Nitzsch, 1818 : 291 Kéler, 1936a : 257 Ansari, 1947 : 290 Traihoriella binhchauensis Brueelia Ansari, 1947 : 290 Traihoriella laticeps Ansari, 1955a: 51 Brueelia Piaget, 1888: 152 Nirmus Carriker, 1954 : 199 Morphological revision of the hyperdiverse Brueelia - complex (Insecta: Phthiraptera: Ischnocera: Philopteridae) with new taxa, checklists and generic key Bush, Sarah E. Zootaxa 2017 2017-08-31 4313 1 1 443 Ansari, 1947 Ansari 1947 [151,462,692,718] Insecta Philopteridae Traihoriella GBIF Animalia Phthiraptera 162 163 Arthropoda genus     Nirmus  Nitzsch, 1818: 291( in partim).  Brueelia  Kéler, 1936a: 257( in partim).  Traihoriella  Ansari, 1947: 290.   Type species.  Traihoriella punjabensis Ansari, 1947: 290, by original designation.   Diagnosis.The male genitalia of  Traihoriella( Figs 266–268, 273–275) are similar to those of  Harpactrox  n. gen.( Figs 249–251, 256–258), and may not be separable when more species of both genera have been examined and described. However, species in these genera are readily separated by non-genitalic characters; see  Harpactrox(above) for a more detailed comparison of unique non-genitalic characters in these two genera.   Traihoriellashare the following set of characters with  Corvonirmus, Priceiella  n. gen., and  Olivinirmus: parameral heads folded medianly; marginal carina uninterrupted but displaced at osculum and frons hyaline. These genera, however, differ in several characteristics.  In  Priceiella( Figs 284, 291, 306, 314) the female subgenital plate flares into a cross-piece at vulval margin, which is not present in  Traihoriella( Figs 269, 276). Furthermore, in  Priceiella( Figs 282, 289, 304, 312) the gonopore is located in the central part of the ventral side, whereas in  Traihoriellait is subterminal ( Figs 267, 274). In  Olivinirmus( Figs 327–328),  tpsare absent and  apsare present in roughly the same segments as in  Traihoriella, and the preantennal structure is largely the same in both genera, including the absence of dorsal preantennal sutures ( Figs 265, 272, 329). The female subgenital plate does not flare into a cross-piece in either genus ( Figs 269, 276, 333). The mesosome is more complex in  Olivinirmus( Figs 334–337) than in  Traihoriella( Figs 266–268, 273–275); gonopore is terminal in  Traihoriellabut central in  Olivinirmus; rugose nodi are typically present in  Olivinirmusbut absent in  Traihoriella.   Corvonirmusis similar to  Traihoriellain the preantennal area, but there are few similarities elsewhere. In most  Corvonirmus( Fig. 321) the antennae are sexually dimorphic, but this is never the case in  Traihoriella( Figs 265, 272). Both ps, aps, psps, and ssoccur in more anterior segments in  Corvonirmusthan in  Traihoriella( Table 2), and  tpsare present in male  Corvonirmus( Fig. 319) but absent in male  Traihoriella( Figs 263, 270); in addition, both sexes of  Corvonirmushave multiple stson at least segments IV–VI, whereas no segments have more than one stsin  Traihoriella. Pleurites continue to ventral side of abdomen, and are associated with broad pleural incrassations in  Traihoriella( Figs 263–264, 270–271) but do not or only barely reach lateral margins, but not the ventral side, and are much reduced, generally without incrassations, in  Corvonirmus( Figs 319–320). The female subgenital plate of  Corvonirmusflares into a cross-piece or have lateral submarginal extensions ( Fig. 326), but this is not the case in  Traihoriella( Figs 269, 276). The male genitalia of both genera ( Figs 266–268, 273–275, 323–325) have medianly folded parameral heads, pst1–2as sensilla, and relatively simple mesosomes. Parameral blades in  Corvonirmus( Fig. 325) are much elongated, and connected to parameral heads by a narrow neck, whereas in  Traihoriella( Figs 268, 275) the parameral blades do not narrow near the parameral heads, and are not as elongated.  