Phthiraptera: Amblycera: Menoponidae) from Chinese Babblers (Passeriformes: Leiothrichidae, Paradoxornithidae, Timaliidae)

Zootaxa 4878 (1): 103–128 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4878.1.4 http://zoobank.org/urn:lsid:zoobank.org:pub:67B9660E-6FC1-414A-A50E-68767C302C46

Four new species of Myrsidea (Phthiraptera: Amblycera: Menoponidae) from Chinese babblers (Passeriformes: Leiothrichidae, Paradoxornithidae, Timaliidae)

LUJIA LEI1,2,6, XINGZHI CHU1,3, BILAL DIK7, FASHENG ZOU1,4, HAITAO WANG6 & DANIEL R. GUSTAFSSON1,5* 1Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Con- servation and Utilization, Guangdong Institute of Zoology, Guangzhou, Guangdong, China. 2 https://orcid.org/0000-0003-3310-1476 3 https://orcid.org/0000-0002-6659-6279 4 https://orcid.org/0000-0002-8913-5651 5 https://orcid.org/0000-0001-8868-8310 6Jilin Engineering Laboratory for Avian Ecology and Conservation Genetics, School of Life Sciences, Northeast Normal University, 5268 Renmin Street, Changchun, 130024, Jilin, China. https://orcid.org/0000-0003-2531-149X 7Department of Parasitology, Faculty of Veterinary Medicine, Selçuk University, Selçuklu, Konya, Turkey. https://orcid.org/0000-0002-7553-5611 *Corresponding author. �[email protected]

Abstract

Four new species of amblyceran chewing lice of the genus Myrsidea Waterston, 1915 are described from hosts of the babbler families Leiothrichidae, Paradoxornithidae and Timaliidae in China. They are: Myrsidea attenuata n. sp. from Garrulax maesi maesi (Oustalet, 1890), Myrsidea zhangae n. sp. from Ianthocincla berthemyi (Oustalet, 1876), Myrsidea liopari n. sp. from Lioparus chrysotis amoenus (Mayr, 1941) and L. chrysotis swinhoii (Verreaux, 1871), and Myrsidea suthorae n. sp. from Suthora verreauxi verreauxi Sharpe, 1883. A checklist of host-louse associations for identified and unidentified Myrsidea species known from babblers is provided.

Key words: Phthiraptera, Amblycera, Menoponidae, Myrsidea, new species, Leiothrichidae, Paradoxornithidae, Timaliidae, babblers, China

Introduction

The genus Myrsidea Waterston, 1915, is the most speciose genus of chewing lice, with over 350 described species (Price et al. 2003; Valim & Weckstein 2013). Myrsidea is a cosmopolitan genus, occurring primarily on perching birds (Passeriformes). Most species are known from a single host species, and species associated with multiple host species typically occur on closely related hosts (Price et al. 2003; but see Sychra et al. 2014). The term “babblers” includes a diverse assemblage of bird species, covering members of several closely related families (Leiothrichidae, Paradoxornithidae, Pellorneidae, Timaliidae), most of which are found in tropical and sub- tropical Asia, including China (e.g. Arlott 2017). Collectively, these families comprise about 300 species (Clements et al. 2018). However, only 28 species of Myrsidea are presently known from hosts of these four families (Price et al. 2003, 2006; Khan et al. 2009; Najer et al. 2012). The known species of Myrsidea are distributed across most of the babbler diversity, but none had been described from parrotbills and allies (Paradoxornithidae). We here describe the first two species of Myrsidea from parrotbills and allies, as well as two new species from other Chinese babblers.

Accepted by R.L. Palma: 28 Sept. 2020; published: 12 Nov. 2020 103 Material, methods and conventions

Most specimens were collected following the method described by Gustafsson et al. (2019); however, collections of lice have been carried out by several people over nearly a decade and, therefore, the methods used varied. Lice have been slide-mounted following the technique in Gustafsson et al. (2019). All specimens are deposited in the collection of the Guangdong Institute for Applied Biological Resources, Guangzhou, Guangdong, China (GIABR). Morphological characters and measurements follow the methodology outlined by Valim & Weckstein (2013). Abdominal setal counts of species of Myrsidea are not consistent in the literature, with some authors (e.g. Price et al. 2006; Najer et al. 2012) including the setae of the post-spiracular complex among the tergal setae, while other authors (e.g. Tandan 1972) excluding them. We here follow Clay (1966) and Valim & Weckstein (2013) by counting all marginal setae except the post-spiracular setae on tergum I, but only counting the variable setae between the setae of the post-spiracular complex on subsequent segments. We also follow Clay (1966) and Valim & Weckstein (2013) when counting sternal setae, excluding the two antero-lateral setae on each side of sternum II. In most species treated here, the setae of the lateral sternal brushes are indistinguishable from those on the posterior margin. Therefore, we follow Clay (1966) including all sternal setae on the posterior margin in the total for posterior marginal setae, with all anterior setae (median + brush setae) included within parentheses after the number of posterior marginal setae. All measurements are given in millimeters, as ranges (mean value, when n ≥ 10). Abbreviations used are: ANW = female anus width; AW = abdominal width (taken at segment IV); dhs = dorsal head seta (sensu Clay 1969); GL = male genitalia length; GSL = male genital sclerite length; HL = head length (taken at midline); ls5 = labial seta 5 (sensu Clay 1966); MSPL = metasternal plate length (taken at midline); MSW = mesothorax width; MTW = metathorax width; POW = preocular width; PRW = prothorax width; PSPL = prosternal plate length (taken at midline); TL = total length (at midline); TW = head width (taken at temples). Host taxonomy follows Clements et al. (2018).

Systematics

Phthiraptera Haeckel, 1896: 703.

Amblycera Kellogg, 1896: 68.

Menoponidae Mjöberg, 1910: 26.

Myrsidea Waterston, 1915: 51

Acolpocephalum Ewing, 1927: 88. Australmenopon Conci, 1942: 30. Allomyrsidea Conci, 1942: 31. Corvomenopon Conci, 1942: 31. Ramphasticola Carriker, 1949: 305. Alcediniphilus Ansari, 1951: 189. Myrsidella Eichler, 1951: 49. Densidea Złotorzycka, 1964: 171. Vulgidea Złotorzycka, 1964: 172. Liquidea Złotorzycka, 1964: 173. Lanimenopon Złotorzycka, 1964: 177. Eichlerinopon Złotorzycka, 1964: 179. Neomyrsidella Złotorzycka, 1964: 182. Wolfdietrichia Złotorzycka, 1973: 51.

Type species: Myrsidea victrix Waterston, 1915: 13, by original designation. Type host: Ramphastos ambiguous swainsonii Gould, 1833.

Myrsidea sp. 6; Chu et al. 2019: 338.

