Brueelia Kéler, 1936

Bush, Sarah E., 2017, Morphological revision of the hyperdiverse Brueelia - complex (Insecta: Phthiraptera: Ischnocera: Philopteridae) with new taxa, checklists and generic key, Zootaxa 4313 (1), pp. 1-443 : 35-41

Brueelia Kéler, 1936

Philopterus Nitzsch, 1818: 288 (in partim). Nirmus Nitzsch, 1818: 291 (in partim).

Degeeriella Neumann, 1906: 60 (in partim). Brueelia Kéler, 1936a: 257 .

Painjunirmus Ansari, 1947: 285 .

Allobrueelia Eichler, 1951b: 36 (in partim). Nigronirmus Złotorzycka, 1964a: 248 .

Spironirmus Złotorzycka, 1964a: 261 .

Serinirmus Soler-Cruz, Rodríguez, Florido-Navío & Muñoz-Parra, 1987: 244 .

Type species. Brueelia rossittensis Kéler, 1936a: 257 [= Nirmus brachythorax Giebel, 1874: 134 ] (by original designation).

Diagnosis. Brueelia s. str. is part of a group of genera including Acronirmus, Teinomordeus n. gen., and Mirandofures n. gen., Sychraella n. gen., and Anarchonirmus n. gen., that share the following characters: absence of pns, as3, fII-v2, fIII-v2; parameral heads not folded medianly; absence of ames (except in species from Turdoides spp.); distal margin of mesosomal lobes at least partially rugose. All known Brueelia s. str. except Br. phasmasoma n. sp. ( Fig. 58 ) and Br. audax ( Fig. 70 ) lack a dorsal preantennal suture and have an uninterrupted marginal carina, which separates the genus from the otherwise similar genera Acronirmus ( Fig. 85 ) and Mirandofures ( Fig. 92 ) in all of which the marginal carina is interrupted at least laterally and dorsal preantennal suture is present. Antennae are sexually dimorphic in Teinomordeus ( Figs 77–78 ), Mirandofures ( Figs 92–93 ), Sychraella ( Figs 108–109 ), and Anarchonirmus ( Figs 116–117 ), but monomorphic in Brueelia s. str. ( Fig. 44 ) except in species on Turdoides spp.

Female Brueelia s. str. ( Fig. 48 ) is separated from females of other genera in this group on the structure of the subgenital plate, which always flares into a cross-piece in Brueelia s. str. In females of Teinomordeus ( Fig. 82 ) and Mirandofures ( Figs 97 , 105 ) there is no cross-piece, even in species where the subgenital plate reaches the vulval margin. Females of both Sychraella ( Fig. 113 ) and Anarchonirmus ( Fig. 121 ) have cross-pieces, but these are laterally submarginal and not connected to the subgenital plate in Sychraella , and medianly interrupted in Anarchonirmus . The subgenital plate of female Acronirmus ( Fig. 89 ) has a cross-piece as in Brueelia s. str., but most members of these two genera can be separated by non-genital characters, such as the presence in Acronirmus of a dorsal preantennal suture which is absent in most Brueelia s. str. In Brueelia s. str. species like Br. phasmasoma n. sp. where a dorsal preantennal suture is present, females can be separated from female Acronirmus by the following characters: as2 and pos present in Brueelia s. str ( Fig. 58 ), but absent in Acronirmus ( Fig. 85 ); ss present on tergopleurites VII–VIII in Acronirmus ( Fig. 82 ), but absent in Brueelia s. str. ( Fig. 57 ).

