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 Rostrinirmus Złotorzycka, 1964 Nirmus Nitzsch, 1818 : 291 ( in partim ). Degeeriella Neumann, 1906 : 60 ( in partim ). Penenirmus Clay & Meinertzhagen, 1938 : 73 ( in partim ). Sturnidoecus Eichler, 1944 : 81 ( in partim ). Rostrinirmus Złotorzycka, 1964a : 276 . Type species. Rostrinirmus refractoriolus Złotorzycka, 1964a : 277 [= Nirmus ruficeps Nitzsch [in Giebel], 1866 : 367], by original designation. Diagnosis. In general morphology, Rostrinirmus ( Figs 437–450 ) is most similar to Sturnidoecus ( Figs 377–426 ) and Buphagoecus n. gen. ( Figs 427–436 ), and was previously considered part of Sturnidoecus . Like these two genera, the dorsal anterior plate of Rostrinirmus ( Fig. 439 ) is completely separated from the main head plate. However, the posterior margin of the dorsal anterior plate is rounded in Rostrinirmus ( Fig. 439 ) and the dorsal preantennal suture does not extend towards the preantennal nodi; in Sturnidoecus (e.g. Fig. 279 ) and Buphagoecus ( Fig. 429 ) the posterior margin of the dorsal anterior plate is more or less flat and one branch of the dorsal preantennal suture extends towards the preantennal nodi. The lateral section of the marginal carina is ventrally uninterrupted in both Rostrinirmus ( Fig. 439 ) and Buphagoecus ( Fig. 429 ), but partially interrupted ( Fig. 415 ) or at least displaced ( Fig. 379 ) laterally in Sturnidoecus (e.g. Fig. 379 ). The pns is present in Rostrinirmus ( Fig. 439 ) and Sturnidoecus (e.g. Fig. 379 ), but absent in Buphagoecus ( Fig. 429 ), and as2 is absent in all Rostrinirmus , but present in Buphagoecus and most Sturnidoecus (see this genus). Male genitalia of Rostrinirmus ( Figs 441–443, 445–450 ) most similar to those of Bizarrifrons ( Fig. 478 ) or Schizosairhynchus n. gen. ( Figs 466–468 , 474–476 ), but unlike those of any species-group in Sturnidoecus ( Figs 384–398 ) and Buphagoecus ( Figs 431–436 ). In Rostrinirmus (e.g. Fig. 449 ) and Schizosairhynchus (e.g. Fig. 467 ) the mesosomal lobes are fused distally, and the gonopore is ventral, whereas in Bizarrifrons ( Fig. 478 ) the mesosomal lobes are not fused and the gonopore is terminal. The male genitalia of Schizosairhynchus and Rostrinirmus can be separated by the following characters: ventral sclerite present in Schizosairhynchus (VS in Fig. 467 ) but absent in Rostrinirmus ( Fig. 449 ); parameral heads with angular median folds in Rostrinirmus ( Fig. 450 ) but with widened blunt proximal ends in Schizosairhynchus (e.g. Fig. 468 ); rugose nodi marginal in Rostrinirmus ( Fig. 449 ) but submarginal in Schizosairhynchus ( Fig. 467 ); pmes as sublateral sensilla in Rostrinirmus ( Fig. 449 ) but as lateral microsetae in Schizosairhynchus ( Fig. 467 ); marginal thickenings of mesosomal lobes continuous distally in Schizosairhynchus ( Fig. 467 ) but interrupted medio-distally in Rostrinirmus ( Fig. 449 ). However, these two genera are best separated by the following non-genitalic characters: marginal carina interrupted laterally in Schizosairhynchus ( Fig. 465 ) but not in Rostrinirmus ( Fig. 439 ); dorsal anterior plate short with rounded posterior margin in Rostrinirmus ( Fig. 439 ) but long with pointed posterior margin in Schizosairhynchus ( Fig. 465 ); as2 absent in Rostrinirmus ( Fig. 439 ) but present in Schizosairhynchus ( Fig. 465 ); accessory sternal plates present on at least some segments in both sexes in Schizosairhynchus ( Figs 463–464 ), but never present in Rostrinirmus ( Figs 437–438 ); sternal plate II of Rostrinirmus ( Figs 437–438 ) not modified as in Schizosairhynchus ( Figs 463–464 ); ppss on medio-posterior margin of prothorax in Schizosairhynchus ( Figs 463–464 ), but on postero-lateral corners in Rostrinirmus ( Figs 437–438 ). Description. Both sexes . Head bulb-shaped ( Fig. 439 ). Marginal carina interrupted only submedianly. Frons hyaline. Dorsal preantennal suture continuous with hyaline margin, reaching ads , dsms , and lateral margins of head, and separating dorsal anterior plate from main head plate posteriorly. Posterior margin of dorsal anterior plate rounded. Ventral carinae diffuse anterior to pulvinus, not clearly continuous with marginal carina. Ventral anterior plate present, broadly crescent-shaped. Head setae as in Fig. 439 ( Rostrinirmus ruficeps species group) or with setae along temporal margin as in