Description. Both sexes. Head trapezoidal (  Traihoriella punjabensisspecies-group, Fig. 265) or indenteddome shaped (  Tr. laticepsspecies-group, Fig. 272). Marginal carina uninterrupted, deeply displaced dorsally and posteriorly at osculum. Dorsal preantennal suture, dorsal anterior plate, and ventral anterior plate absent. Ventral carinae diffuse anterior to pulvinus, and not clearly continuous with marginal carina. Head setae as in Figs 265, 272; as3absent; pnssensillus, often hard to see. Preantennal nodi conspicuous in Tr. laticepsspecies-group, less so in  Tr. punjabensisspecies-group. Coni short, slender. Antennae monomorphic. Temporal carinae not visible;  mts3only macrosetae. Gular plate broadly spade-shaped. Prothorax rounded rectangular ( Figs 263–264, 270–271); ppsson postero-lateral corner. Proepimera with hammer-shaped median ends. Pterothorax pentagonal; lateral margins divergent; posterior margin convergent to median point or rounded; mmswidely interrupted medianly. Meso- and metasterna not fused; mesosternum with 1 seta on postero-lateral corner on each side; metasternum with 1–2 setae on each postero-lateral corner. Metepisterna with large, blunt median ends. Leg chaetotaxy as in Fig. 25, except fI-v4, fI-p2absent, except in  Traihoriella latifrons; tI-v2and tIp1–2dorsal. Abdomen ( Figs 263–264, 270–271) oval, more rounded in  Traihoriella laticepsspecies-group than in  Tr. punjabensisspecies-group. Abdominal chaetotaxy as in Tables 2and 7. Tergopleurites and sternal plates generally weakly pigmented, rounded rectangular; tergopleurites II–IX+X in male and tergopleurites II–VIII in female narrowly to moderately divide medianly. Sternal plates rectangular, not approaching pleurites. Pleural incrassations broad, in some species reaching median to spiracle openings ( Figs 270–271). Re-entrant heads large in  Tr. punjabensisspecies-group ( Figs 263–264), but small in Tr. laticepsspecies-group ( Figs 270–271). Malesubgenital plate roughly triangular, reaching posterior margin of abdomen. Female subgenital plate pentagonal, reaching vulval margin but not flaring into cross-piece ( Figs 269, 276). Vulval margin ( Figs 269, 276) with slender vms, thorn-like vss; vossituated on subgenital plate, following lateral margins; distal vosmedian to vss. Basal apodeme ( Figs 266, 273) rounded trapezoidal, narrowing distally. Shape of proximal mesosome varies between species. Gonopore ( Figs 267, 274) terminal, prominent, either closed or only narrowly open distally. Mesosomal lobes slight, rounded or angular; 2–3 pmesmicrosetae lateral to gonopore on each side. Parameral heads ( Figs 268, 275) large, irregularly shaped. Parameral blades slender; pst1–2sensilla, near distal tip.   TABLE 7.Chaetotaxy of abdominal segments II–VIII of the three species of  Traihoriella. Trichoid setae of segment VIII are present in all species, and are not listed. Sets of setae differing from those of  Tr. punjabensisare highlighted in bold. Material examined from all species is from their respective type hosts. Abbreviations:  aps= accessory postspiracular seta; psps= principal post-spiracular seta; ps= paratergal seta; ss= sutural seta; sts= sternal seta;  tps= tergal posterior seta.  Species Sex ps apspsps tpsss sts   Tr. punjabensisM IV–VIII V–VII IV–VII – V–VIII II–VI F IV–VIII – IV–VII – VII–VIII II–VI  Tr. binhchauensisM IV–VIII – IV–VII – VI–VIIIII–VI F IV–VIII – IV–VII – VI–VIIIII–VI  Tr. laticepsM IV–VIII – IV–VII – II–VIIIII–VI F IV–VIII – IV–VII – II–VIIIII–VI Species-group characters:Two species groups are recognised based on abdominal chaetotaxy, head shape, and host relationships, as follows:    Traihoriella punjabensisspecies-group( Figs 263–269). Head trapezoidal; fI-v4absent; ssabsent on abdominal segments II–IV in both sexes; re-entrant heads of pleurites present. Hosts: Asian Megalaimidae.    