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 Priceiella Gustafsson & Bush , new genus Brueelia Kéler, 1936a : 257 ( in partim ). Allobrueelia Eichler, 1951b : 36 ( in partim ). Type species. Brueelia sternotypica Ansari, 1956a : 148 Diagnosis. The phylogenetic reconstruction of Bush et al . 2016 placed Priceiella n. gen. close to Guimaraesiella s. str., and Priceiella was nested inside Guimaraesiella s. lat. (see below). Although there is variation in the abdominal chaetotaxy between different groups of Guimaraesiella ( Table 10 ) and between the subgenera of Priceiella ( Table 8 ), there are similarities between these two genera: tps are absent in both genera, both sexes of both genera have psps on tergopleurite IV–VII, and ps are present only on segments IV–VIII in both sexes [except for females of subgenus Pr. ( Priceiella ) n. subgen. ( Fig. 278 )]. There are several differences between these two genera in non-setal characters: in Guimaraesiella the dorsal preantennal suture interrupts the marginal carina at least submedianly ( Figs 361–364 , 372 ) [except Gu. cicchinoi ( Valim & Weckstein, 2011 ) and relatives], whereas in Priceiella the dorsal preantennal suture, when present, does not interrupt the marginal carina (e.g. Pr. alliocephala n. sp. , Fig. 302 ). The female subgenital plate flares into a cross-piece at vulval margin in Priceiella ( Figs 284 , 291 , 306 , 314 ) but not in Guimaraesiella ( Figs 360 , 376 ) [except in the Gu. sehri species group; not illustrated]. The mesosome differs within Guimaraesiella s. lat. ( Figs 358 , 365–369 , 374 ) and within Priceiella ( Figs 282 , 289 , 304 , 312 ), which makes it difficult to identify distinct genitalic characters that distinguish between these two genera. A comparison between the male genitalia of Guimaraesiella s. str. (here represented by the type species, Figs 357– 359 ) and Priceiella ( Figs 282–284 , 289–291 , 304–306 , 311–313 ) reveals the following consistent differences: gonopore terminal in Guimaraesiella s. str. ( Fig. 358 ), but ventral in Priceiella ( Figs 283 , 290 , 305 , 312 ); mesosomal lobes small and not extending distal to gonopore in Guimaraesiella s. str. ( Fig. 358 ), but large and wide in Priceiella , fused distal to gonopore ( Figs 283 , 290 , 305 , 312 ); distal margin without thickening or ornamentation found in Guimaraesiella s. str. ( Fig. 358 ; thickening present in other groups of Guimaraesiella s. lat.), but with thickening on at least part of the distal margin in Priceiella ( Figs 283 , 290 , 305 , 312 ); parameral heads with simple, small median folds in Guimaraesiella s. str. ( Fig. 359 ), but large, complex median folds in Priceiella ( Figs 284 , 291 , 306 , 313 ). Antennae may be sexually dimorphic in both genera (see e.g. Gu . menuraelyrae Coinde, 1859 ), but this is rare in Guimaraesiella s. str., and not found in Pr. ( Thescelovora ) n. subgen. ( Fig. 302 ) or many Pr. ( Camurnirmus ) n. subgen. ( Fig. 287 ). TABLE 8. Chaetotaxy of abdominal segments II–VIII of some Priceiella . Trichoid setae of segment VIII are present in all species, and are not listed. Sets of setae differing from those of Pr. sternotypica are highlighted in bold . Some undescribed species have slightly different patterns. 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 Pr. ( Priceiella ) sternotypica )	Sex M F	ps III–VIII III–VIII	aps III–VII –	psps IV–VIII IV–VIII	tps – –	ss II–VIII II–VIII	sts II–VI II–VI
Pr. ( Camurnirmus ) hwameicola	M F	IV–VIII IV–VIII	IV–VIII –	IV–VIII IV–VIII	– –	II–VIII II–VIII	II–VI II–VI
Pr. ( Thescelovora ) alliocephala	M F	IV–VIII IV–VIII	– –	IV–VIII IV–VII	– –	II–VIII II–VIII	II–VI II–VI
Pr. ( Torosinirmus ) koka	M F	III–VIII III–VIII	IV–VII –	IV–VIII IV–VIII	– –	II–VIII II–VIII	II–VI II–VI