Description. Both sexes. Head shape variable, often flat-dome ( Fig. 4 ) or convex-dome ( Fig. 2 ), but may be narrow-triangle ( Fig. 5 ), drop-shaped ( Fig. 6 ), pentagonal ( Fig. 8 ) or trapezoidal ( Fig. 9 ). Marginal carina uninterrupted (except in Brueelia phasmasoma , Fig. 58 ); displaced dorsally and posteriorly at osculum; in some species narrow and translucent at osculum [e.g. Br. acutangulata ( Piaget, 1880) ; not illustrated]; median section possibly absent in species with a deeply concave frons (e.g. Br. chalcomitrae Najer & Sychra [in Najer et al.], 2012a; not illustrated). Displaced section in some species extending posteriorly into marginal carinal plate. Dorsal preantennal suture absent (except in Br. phasmasoma , Fig. 58 ). Ventral anterior plate present or absent. Ventral carinae typically clearly continuous with marginal carina, but diffuse anteriorly in some species. Finger-like extensions of ventral carina into clypeo-labral suture present in some species (e.g. Br. pseudognatha n. sp., Fig. 65 ). Head setae as in Fig. 44 : as3 absent or present; pas minute, possibly absent in some species; pns absent; s3 typically absent. Coni small. Antennae monomorphic, except in species parasitising Turdoides spp. where the males scape is slightly swollen compared to the female scape (not illustrated). Temporal carinae not visible; mts 3 only macrosetae. Gular plate generally triangular.

Prothorax ( Figs 42–43 , 49–50 , 56–57 , 63–64 ) rectangular; ppss on postero-lateral corner. Median ends of proepimera variable. Pterothorax bell-shaped or pentagonal; lateral margins weakly divergent; posterior margin rounded or convergent to median point; mms widely separated medianly. Meso- and metasterna not fused; one seta on postero-lateral corner on each side of each plate. Median ends of metepisterna variable, in some species associated with sublateral nodi. Leg chaetotaxy as in Fig. 25 , except cIv 3, fI-d1, fI-p2, fI-v4, fII-v2, fIII-v2 absent; fIm4 absent or present. Brueelia audax has aberrant leg chaetotaxy for genus (see below).

Abdomen oblong or elongated oval ( Figs 42–43 , 49–50 , 56–57 , 63–64 ), chaetotaxy as in Table 2 (but see Table 3 for some known exceptions). Tergopleurites rectangular to quadratic; tergopleurites II–IX+X in male and tergopleurites II–VIII in female narrowly to moderately divided medianly. Sternal plates medianly continuous, rectangular, may approach but not reach pleurites. Pleural incrassations slender; re-entrant heads typically long, slender.

Male. Subgenital plate trapezoidal or pentagonal, reaching posterior margin of abdomen. Genitalia ( Figs 45– 47 , 52–54 , 59–61 , 66–68 ) roughly homogeneous throughout genus. Basal apodeme small, oblong, and constricted along medial-lateral margins. Proximal mesosome small, typically rounded but elongated in some species (not illustrated). Gonopore ( Figs 46 , 53 , 60 , 67 ) slender to broad, widely open distally; gonopore often comparatively large. Mesosomal lobes small, rounded or angular; distal margin serrated, papillate, fringed, or rugose. Typically 2 pmes on each side on lobes just lateral to gonopore. Parameral heads ( Figs 45 , 54 , 61 , 68 ) cup-shaped, bifid, or blunt. Parameral blades broadly triangular, may be elongated distally; pst1 sensillus, typically near median margin; pst2 microseta, lateral near distal end.

Female. Subgenital plate pentagonal, reaching vulval margin where it flares into cross-piece (may be absent in Brueelia audax ; see below). Vulval margin ( Figs 48 , 55 , 62 , 69 ) with slender vms, thorn-like vss; vos follows lateral margins of subgenital plate; distal vos may be median to vss.

Host distribution. Brueelia s. str. occurs on a wide range of host families, but some host association patterns are discernable. The genus appears most widely distributed on host families that are largely or wholly Holarctic, or that have widely distributed representatives in the Holarctic area. Conversely, whereas some Brueelia are known from the tropics, the genus appears to be absent on most host families that are entirely tropical or subtropical. Species of Brueelia from the tropics are most often found on host families that also occur in the Holarctic. This pattern may be due to sampling bias, however the large material examined by us from museums across the world included many samples from the tropics, and the pattern appears to hold.

Geographical range. Species of Brueelia sensu stricto have a world-wide distribution, but are only known from introduced hosts in Australasia, where other genera of the Brueelia -complex occur.

Remarks. The circumscription of Brueelia proposed here removes roughly half the species of this genus listed in Price et al. (2003) to other genera, but makes the definition of Brueelia s. str. more homogeneous. However, the placement of Br. phasmasoma deep within Brueelia s. str. in the phylogeny of Bush et al. (2016; specimen #24) indicates that the genus still contains some variety in preantennal structure. Brueelia s. str. includes an enormous range of intrageneric variation: there is considerable variation in abdominal chaetotaxy, even between closely related species, and the range of pigmentation patterns is unequalled in other genera within the Brueelia -complex.

