Redescriptions of thirteen species of chewing lice in the Brueelia-complex ( Phthiraptera , Ischnocera , Philopteridae ) , with one new synonymy and a neotype designation for Nirmus lais Giebel , 1874

Thirteen species of chewing lice in the Brueelia-complex are redescribed and illustrated. They are: Brueelia blagovescenskyi Balát, 1955, ex Emberiza schoeniclus (Linnaeus, 1758); B. breueri Balát, 1955, ex Chloris chloris (Linnaeus, 1758); B. conocephala (Blagoveshchensky, 1940) ex Sitta europaea (Linnaeus, 1758); B. ferianci Balát, 1955, ex Anthus trivialis (Linnaeus, 1758); B. glizi Balát, 1955, ex Fringilla montifringilla Linnaeus, 1758; B. kluzi Balát, 1955, ex Fringilla coelebs Linnaeus, 1758; B. kratochvili Balát, 1958, ex Motacilla flava Linnaeus, 1758; B. matvejevi Balát, 1981, ex Turdus viscivorus Linnaeus, 1758; B. pelikani Balát, 1958, ex Emberiza melanocephala Scopoli, 1769; B. rosickyi Balát, 1955, ex Sylvia nisoria (Bechstein, 1792); B. vaneki Balát, 1981, ex Acrocephalus schoenobaenus (Linnaeus, 1758); Guimaraesiella haftorni (Balát, 1958) ex Turdus iliacus Linnaeus, 1758; G. lais (Giebel, 1874) ex Luscinia megarhynchos (Brehm, 1831). Redescriptions are made from type material where available. Holotypes are identified in Balát’s material when possible, and lectotypes are designated for B. blagovescenskyi, B. breueri, B. glizi, B. ferianci, B. kluzi, B. kratochvili, B. pelikani, and B. rosickyi; a neotype of Nirmus lais Giebel, 1874 is designated. Brueelia weberi Balát, 1982, is placed as a synonym of Brueelia conocephala (Blagoveshchensky, 1940).


Introduction
Correct identification of chewing lice (Phthiraptera) to species level is often hampered by inadequate species descriptions.During work on a recent revision of the species-rich Brueelia-complex (Gustafsson and Bush 2017), it became apparent that the majority of the described species in this group are impossible to identify without comparison with type material.Several recent publications have provided redescriptions of some key taxa (e.g., Mey and Barker 2014, Valim and Cicchino 2015, Gustafsson and Bush 2017, Mey 2017, Gustafsson et al. 2018a); however, the majority of the proposed species in this complex are still poorly described and only partially illustrated.
To partially address the difficulties in identifying lice in this complex, we here redescribe 13 species of chewing lice in the Brueelia-complex: 10 species in the genus Brueelia Kéler, 1936 and two species in the genus Guimaraesiella Eichler, 1949.Redescriptions of 10 of these species are based on type material, complemented in some cases by non-type material.In most species, the present status of Balát's specimens is addressed, including notes on specimens that must be regarded as lost.To stabilize the nomenclature and anchor the descriptions and illustrations here to specific specimens, we also designate a number of lectotypes and paralectotypes from Balát's syntype series.
In addition, we redescribe Nirmus lais Giebel, 1874, based on specimens in Balát's collection, and designate one of these as the neotype of this species.Moreover, we here consider one proposed species name, Brueelia weberi Balát, 1982, as a synonym of an older species name, Degeeriella conocephala Blagoveshtchensky, 1940.We take the opportunity to redescribe D. conocephala as well, based on non-type specimens in Balát's collection.
With these redescriptions, only one species of Brueelia and Guimaraesiella described by Balát remain without modern redescriptions: Guimaraesiella tovornikae (Balát, 1981).We were unable to find any specimens of G. tovornikae at the Moravian Museum, and the types must therefore be assumed to be lost.Gustafsson and Bush (2017) saw specimens identified as this species in the Brelih Collection at the Slovenian Museum of Natural History (Ljubljana, Slovenia) but did not redescribe this species.

