Chewing lice from wild passerines (Aves: Passeriformes) from Vietnam, with description of a new species of genus the Brueelia (Phthiraptera

Abstract Seven species of chewing lice (Insecta: Phthiraptera) were found on six species of passerine birds (Passeriformes) in northern Vietnam. A description and illustrations of Brueelia alophoixi sp. nov. ex Alophoixus pallidus (Pycnonotidae) are given. New host records are: Abroscopus superciliaris (Sylviidae) for Myrsidea sp.; and Orthotomus sutorius (Sylviidae) and Lonchura striata (Estrildidae) for Brueelia spp

Apomyrsidea klimesi Kolencik & Sychra & Allen 2021

<i>Apomyrsidea klimesi</i> (Sychra in Sychra <i>et al.</i>, 2006) gen. et comb. nov. <p>Figs 1–7, 10</p> <p> <i>Myrsidea klimesi</i> Sychra in Sychra <i>et al.</i>, 2006: 55, figs 10–11, 14–15 (type host: <i>Formicarius analis</i> (d’Orbigny & Lafresnaye, 1837)).</p> Material examined <p> <b>Holotype</b></p> <p> COSTA RICA • ♀; Hitoy Cerere BR, Provincia Limón; 9°40′ N, 85°27′ W; 100 m a.s.l.; 27 Aug. 2004; Literak, Capek and Havlicek leg.; ex <i>Formicarius analis</i> (d’Orbigny & Lafresnaye, 1837); INBio O.Sychra CR15.</p> <p> <b>Allotype</b></p> <p>COSTA RICA • ♂; same collection data as for holotype; INBio O.Sychra CR15.</p> <p> <b>Paratypes</b> (1 ♀, 1 ♂)</p> <p>COSTA RICA • 1 ♀, 1 ♂; same collection data as for holotype; 27 and 31 Aug. 2004; INBio O.Sychra CR14, CR16.</p> <p> <b>Other material</b></p> <p> COSTA RICA • 1 ♂; Zona Protectora Las Tablas on the Pacific slope of the Cordillera de Talamanca; 8°54′ N, 82°47′ W; 1300 m a.s.l.; 21. Aug. 2010; Sychra and Literak leg.; ex <i>Formicarius analis</i>; MMBC O.Sychra CR226.</p>Published as part of <i>Kolencik, Stanislav, Sychra, Oldřich & Allen, Julie M., 2021, Another puzzle piece in the systematics of the chewing louse genus Myrsidea, with a description of a new genus Apomyrsidea, pp. 36-50 in European Journal of Taxonomy 748</i> on page 44, DOI: 10.5852/ejt.2021.748.1339, <a href="http://zenodo.org/record/4736151">http://zenodo.org/record/4736151</a&gt

Apomyrsidea isacantha Kolencik & Sychra & Allen 2021, gen. et comb. nov.

<i>Apomyrsidea isacantha</i> (Valim & Weckstein, 2013) gen. et comb. nov. <p>Figs 9–10</p> <p> <i>Myrsidea isacantha</i> Valim & Weckstein, 2013: 381, figs 1–2, 11–12, 16, 18, 20–21 (type host: <i>Chamaeza nobilis</i> Gould, 1855).</p> Material examined <p> <b>Holotype</b></p> <p> BRAZIL • ♀; Rio Acanauí, Município Japurá, Amazonas; 2°01′38″ S, 66°40′28″ W; 18 Jul. 2007; Weckstein leg.; ex <i>Chamaeza nobilis</i> Gould, 1855; MZUSP 2310.</p> <p> <b>Paratypes</b> (2 ♀♀, 3 ♂♂)</p> <p>BRAZIL • 1 ♂; same collection data as for holotype; FMNH-INS 94000 • 1 ♀; same collection data as for holotype; DNA voucher Mysp.Chno.1.4.2011.4; FMNH-INS 94001 • 2 ♂♂; same collection data as for holotype; MZUSP 2312, MZUSP 2313 • 1 ♀; same collection data as for holotype; MZUSP 2311.</p>Published as part of <i>Kolencik, Stanislav, Sychra, Oldřich & Allen, Julie M., 2021, Another puzzle piece in the systematics of the chewing louse genus Myrsidea, with a description of a new genus Apomyrsidea, pp. 36-50 in European Journal of Taxonomy 748</i> on page 44, DOI: 10.5852/ejt.2021.748.1339, <a href="http://zenodo.org/record/4736151">http://zenodo.org/record/4736151</a&gt