Fig. 444 ( Ro . raji species group); as3 and as1 absent; pas spine-like; os macrosetae in Ro . ruficeps species-group ( Fig. 439 ) but microsetae in Ro . raji species group ( Fig. 444 ). Coni broad, reaching to or slightly beyond distal margin of scapes. Antennae monomorphic. Temporal carinae not visible; mts 3 only macrosetae. Gular plate spade-shaped or triangular. Prothorax rectangular to trapezoidal, widening posteriorly ( Figs 437–438 ); ppss on postero-lateral corners. Proepimera hook-shaped, curling around coxae II. Pterothorax pentagonal; lateral margins divergent; posterior margin convergent to rounded median point; mms narrowly interrupted medianly. Meso- and metasterna not fused; 1 seta on postero-lateral corner on each side of each plate. Metepisterna either slender ( Rostrinirmus raji speciesgroup) or widening ( Ro . ruficeps species-group) medianly. Leg chaetotaxy as in Fig. 25 , except fI-p2 absent; fII-v2, fIII-v2 absent in Ro . ruficeps but present in members of the Ro . raji species group (but see remarks under Ro . ruficeps ). Abdomen ( Figs 437–438 ) oblong in female, broadly oval in male. Tergopleurites triangular in male, more rectangular in female; tergopleurites II–IX+X in male and tergopleurites II–VIII in female widely separated medianly. Sternal plates broad, rectangular, do not approach pleurites. Pleural incrassations slight. Tergopleurites barely reach ventral surface of abdomen, often absent ventrally in females. Re-entrant heads wide, blunt, short, present mainly in anterior segments of male. Male subgenital plate triangular, with irregular lateral margins, reaching posterior end of abdomen. Female subgenital plate triangular, reaching or approaching vulval margin but not flaring into cross-piece ( Fig. 440 ). Abdominal chaetotaxy variable between species groups ( Table 12 ). Vulval margin ( Fig. 440 ) with slender vms , thorn-like vss ; vos follow lateral margins of subgenital plate; distal vos situated median to vss . TABLE 12. Chaetotaxy of male abdominal segments II–VIII of the species groups of Rostrinirmus . Trichoid setae of segment VIII are present in all species, and are not listed. Sets of setae differing from those of Ro. ruficeps are highlighted in bold . Material examined from all species is from their respective type hosts. Abbreviations: aps = accessory post-spiracular seta; psps = principal post-spiracular seta; ps = paratergal seta; ss = sutural seta; sts = sternal seta; tps = tergal posterior seta.

Species	Sex	ps	aps	psps	tps	ss	sts
Ro. ruficeps	M	IV–VIII	II–VII	II–VIII	VII–VIII	II–VIII	II–VI
Ro. buresi (representing the Ro. raji species group)	M	IV–VIII	II–VII	II–VIII	–	II–VIII	II–VI

Male genitalia different between species groups ( Figs 441–443, 445–450 ). Basal apodeme rectangular, anterior margin flat ( Rostrinirmus ruficeps species-group, Figs 445, 448 ) or rounded ( Ro . raji species-group, Fig. 441 ). Proximal mesosome broad, flat. Gonopore small, open distally, associated with subparallel club-like thickenings in Ro . raji species-group ( Fig. 442 ). Rostrinirmus ruficeps ( Fig. 445 ) with gonopore minute, and sometimes cannot be seen. Some Ro . ruficeps with asymmetrical mesosome as in Fig. 449 . Mesosomal lobes broad, fused distally; rugose nodi near distal end, more extensive medianly in Ro . raji species group ( Fig. 442 ) than in Ro . ruficeps species group ( Fig. 446 ); 2 ames microsetae sublaterally on each side anterior to gonopore; 1 pmes sensillus on each side near rugose nodi, distal to gonopore. Parameral heads ( Figs 443, 447, 450 ) folded, but differ between species-groups. Parameral blades broad; pst1 sensillus, central; pst2 microseta, central, near pst1 . Species-group characters. Two species-groups recognised on the basis of setae of the temporal margin of the head and abdomen and the shape of the male genitalia. Rostrinirmus raji species-group. os microsetae ( Fig. 444 ). Male abdominal chaetotaxy as in Table 12 ; tps present on tergopleurites VII–VIII. Basal apodeme rounded anteriorly ( Fig. 441 ). Gonopore prominent ( Fig. 442 ). Mesosomal lobes with distinct nodi in distal end and large oblique rugose nodi anterior to these ( Fig. 442 ). Parameral heads ( Fig. 443 ) folded, irregularly shaped, short. Rostrinirmus ruficeps species-group. os macrosetae ( Fig. 440 ). Male abdominal chaetotaxy as in Table 12 ; tps absent. Basal