Several groups that we include in Brueelia s. str. may ultimately warrant recognition as subgenera. These include species formerly recognised in the genera Painjunirmus Ansari, 1947 , and Spironirmus Złotorzycka, 1964a , as well as the Br. ornatissima species group. Additional morphological and molecular work is needed to determine whether the monophyly of these groups are supported. We do not recognise or propose any new subgenera within Brueelia s. str. at this time.

We consider the following proposed genera (in totum) as synonyms of Brueelia s. str.: Painjunirmus Ansari, 1947 , Nigronirmus Złotorzycka, 1964a , Spironirmus Złotorzycka, 1964a , and Serinirmus Soler-Cruz, Rodríguez, Florido-Navío & Muñoz Parra 1987 .

Painjunirmus was erected by Ansari (1947: 285) for the interrupto-fasciata group ( sensu Piaget, 1880 ) of Degeeriella , and thus repeated the taxonomic acts of Kéler (1936a). The type species of Painjunirmus , P. pengya Ansari, 1947 , is an atypical Brueelia s. str., with male genitalia and abdominal chaetotaxy that differs from Br. brachythorax (see Table 3). Ultimately, with the examination of more material, species previously considered to be Painjunirmus may form a species group or subgenus within Brueelia s. str. This group would include the species Br. pengya , Br. magnini Ansari, 1956a, Br. chilchil Ansari, 1955b , and Br. brevipennis Ansari, 1956a .

1 All examined females poorly preserved, and tps and ss, if present, are not visible.

2 Chaetotaxy taken from original illustrations (Cicchino & Castro 1980). No material examined . Male unknown.

Nigronirmus was erected by Złotorzycka (1964a: 248); inclusion of species in this genus was largely based on pigmentation differences among these and species of Brueelia sensu Złotorzycka (1964a) . Structurally, these lice are all similar, and given the variability in colour among closely related lice in Brueelia s. str., we do not presently consider these differences in coloration to be sufficient justification for separating Nigronirmus from Brueelia s. str.

Spironirmus was erected by Złotorzycka (1964a: 261) and it included four species that were vaguely unified by “pleural loops” and male genitalia. Recent molecular data indicates that the type species, Brueelia nebulosa ( Burmeister, 1838) is nested deeply inside Brueelia s. str. (Bush et al. 2016), and most morphological characters used to separate Spironirmus from Brueelia s. str. are actually within the variation of Brueelia s. str. as circumscribed here. Thus, we retain Spironirmus as a synonym of Brueelia s. str. It should be noted that “pleural loops” are not found in three of the four species Złotorzycka (1964a) included in Spironirmus .

Serinirmus was established as a monotypic genus by Soler-Cruz et al. (1987) on the basis that Serinirmus sexytanum Soler-Cruz, Rodríguez, Florido-Navío & Muñoz Parra, 1987 , did not fit any of the five groups of genera proposed by Złotorzycka (1964a). We have not examined any specimens of Serinirmus sexytanum , but the original description and illustrations of Soler-Cruz et al. (1987) provide no morphological justification for the separation of Serinirmus from Brueelia s. str.

Included species

Brueelia abrupta ( Osborn, 1896: 229) [in Nirmus ] Brueelia acuminata Cicchino, 1982: 281

* Brueelia acutangulata ( Piaget, 1880: 156) [in Nirmus ] Brueelia elbeli Ansari, 1957b: 185 new synonymy [1]

* Brueelia aguilarae new species

* Brueelia albida ( Rudow, 1869: 21) [in Nirmus ]

* Brueelia alexandrii Eichler, 1953: 338

Brueelia alophoixi Sychra [in Sychra et al.], 2009: 155

* Brueelia altaica Mey, 1982b: 168

* Brueelia amazonae Stafford, 1943: 52 [2]

* Brueelia americana Cicchino & Castro, 1996: 21

* Brueelia anamariae Cicchino, 1980: 5

* Brueelia angustifrons ( Carriker, 1902: 221) [in Nirmus ]