Another puzzle piece in the systematics of the chewing louse genus Myrsidea, with a description of a new genus Apomyrsidea

A new avian chewing louse genus Apomyrsidea gen. nov. is described based on species parasitizing birds in the family Formicariidae. Diagnostic characteristics and phylogenetic analyses were used to evaluate and confirm the generic status and merit its recognition as unique and different from Myrsidea Waterston, 1915. Three species previously belonging to the genus Myrsidea are placed in the new genus Apomyrsidea gen. nov. and are discussed: Apomyrsidea circumsternata (Valim & Weckstein, 2013) gen. et comb. nov., Apomyrsidea isacantha (Valim & Weckstein, 2013) gen. et comb. nov. and Apomyrsidea klimesi (Sychra in Sychra et al., 2006) gen. et comb. nov

Apomyrsidea circumsternata Kolencik & Sychra & Allen 2021, gen. et comb. nov.

<i>Apomyrsidea circumsternata</i> (Valim & Weckstein, 2013) gen. et comb. nov. <p>Figs 8, 10</p> <p> <i>Myrsidea circumsternata</i> Valim & Weckstein, 2013: 383, figs 3–4, 13–15, 17, 19, 22 (type host: <i>Formicarius colma</i> Boddaert, 1783).</p> Material examined <p> <b>Holotype</b></p> <p> BRAZIL • ♀; Rio Acanauí, Município Japurá, Amazonas; 2°01′38″ S; 66°40′28″ W; 20 Jul. 2007; Weckstein leg.; ex <i>Formicarius colma</i> Boddaert, 1783; MZUSP 2314.</p> <p> <b>Paratypes</b> (2 ♀♀, 3 ♂♂)</p> <p>BRAZIL • 1 ♂; same collection data as for holotype; FMNH-INS 94002 • 1 ♀; same collection data as for holotype; DNA voucher Mysp.Foco.1.4.2011.3; FMNH-INS 94003 • 2 ♂♂; same collection data as for holotype; MZUSP 2316, MZUSP 2317 • 1 ♀; same collection data as for holotype; MZUSP 2315.</p>Published as part of <i>Kolencik, Stanislav, Sychra, Oldřich & Allen, Julie M., 2021, Another puzzle piece in the systematics of the chewing louse genus Myrsidea, with a description of a new genus Apomyrsidea, pp. 36-50 in European Journal of Taxonomy 748</i> on pages 43-44, DOI: 10.5852/ejt.2021.748.1339, <a href="http://zenodo.org/record/4736151">http://zenodo.org/record/4736151</a&gt

