Dtsch. Entomol. Z. 71 (1) 2024, 85-109 | DOI 10.3897/dez.71.111874 yee BERLIN New genus and species of lice in the Oxylipeurus-complex (Phthiraptera, Ischnocera, Philopteridae), with an overview of the distribution of ischnoceran chewing lice on galliform hosts Daniel R. Gustafsson’, Chunpo Tian! *, Mengjiao Ren‘, Zhu Li’, Xiuling Sun*, Fasheng Zou! 1 Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Library of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, 105 Xingang West Road, Haizhu District, Guangzhou, 510260, Guangdong Province, China 2 College of Life Sciences, Shaanxi Normal University, 620 West Chang’an Street, Chang'an District, Xi’an City, 710119, Shaanxi Province, China 3 Department of Life Sciences, National Natural History Museum of China, 126, Tiangiao South St. Dongcheng District, Beijing, 100050, China 4 Collections Department, National Natural History Museum of China, 126, Tiangiao South St. Dongcheng District, Beijing, 100050, China https://zoobank. org/AF'C4DD5 7-276 1-424D-B766-FE9696EF F040 Corresponding author: Daniel R. Gustafsson (kotatsu@fripost.org) Academic editor: Viktor Hartung # Received 30 August 2023 # Accepted 31 January 2024 # Published 12 March 2024 Abstract Here, we describe a new genus of lice (Phthiraptera, Ischnocera) in the Oxylipeurus-complex, parasitising galliform hosts in the genera 7ragopan Cuvier, 1829. This genus, Pelecolipeurus gen. nov., is separated from other members of the complex by the unique shape of the male subgenital plate and stylus, the male genitalia and other characters. The only previously-known species in the genus 1s Lipeurus longus Piaget, 1880, which is here tentatively re-described as Pelecolipeurus longus (Piaget, 1880), based on specimens from a non-type host, 7ragopan temminckii (Gray, 1831). In addition, we describe a new species, Pelecolipeurus fujianensis sp. nov., based on specimens from 7ragopan caboti (Gould, 1857). An overview of the distribu- tion patterns of ischnoceran lice on galliforms is presented, which suggests that host phylogeny, host biogeography and host biotope, as well as elevation of host range, may all be important factors that have structured louse communities on landfowl. We transfer the genus Afrilipeurus from the Oxylipeurus-complex to the Lipeurus-complex and include an emended key to the Oxylipeurus-complex. Key Words chewing lice, Galliformes, new genus, Oxylipeurus-complex, Phthiraptera Introduction Chewing lice (Phthiraptera) in the Oxylipeurus-complex mainly parasitise gamefowl (Galliformes; Price et al. (2003)) and most species are known from Asian galli- forms. Traditionally, most of the species have been placed in the one genus, Oxylipeurus (e.g. Clay (1938a); Hop- kins and Clay (1952); Price et al. (2003)). However, this classification was challenged by, for example, von Kéler (1958) and Carriker (1967), who considered several groups of Oxylipeurus to be sufficiently distinct to form separate genera. Mey (2009) considered several of these genera valid and, since then, a large number of new ge- nus-level taxa within this complex have been established (Gustafsson and Zou 2020a, b, 2023; Gustafsson et al. 2020a, b). Assessing taxon limits in this complex is difficult, as the overall chaetotaxy and morphology, including that of the male genitalia, are conserved in many genera and species are often delimited by more nebulous characters, Copyright Daniel R. Gustafsson et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 86 such as head shape, mesosome shape and degree of re- ticulation of the cuticle (e.g. Gustafsson et al. (2020a)). Moreover, many species are poorly known and have not been fully described or illustrated; the last detailed revi- sions of the complex were published by Clay (1938a) and von Kéler (1958). Gustafsson et al. (2020a) tentatively considered Lipeurus longus Piaget, 1880, to belong to the genus Reticulipeurus Kéler, 1958, based on its placement by Clay (1938a), von Kéler (1958) and Zlotorzycka (1966); however, they stated that they had not examined any specimens and that some aspects of the morphology of this species were aberrant for Reticulipeurus. Here, we describe this group as a separate genus, Pelecoli- peurus gen. nov., based on specimens examined from two hosts in China and examination of photos and il- lustrations of Lipeurus longus Piaget, 1880. We tenta- tively re-describe the only previously-known species (L. longus) and add a second species, Pelecolipeurus fujianensis sp. nov. Given that this new genus is the third Oxylipeu- rus-complex genus to be described in recent years from the same host group, we also take this opportuni- ty to summarise what is known about host-associations amongst ischnoceran lice parasitising galliform hosts. Finally, we update the key to the genera of the Oxyli- peurus-complex previously published by Gustafsson et al. (2020b). Materials and methods Previously, slide-mounted specimens deposited at the National Natural Museum of Natural History, Chi- na (NNHM) were examined with a Nikon Eclipse Ni (Nikon Corporation, Tokyo, Japan), with a drawing tube attached for making illustrations. Drawings were scanned, then compiled and edited in GIMP 2.10 (www. gimp.org). Measurements (all in mm) were made from slide-mounted specimens in the digital measuring soft- ware ImageJ 1.48v (Wayne Rasband; imagej.net): AW = abdominal width (at segment V); HL = head length (at mid-line); HW = head width (at widest point of tem- ples); PRW = prothoracic width; PTW = pterothoracic width; TL = total length (at mid-line). Host taxonomy follows Clements et al. (2022). Ter- minology for chaetotaxy and other structures of the lice follows Clay (1951), Mey (1994) Gustafsson and Bush (2017) and Gustafsson et al. (2020a). Abbreviations used in the text follow Gustafsson and Bush (2017) and Gustafsson et al. (2020a) and include: mds = man- dibular seta; mms = marginal mesometathoracic setae; mths = metathoracic thorn-like seta; mtrs = metathorac- ic trichoid seta; mts/—3 = marginal temporal setae 1-3; os = ocular seta; pos = preocular seta; ps = paratergal seta; pstl—2 = parameral setae 1—2; s/—8 = sensilla 1—8 of dor- sal head; sts = sternal seta; vms = vulval marginal setae; vss = vulval submarginal setae. dez.pensoft.net Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Systematics PHTHIRAPTERA Haeckel, 1896 Phthiraptera Haeckel, 1896: 703. Ischnocera Kellogg, 1896 Ischnocera Kellogg, 1896: 63. Philopteridae Burmeister, 1838 Philopteridae Burmeister, 1838: 422. Oxylipeurus-complex Included genera: Calidolipeurus Gustafsson et al., 2020b: 2. Cataphractomimus Gustafsson et al., 2020a: 206. Chelopistes Kéler, 1940: 180. Virgula Clay, 1941: 119. Eiconolipeurus Carriker, 1945: 91. Epicolinus Carriker, 1945: 104. Gallancyra Gustafsson & Zou, 2020a: 11. Megalipeurus Kéler, 1958: 327. Oxylipeurus Mjoberg, 1910: 91. Pelecolipeurus gen. nov. Reticulipeurus Kéler, 1958: 332. Subgenus: Reticulipeurus (Forcipurellus) Gus- tafsson & Zou, 2023:497. Subgenus: Reticulipeurus (Reticulipeurus) Kéler, 1958: 332. Sinolipeurus Gustafsson et al., 2020a: 229. Splendoroffula Clay & Meinertzhagen, 1941: 343. Splendopeurus Kéler, 1958: 309. Talegallipeurus Mey, 1982: 242. Trichodomedea Carriker, 1946: 365. Valimia Gustafsson & Zou, 2020b: 490. Pelecolipeurus gen. nov. https://zoobank.org/FBCCEB4D-7E49-4BFE-88D7-A7 1559102743 Type species. Pelecolipeurus fujianensis sp. nov. Diagnosis. Pelecolipeurus gen. nov. keys to Retic- ulipeurus Kéler, 1958, in the key of Gustafsson et al. (2020b). Species of Pelecolipeurus can be separated from Reticulipeurus and all other members of the Oxy/i- peurus-complex by the following combination of char- acters: frons rounded to slightly flattened (Figs 3, 17); dorsal pre-antennal suture present, transversal, but not reaching lateral margins of head (Figs 3, 17); mms gath- ered into a single sublateral bunch (Figs 1, 2, 15, 16); male tergopleurites H—VII medianly interrupted and in- tertergal sclerites absent (Figs 1, 15); male tergopleurites IX—XI fused to form single plate (Figs 1, 15); female Dtsch. Entomol. Z. 71 (1) 2024, 85-109 tergopleurites [X—XI fused laterally, but not medianly, forming two distinct plates (Figs 2, 16); male subgenital plate of unique shape, with lateral extensions at base of stylus (Figs 7, 21); stylus subterminal, elongated to reach beyond distal margin of abdomen (Figs 7, 21); female vulval margin narrowly concave, without lateral acces- sory vulval plates (Figs 8, 22); male genitalia very long, reaching anteriorly to abdominal segment III (Figs 5, 19); denticulate genital sac present in male genitalia (Figs 5, 19); male genitalia symmetrical, with parameres present, mesosome dominated by large gonopore (Figs 6, 20). Description. Both sexes. Male longer than female (Ta- ble 1). Head longer than wide, frons rounded to slightly flattened (Figs 3, 17). Dorsal pre-antennal suture present, but often not well-defined and visible as pale band across head; suture not reaching lateral margins of head. Interior thickenings of pre-antennal head present as double, un- dulating carinae anterior to suture. Head chaetotaxy as in Figs 3, 17; mds may be absent in female; s2 (?) located median to s/; s5 absent; s6—8 present; mts3 only tempo- ral mesoseta, but os may be longer than pos and mts/—2 in males. Antennae sexually dimorphic, with male scape and pedicel elongated and swollen compared to female (cf. Figs 3, 4, 17, 18); male flagellomere I with distal, finger-like extension and intensely scaly inner surface (Figs 3, 17). Temples rounded, somewhat bulging. Tho- racic and abdominal segments as in Figs 1, 2, 15, 16. Legs II and III much longer than legs I; coxae I-II close to- gether. Meso- and metasterna fused. Metepisternum long, reaching almost to mesometasternum. Pronotum with lateral and posterior setae; pteronotum with microsetae in antero-lateral corners and short seta submedianly in distal half; mms in single sublateral bunch; mths and mtrs roughly dorsal. Tergopleurites II-VI in both sexes medi- anly interrupted; male tergopleurites IX—XI fused into a medianly continuous plate; female