Morphological revision of the hyperdiverse Brueelia - complex (Insecta: Phthiraptera: Ischnocera: Philopteridae) with new taxa, checklists and generic key Author Bush, Sarah E. text Zootaxa 2017 2017-08-31 4313 1 1 443 journal article 32249 10.11646/zootaxa.4313.1.1 d8cc2cd8-8410-49aa-a75d-7a41d9f52b26 1175-5326 883161 A5Fdfba5-F992-44A8-84C2-1756C943C19B The Brueelia -complex Ansari (1956a) stated that Brueelia Kéler, 1936a , was a group unified primarily by abdominal characters. He considered the morphological characters of the head, male genitalia, and abdominal chaetotaxy to be “ minor ” or “insignificant” for dividing the genus into smaller groups. However, the phylogeny of Bush et al . (2016) shows that many characters previously treated as “ minor ” or “superficial” actually contain phylogenetic signal. Indeed, genital and preantennal characters, as well as the chaetotaxy of the abdomen, head, and legs appear to be shared derived characters that accurately delimit monophyletic clades of lice presently placed in the genus Brueelia . For Brueelia s. str. , the most comprehensive morphological characterization is that of Mey & Barker (2014) , who listed 22 different morphological and ecological characters. Their concept of Brueelia s. str. also includes species that here we place either in Olivinirmus Złotorzycka, 1964 or in Guimaraesiella Eichler, 1949 . Thus, their characters are most appropriate for delimiting the core genera within the Brueelia -complex. Part of the difficulty in clearly and unambiguously circumscribing either the Brueelia -complex or Brueelia s. str. lies in the large number of species in these groups, and in the great morphological variation between and within some groups. Clay & Tandan (1967) made the first attempt to characterise the Brueelia -complex as a whole. They used seven characters that are found in all known members of this complex, and they listed additional characters that are found in most of the genera they included in the Brueelia -complex. Here we outline the unique characters that keep the Brueelia -complex together, as well as some of the large variation in morphology within this complex. Diagnosis. The Brueelia -complex can be identified by the combined presence of the following 17 morphological characters: 1) ventral carina interrupted medially and continuing anteriorly towards the frons forming a clypeo-labral suture; 2) avs3 situated near bend of ventral carinae, more or less median to avs2, and not more anteriorly near vsms2 ; 3) trabecula absent; 4) s5–7 absent; 5) pronotum with 1 seta on each side; 6) prosternal plate absent; 7) metepisternum fully sclerotized laterally; 8) pleurites, if present, fused to tergites; 9) first apparent tergopleural plate (II) without setae on anterior margin; 10) no sts on anterior end of abdominal segment II; 11) no sts on male subgenital plate (abdominal segments VII–XI) (but sternal plate VII has setae if male subgenital plate is divided into sternal plate VII + subgenital plate as in e.g. Turdinirmoides n. gen. , Fig. 128 ); 11) no sts on female abdominal segments VII–XI, except when sternal plate VII is not modified into subgenital plate (as in Mirandofures kamena n. sp. , Fig. 97 ); 12) male tergopleurite IX+X separated from XI by definite suture; 13) male anogenital opening and anal setae dorsal; 14) subvulval plates absent; 15) female subgenital plate present at least as sternal plate VII; 16) endomeres, hypomere, and telomeres fused entirely into mesosomal complex; 17) parameres separate from basal apodeme, but may be fused to mesosome (as in Couala n. gen. , Figs 512 , 522 ). The following characters are found in most genera within the Brueelia -complex: 18) if dorsal preantennal suture is present, ads is located in or posterior to suture (except in Meropoecus Eichler, 1940 : Figs 494 , 499 ); 19) mts 3 only temporal macroseta [except in Titanomessor n. gen. , Fig. 212 ; Harpactrox n. gen. , Fig. 248 ; Rostrinirmus ruficeps (Nitzsch [in Giebel], 1866 ), Fig. 439 ; Motmotnirmus Mey & Barker, 2014 , Fig. 504 ]; 20) medianly divided tergopleural plates II–IX+X in male and II–VIII in females (except some Hecatrishula n. gen. ); 21) abdominal sternites III–VI in form of central plates (but may be absent in at least some segments as in some Sturnidoecus Eichler, 1944 , Figs 377–378 ; Manucodicola n. gen. , Figs 451–452 , 457–458; Motmotnirmus , Figs 502–503 ); 22) female subgenital plate reaches to vulval margin (except in e.g. Osculonirmus , Fig. 129 ; some Mirandofures n. gen. , Fig. 97 ; Turdinirmoides , Fig. 181 ); 23) lateral setae posterior to vulval margin on raised area of sternal plate (except in Buerelius Clay & Tandan, 1967 , Fig. 510 ); 24) vss thorn-like (except in Buerelius , Fig. 511 ). The phylogeny of Bush et al . (2016) showed that the closest relatives of the Brueelia -complex are a grade of genera primarily occurring on parrots and South American passerines. This group includes Neopsittaconirmus , Formicaphagus , Formicaricola , Paragoniocotes , Psittaconirmus , Theresiella , Forficuloecus , Psittoecus and Nyctibicola . These genera share the following characters with the Brueelia -complex: ventral carina interrupted medially and continuing anteriorly towards the frons forming a clypeo-labral suture; pronotum with 1 seta on each side; meso- and metanotum fused to form pterothorax; male anogenital opening and anal setae dorsal; trabecula absent; if dorsal preantennal suture is present, ads is located in or posterior to suture; prosternal plate absent; endomeres, hypomere, and telomeres fused entirely into mesosomal complex; parameres separate from basal apodeme; avs3 not situated near vsms2 ; pleurites, if present, fused to tergites; mts 2 only macroseta. However, members of all these genera can be separated from members of the Brueelia -complex by the presence of setae on the anterior margin of tergopleurite II; these setae are absent in the Brueelia -complex. The sclerotization of the metepisternum is always continuous from the lateral margin of the pterothorax to near the metasternum in the Brueelia -complex, but the extent of sclerotization varies among genera in this sister grade. In all these genera except Nyctibicola and Forficuloecus the tergopleurites are complete medianly, whereas in all Brueelia -complex lice except some Hecatrishula n. gen. tergopleurites II–IX+X in males and II–VIII in females are interrupted medianly. Sensilla s5–7 are absent in all Brueelia -complex genera, but present in some of these closely related genera. For these reasons, we do not presently consider any of these closely related louse genera to be members of the Brueelia -complex. However, future studies of this grade may suggest that a more natural circumscription of the Brueelia -complex must contain these taxa. Members of the Brueelia -complex share some characters with members of the Philopterus -complex, which often occur on the same hosts. Like the Philopterus -complex ( Mey 2004 ; Philopterinae, sensu Eichler 1963 ), the genera of the Brueelia -complex have a medianly interrupted ventral carina that extends anteriorly, and medianly divided tergopleurites II–IX+X in male and II–VIII in females (except some Hecatrishula ). However, the two complexes can be separated by the following characters: trabecula are present in the Philopterus -complex but are absent in the Brueelia -complex; metepisterna do not extend medianly in the Philopterus -complex, but do extend medianly to near the metasternum in the Brueelia -complex; prosternal plate is present in the Philopterus -complex but absent in the Brueelia -complex; subvulval plates present in Philopterus -complex but absent in Brueelia - complex; parameres fused to basal apodeme in Philopterus -complex, but separate from basal apodeme in Brueelia - complex. The following characters also separate most of the species in the two complexes: os macrosetae in Philopterus - complex (except Cincloecus Eichler, 1951a ), but microseta in the Brueelia -complex (except Rostrinirmus ruficeps , Fig. 450 ); pos macro- or mesoseta in several Philopterus -complex genera, but microsetae or absent in Brueelia - complex; ads is located on dorsal anterior plate in Philopterus -complex, but in or posterior to dorsal anterior suture (if present) in Brueelia -complex (except Meropoecus ); mts 1 macroseta and mts 2 often macroseta in Philopterus - complex, but both are microsetae in Brueelia -complex (except Motmotnirmus , Fig. 504 ). Mey & Barker (2014) included the Penenirmus -complex ( Penenirmus Clay & Meinertzhagen, 1938 , and Picophilopterus Ansari, 1947 ) within the Brueelia -complex. These two complexes are similar in that both lack trabecula and both have a medianly interrupted, anteriorly bent ventral carina. In addition, the metepisternum extends medianly to near the metasternum in both complexes. However, there are distinct differences. In the Penenirmus -complex, the parameres are fused to the basal apodeme and the anogenital opening is ventral/terminal; in the Brueelia -complex the parameres are articulated with the basal apodeme (but may be fused to the mesosome as in Couala ), and the anogenital opening is dorsal. Males have setae on the subgenital plate and both sexes have setae on the anterior margin of tergopleurite II in the Penenirmus -complex, but not in the Brueelia -complex. Other characters, with few exceptions, delimit the members of the Penenirmus - and Brueelia -complexes: tergopleurites II–VII are fused medianly in the Penenirmus -complex (except Pe . zeylanicus Dalgleish 1967 ), but not in the Brueelia -complex [except Hecatrishula docilis n. comb. ( Ansari 1956b )]. Setae mts 1 and sometimes os are macrosetae in the Penenirmus -complex, but these are microsetae in the Brueelia -complex [except in Ro . ruficeps (Nitzsch [in Giebel], 1866 ), Fig. 439 ]. Description. Apart from the characters listed above, no other morphological characteristics have been identified that are shared among all lice in this complex. The genera of the Brueelia -complex even vary enormously in gross morphology. For example, all four feather louse “ecomorphs” ( Johnson et al . 2012 ) known to occur on birds are found within the Brueelia -complex: “head lice”, “wing lice”, “body lice”, and “generalists”. Each of these ecomorphs vary in morphological traits that are associated with the different microhabitats of the body where they are most commonly found. Head lice have large, triangular heads, round bodies, and usually a widely interrupted marginal carina and derivate preantennal area (e.g. Schizosairhynchus n. gen. , Figs 463–464 ; Sturnidoecus Eichler, 1944 , Figs 377–378 ). Wing lice have long, slender bodies and smaller, triangular or trapezoidal, heads (e.g. Acronirmus Eichler, 1953 , Figs 83–84 ; Aratricerca n. gen. , Figs 168–169 ). Body lice have broadly rounded heads and rounded bodies (e.g. Saepocephalum n. gen. , Figs 238–239 ; Couala n. gen. , Figs 512– 513 ). Generalists have intermediate body forms and are commonly found roaming all over the body of the host (e.g. Mirandofures n. gen. , Figs 90–91 ). These ecomorphs are convergent forms that have evolved repeatedly within the Brueelia -complex (Bush et al . 