Material and methods
We examined slide-mounted specimens in František Balát's collection deposited at the Moravian Museum, Brno (MMBC).In addition, we examined some specimens from the Natural History Museum, London, United Kingdom (NHML), the Slovenian Museum of Natural History, Ljubljana, Slovenia (PMSL), and the Museum of Natural History, University of Wrocław, Poland (MNHW).We typically only illustrated and measured specimens at the MMBC; other specimens were only examined visually.In some cases, we were unable to illustrate, for example, male genitalia accurately, even if specimens in other collections were better preserved than the ones at the MMBC.Specimens were examined in an Olympus CX31 microscope.Illustrations were drawn by hand, using a drawing tube fitted to the microscope.Line drawings were scanned, collated, and edited in GIMP (http://www.gimp.org).Grey lines in all illustrations denote the approximate extent of dark pigmentation on heads, tergopleurites, and female subgenital plates; these patterns typically differ slightly between specimens of the same species and sometimes between sides of the same specimen.
Host taxonomy follows Clements et al. (2018).The species treated here are ordered according to host family.

Note on Balát's type series
In the original descriptions of most of the species redescribed here, Balát explicitly mentioned a single male and a single female as type specimens but listed all other specimens examined as "other material".Article 72.4.6 of the International Code of Zoological Nomenclature (1999) states that if an author establishing a new species-group taxon uses the term "type" or its equivalents for some specimens, but also lists other specimens, these additional specimens are excluded from the type series.Balát appears to have been unaware of this, and labeled several non-type slides as "paratypes", including some slides deposited in other collections.These specimens have no special status, and are not either paratypes or paralectotypes.

Female. Thoracic and abdominal chaetotaxy as in
Remarks.Balát (1955) mentioned a type male and female on slide no.404.The same handwritten notes are on slides 404a and 404b.As Balát (1955) did not explicitly designate a holotype, both examined type specimens mentioned in original description represent syntypes.We hereby designate the male on slide 404a as the lectotype of B. blagovescenskyi.The other syntype becomes a paralectotype.In addition, Balát (1955) mentioned two females and 15 nymphs from the same host species as other (non-type) material.Except one female on slide  404c, these have not been found in the MMBC collection, and must be assumed to be lost.Our redescription of this species is therefore based only on the lectotype and paralectotype, and the single non-type female.
The lectotype male and paralectotype female (404ab) are mounted on slides using a second slide used as a cover slide, which blurs the outline of the thoracic and abdominal plates and prevents using higher magnifications.Accurate illustration of the male genitalia is impossible without remounting the specimen, which was not attempted; the genitalia are therefore illustrated approximately.Moreover, smaller setae are very hard to see, and especially smaller abdominal setae of the male may have been overlooked.The female 404b lacks a subgenital plate.For the head and female illustrations, the non-type female specimen (slide 404c) was used.Fresh collections are needed to establish the correct abdominal and leg chaetotaxy of males of this species, as well as the shape of the male genitalic elements.Brueelia pelikani Balát, 1958: 414.Type host.Emberiza melanocephala Scopoli, 1769, black-headed bunting (Emberizidae).
Remarks.Balát (1958) did not designate a holotype for B. pelikani, but he mentioned that he had examined 3 males, 8 females, and 12 nymphs from 3 hosts; collectively these form the syntype series.The text "Type male and female" is written by hand on the label of slide 969a, and "paratypes" on slides 969b-c.Another slide (969d) with 1♂, 1♀ deposited at the NHML (Brit.Mus. 1958-452) is marked "paratypes".Presently, five slides with a total of two males, six females, and one nymph are de- posited at the MMBC.All specimens other than these and the two specimens at the NHML have been lost.To settle the identity of this species, we hereby designate the male on slide 969a as the lectotype of B. pelikani.The other syntypes become paralectotypes.