Brueelia bicurvata

<i>Brueelia bicurvata</i> (Piaget, 1880) <p>Figs 16–22</p> <p> <i>Nirmus bicurvatus</i> Piaget, 1880: 159, pl. 13, fig. 8.</p> <p> <i>Degeeriella bicurvata</i> – Harrison 1916: 109.</p> <p> <i>Brueelia bicurvata</i> – Hopkins & Clay 1952: 53. — Gustafsson & Bush 2017: 38.</p> Type host <p> <i>Vidua paradisaea</i> (Linnaeus, 1758) – long-tailed paradise whydah (Ploceidae).</p> Type locality <p>Original material from the Leiden Museum, but no type locality given by Piaget (1880).</p> Material examined <p> ZAMBIA • 2 ♂♂, 2 ♀♀, ex <i>Vidua paradisaea obtusa</i>; North Rhodesia [= Zambia], Luanshya; 26 May 1955; British Museum; NHML 1955-486, ML /121.</p> Description <p>Head rounded trapezoidal (Fig. 18), lateral margins of preantennal area slightly convex proximally, but slightly concave distally, frons broadly flattened to slightly concave. Marginal carina slender, deeply displaced and much widened at osculum, and with median margin slightly undulating. Ventral anterior plate oblong, with rounded posterior margin. Head chaetotaxy and pigmentation pattern as in Fig. 18. Preantennal nodi elongated. Preocular nodi larger than postocular nodi. Marginal temporal carina slender, with slightly undulating median margin. Gular plate slenderly lanceolate. Thoracic and abdominal segments and pigmentation patterns as in Figs 16–17.</p> <p> <b>Male</b></p> <p> Male subgenital plate does not reach terminal end of abdomen (Fig. 16). Thoracic and abdominal chaetotaxy as in Fig. 16. Basal apodeme broad, with concave lateral margins (Fig. 19). Proximal mesosome broad, convergent to median point, and with lateral extensions, making entire structure somewhat arrow-shaped (Fig. 20). Mesosomal lobes long and broad, with almost parallel lateral margins, rounded postero-lateral corners, and extensive rugose area along distal margin. Gonopore roughly semioval, about as long as wide. Penile arms long, reaching beyond distal margin of mesosome. Parameres long and broad, distal section elongated, with <i>pst1–2</i> as in Fig. 21. Measurements (n = 2): TL = 1.48– 1.51; HL = 0.36; HW = 0.30; PRW = 0.20; PTW = 0.29–0.31; AW = 0.40–0.41.</p> <p> <b>Female</b></p> <p> Thoracic and abdominal chaetotaxy as in Fig. 17; <i>psps</i> present on tergopleurites IV–VII. Subgenital plate shaped as in Fig. 22, with broad connection to cross piece. Vulval margin somewhat convergent to median point, with 3–5 short, slender <i>vms</i> and 5–8 short, thorn-like <i>vss</i> on each side; 2–4 short, slender <i>vos</i> on each side of subgenital plate; distal 1 <i>vos</i> median to <i>vss</i>. Measurements (n = 2): TL = 1.76–1.81; HL = 0.37–0.39; HW = 0.31–0.34; PRW = 0.20–0.21; PTW = 0.30–0.33; AW = 0.44–0.47.</p> Remarks <p> The presence of <i>psps</i> on the female tergopleurites IV–V is unusual in <i>Brueelia</i>. No examples of females with <i>psps</i> on these segments were included in the list of variation in abdominal chaetotaxy of <i>Brueelia</i> published by Gustafsson & Bush (2017: table 3). We know of no other species of <i>Brueelia</i> in which <i>psps</i> are present on the female tergopleurite IV, but <i>psps</i> are present on the female tergopleurite V in several species of <i>Brueelia</i> found on icterid hosts (Cicchino & Castro 1996). Carriker (1963) illustrated setae on female tergopleurites III–VII in the position of <i>psps</i> in <i>B. mirabile</i> Carriker, 1963, but did not illustrate any <i>sts</i>. It is therefore doubtful whether these setae represent <i>psps</i> or <i>sts</i> translocated to the dorsal side; we have not examined Carriker’s material. In females of most genera in the <i>Brueelia</i> complex, <i>psps</i> are absent on tergopleurites IV–V, and the general absence of these setae in <i>Brueelia</i> (except in <i>B. bicurvata</i>) is unusual for the complex (Gustafsson & Bush 2017: table 2).</p> <p> A specimen of <i>Brueelia</i> from <i>Vidua macroura</i> was included in the phylogeny of Bush <i>et al</i>. (2016), but its placement as sister to the remaining <i>Brueelia</i> s. str. received no statistical support. As <i>psps</i> are</p> <p> commonly found on female tergopleurites IV–V in many other genera of the <i>Brueelia</i> complex (see Gustafsson & Bush 2017), it is possible that this represents the ancestral condition in the <i>Brueelia</i> complex, and that this placement as sister to the remaining <i>Brueelia</i> is correct.</p> <p> Notably, these setae are present in both <i>Formicaphagus</i> Carriker, 1957 (see Price & Clayton 1996) and <i>Formicaricola</i> Carriker, 1957 (see Price & Clayton 1995), two of the genera most closely related to the <i>Brueelia</i> complex in the phylogeny of Bush <i>et al</i>. (2016). In the closest relative, <i>Neopsittaconirmus</i> Conci, 1942, the distribution of <i>psps</i> varies, e.g., present on III–VII in <i>N. albus</i> (Le Souëf & Bullen, 1902) (Price & Emerson 1978), but present only on IV–V in many species (Guimarães 1974) and only on IV in <i>N. gracilis</i> Guimarães, 1974 (see also Sychra 2005).</p>Published as part of <i>Gustafsson, Daniel R., Zou, Fasheng, Oslejskova, Lucie, Najer, Tomas & Sychra, Oldřich, 2019, Four new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera) from African hosts, with a redescription of Nirmus bicurvatus Piaget, 1880, pp. 1-48 in European Journal of Taxonomy 507</i> on pages 16-20, DOI: 10.5852/ejt.2019.507, <a href="http://zenodo.org/record/2612931">http://zenodo.org/record/2612931</a&gt