* Brueelia antimarginalis Eichler, 1951b: 40 [page given as 12] Brueelia anumbii Cicchino, 1981: 37

* Brueelia argentina Cicchino, 1981: 31

Brueelia atricapillae Soler-Cruz, Alcántara-Ibañez & Florido-Navío, 1984: 145 [3] Brueelia paratricapillae Price, Hellenthal & Palma, 2003: 157 new synonymy

* Brueelia audax (Kellogg, 1899: 25) [in Nirmus ] [4] Brueelia badia Cicchino & Castro, 1996: 23

Brueelia balati Kristofík, 1999: 139

* Brueelia bicurvata ( Piaget, 1880: 159) [in Nirmus ]

* Brueelia blagovescenskyi Balát, 1955: 504

Brueelia boae Cicchino & Castro, 1996: 7

* Brueelia bonariensis Cicchino & Castro, 1996: 22 Brueelia borini Lunkaschu, 1970: 53

* Brueelia brachythorax ( Giebel, 1874: 134) [in Nirmus ] Brueelia rossittensis Kéler, 1936a: 257 [Ref.: Hopkins & Clay 1952: 61]

* Brueelia breueri Balát, 1955: 505

* Brueelia brevicolor Ansari, 1956c: 110

* Brueelia brevipennis Ansari, 1956a: 159

Brueelia calandrellae Fedorenko, 1975: 46

Brueelia cambayensis Ansari, 1955b: 54

Brueelia cantans Sychra [in Sychra et al.], 2010a: 62

* Brueelia cedrorum ( Piaget, 1880: 151) [in Nirmus ]

* Brueelia cela Stafford, 1943: 53 [2]

Brueelia chalcomitrae Najer & Sychra [in Najer et al.], 2012a: 92

* Brueelia chayanh Ansari, 1955b: 55

* Brueelia chelydensis Hopkins, 1951: 377

Nirmus vulgatus galapagensis Kellogg & Kuwana, 1902: 474 [Ref.: Hopkins & Clay 1952: 56]

* Brueelia chilchil Ansari, 1955b: 53

Brueelia chopi Valim & Cicchino, 2015: 509

* Brueelia chrysomystris ( Blagoveshtchensky, 1940: 60) [in Degeeriella ]

* Brueelia clara Gustafsson & Bush 2015: 505

* Brueelia conocephala ( Blagoveshtchensky, 1940: 87) [in Degeeriella ] Brueelia coquimbana Cicchino & González-Acuña, 2008: 302

* Brueelia corydalla Timmerman, 1950: 5

* Brueelia coryliventer Gustafsson & Bush 2015: 513

* Brueelia cruciata ( Burmeister, 1838: 429) [in Nirmus ] Brueelia cucullata Cicchino, 1982: 284

* Brueelia currucae Bechet, 1961: 153

Brueelia cyanopa Cicchino, 2004: 73

* Brueelia cyclothorax ( Burmeister, 1838: 429) [in Nirmus ] [5] Nirmus subtilis Nitzsch [in Giebel], 1874: 137 new synonymy Brueelia subtilis obligatus Eichler, 1954: 63 [Ref.: Emerson 1972: 34] Brueelia decumana Cicchino & Castro, 1996: 12

* Brueelia deficiens ( Piaget, 1885: 23) [in Nirmus ]

Nirmus ampullatus Piaget, 1885: 25 [Ref.: Williams 1986: 431]

* Brueelia delicata (Nitzsch [in Giebel], 1866: 368) [in Nirmus ]

* Brueelia densilimba (Nitzsch [in Giebel], 1866: 368) [in Nirmus ] Brueelia diucae Cicchino & González-Acuña, 2009: 505

* Brueelia dorsale Williams, 1983: 600

* Brueelia ductilis ( Kellogg & Chapman, 1899: 89) [in Nirmus ] Brueelia embernagrae Cicchino & Castro, 1980: 83 Brueelia emersoni Cicchino & Castro, 1996: 18

Brueelia exigua (Nitzsch [in Giebel] 1866: 366) [in Nirmus ]