tergopleurites [X—XI fused laterally, but not medianly (Figs 2, 16). Male subge- nital plate with lateral extensions in distal section (Figs 7, 15); stylus slender, elongated, tapering, attached subter- minally and extending beyond distal margin of abdomen. Female subgenital plates reduced to near vulval margin; exact extent of these often not clearly visible. Leg chae- totaxy as in Figs 9-14. Male. Male scape, pedicel and flagellomere I modi- fied compared to female. Male genitalia very long (Figs 5, 19), with basal apodeme reaching to at least abdominal segment HI, but diffuse anteriorly. Genital sac present, irregularly, but densely denticulate (Figs 6, 20). Distal third of basal apodeme with irregularly thickened later- al margins articulating with parameral heads. Mesosome simple, with central sclerite on ventral surface associat- ed with 2-3 sensilla; three additional sensilla in oblique, distally divergent rows lateral to this sclerite. Gonopore large, dominating mesosome. Parameres short, slender, pst! sensilla in distal third, pst2 microsetae, situated more or less apically. Female. Vulval margin deeply and narrowly concave (Figs 8, 22). Three sets of genital setae: long, slender vms, 87 the more median setae shorter than the more lateral setae; short, slender or lightly stout vss in median part of vulval margin; single seta on each side situated further submar- ginally and apart from vss. Subvulval sclerites present, slender and elongated, reaching to vulval margin. Host distribution. Presently known only from trago- pans (genus 7ragopan Cuvier, 1829), Phasianidae, Gal- liformes. Some specimens from other hosts (see below) may represent stragglers or contaminations. Geographical range. All known species are from Chi- na or the Himalayas, corresponding roughly to the com- bined range of the known hosts. Etymology. The name Pe/ecolipeurus is derived from “pélekus’, Greek for “two-headed axe” and the tradition- al name for long slender lice, Lipeurus Nitzsch, 1818. This refers to the shape of the male subgenital plate. Remarks. Gustafsson et al. (2020a) tentatively in- cluded Lipeurus longus Piaget, 1880, in Reticulipeurus Kéler, 1958, following von Kéler (1958) and Zitotorzy- cka (1966). They noted that they had not examined any specimens and that this placement was doubtful, based on the illustrations published by Clay (1938a) and von Kéler (1958). The examined collection at NNHM in- cludes two different species belonging to the same mor- phological group as L. /ongus and these are sufficiently different morphologically from all other members of the Oxylipeurus-complex that the erection of a separate ge- nus is warranted. Unfortunately, no specimens from the type host of Li- peurus longus were found at NNHM and no specimens of this species have been examined from other collec- tions. A lectotype and five paratypes are available at the Natural History Museum, London (NHML), but we had no opportunity to examine or borrow these. A photo of the lectotype female at the NHML homepage (https:// data.nhm.ac.uk/dataset) confirms that this species be- longs to Pelecolipeurus, but is insufficiently detailed to compare adequately with the specimens we have ex- amined at the NNHM. Only two modern illustrations of L. longus have been published (Clay 1938a; von Kéler 1958), both of which depict the ventral view of the distal end of the male abdomen. Allowing for individ- ual variation and differences in illustration techniques, we cannot separate the specimens illustrated in these publications from specimens we have seen from 7ra- gopan temminckii (Gray, 1831) (see below) and these specimens are here tentatively considered conspecific with L. Jongus; however, this will need to be confirmed by comparison with type specimens of L. /Jongus and a re-description of this species. As the type specimens of L. /ongus could not be exam- ined, we select the species that could be examined as the type species of Pelecolipeurus. Included species. Pelecolipeurus fujianensis sp. nov. Type host: Tragopan caboti (Gould, 1857). Pelecolipeurus longus (Piaget, 1880: 370) [in Lipeurus]. Type host: Tragopan satyra (Linnaeus, 1758). dez.pensoft.net 88 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Table 1. Measurements of the species of Pelecolipeurus. Measurements (all in mm) were made in the digital measuring software ImageJ 1.48v (Wayne Rasband; imagej.net): AW = abdominal width (at segment V); HL = head length (at mid-line); HW = head width (at widest point of temples); PRW = prothoracic width; PTW = pterothoracic width; TL = total length (at mid-line). Species Host Sex N TL Pelecolipeurus Tragopan caboti fujianensis Pelecolipeurus longus Tragopan temminckii M_ 6 3.56—4.40 s. lat. F 'N = 15 for TL; N = 18 for AW. ?'N = 24 for TL; N = 29 for PTW; N = 38 for AW. Pelecolipeurus fujianensis sp. nov. https://zoobank.org/1 CEB3DA A-D063-4616-A4A7-703CE2B2544F Figs 1-14 Type host. Zragopan caboti (Gould, 1857) — Cabot’s tra- gopan. Type locality. Fujian Province, China. Specimens examined. Type material. Ex 7Zrago- pan caboti: CHINA * Holotype 3; Fujian Province; 29 Sep 1990: collector unknown; box E0026199, slide 65 (NNHM) [Male in lower right corner, near where cover glass is broken, marked with black dot on slide]. Paratypes 73,92, 8 nymphs; Fujian Province; 29 Sep 1990; collec- tor unknown; box E0026199, slides 64-66, 95 (NNHM). 14, 39; Fujian Province; 16 Dec 1988; collector un- known; box E0026199, slide 68 (NNHM). 192, 3 nymphs; Fujian Province, Jianou; 7 Jan 1997; collector unknown; box E0026195, slide 3 (NNHM). 1¢', 29, 6 nymphs; Fu- jian Province, Wuyi Mountain; Dec. 1989; collector un- known; box E0026011, slide 15, box E0026198, slide 74 (NNHM). 116, 152, 11 nymphs; Zhejiang Province; 8 Dec 1980; collector unknown; box E0026010, slide 76, box E0026199, slides 88—92 (NNHM). Diagnosis. Due to the limited illustrations published for Pelecolipeurus longus from the type host (see above), we here compare P. fujianensis sp. nov. with the spec- imens tentatively identified as P Jongus from T. tem- minckii, which we consider conspecific with the species illustrated by Clay (1938a) and von Kéler (1958). A re-description of P. Jongus from the type host is necessary to determine additional characters separating this species from P. fujianensis. Pelecolipeurus fujianensis can be separated from P. longus as illustrated by Clay (1938a) and von Keéler (1958) by the following characters: male fused abdom- inal segment [X—XI with more or less straight lateral margins in P. /Jongus, but with concave lateral margins in P. fujianensis (Fig. 7); proximal mesosome of P. /ongus with flattened anterior margin, but with medianly point- ed anterior end in P. fujianensis (Fig. 6); parameres more curved in P. Jongus than in P. fujianensis (Fig. 6). In addition, P. fujianensis can be separated from the population from 7: temminckii described above by the fol- lowing characters: frons more flattened in P. /ongus s. lat. (Fig. 17) than in P. fujianensis (Fig. 3); male sternal plate VI with 2 sts of more or less equal length in P. /Jongus s. lat. (Fig. 15), but with lateral seta on each side much shorter than median seta on each side in P. fujianensis dez.pensoft.net HL HW M 20! 4.00-4.41 (4.20) 0.78-0.93 (0.85) 0.50—0.63 (0.57) 0.39-0.58 (0.48) 0.59-0.84 (0.71) 0.63-0.90 (0.77) F 30? 3.45—-4.05 (3.74) 0.81-0.91 (0.86) 0.55-0.67 (0.61) 0.40-0.58 (0.49) 0.62-0.85 (0.73) 0.69-1.09 (0.89) 0.70-0.90 15 3.24-3.94 (3.59) 0.76-0.91 (0.83) 0.55-0.71 (0.63) 0.38-0.57 (0.47) 0.64-0.84 (0.74) 0.80-1.12 (0.96) PRW PTW AW 0.48-0.71 0.41-0.61 0.58—0.80 0.58-0.91 (Fig. 1); female sternal plate VI with 1 sts on each side and sternal plate VI with 3 medium-length setae and up to 2 microsetae on each side in P. fujianensis (Fig. 2), but sternal plate VI with 2 sts on each side and sternal plate VII without microsetae in P. /ongus s. lat. (Fig. 16); male subgenital plates of different shape (cf. Figs 7, 21) and stylus evenly tapering distally in P. /ongus s. lat. (Fig. 21), but with convex lateral margins in distal half in P. fujian- ensis (Fig. 7); female subgenital plate medianly continu- ous in P. Jongus s. lat. (Fig. 22), but medianly interrupted in P. fujianensis (Fig. 8); proximal mesosome with flat- tened to slightly concave anterior margin in P. /ongus s. lat. (Fig. 20), but with pointed anterior margin in P. fuji- anensis (Fig. 6); ventral sclerite of mesosome and shape of gonopore and distal mesosome also differ between species (cf. Figs 6, 20). Male antennal characters may be more similar in these two species than illustrated here (Figs 3, 17), as their shape is affected by mounting. How- ever, scape appears to be broader and the distal process of flagellomere I appears to be longer in P. fujianensis (Fig. 3) than in P. Jongus s. lat. (Fig. 17). Description. Both sexes. Head shape and structure as in Fig. 3; frons gently rounded. No prominent reticulation on head. Marginal carina of moderate width, not widen- ing posteriorly. Dorsal pre-antennal suture prominent, not reaching marginal carina laterally. Head chaetotaxy as in Fig. 3; most dorsal sensilla visible as microsetae in most ex- amined specimens. Antennae sexually dimorphic. Thoracic and abdominal segments and chaetotaxy as in Figs 1, 2. Male. Antennae as in Fig. 3; scape, pedicel and flag- ellomere I swollen and modified in shape compared to female; scape with slight process in proximal third; flag- ellomere I with prominent distal projection and restricted rugose area, which does not extend to proximal bulbous process of segment. Abdominal chaetotaxy as in Fig. 9; inner ventral ps present on segments V—VIII; median sts on sternite VI much longer than lateral sts. Subgenital plate, stylus and terminalia as in Fig. 7; stylus broad- ening in distal half, not tapering evenly. Genitalia as in Figs 5, 6. Proximal mesosome with narrow median point, widening distally. Ventral sclerite small, roughly round- ed-rectangular, with minute postero-lateral extensions; | sensillum on each side associated with sclerite; 3 sensilla on each side lateral to ventral sclerite, forming distally divergent rows. Distal mesosome oval, dominated by large oval gonopore. Parameres curved slightly medianly, with median and lateral fingers of parameral head roughly equal in size. Measurements as in Table 1. Dtsch. Entomol. Z. 71 (1) 2024, 85-109 89 Figure 1. Pelecolipeurus fujianensis sp. nov. ex Tragopan caboti (Gould, 1857). Male habitus, dorsal and ventral views. Figure 2. Pelecolipeurus fujianensis sp. nov. ex Tragopan caboti (Gould, 1857). Female habitus, dorsal and ventral views. Female. Antennae as in Fig. 4. Abdominal chaetotaxy as in Fig. 2; sternal plate VI with 1 sts on each side. Sub- genital plate, vulval margin and terminalia as in Fig. 8; subgenital plate divided medianly. Vulval margin with 17-23 medium-length, slender vms and 6—10 short, slen- der vss on each side; median vms shorter than lateral vms. Measurements as in Table 1. Etymology. The specific name is derived from the type locality. dez.pensoft.net 90 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Figures 3-6. Pelecolipeurus fujianensis sp. nov. ex Tragopan caboti (Gould, 1857). 