of diversity. σ2 of genu I absent. Ventral membrane of tarsus I Representatives of three closely related pteron- about ½ length of the segment. Coxal fields I, II with yssid genera also associated with passerine hosts, striated tegument. Scutulanyssus dasyrhitidis (Gaud & Atyeo, 1981) from Male. – Opisthosomal lobes not expressed, Psalidoprocne fuliginosa Shelley, 1887 (Hirundinidae), opisthosoma gradually attenuate to posterior end Sturnotrogus subtruncatus (Trouessart, 1885) from (figs. 1a, 4a, 18a), or with strongly convex postero- Gracula religiosa Linnaeus, 1758 (Sturnidae), and lateral margins (plocepasseri group) (figs. 13a, 15a, b). Pteronyssoides (Holonyssoides) holoplax (Gaud et Mou- Most posterior end of opisthosoma between setae h3 chet, 1959) from Pycnonotus barbatus (Desfontaines, with a pair of small tongue-like membranous exten- 1789) (Pycnonotidae) were used as outgroups to test sions (terminal lobules), with or without subterminal monophyly of the genus Metapteronyssus. membranes between setae h2 and h3 (figs. 2a, 13a), In total, 18 taxa and 34 characters, two of which or posterior end of opisthosoma bluntly rounded and represent autapomorphies in outgroups and one in bears entire opisthosomal membrane between bases the ingroup (Metapteronyssus spp.), were included of setae ps2 (some species of the angolensis group) in the analysis (tables 1, 2). Constructing and edit- (figs. 18a, b, e). Terminal incision between terminal ing of the data matrix was done using NEXUS Data lobules small, usually slit-like. Supranal concavity long Editor 0.5.0 (Page 2001). All characters were treated and narrow, with well-sclerotized margins, open pos- as unordered; characters having multiple states were teriorly. Setae c2 on inner margin of humeral shields. interpreted as polymorphic and were not modified Setae ps1 anterior to setae h3. Hysteronotal shield free into binary characters. Reconstruction of phyloge- from humeral shields, with pair of bow-shaped heav- netic relationships was performed with PAUP 4.0 ily sclerotized ridges, anterior ends of which reach beta version for Windows 95/NT (Swofford 1998). antero-lateral angles of the shield, and with median The branch and bound algorithm was used for sclerotized ridge extending anterior from supranal the maximum parsimony analysis and reconstruc- concavity (fig. 1a). Hysteronotal gland openings gl in tion of phylogeny. For optimisation of character posterolateral incisions of hysteronotal shield. Coxal states, we used the delayed transformation option fields III closed. Transventral sclerite present, anterior (DELTRAN), which favours parallelism over reversal margin concave, tips of epiandrium short or scarcely when the choice is equally parsimonious. Bremer in- expressed. Anal discs circular. Adanal shield T-shaped, dices used for estimating support for branches were or split into three separate pieces, or represented only calculated with the program Autodecay (Eriksson by median or transverse fragment (figs. 2a, 9d, 11a, 1998). Drawings and editing of tree were accom- 13a). Adanal membranes present. Setae h3 setiform, plished with Winclada, version 1.0 (Nixon 1999). long. Cupules ih indistinct. Tarsus III straight or with slightly curved apical part, with bidentate apical proc- ess; seta r shorter or subequal in length to tarsus III. Systematics Tarsus IV without dorsobasal tooth. Female. – Idiosoma moderately elongated. Arrange- Family Pteronyssidae Oudemans, 1941 ment of hysteronotal shield fragments commonly consists of large central sclerite of ovate or bar-shaped Genus Metapteronyssus Gaud, 1981 form occupying median part of hysterosoma, pair of Metapteronyssus Gaud in: Faccini & Atyeo 1981: 34-36; lateral sclerites, and pair of small pygidial sclerites Mironov & Kopij 2000: 100; Mironov: 2002: 181. ����Type (fig. 3a); rarely central sclerite lost (M. anoplonotus) species:� Pteronyssus glossifer Gaud, 1953, ����������������by original des- ignation. (fig. 12b). Hysteronotal gland openings gl on anterior end or antero-lateral margin of lateral sclerites. Poste- rior margin of opisthosoma may have small lobe-like Diagnosis extensions bearing setae h3. External copulatory tube Both sexes. – Epimerites I fused as a narrow U, usually present, commonly cone-shaped, occupies connection of posterior tips may be very thin. Un- terminal position (figs. 3a, b, 6a-c); the tube absent in paired vertical seta vi absent. Prodorsal shield occu- M. plocepasseri (fig. 14b). Epigynium semicircular or pies antero-median part of pronotum, not developed bow-shaped, thick; sclerotized folds of oviporus short, posterior to scapular setae, postero-lateral extensions not extending beyond epimerites IIIa (fig. 3b). present or absent, setae se on these extension or on The genus currently includes 15 species arranged striated tegument. Setae c2 setiform, short. Setae c3 into three species groups (angolensis, glossifer, and commonly lanceolate. Setae dp2 of palpae simple, plocepasseri). hair-like. Setae ba of tarsi I, II as small spine, rarely
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Key to Metapteronyssus species margin of opisthosoma between setae h2 notice- ably convex (figs. 4 a, b, e) ...... 11 Males 9. Epiandrium tips extending to level of genital arch 1. Posterior margin of opisthosoma with slightly apex (fig. 5a). Setae e1 slightly posterior to open- convex opisthosomal membrane between setae ings gl. Length of idiosoma more than 340 ...... ps2 (fig. 18d) ...... M. lonchurae ...... M. bubalornis – Posterior margin of opisthosoma with a pair of – Epiandrium tips short, not extending to level of small lobules between setae h3 or with large bilo- genital arch apex (figs. 2a, 7 b). Setae e1 anterior bate membrane between setae ps2 (figs. 2a, 15a, to openings gl. Length of idiosoma less than 300 16a, 18a) ...... 2 ...... 10 2. Posterior margin of opisthosoma with large bilo- 10. Setae c3 lanceolate, 24-27 in length (fig. 2c) ...... bate membrane between setae ps2 (figs. 18a, b) ...... M. glossifer ...... M. angolensis – Setae c3 with setiform apex, 32-35 in length – Posterior margin of opisthosoma with a pair of (fig. 7c) ...... M. hordacei sp.n. small lobules between setae h3, margins of these 11. Prodorsal shield with rounded posterior end, lobules membranous (figs. 2a, 15a, 16a) ...... 3 without extending posterior angles, scapular setae 3. opisthosomal lobules finger-shaped, 2.5-3 times si, se on striated tegument. Adanal shield repre- longer than wide (figs. 16a, b) ...... sented by single median sclerite (fig. 9d) ...... M. amadinae sp.n...... M. plocei – opisthosomal lobules short, tongue-like, approx- – Prodorsal shield with extending posterior angles imately as long as wide (figs. 2b, 4b, 15b) ...... 4 encompassing bases of scapular setae si, se. Adanal 4. Postero-lateral margins of opisthosoma strongly shield consisting of three fragments (figs. 4a, 5d) convex (figs. 11d, 13a, d) ...... 5 ...... 12 – opisthosoma moderately or strongly attenuate 12. Transventral sclerite thin, about 4-6 along mid- posteriorly (figs. 2a, 4a) ...... 8 line of the mite body, tips of epiandrium scarcely 5. adanal shield T-shaped or cross-like (fig. 13d). expressed (fig. 5d). Length of idiosoma 330-360 Genital setae g at base of genital apparatus or pos- ...... M. capensis terior to it ...... 6 – transventral sclerite about 8-10 along midline, – adanal shield as transverse band with very thin with short epiandrium tips on posterior mar- median part (fig. 13a). Genital setae g approxi- gin (figs. 4a, 9a, 11a). Length of idiosoma less mately at midlevel of genital apparatus ...... 7 than 330 ...... 13 6. Setae s and f of tarsus III setiform. Tips of epian- 13. Genital arch 22-26 in length. Lateral fragments drium short blunt-angular. Seta ba of tarsi I, II of adanal shield with acute posterior ends, almost present. Subapical membranes between setae ps2 extending to adanal membranes (fig. 11a) ...... and h2 absent (figs. 13d-f) ...... M. plocepasseri ...... M. puylaerti – Setae s and f of tarsus III with lanceolate apical – Genital arch 16-18 in length. Lateral fragments part. Tips of epiandrium acute. Seta ba of tarsi I, of adanal shield of irregular form, not extending II absent. Subapical membrane between setae h2 to anterior end of adanal membranes ...... 14 and h3 present (figs. 15a-e) ...... 