The abdomen of this lectotype male is unfortunately disrupted distally, which has affected the genitalia.In the paralectotype male 934, the mesosome is partially obscured by gut content, and the shape of the proximal mesosome cannot be seen clearly.We have illustrated the mesosome as seen in the lectotype, but the other genital elements as seen in the paralectotype male (934).Balát, 1955 Figs 13-19 Brueelia breueri Balát, 1955: 505.Type host.Chloris chloris (Linnaeus, 1758), European greenfinch (Fringillidae).
Description.Both sexes.Head flat dome-shaped (Fig. 15), lateral margins of preantennal area slightly convex, frons broadly concave.Marginal carina narrow, deeply displaced and widened at osculum, median margin undulating.Ventral anterior plate small, shieldshaped.Head chaetotaxy and pigmentation patterns as in Figure 15; head sensilla and pts not visible in examined specimens.Preantennal nodi not bulging.Preocular nodi much larger than postocular nodi.Marginal temporal carina moderate in width, with undulating median margin.Gular plate lanceolate, slender.Thoracic and abdominal segments and pigmentation patterns as in Figures 13, 14.
Remarks.Balát (1955) designated a male from Gabčíkovo and a female from Podunajské Biskupice as types; these are therefore syntypes.The word "type" is written in pencil on the slides 676 and 1118a in the MMBC collection.In addition, Balát mentioned seven females and one male from the same two hosts, and one female from Tormafölek (Zala m., Hungary, 4 Apr. 1952, leg. Georg Breuer; not at MMBC) as "other material", which do not comprise type material.Presently, there are 4 slides of B. breueri with a total of two males and four females deposited at MMBC.The other four females, including that from Hungary, are lost.Therefore, to settle the identity of this species, we hereby designate the male on slide 676 as the lectotype of B. breueri.The other syntypes become paralectotypes.Some specimens deposited in the NHML are called "paratypes", are not type specimens, as they are not referred to as such in the original publication.
Both antennae of the lectotype male are folded underneath the head and seemingly squashed.We have here reversed the dorsal view of the antenna and illustrated it in a more natural position; the antenna in the ventral view is illustrated as in the specimen.However, in both cases the antennae are likely narrower than illustrated here.As both antennae are displaced, the precise location of antennal setae cannot be established, and these have therefore not been illustrated here.Additional material is needed to fully redescribe B. breueri.Balát, 1955 Figs 20-26 Brueelia glizi Balát, 1955: 509.Type host.Fringilla montifringilla Linnaeus, 1758, brambling (Fringillidae).
Remarks.Balát (1955) designated one male and one female on slide 672 as types, but did not explicitly designate either of these as holotype; these therefore constitute the syntype series.The specimens are designated as "types" on the handwritten label.Another nine females and two nymphs were mentioned from the same host specimen, and one female from a different host specimen.Presently, four slides with one male, seven females and one nymph are present at the MMBC; the remaining specimens must be regarded as lost.To settle the identity of B. glizi, we hereby designate the male on slide 672a as the lectotype, and the female on the same slide as paralectotype.Specimens deposited at the NHML and MNHW are labeled "paratypes", but these are not mentioned as paratypes in the original description, and thus do not have type status.
Additional material is necessary to describe the male genitalia accurately.
Type locality.Lednice, Czechia.Description.Both sexes.Head flat-dome shaped (Fig. 29), lateral margins of preantennal area convex, frons rounded to slightly flattened.Marginal carina moderate in width, shallowly displaced and widened at osculum, median margin undulating.Ventral anterior plate small, shield-shaped.Head chaetotaxy and pigmentation patterns as in Figure 29; head sensilla and pts not visible in examined specimens.Preantennal nodi with slight median bulge.Preocular nodi larger than post-ocular nodi.Marginal temporal carina moderate in width, undulating.Gular plate not entirely clear in examined specimens, but roughly lanceolate.Thoracic and abdominal segments and pigmentation patterns as in Figures 27, 28.
Remarks.Balát (1955) did not explicitly designate a holotype, but mentioned one male and one female on slide 15/53 (= number on host's ring, current slide number 1138) as types; these comprise the syntype series.This is confirmed by Balát's handwritten notes on the slide label.In addition, he mentioned three males and eight females from the same host specimen, and one female from a different host as additional material.These are all present in the Balát collection at the MMBC.We hereby select the male on slide 1138 as the lectotype, and one of the females on the same slide as a paralectotype.These have been marked on the slide with dark spots.