3. Male head, dorsal and ventral views; 4, Female antenna, ventral view; 5. Male genitalia, ventral view; 6. Distal male genitalia, ventral view. dez.pensoft.net Dtsch. Entomol. Z. 71 (1) 2024, 85-109 0.1 mm 0.1 mm Figures 7, 8. Pelecolipeurus fujianensis sp. nov. ex Tragopan caboti (Gould, 1857). 7. Male subgenital plate and abdominal segments VIII—XI, ventral view; 8. Female subgenital plate and abdominal segments VIII—XI, ventral view. 91 Pelecolipeurus longus (Piaget, 1880), comb. nov. Figs 15-22 Lipeurus longus Piaget, 1880: 370. Oxylipeurus longus (Piaget), 1880; Clay, 1938a: 171. Reticulipeurus longus (Piaget, 1880); Kéler, 1958: 332. Type host. 7ragopan satyra (Linnaeus, 1758) — satyr tra- gopan. Type locality. The Hague, Netherlands (captive bird; host is limited to the Himalayas). Other hosts. 7ragopan temminckii (Gray, 1831) — Temminck’s tragopan [tentative]. Tragopan melanoceph- alus (Gray, 1829) — western tragopan [uncertain; Clay 1938a: 172]. Specimens examined. Ex TZragopan temminckii: CHINA * 23), 29; Shanghai, Shanghai Zoo; 12 Sep. 1988; Shi Xinquan leg.; box E0026199, slides 73—76 (NNHM). 52; Beying, Beijing Zoo; 10 Oct 1973; collector un- known; box E0026199, slides 78-82 (NNHM). 4, 69; Sichuan Province, Beichuan; 4 May 1984; collector un- known; box E0026199, slides 84—87 (NNHM). Ex Cros- soptilon auritum [straggler?]: CHINA * 19; no locality; 30 Oct 1990; collector unknown; box E0026199, slide 67 (NNHM). Ex Lophura nycthemera fokiensis |strag- gler? |: Cumna * 14; Fujian Province; Dec 1990; collector unknown; box E0026199, slide 71 (NNHM). Ex Trago- pan sp.: CHINA * 19; no collection data; box E0026199, slide 83 (NNHM). Diagnosis. Both specimens from 7! temminckii and those illustrated from the type host by Clay (1938a) and von Kéler (1958) can be separated from P. fujianensis sp. nov. by the following characters: male fused abdominal segment IX—XI with more or less straight lateral margins in P. Jongus, but with concave lateral margins in P. fujian- ensis (Fig. 7); proximal mesosome of P. /ongus with flat- tened anterior margin, but with medianly pointed anterior end in P. fujianensis (Fig. 6); parameres more curved in P. longus than in P. fujianensis (Fig. 6). Specimens from 7. temminckii can be further separated from P. fujianen- sis by the characters listed under this species above, but examination of specimens from the type host of P. Jongus is necessary to establish whether the population on this host can also be separated from P. fujianensis by the same characters and whether the populations on 7. satyra and T. temminckii are conspecific. Description (of specimens from Tragopan temminckii). Both sexes. Head shape, structure and reticulation pattern as in Fig. 17; frons somewhat flattened. Marginal carina of moderate width, widening posteriorly. Dorsal pre-antennal suture prominent, reaching to or nearly to marginal carina laterally. Head chaetotaxy as in Fig. 17; many dorsal sensilla very small and difficult to see. Antennae sexually dimorphic. Thoracic and abdominal segments and chaetotaxy as in Figs 15, 16. dez.pensoft.net 92 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice 10 @ 0.5 mm Figures 9-14. Pelecolipeurus fujianensis sp. nov. ex Tragopan caboti (Gould, 1857). 9. Male leg I, dorsal side; 10. Male leg I, ventral side; 11. Male leg II, dorsal side; 12. Male leg II, ventral side; 13. Male leg ITI, dorsal side; 14. Male leg HI, ventral side. Male. Antennae as in Fig. 17; scape, pedicel and flag- ellomere I swollen and modified in shape compared to female; scape with seemingly hyaline, broad process in proximal third; flagellomere I with intensely rugose sur- face and intensely rugose bulbous process near proximal base. Abdominal chaetotaxy as in Fig. 15; inner ventral ps absent on all tergopleurites; sts on sternite VI of about equal length. Subgenital plate, stylus and terminalia as in Fig. 21; stylus tapering more or less evenly towards distal dez.pensoft.net end. Genitalia as in Figs 19, 20. Proximal mesosome flat- tened to slightly concave, with short, stout antero-lateral extensions bent slightly anteriorly. Ventral sclerite inverse V-shaped, with up to 3 sensilla on each side associated with its distal margin; 3 sensilla on each side lateral to ven- tral sclerite, forming distally divergent rows. Distal me- sosome rounded rectangular, dominated by large, roughly rounded-trapezoidal gonopore. Parameres roughly paral- lel; pst/—2 as in Fig. 20. Measurements as in Table 1. Dtsch. Entomol. Z. 71 (1) 2024, 85-109 Figure 15. Pe/ecolipeurus cf. longus (Piaget, 1880) ex Trago- pan temminckii (Gray, 1831). Male habitus, dorsal and ventral views. a3 Figure 16. Pelecolipeurus cf. longus (Piaget, 1880) ex Tra- gopan temminckii (Gray, 1831). female habitus, dorsal and ventral views. Female. Antennae as in Fig. 18. Abdominal chae- totaxy as in Fig. 16; sternal plate VI with 2 sts on each side. Subgenital plate, vulval margin and terminalia as in Fig. 22; subgenital plate continuous medianly. Vulval margin with 17—22 medium-length, slender vms and 8—12 short, slender vss on each side; median vms shorter than lateral vms. Measurements as in Table 1. Remarks. We have not seen any specimens of L. /on- gus from the type host. The original illustrations (Piaget dez.pensoft.net 94 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Figures 17-20. Pelecolipeurus cf. longus (Piaget, 1880) ex Tragopan temminckii (Gray, 1831). 17. Male head, dorsal and ventral views; 18. Female antenna, ventral view; 19. Male genitalia, ventral view; 20. Distal male genitalia, ventral view. dez.pensoft.net Dtsch. Entomol. Z. 71 (1) 2024, 85-109 21 0.1mm iis 0.1mm Figures 21, 22. Pelecolipeurus cf. longus (Piaget, 1880) ex Tra- gopan temminckii (Gray, 1831). 21. Male subgenital plate and abdominal segments VIII—XI, ventral view; 22. Female subgen- ital plate and abdominal segments VIII—XI, ventral view. 9D 1880; figs 8, 8a, 8b) are insufficiently detailed to es- tablish its identity beyond placing it in Pelecolipeurus. Clay (1938a: 171, figs 33b, 35a) and Kéler (1958: fig. 34) illustrated the male terminalia and male genitalia of this spe- cies, confirming our placement of L. /ongus in the new ge- nus Pelecolipeurus. Kéler (1958: 327-333) provided some additional morphological details, but did not consider it pos- sible to separate this species from Reticulipeurus, even sub- generically. Zlotorzycka (1966) placed P. /ongus in Reticu- lipeurus; this was followed tentatively by Gustafsson et al. (2020a), but they did not examine any specimens and noted that the species may need to be moved to a separate genus. Piaget (1880) treated specimens from both hosts as conspecific. Specimens of Pelecolipeurus from Tragopan temminckii are similar to P. Jongus (Piaget, 1880) as illus- trated by Clay (1938a) and Kéler (1958), but published illustrations and descriptions (Piaget 1880; Clay 1938a; von Keéler 1958) are insufficient to establish the status of these populations adequately. As in illustrations of L. /on- gus, the male specimens from 7. temminckii have largely flat lateral margins of abdominal segment [X—XI, sug- gesting they are conspecific. However, Clay’s illustration of the male mesosome of L. /ongus (Clay 1938a: fig. 35a) indicates that there may be differences in the shape of the mesosome between L. /ongus and specimens we have ex- amined. In our previous experience with Clay’s illustra- tions in this publication (e.g. Gustafsson et al. (2020a)), details such as these are not always reliable when com- pared to specimens examined by Clay, presumably due to limitations of the microscopy and illustration techniques of the time. Clay (1938a: 172) stated that she had also ex- amined specimens from 7! temminckii and Tragopan me- lanocephalus and that it was “impossible to say whether [P. longus| normally occurs on these two hosts or whether Piaget’s specimens are stragglers”. A photo of the lectotype female of P. /ongus is available online at the NHML’s homepage (https://data.nhm.ac.uk/ dataset). In this photo, the distal claspers of the abdomen are more attenuated than illustrated here and the scleroti- sations following the vulval margin may be narrower, but this is not clear in the photo. Moreover, these characters may be affected by mounting or be subject to individual variation within the Oxylipeurus-complex and cannot be used alone as reliable indicators of species identity. Other characters, such as vulval chaetotaxy, cannot be seen in the photo. A re-examination of the lectotype and the six paralectotypes of P. /ongus will be necessary to establish the identity of specimens from 7! temminckii (and other hosts) listed here, but, unfortunately, we were not able to either examine the specimen at the NHML, nor borrow this specimen. We presently consider populations from both 7) satyra and 7) temminckii to be conspecific, but note that P. /on- gus from the type host is in need of re-description and that such a re-description may warrant the recognition of the specimens described here as a separate