Maculinirmus Złotorzycka, 1964

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 : 125-127

Maculinirmus Złotorzycka, 1964

Nirmus Nitzsch, 1818: 291 (in partim).

Degeeriella Neumann, 1906: 60 (in partim). Brueelia Kéler, 1936a: 257 (in partim).

Maculinirmus Złotorzycka, 1964a: 247 .

Type species. Nirmus mundus Nitzsch [in Giebel], 1866: 366, by original designation.

Diagnosis. Maculinirmus is most similar to Turdinirmus ; for a comparison with this genus, see the diagnosis of Turdinirmus . Maculinirmus also resembles Titanomessor n. gen., and, superficially, Indoceoplanetes n. gen.

Apart from Turdinirmus , Maculinirmus is most similar to Titanomessor . Both genera contain pale species with roughly rectangular female subgenital plates that do not approach the vulval margin, short mts 3, slender marginal temporal carinae, and the presence of both pos and pns. Abdominal chaetotaxy ( Table 2) is largely identical between the two genera, but male Titanomessor ( Fig. 210 ) have tps on tergopleurites VII–VIII, but these are absent in Maculinirmus ( Figs 196 , 203 ). In Maculinirmus ( Figs 196–197 , 203–204 ), all sternal plates have more than one sts on each side in both sexes, but these are absent on male sternal plates II–V ( Fig. 210 ) and female sternal plates II–IV ( Fig. 211 ) in Titanomessor . In Titanomessor ( Fig. 212 ), the dorsal preantennal suture does not reach the margin of the head either submedianly or laterally, and often does not reach either dsms or ads; in Maculinirmus ( Figs 198 , 205) the suture always reaches ads, dsms, and both the lateral margin of the head and the hyaline margin. The marginal carina is uninterrupted in Titanomessor ( Fig. 212 ), but interrupted submedianly in Maculinirmus ( Figs 198 , 205). In both Maculinirmus ( Figs 198 , 205) and Titanomessor n. gen ( Fig. 212 ), mts 3 is shorter than in most other genera in the Brueelia -complex; however, in Titanomessor mts 4–5 of males and mts 4 of females are mesosetae, but in Maculinirmus these are microsetae in both sexes. There are also large differences between Titanomessor ( Figs 213–215 ) and Maculinirmus ( Figs 199–201 , 206–208) in the male genitalia. In Titanomessor the parameres are rounded and highly convergent, whereas in Maculinirmus they are broad and curved.

The pigmentation patterns of Maculinirmus are similar to those of Indoceoplanetes , and the male genitalia of both genera are structurally similar. Maculinirmus can be separated from members of the Indoceoplanetes by the following characters: dorsal preantennal suture is absent entirely in In. ( Indoceoplanetes ) ( Fig. 219 ), and in In. (Capnodella) ( Figs 226 , 233 ) the suture is present but does not reach either the lateral margin of the head or median of ads, whereas in Maculinirmus ( Figs 198 , 205) the suture reaches both the lateral margin of the head and much median to the ads; pns are mesosetae in both subgenera of Indoceoplanetes ( Figs 219 , 226 , 233 ), but sensilla in Maculinirmus ( Figs 198 , 205); mts 3 are macrosetae in Indoceoplanetes ( Figs 219 , 226 , 233 ), but mesosetae in Maculinirmus ( Figs 198 , 205); tps are present on some tergopleurites of males in the In. ( Indoceoplanetes ) ( Fig. 217 ) but absent in In. (Capnodella) ( Figs 224 , 231 ) and Maculinirmus ( Figs 196 , 203 ); female tergopleurite IX+X is fused with tergopleurite XI in In. (Capnodella) ( Figs 225 , 232 ), but this is not the case in Maculinirmus ( Figs 197 , 204 ) or In. ( Indoceoplanetes ) ( Fig. 218 ); mesosomal lobes have a substantial papillate section in In. ( Indoceoplanetes ) ( Figs 220–221 ), but mesosomes have no papillate section in Maculinirmus ( Figs 199–200 , 206– 207); ames are visible as microsetae in In. ( Indoceoplanetes ) ( Figs 220–221 ), but sensilla in Maculinirmus ( Figs 199–200 , 206–207); the dorsal submedian fingers of the mesosomal lobes found in In. ( Indoceoplanetes ) ( Figs 220–221 ) are absent in Maculinirmus ( Figs 199–200 , 206–207).