Description. Both sexes . Head shape variable ( Figs 279 , 287 , 294 , 302 , 309 ). Frons often concave. Marginal carina uninterrupted, deeply displaced posteriorly and dorsally at osculum; displaced section filled by hyaline margin. Dorsal preantennal suture absent, or in a few species present as a small dot around dsms ( Fig. 287 ), rarely reaching farther posterior, or ads ( Fig. 279 ). In some Priceiella ( Thescelovora ) ( Fig. 302 ) dorsal preantennal suture reaches both dsms and ads , but does not reach margin of head, and oes not interrupt marginal carina. Ventral carinae diffuse anterior to pulvinus, and not clearly connected to marginal carina. Ventral anterior plate absent. Head setae as in Figs 279 , 287 , 302 , 309 , similar between subgenera. as3 absent. Coni small, blunt. Antennae sexually dimorphic in some species, with male scapes larger than female scapes. Temporal carinae not visible; mts 3 only macrosetae. Gular plate spade-shaped. Prothorax rectangular ( Figs 277–278 , 285–286 , 292–293 , 300–301 , 307–308 ); ppss on postero-lateral corner. Proepimera hook- or hammer-shaped medianly, curling around coxae II. Pterothorax pentagonal; lateral margins widely divergent; posterior margin converging on rounded median point; mms moderately to widely separated medianly. Meso- and metasterna not fused; 1 seta on postero-lateral corner on each side of each plate. Metepisterna blunt, widening medianly. Leg chaetotaxy as in Fig. 25 , except fI-p2 absent. Abdomen ( Figs 277–278 , 285–286 , 292–293 , 300–301 , 307–308 ) oblong in female, more oval in male. Terminal segment of male typically protruding. Abdominal chaetotaxy as in Tables 2 and 8 , differing between subgenera. Tergopleurites rectangular in both sexes, but more posterior tergopleurites of male triangular; tergopleurites II–IX+X in male and tergopleurites II–VIII in female moderately divided medianly. Tergopleurites typically with pre-spiracular ridges ( psr in Fig. 285 ), but these are less conspicuous in Priceiella ( Priceiella ) ( Figs 277–278 ) and Pr. ( Torosinirmus ) n. subgen. ( Figs 307–308 ) compared to Pr. ( Camurnirmus ) ( Figs 285–286 ); ridges absent in Pr . alliocephala ( Figs 300–301 ), but are present in other members of Pr. ( Thescelovora ), including undescribed species. Sternal plates medianly continuous, but do not reach pleurites. Lateral margins of sternal plates concave, in Pr. ( Priceiella ) with elongated, thickened antero-lateral corners ( Figs 277–278 ). Pleural incrassations complex, differing between subgenera ( Figs 315–318 ). Reentrant heads typically slight or absent ( Figs 315, 317 ), but pronounced in Pr. ( Torosinirmus ) ( Fig. 318 ). Male subgenital plate roughly triangular distally, but anterior end often expanded laterally ( Fig. 285 ); in Pr. ( Priceiella ) ( Fig. 277 ) with anterior corners as preceding sternal plates. Female subgenital plate triangular, reaching vulval margin, where it flares into cross-piece ( Figs 284 , 291 , 299 , 306 , 314 ). Vulval margin ( Figs 284 , 291 , 299 , 306 , 314 ) with slender vms , numerous thorn-like vss ; vos follows lateral margins of subgenital plate; distal vos situated near vss . Basal apodeme ( Figs 281 , 288 , 296 , 303 , 311 ) generally rounded, more or less constricted at mid-length, in some species with anterior end much expanded laterally. Details of male genitalia differ between subgenera (see below). Proximal mesosome overlaps with distal basal apodeme. Gonopore ( Figs 282 , 289 , 297 , 304 , 312 ) open distally, diffuse or open proximally in some Priceiella ( Thescelovora ) ( Fig. 304 ). Mesosomal lobes fused distally. Distal margin of mesosomal lobes thickened; pmes and ames variable between