Traihoriella latifronsspecies-group( Figs 270–276). Head indented-dome shaped; fI-v4present; sspresent on abdominal segments II–IV in both sexes; re-entrant heads of pleurites absent. Hosts: Neotropical Ramphastidae.   Host distribution.Occurs disjunctly on two families of tropical Piciformes, the South American Ramphastidaeon the genera  Aulacorhynchusand  Andigenaand the Asian Megalaimidaeon the genus  Psilopogon[given as  Megalaimain Bush et al. (2016)]. There are no known records of this genus—or any other genus in the  Brueelia-complex—from South American Barbets( Capitonidae) or African Barbets( Lybiidae).   Geographical range. Disjunctly in South Americaand South-EastAsia.   Remarks.Three species of  Traihoriellawere included in the phylogeny of Bush et al. (2016). Two other genera (  Saepocephalum  n. gen.and  Bizarrifrons Eichler, 1938) appeared within the clade containing  Traihoriellain the phylogeny of Bush et al. (2016). However, these phylogenetic relationships were not statistically supported, and they are not supported by morphological characters. All three genera have very different abdominal chaetotaxy, preantennal structure, male genitalia, and other characters. Within  Traihoriellathere is substantial variation in head shape and abdominal chaetotaxy, yet the species in this genus are strongly allied by similarities in the male genitalia and structural similarities in their preantennal areas. Amore comprehensive understanding of the relationships of these three genera will require additional material and research. Included species   Traihoriella punjabensisspecies-group    * Traihoriella binhchauensis(Najer & Sychra [in Najer et al.], 2014): 423 n. comb.[in  Brueelia] *  Traihoriella punjabensis  Ansari, 1947: 290   Traihoriella laticepsspecies-group    Traihoriella carrikeri(  Ansari, 1955a: 51)  n. comb.[in  Brueelia] [1] *  Traihoriella laticeps(  Piaget, 1888: 152)  n. comb.[in  Nirmus]  Brueelia laticeps prasinus  Carriker, 1954: 199   [1] Mey & Barker (2014)suggested that  Brueelia carrikeri Ansari, 1955b, may be a member of  Traihoriella. The illustrations that Ansari (1955b)provided indicate that the species belongs in  Traihoriella, and in the  laticepsspecies group. However, the illustration of the full-body and vulval margin are from a different host species than the holotype of  Br. carrikeri, and Ansari (1955a: 52)was himself not certain that the two samples were the same species. His illustrations of the male genitalia ( Ansari, 1955a, figs 1b–d) are very similar to those of  Tr. laticeps, but as no clear illustration of the male was provided. Therefore, we do not presently place the name  Br. carrikerias a synonym of  Tr. laticeps. The illustrated female [ex  Turdus olivater sanctaemartae(Todd, 1913)] is indistinguishable from  Tr. laticepsin abdominal and vulval chaetotaxy, head shape, and the wide pleural incrassations, but has a cross-piece at the vulval margin, which is not seen in  Traihoriella. We have not examined any material of  Br. carrikeri, and cannot confirm whether this cross-piece exists in the material Ansari studied, nor can we confirm to which genus the holotype belongs. We tentatively agree with Mey & Barker (2014)and place  Br. carrikeriin  Traihoriellabased on the original description and illustrations, as well as Ansari’s (1955a) statement that the illustrated female “closely resembles” the male. 1587279388 [199,1014,1993,2018] Disjunctly in South America 163 164 1 South-East