Description. Both sexes. Head trapezoidal ( Figs 198 , 205). Marginal carina interrupted submedianly. Hyaline margin continuous with dorsal preantennal suture; suture reaches dsms and ads; separating dorsal anterior plate partially, in some undescribed species entirely, from main head plate. Suture reaches lateral margins of head but does not completely interrupt marginal carina. Ventral anterior plate present, roughly triangular. Ventral carinae diffuse anteriorly, not clearly continuous with marginal carina. Head setae as in Figs 198 , 205. Coni short, stout. Antennae monomorphic. Temporal carinae not visible; mts 3 longer than other mts but not macrosetae. Gular plate broadly triangular. Typically, only mandibles, head nodi and gular plate pigmented.

Prothorax rectangular to square-shaped ( Figs 196–197 , 203–204 ). ppss on postero-lateral corner. Proepimera variable. Pterothorax pentagonal; lateral margins widely divergent; posterior margin convergent to median point; mms widely interrupted medianly. Meso- and metasterna not fused; 1 seta on postero-lateral corner on each side of each plate. Metepisterna slender, median ends widened, blunt. Typically, only proepimera and metepisterna darkly pigmented. Leg chaetotaxy as in Fig. 25 , except fI-v4, fI-p2 absent.

Abdomen elongated oval ( Figs 196–197 , 203–204 ). Abdominal chaetotaxy as in Table 2. Tergopleurites rounded rectangular; tergopleurites II–IX+X in male and tergopleurites II–VIII in female moderately interrupted medianly. Sternal plates translucent and margins cannot be seen clearly, not approaching pleurites. Pleural incrassations moderate. Re-entrant heads slight. Typically, only incrassations darkly pigmented. Male subgenital plate slender, elongated, reaching distal margin of abdomen. Female subgenital plate translucent, anterior and lateral margins hard to see, not approaching vulval margin ( Figs 202 , 209). There is no cross-piece. Vulval margin ( Figs 202 , 209) with slender vms, thorn-like vss; vos follows lateral margins of subgenital plate, with distal setae approaching or median to vss.

Basal apodeme roughly rectangular ( Figs 199 , 206) Proximal mesosome of varying shape, may overlap with distal basal apodeme. Gonopore ( Figs 200 , 207) as pair of subparallel or distally convergent thickenings protruding beyond distal margin of mesosome. Mesosomal lobes of varying shape, generally rounded, wide, overlapping with parameres laterally; distal margin smooth or serrated, often with sublateral hook; 2–3 pmes sensilla visible lateral to gonopore. Distal sensillus typically separated from anterior sensilla, and sometimes with visible microseta. Parameral heads ( Figs 201 , 208) with U-shaped folds. Parameral blades large, roughly oval; pst1 sensillus, central, aperture often large; pst2 microseta, lateral near distal tip.

Host distribution. Maculinirmus is presently known from the host family Oriolidae , and we have examined material from several additional host species [ Oriolus auratus Lichtenstein, 1823 ; O. flavocinctus (King, 1826) ; O. larvatus Lichtenstein, 1823 ; O. tenuirostris Blyth, 1846 ; O. xanthonotus Horsfield, 1821 ; Specotheres vieilloti flaviventris Gould, 1850], suggesting that the genus is prevalent throughout members of the Oriolidae . In the phylogeny of Bush et al. (2016), material from Cinclocoma punctatum (Shaw, 1784) —a member of the host family Cinclosomatidae—was nested within Maculinirmus . This material is very aberrant, but recognizably as a member of Maculinirmus . The relationship between the Oriolidae and Cinclosoma is not clear ( Norman et al. 2009).

Geographical range. Africa, Eurasia, and Oceania.

Remarks. Złotorzycka’s (1964a) original description of Maculinirmus is not useful, focusing almost entirely on pigmentation patterns rather than morphological structures. Apart from photos, the only illustration is of the male genitalia, and these are unrecognizable. While pigmentation patterns are a good indicator of species differences within Maculinirmus and many other genera treated here, they are generally poor genus-level traits.