2016). Coloration. Coloration within the Brueelia -complex varies from very light and almost translucent to heavily sclerotized and very dark. Extreme colour variation even occurs within some genera (e.g. Brueelia Kéler, 1936a ). Not all lice in the complex are uniformly pigmented, and in some species pigmentation is restricted to parts of the head, thoracic, and abdominal plates, which gives the lice a striated or otherwise distinct appearance (e.g. Corvonirmus Eichler, 1944 , Hecatrishula n. gen. , some Brueelia s. str. ). Louse colour is not necessarily phylogenetically informative. Bush et al . (2010) showed that colour variation among congeneric lice was associated with the colour of the host. Lice that are similar in colour to the hosts’ feathers are cryptic, and are less likely to be removed when the bird preens, which is a bird’s principal defense against ectoparasitic lice ( Bush et al . 2010 ). The potential for repeated evolution of cryptic evolution among congeneric lice suggests that the use of pigmentation patterns for generic delimitation (as in Złotorzycka 1964a ) may be misleading and should be used cautiously. colour may be informative at the species level, but if pigmentation patterns are used, this should be done cautiously and in combination with other characters, until more is known about colour variation within and between species of lice. Head. Head shape variable among genera within the complex ( Figs 1–9 ), as well as within some genera (e.g. Brueelia s. str. and Guimaraesiella Eichler, 1949 ). Marginal carina —variable ( Figs 10–18 ). Marginal carina may be uninterrupted ( Figs 10–11 ), interrupted submedianly ( Figs 12–13 ), laterally ( Figs 14– 15 ), or both ( Fig. 16 ). Laterally interrupted marginal carinae may be divided into pre- and postmarginal carinae ( sensu Clay 1951 , Fig. 85 ), or premarginal carina may be absent ( Figs 17–18 ). The section of the marginal carina between the submedian interruptions may be present ( Fig. 13 ) or absent ( Fig. 16 ). Median section of the marginal carina often displaced posteriorly and dorsally at osculum, in which case frons is hyaline ( Figs 11, 15 ). In some genera marginal carina is not displaced, and there is no hyaline margin ( Fig. 10 ). Displaced section of marginal carina may be present only as a discrete lateral thickening of the dorsal anterior plate (e.g. Aratricerca , Fig. 170 ) or as a medianly continuous sinuous thickening of the dorsal anterior plate (e.g. Resartor n. gen. , Fig. 163 ). Displaced section of marginal carina may extend posteriorly into a distinct marginal carinal plate (e.g. Mirandofures kamena n. sp. , MCP in Fig. 92 ). Hyaline margin —Hyaline margin generally confined to median section ( Fig. 11 ), but may extend along lateral sides of head ( Fig. 18 ). In Schizosairhynchus hyaline margin forms fleshy lobes on antero-lateral ends of head ( Figs 465 , 472 ). Dorsal preantennal sutures —may arise from either submedian or lateral interruptions of marginal carina, or both. Extent of dorsal preantennal sutures variable, even within genera (e.g. Guimaraesiella , Figs 361–364 ). Sutures range from minute dots near base of dsms (some Priceiella n. gen. , Fig. 287 ) to transversally and longitudinally continuous bands delimiting dorsal anterior plate (e.g. Sturnidoecus , Fig. 379 ). In Priceiella ( Thescelovora ) n. gen et n. subgen. ( Fig. 302 ) sutures are longitudinal and may reach both dsms and ads , but sutures do not connect medianly and do not reach margins of head. In Titanomessor n. gen. ( Fig. 212 ) and Harpactrox n. gen. ( Figs 248 , 255, 260) dorsal preantennal suture is transversal but does not reach the margins of the head. In Mirandofures ( Fig. 92 ) dorsal preantennal suture is transversally continuous but does not connect to the hyaline margin. Several genera (e.g. Corvonirmus , Fig. 321 ) lack dorsal preantennal sutures entirely. Dorsal anterior plate —present or absent, variable between and within genera. Plate, when present, may be completely separated from main head plate as in Sturnidoecus ( Fig. 379 ), but is often continuous with main head plate at least posteriorly as in many Guimaraesiella ( Fig. 356 ). In Priceiella ( Thescelovora ) ( Fig. 302 ) doral anterior plate is continuous with main head plate both antero-laterally and posteriorly. In some genera ( Titanomessor , Fig. 212 ; Harpactrox , Fig. 248 ) dorsal anterior plate is separated posteriorly but continuous anterolaterally. Posterior margin of dorsal anterior plate may be extended into spur overlapping with main head plate as in Schizosairhynchus n. gen. ( Fig. 465 ). Ventral carina and clypeo-labral suture —ventral carina interrupted medianly, bending anteriorly towards frons, thereby forming clypeo-labral suture medianly ( Fig. 20 ). Clypeo-labral suture reaches anterior margin of head in all genera except Anarchonirmus n. gen. ( Fig. 116 ), Harpactrox ( Fig. 248 ), and Saepocephalum ( Fig. 240 ). Ventral carinae displaced medianly at avs3 ( Fig. 20 ); may have median finger-like extension on each side near anterior end of pulvinus (e.g. Brueelia pseudognatha n. sp. , Fig. 65 ; Resartor , Fig. 163 ). In Harpactrox loeiensis n. sp. ( Fig. 248 ) lateral margins of clypeo-labral suture are not clearly delimited anterior to pulvinus. Ventral anterior plate —present or absent. When present generally crescent- or square-shaped. In groups where clypeo-labral suture does not reach anterior margin of head, ventral anterior plate not laterally separated from main head plate (e.g. Harpactrox , Fig. 255). Coni —small ( Figs 10–11 ) to large ( Fig. 18 ). Preantennal nodi —variable between species and genera. Preantennal nodi typically rounded, not extending median to the antennal socket, with defined lateral margin (e.g. Manucodicola n. gen. , Fig. 453 ). In many Brueelia s. str. (e.g. Br . pseudognatha , Fig. 65 ) lateral margin diffuse or absent. Preantennal nodi prominent in some genera (e.g. Traihoriella Ansari, 1947 , Fig. 265 ), and extend almost to base of mandibles in some species of Couala ( Fig. 520 ). Small accessory nodi median to preantennal nodi present in Harpactrox ( Fig. 248 ). Pre- and postocular nodi —variable between and within genera. Preocular nodi slightly to considerably larger than postocular nodi. Ocular nodi almost entirely enclose the eye in Nemuus n. gen. ( Fig. 340 ); nodi almost entirely absent in Acronirmus ( Fig. 85 ). Dorsal postantennal suture —present only in Meropoecus Eichler, 1940 ( Figs 494 , 499 ), where it is medianly continuous but often diffuse. Antennae— monomorphic in most genera. Sexually dimorphic antennae do, however, occur sporadically throughout the Brueelia -complex, and may differ even within genera. Among species with dimorphic antennae the male scapes and sometimes the male pedicels larger than those of females (e.g. Anarchonirmus , Figs 116–117 ; Ceratocista n. gen. , Figs 155–156 ). Flagellomeres generally constant in length and width between sexes even in genera where scapes and pedicel are dimorphic. In Melibrueelia Valim & Palma, 2015 , and Ceratocista ( Fig. 155 ) flagellomeres of male are wider than those of females. Male flagellomeres are large and bulbous in Anarchonirmus ( Fig. 116 ) and flagellomere II is extended distally along posterior margin in male Osculonirmus Mey, 1982a ( Fig. 124 ). Head chaetotaxy. Setae of the head generally as in Fig. 19 . In many genera, all setae named by Clay (1951) are present, but presence or absence of specific head setae are important in genus delimitations. Position of dsms and ads more or less constant throughout Brueelia -complex. In species with dorsal preantennal suture, dsms may be situated in (e.g. Guimaraesiella , Fig. 356 ), anterior to (e.g. Titanomessor , Fig. 212 ), or lateral to (e.g. Turdinirmoides australissimus n. sp. , Fig. 191 ) suture, but never median to suture; ads is situated in (e.g. Osculonirmus , Fig. 124 ) or posterior to suture (e.g. Brueelia phasmasoma n. sp. , Fig, 58), but never anterior to suture or on anterior margin or suture. Position of pas more or less constant throughout complex, as in Fig. 19 . Presence or absence and length of pns, pts and s1–4 diagnostic for some genera; s5–7 absent in all genera of the Brueelia -complex. In most genera pns, pts , and s1–4 are present, but pns is absent in e.g. Ceratocista ( Fig. 155 ) and both pns and s4 are absent in Brueelia s. str. ( Fig. 44 ). In Ro . ruficeps (Nitzsch [in Giebel], 1866 ) os is macroseta ( Fig. 439 ), but this is not the case in other species of this genus [e.g. Ro . buresi ( Balát, 1958 ) , Fig. 444 ], and not the case for any other genus treated here. Position of pos variable, either on eye or posterior to eye. In the genera Acronirmus ( Fig. 85 ), Ceratocista ( Fig. 155 ), Resartor ( Fig. 163 ), Aratricerca ( Fig. 170 ), and Turdinirmoides n. gen. ( Fig. 177 ) pos absent. All five mts are present in all genera of the Brueelia -complex except Acronirmus ( Fig. 85 ), in which mts 1 is absent. The only temporal macrosetae for most genera in complex is mts 3 . In Couala ( Fig. 514 ) mts 2 is ventral, thick, and may be curved ( Fig. 520 ). In Motmotnirmus Mey & Barker, 2014 ( Fig. 514 ) mts 2 is macroseta. In Harpactrox ( Fig. 249 ) all mts are microsetae. In Titanomessor ( Fig. 212 ) mts 3–5 are of roughly equal length in males, and mts 3–4 are of roughly equal length in females. If present, as1–3 marginal or submarginal. In many genera (e.g. Brueelia s. str. , Fig. 19 ) as3 absent; as2–3 absent in Acronirmus ( Fig. 85 ). In species with laterally interrupted marginal carina, as3 is situated either anterior to suture (e.g. Mirandofures , Fig. 92 ) or posterior to suture (e.g. Sturnidoecus , Fig. 379 ). Location of vsms2 variable, typically on lateral margins of clypeo-labral suture ( Fig. 19 ), but may be inside clypeo-labral suture [e.g. Br . brachythorax ( Giebel, 1874 ) , Fig. 44 ]; vsms1 typically lateral or slightly posterior to vsms2 . In all Brueelia -complex genera, avs3 is situated near median interruption of ventral carina, clearly separated from vsms2 ; avs1–2 typically close together, near anterior margin of preantennal nodi. In most genera avs1 is submarginal ( Fig. 20 ), but may be marginal ( Fig. 19 ). Position of pcs and mds more or less constant throughout complex, as in Fig. 19 . Thorax. Thoracic segments largely similar throughout subfamily. Prothorax —usually rectangular, broader than long. Proepisternum slender to broad; pleural ridges prominent in some genera (e.g. Resartor , Figs 161–162 ). Prosternal plate absent. Proepimera generally slender, oblique. Median section of proepimera generally widened anteriorly and posteriorly; posterior extension may be long and curl around coxa II (e.g. Anarchonirmus , Fig. 114 ). One pronotal post-spiracular seta ( ppss ) present on each side on postero-lateral corner of pronotum ( Fig. 26 ) or rarely on posterior margin of pronotum ( Schizosairhynchus , Figs 463–464 ); absent in Meropsiella Conci, 1941b ( Figs 479–480 ). Pterothorax— typically short and wide, roughly pentagonal in shape (see e.g. Brueelia s. str. , Figs 42–43 ); hexagonal in Aratricerca ( Figs 168–169 ). Meso- and metasternal plates may or may not be fused. Pteronotum not medianly divided, except partially in posterior end in Schizosairhynchus ( Figs 463–464 ). Metepisternum continuous from metasternum to lateral margin of pterothorax. Metepimeron much thickened in some genera (e.g. Indoceoplanetes n. gen. , Figs 217–218 ). Postero-lateral corner of pterothorax with 1 