Type host: Garrulax maesi maesi (Oustalet, 1890) – grey laughingthrush (Leiothrichidae). Type locality: Nanling Reservation, Ruyang County, Guangdong Province, China. Diagnosis. Myrsidea attenuata is identified as M. assamensis Tandan, 1972, in the keys to males in Tandan (1972). The same type of genital sclerite, with a single median attenuation, is also found in Myrsidea agarwali Khan et al. 2009. These species can be separated by the thoracic and abdominal chaetotaxy (Table 1), particularly the number of setae on the metanotum and on terga I–II (and in male also tergum III). In addition, Myrsidea attenuata can be separated from M. assamensis by number of setae in the femoral brush of femur III: 19–23 setae in female and 16–21 setae in male in M. attenuata (Figs 1–2), but 24–27 setae in female and 22–25 setae in male in M. assamensis. Males of Myrsidea attenuata can be separated from those of M. agarwali by having distinct gaps between the two median-most seta on each side of terga III–V that are distinctly larger than the gaps between the more lateral tergal setae on the same segments (Fig. 1); in M. agarwali, the tergal setae on these segments are evenly spaced. Also, the genital sclerite of M. agarwali appears to lack the distal attenuated section of the median part, which is present in M. attenuata (Fig. 5). Description. Female (n=3). Habitus as in Fig. 2. Setal measurements: dhs10 = 0.044–0.050; dhs11 = 0.120– 0.137; ratio dhs10/11 = 0.35–0.41; ls5 = 0.044–0.071 Ventrolateral fringe with 10–12 setae. Gula with 4–5 setae on each side. Hypopharynx similar to Fig. 4. First tibia with 2–3 outer lateral ventral and 4–5 dorsal setae. Metanotum with posterior margin straight and 2 setae on each side. Metasternal plate with 4–5 setae on each side. Femoral brush with 19–23 setae. Metanotum and abdomen as in Fig. 2. Posterior margin of tergites straight. Se- tal rows continuous on segments II–III, with small median gap on segments IV–VIII. Anterior setae absent on terga. Tergal setae: I, 2–4; II, 8–11; III, 12–13; IV, 12–13; V, 10–15; VI, 11; VII, 9–10; VIII, 4–5. Tergum IX with 4–5 medium-sized inner posterior setae on each side. Anterior setae present on sterna II–VI. Sternal setae: I, 0; II, each aster of 3–4 setae, posterior margin with 8–10 setae and anteriorly with 9–10 setae on one side; III, 12–14 (2–3); IV, 16–22 (11–16); V, 20–22 (20–24); VI, 20–21 (18–26); VII, 8–12 (6–10); VIII–IX, 10–15; vulval margin with 12 setae. Anus with 32–38 setae in ventral fringe and 32–38 setae in dorsal fringe; dorsal fringe with small number of short setae (about 1/2 length of other setae) interspersed centrally; all central setae of ventral fringe at most 1/3 of length of lateral setae. Pleural setae: I, 7–10 (0–1); II, 8–11 (1–3); III, 8–10 (4–6); IV, 8–9 (3–5); V, 6–8 (3–5); VI, 3–7 (1–3); VII, 5–6 (1–2), VIII: 3–5 (0–1). Measurements as in Table 2. Male (n=4). Much as in female, habitus as in Fig. 1, head shape as in Fig.3. Setal measurements: dhs10 = 0.038–0.056; dhs11 = 0.109–0.127; ratio dhs10/11 = 0.33–0.51; ls5 = 0.040–0.075. Gular plate with 4 setae on each side (1 male with 5 on one side). Hypopharynx as in Fig. 4. Metanotum with 4–6 setae on posterior margin. Femoral brush with 16–21 setae. Anterior setae absent on terga. Tergal setae: I, 2–4; II, 6; III, 7–8; IV, 10–11; V, 10–11; VI, 8–10; VII, 6–8; VIII, 4. Setal rows with small median gap in at least segments II–VIII. Inner posterior setae present on tergum IX. Anterior setae present on sternites II–VII. Sternal setae: I, 0; II, each aster with 4–5 setae, posterior margin with 15–18, and anteriorly with 23 setae; III, 21–23 (3–4); IV, 21–22 (10–16); V, 20–24 (13–17); VI, 22–24 (17–18); VII, 16 (8–12); VIII–IX, 22–23 (holotype with 8 visible, but many apparently absent). Anus with 8 internal setae on ventral side and 4–5 on dorsal side. Pleural setae: I, 5–6 (0); II, 6–8 (0–1); III, 5–9 (0–3); IV, 7–8 (1–3); V, 6–7 (1–3); VI, 6–7 (1–2); VII, 4–6 (0–2); VIII, 3–4 (0). Genitalia and genital sac sclerite as in Fig. 8. Measurements as in Table 2. Type material: Holotype ♂, Nanling Reservation, Ruyang County, Guangdong Province, China, 11 Apr. 2013, Q. Zhang & Y. Zhang, bird J0819, GD-PHTH-00057 (GIABR). Paratypes: 2♀, Ledongjianfengling, Ledong Coun- ty, Hainan Province, China, 24 Mar. 2015, X. Che & X. Chu, GD-PHTH-00059 and GD-PHTH-000062 (GIABR); 3♂, same data as previous sample, except bird J2570, GD-PHTH-00056 (GIABR); 1♀, same data as previous sample, except bird J2569, GD-PHTH-00063 (GIABR). Non-type specimens: 2 nymphs, same data as holotype, GD-PHTH-00058; 3 nymphs, Ledongjianfengling, Ledong County, Hainan Province, China, 24 Mar. 2015, X. Che & X. Chu, bird J2568, GD-PHTH-00059–61 (GIABR).

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 105 FIGURE 1. Myrsidea attenuata n. sp., male habitus, dorsal and ventral views. Tarsi of leg I broken or distorted in all examined males, and not illustrated.