14. Setae c3 narrowly lanceolate, about 24-26 in ������������������������������������������� M. pseudonigritae sp. n. length (fig. 9b) ...... M. gaudi 7. Length of idiosoma 295-340. Anterior end of su- – Setae c3 with setiform apical part, over 30 in pranal concavity extends to the level of setae ps3. length (fig. 4c) ...... M. anaplecti sp. n. Setae c3 18-20 in length (figs. 11d-f) ...... M. anoplonotus Females – Length of idiosoma more than 350. Anterior end 1. Central hysteronotal sclerite absent (fig. 12b) ..... of supranal concavity not extending to the level of ����������������������������������������������� M. anoplonotus sp. n. setae ps3. Setae c3 longer than 20 (figs. 13a-c) .... – Central hysteronotal sclerite present, represented ...... M. daberti by one or two pieces (figs. 3a, 6a-c, 12a) ...... 2 8. adanal shield cross-like or T-shaped. Setae 3a 2. Central hysteronotal sclerite fused with pygidial situated on transventral sclerite. Tarsus III with sclerites forming a fish-shaped shield (fig. 10a) ... straight apex. Posterior margin of opisthosoma ...... M. gaudi sp. n. between setae h2 blunt (figs. 2a, b, f) ...... 9 – Central hysteronotal sclerites separated from – adanal shield split into median and pair of lateral paired pygidial sclerites (figs. 3a, 8a, b, 10b) ... 3 fragments, or represented only by single median 3. Central hysteronotal sclerite as longitudinal oval. sclerite. Setae 3a situated posterior to transven- Lateral sclerites as narrow longitudinal bands tral sclerite. Apex of tarsus III curved. Posterior (figs. 14a, b) ...... 4
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– Central hysteronotal sclerite represented by entire (fig. 19b) ...... M. lonchurae median band greatly variable in width (figs. 3a, – Width of anterior end of central sclerite 25- 8b, 17a) or split into anterior and posterior frag- 30. External copulatory tube is a short cone ments (figs. 6b, 12a). Lateral sclerites of another (fig. 19a) ...... M. angolensis form ...... 5 11. Anterior end of central hysteronotal shield not 4. Posterior margin of opisthosoma with extending reaching setae d1, these setae situated on stri- terminal region bearing two short lobes. External ated tegument (fig. 8a). External copulatory tube copulatory tube present, finger-shaped. Small ad- finger-like, 34-42 in length ...... M. capensis ditional sclerites of ovate form near bases of setae – anterior end of central hysteronotal shield en- d2 present (fig. 14a) ...... M. daberti compassing bases of setae d1 (fig. 9b). External – Posterior margin of opisthosoma bluntly round- copulatory tube conical, shorter than 25 ...... 12 ed, without terminal extension and lobes. Exter- 12. Central hysteronotal sclerite almost rod-shaped, nal copulatory tube absent. Additional sclerites width of anterior end 14-20; setae e1 commonly near lateral margins of central sclerites absent off this sclerite (fig. 8b) ...... M. hordacei sp.n. (fig. 14b) ...... M. plocepasseri – Central hysteronotal sclerite as longitudinal bar, 5. anterior end of central hysteronotal sclerite ex- width of anterior end over 30; setae e1 on margin tending to the level of setae c2; its width more of this sclerite (figs. 3a, 10b) ...... 13 than one third of greatest idiosoma width. Lat- 13. Posterior margin of prodorsal shield extending eral sclerites ovate or kidney-shaped (fig. 17a) .... beyond the level of scapular setae; anterior end ...... M. pseudonigritae sp. n. of central hysteronotal sclerite extending beyond – Central hysteronotal sclerite (or its anterior frag- the level of setae d1. Setae e1 approximately at the ment, if this sclerite is split into pieces) not ex- level of openings gl (fig. 3a) ...... M. glossifer tending to the level of setae c2; width of its an- – Posterior margin of prodorsal shield not extend- terior end less than one third of idiosoma width. ing beyond the level of scapular setae; anterior Lateral sclerites as longitudinal plate of irregular end of central sclerite not extending beyond the form (figs. 3a, 6a, b, 8a, b) ...... 6 level of setae d1. Setae e1 anterior to the level of 6. Central hysteronotal sclerite entire, represented openings gl (fig. 10b) ...... M. plocei by entire longitudinal sclerite, anterior end of 14. Central hysteronotal shield represented by which extends to the level of trochanters III two pieces, anterior and posterior fragments (figs. 3a, 5a, 6c, 8a) ...... 7 (fig. 11a) ...... 15 – Central hysteronotal sclerite split into anterior – Central hysteronotal sclerite represented by sin- and posterior fragments separated by large area gle fragment in posterior third of hysterosoma of striated tegument (figs. 6b, 12a), or only pos- (fig. 6a) ...... M. anaplecti sp. n. (part) terior fragment is developed in posterior third of 15. Anterior fragment of central hysteronotal sclerite hysterosoma (fig. 6a) ...... 14 roughly rectangular. Sclerotization around setae 7. External copulatory tube as short apically round- se, si poorly developed (fig. 6b) ...... ed cone or cylinder, 5-10 in length (figs. 6c, 17b) ...... M. anaplecti sp. n. (part) ...... 8 – anterior fragment of hysteronotal sclerite as ir- – External copulatory tube elongated, cone- or regular polygon. Posterior angles of prodorsal finger-shaped, 12-45 in length (figs. 8a, b) ..... 11 shield encompassing bases of setae se, si well scle- 8. Central hysteronotal sclerite almost parallel sided, rotized (fig.11a) ...... M. puylaerti its posterior end with narrow and deep incision (figs. 6c, 17b) ...... 9 Group glossifer – Central hysteronotal sclerite strongly narrowed to posterior end, without incision in posterior end Diagnosis (figs. 19a, b) ...... 10 Male. – Opisthosoma monotonously attenuate to 9. Posterior margin of prodorsal shield rounded. posterior end, its margin between setae ps2 blunt or Setae e1 mesal from margins of central sclerite convex (figs. 2a, 4a); posterior margin of part between (fig. 17b). Seta ba of tarsi I, II absent. Length of setae h3 with pair of small terminal lobules having idiosoma less than 400 ...... M. amadinae sp.n. membranous margins; opisthosomal margins be- – Posterior margin of prodorsal shield as short blunt tween bases of setae h3 and h2 with small subterminal angle. Setae e1 on lateral margins of central scle- membranes. Adanal shield varies in form from being rite (fig. 6c). Seta ba of tarsi I, II present. Length T-shaped, split into 3 pieces (median and a pair of of idiosoma 420-460 ...... M. bubalornis lateral fragments), or represented by median fragment 10. Width of anterior end of central sclerite 45-50. Ex- only. Tarsus III curved or straight. ternal copulatory tube button-like, hemispherical Female. – Central sclerite commonly represented
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a b
se 1a si
c1
d1 c2 cp cp 3a 3b c3 c3 d2 g e1 4a gl
ps3 e2
f 2 ps2 ps2
ps1 m μ h3 h3 100
h2 h2
Fig. 1. Metapteronyssus glossifer, male. – a, dorsal view; b, ventral view.
by entire median longitudinal band, width of which Metapteronyssus glossifer: Faccini & Atyeo 1981: 36 (part.); is less than one third of idiosoma width (figs. 3a, 8a, Mironov, 2002: 197 (part.) b); rarely (M. anaplecti, M. puylaerti) central sclerite split into anterior and posterior fragments (figs. 6b, Material examined. – central african republic: male 12a). Lateral shields as longitudinal plate of various lectotype (here designated), 5 male and 1 female paralec- totypes ex Euplectes franciscanus (Isert, 1789) (Ploceidae), shape, commonly with narrowed or acute posterior Bougouni, October 1950, coll. unknown (mrac 179 994); end. 9 male and 1 female paralectotypes, same data (mrac 179 991). 1. Metapteronyssus glossifer (Gaud) (figs. 1a, b, 2a-g, 3a, b) Diagnosis Pteronyssus glossifer Gaud, 1953: 220, fig. 11. Setae ba of tarsi I, II present (figs. 2d, e). In males, Pteronyssoides glossifer: Gaud & Mouchet 1959: 510; Gaud opisthosoma with blunt posterior end, setae ps2 lat & Till 1961: 274. eral to setae h2; terminal lobules tongue-like, terminal
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a b ts e2 3b 3a
g c3
4a ps1 ad f2 ps3 sm ps2 tm lb h3 h2 am m μ
ps2 c f 100
h3 r w m μ h2 cp 50 e d f s
c3 d ω1 e ω3 ω1 g kT ω3 f f r d ba d ba w la e e e d
e s e s vm φ f wa ra wa ra
Fig. 2. Metapteronyssus glossifer, male. – a, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, humeral shield and setae c3, ventral view; d, tarsus I, dorsal view; e, tarsus I, ventral view; f, tarsus III, dorsal view; g, tarsus IV, dorsal view. ad - adanal shield, am - adanal membrane, lb - lobules, sm - subterminal membrane, tm - terminal membrane, ts - transventral sclerite, vm - ventral membrane of tarsus I.