All examined specimens in Brno are poorly cleared, and many are still attached to feather fragments that further obscure the morphology.As a result, thoracic and abdominal chaetotaxy and plates are not always clearly visible, and are here illustrated as accurately as possible.Vulval setae only clearly visible in one female, and range of variation may be greater than given above if more specimens are examined.
Remarks.Balát (1982) explicitly designated the female on slide 1448 as the holotype Brueelia weberi, and several other specimens as paratypes.This is confirmed in his handwritten notes on the slides.All specimens are present in the MMBC collection, with the exception that there is only one slide marked "Pfl90".However, this female and one of the paratype males (slide 1411) represent a separate species (see below) and have, therefore, been excluded from the paratypes.
We have examined Balát's type and non-type material identified as B. weberi, and compared these with his extensive collection of B. conocephala from Sitta europaea caesia.No diagnostic characters that could separate these two species have been found, and most measurements for specimens from P. major fall within the range of the measurements for specimens from S. europaea.We therefore consider B. weberi to be a synonym of B. conocephala.There is enough variation in the head shape and measurements of Balát's specimens of B. conocephala to accommodate the perceived differences in dimensions reported by Balát (1982), and the reported differences in the shape of the parameres can be ascribed to individual variation or artificial differences due to mounting.Balát collected B. weberi from several localities, and it would appear that this species is well established on the host, Parus major.This is in contrast to the only other material known from birds in the P. major-complex reported by Gustafsson et al. (2018b).They described two species of Brueelia (B.picea Gustafsson et al. 2018b andB. nazae Gustafsson et al. 2018b) which they did not consider to be closely related to B. conocephala.However, all material Gustafsson et al. (2018b) examined was from non-European members of the P. major-complex.
Interestingly, the "paratype" male on slide 1411 (Břeclav -Kančí obora, Czechia, 5 Mar.1954, F. Balát, 1411, MMBC) and "paratype" female on slide Pfl90 (Chuchle, Czechia, 28 Jan. 1938, K. Pfleger, Pfl90, MMBC) represent a different, undescribed, species of Brueelia.The male specimen is similar to B. nazae in head shape, but more similar to B. picea in the shape of the genitalia; the abdominal chaetotaxy is different from both species, with aps on abdominal segment IV (absent in both B. picea and B. nazae).The female specimen is slightly different in head shape from the male specimen, and may represent a different species.We do not describe this species here, as more material is needed to sort out whether both spe- cies of Brueelia actually occur on P. major in Europe, or whether Pfleger's and Balát's material originated in contaminations or stragglers.
Non-type material.1♂, 6♀, same data as holotype, F. Balát, 1062, 1127, 1177 (MMBC).1♀, same data as holotype, Brit.Mus. 1955-662 (NHML) Remarks.Balát (1955) did not explicitly designate a holotype for B. ferianci, but mentioned a male and a female as types.On slide no.1062, which contains three specimens, the male is circled; on the label, the ♂ is circled within a box that reads "Typ ♂ a ♀".We therefore consider this to be an indication that Balát considered this to be the holotype.However, as he did not explicitly name it as such in the original publication, it is a syntype, not a holotype.We hereby designate this male the lectotype, and  Balát (1955) is at the MMBC, except one female at the NHML and one female we have not been able to locate; this specimen must be regarded as lost.Another three females (slide no.1177) were collected from the same host species at the same day on the same location as holotype, but according to ring number (42/53) these lice are from another host specimen that is not mentioned in original paper.The specimen deposited at NHML is labeled "paratype", but has no type status.
The width of the frons differs somewhat between different specimens.The head is here illustrated from the holotype, whereas the full-body illustration is from a more narrow-headed specimen, to illustrate the variation in this species.Most specimens examined are more similar to the narrow-headed illustration.We do not presently consider these differences to be of any taxonomic importance, as the specimens we have examined are otherwise similar.However, fresh material from a number of host subspecies and populations may reveal that the material we have examined represents multiple species.Antennae in holotype and paratype males folded under the head, and here illustrated based on non-type material.