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 107 Table 1. Comparison of tergal and sternal setae of Myrsidea attenuata n. sp., Myrsidea agarwali Khan et al. 2009, and Myrsidea assamensis Tandan, 1972, with data from the two latter species taken from their original descriptions. Numbers refer to total number, with both sides of the specimen put together, with the exception of asters, which are counted separately. For setae on sterna, anterior setae are given within parentheses for all segments except sternum II, for which each set of setae is given separately. Pleural setae are not given in full by Tandan (1972) and Khan et al. (2009), and therefore omitted from this list. Myrsidea attenuata n. sp. Myrsidea agarwali Khan et al. Myrsidea assamensis Tandan, 2009 1972 Male Female Male Female Male Female Posterior 4–6 4 6 8 4 4 metanotum Tergum I 2–4 2–4 6–7 8–10 4 4–5 Tergum II 6 8–11 9–12 10–12 7–8 12–13 Tergum III 7–8 12–13 12–14 12–14 10–11 13–14 Tergum IV 10–11 12–13 10–14 17–18 9–11 12–14 Tergum V 10–11 10–15 10–12 12–14 10–11 12–14 Tergum VI 8–10 11 10–12 12–13 9–10 10–13 Tergum VII 6–8 9–10 6–8 6–8 7–8 8–9 Tergum VIII 4 4–5 4–6 7–8 4–5 5–6 Sternum I 0 0 0 0 0 0 Sternum II 23 9–10 12–17 18–20 28–34 11–13 (anterior) Aster 4–5 3–4 3–4 3–4 4–5 3–5 Sternum II 15–18 8–10 14–16 8–9 14–16 9 (posterior) Sternum III 11 (0–1) 7–8 (0) 8–10 (1–2) 8–10 (0–4) 11–14 (4–7) 7–9 (0) (central) Sternum III 4–5 (1–2) 3–4 (0–1) 3–4 (1–3) 2–3 (3–5) 45–5 (2–4) 4–5 (2–4) (brush) Sternum IV 10–12 (3–4) 10–12 (1–2) 8–10 (2–5) 8–10 (7–10) 12–13 (7–11) 9–11 (3–6) (central) Sternum IV 5–6 (4–6) 4–5 (4–7) 4–5 (3–5) 3–4 (4–6) 6–7 (5–8) 5–9 (6–5) (brush) Sternum V 9–13 (4–6) 12–13 (3–4) 7–9 (4–5) 8–10 (7–11) 10–13 (6–10) 9–11 (4–7) (central) Sternum V 5 (5–7) 4–5 (7–10) 6–7 (3–5) 3–4 (8–10) 6–7 (5–10) 6–7 (8–11) (brush) Sternum VI 10–11 (4–5) 9–12 (3–4) 7–10 (3–5) 8–10 (8–10) 11–12 (6–10) 9–10 (5–9) (central) Sternum VI 5–6 (6–8) 4–6 (7–9) 5–6 (6–7) 3–5 (4–6) 5–6 (5–8) 5–7 (6–11) (brush) Sternum VII 9–12 (3–5) 4 (2–3) 6–7 (4–3) 7–10 (6–8) 7–10 (7–9) 7–8 (4–9) (central) Sternum VII 2–3 (2–4) 3–4 (2) 3–4 (1–2) 2–3 (2–4) 3–4 (1–4) 1–3 (2–4) (brush) Vulval margin --- 12 --- 42–50 --- 15–16

Etymology. The specific epithet is derived from “attenuare”, Latin for “to make slender”, referring to the middle section of the genital sclerite. Remarks. In all females, the dorsal side of the abdomen has been displaced slightly to one side due to mounting, so that the pleurites are visible only on the ventral side. Our illustration of the abdomen is as seen on the slides, without any symmetrical reconstruction.

Myrsidea zhangae Lei, Chu, Dik, Zou, Wang & Gustafsson, new species (Figs 5, 9, 12–14)

Myrsidea sp. 7; Chu et al. 2019: 338.

Type host: Ianthocincla berthemyi (Oustalet, 1876) – rusty laughingthrush (Leiothrichidae). Given as “Garrulax caerulatus” in Chu et al. (2019: 338), see below. Type locality: Badagongshan, Sangzhi County, Hunan Province, China. Diagnosis. Myrsidea zhangae is identified as M. monilegeri in the key of Tandan (1972). Females of M. monilegeri are unknown, but males of these two species can be separated by the following characters: femoral brush with 26–29 setae in M. monilegeri, but with 18–22 setae in M. zhangae; tergum I with 7 setae in M. monilegeri, but with 3–4 setae in M. zhangae (one specimen of M. zhangae has 9 setae on tergum I); tergum III with 13 setae in M. monilegeri, but with 6–9 setae in M. zhangae; anterior pleural setae present on segments III–VI in M. monilegeri, but only rarely present, and then only on one side, on segments III and V–VI in M. zhangae; sternum II with 27 anterior setae and 20 marginal setae in M. monilegeri, but with 6–11 anterior setae and 10–14 marginal setae in M.

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 109 zhangae; genital sclerite with median section longer than lateral sections and lateral sections slender distally in M. monilegeri, but genital sclerite with median section shorter than lateral sections, and lateral sections conspicuously widened distally in M. zhangae.

FIGURES 4–7. Hypopharyngeal sclerites. 4, Myrsidea attenuata n. sp. 5, Myrsidea zhangae n. sp. 6, Myrsidea liopari n. sp. 7, Myrsidea suthorae n. sp. All illustrations drawn from male specimens, and to same scale.

Tandan (1972) described two females from Garrulax monileger schauenseei that could not be placed unambig- uously in M. monilegeri, and he considered that future collections may resolve this issue. Assuming that Tandan’s females are conspecific with M. monilegeri, the female M. zhangae can be separated M. monilegeri by the following characters: femoral brush with 27–33 setae in specimens from G. m. schauenseei, but with 23–25 setae in M. zhangae; tergum I with 11 setae in specimens from G. m. schauenseei, but with 4 setae in M. zhangae; each aster of sternum II with 5 setae in material from G. m. schauenseei, but with 4 setae in M. zhangae; sternum II with 2–4 anterior setae and 15 marginal setae in material from G. m. schauenseei, but without anterior setae and with 9–12 marginal setae in M. zhangae. However, much of sternum II is obscured by gut contents in all females examined, and the anterior setae of G. m. schauenseei are said to be medio-lateral, which may suggest that those of M. zhangae are located very medianly and thus overlooked.

110 · Zootaxa 4878 (1) © 2020 Magnolia Press lei et al. FIGURES 8–11. Male genitalia, dorsal and ventral views. 8, Myrsidea attenuata n. sp. 9, Myrsidea zhangae n. sp. 10, Myrsidea liopari n. sp. 11, Myrsidea suthorae n. sp.

Description. Female (n=5). Habitus as in Fig. 13. Setal measurements: dhs10 = 0.057–0.072; dhs11 = 0.102– 0.117; ration dhs10/11 = 0.52–0.66; ls5 = 0.062–0.073. Ventrolateral fringe with 9–12 setae. Gular plate with 3–4 setae on each side. Hypopharynx similar to Fig. 5. First tibia with 2–3 outer lateral ventral and 2–5 dorsal setae. Metanotum with posterior margin straight and with 4–6 setae on posterior margin. Metasternal plate with 8–10 setae. Femoral brush with 23–25 setae. Metanotum and abdomen as in Fig. 13. Posterior margin of tergites straight. Setal rows continuous on terga I–III or with gaps on terga IV–VIII. Anterior tergal setae absent. Tergal setae: I, 4; II, 9–10; III, 12–16; IV, 12–14; V, 11–14; VI, 9–12; VII, 8–11; VIII, 4–6. Inner posterior setae present on tergite IX. Sternal setae: I, 0; II, each aster of 4 setae, posterior margin with 9–12 setae and anteriorly with 0–1 setae; III, 16–18 (3–7); IV, 21–23 (22–31); V, 21–23 (24–35); VI, 19–20 (27–29); VII, 9–12 (6–12); VIII–IX, 11–16; Vulval margin with 5–14 setae. Pleural setae: I, 7–9 (0); II, 8–11 (0–2); III, 8–10 (1–3); IV, 6–10 (1–2); V, 6–8 (1–3); VI, 6–7 (0–2); VII, 4–6 (0–2); VIII, 3–4 (0). Anus with 30–37 setae in ventral fringe and 30–33 setae in dorsal fringe. About every third central setae of dorsal fringe much shorter than others (about 1/2 length). Measurements as in Table 2. Male (n=9). Much as in female, habitus as in Fig. 12, head shape as in Fig.14. Setal measurements: dhs10 = 0.058–0.096; dhs11 = 0.096–0.114; ratio dhs10/11 = 0.55–0.93; ls5 = 0.053–0.080. Gular plate with 3–4 setae on each side. Hypopharynx as in Fig. 5. Metanotum with 2–6 setae on posterior margin. Femoral brush with 18–22 setae. Anterior setae absent on terga. Tergal setal rows with median gap in all segments. Tergal setae: I, 3–4 (one specimen with 9); II, 5–11; III, 6–9; IV, 8–11; V, 8–10; VI, 8–11; VII, 6–8; VIII, 4. Inner posterior setae present. Anterior setae present on sterna II–VII. Sternal setae: I, 0; II, each aster of 4–5 setae, with 10–14 marginal setae and 6–11 anterior setae; III, 16–18 (3–6); IV, 19–22 (19–26); V, 20–22 (25–26); VI, 17–22 (20–25); VII, 11–14 (7–12); VIII–IX, 14–25. Anus with 6–8 internal setae on ventral side and 3–5 on dorsal side. Pleural setae: I, 5–7 (0–2); II, 6–8 (0); III, 6–7 (0–2); IV, 6–7 (0–1); V, 5–7 (0–1); VI, 5–6 (0–1); VII, 4–5 (0); VIII, 3 (0). Genitalia and genital sac sclerite as in Fig. 9. Measurements as in Table 2.