* Brueelia ferianci Balát, 1955: 508

* Brueelia flinti Cicchino & Castro, 1996: 27

* Brueelia fulmeki Eichler, 1957: 580

* Brueelia fuscopleura ( Blagoveshtchensky, 1951: 303) [in Degeeriella ] [6] Brueelia gulabitilyar Ansari, 1955a: 54 new synonymy

* Brueelia ginginianus Ansari, 1955b: 55

* Brueelia glizi Balát, 1955: 509

* Brueelia gobiensis Mey, 1982b: 170

* Brueelia goertae Dalgleish, 1971: 141

* Brueelia guldum Ansari, 1955b: 54

* Brueelia iliaci ( Denny, 1842: 51) [in Nirmus ]

Brueelia inornata Timmermann, 1950: 3 [Ref.: Ansari 1956c: 110] Brueelia iliaci indiensis Ansari, 1956c: 111 [Ref.: Price et al. 2003: 155]

* Brueelia immaculata ( Piaget, 1890: 230) [in Nirmus ]

* Brueelia imponderabilica Eichler, 1954: 61

* Brueelia infrequens ( Carriker, 1902: 220) [in Nirmus ]

* Brueelia infuscata Cicchino, 1979: 92

* Brueelia intermedia (Nitzsch [in Giebel], 1866: 366) [in Nirmus ] [7]

* Brueelia jacarinae Valim & Palma, 2006: 28

* Brueelia jacobi Eichler, 1951b: 10

* Brueelia juno ( Giebel, 1874: 137) [in Nirmus ]

Brueelia kalkalichi ( Ansari, 1955b: 58) [in Philopterus ] [8]

* Brueelia kistiakowskyi Fedorenko, 1975: 48

* Brueelia kluzi Balát, 1955: 512

* Brueelia kratochvili Balát, 1958: 413

Brueelia latiuscula ( Kellogg & Chapman, 1899: 90) [in Nirmus ] [9]

* Brueelia limbata ( Burmeister, 1838: 429) [in Nirmus ] Docophorus serenus Rudow, 1869: 16 [Ref.: Hopkins & Clay 1952: 61] Brueelia limnothlypiae Valim & Reiley, 2015: 855

* Brueelia locustellae Fedorenko, 1975: 49

Brueelia longifrons Carriker, 1956a: 81

* Brueelia longipes ( Piaget, 1880: 160) [in Nirmus ]

* Brueelia lullulae Bechet, 1961: 154

* Brueelia magnini Ansari, 1956a: 161

Brueelia marcoi Cicchino & Castro, 1996: 20

* Brueelia matvejevi Balát, 1981a: 278

Brueelia mauroi Cicchino & Castro, 1996: 17

Brueelia mediterranea Cicchino, 1981: 34

* Brueelia melanocoryphae Bechet, 1966: 79

Brueelia mimas Cicchino & Castro, 1996: 28

* Brueelia minor Lunkaschu, 1970: 55

* Brueelia mirabile Carriker, 1963: 306

* Brueelia modularis ( Piaget, 1880: 151) [in Nirmus ]

* Brueelia mongolica Mey, 1982b: 172

* Brueelia moreli Ansari, 1957b: 182

* Brueelia museiberolinensis ( Eichler, 1957: 579) [in Allobrueelia ]

* Brueelia nebulosa ( Burmeister, 1838: 429) [in Nirmus ] Docophorus ochroleucus Nitzsch [in Giebel], 1874: 90 [Ref.: Hopkins & Clay 1952: 59] Brueelia chitlatilyar Ansari, 1955b: 55 [Ref.: Emerson 1972: 31]

* Brueelia nivalis ( Giebel, 1874: 140) [in Nirmus ]

* Brueelia ornatissima ( Giebel, 1874: 144) [in Nirmus ] Nirmus illustris Kellogg, 1896: 494 [Ref.: Hopkins & Clay 1952: 56] Brueelia oxypyga ( Giebel, 1874: 135) [in Nirmus ]

* Brueelia pagodarum Ansari, 1955b: 55

* Brueelia pakistanaise Ansari, 1955b: 52

* Brueelia parabolocybe ( Carriker, 1903: 137) [in Nirmus ] Brueelia paradoxa Valim & Cicchino, 2015: 515