membranes poorly sclerotized, not clearly demarked Description from basal part of lobules; adanal shield T-shaped; Male (lectotype). – Idiosoma lengthwidth transventral sclerite narrow, with slightly expressed 310177 (idiosomal size in 14 paralectotypes epiandrium tips, setae 3a off transventral sclerite; se- 290-315165-180). Length of hysterosoma 172. tae c3 narrowly lanceolate in form, relatively short, Prodorsal shield: size 7856, without posterolateral 24-27 in length; tarsus III with curved apex (figs. 2a- extension, setae se separated by 67. Setae c3 narrowly c, f). In females, central sclerite as longitudinal band lanceolate, 24 in length. Hysteronotal shield: anterior extending to the level of trochanters III, greatest width margin slightly convex in median part, anterior an- 30-40, posterior end with deep and narrow incision; gles greatly extended laterally, length along midline setae d1, e1 on central sclerite; external copulatory 158, width at anterior margin 120. Distance between tube cone-shaped, short, 15-18 long (figs. 3a, b). prodorsal and hysteronotal shield along midline
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a b m μ 200
se 1a si c1 c2 3a cp 3b cp c3 g c3 d1 4a d2 cs e1 gl ls e2
py f 2 f 2 ct ps1 ps2 ps3 ps2
h2 h2
h3 h3
Fig. 3. Metapteronyssus glossifer, female. – a, dorsal view; b, ventral view. ct - external copulatory tube, cs - central sclerite, ls - lateral sclerite, py - pygidial sclerite. about 70. Opisthosoma with blunt posterior end, ps3 27, ps3:h3 34. Setae ba of tarsi I, II small spini- setae ps2 lateral to setae h2. Terminal lobules small form. Tarsus III 43 in length, with straight apex; tarsal tongue-like, terminal membranes not clearly de- setae f, w setiform, setae s slightly enlarged in medial marked from the lobules, length of incision between part; setae r slightly longer than segment. them 8; margins of opisthosoma between setae h2 Female (paralectotype). – Idiosoma lengthwidth and h3 with short subterminal membranes. Dorsal 383182 (in other paratype 380185). Length of measurements: c2:d2 52, d2:e2 58, d2:gl 27, e1:gl 5-7, hysterosoma 248. Prodorsal shield as in the male, e2:h3 43, e2:e2 51, h2:h2 32. Transventral sclerite 8 8460, setae se separated by 73. Setae c3 narrowly along midline; tips of epiandrium short, not extend- lanceolate, 20 in length. Arrangement of hysteronotal ing to genital apparatus apex; genital arch 1613; shields: central sclerite, a pair of lateral sclerites and setae g almost at midlevel of genital apparatus pair of pygidial sclerites. Central sclerite as longitu- (figs. 1b, 2a). Adanal shield T-shaped. Diameter of dinal bar (fig. 3a) extending by anterior end to the anal discs 11. Ventral measurements: ps2:ps2 48, ps3: level of trochanters III, length 176, greatest width
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a b
c
e d m m μ μ 50 100
Fig. 4. Metapteronyssus anaplecti, male. – a, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, setae c3; d, tarsus I, ventral view; e, tarsus III, dorsal view.
30 (in other paratype 17040). Lateral sclerites with fig. 11) are slightly schematic, they correspond well acute anterior end and narrowed posterior end, 67-72 to the specimens from the type host, from which we in length, with opening gl on inner margin. Pygidial select a lectotype here to firmly establish its identity. sclerites small triangular, encompassing bases of se- In the generic revisions of pteronyssids, Faccini and tae h2, h3, ps1. Setae d1 on lateral margins of central Atyeo (1981) pointed out that they re-examined only sclerite; setae d2, e1, e2 on striated tegument; setae the material of Gaud from both Euplectes species. e1 slightly posterior to level of openings gl. External Nevertheless, these authors apparently used mate- copulatory tube a cone-shaped process slightly curved rial from other hosts as well, because the drawings of downward, 16 in length, 12 in width at base. Dor- ‘Metapteronyssus glossifer’ given in the revision (Faccini sal measurements: c2:d2 94, d2:e2 95, d2:gl 52, e1: & Atyeo 1981: figs. 17-20), correspond neither to the gl 8-19, e2:h3 50, e2:e2 55, h2:h2 88, h3:h3 72. specimens from the type host, E. franciscanus, nor to Epigynium semicircular, 3275. ones from E. hordaceus (specimens from the latter host we consider a new species and describe below). Based Remarks on several clearly visible features in the figures given According to the original description by Gaud by Faccini and Atyeo, namely, the shape of prodorsal (1953), this species was recorded from Euplectes shield in both sexes, structure of terminal membrane franciscanus (type host), E. hordaceus Linnaeus, 1758, in the male, and the shape of hysteronotal sclerites Ploceus cucullatus (Statius Müller, 1776) (Ploceidae), and length of external copulatory tube in the female, and Uraeginthus bengalus (Linnaeus, 1776) (Estrildi- we suggest that these authors used for drawing the dae). This wide host range suggests that the author specimens from U. bengalus, i.e. they actually figured apparently dealt with a complex of closely related M. angolensis (see below, angolensis species group). species. Although original drawings (Gaud 1953:
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a d m μ 100
b
c
e f m μ 50
Fig. 5. Metapteronyssus males. – a, Metapteronyssis bubalornis, ventral view of hysterosoma; b, setae c3; c, tarsus III, dorsal view; d, M. capensis, ventral view of hysterosoma; e, setae c3; f, tarsus III, dorsal view.
2. Metapteronyssus anaplecti sp. n. Diagnosis (figs. 4a-e, 6a, b) Setae ba of tarsi I, II present. In males, opistho- soma with extending convex posterior end, setae ps2 Type material. – rwanda: male holotype and 4 anterior to setae h2; terminal lobules short, terminal male, 11 female paratypes ex Anaplectes rubriceps membranes semicircular, clearly demarked from lob- (Sundevall, 1850) (Ploceidae), Akanyaru, 28 October ules; adanal shield split into median and pair of lateral 1955, A. Fain (irsnb). fragments; transventral sclerite narrow, with slightly
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a c m μ 200
b
Fig. 6. Metapteronyssus females, dorsal view. – a, Metapteronyssus anaplecti, idiosoma; b, same, opisthosoma; c, M. bubalornis, idiosoma. expressed epiandrium tips, setae 3a off transventral short incision; setae d1 on anterior fragment (if it sclerite; setae c3 narrowly lanceolate in basal part, is present), setae e1 on posterior fragment; external with setiform apex, 33-38 in length; tarsus III with copulatory tube cone-shaped, long, 28-32 in length curved apex (figs. 4a-e). In females, central sclerite (figs. 6a, e). represented either by two fragments (anterior frag- ment of square form at the level of trochanters III Description and posterior one in posterior part of opisthosoma) Male (holotype). – Idiosoma lengthwidth or only by the posterior fragment, greatest width of 305177 (in 4 paratypes 300-310160-190). the posterior fragment 35-45, its posterior end with Length of hysterosoma 172. Prodorsal shield: without
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3. Metapteronyssus bubalornis Mironov posterolateral extensions (in some specimens they are (figs 5a-c, 6c) poorly expressed), size of proper shield 6445, setae se separated by 54. Setae c3 lanceolately enlarged in Metapteronyssus bubalornis Mironov, 2002: 188, fig. 12-17. basal part, 37 in length. Hysteronotal shield: anterior margin slightly convex in median part, anterior an- Material examined. – cameroon: male holotype and 2 gles acute, greatly extended, size 164127. Distance female paratypes ex Bubalornis albirostris (Vieillot, 1817) (Ploceidae), Maroua, 1949, coll. unknown (mrac 180601); between prodorsal and hysteronotal shields about 65. 2 male, 4 female paratypes, same data (mrac 180602 Opisthosoma with convex posterior end, setae ps2 an- – 180604). terior to setae h2. Most terminal part of opisthosoma between setae h3 with pair of small terminal lobules; terminal membranes semicircular, clearly demarked Diagnosis from lobules; margins of opisthosoma between se- Setae ba of tarsi I, II present. In males, opisthosoma tae h2 and h3 with subterminal membranes. Dorsal with blunt posterior end; setae ps2 lateral to setae h2; measurements: c2:d2 49, d2:e2 75, d2:gl 35,gl:e112, terminal lobules tongue-like, terminal membranes e2:h3 36, e2:e2 49, h2:h2 33. Transventral sclerite short, poorly sclerotized, not clearly demarked from 5 in length; tips of epiandrium short, not extending lobules; adanal shield T-shaped; transventral sclerite to genital apparatus apex; genital arch 1611; setae wide, with short epiandrium tips not extending to g slightly posterior to genital arch base. Adanal shield genital arch apex, setae 3a on transventral sclerite; consisting of 3 fragments: median fragment and setae c3 narrowly lanceolate, with thin apex, 27-30 pair of small lateral fragments of irregular form. Di- in length; tarsus III with straight apex (figs. 5a-c). In ameter of anal discs 10. Ventral measurements: ps2: females, central sclerite as longitudinal bar extend- ps2 40, ps3:ps3 22, ps3:h3 43. Setae ba of tarsi I, II ing to level of trochanters III, greatest width 25-32, small spiniform. Tarsus III 40 in length, with curved its posterior end with long narrow incision, setae d1, apex; tarsal setae f, w setiform, seta s slightly thick- e1 on margins of central sclerite; external copula- ened in basal part; seta r shorter than segment. tory tube button-shaped, short, about 7 in length Female (paratype). – Idiosoma lengthwidth (fig. 6c). 