Remarks.Balát (1958) did not designate any type specimens, and all specimens he mentioned are therefore syntypes.The words "Type male and female" is handwritten on the label of slide 917a, and we therefore designate the male on this slide as the lectotype (this male has been marked with a dark spot on the slide); all other specimens mentioned by Balát (1958) thus become paralectotypes.All material is present at the MMBC except for the slides at the NHML and PSML listed above, as well as a slide with a single male we have been unable to trace; it should be regarded as lost.
In addition, there are two slides at MMBC (1485 and 1486) marked 'Type male" and "Type female" on the labels.However, these were collected a year after the publication of B. kratochvili, and can thus not be part of the type series.The slides from Motacilla alba are also labeled "Type male" (slide no.1215), "Paratype male" (slide no.1216), and "Type female" (slide no.1217), but  no species name based on these specimens have ever been published.The specimens from Motacilla alba are here deemed to be conspecific with M. kratochvili from M. flava.There are no significant differences in head shape, male genitalia, or abdominal chaetotaxy between material from the two host species, but females from M. alba have slightly different vulval chaetotaxy from that described above (4 vos, 3 or 4 vms, 5-7 vos on each side).These setal numbers overlap, and we therefore consider M. alba to be a new host record of B. kratochvili.

Brueelia rosickyi Balát, 1955
Remarks.Balát (1955) did not designate any holotype, but mentioned a male and a female as "types"; these two specimens comprise the syntype series.The specimens on slides 1133a and 1133b are marked accordingly in handwriting, and the male is here designated the lectotype with the female becoming the paralectotype.All other specimens mentioned by Balát as additional specimens have no type status.Slide 1133d is marked "allotype female" and slides 1133e-n are marked "paratypes", but this does not seem to be in Balát's hand.Presently, 20 slides with a total of four males and 18 females are deposited at the MMBC.We have been unable to trace the remaining one male and three females and consider them to be lost.
Type locality.Velký Dvůr u Pohořelic, Czechia.Description.Both sexes.Head elongated, rounded-trapezoidal (Fig. 64), lateral margins of preantennal area convex proximally and concave distally, frons narrowly concave.Marginal carina moderate in width, with undulating median margin, deeply displaced at osculum.Ventral anterior plate small, shield-shaped.Head chaetotaxy and pigmentation patterns as in Figure 64.Preantennal nodi not bulging.Pre-and postocular nodi of roughly equal size.Marginal temporal carina of moderate width, median margin undulating.Gular plate lanceolate.Thoracic and abdominal segments and pigmentation patterns as in Figures 62 and 63.
Female.Thoracic and abdominal chaetotaxy as in Figure 63; holotype has 5 mms on one side and 7 mms on the other; we have here illustrated only 5, as this is the normal amount in Brueelia and the number found in the examined non-type females.Female subgenital plate is almost completely translucent and exact limits very hard to ascertain; apparently broadly pentagonal, with connection to cross-piece moderate in width (Fig. 68).Vulval margin rounded, with slight bulge in median section; 3  or 4 short, slender vms and 3 or 4 short, thorn-like vss on each side; 3 short, slender vos on each side of subgenital plate; distal 1 vos median to or only slightly anterior to vss.Measurements (n = 2): TL = 1.42-1.72;HL = 0.34-0.38;HW = 0.25-0.27;PRW = 0.17-0.19;PTW = 0.24-0.26;AW = 0.34-0.39.
Remarks.Balát (1981) explicitly designated a holotype (female on slide 1519), which is also marked accordingly on the label in handwriting.All other specimens were explicitly designated paratypes.Presently, there are four slides with this material at the MMBC, comprising one male and three females; the remaining two males and two nymphs mentioned by Balát are not in the MMBC, and must be regarded as lost.In addition, slide 1520, which supposedly contained a male of this species, is empty.