114 · Zootaxa 4878 (1) © 2020 Magnolia Press lei et al. Type material: Holotype ♂, Badagongshan, Sangzhi County, Hunan Province, China, 24 Jul. 2013, D. Su & J. Ma. Bird J1531, GD-PHTH-00020 (GIABR). Paratypes: 8♂, 5♀, same data as holotype, GD-PHTH-00020, 00022, 00027–31 (GIABR). Non-types examined: 9 nymphs, same data as holotype, GD-PHTH-00021, 00023–26, 00029 (GIABR). Etymology. Myrsidea zhangae is named in honour of Dr Zhang Min (GIABR), good friend and colleague of the authors, who was instrumental in helping D.R.G. to settle in China. Remarks. The host of this louse species cannot be confirmed with certainty. As reported by Chu et al. (2019: 338), the host given in the collection data associated with specimens of Myrsidea sp. 7 (now M. zhangae) is Garrulax caerulatus (Hodgson, 1836). However, this bird species is restricted to western Yunnan and north Myanmar (Arlott 2017). It appears that the collectors of the lice misidentified the host bird, which was likely Ianthocincla berthemyi, a species that does occur in Hunan, and which is morphologically similar to Garrulax caerulatus. More collections of lice are needed from both bird species to establish the correct identity of the type host of M. zhangae.

Table 2. Measurements of Myrsidea attenuata and Myrsidea zhangae. Abbreviations: ANW = female anus width; AW = abdominal width (at segment IV); GL = male genitalia length; GSL = male genital sclerite length; HL = head length (at midline); MSPL = metasternal plate length (at midline); MSW = mesothorax width; MTW = metathorax width; POW = preocular width; PRW = prothorax width; PSPL = prosternal plate length (at midline); TL = total length (at midline); TW = head width (at temples). Post-spiracular setae are measured on one side of the abdomen only. Roman numbers refer to abdominal segments. Species Myrsidea attenuata Myrsidea zhangae Sex (No) Male (4) Female (3) Male (9) Female (5) TL 1.54–1.72 1.78–2.07 1.34–1.65 1.75–1.89 HL 0.31–0.34 0.33–0.34 0.28–0.34 0.32–0.34 POW 0.31–0.33 0.34 0.31–0.34 0.33–0.36 TW 0.47–0.51 0.49–0.53 0.45–0.49 0.49–0.53 PRW 0.33–0.35 0.36–0.37 0.29–0.34 0.31–0.38 PSPL 0.14–0.15 0.14–0.16 0.12–0.13 0.14–0.15 MSW 0.38–0.41 0.41–0.47 0.30–0.39 0.36–0.43 MTW 0.47–0.54 0.60–0.64 0.41–0.47 0.53–0.59 MSPL 0.18 0.23–0.24 0.151 0.22 1 AW 0.61–0.67 0.70–0.75 0.51–0.58 0.65–0.71 ANW - 0.27–0.31 - 0.25–0.28 GL -2 - 0.37–0.45 - GSL 0.11–0.13 - 0.056–0.075 - Postspiracular setae I 0.27–0.30 0.34–0.35 0.29–0.37 3 0.27–0.41 II 0.40–0.50 0.39–0.42 0.41–0.53 0.45–0.51 III 0.19–0.21 0.21–0.22 0.19–0.23 0.22–0.27 IV 0.40–0.51 0.45–0.50 0.38–0.50 0.30–0.55 V 0.13–0.15 4 0.12–0.15 0.07–0.15 0.13–0.18 VI 0.33–0.39 5 0.36–0.39 0.30–0.42 0.32–0.40 VII 0.37–0.41 0.39–0.41 0.34–0.43 0.29–0.40 VIII 0.36–0.51 0.48–0.50 0.39–0.43 0.40–0.46 1 Only measured in one specimen of each sex, because it is either obscured by gut contents or disrupted during mounting in all other specimens examined. 2 Anterior end of genitalia not clearly visible, and not measured. 3 Only present and unbroken in two specimens. 4 On one side of one specimen this seta is 0.32, but absent on the other side. 5 On one side of one specimen this seta is 0.24, but absent on the other side.

Myrsidea sp. 10; Chu et al. 2019: 338.