384197 (in other 10 paratypes 385-405195- 215). Length of hysterosoma 252. Prodorsal shield as Remarks in the male, 7756, setae se separated by 67. Setae This species is known only from the white-billed c3 narrowly lanceolate, 30 in length. Arrangement of buffalo weaver Bubalornis albirostris. hysteronotal shields: central sclerite, pair of opistho- somal sclerites and pair of pygidial sclerites. Central 4. Metapteronyssus capensis Mironov sclerite as narrow longitudinal bar in posterior half (figs. 5d-f, 8a) of hysterosoma (fig. 6a), length 78, greatest with 38 (in all other paratypes 75-8035-42; in 3 paratypes, Metapteronyssus capensis Mironov, 2002: 187, fig. 16. central sclerite represented also by additional anterior fragment of approximately square form, 52-5556- Material examined. – south africa: male holotype and 60, situated at level of trochanters III, IV, – fig. 6b). 1 male, 5 female paratypes ex Ploceus capensis (Linnaeus, 1766) (Ploceidae), Cape Province, East London, October Lateral sclerites as longitudinal plates slightly attenu- 1966, coll. unknown (mrac 180635); 3 males, 7 female ated to posterior end, which is curved to ventral side paratypes, same data (mrac 180618). of the body 64-66 in length; openings gl in anterior end. Pygidial sclerites small triangular, encompassing bases of setae h2, h3, ps1. Setae d1 on lateral margins Diagnosis of central sclerite; setae d2, e1, e2 on striated tegu- Setae ba of tarsi I, II present. In males, opisthosoma ment (setae d1 in anterior fragment of central sclerite, with convex posterior end, setae ps2 anterior to setae when it present); setae e1 slightly anterior to level of h2; terminal lobules short, terminal membranes semi- openings gl. External copulatory tube a cone-shaped circular, clearly demarked from lobules; adanal shield process slightly curved downward, 30 in length, 20 in split into median sclerite and pair of lateral fragments; width at base. Dorsal measurements: c2:d2 83, d2:e2 transventral sclerite narrow, without epiandrium tips; 113, d2:gl 73, e1:gl 14-17, e2:h3 45, e2:e2 57, h2:h2 setae 3a off transventral sclerite; setae c3 narrowly lan- 80, h3:h3 70. Epigynium bow-shaped, 2978. ceolate, 30-32 in length; tarsus III with curved apex (figs. 5d-f). In females, central sclerite as longitudi- Etymology nal bar extending to level of trochanters IV, greatest The specific epithet derives from the generic name width 30-35, its posterior end with long narrow inci- of the type host and is a noun in the genitive case. sion, setae e1 on margins of central sclerite, d2 off this
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a b
c d m μ m μ 100 50
Fig. 7. Metapteronyssus hordacei, male. – a, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, setae c3; d, tarsus III, dorsal view. sclerite; external copulatory tube cone-shaped, long, Diagnosis 34-42 in length (fig. 8a). Setae ba of tarsi I, II present. In males, opisthosoma with convex posterior end, setae ps2 anterior to setae Remarks h2, terminal lobules short, terminal membranes semi- This species is known only from the type host, cape circular, clearly demarked from lobules; adanal shield weaver Ploceus capensis, in South Africa (table 3). split into median sclerite and pair of lateral fragments; transventral sclerite narrow, with short epiandrium tips; setae 3a off transventral sclerite; setae c3 narrow- 5. Metapteronyssus gaudi Mironov ly lanceolate, 24-26 in length; tarsus III with curved (figs. 9a-c, 10a) apex (figs. 9a-c). In females, central sclerite fused with Metapteronyssus gaudi Mironov, 2002: 185, fig. 8, 10. pygidial shields forming fish-shaped shield, anterior end of this shield extending to level of trochanters III, Material examined. – nigeria: Male holotype and greatest width 55-60, setae d1, e1 on central sclerite, 1 female paratype ex Ploceus aurantius aurantius (Vieillot, external copulatory tube cone-shaped, long, 28-30 in 1805) (Ploceidae), S. Nigeria, Degema, 14 January 1902, W.J. Ansorge (mrac 180 639). – cameroon: 2 males, length (fig. 10a). 2 females ex P. bicolor amaurocephalus (Vieillot, 1819), S. Cameroon, November 1955, coll. unknown (mrac Remarks 180 636, 180 637); 4 males, 5 females ex P. nigerrimus Metapteronyssus gaudi is known from four species of (Vieillot, 1819), S. Cameroon, August 1955, coll. unknown weavers of the genus Ploceus Cuvier, 1816 in west and (mrac 180 631, 180 632); 2 females, same host, Yaoundé, north of Sub-Saharan Africa (table 3). This species July 1955, coll. unknown (mrac 180 633); 2 males, differs from all species of the genus 2 females ex P. nigricolliis brachypterus (Swainson, 1834), Metapteronyssus S. Cameroon, August 1955, coll. unknown (mrac by the unique shape of the central sclerite in females 180 630). (fig. 10a).
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a b m μ m μ 200 200
Fig. 8. Metapteronyssus females, dorsal view of idiosoma. – a, Metapteronyssus capensis; b, M. hordacei.
6. Metapteronyssus hordacei sp. n. epiandrium tips; setae 3a off transventral sclerite; se- (figs. 7a-c, 8b) tae c3 setiform, slightly thickened in basal part, 32- 35 in length; tarsus III with curved apex (figs. 7a-c). Type material. – central african republic: male In females, central sclerite as very narrow median holotype and 2 male, 3 female paratypes ex Euplectes bar extending to the level of trochanters III, greatest hordaceus (Linnaeus, 1758) (Ploceidae), Bossangoa, width 14-18; setae d1, e1 on central sclerite; external July 1951, coll. unknown (mrac 179 896); 3 male, copulatory tube cone-shaped, short, 13-15 in length 4 female paratypes, same data (mrac 179 996). Holo- (fig. 8b). type and all paratypes – mrac. Description. Diagnosis Male (holotype). – Idiosoma lengthwidth Setae ba of tarsi I, II present. In males, opisthosoma 294164 (in 5 paratypes 270-295144-158). with blunt posterior end, setae ps2 lateral to setae h2, Length of hysterosoma 179. Prodorsal shield: size terminal lobules tongue-like, terminal membrane 7049, without posterolateral extensions, setae not clearly demarked from lobules; adanal shield se separated by 56. Setae c3 setiform, slightly thick- T-shaped; transventral sclerite narrow, with short ened in basal part, 33 in length, 1.5 in width.
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a d m μ 100
c b f e m μ 50
Fig. 9. Metapteronyssus males. – a, M. gaudi, ventral view of hysterosoma; b, setae c3; c, tarsus III, dorsal view; d, M. plocei, ventral view of hysterosoma; e, setae c3; f, tarsus III, dorsal view.
Hysteronotal shield: anterior margin slightly convex of epiandrium short, not extending to genital ap- in median part, anterior angles extended, relatively paratus apex; genital arch 169; setae g at base of short, size 16295. Distance between prodorsal genital apparatus. Adanal shield T-shaped, with very and hysteronotal shields about 70. Opisthosoma weakly sclerotized lateral parts. Diameter of anal discs with blunt posterior end, setae ps2 lateral to setae h2. 11. Ventral measurements: ps2:ps2 40, ps3:ps3 25, ps3: Terminal lobules small tongue-like, terminal mem- h3 38. Setae ba of tarsi I, II small spiniform. Tarsus brane not clearly demarked from lobules, length of III 38 in length, with straight apex; tarsal setae f, w incision between membranes 8. Margins of opistho- setiform, setae s slightly thickened in basal part; setae soma between setae h2 and h3 with short subtermi- r longer than segment. nal membranes. Dorsal measurements: c2:d2 50, d2: Female (paratype). – Idiosoma lengthwidth e2 62, d2:gl 35, e1:gl 3-9, e2:h3 45, e2:e2 49, h2:h2 366164 (in 6 paratypes 355-375150-175). 30. Transventral sclerite narrow, 6 along midline; tips Length of hysterosoma 250. Prodorsal shield as in
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a b m μ 200
Fig. 10. Metapteronyssus females, dorsal view of idiosoma. – a, Metapteronyssus gaudi; b, M. plocei. the male, 7653, setae se separated by 65. Setae measurements: c2:d2 83, d2:e2 112, d2:gl 60, c3 narrowly lanceolate, 24 in length. Arrangement gl:e1 10-12, e2:h3 47, e2:e2 56, h2:h2 59, h3:h3 51. of hysteronotal shields: central sclerite, and pair Epigynium semicircular, 4072. of opisthosomal sclerites and pair of pygidial scler- ites. Central sclerite as very narrow median sclerite Etymology (fig. 8b) extending by anterior end to level of tro- The specific epithet derives from the species name chanters III, length 180, greatest width 18 (in other of the type host and is a noun in the genitive case. paratypes 170-18014-18). Lateral sclerites with acute anterior end and narrowed posterior end, with 7. Metapteronyssus plocei Mironov opening gl on inner margin, 92-94 in length. Pygidial (figs. 9d-f, 10b) sclerites small triangular, encompassing bases of se- tae h2, h3, ps1. Setae d1 on lateral margins of central Metapteronyssus plocei Mironov, 2002: 182, fig. 1-7. sclerite; setae d2, e1, e2 on striated tegument; setae e1 slightly posterior to the level of openings gl. Ex- Material examined. – central african republic: ternal copulatory tube cone-shaped, slightly curved male holotype and 3 male, 1 female paratypes ex Ploceus downward, 13 in length, 12 in width at base. Dorsal cucullatus (Statius Müller, 1776) (Ploceidae), Oubangai,
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a d m μ 100
b c e f m μ 50
Fig. 11. Metapteronyssus males. – a, Metapteronyssus puylaerti, ventral view of hysterosoma; b, setae c3; c, tarsus III, dorsal view; d, M. anoplonotus, ventral view of hysterosoma; e, setae c3; f, tarsus III, dorsal view.