apodeme flat anteriorly ( Figs 445, 448 ). Gonopore not prominent ( Figs 446, 449 ). Mesosome symmetrical ( Fig. 446 ) or asymmetrical ( Fig. 449 ). Mesosomal lobes irregularly shaped, with small rugose nodi on posterior margin. Parameral heads with unique shape, much elongated ( Figs 447, 450 ). Host distribution. Most described species and our material examined are from members of either Passeridae or Emberizidae , but described and undescribed material from a few other families, such as Pycnonotidae and Sylviidae , suggest that the host range of Rostrinirmus may be wider. Also, some of the Sturnidoecus we listed as incertae sedis may ultimately belong to Rostrinirmus . These taxonomic decisions require examination of additional material. We have tentatively included all species originally described in Rostrinirmus , but we encourage future collections from these hosts to ascertain whether this is the proper placement of these taxa. Geographical range. Widely distributed across the Old World. Remarks. In the original description of Rostrinirmus , Złotorzycka (1964a: 276) only mentions the rounded dorsal anterior plate, but no other characters for differentiating her new genus from Sturnidoecus . Balát (1981b: 165) added that the male genitalia had a “peculiar shape”. Included species Rostrinirmus raji species-group: * Rostrinirmus boevi ( Balát, 1958: 418 ) n. comb. [in Penenirmus ] * Rostrinirmus buresi ( Balát, 1958: 416 ) n. comb. [in Penenirmus ] [1] Rostrinirmus rostratus Mey, 1982b : 181 new synonymy [2] Rostrinirmus carpodaci Balát, 1981b : 165 Rostrinirmus hudeci Balát, 1981b : 166 Rostrinirmus pflegeri Balát, 1981b : 166 * Rostrinirmus raji ( Ansari, 1947: 269 ) n. comb. [in Penenirmus ] Rostrinirmus tulackovae Balát, 1981b : 167 Rostrinirmus ruficeps species-group: * Rostrinirmus ruficeps (Nitzsch [in Giebel], 1866 : 367 ) [in Nirmus ] Philopterus suzume Uchida, 1949 : 548 Rostrinirmus refractoriolus Złotorzycka, 1964a : 277 [3] [1] Balát (1958) stated that Ro . buresi is close to Penenirmus [here Rostrinirmus ] ruficeps ( Nitzsch, 1866 ) , but did not illustrate it. Złotorzycka (1964a , 1968b , 1977 , 1997 ) and Balát (1981b) placed Ro . buresi in Rostrinirmus , together with Ro . ruficeps and Ro . refractoriolus . Złotorzycka (1997: 217) provided some illustrations of the male genitalia, which are similar to those of other Rostrinirmus species, but unlike those of Penenirmus . We agree with this placement, rather than the treatment of Ro . buresi as Penenirmus by Price et al . (2003 : 210). [2] Based on the few specimens of Ro. rostratus we have been able to examine, we find them indistinguishable from Ro. buresi . Mey (1982b: 181) used differences in the shape of the body and the female subgenital plate to differentiate these two species, but the shape of the lateral sides of most sternal plates and the subgenital plate for both Ro . buresi and Ro . ruficeps are subject to individual variation, with some specimens having flat lateral margins on one side and concave margins on the other. Therefore, this character is probably not reliable, given large enough samples. The measurements given by Mey (1982b) for Ro . rostratus all fall within the ranges given by Balát (1958) for Ro . buresi . We consider the two names synonymous. It is interesting to note that the two host species, Emberiza bruniceps Brandt, 1841 and E. melanocephala Scopoli, 1769 , readily hybridise where ranges overlap ( Byers et al . 1995 : 179, 182). [3] Rostrinirmus refractoriolus was first synonymized under Ro . ruficeps by Machácek (1977) , who noted that the material of Rostrinirmus he examined from Passer montanus and P. domesticus were indistinguishable. Złotorzycka (1964a: 277) did not have any material of Ro . ruficeps when she described Ro . refractoriolus , and did not provide any characters to differentiate the two species in her short description, only mentioning Ro . ruficeps briefly, and noting that it probably belonged to Sturnidoecus . Złotorzycka (1997: 216) later illustrated the male genitalia of Ro. refractoriolus and, despite the poor quality of the illustrations, the male genitalia are indistibguishable from those of Rostrinirmus from Passer montanus . We accept Machácek’s (1977) synonymization of these two species, based on the similarity of the male genitalia and the lack of distinguishing characters.