thorn-like (here pths ) and 1 trichoid seta (here ptrs ), typically on ventral side; ptrs absent in Osculonirmus ( Figs 122–123 ). Marginal mesometanotal setae ( mms ) variable, typically forming sparse rows along outer third of pterothorax on each side (e.g. Titanomessor , Figs 210–211 ). In some genera mms clustered near poster-lateral corners (e.g. Aratricerca , Figs 168–169 ), in other genera mms form continuous row along posterior margin (e.g. Sturnidoecus , Figs 377–378 ). Numbers of setae on pteronotum vary among genera and species. Leg chaetotaxy —illustrated for coxae, trochanters, and femora in Fig. 25 ; cI-d1, cI-a1–4, cI-v3, tI-v2, fI-p2–3, cII-d1, tII-a1, tII-d1, cIII-d1, tIII-a1, tIII-d1 are typically very small and inconspicuous, especially if specimens are small, darkly pigmented, or poorly mounted. Leg setae are generally constant throughout a given genus, and are often informative for delimitations of genera and higher levels. However, some groups show slight differences in leg chaetotaxy between species groups. Abdomen. Abdominal shape varies from slender (e.g. Acronirmus , Figs 83–84 ) to rotund (e.g. Schizosairhynchus , Figs 463–464 ). Typically, males shorter and broader than females. Tergopleurites— tergites and pleurites are fused into tergopleurites. Male tergopleurites II–IX+X divided medianly ( Fig. 26 a) except for male segment IX+X in some Hecatrishula ; in other Hecatrishula all tergopleurites connected medianly by narrow bridge (not illustrated). Female tergopleurites II–VIII divided medianly, tergopleurite IX+X transversally continuous ( Fig. 26 b), except in Aratricerca ( Fig. 169 ). Rarely, female tergopleurite IX+X fused with tergopleurite XI [e.g. Resartor , Fig. 162 ; Indoceoplanetes ( Capnodella ) n. gen. & n. subgen. , Fig. 225 ; Melibrueelia ]. Some genera (e.g. Priceiella , Figs 285–286 ) with conspicuous ridges anterior to spiracle openings (here pre-spiracular ridges, PSR in Fig. 285 ). In some genera tergopleurites do not reach lateral margins of abdomen (e.g. Anarchonirmus , Figs 114–115 ). Typically tergopleurites extend to ventral side of abdomen, but this is not the case in e.g. some Corvonirmus ( Figs 319–320 ). Pleural incrassations generally present at least in anterior segments, but often absent on segments VII–IX+X. Pleural incrassations absent in some genera (e.g. most Corvonirmus , Figs 319–320 ). Pleurites II with median anterior extensions in some genera (e.g. Meropsiella , Figs 479–480 ). Re-entrant heads of pleurites present or absent. Sternal plates —if present medianly continuous, typically rectangular, and not approaching ventral section of tergopleurites. Sternal plates absent in anterior segments of Manucodicola ( Figs 451–452 ) and all segments of some Sturnidoecus ( Figs 377–378 ) and Motmotnirmus ( Figs 502–503 ). Females with no sternal plates posterior to vulval margin. Lateral margins may be concave ( Anarchonirmus , Figs 114–115 ). In some genera sternal plates are crescent-shaped (e.g. some Sturnidoecus , Figs 420–421 ). Sternal plate II of Schizosairhynchus ( Figs 463–464 ) modified into transversal band. Accessory sternal plates may be present lateral to medial sternal plates or subgenital plate and pleurites in males [e.g. Brueelia audax ( Kellogg, 1896 ) , Fig. 71 ], females ( Meropoecus , Fig. 493 ), or both sexes ( Schizosairhynchus , Figs 463–464 ). In some Priceiella ( Fig. 277 ), accessory sternal plates are present lateral to male subgenital plate. In Aratricerca ( Figs 168–169 ) at least sternal plate II with thickened lateral and anterior margins. Male subgenital plate —typically from segment VII to posterior margin of abdomen; does not reach posterior margin in, e.g. Meropoecus ( Fig. 492 ); divided into sternal plate VII and vaguely delimited subgenital plate in Aratricerca ( Fig. 168 ) and Turdinirmoides ( Fig. 175 ). Male subgenital plate may continue around terminal end of the abdomen, forming small dorsal plate (e.g. Hecatrishula , Fig. 138 ). Male genital opening is dorsal, on segment IX+X, usually with three anal setae on each side. Female subgenital plate —variable, typically covering segments VII–VIII. Figures 27–41 introduces the terminology used for the shape of the female subgenital plate within the Brueelia -complex. If at least the median section of the distal margin of the subgenital plate is continuous with the vulval margin, the subgenital plate is said to reach the vulval margin ( Figs 29 , 35–38 ). If no section of the distal margin of the subgenital plate is continuous with the vulval margin, but at least the median section is close to the vulval margin (typically reaching to near the vss , e.g. Guimaraesiella , Fig. 360 ), the submarginal plate is said to approach the vulval margin ( Figs 30 , 33 ). In some genera (e.g. Osculonirmus , Fig. 89 ) the subgenital plate does not approach the vss ( Figs 27, 29 , 31 , 40 ). The distal end of the subgenital plate may be unmodified ( Figs 27–30 ), but in many genera the distal end is widened laterally and follows the vulval margin. Ansari (1956a) referred to this structure as a “cross-piece”. The term “cross-piece” is here limited to those cases where the widened section of the distal subgenital plate is continuous with the vulval margin ( Figs 35–40 ). The cross-piece may be partial ( Fig. 35 ) in which case it does not reach the lateral ends of the vulval margin, but in most species with a cross-piece, this structure reaches the lateral ends of the vulval margin, in which case it is said to be complete . The cross-piece may be narrow ( Fig. 36 ) or broad (37). In Anarchonirmus ( Fig. 83 ) the cross-piece is medianly displaced ( Fig. 38 ), as the median