July 1951, coll. unknown (mrac 180 622); 5 male, 2 female Diagnosis paratypes, same data (mrac 180 621); 3 male paratypes, Setae ba of tarsi I, II present. In males, opisthosoma same host, Bossangoa, July 1951, coll. unknown (mrac with convex posterior end; setae ps2 antero-lateral to 180 634); 15 male, 13 female paratypes, same host, Bamako, October 1950, coll. unknown (mrac 180 623 – 180 625); setae h2; terminal lobules short; terminal membranes 14 males, 8 females ex P. nigricolliis brachypterus (Swain- semicircular, clearly demarked from lobules; adanal son, 1834), Bobodiolasso, October 1950, coll. unknown shield represented by longitudinal median plate, (mrac 180 626 – 180 629). – south africa: 1 female ex transventral sclerite narrow, with very short epian- P. xanthops (Hartlaub, 1862), Transvaal, Haenertsburg, drium tips, setae 3a off transventral sclerite; setae c3 26-27 November 1961, S.A.I.M.R. (mrac 180 638). lanceolate, with thin apex, 30-38 in length; tarsus III with curved apex (figs. 9d-f). In females, central
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a b m μ 200
Fig. 12. Metapteronyssus females, dorsal view of idiosoma. – a, Metapteronyssus puylaerti; b, M. anoplonotus. sclerite as longitudinal bar extending to the level of Diagnosis trochanters III, greatest width 45-52, posterior end of Setae ba of tarsi I, II present. In males, opisthosoma this sclerite with short incision; setae d1, e1 on cen- with convex posterior end; setae ps2 anterior to setae tral sclerite; external copulatory tube cone-shaped, h2; terminal lobules short, terminal membranes clear- medium-sized, 15-18 in length (fig. 10b). ly demarked from lobules; adanal shield represented by median part and a pair of lateral fragments; trans- Remarks ventral sclerite narrow, with short epiandrium tips, se- This species is known from three weaver species tae 3a off transventral sclerite; setae c3 lanceolate with of the genus Ploceus in central and southern Africa thin apex, 30-35 in length; tarsus III with curved apex (table 3). (figs. 11a-c). In females, central sclerite split into two fragments (anterior piece of irregular form situated at the level of trochanters III, IV and longitudinal plate 8. Metapteronyssus puylaerti Mironov in posterior third of opisthosoma), greatest width of (figs. 11a-c, 12a) posterior fragment 55-60, its posterior end with short Metapteronyssus puylaerti Mironov, 2002: 185, fig. 9, 11. incision, setae d1 and e1 on anterior and posterior fragments of the central sclerite, respectively; exter- Material examined. – south africa: male holotype and nal copulatory tube cone-shaped, elongated, 34-42 in 3 male, 3 female paratypes ex Ploceus velatus (Vieillot, 1819) length (fig. 12a). (Ploceidae), Transvaal, Potchefstroom, 26 December 1952, S.A.I.M.R. (mrac 180 045), 4 male, 5 female paratypes, same data (mrac 180 042, 180 046).
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Remarks (Ploceidae). Taking into consideration that all other This species is known only from the type host, members of the plocepasseri group are associated southern masked-weaver Ploceus velatus, in South exclusively with birds of the family Passeridae, we Africa (table 3). do not exclude that the record of this species on Q. quelea could be the result of contamination. Group plocepasseri 10. Metapteronyssus daberti Mironov Diagnosis (figs. 13a-c, 14a) Male. – Opisthosoma with strongly convex pos- tero-lateral margin and with trapezoidal median ex- Metapteronyssus daberti Mironov, 2002: 191, fig. 15, 19. tension between setae ps2, the terminal part of this extension between setae h3 with pair of small termi- Material examined. – zambia: male holotype and fe- nal lobules bearing short membranes; subapical mem- male paratype ex Plocepasser rufoscapulatus Büttikofer, 1888 (Passeridae), North Rhodesia (sic), Luanshya, 22 March branes absent or present. Adanal shield T-shaped or 1953, E.L. Haydock (uga 3044 usnm 46038). Holotype represented by only a transverse bar-like plate. Tarsus and paratype are deposited in usnm. III with slightly curved apex. Female. – Central sclerite ovate, or represented by wide longitudinal bar-like plate extending to sejugal Diagnosis region (M. pseudonigritae), or absent (M. anoplonotus). Setae ba of tarsi I, II present. In males, opisthosoma Lateral shields band-like or kidney-shaped. with convex lateral margins and trapezoidal median extension having pair of short terminal lobules; setae ps2 anterior to setae h2, terminal membranes semicir- 9. Metapteronyssus anoplonotus Mironov cular, short, clearly demarked from lobules; supranal (figs. 11d-f, 12b) concavity not extending to the level of setae ps3; adan- Metapteronyssus anoplonotus Mironov, 2002: 189, fig. al shield represented by transverse plate; transventral 14, 18. sclerite with scarcely expressed epiandrium tips; se- tae 3a on transventral sclerite; setae c3 lanceolate, Material examined. – zimbabwe: 13 males, 15 females 24-28 in length, tarsus III with slightly curved apex ex Plocepasser mahali Smith, 1836 (Passeridae), South (figs. 13a-c). In females, central sclerite represented Rhodesia (sic), Kariba, February 1964, coll. unknown (mrac by one large, ovate shield extending to the level of tro- 180 615 – 180 619); 12 males, 15 females ex Quelea quelea (Linnaeus, 1758) (Ploceidae), South Rhodesia (sic), Kariba, chanters III, greatest width 80-85, and a pair of small February 1964, coll. unknown (mrac 180 609 – 180 614). additional sclerites situated near its lateral margins – south africa: 2 females ex P. mahali, Transvaal, Pieters- at the level of trochanters IV, setae d1, e1 on central burg, 10 December 1967, coll. unknown (mrac 180620). sclerite; copulatory tube cone-shaped, long, 40-46 (fig. 14a). Diagnosis Setae ba of tarsi I, II present. In males, opistho- Remarks soma with convex lateral margins and trapezium- Known only from the type host, red-backed spar- shaped median extension having pair of short termi- row weaver Plocepasser rufoscapulatus, in Zambia nal lobules; setae ps2 anterior to setae h2; terminal (table 3). membranes semicircular, short, clearly demarked from lobules; supranal concavity extending to the 11. Metapteronyssus plocepasseri Mironov level of setae ps3; adanal shield represented by trans- (figs. 13d-f, 14b) verse plate; transventral sclerite with epiandrium tips scarcely marked or indistinct; setae 3a on transventral Metapteronyssus plocepasseri Mironov, 2002: 194, fig. sclerite; setae c3 lanceolate, 28-30 in length; tarsus 13, 20. III with curved apex (figs. 11d-f). In females, central sclerite absent, setae d1, e1 on striated integument; Material examined. – zimbabwe: male holotype and 3 external copulatory tube cone-shaped, elongated, male paratypes ex Plocepasser mahali Smith, 1836, (Passeri- dae), South Rhodesia (sic), Chirundu, February 1964, coll. 26-30 in length (fig. 12b). unknown (mrac 179 999); 10 male, 4 female paratypes, same data (mrac 179 997, 179 998, 180 600). Remarks This species is known from two passeridan hosts of different families in South Africa (Mironov 2002): Diagnosis white-browed sparrow weaver, Plocepasser mahali Setae ba of tarsi I, II present. In males, opisthosoma (Passeridae), and red-billed quelea, Quelea quelea with convex lateral margins and trapezium-shaped
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a d m μ 100
b c e f m μ 50
Fig. 13. Metapteronyssis males. – a, Metapteronyssus daberti, ventral view of hysterosoma; b, setae c3; c, tarsus III, dorsal view; d, M. plocepasseri, ventral view of hysterosoma; e, setae c3; f, tarsus III, dorsal view. median extension having pair of short terminal setae d1 off central sclerite, e1 on this sclerite; external lobules; setae ps2 anterior to setae h2, terminal mem- copulatory tube absent (fig. 14b). branes semicircular, short, clearly demarked from lob- ules; supranal concavity not extending to the level of Remarks setae ps3; adanal shield T-shaped, transventral sclerite Metapteronyssus plocepasseri is known only from the with well-expressed epiandrium tips not extending to type host, white-browed sparrow weaver Plocepasser the level of genital arch apex; setae 3a on transventral mahali, in Zimbabwe. It is interesting to note that sclerite; setae c3 narrowly lanceolate, 18-20 in length; one more species, M. anoplonotus (see above), was tarsus III with slightly curved apex (figs. 13d-f). In fe- also recorded from this host in South Africa; however males, central sclerite narrowly ovate, scarcely extend- both mite species have not been recorded in samples ing to level of trochanters III, greatest width 58-65, from the same host specimens (Mironov 2002).
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a b m μ 200
Fig. 14. Metapteronyssus females, dorsal view of idiosoma. – a, Metapteronyssus daberti; b, M. plocepasseri.