Type locality.Zabljak, Montenegro.Description.Both sexes.Head flat dome-shaped (Fig. 71), lateral margins of preantennal area convex, frons flat to slightly concave.Marginal carina moderate in width, median margin slightly undulating, deeply displaced and widened at osculum.Ventral anterior plate small, shieldshaped with concave anterior margin.Head chaetotaxy and pigmentation patterns as in Figure 71; pigmentation very uniform, and difference between different areas slight.Preantennal nodi slightly bulging.Pre-and postocular nodi large.Marginal temporal carina wide, with undulating median margin.Gular plate broad, with concave lateral margins.Thoracic and abdominal segments as in Figures 69 and 70.Thoracic and abdominal pigmentation more or less uniform, and not denoted in Figures 69, 70.
Remarks.Balát (1981) explicitly designated the male on slide 1523 (Brelih's collection number 6342) as the holotype, and this is confirmed by the handwritten note on the slide label.Another female from the same host specimen (slide no.1524; Brelih's collection number 6344), and 20 males, 26 females, and 6 nymphs were designated as paratypes.Presently, six slides with the holotype and five paratypes are at the MMBC.All other specimens are missing from the MMBC, and must be regarded as lost.
Remarks.Balát (1981) explicitly designated the female on slide 1242 as the holotype, and the specimens on slides 1240 and 1241 as paratypes.This is confirmed by the handwritten notes on the slide labels.All specimens are present in the MMBC.Balát (1981) stated that both paratype males were immature.This is incorrect, as both males are adult.However, all three known specimens are poorly cleared, and many details cannot be seen properly, including the meso-and metasterna, metepisterna, proepimera, the gular plate, many leg setae, and the distal section of the subgenital plate of both sexes.More specimens of G. haftorni are needed to completely redescribe and reillustrate this species.
The Guimaraesiella of European thrushes are all morphologically very similar, differing mainly in the male genitalia and the head shape.Moreover, we have seen some specimens of Guimaraesiella from non-type host species in European material (D.Gustafsson unpublished data).Unless these records are the result of contamination or misidentification of the host, this may suggest that at least some European species of Guimaraesiella occur on more than one host species.Relying on host relationships to obtain the species identity of Guimaraesiella samples from thrushes may thus be unreliable.However, almost all species of Guimaraesiella, including those from thrushes, are poorly described, and presently unidentifiable.Redescriptions of Guimaraesiella amsel (Eichler, 1951), Guimaraesiella marginata (Burmeister, 1838), Guimaraesiella turdinulae (Ansari, 1956), and Guimaraesiella viscivori (Denny, 1842) are urgently needed to establish the species limits in this group.Type locality.None given in original, but likely Germany.Neotype (designated herein) is from Nejdek u Lednice, Czechia.

Guimaraesiella lais (Giebel, 1874)
Description.Both sexes.Head broad, rounded pentagonal (Fig. 85), lateral margins of preantennal area convex, frons broadly concave.Marginal carina moderate in width, with undulating median margin.Exact posterior extent of dorsal preantennal suture not clear in examined specimens, but suture does not appear to reach ads.Ventral anterior plate with deeply concave anterior margin.Head chaetotaxy and pigmentation patterns as in Figure 85; pigmentation of preantennal head rather uniform.Preantennal nodi with slight median bulge.Preocular nodi larger than postocular nodi.Marginal temporal carina thin, of more or less equal width.Gular plate short, broad, with median point.Thoracic and abdominal segments and pigmentation patterns as in Figures 83, 84.