* Brueelia parae Ansari, 1955b: 57

* Brueelia parviguttata ( Blagoveshtchensky, 1940: 62) [in Degeeriella ]

* Brueelia pelikani Balát, 1958: 414

* Brueelia pengya ( Ansari, 1947: 285) [in Painjunirmus ]

* Brueelia peninsularis (Kellogg, 1899: 21) [in Nirmus ]

* Brueelia phasmasoma new species

* Brueelia picturata ( Osborn, 1896: 226) [in Nirmus ]

* Brueelia piechockii Mey, 1982b: 157

Brueelia plocea ( Lakshminarayana, 1968: 99) [in Nigronirmus ] Brueelia priniae Najer & Sychra [in Najer et al.], 2012a: 94

* Brueelia propinqua ( Giebel, 1874: 136) [in Nirmus ]

* Brueelia pseudognatha new species

* Brueelia pseudopicturata Cicchino, 1986a: 8

* Brueelia pyrrhularum Eichler, 1954: 62

* Brueelia quelea Sychra & Barlev [in Sychra et al.], 2010b: 18

* Brueelia rhipidura ( Thompson, 1941: 533) [in Degeeriella ]

* Brueelia rigbyi Gustafsson & Bush 2015: 510

* Brueelia rosickyi Balát, 1955: 517

* Brueelia rotundifrons Cicchino, 1981: 38

Brueelia montana Williams, 1983: 600

* Brueelia ruficapilla Cicchino, 1990: 22

Brueelia sallei Carriker, 1963: 307

Brueelia sayacae Cicchino, 1982: 284

Brueelia scotocercae ( Blagoveshtchensky, 1951: 304) [in Degeeriella ] Brueelia senegala Sychra [in Sychra et al.], 2010a: 65 Brueelia sexytana (Soler-Cruz, Rodríguez, Florido-Navío & Muñoz Parra, 1987: 244) [in Serinirmus ]

* Brueelia sibirica Mey, 1982b: 174

* Brueelia solitari a Cicchino, 1990: 22

* Brueelia stadleri Eichler, 1954: 61

* Brueelia straminea ( Denny, 1842: 53) [in Nirmus ] Degeeriella sublucida Blagoveshtchensky, 1940: 58 [Ref.: Hopkins & Clay 1952: 62] Brueelia fixa Złotorzycka, 1964a: 256 [Ref.: Dalgleish 1971: 144] Brueelia tersinae Cicchino, 1982: 279

Brueelia thilia Cicchino, 2004: 75

* Brueelia tkachi Gustafsson & Bush 2015: 510

* Brueelia trinidadensis Cicchino & Castro, 1996: 21 Brueelia trithorax ( Burmeister, 1838: 429) [in Nirmus ]

* Brueelia vaneki Balát, 1982: 277

Brueelia virgata (Kellogg, 1899: 19) [in Nirmus ]

* Brueelia vulgata ( Kellogg, 1896: 496) [in Nirmus ] Brueelia weberi Balát, 1982: 44

* Brueelia xanthocephali ( Osborn, 1896: 224) [in Nirmus ] Brueelia xanthocollis Ansari, 1955b: 56

Brueelia yal Cicchino & González-Acuña, 2008: 303

* Brueelia zavattariornis Ansari, 1956b: 387

* Brueelia zohrae Ansari, 1956b: 389

[1] There are no apparent differences between Br. acutangulata and Br. elbeli in the material we have examined (Appendix III), and the original description of Br. elbeli Ansari 1957b does not provide sufficient diagnostic characters to separate Br. elbeli from Br. acutangulata .

[2] Having examined type material of both species (Appendix III), we agree with the treatment of Br. amazonae and Br. cela as separate species following Cicchino & Castro (1996), despite the treatment of these two as conspecific by Hopkins & Clay (1952), Carriker (1955), and Price et al. (2003).

[3] As noted by Martín-Mateo (2009: 340), the difference of a single letter in a species epithet prevents homonymy according to article 57.6 of the I.C.Z.N. (International Commission on Zoological Nomenclature, 1999); thus, the erection of a new name for this species by Price et al. (2003: 157) was unjustified.

[4] Brueelia audax is a very aberrant member of Brueelia s. str. (see below), and most of the material examined (including all females) are poorly preserved. Examinations of future collections of this species may warrant the erection of a new subgenus, or even genus, for this species.