12. Metapteronyssus pseudonigritae sp. n. tips of epiandrium thin and acute, not extending to (figs. 15a-e, 17a) the level of genital arch apex; setae 3a on transventral sclerite; setae c3 narrowly setiform, slightly thickened Type material. – kenya: male holotype and 1 male, in basal part, about 24 long; tarsus III with slightly 1 female paratypes ex Pseudonigrita cabanisi (Fischer et bent apex (figs. 15a-e). In females, central sclerite a Reichenov, 1884) (Passeridae), Waso, 12 June 1923, wide almost rectangular longitudinal plate extending K. Caldwell (uga 5914 amnh 210 302). Holotype, to the level of setae c2, posterior margin without inci- 1 male paratype – amnh; 1 male paratype – zisp. sion, greatest width about 60, setae d1, e1 on central sclerite; external copulatory tube small, cylindrical in Diagnosis form, about 9 long (fig. 17a). Setae ba of tarsi I, II absent. In males, opistho- soma with convex lateral margins and trapezium- Description shaped median extension with a pair of short Male (holotype). – Idiosoma lengthwidth terminal lobules; setae ps2 anterior to setae h2, 298172 (in paratype 300180). Length of hys- terminal membranes semicircular, not clearly de- terosoma 174. Prodorsal shield with posterolateral marked from lobules; subterminal membranes present; extensions, size 7049, setae se separated by 56. supranal concavity not extending to the level of setae Setae c3 setiform, slightly thickened in basal part, ps3; adanal shield T-shaped; transventral sclerite wide, 24 in length, 1.5 in width. Hysteronotal shield:
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a b
c m μ
100 d e m μ 50
Fig. 15. Metapteronyssus pseudonigritae, male. – a, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, seta c3; d, tarsus I, ventral view; e, tarsus III, dorsal view. anterior margin strongly convex in median part, size Diameter of anal discs 11. Ventral measurements: ps2: 16295. Distance between prodorsal and hyster- ps2 33, ps3:ps3 26, ps3:h3 40. Setae ba of tarsi I, II onotal shields about 50. Postero-lateral margins of absent. Tarsus III 41 in length, with slightly curved opisthosoma greatly convex; setae ps2 antero-lateral apex; tarsal setae f, w, s lanceolate in distal part; setae to setae h2, terminal region of opisthosoma between r longer than segment. setae ps2 with trapezoidal extensions having short Female (paratype). – Idiosoma lengthwidth lobules ending by terminal membranes; incision be- 374172. Length of hysterosoma 245. Prodorsal tween lobules 6 long; terminal membranes semicir- shield as in the male, 6473, setae se separated by cular, not clearly demarked from lobules (fig. 15b). 64. Setae c3 spiculiform, 18 in length, 1.5 in width. Opisthosomal margins between setae h2 and h3 with Arrangement of hysteronotal shields: central sclerite, semicircular subterminal membranes. Supranal con- pair of lateral sclerites and pair of pygidial sclerites. cavity not extending to the level of setae ps3. Dor- Central sclerite as wide longitudinal bar (fig. 17a) ex- sal measurements: c2:d2 54, d2:e2 65, d2:gl 30, e1:gl tending by anterior end to level of setae c2, length 7-24, e2:h3 32, e2:e2 46, h2:h2 29. Transventral scler- 194, greatest with 62. Lateral sclerites kidney-shaped, ite thick, 20 in length including median extension on 56-60 long, with opening gl on anterior half. Pygidial posterior margin; tips of epiandrium acute, not ex- sclerites small triangular, encompassing bases of setae tending to genital apparatus apex; genital arch 189; h2, h3, ps1. Setae d1 and e1 on central sclerite; se- setae g at base of genital arch. Adanal shield T-shaped. tae d2, e2 on striated tegument; setae e1 anterior to
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a b
c m μ 100 m μ 50
e d m μ 50
Fig. 16. Metapteronyssus amadinae, male. – a, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, seta c3; d, tarsus I, ventral view; e, tarsus III, dorsal view. level of openings gl. External copulatory tube a small opisthosoma and lobules (figs. 16a, b). Adanal shield cylindrical process, 9 in length, 5 in width. Dorsal T-shaped. Tarsus III straight, cone shaped. measurements: c2:d2 70, d2:e2 124, d2:gl 93, e1:gl Female. – Central sclerite as median bar-like plate 17-19, e2:h3 44, e2:e2 50, h2:h2 60, h3:h3 49. Epigy- extending to the level of trochanters III (figs. 17a, nium bow-shaped, 2265. 19a, b); lateral sclerites as longitudinal plates of ir- regular form. Etymology The specific epithet derives from the generic name 13. Metapteronyssus amadinae sp. n. of the type host and is a noun in a genitive case. (figs. 16a-e, 17b) Type material. – ethiopia: male holotype and Group angolensis 3 male, 2 female paratypes ex Amadina fasciata Diagnosis alexanderi Neumann, 1908 (Estrildidae), SW Male. – Opisthosoma moderately attenuate to Ethiopia, Indunumra Mountains, 15 July 1912, posterior end, posterior margin bluntly rounded or E.A. Mearns (uga 3055 usnm 247 105). Holotype, with pair of small finger-like lobules between setae h3 2 male and 1 female paratypes – usnm, 1 male, 1 fe- (M. amadinae). Posterior margin of opisthosoma male paratypes – zisp. bears entire opisthosomal membrane between bases of setae ps2 (with free margin of this membrane slightly Diagnosis convex or bilobate) (figs. 18a, b, f); if opisthosoma Setae ba of tarsi I, II absent. In males, opistho- with lobules, the opisthosomal membrane is repre- soma with short median extension between setae h3 sented by a narrow band along the entire margin of bearing pair of finger-shaped terminal lobules, setae
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a b m μ 200
Fig. 17. Metapteronyssus females, dorsal view of idiosoma. – a, M. pseudonigritae; b, M. amadinae. ps2 antero-lateral to setae h2, opisthosomal mem- passing scapular setae, setae se separated by 56. Setae brane along entire margin of opisthosoma and lob- c3 narrowly lanceolate, 22 in length. Hysteronotal ules narrow, 2-3 in width; adanal shield T-shaped, shield: anterior margin convex in median part, ante- transventral sclerite wide; tips of epiandrium thin rior angles greatly extended laterally, size 156127. and acute, not extending to the level of genital arch Distance between prodorsal and hysteronotal shields apex; setae 3a on transventral sclerite; setae c3 nar- about 65. Opisthosoma with rounded postero-lateral rowly lanceolate, 22-24 in length, tarsus III with angles; posterior margin of opisthosoma with median straight apex (figs. 16a-e). In females, central sclerite extension between setae h2 bearing a pair of finger- as longitudinal parallel-sided plate extending to the shaped lobules, about 258; setae ps2 antero-lateral level of trochanters III, posterior margin with deep to h2; opisthosomal membrane flanking posterior narrow incision, greatest width 40-43; setae d1, e1 on margin of opisthosoma and margin of lobules very central sclerite; external copulatory tube a small cone narrow, about 2-3 in width; incision between lobules rounded apically, 6-8 in length (fig. 17b). slit-shaped, 22 in length (figs. 16b). Dorsal measure- ments: c2:d2 56, d2:e2 64, d2:gl 28, e1:gl 7-12, e2:h3 Description 43, e2:e2 50, h2:h2 30. Transventral sclerite thick, 16 Male (holotype). – Idiosoma lengthwidth in length; tips of epiandrium not extending to genital 300184 (in 3 paratypes 290-300170-180). apparatus apex; genital arch 1411; setae g at base Length of hysterosoma 174. Prodorsal shield: size of genital arch. Adanal shield T-shaped. Diameter of 6866, without posterolateral extension encom- anal discs 13. Ventral measurements: ps2:ps2 46, ps3:
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ps3 24, ps3:h3 41. Setae ba of tarsi I, II absent. Tarsus (figs. 18a-e). In females, central sclerite as a narrow III cone-shaped, 38 in length, with straight apex; tar- bar slightly attenuated posteriorly, anterior end ex- sal setae f, w setiform, setae s slightly thickened, setae tending to the level of trochanters III, posterior mar- r subequal in length to the segment. gin without incision, greatest width 24-27; setae d1, Female (paratype). – Idiosoma lengthwidth e1 on margins of central sclerite; external copulatory 373174 (in other paratype 375180). Length tube small, cylindrical, 6-8 long (fig. 19a). of hysterosoma 245. Prodorsal shield as in the male, 7773, setae se separated by 67. Setae c3 narrowly Remarks lanceolate, 19 in length. Arrangement of hyster- This species was originally described from the onotal shields: central sclerite, pair of opisthosomal blue waxbill (or cordon bleu) Uraeginthus angolen- sclerites and pair of pygidial sclerites. Central scler- sis in South Africa (Mironov & Kopij 2000); in the ite as wide longitudinal band (fig. 17b) extending by course of the present study it was also recorded from anterior end to level of trochanters III, with narrow the red-cheeked blue waxbill U. bengalus ugogoen- incision in posterior end, length 168, greatest width sis in Rwanda. The figures of ‘M. glossifer’ given by 43 (in other paratype 16540). Lateral sclerites as Faccini & Atyeo (1981: figs. 17-20) in the generic longitudinal plates of irregular form, with opening revision of pteronyssids correspond well to specimens gl on their anterior ends, 70-72 in length. Pygidial of Metapteronyssus angolensis rather than to those of sclerites small triangular, encompassing bases of setae M. glossifer from its type host Euplectes franciscanus, h2, h3, ps1. Setae d1 and e1 on central sclerite; setae and we suggest that they were probably based on d2, e2 on striated tegument; setae e1 approximately the material of Gaud (1953) from U. bengalus from at level of openings gl. External copulatory tube a Sudan (see above, remarks to M. glossifer). very short cone rounded apically 8 in length, 10 in width at base. Dorsal measurements: c2:d2 76, d2:e2 15. Metapteronyssus lonchurae Mironov 107, d2:gl 56, e2:h3 49, e2:e2 49, h2:h2 72, h3:h3 53. (figs. 18f-h, 19b) Epigynium semicircular, 3370. Metapteronyssus lonchurae Mironov, 2002: 197, fig. 21, 22. Etymology The specific epithet derives from the generic name Material examined. – somali: male holotype from of the type host and is a noun in a genitive case. Lonchura cantans orientalis (Gmelin, 1789) (Estrildidae), Bikendula, 16 March 1919, coll. unknown (mrac 180 047); female paratype, same data (mrac 180 048). 14. Metapteronyssys angolensis Mironov et Kopij (figs. 18a-e, 19a) Diagnosis Metapteronyssys angolensis Mironov & Kopij, 2000: 100, Setae ba of tarsi I, II present. In males, opisthosoma figs. 1-3; Mironov 2002: 198, fig. 23-25. bluntly rounded; setae ps2 lateral to setae h2; opistho- [Metapteronyssus glossifer: sensu Faccini & Atyeo 1981 (par- tim): 36, fig. 17-20; Mironov 2002: 197, fig. 26, 27. somal membrane on posterior margin of opisthosoma Misidentifications] between bases of setae ps2 with slightly convex free margin; adanal shield T-shaped, transventral sclerite wide; tips of epiandrium acute, not extending to the Material examined. – south africa: Male holotype and level of genital arch apex; setae 3a on transventral 1 male paratype ex Uraeginthus angolensis (Linnaeus, 1758) sclerite; setae c3 lanceolate, about 24 long; tarsus III (Estrildidae), Northern Province, Kruger National Park, Pafuri, 10 January 1989, R.A. Earlé (nmb 00189). – rwan- with straight apex (figs. 18f-h). In females, central da: 4 males and 4 females ex U. bengalus ugogoensis Reiche- sclerite as a narrow bar attenuated posteriorly, ante- now, 1911, Astrida, 7 November 1955, A. Fain (irsnb). rior end extending to the level of trochanters III, pos- terior margin without incision, greatest width about 47; setae d1, e1 on margins of central sclerite; exter- Diagnosis nal copulatory tube small, hemispherical, 6 in length Setae ba of tarsi I, II present. In males, opisthosoma (fig. 19b). bluntly rounded; setae ps2 lateral to setae h2; opistho- somal membrane along its margin between setae ps2 Remarks large, with pair of wide and rounded extensions, This species is known only from the African sil- greatest length of membrane 14-18; adanal shield verbill Lonchura cantans orientalis in Somali (table T-shaped, transventral sclerite wide; tips of epian- 3). Metapteronyssus lonchurae clearly differs from drium acute, extending to the level of genital arch other known species of the genus by having an apex; setae 3a on transventral sclerite; setae c3 lan- opisthosomal membrane with a slightly convex free ceolate, 21-25 in length; tarsus III with straight margin in the males.