Male.Sternites II-IV partially ruptured and displaced in neotype, and here illustrated approximately.Thoracic and abdominal chaetotaxy as in Figure 83; neotype has no setae on dorsal side of abdominal segment XI, but this is likely an anomaly.Male genitalia partially obscured by gut content.Basal apodeme widens proximally, with slightly concave lateral margins in distal half (Fig. 86).Proximal mesosome widening proximally, with concave lateral margins (Fig. 87).Ventral sclerite obscured by gut  Remarks.Gustafsson and Bush (2017) included Nirmus lais Giebel, 1874, in Guimaraesiella Eichler, 1949, without comment; they did not examine any specimens of this species.The placement of this species in Guimaraesiella followed Balát (1955), who placed it in Allobrueelia Eichler, 1951, a synonym of Guimaraesiella, andZłotorzycka (1977), who placed it in Allonirmus Złotorzycka, 1964, also a synonym of Guimaraesiella.However, they overlooked that Giebel (1874) stated that this species was close to Nirmus intermedius Nitzsch [in Giebel], 1866, which Gustafsson andBush (2017) placed in Brueelia Kéler, 1936.This apparent contradiction requires some additional discussion.
Giebel's description of N. lais was based on a single female, and does not contain any specific character that can be used to place N. lais in either Brueelia or Guimaraesiella with certainty.Giebel (1874) did not illustrate this species.Giebel (1866) recorded lice from the same host merely as "N …" [species 25 under the genus Nirmus], but lists specimens from Erithacus rubecula (Linnaeus, 1758) under the same heading; the lice from E. rubecula were later (Giebel 1874) described as Nirmus tristis Giebel, 1874, which was also placed in Guimaraesiella in the revision of Gustafsson and Bush (2017).
Giebel's (1874) statement that N. lais is similar to N. intermedius is unreliable, as his other statements about similarity between louse species are often confusing.For instance, on the page before the description of N. lais, Giebel (1874: 142) stated that Nirmus intermedius is similar to Nirmus ruficeps Nitzsch [in Giebel], 1866, and N. limbatus Burmeister, 1838.The former species is a head louse, now placed in the genus Rostrinirmus Złotorzycka, 1964, whereas the latter is an uncommonly wide-headed and large-bodied member of Brueelia s. str.Brueelia intermedia, by contrast, is a slender-headed species of Brueelia, quite unlike both N. ruficeps and N. limbatus.This issue is further confused by Giebel's statement that N. intermedius is similar to N. merulensis Denny, 1842, differing only in the proportions of the antennae and the prothorax.Gustafsson and Bush (2017)  Apart from the specimens listed here, we have been unable to locate any specimens of Brueelia-complex lice from L. megarhynchos in any of the museum collections we have searched (see list in Gustafsson and Bush 2017).In particular, Giebel's original specimen appears to have been destroyed in the war (Clay and Hopkins 1955).Moreover, Balát's (1955) report appears to be the only subsequent report of any species of louse in the Brueelia-complex from L. megarhynchos.Eichler [in Niethammer] (1937; not seen) and Séguy (1944) reported N. lais from Luscinia luscinia (Linnaeus, 1758); we have not seen these specimens.It is not clear from Séguy's (1944) short description whether his specimens represent the same species as Giebel's N. lais, or whether this identity is assumed based on the close relationship between the host species.Złotorzycka (1977: figs 149-152) illustrated the head, ventral anterior plate, male genitalia, and pleurites of N. lais, but indicated that this species was not known from Poland (ibid.: 10).It is therefore uncertain where the material she based her illustration on originated, nor where this specimen is located today.Złotorzycka's illustrations are rarely very informative, especially those of male genitalia.However, the specimens we have examined are largely concordant with the illustrations of Złotorzycka (1977).
To stabilize the nomenclature of the lice found on thrushes and flycatchers, we here designate a neotype for Nirmus lais Giebel, 1874, from Balát's specimens.These specimens all belong to Guimaraesiella (sensu Gustafsson and Bush 2017), and our neotype designation thus conforms to the placement of this species in Guimaraesiella by Gustafsson and Bush (2017), in Allobrueelia [= Guimaraesiella] by Balát (1955) and Mey (2017), and in Allonirmus by Złotorzycka (1977).More- over, this conforms to Giebel's (1866) earlier placement of Nitzsch's material from L. luscinola [= L. megarhynchos; but given as Sylvia luscinia by Giebel (1866)] with his material from E. rubecula, which represents Guimaraesiella tristis.