species. dez.pensoft.net 96 Discussion Galliforms have some of the most diverse chewing louse faunas of any bird orders. Price et al. (2003) recognised 21 genera of lice from galliform hosts, whereas Mey (2009) recognised a total of 64 genera from the same hosts. In- cluding the new genus described here, an additional eight genera of lice have been described from galliform hosts since 2009, all except two in the Oxylipeurus-complex (Mey 2010, 2013; Gustafsson and Zou 2020a, b, 2023; Gustafsson et al. 2020a, b, 2023). In the checklist of Price et al. (2003), lice on galliform hosts represent almost 10% (21 of 212) of all avian louse genera accepted as valid; if genera accepted by Mey (2009) and those described from other host groups since 2003 are added (in total 49 genera; for example, Mey (2004); Gustafsson and Bush (2017); Gustafsson et al. (2020a, b)), this would imply that over 23% (72 of 302) of the known louse genera occur on galliform hosts, despite Galliformes itself com- prising ~ 2.66% of bird diversity (290 of 10906 spp.; Cle- ments et al. (2022)). Clearly, the diversity of lice on galliform hosts is disproportionate to the diversity of host species in this group. The reasons for this over-diversity are unclear. Galliformes constitutes an ancient lineage of birds, with fossil records going back to perhaps the late Cretaceous (Clarke 2004; Agnolin et al. 2006). However, age itself does not necessarily indicate that a host group should have a diverse louse fauna. For instance, the closely-re- lated anseriforms comprise 180 species (~ 1.65% of bird diversity; Clements et al. (2022)), but are only parasitised by 11 (Price et al. 2003) or 14 (Mey 2009) louse genera, constituting 3.61% or 4.61% of louse genera, respective- ly; note that taxonomic richness of lice is lower in diving than in non-diving birds (Fels6 and R6zsa 2006), which may affect this comparison. Probably, as more becomes known of the lice of galli- form hosts, clear patterns may emerge in the distribution of these louse genera that could explain the unexpectedly high diversity of lice on landfowl. However, some pat- terns are already dimly visible in the known distribution of lice on these hosts. In Table 2, we list the distribution of ischnoceran lice parasitising galliforms, roughly fol- lowing the classification of Mey (2009), but with some modifications, based on our own examinations of spec- imens (DRG, unpublished data). It should be noted that no characters are known that can consistently separate the Goniodes- and Goniocotes-complexes as currently circumscribed (Gustafsson et al. 2021) and Johnson et al. (2011) found both complexes to be paraphyletic. The structure of the male genitalia may ultimately be useful for defining the Goniocotes-complex, but this complex is likely nested inside the Goniodes-complex as defined by Price et al. (2003; as Goniodes), Mey (2009) and here. Each of the groups of lice included in Table 2 show dif- ferent patterns of distribution and the Ischnocera of galli- form hosts include both generalist genera occurring over several of the major radiations and genera that are more dez.pensoft.net Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice restricted. Of particular interest are the lice of tragopans and allies (genera Lophophorus Temminck, 1813, Tetra- ophasis Elliot, 1871 and Tragopan). These three genera together form a monophyletic clade, with no close rela- tives (Meng et al. 2008; Bao et al. 2010; Liu et al. 2014; Wang et al. 2014, 2017; Kimball et al. 2021). We here refer to this group as the “tragopan group” for simplicity. Oxylipeurus-complex The Oxylipeurus-complex is widely distributed across galliforms, being absent only from numidid hosts (Table 2); the genus Afrilipeurus Mey, 2010, was originally de- scribed from species known from numidid hosts, but this genus appears to be a member of the Lipeurus-complex (see below) and only superficially similar to lice in the Ox- ylipeurus-complex. Amongst the genera within the Oxyli- peurus-complex, most are restricted to certain host groups. There is a clear division in the Oxylipeurus-com- plex between genera occurring mainly on New World host groups and those occurring mainly on Old World host groups. With the exception of Chelopistes lervico- la (Clay, 1941), all members of the genera Chelopistes Kéler, 1940, Eiconolipeurus Carriker, 1945, Epicolinus Carriker, 1945, Labicotes Kéler, 1940, Trichodomedea Carriker, 1946 and Valimia Gustafsson & Zou, 2020b, are found only on New World hosts. In contrast, the genera Megalipeurus Kéler, 1958, Pelecolipeurus gen. nov., Ca- lidolipeurus Gustafsson et al., 2020b, Cataphractomimus Gustafsson et al., 2020a, Gallancyra Gustafsson & Zou, 2020a,. and Sinolipeurus Gustafsson et al., 2020a, are only found on Old World hosts. The genera Oxylipeurus Mjoberg, 1910 and Jalegallipeurus Mey, 1982, are exclu- sively known in Australia and Wallacea. That leaves only One cosmopolitan genus, Reticulipeurus Kéler, 1958, which is known both in the Old and New Worlds. Reticulipeurus also has wider host associations than most other genera in this complex, being known from both Cracidae and Phasianidae II-III. However, the species known from Phasianidae III belong to a different subge- nus (Gustafsson and Zou 2023). The species from cracid hosts have not been revised in recent years and may rep- resent a separate radiation. Reticulipeurus, as currently understood, may represent a plesiomorphic morphotype, from which other, morphologically more distinct, groups of Oxylipeurus-complex lice, have evolved. If not, the distribution of Reticulipeurus on two distinct host groups — mainly Asian phasianids and almost entirely Neotropi- cal cracids — requires further study to understand. Similarly, Gustafsson et al. (2020a) noted that the widely distributed genus Megalipeurus slightly differs morphologically amongst different host groups and may also represent several distinct lineages. Most other gen- era are more restricted: Eiconolipeurus and Epicolinus on odontophorid hosts, Labicotes on cracid hosts, Tricho- domedea on cracid and odontophorid hosts and Calidoli- peurus, Gallancyra and Valimia being known from one Dtsch. Entomol. Z. 71 (1) 2024, 85-109 97 Table 2. Distribution of ischnoceran lice across different galliform hosts. The host groupings are based on Kimball et al. (2021); host taxonomy follows Clements et al. (2022). Associations are based on Mey (2006, 2009, 2010, 2013), Gustafsson and Zou (2020a, b, 2023), Gustafsson et al. (2020a,b, 2023) and here; note that the Goniodes-, Goniocotes- and Lipeurus-complexes have not been comprehensively revised since Clay (1938a, 1940) and von Kéler (1940) and some of these taxa may not form meaningful groups. In Goniocotes Burmeister, 1838 (sensu Price et al. (2003)), four morphologically distinct groups are denoted by Roman numerals; due to the lack of detail in original descriptions of many species in this genus, it is not possible to assess whether these groups represent distinct genera or just well-marked species groups. In Goniodes Nitzsch, 1818 (sensu Price et al. (2003)), nine morphologically distinct groups are denoted by Arabic numerals; 1f Goniodes 1s divided as suggested by Mey (2009), these groups would represent separate genera for which no genus name has ever been published. Note that Price et al. (2003) used a more conservative classifi- cation, in which all genera were placed as synonyms of Goniocotes, Goniodes, Lipeurus Nitzsch, 1818 and Oxylipeurus Myjoberg, 1910, except for Pachyskelotes Kéler, 1940 and Passonomedea Carriker, 1944. Some genera treated as synonyms of Goniodes by Price et al. (2003) are probably closer to Goniocotes (see Mey (1997)), based on the morphology of the male genitalia. Conversely, it seems likely that Pavoniocotes Gustafsson et al., 2023 and the groups denoted Goniocotes III-IV here are more closely related to the Goniodes-complex than to the Goniocotes-complex. For ease of reference, the position of these species follows Price et al. (2003); these genera are marked with an asterisk (*) in the list. A few species of Goniodes cannot be identified from their available illustrations and descriptions and are here entered as “unknown”. and genus Megapodiidae Aepypodius Oxylipeurus Homocerus*, Weelahia* Megathellipeurus Megapodiella Alectura Oxylipeurus Homocerus*, Weelahia* Megathellipeurus Eulipoa Leipoa Leipoiella*, Megatheliella* Megathellipeurus Megapodiella Macrocephalon Goniocotes I Megathellipeurus Megapodius Oxylipeurus, Talegallipeurus Euligoniodes*, Lobicrotaphus*, Malaulipeurus Maleoicus* Talegalla Talegallipeurus Homocerus*, Maleophilus* Lipeuroides, Megathellipeurus | Megapodiella Cracidae Aburria Labicotes Chamaepetes Labicotes, Trichodomedea Crax Labicotes, Reticulipeurus, Trichodomedea Mitu Reticulipeurus, Trichodomedea Nothocrax Oreophasis Trichodomedea Ortalis Reticulipeurus, Trichodomedea Pauxi Reticulipeurus, Trichodomedea Penelope Reticulipeurus, Trichodomedea Penelopina Labicotes, Trichodomedea Pipile Numididae Acryllium Goniocotes IT Lipeurus Agelastes Stenocrotaphus Lipeurus Guttera Goniocotes IT Clayarchigoniodes, Stenocrotaphus Afrilipeurus, Lipeurus Numida Goniocotes I, Clayarchigoniodes, Stenocrotaphus Lipeurus, Numidilipeurus Goniocotes IT Odontophoridae Callipepla Epicolinus Genus 8, Unknown Colinicola Colinus Epicolinus Solenodes?