Brueelia Keler 1936

Genus <i>Brueelia</i> Kéler, 1936 <p> <i>Philopterus</i> Nitzsch, 1818: 288 partim.</p> <p> <i>Nirmus</i> Nitzsch, 1818: 291 partim.</p> <p> <i>Degeeriella</i> Neumann, 1906: 60 partim.</p> <p> <i>Brueelia</i> Kéler, 1936: 257.</p> <p> <i>Painjunirmus</i> Ansari, 1947: 285.</p> <p> <i>Allobrueelia</i> Eichler, 1951: 36 partim.</p> <p> <i>Nigronirmus</i> Złotorzycka, 1964: 248.</p> <p> <i>Spironirmus</i> Złotorzycka, 1964: 261.</p> <p> <i>Serinirmus</i> Soler Cruz <i>et al.</i>, 1987: 244.</p> Type species <p> <i>Brueelia rossittensis</i> Kéler, 1936: 257 (= <i>Nirmus brachythorax</i> Giebel, 1874: 134) by original designation.</p> Remarks <p> Clay (1954) discussed the use of the <i>post-spiracular sensillum</i> in determining homology in the abdominal chaetotaxy of Ischnocera. She stated that in <i>Brueelia</i>, these sensilla are known from segments III–VII, whereas in all other groups of ischnoceran lice, they are never found posterior to segment V. Gustafsson & Bush (2017) included these sensilla in their illustrations, but neglected to discuss their importance in the text. Based on our investigation of several hundred species of lice in the <i>Brueelia</i> complex, it seems that these sensilla occur on segments II–III only in the following genera: <i>Brueelia</i>, <i>Teinomordeus</i> Gustafsson & Bush, 2017, <i>Acronirmus</i> Eichler, 1953 and <i>Sychraella</i> Gustafsson & Bush, 2017. In all other genera of the <i>Brueelia</i> complex, these sensilla only occur on segments IV–V. However, they are typically very hard to see, especially in species with reduced tergopleurites.</p> <p> Gustafsson & Bush (2017) also neglected to explicitly state that it is the position of post-spiracular setae in relationship to this sensillum that determines whether they are <i>psps</i> or <i>aps</i>. Any setae positioned laterally to the sensillum are <i>aps</i>, whereas any setae situated immediately median to this sensillum are <i>psps</i>. Note that <i>aps</i> and <i>psps</i> on the first abdominal segment bearing post-spiracular setae (often segment V or VI in <i>Brueelia</i>) may be similar in length. Moreover, in some species of, e.g., <i>Olivinirmus</i> Złotorzycka, 1964 there may be more than one <i>psps</i> per side on some segments. To our knowledge, no species in the <i>Brueelia</i> complex has more than one <i>aps</i> per side on any segment.</p> <p> Several of the species here belong to a group of pied <i>Brueelia</i> species found mainly on African hosts in the families Ploceidae, Estrildidae and Paridae. The only species of this group known from hosts outside Africa is <i>Brueelia plocea</i> (Lakshminarayana, 1968), from India. We have seen many additional species in this group, all from African hosts; however, suitable hosts in the same genera are found in South Asia. We here refer to this group as the “African pied <i>Brueelia</i> ” group, to distinguish it from the New World <i>ornatissima</i> group, which have similar pigmentation patterns. This group comprises the following species: <i>Brueelia plocea</i> (Lakshminarayana, 1968); <i>B. queleae</i> Sychra & Barlev in Sychra <i>et al</i>., 2010a; <i>B. cantans</i> Sychra in Sychra <i>et al.</i>, 2010b; <i>B. aguilarae</i> Gustafsson & Bush, 2017; <i>B. mpumalangensis</i> Gustafsson <i>et al</i>., 2018; <i>B. semiscalaris</i> sp. nov.; <i>B. terspichore</i> sp. nov.; <i>B. sima</i> sp. nov.</p> <p>The main characteristic of this group is the striking pigmentation pattern. This varies slightly between species, but typically includes having dark pigmentation on the anterior and posterior margins of sternites III–VI, the female tergopleurite IX +X, along the lateral margins of the abdomen, around the distal section of femora I–III and on the subgenital plates. The dark areas are generally at least dark brown, but may appear black in some species; both sternal and subgenital plates typically have distinct translucent fenestrae in both sexes.