Type host: Lioparus chrysotis amoenus (Mayr, 1941) – golden-breasted fulvetta (Paradoxornithidae). Type locality: Ailaoshan, Jingdong County, Yunnan Province, China. Other hosts: Lioparus chrysotis swinhoii (Verreaux, 1871). Diagnosis. The male genital sclerite and lack of hypopharyngeal sclerite of M. liopari places this species close to M. bharat Tandan & Clay, 1971, and M. sabthai Tandan & Clay 1971. As female terga I–II are enlarged in M. liopari, this places this species closest to M. bharat in the key of Tandan & Clay (1971). These two species can be separated by the following characters: female tergum I with 10–11 setae in M. liopari (Fig. 16), but with only 2 setae in M. bharat; female terga IV–V with prominent gaps in the setal rows separating the 2 median setae from the other setae in M. liopari (Fig. 16), but with complete or near complete setal rows in M. bharat; tergum IX with 8–12 inner posterior setae in M. liopari (Fig. 16; see below), but with only 2 inner posterior setae in M. bharat. Males of M. liopari differ from those of M. sabthai by having fewer setae on most terga; there is some overlap on terga I–III, but terga IV–VI have a maximum of 12 setae each in M. liopari, whereas these terga have a minimum of 14 setae each in M. sabthai. Males of M. liopari may have 4 inner posterior setae on tergum IX (Fig. 15), whereas in M. bharat there appears to be only 2; however, some examined specimens of M. liopari have only 2 inner posterior setae in total, or have 1 on one side and 2 on the other. Description. Female (n=8). Habitus as in Fig. 16. Setal measurements: dhs10 = 0.030–0.056; dhs11 = 0.091– 0.115; ratio dhs10/11 = 0.27–0.53; ls5 = 0.013–0.022. Ventrolateral fringe with 8–10 setae. Gular plate with 4 setae on each side. Hypopharynx similar to Fig. 6. First tibia with 3 outer lateral ventral and 3 dorsal setae. Metanotum with posterior margin more or less straight and with 6–12 setae on posterior margin. Metasternal plate with 6–7 setae. Femoral brush with 13–17 setae. Metanotum and abdomen as in Fig. 16. Posterior margin of tergites straight except terga II–III which are arched as in Fig. 16. Setal rows with median gaps on all terga; terga IV–V also with prominent submedian (sensu Smith, 2000) gaps separating single median seta on each side from lateral setae. Ante- rior tergal setae absent. Tergal setae: I, 10–11; II, 15–17; III, 13–15; IV, 8–13; V, 8–13; VI, 11–13; VII, 11–13; VIII, 10–12; 8–12 inner posterior setae on tergum IX. Anterior sternal setae present on sterna III–VI. Sternal setae: I, 0; II, each aster of 4 setae, posterior margin with 11–15 setae and anteriorly with 8–11 setae; III, 19–24 (3–10); IV, 22–26 (14–23); V, 20–24 (16–23); VI, 18–22 (10–18); VII, 10–11 (7–11); VIII–IX, 17–29; Vulval margin deeply serrated, with 8–12 setae on margin. Anterior pleural setae absent. Pleural setae (on segments I–II thorn-like, but more median setae on ventral side slender; on segment III slender but of roughly same length as on segments I–II; on segments IV–VIII long and slender, similar to setae of sternal brushes): I, 5–8; II, 6–9; III, 6–8; IV, 5–7; V, 4–6; VI, 4–5; VII, 3–4; VIII, 3. Anus with 29–33 setae in ventral fringe and 20–33 setae in dorsal fringe. In median section about 1/3 of ventral setae much shorter than others (about 1/3 of length); all setae of dorsal fringe of roughly similar length. Measurements as in Table 3. Male (n=5). Much as in female, habitus as in Fig. 15, head shape as in Fig. 17. Setal measurements: dhs10 = 0.033–0.044; dhs11 = 0.083–0.109; ratio dhs10/11 = 0.38–0.49; ls5 = 0.012–0.018. Gular plate with 4 setae on each side. Hypopharynx as in Fig. 6. Metanotum with 6–8 setae on posterior margin. Femoral brush with 13–16 setae. Anterior setae absent on terga. Tergal setal rows with pronounced median gaps. Tergal setae: I, 6–8; II, 10–14; III, 11–14; IV, 7–12; V, 9–12; VI, 10–12; VII, 7–10; VIII, 6–7; 2–4 inner posterior setae present on tergum IX. Anterior setae present on sterna II–VII. Sternal setae: I, 0; II, each aster with 3–4, posterior margin with 11–12 setae, and anteriorly with 10–12 setae; III, 14–18 (0–6); IV, 16–22 (9–17); V, 19–20 (14–16); VI, 15–17 (11–15); VII, 9–14 (8–10); VIII–IX, 15–29. Anus with 7–9 internal setae on ventral side and 2–3 on dorsal side. Anterior pleural setae absent. Pleural setae: I, 4–6; II, 5–7; III, 6–7; IV, 5–6; V, 4–6; VI, 3–5; VII, 3; VIII, 3. Genitalia and genital sac sclerite as in Fig. 10. Measurements as in Table 3. Type material: Holotype ♂, Ailaoshan, elev. 2420 m, Jingdong County, Yunnan Province, China, 4 Sep. 2018, D.R, Gustafsson & L. Lei, bird J3745, GD-PHTH-00032 (GIABR). Paratypes: 2♀, same locality and collectors as holotype, 2 Sep. 2018, bird J3735, GD-PHTH-00035–00036 (GIABR). 1♂, 1♀, same locality and collectors as holotype, 29 Aug, 2018, bird J3698, GD-PHTH-00040–41 (GIABR). 2♂, 2♀, Laojunshan, Pingshan County, Sichuan Province, China, 10 Aug. 2014, Y. Zhao, Y. Zhang, X. Chu, W. Li, bird J2261, GD-PHTH-00042–00044 (GIABR). 1♀, same data except bird J2259, GD-PHTH-00042 (GIABR). 1♂, same data except bird J2266, GD-

FIGURE 15. Myrsidea liopari n. sp., male habitus, dorsal and ventral views.

Non-types examined: 2 nymphs, same data as holotype, GD-PHTH-00033–00034 (GIABR). 2 nymphs, same locality and collectors, 2 Sep. 2018, bird J3735, GD-PHTH-00037–38 (GIABR). 2 nymphs, same locality and collectors, 28 Aug. 2018, bird J3698, GD-PHTH-00041 (GIABR). 1 nymph, same locality and collectors, 2 Sep. 2018, bird J3731, GD-PHTH-00039 (GIABR). 1 nymph, Laojunshan, Pingshan County, Sichuan Province, China, 10 Aug. 2014, Y. Zhao, Y. Zhang, X. Chu, W. Li, bird J2261, GD-PHTH-00045 (GIABR). 3 nymphs, same data, except bird J2267, GD-PHTH-00048–00049 (GIABR). 1 nymph, same data except bird J2269, GD-PHTH-00051 (GIABR). Etymology. The specific epithet is derived from the genus of the type host. Remarks. The female head shape of Myrsidea liopari appears to differ among specimens, including those collected from the same host individual, with some of the females having a head shaped as that illustrated for the male (Fig. 15). We have illustrated a female with a rounder anterior head (Fig. 16) to highlight the morphological variation within this species. Specimens from Sichuan (host subspecies L. chrysotis swinhoii) differ from specimens from the type host subspecies by females having 10–12 inner posterior setae on tergum IX (8–9 in specimens from type host). There appears to be no correlation between head shape and locality. Moreover, other characters, including setal counts, are similar among specimens with different head shapes. More specimens and genetic data are needed to confirm whether these specimens represent cryptic species or natural variation in head shape. Since there are no significant morphological differences among specimens collected from the two host subspecies, all specimens are considered conspecific.