subgenera. Parameral heads folded medianly, rectangular or irregularly shaped. Parameral blades ( Figs 283 , 290 , 298 , 305 , 313 ) variable between subgenera; pst1–2 sensilla, central, near distal tip of parameres. Host distribution. Species of Priceiella partasitise mainly “babblers” of the families Leiotrichidae, Pellorneidae and Timaliidae , with the exception of Priceiella ( Th. ) alliocephala n. sp. which occurs on a host species that is not closely related to babblers: Platylophus galericulatus ardesiacus (Bonaparte, 1850) and traditionally considered a member of the family Corvidae (e.g. Madge & Burn 1999 ; Clements et al . 2015 ). However, this placement has been questioned (e.g. Ericson et al . 2005 ; Manegold, 2008 ), and genetic data places this species closer to shrikes ( Laniidae ) ( Jønsson et al. 2008 ). No data known to us suggests that this host species is particularly close to the “babbler” families. Geographical range. Old World Tropics, from Indonesia to Sub-Saharan Africa. Etymology. Priceiella is named in honour of the phthirapterist Roger D. Price, in recognition of his countless contributions to louse taxonomy. Gender: feminine. Included subgenera Priceiella sensu stricto Camurnirmus n. subgen. Thescelovora n. subgen. Torosinirmus n. subgen. Remarks. In the phylogeny of Bush et al . (2016; Clade B, fig. 3c), Priceiella was represented only by members of Pr . ( Thescelovora ). These were placed inside Guimaraesiella s. lat., but this relationship had low statistical support. The morphological differences between Priceiella and Guimaraesiella are substantial (see above), and we therefore propose Priceiella as a separate genus. Based solely on morphology, it seems probably that Pr . ( Camurnirmus ) and Pr . ( Torosinirmus ) are sister groups, with Pr . ( Priceiella ) the sister to both of these subgenera combined, and Pr . ( Thescelovora ) the sister of the rest of Priceiella . A phylogenetic study with more complete taxon sampling is needed to sort out the relationships within this genus. Included species * Priceiella ( Camurnirmus ) hwameicola new species * Priceiella ( Camurnirmus ) nipalensis ( Ansari, 1956a: 143 ) n. comb. [in Brueelia ] * Priceiella ( Camurnirmus ) paulbrowni new species * Priceiella ( Camurnirmus ) rhinocichlae ( Eichler, 1957: 579 ) n. comb. [in Allobrueelia ] [1] * Priceiella ( Priceiella ) longisterna ( Ansari, 1956a: 146 ) n. comb. [in Brueelia ] * Priceiella ( Priceiella ) mahrastan ( Ansari, 1956a: 164 ) n. comb. [in Brueelia ] * Priceiella ( Priceiella ) sternotransversa ( Ansari, 1956a: 147 ) n. comb. [in Brueelia ] * Priceiella ( Priceiella ) sternotypica ( Ansari, 1956a: 148 ) n. comb. [in Brueelia ] * Priceiella ( Priceiella ) ventrata ( Ansari, 1956a: 150 ) n. comb. [in Brueelia ] * Priceiella ( Thescelovora ) alliocephala new species Priceiella ( Thescelovora ) malacocincla (Najer & Sychra [in Najer et al .], 2014 ) n. comb. [in Brueelia ] [2] * Priceiella ( Torosinirmus ) brueliodes ( Ansari, 1956a: 164 ) n. comb. [in Brueelia ] * Priceiella ( Torosinirmus ) koka new species * Priceiella ( Torosinirmus ) nivea ( Ansari, 1956a: 162 ) n. comb. [in Brueelia ] [1] We tentatively place Allobrueelia rhinocichlae Eichler, 1957 , in Pr. ( Camurnirmus ) based on the close similarity of the poorly preserved holotype, with additional material from another host subspecies [ Ianthocichla mitrata major (Robinson & Kloss, 1919) ] examined by us. [2] In the original illustrations by Najer et al . (2014) , as3 is present, but appears to be located in the same spot as dsms , and may be an unintended inclusion of a dorsal seta on the ventral side. In addition, a sixth mts is illustrated, which is likely one of the pmas ( sensu Mey 1994a ).