[5] We examined material of Br. subtilis and Br. s. obligatus from the two known host species ( Passer montanus and P. domesticus ) (Appendix III). These specimens are inseparable from Br. cyclothorax . Although the poor condition of type material makes direct comparisons difficult, we consider these species to be the same (as in Złotorzycka 1997: 145) despite the treatment of these as separate species by Price et al. (2003: 159).

[6] We consider Br. gulabitilyar to be a synonym of Br. fuscopleura , based on the specimens we examined from Pastor roseus (the type host of Br. gulabitilyar ; Appendix III). The original description of Br. gulabitilyar Ansari (1955b) is also inseparable from the vague description of Degeeriella cruciata fuscopleura by Blagoveshchensky (1951).

[7] We disagree with the synonymization of Br. intermedia with Br. marginata by Price et al. (2003). Based on morphology of material examined and the original descriptions, we place Br. intermedia in Brueelia s. str., whereas we place Br. marginata in Guimaraesiella.

[8] We agree with Mey & Barker (2014: 114) that Philopterus kalkalichi as described by Ansari (1955b, 1958a) and illustrated by Ansari (1958a) is actually a Brueelia s. str. Ansari later redescribed and illustrated this species ( Ansari 1956d) and in doing so contradicted his earlier work, as the illustration in Ansari (1956d) is a Philopterus . It seems that Ansari himself confused two different species found on the same host. As the original description ( Ansari 1955b) refers to characters that are not found in Philopterus but are found in Brueelia s. str., we disagree with Palma & Price (2006) and Sychra et al. (2011), and consider this species a member of Brueelia s. str. We agree with Mey & Barker (2014) that Philopterus kalkalichi sensu Ansari (1956d) needs a new name, but this is outside the scope of this study. Interestingly, no other Brueelia s. str. are known from the Dicruridae , and this host association may be a case of straggling.

[9] We disagree with Price et al. (2003) about the synonymy of Br. latiuscula as Br. ornatissima . Emerson (1972) treated the two as separate (this was mis-cited in Price et al. 2003, and may be the source of error). Brueelia ornatissima has dark ventral carinae, whereas the illustrations of Br. latiuscula has translucent ventral carinae.

Figures

FIGURES 25. Dorso-ventral views of coxae I – III, trochanters I – III, and femora I – III, showing terminology of the leg setae used here. Other parts of the thorax removed for clarity. Setae situated marginally are illustrated and named on both dorsal (lefthand) and ventral (right-hand) sides. Terminology for individual seta as used in the text obtained by combining the leg segment (larger font) and the setal name (smaller font). Abbreviations: a 1 – 5, anterior setae 1 – 5; cI – III, coxae I – III; d 1 – 3, dorsal setae 1 – 3; dm 1 – 4, distal marginal setae 1 – 4; fI – III, femur I – III; p 1 – 3, posterior setae 1 – 3; s 1 – 6, sensilla 1 – 6; tI – III, trochanter I – III; v 1 – 4, ventral setae 1 – 4.

FIGURES 58 – 62. Brueelia phasmasoma n. sp. ex Coereba flaveola luteola: 58, male head, dorsal and ventral views. 59, male genitalia, dorsal view. 60, male mesosome, ventral view. 61, male paramere, dorsal view. 62, female subgenital plate and vulval margin, ventral view.

FIGURES 70 – 74. Brueelia audax (Kellogg, 1899) ex Auriparus flavifrons ornatus: 70, male head, dorsal and ventral views. 71, male habitus, dorsal and ventral views. 72 male genitalia, dorsal view. 73, male mesosome, ventral view. 74, male paramere, dorsal view.

FIGURES 85 – 89. Acronirmus gracilis (Burmeister, 1838) ex Delichon urbicum urbicum: 85, male head, dorsal and ventral views. 86, male genitalia, dorsal view. 87, male mesosome, ventral view. 88, male paramere, dorsal view. 89, female subgenital plate and vulval margin, ventral view. Abbreviations: POMC, postmarginal carina; PRMC, premarginal carina.