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a f m μ
om 100 om
b d g m μ 50
om
c e h
Fig. 18. Metapteronyssus males. – a, Metapteronyssis angolensis, ventral view of hysterosoma; b, dorsal view of opisthosoma; c, seta c3; d, tarsus I, ventral view; e, tarsus III, dorsal view; f, M. lonchurae, ventral view of hysterosoma; g, seta c3; h, tarsus III, dorsal view. om - opisthosomal membrane.
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a b m μ 200
Fig. 19. Metapteronyssus females, dorsal view of idiosoma. – a, M. angolensis; b, M. lonchurae.
Discussion ridge (7.1), and placement of hysteronotal gland openings gl at postero-lateral incisions in the hyster- Phylogeny onotal shield (9.1). Four synapomorphies are female The branch-and-bound search produced a single characters: setae e2 moved relatively close to midline shortest tree having length of 47 steps and standard of the body (23.1), development of central (24.1) and indices, excluding uninformative characters, as fol- lateral (30.1) shields due to splitting of entire hyster- lows: ci=0.8140, ri=0.9048, rc=0.7508 (fig. 20). onotal shield, and development of external copula- Monophyly of the genus Metapteronyssus is supported tory tube (33.1). by eleven unambiguous synapomorphies, four of The Metapteronyssus branch forms three clusters which are characters for both sexes (see table 1): loss corresponding to our taxonomic concept of three of prodorsal shield in posterior part of prodorsum species groups. The first cluster to split off corre- (character 1.1), fusion of epimerites I as a narrow U sponds to the plocepasseri group and is supported by (2.1), loss of unpaired vertical seta vi (3.1), reduction three synapomorphies representing structures of the of seta ba of tarsi I, II to small spine-like seta (4.1). opisthosoma in males: strongly convex posterolateral Three synapomorphies characterizing males represent margins (8.1) and median trapezium-shaped exten- features of the hysteronotal shield: development of a sion between setae h2 (10.1), and a pair of short ter- pair of lateral hysteronotal ridges (6.1) and median minal membranous lobules (13.1). The clade bearing
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Sturnotrogus subtruncatus Mite species Avian host Scutulanyssus dasyritidis groups families 20 30 Pteronyssoides holoplax 18 1 1 5 15 31 1 M. pseudonigritae 1 1 2 8 10 13 33 22 1 1 1 1 M. plocepasseri 1 25 31 0 24 1 1 1 1 19 34 M. anoplonotus* 1 2 asseridae
1 1 P plocepasseri 1 2 3 4 6 7 9 23 24 30 33 M. daberti 10 1 1 1 1 1 1 1 1 1 2 1 M.lonchurae 11 14 2 26 1 1 12 2 M. angolensis 1 5 1 Estrildidae angolensis 21 26 1 M. amadinae 1 1 1 M. bubalornis
13 15 M. glossifer 1 26 1 1 M. hordacei 16 34 2 1 1 1 M. plocei 29 M. capensis
17 20 21 1 glossifer 3 28 Ploceidae 1 1 0 1 M. gaudi
27 32 M. puylaerti 2 29 subgroup plocei 1 1 M. anaplecti 2
Fig. 20. Phylogeny of Metapteronyssus. Single most parsimonious tree. deltran character optimisation. Numbers above dots (black – unique apomorphy, white – homoplasy) refer to characters; numbers under dots refer to a character state achieved in the respective node. Numbers in italics to the right from nodes are Bremer indices. (*) Species is also recorded from Ploceidae, see detailed Remarks in the text.
three derived species of this cluster, M. anaplonotus, The glossifer cluster is supported by two synapo- M. daberti and M. plocepasseri, is characterized by the morphies: the development of membranous terminal ovate form of the central sclerite (25.1) and narrow lobules between setae h3 (13.1) and subapical mem- band-like lateral sclerites (31.1) in females. branes (15.1) in males. The main clade of this cluster, The branch bearing two other clusters (angolensis which is the sister to a sole species M. bubalornis, is and glossifer) is characterised by two synapomorphies: characterized by a narrowed transventral sclerite in straight, almost conical form of tarsus III in males males (16.1) and development of a cone-like and rela- (21.1) and relatively narrow band-like central scle- tively long external copulatory tube in females (34.1). rite in females (26.1). The angolensis cluster is sup- The plocei clade uniting the five most derived spe- ported by the development of entire (11.1) and rela- cies of the glossifer cluster (M. anaplecti, M. capensis, tively wide (14.1) opisthosomal membrane between M. gaud, M. plocei, and M. puylaerti) is characterised setae ps2 in males. In the basal species of the group, by three apomorphies in males: setae 3a are situ- M. lonchurae, the margin of membrane is smooth ated off the transventral sclerite (17.1), lateral parts and slightly convex (fig. 18e), while in the two de- of the adanal shield are separated from the median rived species, the membrane forms a pair of rounded part (20.1), and tarsus III has a curved apex (21.0). extensions as in M. angolensis (figs. 18a, b) or it is sec- In the context of the genus Metapteronyssus, a curved ondarily narrowed and represented by narrow band tarsus III in the plocei clade represents a reversion to a along the margin of the opisthosoma and a pair of plesiomorphic state of this character. Three relatively finger like lobules as in M. amadinae (figs. 17a, b). primitive species of the glossifer cluster (M. bubalornis,
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Table 2. Data matrix of characters states for Metapteronyssus and outgroup taxa. Character states are scored as 0 to 2, inapplicable states as ‘-’.
1 1111111112 2222222223 3333 1234567890 1234567890 1234567890 1234 Sturnotrogus subtrincatus 0000000000 000-0000-- 0000-00-00 --0- Scutulanyssus dasyritidis 0000000000 000-000100 0000-00-00 --0- Pteronyssoides holoplax 0000000000 000-000101 0100-00-01 --0- Metapteronyssus glossifer 1111011010 0-10110100 1111010002 0011 M. hordacei 1111011010 0-10110100 1111020002 0011 M. plocei 1111011010 0-10111101 0111010002 0011 M. capensis 1111011010 0-10111101 0111010012 0011 M. puylaerti 1111011010 0-10111101 0111011002 0111 M. anaplecti 1111011010 0-10111101 0111011022 0111 M. gaudi 1111011010 0-10111101 0111010102 0011 M. bubalornis 1111011010 0-10100100 1111010002 0010 M. plocepasseri 1111011111 0-10000100 0111100002 100- M. anoplonotus 1111011111 0-10000110 0112--0002 1011 M. daberti 1111011111 0-10000110 0111100002 1011 M. pseudonigritae 1111111111 0-10100100 0111000002 2010 M. lonchurae 1111011010 1001000100 1111010002 0010 M. angolensis 1111011010 1101000100 1111020002 0010 M. amadinae 1111111010 1101000100 1111010002 0010
M. glossifer, and M. hordacei) retain setae 3a on the of the opisthosoma between setae ps2 and develop- transventral sclerite and a T-shaped adanal shield, but ment of terminal lobules with membranous margins. have a straight tarsus III (figs. 2a, f, 5a, c) which is the This structure of the opisthosoma was apparently de- derived state of this character. veloped independently in the glossifer (figs. 2a, b) and In mapping the main evolutionary changes in the plocepasseri groups (figs. 13a, d) and in M. amadinae genus Metapteronyssus (fig. 20) it is possible to note from the angolensis group (figs. 16a, b). that its three clusters demonstrate some similar trends in morphological modifications. A strong tendency to Host associations reduce the central sclerite in females is realized in each Representatives of the genus Metapteronyssus lineage independently and in some cases in different (table 3) are restricted to three families of the super- ways. In the glossifer and angolensis groups the central family Passeroidea (parvorder Passerida) distributed sclerite is commonly represented by a narrow longi- in the Old World. Each species group of mites recog- tudinal band, the greatest width of which is less than nised in the course of phylogenetic analysis is restrict- 1/3 of idiosomal width. In M. angolensis (angolensis ed to a particular host family: M. angolensis to estrildid group) and M. hordacei (glossifer group) this sclerite finches (Estrildidae), glossifer to weavers (Ploceidae), is reduced to a narrow, almost rod-shaped sclerite and plocepasseri to sparrows, (Passeridae). Within (figs. 8b, 19a). In two species of the glossifer group all species groups most species are either monoxenous (M. anaplecti and M. puylaerti), the central sclerite is or restricted to a certain bird genus (with the excep- relatively large but split into anterior and posterior tion of one species from the plocepasseri group). For fragments (fig. 12a). In the plocepasseri group, most example, a relatively primitive species of the glossifer species have a relatively wide rectangular or ovate group, M. bubalornis, occurs on the buffalo weaver central sclerite (figs. 14a, b), but in M. anaplonotus Bubalornis albirostris, and two closely related spe- this sclerite is completely lost (fig. 12b). In males the cies M. glossifer and M. hordacei are associated with most characteristic trend independently realized in the bishops Euplectes franciscanus and E. hordaceus, different lineages is the extension of the terminal end respectively. Members of the derived subgroup
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Table 3 Host associations and geographic locations of Metapteronyssus species; (*) Questionable host association.
