Priceiella Gustafsson & Bush, 2017

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 : 173-175

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 ).

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 ).

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 277 – 278. Priceiella (Priceiella) sternotypica (Ansari, 1956 a) n. comb. ex Garrulax pectoralis subfuscus: 277, male habitus, dorsal and ventral views. 278, female habitus, dorsal and ventral views.

FIGURES 361 – 364. Variation of preantennal area in Guimaraesiella Eichler, 1949, dorsal views: 361, Guimaraesiella sueta Valim & Weckstein, 2011 n. comb., ex Pharomachrus pavoninus: 362, Guimaraesiella sp. ex Turdus falcklandii. 363, Guimaraesiella sp. ex Rhipidura javanica. 364, Guimaraesiella antiqua (Ansari, 1956 b) n. comb. ex Catharus guttatus nanus.

FIGURES 372 – 376. Guimaraesiella pandolura n. sp. ex Pericrocotus flammeus semiruber: 372, male head, dorsal and ventral views. 373, male genitalia, dorsal view. 374, male mesosome, ventral view. 375, male paramere, dorsal view. 376, female submarginal plate and vulval margin, ventral view.

FIGURES 302 – 306. Priceiella (Thescelovora) alliocephala n. gen., n. subgen. & n. sp. ex Platylophus galericulatus ardesiacus: 302, male head, dorsal and ventral views. 303, male genitalia, dorsal view. 304, male mesosome, ventral view. 305, male paramere, dorsal view. 306, female subgenital plate and vulval margin, ventral view.

FIGURES 279 – 284. Priceiella (Priceiella) sternotypica (Ansari, 1956 a) n. comb. ex Garrulax pectoralis subfuscus: 279, male head, dorsal and ventral views. 280, female antenna, ventral side. 281, male genitalia, dorsal view. 282, male mesosome, ventral view. 283, male paramere, dorsal view. 284, female subgenital plate and vulval margin, ventral view.

FIGURES 287 – 291. Priceiella (Camurnirmus) hwameicola n. gen., n. subgen. & n. sp. ex Garrulax taewanus: 287, male head, dorsal and ventral views. 288, male genitalia, dorsal view. 289, male mesosome, ventral view. 290, male paramere, dorsal view. 291, female subgenital plate and vulval margin, ventral view.

FIGURES 309 – 314. Priceiella (Torosinirmus) koka n. gen., n. subgen. & n. sp. ex Turdoides tenebrosa: 309, male head, dorsal and ventral views. 310, female antenna, ventral view. 311, male genitalia, dorsal view. 312, male mesosome, ventral view. 313, male paramere, dorsal view. 314, female subgenital plate and vulval margin, ventral view.

FIGURES 356 – 360. Guimaraesiella papuana (Giebel, 1870) ex Paradisaea minor: 356, male head, dorsal and ventral views. 357, male genitalia, dorsal view. 358, male mesosome, ventral view. 359, male paramere, dorsal view. 360, female subgenital plate and vulval margin, ventral view.

FIGURES 365 – 369. Guimaraesiella male genitalia, ventral views: 365, Guimaraesiella cicchinoi Valim & Weckstein, 2011 n. comb., ex Trogon collaris. 366, Guimaraesiella sp. ex Heterophasia melanoleuca. 367, Guimaraesiella sp. ex Dumetella carolinensis. 368, Guimaraesiella sp. ex Dicrurus hottentottus. 369, Guimaraesiella diaprepes (Kellogg & Chapman, 1902) n. comb. ex Ƒestiaria coccinea.

FIGURES 294 – 299. Priceiella (Camurnirmus) paulbrowni n. gen., n. subgen. & n. sp. ex Garrulax leucolophus diardi: 294, male head, dorsal and ventral views. 295, female antenna, ventral view. 296, male genitalia, dorsal view. 297, male mesosome, ventral view. 298, male paramere, dorsal view. 299, female subgenital plate and vulval margin, ventral view.

FIGURES 285 – 286. Priceiella (Camurnirmus) hwameicola n. gen., n. subgen. & n. sp. ex Garrulax taewanus: 285, male habitus, dorsal and ventral views. 286, female habitus, dorsal and ventral views.

FIGURES 292 – 293. Priceiella (Camurnirmus) paulbrowni n. gen., n. subgen. & n. sp. ex Garrulax leucolophus diardi: 292, male habitus, dorsal and ventral views. 293, female habitus, dorsal and ventral views.

FIGURES 300 – 301. Priceiella (Thescelovora) alliocephala n. gen., n. subgen. & n. sp. ex Platylophus galericulatus ardesiacus: 300, male habitus, dorsal and ventral views. 301, female habitus, dorsal and ventral views.

FIGURES 307 – 308. Priceiella (Torosinirmus) koka n. gen., n. subgen. & n. sp. ex Turdoides tenebrosa: 307, male habitus, dorsal and ventral views. 308, female habitus, dorsal and ventral views.

FIGURES 315 – 318. Pleurites IV – V of male, including lateral end of sternite, dorsal view (ventral structures shown with dotted lines): 315, Priceiella (Priceiella) sternotypica (Ansari, 1956 a) n. comb. 316, Priceiella (Camurnirmus) hwameicola n. gen., n. subgen. & n. sp. 317, Priceiella (Thescelovora) alliocephala n. gen., n. sungen. & n. sp. 318, Priceiella (Torosinirmus) koka n. gen., n. sugen. & n. sp.