*, Genus 8 Lipeurus Colinicola, Cuclotogaster Cyrtonyx Colinicola Dactylortyx Eiconolipeurus, Trichodomedea Dendrortyx Eiconolipeurus, Epicolinus, Trichodomedea Odontophorus __Eiconolipeurus, Trichodomedea Passonomedea Oreortyx Genus 8 Colinicola Philortyx Colinicola Ptilopachus Solenodes* Cuclotogaster Rhynchortyx Genus 8 Phasianidae I Afropavo Goniocotes ITI, Archigoniodes Lipeurus Goniocotes IV Alectoris Goniocotes I Solenodes*, Genus 6 Cuclotogaster Ammoperdix Oulocrepis Cuclotogaster Argusianus Pachyskelotes, Unknown Bambusicola Oulocrepis dez.pensoft.net 98 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Host group Oxylipeurus-complex Goniocotes-complex Goniodes-complex Lipeurus-complex Other genera and genus Campocolinus Coturnix Astrocotes Cuclotogaster Francolinus Goniocotes I Lipeurus Cuclotogaster Galloperdix Megalipeurus Goniocotes I Gallus Gallancyra Goniocotes I Oulocrepis, Stenocrotaphus Lipeurus, Numidilipeurus Cuclotogaster, Lagopoecus Haematortyx Margaroperdix Oulocrepis Cuclotogaster Ophrysia Ortygornis Stenocrotaphus Cuclotogaster Pavo Goniocotes I, Goniodes, Genus | Lipeurus Pavoniocotes Peliperdix Cuclotogaster Perdicula Cuclotogaster Polyplectron Megalipeurus Lipeurus Pternistis Goniocotes I Oulocrepis, Stenocrotaphus Lipeurus Cuclotogaster Rheinardia Lipeurus Scleroptila Goniocotes I Oulocrepis, Genus 6 Cuclotogaster, Lagopoecus Synoicus Astrocotes Tetraogallus Oulocrepis Cuclotogaster Tropicoperdix Megalipeurus Lipeurus Phasianidae II Bonasa Oulocrepis Lagopoecus Canachites Catreus Oulocrepis Centrocercus Oulocrepis Lagopoecus Chrysolophus Reticulipeurus Oulocrepis Lipeurus Crossoptilon Reticulipeurus Dictyocotes Genus 5 Lipeurus Lagopoecus Dendragapus Oulocrepis Lagopoecus Falcipennis Ithaginis Reticulipeurus Oulocrepis Lagopoecus Lagopus Oulocrepis Lagopoecus Lerwa Chelopistes Lerwoecus Lophophorus Cataphractomimus Dictyocotes Margaritenes, Genus 2 Lipeurus Lagopoecus Lophura Reticulipeurus Goniocotes I Oulocrepis Lipeurus Cuclotogaster Tyrurus Lagopoecus Meleagris Chelopistes, Valimia Goniocotes I Lipeurus Perdix Goniocotes I Solenodes* Lipeurus Cuclotogaster Phasianus Reticulipeurus Goniocotes I Oulocrepis, Solenodes* Lipeurus Cuclotogaster, Lagopoecus Pucrasia Reticulipeurus Oulocrepis Rhizothera Reticulipeurus Lipeurus Syrmaticus Reticulipeurus Goniocotes I Oulocrepis Lipeurus Lagopoecus Tetrao Reticulipeurus Oulocrepis Lagopoecus Tetraophasis Sinolipeurus Dictyocotes Genus 4 Tetrastes Lagopoecus Tragopan Cataphractomimus, Dictyocotes Genus 3 Lagopoecus Pelecolipeurus, Sinolipeurus Tympanuchus Oulocrepis Lagopoecus Phasianidae II Arborophila Megalipeurus, Reticulipeurus Goniocotes I Astrodes, Kelerigoniodes Cuclotogaster, Galliphilopterus Caloperdix Megalipeurus Goniocotes I Lipeurus Melanoperdix Lipeurus Rollulus Calidolipeurus Astrodes Lipeurus Xenoperdix host genus each. The perplexing distribution of the genus Chelopistes was discussed in detail by Mey (2006). Notably, the genera in the tragopan group are hosts to three genera of Oxylipeurus-complex that are, so far, not known from hosts outside that clade (Cataphractomimus, Pelecolipeurus, Sinolipeurus). The distribution of lice in these genera on the hosts of this radiation is summarised in Table 3. In at least one case, lice from all three genera dez.pensoft.net are known from the same host species, echoing the radi- ation into three congeneric species of the genus Valimia on the same host species (Gustafsson and Zou 2020b). To date, there is no example of all three genera occurring on the same host individual. However, data from any galli- form host are rather limited, not least because many birds in this radiation are protected. Examinations of birds in, for example, rescue centres may be necessary to establish Dtsch. Entomol. Z. 71 (1) 2024, 85-109 99 Table 3. Known distribution of Oxy/ipeurus-complex lice on 7ragopan spp., Tetraophasis spp. and Lophophorus spp. These three host genera form a monophyletic clade with no close relatives (Meng et al. 2008; Bao et al. 2010; Liu et al. 2014; Wang et al. 2014, 2017; Kimball et al. 2021). Dashes (“---“) indicate that no species of lice in this genus has, so far, been described from this host. Records suspected to be stragglers or contaminations (see Pelecolipeurus longus) are not listed here. Host species Lophophorus impejanus (Latham, 1790) Lophophorus lhuysii Geoffroy Saint-Hilaire, 1866 Lophophorus sclateri Jerdon, 1870 Tetraophasis obscurus (Verreaux, 1869) Tetraophasis szechenyii Madarasz, 1885 Tragopan blythii (Jerdon, 1870) Tragopan caboti (Gould, 1857) Tragopan melanocephalus (Gray, 1829) Tragopan satyrus (Linnaeus, 1758) Tragopan temminckii (Gray, 1830) whether the three Oxylipeurus-complex genera on hosts in the tragopan group ever co-occur on the same host individual and, if so, if they then partition the plumage amongst them. Goniocotes-complex Lice in the Goniocotes-complex are conspicuously ab- sent from both the mainly New World host radiations, Odontophoridae and Cracidae, as well as from all New World genera in the other host radiations. The sole ex- ception is the turkey, which is sometimes parasitised by Goniocotes gallinae (Linnaeus, 1758), normally found on domestic chicken. Goniocotes gallinae never seems to be reported from wild turkey in their native range (e.g. Hightower et al. (1953); Kellogg et al. (1969); Nelder and Reeves (2005); Cruz et al. (2013); Camacho-Esco- bar et al. (2014)) and this host association is likely based only on domestic birds which have been in contact with domestic chicken. Based on current knowledge, the Go- niocotes-complex would, thus, seem to be an exclusively Old World radiation. Based on the structure of the male genitalia, lice of the Goniodes-complex, listed from megapodiid hosts in Table 2, are likely more closely related to Goniocotes than to Goniodes. Mey (1997) circumscribed the genera on megapodiid hosts as a distinct group, but excluded the one known Goniocotes species from this group. If this group is considered part of the Goniodes-complex (as by, for example, Price et al. (2003)), it must be considered an aberrant group within this genus. The only known Gonio- cotes species from a megapodiid host may, as Mey (1997) pointed out, be evidence either of a secondary infestation or of a relict association. Goniocotes sensu lato is widely distributed across Old World landfowl (Table 2) and do not show any obvious Cataphractomimus Gustafsson et al., 2020a Cataphractomimus himalayensis (Rudow, 1869) Cataphractomimus ceratornis (Eichler, 1958) Cataphractomimus junae Gustafsson et al., 2020a Pelecolipeurus gen. nov. Sinolipeurus Gustafsson et al., 2020a Cataphractomimus burmeisteri (Taschenberg, 1882) Cataphractomimus mirapelta Gustafsson et al., 2020a Cataphractomimus impervius Gustafsson et al., 2020a Sinolipeurus tetraophasis (Clay, 1938) Pelecolipeurus fujianensis sp. nov. Pelecolipeurus longus (Piaget, 1880)? Pelecolipeurus longus (Piaget, 1880) Pelecolipeurus longus (Piaget, 1880)? Sinolipeurus sichuanensis Gustafsson et al., 2020a patterns of distribution. At least six morphologically dif- ferent groups can be found within Goniocotes, but the re- lationship between these groups is unclear. Only one of these groups, Goniocotes I, is widely distributed across Numididae and Phasianidae I-fII. The poorly-known Goniocotes IT group is only known from numidid hosts, where it may overlap in distribution with species in Go- niocotes I. Goniocotes III-IV are only known from the Congo peafowl and both groups are poorly known and may not be closely related to the rest of Goniocotes (see Clay (1938b)). A fifth group, only known from peafowl, was recently described as the genus Pavoniocotes Gus- tafsson et al., 2023. The distribution patterns of the sixth group, previously called Dictyocotes Kéler, 1940, mirrors that of the three Oxylipeurus-complex genera summarised in Table 3, be- ing found mainly on hosts in the tragopan group. How- ever, some species of Dictyocotes are also known from hosts in the genus Crossoptilon Hodgson, 1838, another high-altitude group of birds, mainly distributed in and around China. The presence of a mesosome in the male genitalia in this group, as well as other morphological characters, suggests that Dictyocotes should be separat- ed from Goniocotes, this will be discussed in more detail elsewhere (DRG, in prep.). Goniodes-complex The Goniodes-complex is by far the most diverse of the ischnoceran louse groups known from galliform hosts and almost half (28 of 60; 46.7%) of the groups identified in Table 2 belong to this complex. Of these, at least eight currently have no genus-level name and, with the excep- tions of Pachyskelotes Kéler, 1940 and Passonomedea Carriker, 1944, all were treated as members of a highly polytypic Goniodes by Price et al. (2003). To discuss the dez.pensoft.net 100 distribution of morphologically distinct groups within this complex, we here follow Mey (2009) in resurrecting numerous older names within this complex and use the numbers 1-8 to denote some groups that have no avail- able genus names. We deviate from Mey (2009) only in considering Z/otorzyckella Eichler [in Eichler and Vas- jukova 1981], 1981, as a synonym of Oulocrepis Kéler, 1940. Note that, as some species in this complex have never been adequately described or illustrated, the exact limits of these proposed genera and groups is in some cases tentative. A small number of species are so poorly described that they are noted as “Unknown” genera in Ta- ble 2 and not discussed further here. Goniodes-complex lice are unknown from cracid hosts and if the Goniodes-complex genera parasitising megapodiid hosts discussed above are moved to the Go- niocotes-complex, no Goniodes-complex lice would be known from members of this host family either. Other- wise, lice in the Goniodes-complex occur across all ma- jor radiations of galliforms. However, only three groups within this complex could reasonably be said to be wide- ly distributed: Oulocrepis Kéler, 1940, Solenodes Keéler, 1940 and Stenocrotaphus Kéler, 1940. The remaining genera and groups in this complex are known only from single host families or even single host genera (Table 2). Stenocrotaphus is mainly known from numidid hosts and African and South Asian francolins and spurfowl, but has, secondarily, also become established on chicken. Oulocrepis is more widely distributed, occurring on many different host genera in Phasianidae J—II. Morpho- logical variation, above all, in the male genitalia in this group is large (see, for example, Clay (1940)) and, above all, the type species (Goniodes dissimilis Denny, 1842) is somewhat different from all other species in the group with regards to head shape and male genitalia; however, other characters, such as female genitalia, indicate a close relationship. The genus as circumscribed here seems to be established on hosts in