section of the cross-piece is not continuous with the vulval margin. If the lateral widenings of the subgenital plate are not marginal, these are referred to as lateral submarginal extensions ( Figs 32–34 ). Lateral extensions may be present as small bulges ( Fig. 32 ) that are limited to the median section, or reach the lateral margins of the vulval margin ( Figs 33–34 ), and may be narrow ( Fig. 33 ) or broad ( Fig. 34 ). In Saepocephalum ( Fig. 181 ) the sclerotized sections of the vulval margin are separated from the subgenital plate submedianly, and are referred to as lateral marginal plates ( Fig. 39 ). In Turdinirmoides ( Fig. 181 ) and Sychraella ( Fig. 113 ) the vulval margin is scleritozed but the subgenital plate does not reach the sclerotized section, which is referred to as a detached cross-piece ( Fig. 40 ). In Sychraella ( Fig. 41 ) the subgenital plate is detached from the sclerotized median section of the vulval margin, which is connected to lateral extensions that reach the lateral ends of the vulval margins; this structure is here termed a laterally submarginal cross-piece . Abdominal chaetotaxy —variable among genera ( Table 2 ) and within some genera ( Tables 3–12 ). Some setal characters are more or less constant throughout complex. No setae in anterior end of tergopleurite II. Typically no setae on subgenital plates of either sex, except oblique set of short setae in females ( Fig. 26 b). Trichoid seta of tergopleurite VIII often located submarginally in grove on tergopleurite. Presence or absence of groups of setae are very useful characters for delimiting genera as discussed below (see also Tables 3–12 ). The post-spiracular sensillus ( Clay 1954; here pss ) is often visible only in segments IV–V, but can be seen in more anterior segments in some genera (e.g. Anarchonirmus , Figs 114–115 ; Osculonirmus , Figs 122–123 ) where it is associated with psps in these segments. In Brueelia s. str. ( Figs 42–43 ), Acronirmus ( Figs 83–84 ), and some other genera, pss is present also on segments VI–VII, but this is generally not the case in the Brueelia -complex. The pss is lateral to and typically contiguous with the aperture of the psps , but this is not the case in e.g. Anarchonirmus ( Figs 114–115 ) or Corvonirmus ( Figs 319–320 ) where the pss does not touch the aperture of the psps . The pss may not differentiate genera within the Brueelia -complex, but may be useful in differentiating higher taxonomic levels. TABLE 2. Summary of abdominal chaetotaxy of segments II–VIII for the Brueelia -complex, including both genera and subgenera. In genera where chaetotaxy differs between species group (marked with *), only that of the type species is given here, and more detailed accounts are given separately under the respective genus entries and in Tables 3–12. Trichoid setae of segment VIII are present in all genera, and are not listed. Abbreviations: ps = paratergal seta; aps = accessory post-spiracular seta; psps = principal post-spiracular seta; tps = tergal posterior seta; ss = sutural seta; sts = sternal seta. Genus Sex ps aps psps tps ss sts Brueelia * M III–VIII V–VII VI–VIII – V–VIII II–VI F IV–VIII – VI–VII – – II–VI Teinomordeus M III–VIII III–VII V–VIII V–VIII III–VIII II–VI F II–VIII – V–VII – VIII II–VI Acronirmus M III–VIII VI–VIII VII–VIII VII–VIII V–VIII II–VI F III–VIII – VII–VIII – VII–VIII II–VI Mirandofures * M IV–VIII – – IV–VIII V–VIII II–VI F III–VIII – – – – II–VI Sychraella M III–VIII IV–V VI–VIII IV–VIII IV–VIII II–VI F III–VIII – – – – II–VI Anarchonirmus M IV–VIII II–V II–VI II–VIII II–VIII II–VI F IV–VIII – II–VI – II–VIII II–VI Osculonirmus M III–VIII – II–VIII – II–VIII II–VI F III–VIII – II–VIII – II–VIII II–VI Hecatrishula M III–VIII – III–VII II–VIII II–VIII II–VI F III–VIII – III–VII II–VIII II–VIII II–VI Psammonirmus . M IV–VIII – IV–VIII V–VIII II–VIII II–VI F IV–VIII – V–VIII V–VIII – II–VI Ceratocista M IV–VIII – V–VII VI–VIII VII–VIII II–VI F II–VIII – V–VIII – – II–VI Resartor * M IV–VIII – IV–VII VI–VIII II–VIII II–VI F IV–VIII – IV–VII – – II–VI ......continued on the next page Genus Sex ps aps psps tps ss sts Buphagoecus M IV–VIII IV–VII III–VIII VIII II–VIII II–VI F IV–VIII – IV–VIII – II–VIII II–VI Rostrinirmus * M IV–VIII II–VII II–VIII VII–VIII II–VIII II–VI F IV–VIII II–VII II–VIII – II–VIII II–VI Manucodicola M IV–VIII VI–VIII IV–VIII VI–VIII II–VIII II–VI F IV–VIII – IV–VIII – II–VIII II–VI Schizosairhynchus M IV–VIII IV–VIII IV–VIII II–VIII II–VIII II–VI F IV–VIII – IV–VIII II–VIII II–VIII II–VI Bizarrifrons M IV–VIII V–VII IV–VIII VI–VIII II–VIII II–VI F IV–VIII – IV–VII – II–VIII II–VI Meropsiella M IV–VIII – IV–VII VI–VIII II–VIII II–VI F IV–VIII – IV–VII – II–VIII II–VI Meropoecus M IV–VIII – II–VIII II–VIII II–VIII II–VI F IV–VIII – II–VIII II–VIII II–VIII II–VI Motmotnirmus M 3 IV–VIII – III–VIII VII–VIII II–VIII II–VI F 4 IV–VIII – III–VIII VIII II–VIII II–VI Buerelius M IV–VIII – IV–VII – II–VIII II–VI F IV–VIII – IV–VII – II–VIII II–VI Couala M III–VIII – VI–VII VII–VIII II–VIII II–VI F III–VIII – VI–VII – II–VIII II–VI Aporisticeras M IV–VIII – IV–VII VI–VIII II–VIII II–VI F IV–VIII – IV–VII – II–VIII II–VI

TABLE 2. (Continued) Genus Aratricerca	Sex M	ps III–VIII	aps –	psps IV–VII	tps ss – II–VIII	sts II–VII
Turdinirmoides * Turdinirmus Maculinirmus	F M F M F M	III–VIII III–VIII III–VIII IV–VIII IV–VIII IV–VIII	– – – – – –	V–VII IV–VIII IV–VIII IV–VII IV–VII IV–VII	– II–VIII V–VIII II–VIII – II–VIII – II–VIII – II–VIII – II–VIII	II–VI II–VII II–VI II–VI II–VI II–VI
Titanomessor Indoceoplanetes (Capnodella) Indoceoplanetes (Indoceoplanetes)	F M F M F M	IV–VIII IV–VIII IV–VIII IV–VIII IV–VIII IV–VIII	– – – – – –	IV–VII IV–VII IV–VII IV–VIII IV–VII IV–VIII	– II–VIII VII–VIII II–VIII – II–VIII – II–VIII – II–VIII IV–VIII II–VIII	II–VI II–VI II–VI II–VI II–VI II–VI