</p> <p> In the phylogeny of Bush <i>et al</i>. (2016), members of this group (e.g., <i>Brueelia queleae</i> and <i>Brueelia</i> sp. (= <i>B. mpumalangensis</i>) ex <i>Melaniparus niger</i>) were placed in different parts of the tree, suggesting that they do not form a natural group; however, these placements received no statistical support. Apart from pigmentation patterns, the morphological characters of this group are also very diverse, suggesting that the division of this group in the phylogeny of Bush <i>et al.</i> (2016) may be correct.</p> <p> Nevertheless, for the purposes of identification and keying, we consider the ʻAfrican pied <i>Brueelia</i> ʼ group a useful grouping to help sort out the vast diversity of species of <i>Brueelia</i> on African hosts. As more species of <i>Brueelia</i> from African hosts become known, the relationships of the species in this informal group may have to be revised, and the group may be found to be artificial. We provide a key to the described species in this group below.</p> Key to the ‘African pied <i>Brueelia</i> ’ <p> Note that the dorsal abdominal setae in the original illustration of <i>Brueelia plocea</i> have been translocated to the ventral side (Lakshminarayana 1968). No dorsal setae are given in the original description (ibid.: table II), but multiple setae are illustrated on some segments; we interpret all setae on these segments except the <i>sts</i> as dorsal setae. The female of <i>B. plocea</i> is undescribed.</p> <p>1. Male....................................................................................................................................................2 – Female................................................................................................................................................9</p> <p> 2. <i>Accessory post-spiracular setae</i> present on tergopleurite V (Fig. 9)..............................................3 – <i>Accessory post-spiracular setae</i> absent on tergopleurite V (Fig. 23).............................................6</p> <p> 3. <i>Tergal posterior setae</i> present on tergopleurites V–VI....................................................................4 – <i>Tergal posterior setae</i> absent on tergopleurites V–VI (Fig. 9)........................................................5</p> <p> 4. Frons rounded; <i>aps</i> present on tergopleurite IV.................................................................................................................................................... <i>Brueelia queleae</i> Sychra & Barlev in Sychra <i>et al</i>., 2010a</p> <p> – Frons flattened; <i>aps</i> absent on tergopleurite IV...... <i>Brueelia cantans</i> Sychra in Sychra <i>et al</i>., 2010b</p> <p> 5. <i>Tergal posterior setae</i> present on tergopleurite VII (Fig. 9); dark pigmentation of subgenital plate limited to anterior margin (Figs 9, 40).................................... <i>Brueelia semiscalaris</i> sp. nov.</p> <p> – <i>Tergal posterior setae</i> absent on tergopleurite VII; dark pigmentation of subgenital plate extensive along lateral margins, reaching distal end of subgenital plate............................................................................................................................. <i>Brueelia aguilarae</i> Gustafsson & Bush, 2017</p> <p> 6. Preantennal head narrowly rounded............................... <i>Brueelia plocea</i> (Lakshminarayana, 1968) – Preantennal head broad, frons either rounded or flattened (Fig. 25).............................................7</p> <p> 7. <i>Tergal posterior setae</i> present on tergopleurite VI (Fig. 23) (absent in single examined specimen from <i>E. p. delamerei</i>); 2 <i>ps</i> on each side of abdominal segment IV (Fig. 23)................................................................................................................................................ <i>Brueelia terpsichore</i> sp. nov.</p> <p> – <i>Tergal posterior setae</i> absent on tergopleurite VI (Fig. 30); 1 <i>ps</i> on each side of abdominal segment IV (Fig. 30)..........................................................................................................................8</p> <p> 8. Preantennal area roughly trapezoidal, with flattened frons (Fig. 32); <i>tps</i> present on tergopleurite VII (Fig. 30)...................................................................................................... <i>Brueelia sima</i> sp. nov.