Mite species host species Locality Source glossifer group M. anaplecti Anaplectes rubriceps Rwanda Present study M. bubalornis Bubalornis albirostris Cameroon Mironov 2002 M. capensis Ploceus capensis South Africa Mironov 2002 M. gaudi Ploceus aurantius aurantius Nigeria Mironov 2002 P. bicolor amaurocephalus Cameroon Mironov 2002 P. nigerrimus Cameroon Mironov 2002 P. nigricollis brachypterus Cameroon Mironov 2002 M. glossifer Euplectes franciscanus Sudan Gaud 1953 M. hordacei E. hordaceus Central African Republic Present study M. plocei Ploceus cucullatus Central African Republic Mironov 2002 P. nigricolliis brachypterus Central African Republic Mironov 2002 P. xanthops South Africa Mironov 2002 M. puylaerti Ploceus velatus South Africa Mironov 2002 plocepasseri group M. anoplonotus Plocepasser mahali South Africa, Zimbabwe Mironov 2002 Quelea quelea* Zimbabwe Mironov 2002 M. daberti Plocepasser rufoscapulatus Zambia Mironov 2002 M. plocepasseri. Plocepasser mahali Zimbabwe Mironov 2002 M. pseudonigritae Pseudonigrita cabanisi Kenya Present study angolensis group M. angolensis Uraeginthus angolensis South Africa Mironov & Kopij 2000 U. bengalus Sudan Gaud 1953 Uraeginthus bengalus ugogoensis Rwanda Present study M. amadinae Amadina fasciata alexanderi Ethiopia Present study M. lonchurae Lonchura cantans orientalis Somali Mironov 2002
plocei are associated with weavers of the genera Ploceus Payne 2001, Ericson et al. 2002, Ericson & Johanson Cuvier, 1816 and Anaplectes Reichenbach, 1863. 2003, Barker et al. 2004), it is possible to propose The records of M. anoplonotus (plocepasseri group) a very provisional co-phylogenetic hypothesis for on hosts belonging to different families, Plocepasser Metapteronyssus species and their hosts. mahali (Passeridae) and Quelea quelea (Ploceidae) are exceptions and could be the result of natural or ac- We hypothesize that the genus Metapteronyssus cidental contamination. originated on the common ancestor of the core of Associations with particular host taxa noted at spe- Passeroidea and owing to co-speciation has become cies and species group levels give clear evidence of distributed among three avian families of this vast co-speciation of the genus Metapteronyssus with their grouping of sparrow-like passerines. However, tak- hosts. We realise that species diversity of this genus ing into consideration its numerous derived features is so far incompletely known, because many avian and significant morphological differences from re- species of the three host families, which potentially lated genera, it is possible that Metapteronyssus origi- might bear representatives of this genus, still remain nated earlier, even on the ancestor of the parvorder unexplored (almost 300 avian species). Neverthe- Passerida, where it split off from a common ancestor less, based on the phylogenetic relationships within with the genus Pteronyssoides. However, in contrast Metapteronyssus (fig. 20), currently known host asso- to the closely related genus Pteronyssoides, which is ciations (table 3), a current phylogenetic hypothesis distributed on twelve families of Passerida includ- for suprageneric taxa of passerine hosts (Sorenson & ing nine families of Passeroidea (Mironov & Wauthy
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2005a), Metapteronyssus species occur only on rep- Acknowledgements resentatives of three families (Estrildidae, Ploceidae, and Passeridae) belonging to the core Passeroidea. The authors wish to thank Prof. Dr. A. Fain In turn, within the core Passeroidea, these three (I������������������������������������������������nstitut royal des Sciences naturelles de Belgi- families belong to two different major branches. The que,���������������������������������������������� Bussels, Belgium), Dr. H. André (Musée royal smaller of these branches unites the estrildid finches de l’Afrique centrale, Tervuren, Belgium), and (Estrilduidae), weavers (Ploceidae), and whydahs and Dr. W.T. Atyeo (The University of Georgia, Athens, indigobirds (Viduidae); the larger branch incorporates USA) for loaning material used in the present study, the rest 10 families of the core Passeroidea includ- and Dr. H.C. Proctor (The University of Alberta, ing sparrows (Passeridae) (Sorenson & Payne 2001, Edmonton, Canada) for critically reviewing the man- Sorenson et al. 2003, Barker et al. 2004). uscript. This research is supported by a grant from the The plocepasseri group of Metapteronyssus probably Belgian Scientific Policy to SVM. originated on the ancestors of the larger branch, but was retained only on representatives of Passeridae. References The absence of Metapteronyssus on other avian fami- lies of this major branch of the core Passeroidea has Barker, F.K., A. Cibois, P Schikler, J. Feinstein & J. Cracraft, no clear explanation and is open for speculation. One 2004. Phylogeny�������������������������������������������������� and diversification of the largest avian radiation. – Proceedings of the National Academy of of possible suggestions is that the representatives of Sciences of the USA 101: 11040-11045. Metapteronyssus have gone extinct on other familial Dickinson, E.C., 2003. The Howard and Moore complete lineages of this passeroidean branch. checklist of the birds of the world, 3rd Edition. Princeton Two other species groups, angolensis and glossifer, University Press, Princeton, N.J., 1056 pp. evolved in association with two families of the second Ericson, P.G. P. & U.S. Johanson, 2003. Phylogeny of major branch of Passeroidea, Estrildidae and Plocei- Passerida (Aves: Passeriformes) based on nuclear and mi- tochondrial sequence data. – Molecular Phylogenetics dae respectively. It is interesting to note that repre- and Evolution 29: 129-138. sentatives of Metapteronyssus are not known from Ericson, P.G.P., L. Christidis, A. Cooper, M. Irestedt, the third family of this branch, Viduidae, which is J. Jackson, U.S. Johansson & J.A. Norman, 2002. phylogenetically closest to Estrildidae (Sorenson et al. A Gondwanan origin of passerine birds supported by 2003). 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Mironov, S.V., 2002. New species of the feather mite genus Nixon, K.C., 1999. Winclada. Version 1.0. – Published by Metapteronyssus Gaud, 1981 (Astigmata Analgoidea Pte- the author. Ithaca, New York. ronyssidae) from African passerines (Aves Passeriformes). Page, R.D.M., 2001. nde: nexus Data Editor 0.5.0. – Uni- – Bulletin de l’Institut Royal des Sciences Naturelles de versity of Glasgow, Glasgow, England. Belgique, Entomologie 72: 181-199. Sorenson, M.D. & Payne, R.B., 2001. A��������������������� single ancient ori- Mironov, S.V. & G. Kopij, 2000. A new feather mite species gin of brood parasitism in African finches: implication for of the genus Metapteronyssus Gaud, 1981 (Analgoidea: host-parasite coevolution. – Evolution 55: 2550-2567. Pteronyssidae). – Genus 11: 99-103. Sorenson, M.D., K.M. Sefc & R.B. Payne, 2003. Speciation���������� Mironov, S.V. & G. Wauthy, 2005a. A review of the feather by host switch in brood parasitic indigo birds. – Nature, mite genus Pteronyssoides Hull, 1931 (Astigmata: Ptero- 424: 928-931. nyssidae) from African and European passerines (Aves: Swofford, D.L., 1998. paup. Phylogenetic Analysis Us- Passeriformes) with analysis of mite phylogeny and host ing Parsimony (*and Other Methods), Beta version 4. associations. – Bulletin de l’Institut Royal des Sciences – Sinauer Associates, Sunderland, Massachusetts. Naturelles de Belgique, Entomologie 75: 155-214. Mironov, S.V. & G. Wauthy, 2005b. A new feather mite species of the genus Hyonyssus Gaud and Mouchet, 1959 (Astigmata: Pteronyssidae) from the Tit-hylia Pholidornis rushiae (Passeriformes: Remizidae). – International Jour- nal of Acarology 31: 101-106. Mironov, S.V. & G. Wauthy, 2005c. A new species of the feather mite genus Mouchetia Gaud, 1961 (Astigmata: Pteronyssidae) from the greencap eremomela Eremomela scotops (Passeriformes: Sylviidae) and taxonomic notes to Received: 29 March 2006 species of the genus. – Acarina 13: 3-14. Accepted: 3 April 2006
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