Saepocephalum Harpactrox Traihoriella *	F M F M F M	IV–VIII III–VIII III–VIII III–VIII III–VIII IV–VIII	– – – – – V–VII	IV–VII III–VII IV–VII IV–VIII IV–VIII IV–VII	– II–VIII IV–VII II–VIII – II–VIII – (VIII) II–VIII – II–VIII – V–VIII	II–VI II–VI II–VI II–VI II–VI II–VI
Priceiella (Priceiella) Priceiella (Camurnirmus) Priceiella (Thescelovora) Priceiella (Torosinirmus)	F M F M F M1 F M	IV–VIII III–VIII III–VIII IV–VIII IV–VIII IV–VIII IV–VIII III–VIII	– III–VII – IV–VIII – – – III–VII	IV–VII IV–VIII IV–VIII IV–VIII IV–VIII IV–VIII IV–VII IV–VIII	– VII–VIII – II–VIII – II–VIII – II–VIII – II–VIII – II–VIII – II–VIII – II–VIII	II–VI II–VI II–VI II–VI II–VI II–VI II–VI II–VI
Corvonirmus	F M	III–VIII III–VIII	– IV–VII	IV–VII II–VIII	– II–VIII III–VIII II–VIII	II–VI II–VI
Olivinirmus * Nemuus Melibrueelia Guimaraesiella * Sturnidoecus *	F M F M F2 M F M F M F	III–VIII IV–VIII IV–VIII II–VIII II–VIII IV–VIII IV–VIII IV–VIII IV–VIII IV–VIII IV–VIII	– IV–VII – II–VII II–VII III–VII – IV–VII – II–VIII II–VIII	III–VIII IV–VIII IV–VII (VIII) II–VII II–VII V–VII V–VII IV–VIII IV–VIII II–VIII II–VIII	– II–VIII – II–VIII – II–VIII IV–VIII II–VIII VI–VIII II–VIII VIII II–VIII – II–VIII – II–VIII – II–VIII II–VIII II–VIII II–VIII II–VIII ......continued on the	II–VI II–VI II–VI II–VI II–VI II–VI II–VI II–VI II–VI II–VI II–VI next page

TABLE 2. (Continued) 1 Males of some undescribed species of Thescelovora have aps on tergites VI–VII. 2 Nemuus hoedhri n. sp. has tps also on tergites II–V. 3 An undescribed species from Eumomota superciliosa australis Bangs, 1906 , has 1–3 tps on tergite VI. 4 Motmotnirmus humphreyi lacks tps on tergite VII. Male genitalia. Male genitalia are variable among genera ( Figs 21–24 ), and within some genera, where they form the basis of delimited species-groups (e.g. Olivinirmus Złotorzycka, 1964a , Figs 334–337 ; Sturnidoecus , Figs 384–398 ). In other genera ( Brueelia s. str. , Fig. 45 ; Acronirmus , Fig. 86 ) male genitalia are more or less identical in structure throughout genus, and only the shape of the genitalic elements differ between species. Most genera within the Brueelia -complex have similar genitalic structures that are described here and illustrated in Figs 21–24 . The structures of two genera with aberrant genitalia ( Meropoecus , Fig. 495 ; Aporisticeras n. gen. , Fig. 527–530 ) are not treated here, instead see relevant genus and species descriptions. All other genera treated here follow a general pattern. Basal apodeme —present, usually large. Lateral margins may be folded medianly in distal end. Lower endomere absent; however several genera have a ventral sclerite which may be remnant of lower endomere (e.g. Hecatrishula , Fig. 135 ; Corvinirmus , Fig. 324 ; Schizosairhynchus , Fig. 467 ). Male genitalia of Aporisticeras (Fig. 538) contain structures that may correspond to the lower endomere. In many genera extrusor muscles well defined as convergent ventral ridges (VR in Figs 22, 24 ). Mesosome— the endomeres, hypomere, and telomeres are fused entirely into mesosomal complex; distinct parts cannot be discerned clearly in any genus. To avoid any positive statements of homology we use terms that divide the mesosome into three clearly visible parts: proximal mesosome, mesosomal lobes, and gonopore (see Figs 21–24 ). Proximal mesosome —this term refers to the proximal end of the mesosome, which often overlaps with the posterior end of the basal apodeme and/or the parameres. Proximal mesosome may be homologous to antero-dorsal endomere ( sensu Ewing 1932 ) but is completely fused to other parts of mesosomal complex and this cannot be established with certainty. Shape typically variable between species in same genus. Ventral ridges distinctly convergent and wing-shaped (e.g. Rostrinirmus Złotorzycka, 1964a , Figs 442, 446, 449 ) or band-shaped (e.g. Guimaraesiella , Fig. 357 ). When present, ventral ridges usually reach or are continuous with gonopore distally and extrusor muscles proximally. Anterior margin of proximal mesosome may be thickened (e.g. Schizosairhynchus , Figs 467 , 474 ). In Couala ( Figs 515 , 521 ) proximal mesosome is fused to basal apodeme. Gonopore— present in most genera, but highly modified in some Sturnidoecus ( Figs 384–398 ). Gonopore may be entirely ventral ( Figs 22, 24 ), or may be located on terminal margin of mesosome ( Figs 21, 23 ). Gonopore typically visible as a circle or semicircle, and is typically open distally (e.g. Brueelia s. str. , Fig. 46 ), but may be open proximally as well, present as convergent median ridges (e.g. Turdinirmoides , Fig. 178 ). However, gonopore is usually a ventral dome, in which the ridges or circle are just the outline. Gonopore may be expanded laterally (e.g. Sychraella n. gen. , Fig. 111 ). Distally gonopore may extend posterior to mesosomal lobes (e.g. Hecatrishula atherae , Fig. 135 ). Mesosomal lobes —lateral structures of distal mesosome, variable between and within genera. Lobes may be homologous to endomeres, but fused to rest of mesosomal complex. In many groups the shape of lobes, as well as the shapes of structures (carinae, nodi, ridges) on lobes and the position of setae are one of the best ways to recognise closely related species. In some genera lobes form prominent lateral “wings” that overlap considerably with parameres (e.g. Hecatrishula , Fig. 135 ). Conversely, mesosomal lobes are almost entirely absent in genera like Guimaraesiella s. str. ( Fig. 358 ) and Traihoriella ( Fig. 274 ). Lateral and posterior margins usually with carina, distal margin may be