Mey & Barker (2014: 108) gave four morphological characters to separate Maculinirmus from other genera in the Brueelia -complex. Two of them—lack of sexual dimorphism in size and lack of pigmentation—are less useful than the other two, as all species examined by us (including the type species and several undescribed species) are sexually dimorphic in size (see Figs 196–197 , 203–204 ), and similar pigmentation patterns can be found also in Indoceoplanetes and Guimaraesiella.

Several species of the Oriolidae are hosts to one species of Maculinirmus and one of Guimaraesiella together, both of which are more or less translucent, and have similar preantennal structures. Hence, a study of the male genitalia, chaetotaxy, and other characters is necessary to distinguish them.

Included species

* Maculinirmus ljosalfar new species

* Maculinirmus mundus (Nitzsch [in Giebel], 1866: 366) [in Nirmus ]

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 210 – 211. Titanomessor sexloba n. gen. & n. sp. ex Laniarius erythrogaster: 210, male habitus, dorsal and ventral views. 211, female habitus, dorsal and ventral views.

FIGURES 196 – 197. Maculinirmus mundus (Nitzsch [in Giebel], 1866) ex Oriolus oriolus oriolus: 196, male habitus, dorsal and ventral views. 197, female habitus, dorsal and ventral views.

FIGURES 203 – 204. Maculinirmus ljosalfar n. sp. ex Oriolus chinensis diffusus: 203, male habitus, dorsal and ventral views. 204, female habitus, dorsal and ventral views.

FIGURES 212 – 216. Titanomessor sexloba n. gen. & n. sp. ex Laniarius erythrogaster: 212, male head, dorsal and ventral views. 213, male genitalia, dorsal view. 214, mesosome, ventral view. 215, parameres, dorsal view. 216, female subgenital plate and vulval margin, ventral view. Scale for parameres same as for male genitalia.

FIGURES 198 – 202. Maculinirmus mundus (Nitzsch [in Giebel], 1866) ex Oriolus oriolus oriolus: 198, male head, dorsal and ventral views. 199, male genitalia, dorsal view. 200, male mesosome, ventral view. 201, male paramere, dorsal view. 202, female subgenital plate and vulval margin, ventral view.

FIGURES 219 – 223. Indoceoplanetes (Indoceoplanetes) indonesiana (Eichler, 1947) n. comb. ex Coracina striata difficilis: 219, male head, dorsal and ventral views. 220, male genitalia, dorsal view. 221, male mesosome, ventral view. 222, male paramere, dorsal view. 223, female subgenital plate and vulval margin, ventral view.

FIGURES 226 – 230. Indoceoplanetes (Capnodella) laurocorythes n. gen., n. subgen. & n. sp. ex Edolisoma holopolium holopolium: 226, male head, dorsal and ventral views. 227, male genitalia, dorsal view. 228, male mesosome, ventral view. 229, male paramere, dorsal view. 230, female subgenital plate and vulval margin, ventral view.

FIGURES 233 – 237. Indoceoplanetes (Capnodella) loboccupatrix n. gen., n. subgen. & n. sp. ex Lobotos oriolinus: 233, male head, dorsal and ventral views. 234, male genitalia, dorsal view. 235, male mesosome, ventral view. 236, male paramere, dorsal view. 237, female subgenital plate and vulval margin, ventral view.

FIGURES 217 – 218. Indoceoplanetes (Indoceoplanetes) indonesiana (Eichler, 1947) n. comb. ex Coracina striata difficilis: 217, male habitus, dorsal and ventral views. 218, female habitus, dorsal and ventral views.

FIGURES 224 – 225. Indoceoplanetes (Capnodella) laurocorythes n. gen., n. subgen. & n. sp. ex Edolisoma holopolium holopolium: 224, male habitus, dorsal and ventral views. 225, female habitus, dorsal and ventral views.

FIGURES 231 – 232. Indoceoplanetes (Capnodella) loboccupatrix n. gen., n. subgen. & n. sp. ex Lobotos oriolinus: 231, male habitus, dorsal and ventral views. 232, female habitus, dorsal and ventral views.