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 119 Table 3. Measurements of Myrsidea liopari and Myrsidea suthorae. Abbreviations: ANW = female anus width; AW = abdominal width (at segment IV); GL = male genitalia length; GSL = male genital sclerite length; HL = head length (at midline); MSPL = metasternal plate length (at midline); MSW = mesothorax width; MTW = metathorax width; POW = preocular width; PRW = prothorax width; PSPL = prosternal plate length (at midline); TL = total length (at midline); TW = head width (at temples). Post-spiracular setae are measured on one side of the abdomen only. Roman numbers refer to abdominal segments. Species Myrsidea liopari Myrsidea suthorae Sex (No) Male (5) Female (8) Male (3) TL 1.24–1.38 1.50–1.69 1.29–1.32 HL 0.26–0.29 0.28–0.32 0.26–0.28 POW 0.27–0.30 0.28–0.33 0.29 TW 0.36–0.38 0.39–0.43 0.39–0.42 PRW 0.24–0.28 0.26–0.29 0.27–0.29 PSPL 0.09–0.10 0.09–0.12 0.10–0.11 MSW 0.26–0.30 0.29–0.34 0.30–0.31 MTW 0.33–0.38 0.43–0.48 0.40 MSPL1 0.13 0.16 0.14 AW 0.45–0.52 0.56–0.67 0.49 ANW - 0.13–0.20 - GL 0.33–0.40 - 0.40–0.41 GSL 0.06–0.09 - 0.08 Postspiracular setae I 0.17–0.28 0.16–0.32 0.12–0.19 II 0.37–0.40 0.33–0.44 0.31–0.37 III 0.11–0.13 0.14–0.22 0.08–0.14 IV 0.35–0.40 0.33–0.41 0.21–0.39 V 0.12–0.14 0.14–0.18 0.10–0.14 VI 0.16–0.26 0.16–0.27 0.19–0.22 VII 0.23–0.31 0.21–0.34 0.28–0.30 VIII 0.34–0.41 0.37–0.43 0.40–0.46 1 Only measured in one specimen of each sex of each species, because it is either obscured by gut contents or disrupted during mounting in all other specimens examined.

Myrsidea suthorae Lei, Chu, Dik, Zou, Wang & Gustafsson, new species (Figs 7, 11, 18–19)

Myrsidea sp. 11; Chu et al. 2019: 338.

Type host: Suthora verreauxi verreuaxi Sharpe, 1883 – golden parrotbill (Paradoxornithidae). Type locality: Laojunshan, Pingshan County, Sichuan Province, China. Diagnosis. Males of Myrsidea suthorae key to couplet 6 in the key of Tandan (1972), but it does not fit any of the two given options. One is M. singularis, but M. suthorae differs by the shape of its unique genital sclerite, and by a proportionately broader genitalia than M. singularis (see Fig. 11 and fig. 53 in Tandan 1972); however, the abdominal chaetotaxy of these two species is very similar and may entirely overlap. The second option in couplet 6 leads to M. sikkimensis Tandan, 1972, M. sehri Ansari, 1951 and M. erythrocephali; however, these three species have no more than 4 setae on tergum I, whereas M. suthorae has at least 8. Moreover, M. suthorae can be separated from all these species by the presence of 2 setae on sternite II in M. suthorae (absent in the other species), and the presence in M. suthorae of 8–9 inner posterior setae on tergum IX (only 2 inner posterior setae in the other species).

Description. Male (n=3). Habitus as in Fig. 18 and head shape as in Fig. 19. Setal measurements: dhs10 = 0.044–0.056; dhs11 = 0.081–0.104; ratio dhs10/11 = 0.44–0.64; ls5 = 0.045–0.062. Ventrolateral fringe with 9–10 setae. Gular plate with 5–6 setae on each side. Hypopharynx reduced as in Fig. 7. First tibia with 3 outer lateral ventral and 2–3 dorsal setae. Metanotum with posterior margin straight, with 8–9 setae on posterior margin. Metasternal plate with 7 setae. Femoral brush with 13–15 setae. Metanotum and abdomen as in Fig. 18. Posterior margin of tergites straight, with setal rows interrupted by median gap in at least segments III–IV. Anterior tergal setae absent. Tergal setae: I, 8–9; II, 10–11; III, 10–12; IV, 10–11; V, 10–13; VI, 10–12; VII, 10–12; VIII, 6–8; 8–9 inner posterior setae present on segment IX. Anterior sternal setae present on sterna II–VI. Sternal setae: I, 2; II, each aster of 3 setae, posterior margin with 11–12 setae, and anteriorly with 6 setae; III, 14 (0–2); IV, 18–21 (15–19); V, 21–22 (16–20); VI, 20–21 (16–19); VII, 13–15 (16–22); VIII–IX, 29–33. Anus with 7–9 internal setae on ventral side and 3–4 on dorsal side. Anterior pleural setae absent. More median pleural setae on ventral side very long and slender on segments V–VII, similar to or longer than setae of setal brushes on same segments. Pleural setae: I, 4–5; II, 5–6; III, 6–7; IV, 6; V, 5–6; VI, 4–6; VII, 3–4; VIII, 3. Genitalia and genital sac sclerite as in Fig. 11. Measurements as in Table 3. Female. Unknown. Type material: Holotype ♂, Laojunshan, Pingshan County, Sichuan Province, China, 10 Aug. 2014, Y. Zhao, Y. Zhang, X. Chu, W. Li, bird J2252, GD-PHTH-00055 (GIABR). Paratypes: 1♂, same data as holotype, GD- PHTH-00054 (GIABR). 1♂, same locality and collector as holotype, 7 Aug. 2014, bird J2195, GD-PHTH-00053 (GIABR). Non-type specimens: 3 nymphs, same data as holotype, GD-PHTH-00054–55 (GIABR). Etymology. The specific epithet is derived from the genus of the type host.

A large number of babbler species are found in China (MacKinnon & Phillipps 2016). The chewing lice parasitising these hosts are generally poorly known, with only 28 species of Myrsidea described from babblers worldwide (Tandan & Clay 1971; Tandan 1972; Rai 1978; Price et al. 2006; Khan et al. 2009; Najer et al. 2012). Only five species have been reported from China (Price et al. 2006; Chu et al. 2019) and four more are described in this pa- per, including the first species from hosts of the family Paradoxornithidae. However, this quantity represents only a small part of the true diversity of Myrsidea lice living on Chinese babblers, and additional species are likely to be recorded and described. In Appendix 1, we have summarised the known distribution of Myrsidea species parasitising babblers worldwide. Some species of Myrsidea can be grouped together based on the structure and shape of their genital sclerites. Among the Myrsidea parasitising babblers, species with similar genital sclerites often parasitise closely related hosts. For example, species of Myrsidea with M-shaped genital sclerite (Figs 8–9) mainly parasitise hosts in the Leiothrichidae, particularly the genera Garrulax and Ianthocincla, which form a monophyletic clade in the phylogeny of Cai et al. (2019). However, some Myrsidea species with M-shaped genital sclerite are also found on hosts of the genus Trochalopteron, which is more distantly related to Garrulax and Ianthocincla (Cai et al. 2019). The species included in Garrulax, Ianthocincla and Trochalopteron all have large bodies, which would suggest that host size may also influence host distribution patterns in Myrsidea. By contrast, other types of genital sclerites (e.g. Fig. 10) are more widely distributed among species of Myrsidea from babblers in the families Leiothrichidae, Paradoxornithidae and Timaliidae (e.g. Myrsidea chilchil Ansari, 1951 and Myrsidea satbhai Ansari, 1951, see Tandan & Clay 1971). Furthermore, some types of genital sclerites (e.g. that of Myrsidea falcatae Price et al., 2006) are remarkably similar to those found in Myrsidea parasitising non-babbler hosts (e.g. thrushes, Turdidae; see Clay 1969). This would suggest that host switching of lice among hosts of different families may have played a role in the evolutionary history of Myrsidea parasitising babblers. More research is needed before the significance of the association between different types of genital sclerites in species of Myrsidea parasitising babblers and their host distribution can be assessed adequately, including analysis of molecular data. The finding of a previously unknown type of genital sclerite in Myrsidea suthorae (Fig. 11) sug- gests that much remains to be discovered in this genus of lice.