fringed, scaly, brush-like, or rugose (e.g. Brueelia s. str. , Fig. 46 ; Psammonirmus n. gen. , Fig. 150 ; Ceratocista , Fig. 158 ). Postero-lateral corner in many genera with distinct rugose nodi (e.g. Nemuus , Fig. 351 ). In Couala ( Figs 516 , 522 ) mesosomal lobes are partially fused to parameral heads. Genital chaetotaxy —Up to 6 setae present on each side of mesosome, here tentatively divided into two sets. Anterior mesosomal setae ( ames ) are located ventrally, anterior to gonopore. Posterior mesosomal setae ( pmes ) are located lateral or posterior to gonopore, and may be on lateral margin of mesosome. In some genera (e.g. Priceiella , Fig. 289 ) both ames and pmes are anterior to gonopore; in these cases, we interpret the lateral set, which is typically associated with a marginal thickening of the mesosomal lobes, as ames , and the more median set, typically associated with the gonopore, as pmes . In Buphagoecus husaini ( Ansari, 1968 ) n. comb. both ames and pmes are located distal to gonopore ( Fig. 432 ) Other configurations also exist, e.g. Sturnidoecus ( Figs 384–398 ). Sensilliform pmes and ames are typically very hard to see, especially on mesosomes that are partially papillate, rugose, or with multiple ventral folds or thickenings. Lateral pmes and ames are often easy to overlook in noneverted gentalia. Distal part of paramere with 0–2 setae, here called parameral setae 1–2 ( pst1–2 , Fig. 21 ); pst1 typically sensillus located centrally on paramers; pst2 sensillus or microseta, located either centrally, laterally, or distally on paramere. Meropoecus ( Fig. 495 ) and Aporisticeras ( Fig. 529 ) unique within the Brueelia -complex in having more then two parameral setae. Parameral heads —shapes differ among genera. In some genera (e.g. some Brueelia , Fig. 61 ) heads are bluntly rounded with no modifications. In some genera (e.g. Osculonirmus , Fig. 128 ; Turdinirmus , Fig. 187 ) heads are bifid. In Hecatrishula ( Figs 136 , 144 ) the heads are divided into several anterior points. In Maculinirmus ( Figs 201 , 208) heads are folded laterally into narrow semi-circle. In many genera (e.g. Corvonirmus , Fig. 325 ; Guimaraesiella , Fig. 359 ; Bizarrifrons Eichler, 1938 , Fig. 478 ) heads extend medianly, and often overlap with the proximal mesosome; these median extensions are often rectangular (e.g. Bizarrifrons , Fig. 478 ), triangular (e.g. Nemuus , Fig. 344 ), or oblique finger-like (e.g. some Sturnidoecus , Figs 382 , 385 ), but other shapes occur. Parameral blades —variable among genera. Four basic shapes common: roughly triangular with lateral margin variable, often elongated distally (e.g. Brueelia s. str. , Fig. 47 ); irregularly oval, widening distally (e.g. Maculinirmus , Fig. 201 ); slenderly rounded-rectangular, tapering only in distal end (e.g. many Sturnidoecus , Fig. 382 ); elongatedly triangular (e.g. Nemuus , Fig. 344 ). In some genera, parameral blades are connected to parameral heads by slender parameral neck (e.g. Corvonirmus , Fig. 325 ). Parameral blades may be curled around mesosome (e.g. Schizosairhynchus , Fig. 466 ), highly convergent ( Titanomessor , Fig. 215 ), or highly divergent [ Priceiella ( Camurnirmus ) n. gen et n. subgen. , Fig. 298 ). Some genera have distinct “heel” on median margin just distal to mesosome (e.g. some Mirandofures , Fig. 104 ; Anarchonirmus , Fig. 120 ). Parameral blades of Meropsiella bullockoda ( Williams, 1981 ) have rugose bands ( Fig. 488 ). Female genitalia. Female genitalia more or less constant throughout complex. Vulval margin typically gently rounded, but highly convex in some genera (e.g. Osculonirmus , Fig. 129 ). Vulval chaetotaxy is identified in Fig. 26 b. Vulval chaetotaxy typically consisting of one marginal set of slender setae ( vms ), one submarginal set of thorn-like setae ( vss ), and one oblique set of slender setae ( vos ) following lateral margins of subgenital plate. Number, size, and position of setae vary among genera. In most groups, vss are situated clearly anterior to the vms , but in some groups [e.g. many Priceiella ( Thescelovora ) , Fig. 306 ] both vms and vss form a combined row of setae. In these groups, the number of setae in the two different set can still be counted separately as the vms and vss differ in shape and size. Distally convergent rows of vos typically situated lateral to submarginal plate, but some or all vos may be situated on plate (e.g. Schizosairhynchus , Fig. 469 ). If cross-piece is present, most vos are typically located anterior to cross-piece. However, 1–3 distal vos typically located near distal margin of subgenital plate. These distal vos are typically located median to the vss (e.g. Nemuus imperator n. sp. , Fig. 345 ). In many genera, there is a gap in the row of vos on each side, so that the distal 1–3 vos are separated from the more proximal ones (e.g. Resartor , Fig. 167 ). Typically, the vos distal to the gap are longer than the vms , but the same length as the more proximal vos . In addition, in most cases these distal vos are not situated on the vulval margin, but more proximally, often proximal to the vss . If the subgenital plate reaches the vulval margin, the distal vos are often situated on the surface of the subgenital plate (e.g. Brueelia , Fig. 48 ). If the subgenital plate does not reach the vulval margin, the distal vos are typically situated on the distal margin of the subgenital plate (e.g. Resartor , Fig. 167 ). Due to the difference in length between these setae and the vms , the more proximal placement of these setae, and the fact that in many groups there is no gap in the row of vos (e.g. Nemuus , Fig. 345 ) we treat all vos as one set, separate from the vms .