different geographical regions, from the Arctic to Sub-Saharan Africa and, in many cas- es, seems to occur on the same host species as other Go- niodes-complex lice. Solenodes is a widely distributed group, which as cir- cumscribed here, occurs on hosts from Odontophoridae and Phasianidae I-II. Notably, most of the hosts of spe- cies in Solenodes are associated with drier grasslands. The male genitalia of this group are more reminiscent of those of the Goniocotes-complex than those of any other group of Goniodes-complex lice; however, as these gen- italia are much reduced in complexity, it is possible that the group is artificial and, in reality, comprises several different lineages. Several species here placed in this ge- nus are poorly described and illustrated and a revision of the group is needed to establish its limits. The tragopan group of birds is collectively parasi- tised by four Goniodes-complex genera, of which only one presently has a proposed name: Margaritenes Keéler, 1940; the others are here referred to as Genera 2-4. Amongst these, only Genus 2 and Genus 3 appear to be dez.pensoft.net Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice closely related, sharing similarities in the structure of the male antennae and a unique fusing of the pterono- tum and tergopleurite II. Potentially, as these species are studied in more detail, further similarities may be found, but, at present, there seems to be nothing to indicate that all four genera are part of the same radiation within the Goniodes-complex. Lipeurus-complex Lice in the Lipeurus-complex are the most morpholog- ically homogeneous amongst the groups of ischnoceran lice occurring on galliforms. Lice in this complex are unknown from all New World hosts, except the turkey, which is parasitised by Lipeurus caponis (Linnaeus, 1758) naturally found on domestic chicken. Two genera in this complex are known from numidid hosts only (but secondarily established on domestic chicken) and three genera are unique to the Megapodiidae (Table 2). Based on the structure of the tergopleurites, female genitalia, male subgenital plate, abdominal chaetotaxy and other characters, it seems likely that Afrilipeurus belongs in this complex (see below); thus, three different genera occur on numidid hosts, although only two genera are known to occur on the same host genus. As with the Go- niodes- and Oxylipeurus-complexes, the Lipeurus-com- plex genera known from megapodiid hosts are unique to that radiation, highlighting the distinction of the louse fauna on megapodes. No Lipeurus-complex species have been described from any species of 7Jetraophasis or Tragopan and the only species of the genus Lipeurus known from Lopho- phorus spp. needs verification and may represent a con- tamination. As both Lipeurus- and Oxylipeurus-complex lice are of the wing louse ecomorph, it is conceivable that the multitude of Oxy/ipeurus-complex lice on hosts in the tragopan group have prevented Lipeurus-complex lice from establishing themselves there. However, the louse fauna of many members of the tragopan group re- main poorly known and the absence of Lipeurus-complex species on these hosts needs verification. Moreover, the mechanisms of interspecific competition in lice are poor- ly known and cases are known where the same host spe- Cles iS parasitised by multiple louse species of the same ecomorph (e.g. head lice on common blackbird; Osle- jskova et al. (2020)). Other ischnoceran genera Several smaller groups of ischnoceran louse genera are also known from galliform hosts. Of these, Megapodiella Emerson & Price, 1972, is only known from megapodiid hosts, Colinicola Carriker, 1946, only from odontophor- id hosts, Lerwoecus Mey, 2006, only from Lerwa lerwa (Hodgson, 1833) and Galliphilopterus Emerson & El- bel, 1957, only from Arborophila brunneopectus Blyth, Dtsch. Entomol. Z. 71 (1) 2024, 85-109 1855. It should be noted that Colinicola may be polytyp- ic, based on the structure of the male genitalia and other characters, but this has no major implications for the dis- tribution of this genus. The remaining two genera, Cuclo- togaster Carriker, 1936 and Lagopoecus Waterston, 1922, are more widely distributed. Cuclotogaster 1s known from hosts in Odontopho- ridae and Phasianidae I-III; however, the species from New World odontophorid hosts needs verification and may be an introduction following the European colo- nisation of the Americas. Otherwise, Cuclotogaster is absent from all New World hosts, despite being widely distributed in the Old World. Species of Cuclotogaster from Arborophila spp. are morphologically different from other species, with much narrowed male genitalia and possibly some differences in the tergopleurites and the female genitalia; these characters are poorly stud- ied. The genus has not been thoroughly revised since Clay (1938a) and the overall variation in Cuclotogaster is poorly known. Notably, most known hosts are in Pha- sianidae I and are associated with drier, open country (e.g. savannah, grassland). Species occurring on hosts outside this radiation also often share the same kind of habitat, suggesting that host-switching between sympat- ric host species may have occurred. In contrast, the genus Lagopoecus is mainly known from hosts in the Phasianidae II radiation, with a few species known from hosts in Phasianidae I; at least the association with domestic fowl may be due to straggling in domestic settings. Species of Lagopoecus occur in both the Old and New World and are often associated with more boreal or mountain- or forest-dwelling hosts, but exceptions are known (Table 2). In general, Lagopoe- cus occurs on lowland hosts in the boreal area, but seems more restricted to mountain-dwelling hosts further south and is largely absent south of the Equator. Galliforms in the tragopan group are parasitised by lice in the genus Lagopoecus, but no species of Cuclotogaster are known from these hosts. The Lagopoecus species par- asitising Lophophorus spp. are morphologically distinct, lacking the dorsal pre-antennal suture, but species known from 7ragopan spp. are not similar to these and do not ap- pear to be closely related. The genus Lagopoecus has not been comprehensively reviewed since Clay (1938a) and the patterns of variation are poorly known. Nevertheless, based on our current knowledge, there is nothing to sug- gest that the Lagopoecus species on tragopan group hosts form a unique radiation within this genus. Contrasting and overlapping patterns It is clear from this brief overview that no single factor can be used to explain distribution patterns amongst the Ischnocera that parasitise galliform hosts. Overall, both host phylogeny, host biogeography and host ecology appear to influence the known host associations in the groups included in Table 2. Moreover, in some cases, it 101 is not clear which factors are most important, as several factors overlap. Undoubtedly, host phylogeny is an important factor structuring host associations in louse communities on galliforms. For instance, there appears to be little overlap between the lice of megapodiid hosts and other landfowl (Table 2), likely reflecting that megapodiids are the sis- ter group of all other galliforms (Kimball et al. 2021). Similarly, many of the groups of lice occurring on nu- midid hosts do not occur on other host groups naturally (but some have spread to, for example, domestic chicken in domestic settings). Numerous smaller groups are also limited to one or a few closely-related genera, especially in the Goniodes-complex. There is also a distinct difference between most of the New World and Old World galliforms, with Tricho- domedea being shared by two New World host families, but absent on all Old World hosts and most Cuclotogas- ter and Goniocotes being absent from New World hosts despite being widely distributed across the Old World. Notably, African odontophorids are not parasitised by the same lice as New World members of this family, but by Cuclotogaster, which is widely distributed on other Af- rican hosts. Contrasting with the large-scale biogeographical pat- tern, some patterns may have more to do with host bio- tope than with faunal regions. For instance, even if Cu- clotogaster is largely limited to hosts in Phasianidae I, the genus also occurs on some members of Phasianidae IT that occur in less forested areas, such as Phasianus Lin- naeus, 1758 and Perdix Brisson, 1760 (Table 2). Similar- ly, Oulocrepis is found across both these radiations, often on birds that inhabit more open, grassy areas; Solenodes and Stenocrotaphus also appear to be distributed mainly on hosts in the same type of biotope and include at least some species in other host radiations. Notably, some patterns cannot easily be explained and may be due to gaps in our knowledge or on incorrect classification of known species. It is, for instance, curious that the widely-distributed genus, Reticulipeurus, should occur on both Old World phasianids and New World cracids, despite all other ischnoceran lice on cracids be- ing specific to the New World. Based on published data, there are no obvious morphological differences between the species on these host groups, although the species of cracids have not been revised in recent decades and few detailed illustrations have been published. The distribu- tion of the genus Megalipeurus is also difficult to under- stand, but the genus is morphologically heterogeneous and a revision of the group may reveal that the current circumscription is artificial (Gustafsson et al. 2020a). It is worth noting that elevation may influence distri- bution patterns. Several distinct genera and groups are known only or mainly to infest high-elevation hosts, such as Lerwoecus and many of the unnamed groups within the Goniodes-complex. In contrast, low-elevation hosts are often parasitised by more widely-distributed louse genera (e.g. Lipeurus, Oulocrepis, Goniocotes 1). dez.pensoft.net 102 Lice of the tragopan group Most relevant to the taxa described here are those found in the tragopan group, all of which are high-elevation birds within Phasianidae IT (Kimball et al. 2021). With the exception of an unconfirmed Lipeurus species occur- ring on one of the species of Lophophorus, all of the typ- ically low-elevation groups of Ischnocera are absent from hosts in the tragopan group. However, the diversity of lice in this group is considerable. Despite comprising only ten species in three genera, the species in this group are col- lectively hosts to at least three Oxylipeurus-complex gen- era (Table 3), as well as a morphologically distinct group of Lagopoecus, four genera within the Goniodes-complex and almost all the known species of Dictvocotes. With the exception of Dictyocotes, all these genera and groups are unique to hosts in the tragopan group. This pattern may also be mirrored in the Amblycera. Price and Beer (1964) considered Colpocephalum tetraophasis Price & Beer, 1964, “rather unique”, but did not detail in what way; Amyrsidea impejani Scharf [in Scharf and Price], 1983, was also described as having some distinct morphologi- cal characters, rare for the genus. Emended key to the Oxylipeurus-complex Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Possibly, the relevant factors structuring these host as- sociations are a mixture of host phylogeny — all the hosts, except Crossoptilon are closely related — and habitat fac- tors — all the hosts including Crossoptilon are high-eleva- tion birds. The overlap in range varies between species pairs, but, in at least some cases, the hosts in these gen- era co-occur and may even forage together (Madge and McGowan 2002). More data are needed from all hosts in the genera Lophophorus, Tetraophasis and Tragopan, as well as from other high-altitude galliforms, to evaluate the louse community parasitising these hosts and their re- lationship to related lice on other galliforms. Future collections may also help determine the extent to which the three genera of Oxylipeurus-complex lice on T! temminckii co-occur on the same host population. All three species are known from Sichuan Province, China, but from different collection events and areas within this Province. The co-occurrence of three congeneric species of lice in the Oxylipeurus-complex was recently reported from turkey (Gustafsson and Zou 2020b) and cases such as these could give insights into the process of microhab- itat partitioning in chewing lice and potentially into the process of higher-level radiation of lice on hosts. Here, we update the genus-level key to the Oxylipeurus-complex previously published by Gustafsson et al. (2020b) and emended by Gustafsson and Zou (2023) after the description of the subgenus Reticulipeurus (Forcipurel- lus). We here remove the genus A/rilipeurus from this complex, based on the justification below and include the genus Pelecolipeurus. 1 Broad-headed, with width of head similar to, or wider than, length of head; temples with elongated “horns” (Fig. 23) or with prominent lateral bulges (Fig. 24) ..............ccccceceeeees Slender-headed, with head clearly longer than wide; temples generally rounded, never with prominent bulging (Fig. 25) tari eed A bb det debe decrcte oh isc A iach ed Scencit eeRC dete atoetin tobioge o ta dk choco SN SOAR At oer Peet ere omen de th mage teeamne nade L ner canoe noes Sect: 3 Temporal setae mts1-—2 macrosetae (Fig. 24) ..........cccececcececeececeeeeseeeececeecscaeceseeaeseseeseseeaes Trichodomedea Carriker, 1946 Temporal setae mts1—2 microsetae (FIS. 23) ........ccccccececsececeececeeeeseesececeecsseecsceeseseseeceseeeeeegeeeeaes Chelopistes Kéler, 1939 Borsal.ore-anteniial Suture PresenECRiG P25). ayia eds cebeact ch A MMe LOK cl etek cama cide dade AE Redtch abe Meta NIMi Att. dll pablo pad ea eddie emg 4 Dorsal pre-antennal suture absent or, if present, only visible around aperture of ads and not extending medianly (does 51) eA, beeen hal Pye tL EL ey, 2d OMMOT. A AO MEET en ne De, 28 LL a PE oy Oe SS OL AE 1G). Dorsal pre-antennal suture as median, elongated oval, not expanded laterally (Fig. 27); female terminalia with marginal mesosetae distributed more or less equally around distal margin (Fig. 28); eye very large (Fig. 27) and pre-ocular nodus ASE Riles ciak hat hk eee ee Peer oe Onn ee we eee | ei ee ee ne Calidolipeurus Gustafsson et al., 2020b Dorsal pre-antennal suture transversal, normally reaching apertures of ads (Fig. 25); female terminalia with marginal setae gathered in the same area (Fig. 29); eye not very large (Fig. 26) and pre-ocular nodus present ..............cccceeeeee es 5 Clypeo-labral suture present (Fig. 30); stylus expanded distally, with small “hooks” on lateral margins (Fig. 31)........... Peete ete oe cde eee heel tbs Atti tome tena e aoe fered Re ie ACRE enh ME eee lansoactelsreenee ec cite wee ratte theres eee Gallancyra Gustafsson & Zou, 2020a Clypeo-labral suture absent; stylus differing in shape, but never with lateral “HOOKS” .............cccccceccecc eee eee eee eesensenseneenes 6 Dorsal pre-antennal suture with postero-lateral elongations (“epistomal suture” sensu von Kéler (1958)) extending to- wards preantennal nodi (Fig. 32); hyaline margin present (Fig. 32)............... Splendoroffula Clay & Meinertzhagen, 1941 Dorsal pre-antennal suture without such extensions (Fig. 25); hyaline margin absent (Fig. 25) ...........cc ccc cceeceeeeeeeeen sees 7 Dorsal postantennal suture present (Fig. 33); male genitalia asymmetrical, with mesosome much reduced (Fig. 34) .... U anet wi Re Gees tes ee a eceseni AL ip arta Seiten a en sects ok Resi tg Weevaene eee Mech Ob be aac rsmeh she kane ciR Cette prc Se oy eat Bc Oxylipeurus Mjoberg, 1910 Dorsal postantennal suture absent (Fig. 25); male genitalia symmetrical, with prominent mesosome (variable in dez.pensoft.net Dtsch. Entomol. Z. 71 (1) 2024, 85-109 103 8 10 11 14 15 Removal of Afrilipeurus We believe it is justified to remove the genus A/frilipeurus from the Oxylipeurus-complex and, instead, place it in the Lipeurus-complex, where it is probably closely related to Numidilipeurus Tendeiro, 1955, which occurs on the same host group. This is based on the following morpho- logical comparisons: 1. Amongst Coni elongated (Fig. 27); male mesosome with prominent V-....or Y-shaped thickening in distal half (Fig. 35); proximal margin of mesosome with rounded lateral lobes (Fig. 35); frons convergent to median point in most Species ............... Ae ata ei en nh ea one ta ae re Megalipeurus Kéler, 1958 Coni short (Fig. 25); male mesosome without thickening in distal half; proximal margin of mesosome variable, but never with rounded lateral lobes; frons rounded to flattened (Fig. 25).............ccccccccceceeeeceeseceeeeeeeeegeseeuecesgeaeausaeeesaeaesasenaesanass 9 Male abdominal segments IX+X and XI with prominent postero-lateral extensions (“claspers” sensu Carriker (1945)) Gh REE LSS eee Sena eel Ine hdl oer mua cbt Mite Neoape ae A ble ead em kr aa be ane estan Lh bel lt a Eiconolipeurus Carriker, 1945 Male-abdomelr withoutsuches hUCLUTES. can} ssbveccldides phe degenmctyrs Soodiaate pfosonnbuhiey decuiies pop \Ad an eitrg gotssaen ppgecdin hye decumates 12 Stylus of male subgenital plate about as long as rest of subgenital plate (Fig. 37); male genitalia much elongated, with mesosome-camprisingy<-L/ LOser orale neti (Fle) «teed fe ctcesteie de Cosa Nee oe tect rye WSR ae Bt a dt Pelecolipeurus gen. nov Stylus of male subgenital plate < 1/4 of length of subgenital plate; male genitalia shorter, with mesosome comprising ef UES -Ol COLA OME TM ak LEA Re Lee Lethe eh ik EE eee Wen A Ae Oe ene oh bea OY ie se, Wein vee el, Female with prominent “claspers” formed by extensions of abdominal segment XI (Fig. 39); female vulval margin deeply emarginated, with lateral sections forming rounded lobes that have subparallel median margins and median sections convex (Fig. 39); male stylus terminal (Fig. 40)..............cccccseeeeees Reticulipeurus (Forcipurellus) Gustafsson & Zou, 2023 Female without such claspers (Fig. 41); female vulval margin variably concave, but either with no section of the margin forming lobes or without median section being convex (Fig. 41); male stylus subterminal (Fig. 42)...........cccccccceseeeeeeees EE te IY ne RE Pa aD EN i fee SATYAM CTR nee ROMY PY EC TORRE DES TR ch OEY PPO DAR tenn Ae Reticulipeurus (Reticulipeurus) Kéler, 1958 Frons convergent to median: point:(similar: to’ Fig,. 83) .s 0. GA vel eee eee em ieee see Talegallipeurus Mey, 1982 Proms -OUneCed: LOstatlened CRiest2 oy Oye Meine ust msiee doy, kee te eee I Sle eel heiith eh Ae geble ReG Soe SIT Lecce edie a MS Male parameres strongly S-curved (Fig. 43); stylus arising centrally on abdominal segment IX+X (Fig. 44) .............ccee ee Se eet oe Dn Ble en el AO ee Ee ee re Le ee ee EP oe Dn Be Sinolipeurus Gustafsson et al., 2020 Male parameres not S-curved (Fig. 45); stylus varying in shape, but always arising terminally or subterminally on sub- ecnitall platecBiO SHO 423 Fo jek acta coctiane pepedorb bby edeociliuelsbe dc dred wetey galulene erduadarhupegedelidessvegde dobry geouslane pobidath ha eeesenuidaicr: 14 Male genitalia simple, with parameres fused to basal apodeme and mesosome much reduced (Fig. 46).............cceee eee Re TR RO ee oa, OPM TOT a tee PERRO NY OoPPe nba FET STN EDL TYS aed eee PRET UMN T eve... PPP T ToT EDT Tye norte FTP Epicolinus Carriker, 1945 Male genitalia with parameres articulating with basal apodeme, and mesosome not reduced (Fig. 47) ............ccceee ee 15 Lateral margins of postantennal head with secondary, ventral carina between antennal socket and site of mts2 or mts3 (Fig. 48); area between margin of head and secondary carina, densely reticulated, in some species, including ventral surface of eye (Fig. 48); male parameres with pst1—2 situated close together apically (Fig. 49); female subgenital plate divided medianly and without lateral accessory vulval plates (Fig. 50)..............cccceeeees Valimia Gustafsson & Zou, 2020b Lateral margins of postantennal head without secondary carina and without extensive ventral reticulation; male param- eres with pst1-2 separated and only pst2 apical (Fig. 51); female subgenital plate medianly continuous