Acknowledgements

This study was carried out in strict accordance with the Regulation for the Administration of Laboratory Animals (Decree No. 2, State Science and Technology Commission of the People’s Republic of China, Nov. 14 1988). We obtained approval for this study from the Guangdong Institute of Applied Biological Resources Administrative pan- el on Laboratory Animal Care. This study was supported by the Introduction of Full-Time High-Level Talent Fund of the Guangdong Academy of Sciences grant 2018GDASCX-0809 and GIABR-GJRC201701 to DRG and FZ, and Guangdong Forestry Special Project Grant 1210-1741YDZB0401, GDAS Special Project of Science and Technol- ogy Development 2018GDASCX-0107 and National Natural Science Foundation of China Grant 31961123003 to FZ. These agencies had no hand in the design or execution of this study, and we declare no conflict of interest. We would like to thank two anonymous reviewers for helpful comments on this manuscript.

References

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Appendix 1. Checklist of host associations of Myrsidea species known from “babblers” (Leiothrichidae, Para- doxornithidae, Pellorneidae, Timaliidae and Yuhina sensu lato) arranged alphabetically by host families and genera.

Unidentified species of Myrsidea are included for completeness, with Remarks. Unidentified species reported by Chu et al. (2019) and Gustafsson et al. (2019), which are described herein are not listed separately. References to host records for species described prior to 2003 are included only for those hosts not listed by Price et al. (2003). Host taxonomy follows Clements et al. (2018). Host species Louse species Remarks Leiothrichidae Actinodura cyanouroptera (Hodgson, 1837) Myrsidea sp. [1] [2] Actinodura ramsayi Walden, 1875 Myrsidea sp. [1] Actinodura strigula (Hodgson, 1837) Myrsidea sp. [1] Actinodura strigula yunnanensis (Rothschild, 1921) Myrsidea sp. [3] Alcippe brunneicauda (Salvadori, 1879) Myrsidea sp. [1] Alcippe davidi Styan, 1896 Myrsidea cheni Price et al., 2006 [2] Alcippe fratercula yunnanensis Harington, 1913 Myrsidea sp. [3] Alcippe morrisonia Swinhoe, 1863 Myrsidea cheni Price et al., 2006 [4] Myrsidea sp. [1] [5] Alcippe nipalensis (Hodgson, 1837) Myrsidea sp. [2] Alcippe poioicephala (Jerdon, 1841) Myrsidea cheni Price et al., 2006 Myrsidea sp. [1] Garrulax leucolophus (Hardwicke, 1816) Myrsidea assamensis Tandan, 1972 Myrsidea patkaiensis Tandan, 1972 Myrsidea sp. [1] Garrulax leucolophus belangeri Lesson, 1831 Myrsidea assamensis Tandan, 1972 [Ref: Tandan, 1972: 391] Garrulax leucolophus diardi (Lesson, 1831) Myrsidea assamensis Tandan, 1972 [Ref: Tandan, 1972: 391] Garrulax leucolophus leucolophus (Hardwicke, 1816) Myrsidea assamensis Tandan, 1972 Garrulax leucolophus patkaicus Reichenow, 1913 Myrsidea patkaiensis Tandan, 1972 Garrulax maesi maesi (Oustalet, 1890) Myrsidea attenuata new species ...... continued on the next page

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 125 APPENDIX 1. (Continued) Host species Louse species Remarks Myrsidea sp. [2] Garrulax merulinus Blyth, 1851 Myrsidea thailandensis Tandan, 1972 Myrsidea sp. [1] Garrulax monileger fuscatus Baker, 1918 Myrsidea monilegeri Tandan, 1972 Garrulax monileger schauenseei Delacour & Greenway, Myrsidea sp. [6] 1939 Garrulax strepitans Blyth, 1855 Myrsidea sp. [1] Grammatoptila striata (Vigors, 1831) Myrsidea sp. [1] Grammatoptila striata sikkimensis (Ticehurst, 1924) Myrsidea sikkimensis Tandan, 1972 Heterophasia capistrata (Vigors, 1831) Myrsidea sp. [1] Heterophasia desgodinsi (Oustalet, 1877) Myrsidea sp. [2] [3] Heterophasia melanoleuca (Blyth, 1859) Myrsidea sp. [1] Ianthocincla albogularis (Gould, 1836) Myrsidea macraidoia Tandan, 1972 Ianthocincla albogularis albogularis (Gould, 1836) Myrsidea macraidoia Tandan, 1972 Ianthocincla albogularis whistleri Baker, 1921 Myrsidea macraidoia Tandan, 1972 Ianthocincla berthemyi (Oustalet, 1876) Myrsidea zhangae new species Ianthocincla chinensis (Scopoli, 1786) Myrsidea patkaiensis Tandan, 1972 Myrsidea sp. [1] Ianthocincla chinensis lochmius Deignan, 1941 Myrsidea patkaiensis Tandan, 1972 Ianthocincla mitrata (S. Müller, 1836) Myrsidea patkaiensis Tandan, 1972 Ianthocincla pectoralis (Gould, 1836) Myrsidea orientalis Tandan, 1972 Ianthocincla pectoralis pectoralis (Gould, 1836) Myrsidea orientalis Tandan, 1972 Ianthocincla pectoralis subfusa Kinnear, 1924 Myrsidea orientalis Tandan, 1972 Ianthocincla ruficollis (Jardine & Selby, 1838) Myrsidea bhutanensis Tandan, 1972 Myrsidea sp. [1] Leiothrix argentauris (Hodgson, 1836) Myrsidea argentauris Price et al., 2006 Myrsidea sp. [1] Leiothrix lutea (Scopoli, 1786) Myrsidea sp. [1] Liocichla phoenicea bakeri (Hartert, 1908) Myrsidea ananthakrishnani Rai, 1978 Liocichla ripponi (Oates, 1900) Myrsidea ananthakrishnani Rai, 1978 [2] Myrsidea sp. [1] Liocichla steerii Swinhoe, 1877 Myrsidea sp. [1] Minla ignotincta mariae La Touche, 1921 Myrsidea sp. [3] Trochalopteron erythrocephalum (Vigors, 1832) Myrsidea erythrocephali Tandan, 1972 Myrsidea sp. [1] Trochalopteron lineatum lineatum (Vigors, 1831) Myrsidea sehri Ansari, 1951 Myrsidea agarwali Khan et al., 2009 Trochalopteron subunicolor subunicolor Blyth, 1843 Myrsidea singularis Tandan, 1972 Trochalopteron squamatum (Gould, 1835) Myrsidea manipurensis Tandan, 1972 Turdoides bicolor (Jardine, 1831) Myrsidea sp. [7] Turdoides caudata (Dumont, 1823) Myrsidea sp. [1] Turdoides caudata caudata (Dumont, 1823) Myrsidea chilchil Ansari, 1951 Turdoides earlei (Blyth, 1844) Myrsidea salimalii Tandan & Clay, 1971 ...... continued on the next page