</p> <p> – Preantennal area roughly semioval, with rounded frons; <i>tps</i> absent on tergopleurite VII.......................................................................................... <i>Brueelia mpumalangensis</i> Gustafsson <i>et al</i>., 2018</p> <p>9. Frons rounded.................................................................................................................................10 – Frons flattened (Fig. 31)..................................................................................................................11</p> <p> 10. Subgenital plate largely translucent, with lateral areas of dark pigmentation clearly separated from anterior band of dark pigmentation....... <i>Brueelia queleae</i> Sychra & Barlev in Sychra <i>et al</i>., 2010a</p> <p> – Subgenital plate largely dark, with lateral and anterior sections of dark pigmentation fused.................................................................................... <i>Brueelia mpumalangensis</i> Gustafsson <i>et al</i>., 2018</p> <p> 11. <i>Pleural setae</i> present on abdominal segment IV (Fig. 10)............................................................12 – <i>Pleural setae</i> absent on abdominal segment IV (Fig. 31)............................. <i>Brueelia sima</i> sp. nov.</p> <p>12. Subgenital plate with largely dark pigmentation apart from a central more or less T-shaped translucent fenestra (sections of this fenestra may be interrupted as in Fig. 29).....................13</p> <p> – Subgenital plate largely translucent, but with central arched section of dark pigmentation connected to anterior band of dark pigmentation as in Figs 15, 39............ <i>Brueelia semiscalaris</i> sp. nov.</p> <p> 13. Vulval margin with distinct median point............... <i>Brueelia aguilarae</i> Gustafsson & Bush, 2017 – Vulval margin without distinct median point (Fig. 29)..................................................................14</p> <p> 14. Subgenital plate almost entirely dark, with translucent areas small and isolated from each other (Figs 29, 41)........................................................................................... <i>Brueelia terpsichore</i> sp. nov.</p> <p> – Subgenital plate with clear central T-shaped translucent fenestrum.............................................................................................................................. <i>Brueelia cantans</i> Sychra in Sychra <i>et al.</i>, 2010b</p>Published as part of <i>Gustafsson, Daniel R., Zou, Fasheng, Oslejskova, Lucie, Najer, Tomas & Sychra, Oldřich, 2019, Four new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera) from African hosts, with a redescription of Nirmus bicurvatus Piaget, 1880, pp. 1-48 in European Journal of Taxonomy 507</i> on pages 7-9, DOI: 10.5852/ejt.2019.507, <a href="http://zenodo.org/record/2612931">http://zenodo.org/record/2612931</a&gt

Four new species of Brueelia Kéler, 1936 (Phthiraptera: Ischnocera) from African hosts, with a redescription of Nirmus bicurvatus Piaget, 1880

Four new species of Brueelia Kéler, 1936 are described and illustrated. All of them parasitize African endemic host species in the families Passeridae, Ploceidae, and Estrildidae (Passeriformes). They are: Brueelia pofadderensis sp. nov. ex Passer melanurus damarensis Reichenow, 1902 and P. m. vicinus Clancey, 1958; B. semiscalaris sp. nov. ex Granatina granatina (Linnaeus, 1758); B. sima sp. nov. ex Malimbus nitens (Gray, 1831); B. terpsichore sp. nov. ex Euplectes jacksoni (Sharpe, 1891) and E. progne delamerei (Shelley, 1903). In addition, Brueelia bicurvata (Piaget, 1880) is redescribed and reillustrated from non-type material. A summary of all published records of lice in the Brueelia complex from Africa since 1980 is provided. We also estimate the unknown diversity of African species of Brueelia based on an index of host specificity calculated for each host family independently. The unknown diversity is estimated to be over 1000 species of Brueelia from African hosts, compared to the < 50 species in this genus currently recorded from Africa