Remarks.Balát (1981) reported four males and three females of this species from three localities in Czechia and Yugoslavia.The male on slide 1383 was explicitly designated as holotype, and the other specimens as paratypes.Unfortunately, these slides cannot be found at the MMBC, and we have been unable to trace them elsewhere.The type material of this species must be regarded as lost.This is unfortunate, as A. tovornikae is considered to be a senior synonym of Nigronirmus atricapillae Soler- Cruz et al., 1984, from the same host (Gustafsson and Bush 2017).While this synonymy should not be controversial, considering the morphological similarities of the two species, any neotype designation for A. tovornikae will need to take the synonymy with N. atricapillae into consideration.For this, fresh material is needed.Discussion Dalgleish and Price (2003) stated that the only way to realistically deal with a super-species-rich genus like Myrsidea Waterston, 1915, is to circumscribe each revision to species of lice from the same host family; this practice is generally followed by taxonomists working on Myrsidea (e.g., Price and Johnson 2006, Sychra and Literák 2008, Kounek et al. 2011).Taken as a whole, the Brueelia-complex is more species-rich than Myrsidea, and the host range of the Brueelia-complex is similar to that of the genus Myrsidea.Any approach likely to make species identification and description within the Brueelia-complex easier is thus appealing.Is the approach used for Myrsidea then applicable to the Brueelia-complex as well?
In a wider perspective, using this approach in the Brueelia-complex is not without problems.Gustafsson and Bush (2015) and Gustafsson et al. (2018b) showed several examples of morphologically similar species of Brueelia occurring on different host families, and, conversely, species of Brueelia occurring on the same host family being morphologically different.
The species redescribed here show similar patterns.Most taxa treated here are fairly typical species for their respective host families.For instance, both B. ferianci and B. kratochvili have the head shape typical of Brueelia species parasitizing boreal (but not tropical or southern; Gustafsson and Bush in prep.)motacillids.The extensive dark pigmentation patterns of B. breueri are also typical of the species of Brueelia parasitizing many boreal fringillids.
However, the head shape of B. blagovescenskyi (Fig. 4) is more similar to Brueelia species on boreal motacillids (e.g., Fig. 50) than it is to B. pelikani from another emberizid host (Fig. 8).The same head shape is found in some undescribed species from cisticolid hosts (Gustafsson and Bush in prep.).Similarly, the lack of aps on male tergopleurites VI-VII in B. glizi (Fig. 20) is more similar to some species of Brueelia on North American passerellids (Gustafsson and Bush in prep.)than it is to any species of Brueelia known from fringillids.
Descriptions of new species in large genera like Brueelia and Guimaraesiella thus need to be done with caution, as the close relatives may parasitize different host families (Gustafsson andBush 2015, Bush et al. 2016).A simple comparison of a potential new louse species with only species found on the same host family may therefore not be sufficient.Unfortunately, of the 426 species of lice in this complex recognized by Gustafsson and Bush (2017; additional species have since been described by Mey 2017, Gustafsson et al. 2018a, b, c, 2019), less than half are identifiable from their original descriptions.Moreover, there are no published suggestions for species groups in Brueelia and Guimaraesiella to consult.Apart from species description and illustration, future taxonomic work on the Brueelia-complex should include attempts to delimit species groups within the larger genera of the complex (Brueelia, Guimaraesiella).In addition, it is vital that more already described species within this complex are examined critically and redescribed whenever possible.

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Złotorzycka 1964: 250.much displaced and widened at osculum.Ventral anterior plate small, rounded rectangular, hard to see in many specimens.Head chaetotaxy and pigmentation patterns as in Figure 50.Preantennal nodi moderate, not bulging.Pre-and postocular nodi moderate.Marginal temporal carina of moderate width, with median margin undulating.Gular plate lanceolate.Thoracic and abdominal segments and pigmentation patterns as in Figures 48, 49.
placed N. merulensis in the genus Turdinirmus Eichler, 1951, a genus superficially similar to Guimaraesiella, but very different from species of Brueelia known from thrushes in size, head shape, and head structure.It is therefore not at all clear what specimens Giebel actually examined, and what he means by "similar".