126 · Zootaxa 4878 (1) © 2020 Magnolia Press lei et al. APPENDIX 1. (Continued) Host species Louse species Remarks Turdoides fulva acaciae (Lichtenstein, 1823) Myrsidea meinertzhageni Tandan & Clay, 1971 Turdoides fulva maroccana Lynes, 1925 Myrsidea meinertzhageni Tandan & Clay, 1971 Turdoides jardineii (A. Smith, 1836) Myrsidea breviterga Tandan & Clay, 1971 Turdoides malcolmi (Sykes, 1832) Myrsidea bharat Tandan & Clay, 1971 Myrsidea sp. [1] Turdoides plebejus platycirca (Swainson, 1837) Myrsidea clamosae Tandan & Clay, 1971 Turdoides sharpei sharpei (Reichenow, 1891) Myrsidea clamosae Tandan & Clay, 1971 Turdoides sharpei vepres Meinertzhagen, 1937 Myrsidea satbhai Ansari, 1951 Turdoides squamiceps (Cretzschmar, 1827) Myrsidea chilchil Ansari, 1951 Turdoides squamiceps squamiceps (Cretzschmar, 1827) Myrsidea chilchil Ansari, 1951 [Ref: Tandan & Clay, 1971: 222] Turdoides squamiceps yemensis (Neumann, 1904) Myrsidea chilchil Ansari, 1951 [Ref: Tandan & Clay, 1971: 222] Turdoides striata sindiana (Ticehurst, 1920) Myrsidea satbhai Ansari, 1951 [Ref: Tandan & Clay, 1971: 216] Turdoides striata somervillei (Sykes, 1832) Myrsidea satbhai Ansari, 1951 [Ref: Tandan & Clay, 1971: 216] Turdoides striata striata (Dumont, 1823) Myrsidea satbhai Ansari, 1951

Paradoxornithidae Fulvetta manipurensis tonkinensis Myrsidea sp. [3] (Delacour & Jabouille, 1930) Lioparus chrysotis amoenus (Mayr, 1941) Myrsidea liopari new species Lioparus chrysotis swinhoii (Verreaux, 1871) Myrsidea liopari new species Myzornis pyrrhoura Blyth, 1843 Myrsidea sp. [1] Psittiparus gularis (Gray, 1845) Myrsidea sp. [1] Sinosuthora webbiana (Gould, 1852) Myrsidea sp. [1] Suthora nipalensis (Hodgson, 1837) Myrsidea sp. [1] Suthora nipalensis poliotis Blyth, 1841 Myrsidea sp. [3] Suthora verreauxi verreauxi Sharpe, 1883 Myrsidea suthorae new species

Pellorneidae Gampsorhynchus rufulus Blyth, 1844 Myrsidea sp. [1] Malacopteron cinereum Eyton, 1839 Myrsidea sp. [1] Pellorneum albiventre (Goodwin-Austen, 1877) Myrsidea sp. [1] Pellorneum capistratum (Temminck, 1823) Myrsidea sp. [1] Pellorneum malaccense (Hartlaub, 1844) Myrsidea sp. [1] Pellorneum ruficeps Swainson, 1832 Myrsidea sp. [1] [8] Pellorneum tickelli Blyth, 1859 Myrsidea sp. [1] Ptilocichla falcata Sharpe, 1877 Myrsidea falcatae Price et al., 2006 Schoeniparus castaneceps (Hodgson, 1837 Myrsidea sp. [1] Schoeniparus castaneceps exul Delacour, 1932 Myrsidea sp. [3] ...... continued on the next page

New species of Myrsidea from babblers Zootaxa 4878 (1) © 2020 Magnolia Press · 127 APPENDIX 1. (Continued) Host species Louse species Remarks Schoeniparus dubius (Hume, 1874) Myrsidea sp. [2] Schoeniparus dubius dubius (Hume, 1874) Myrsidea sp. [3] Schoeniparus rufogularis (Mandelli, 1873) Myrsidea annae Najer & Sychra [in Najer et al.], 2012 Turdinus abbotti (Blyth, 1845) Myrsidea sp. [1] Turdinus brevicaudatus (Blyth, 1855) Myrsidea sp. [1]

Timaliidae Cyanoderma chrysaeum (Blyth, 1844) Myrsidea sp. [1] Cyanoderma erythropterum (Blyth, 1842) Myrsidea macronoi Price et al., 2006 Myrsidea sp. [1] Cyanoderma ruficeps (Blyth, 1843) Myrsidea sp. [1] [2] Cyanoderma ruficeps davidi Oustalet, 1899 Myrsidea sp. [3] Cyanoderma rufifrons (Hume, 1873) Myrsidea sp. [1] Mixornis gularis (Horsfield, 1822) Myrsidea macronoi Price et al., 2006 Myrsidea sp. [1] Pomatorhinus erythrogenys Vigors, 1832 Myrsidea sp. [1] [6] Pomatorhinus ferruginosus Blyth, 1845 Myrsidea sp. [1] [6] Pomatorhinus schisticeps Hodgson, 1836 Myrsidea duplicata Tandan, 1972 Myrsidea sp. [1] Stachyris maculata (Temminck, 1836) Myrsidea macronoi Price et al., 2006 Myrsidea sp. [1] Stachyris nigriceps coltarti Harington, 1913 Myrsidea sp. [1] [5] Stachyris nigricollis (Temminck, 1836) Myrsidea macronoi Price et al., 2006 Myrsidea sp. [1] Stachyris poliocephala (Temminck, 1836) Myrsidea sp. [1] Stachyris strialata (S. Müller, 1836) Myrsidea dukguni Price et al., 2006

Zosteropidae Yuhina gularis omeiensis Riley, 1930 Myrsidea sp. [3] [1] Unidentified species of Myrsidea reported by McClure et al. (1973). In many cases, the specimens reported by McClure et al. (1973) are likely conspecific with species of Myrsidea described subsequently from the same host. For instance, M. falcatae was described by Price et al. (2006) from specimens collected by McClure et al. (1973). As we have not seen any of the specimens reported by McClure et al. (1973), we list their unidentified species separately, because we cannot establish whether they represent known or unknown species. [2] Chu et al. (2019). [3] Gustafsson et al. (2019). [4] Alcippe morrisonia was given as the type host of this species by Price et al. (2006), but this host species is now considered to be endemic to Taiwan, with other species in the same complex occurring on the mainland (Zou et al. 2007; Song et al. 2009; Clements et al. 2018). [5] Najer et al. (2012). [6] Tandan (1972). [7] Tandan & Clay (1971). [8] Najer et al. (2014).