FIGURES 92 – 97. Mirandofures kamena n. gen. & n. sp. ex Erythrura trichroa sigillifer: 92, male head, dorsal and ventral views. 93, female antenna, ventral view. 94, male genitalia, ventral view. 95, male mesosome, ventral view. 96, male paramere, dorsal view. 97, female subgenital plate and vulval margin, ventral view. Abbreviation: MCP, marginal carinal plate.

FIGURES 77 – 82. Teinomordeus entelosetus n. gen. & n. sp. ex Eurocephalus rueppelli: 77, male head, dorsal and ventral views. 78, female antennae, ventral view. 79, male genitalia, dorsal view. 80, male mesosome, ventral view. 81, male paramere, dorsal view. 82, female subgenital plate and vulval margin, ventral view.

FIGURES 108 – 113. Sychraella sinsutura n. gen. & n. sp. ex Pomatostomus isidorei isidorei: 108, male head, dorsal and ventral views. 109, female antenna, ventral view. 110, male genitalia, dorsal view. 111, male mesosome, ventral view. 112, male paramere, dorsal view. 113, female subgenital plate and vulval margin, ventral view.

FIGURES 116 – 121. Anarchonirmus albovittatus n. gen. & n. sp. ex Pomatostomus temporalis strepitans: 116, male head, dorsal and ventral view; dorsal sensilla and pts not visible in the material examined. 117, female antenna, ventral view. 118, male genitalia, dorsal view. 119, male mesosome, ventral view. 120, male paramere, dorsal view. 121, female subgenital plate and vulval margin, ventral view (anterior dotted line outline pigmented part of subgenital plate, posterior dotted line extent of marginal thickening).

FIGURES 44 – 48. Brueelia brachythorax (Giebel, 1874) ex Bombycilla garrulus garrulous: 44, male head, dorsal and ventral views. 45, male genitalia, dorsal view. 46, male mesosome, ventral view. 47, male paramere, dorsal view. 48, female subgenital plate and vulval margin, ventral views.

FIGURES 100 – 105. Mirandofures altoguineae n. gen. & n. sp. ex Oreostruthus fuliginosus: 100, male head, dorsal and ventral views. 101, female antenna, ventral view. 102, male genitalia, dorsal view. 103, male mesosome, ventral view. 104, male paramere, dorsal view. 105, female subgenital plate and vulval margin, ventral view.

FIGURES 56 – 57. Brueelia phasmasoma n. sp. ex Coereba flaveola luteola: 56, male habitus, dorsal and ventral views. 57, female habitus, dorsal and ventral views.

FIGURES 1 – 9. Head morphology of the Brueelia - complex (not drawn to scale). Head shapes are: 1, concave dome-shaped; 2, convex dome-shaped; 3, indented dome-shaped; 4, flat dome-shaped; 5, wedge-shaped; 6, drop-shaped; 7, bulb-shaped; 8, pentagonal; 9, trapezoidal.

FIGURES 65 – 69. Brueelia pseudognatha n. sp. ex Pycnonotus nigricans superior: 65, male head, dorsal and ventral views. 66, male genitalia, dorsal view. 67, male mesosome, ventral view. 68, male paramere, dorsal view. 69, female subgenital plate and vulval margin, ventral view.

FIGURES 42 – 43. Brueelia brachythorax (Giebel, 1874) ex Bombycilla garrulus garrulus: 42, male habitus, dorsal and ventral views. 43, female habitus, dorsal and ventral views.

FIGURES 49 – 50. Brueelia aguilarae n. sp. ex Euplectes franciscanus pusillus: 49, male habitus, dorsal and ventral views. 50, female habitus, dorsal and ventral views. Thin dotted lines delimit pigmented areas on head, pterothorax, sternites, and tergite IX + X.

FIGURES 63 – 64. Brueelia pseudognatha n. sp. ex Pycnonotus nigricans superior: 63, male habitus, dorsal and ventral views. 64, female habitus, dorsal and ventral views.

FIGURES 51 – 55. Brueelia aguilarae n. sp. ex Euplectes franciscanus pusillus: 51, male head, dorsal and ventral views. 52, male genitalia, dorsal view. 53, male mesosome, ventral view. 54, male paramere, dorsal view. 55, female subgenital plate and vulval margin, ventral views.