and with lateral accessory vulval plates*present- (Fig. 52)... conc se2es.scis0 i4hs Soetea weds Shade tg Abaca! Cataphractomimus Gustafsson et al., 2020 2. Tergopleurites are medianly divided in all Oxylipeu- rus-complex genera except Jalegallipeurus Mey, 1982, in which tergopleurites VI-—IX+X are appar- ently medianly continuous; moreover, intertergal plates are absent in all Oxylipeurus-complex gen- era. In Lipeurus and Numidilipeurus, tergal plates are medianly continuous and intertergal plates are common in males, but do not occur in all species. In Numidilipeurus, intertergal plates are present on slender-bodied Oxylipeurus-complex at least male segments IN-—V. Afrilipeurus has medi- genera, abdominal chaetotaxy consists of only su- tural setae and principal post-spiracular setae dorsally and only one sternal seta on each side on segments II—V ventrally. In the Lipeurus-complex, multiple sternal setae per segment are the norm and, in the genus Numidilipeurus, both multiple sternal setae and tergal posterior setae are present on tergopleurites H—VIII. Afrilipeurus has multiple sternal setae and tergal posterior setae present on segments II-VI. anly continuous tergopleurites and intertergal plates on male segments ITI-IV. . Female genitalia lack distally convergent rows of vulval oblique setae in the Oxylipeurus-complex and this set of setae is often reduced to one or a few short setae on the posterior margin of abdominal segment VII. In Numidilipeurus, these setae are nu- merous (> 5 per side) and form roughly convergent rows on the ventral side of segments VII-IX+X, which is the same as in Afrilipeurus. dez.pensoft.net 104 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice Figures 23-30. Key characters of the Oxylipeurus-complex. 23. Outline of head and temporal macrosetae (cut off distally) of male Chelopistes meleagridis (Linnaeus, 1758), redrawn from Kéler (1939); 24. Outline of head and temporal macrosetae (cut off distal- ly) of female 7richodomedea setosus Carriker, 1946, redrawn from original description; 25. Outline of head and dorsal preantennal suture of male Reticulipeurus (Reticulipeurus) mesopelios (Nitzsch [in Giebel], 1866), redrawn from Gustafsson et al. (2020a); 26. Outline of head and dorsal preantennal suture of male Cataphractomimus junae Gustafsson et al., 2020, redrawn from original description; 27. Outline of head, dorsal anterior suture and conus of Calidolipeurus megalops (Piaget, 1880), redrawn from Gus- tafsson et al. (2020b); 28. Female terminalia of Calidolipeurus megalops (Piaget, 1880), redrawn from Gustafsson et al. (2020b); vulval margin, lateral macrosetae and subvulval plates not illustrated; 29. Female terminalia of Reticulipeurus (Reticulipeurus) mesopelios (Nitzsch [in Giebel], 1866), redrawn from Gustafsson et al. (2020a); vulval margin, lateral macrosetae and subvulval plates not illustrated; 30. ventral view of pre-antennal area in Gallancyra dentata (Sugimoto, 1934), redrawn from Gustafsson and Zou (2020a). Figures 23—26 and 29-30 reproduced from Gustafsson et al. (2020b), with kind permission of the European Journal of Taxonomy. Abbreviations used: C = conus; CLS = clypeo-labral suture; DPS = dorsal pre-antennal suture; E = eye; HM = hyaline margin; mts3 = marginal temporal seta 3; os = ocular seta. Figures not to scale. dez.pensoft.net V . = ee we hs Figures 31—45. Key characters of the Oxy/ipeurus-complex. 31. Outline of stylus in Gallancyra dentata (Sugimoto, 1934), redrawn from Gustafsson and Zou (2020a); 32. Outline of preantennal area and dorsal pre-antennal suture of Splendoroffula ampullacea Keéler, 1955, redrawn from von Kéler (1958); 33. Outline of head and dorsal post-antennal suture of Oxyvlipeurus inaequalis (Piaget, 1880), redrawn from Mey (1990); original drawing asymmetrical; 34. Male genitalia of Oxvlipeurus inaequalis (Piaget, 1880), redrawn from Mey (1990); some details left out for clarity; 35. ventral view of mesosome of Megalipeurus sinensis Gustafsson et al., 2020a, redrawn from original description; 36. dorsal view of male terminalia of Eiconolipeurus melanotis Carriker, 1945, redrawn from original description; setae not illustrated; 37. Male subgenital plate and stylus of Pelecolipeurus fujianensis sp. nov., redrawn from Fig. 7 and simplified somewhat for clarity; 38. Male genitalia of Pe/ecolipeurus fujianensis sp. nov., redrawn from Fig. 5 and simplified somewhat for clarity; 39. Female terminalia and vulval margin of Reticulipeurus (Forcipurellus) formosanus (Uchida, 1917), redrawn from Gustafsson and Zou (2023); chaetotaxy and other detail omitted for clarity; 40. Male subgenital plate of Reticulipeurus (Forcipurellus) formosanus (Uchida, 1917), redrawn from Gustafsson and Zou (2023); 41. Female terminalia and vulval margin of Reticulipeurus (Reticulipeurus) reevesi (Clay, 1938), redrawn from Gustafsson et al. (2020a); chaetotaxy and other detail omitted for clarity; 42. Male subgenital plate of Reticulipeurus (Reticulipeurus) mesopelios (Nitzsch [in Giebel], 1866), redrawn from Gustafsson et al. (2020a); 43. Outline of male paramere of Sinolipeurus tetraophasis (Clay, 1938), redrawn and sim- plified from Gustafsson et al. (2020a); 44. Outline of male terminalia and stylus of Sinolipeurus tetraophasis (Clay, 1938), redrawn and simplified from Gustafsson et al. (2020a); 45. Outline of male paramere of Reticulipeurus (Reticulipeurus) ithaginis (Clay, 1938), redrawn and simplified from Gustafsson et al. (2020a). Figures 31—36 reproduced from Gustafsson et al. (2020b), with kind permission of the European Journal of Taxonomy. Abbreviations used: AL = anterior lobes; BA = basal apodeme; CL = “claspers”’; DPAS = dorsal post-antennal suture; DPS = dorsal pre-antennal suture; ES = epistomal suture; HM = hyaline margin; MES = meso- some; SGP = subgenital plate; STY = stylus; VM = vulval margin; Y = Y-shaped thickening. Figures not to scale. dez.pensoft.net 106 Daniel R. Gustafsson et al.: New genus of Oxylipeurus-complex chewing lice 46 47 50 52 YG Figures 46-52. Key characters of the Oxy/ipeurus-complex. 46. Distal section of male genitalia of Epicolinus clavatus (McGre- gor, 1917), redrawn from Carriker (1945); 47. Distal section of male genitalia of Cataphractomimus mirapelta Gustafsson et al., 2020a, redrawn from the original description, with some simplification for clarity; 48. Outline of male head of Valimia polytrapezia (Burmeister, 1838), with post-antennal ventral carina and densely reticulated area marked with grey dots; other characters omitted; 49. Male paramere of Valimia corpulenta (Clay, 1938), redrawn from Gustafsson and Zou (2020b); 50. Outline of ventral view of female terminalia of Valimia polytrapezia (Burmeister, 1838); 51. Male paramere of Cataphractomimus mirapelta Gustafsson et al., 2020a, redrawn from the original description; 52. Outline of ventral view of female terminalia of Cataphractomimus impervius Gustafsson et al., 2020a, redrawn and simplified from the original description. Figs 46, 48, 50, 52 reproduced from Gustafsson et al. (2020b), with kind permission of the European Journal of Taxonomy. Abbreviations used: BA = basal apodeme; LAVP = lateral accessory vulval plates; MES = mesosome; PM = parameres; ps//—2 = parameral setae /—2; SGP = subgenital plate; SMC = second- ary marginal carina. Figures not to scale. 4. Marginal temporal seta | is at least a mesoseta in otes Kéler, 1940, in which the stylus is absent. The Afrilipeurus and Numidilipeurus, but always a mi- croseta in the Oxylipeurus-complex. . Despite considerable variation amongst genera, the male terminalia in the Oxylipeurus-complex are rather uniform in their basic structure, with a gener- ally rounded ano-genital opening, anterior to which may be a transverse sclerotisation that may be con- tinuous with the subgenital plate; several setae of varying length are situated anterior to this opening and a maximum of one seta on each side (typically none) is situated on the ventral side of the poor- ly-sclerotised areas postero-lateral to the ano-geni- tal opening. Even in genera such as Pelecolipeurus, where the ano-genital opening is not clearly visi- terminalia of the Lipeurus-complex males are more variable, but do not include a transverse sclerotisa- tion and the non-sclerotised areas distal to the sub- genital plate may form a longitudinal groove, with multiple setae on each side; this is, for instance, the case in some Numidilipeurus. In Afrilipeurus, there is no transverse sclerotisation and there are multiple small setae on each side lateral to a longi- tudinal groove. . The female subgenital plate is never extended much distal to the row of sternal setae of segment VII in the Oxylipeurus-complex, but is extended distal to this row in Lipeurus-complex and in Afrilipeurus. ble, its position can be judged by the distribution of setae and the anterior sclerotisation and this struc- ture appears to be found even in the genus Labic- For these reasons, we here exclude Afrilipeurus from the Oxylipeurus-complex and transfer it to the Lipeu- rus-complex, where it is probably close to Numidilipeurus. dez.pensoft.net Dtsch. Entomol. Z. 71 (1) 2024, 85-109 Acknowledgements Most of the Phthiraptera specimens in the collection of the National Natural History Museum of China were accumulated over the years by the late research- er Liu Sikong and these specimens provided the type specimens for comparison and additional research ma- terials for this study, for which we express our grati- tude. This research was funded by the Introduction of Full-Time High-Level Talent Fund of the Institute of Zoology, Guangdong Academy of Sciences (grant GIABR-GJRC201701), the National Natural Science Foundation of China (grants 31961123003, 32001098), the Foreign Young Talent Plan (QN20200130012) and the Pearl River Talent Recruitment Program of Guang- dong Province (Grant 2019QNO1N968). These agencies had no influence over the design and execution of this study. We are very grateful for the comments and sug- gestions made by two reviewers, which helped improve this manuscript. 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