Philopteroides Mey, 2004

Ren, Mengjiao, Tian, Chunpo, Grossi, Alexandra A., Zou, Fasheng & Gustafsson, Daniel R., 2024, Cryptic genera, cryptic species: phylogeny of the genus Philopteroides Mey, 2004, sensu lato, with descriptions of two new genera and one new species, Arthropod Systematics & amp; Phylogeny 82, pp. 585-605 : 585-605

Philopteroides Mey, 2004

Philopterus Nitzsch, 1818: 288 in partim.

Docophorus Nitzsch, 1818: 289 in partim.

Bitrabeculus Uchida, 1948: 317 in partim.

Philopteroides Mey, 2004: 173.

Tyranniphilopterus Mey, 2004: 178 in partim.

Type species.

Philopteroides novaezelandiae Mey, 2004: 174 , by original designation.

Diagnosis.

With the data provided herein, a redefinition of the genus Philopteroides is necessary. Essentially, most of the characters used by Mey (2004) to diagnose the genus are still valid, but here we restrict the genus to contain only those species in which the genitalia are of the same type as in the type species. Specifically: species in which the mesosome is broad, rectangular (or at least widely following distal margin of basal apodeme) (Figs 2–5 ); gonopore with distal extensions that protrude beyond the distal margin of the mesosome; parameres not densely sclerotized, somewhat lobe-like, not strongly convergent, and without apical mesoseta; in some species parameres are apparently absent (see below) (Figs 6–8 ).

In general, known species of Philopteroides all seem to have dorsal preantennal suture not reaching lateral margin of the head, and marginal carina being indented but not interrupted laterally, but this needs to be confirmed for some species. Females of Philopteroides have a large number of short, central setae (vss?) on the vulval margin, and numerous longer setae (vms?) sublaterally. The homology of these setae compared to the rest of Ischnocera are uncertain, and require further study.

Host associations.

Known from numerous host families (see Table 3 ).

Geographical range.

All known species are Australo-Papuan or Indo-Malayan.

Included species.

See Table 3 .

Remarks.

As noted by Mey (2004), Tandan (1955) illustrated the genitalia of Philopterus sclerotifrons Tandan, 1955 , without parameres (Fig. 8 ), which is unlike the type species of Philopteroides , but similar to the species described by Najer et al. (2016) (Figs 6 , 7 . We have not examined any specimens of Ph. sclerotifrons , but have seen a single male of another undescribed species from a sunbird, which is similar to Ph. sclerotifrons . In this male, the genitalia are partially obscured by gut content, but appear to be lacking clear parameres. Specimens from two species of sunbirds were nested inside Philopteroides s. str. in our phylogeny. The lack of parameres needs to be confirmed with additional samples, and the relationship between the paramere-less species and the paramere-bearing species needs further evaluation. For the present, we retain the paramere-less species in Philopteroides , but consider them atypical, and do not include characters from these species in the genus-level comparisons below.

Philopteroides pilgrimi Valim & Palma, 2013 , has male genitalia of the same type as the type species of Philopteroides (cf. Figs 2–5 , 9 ), and probably belongs to this genus. However, the female genitalia lack the central short setae (vss?; see Valim & Palma 2013: fig. 9), which is more typical of the genus Coronedax (see below). If these are very short in this species, they may have been overlooked; no specimen of Ph. pilgrimi was examined. We here retain Ph. pilgrimi in Philopteroides , but note that a reexamination of the species is necessary.

Philopteroides beckeri ( Mey, 2004) , originally placed in the genus Tyranniphilopterus is also here retained in the genus Philopteroides ; however, this placement is more tentative. Mey (2004) illustrated the male genitalia of this species without parameres, similar to e. g., Ph. sclerotifrons (cf. Fig. 8 with Mey 2004: fig. 29 d). However, when Valim & Palma (2013) reexamined the type specimens, they found that the parameres are present in this species but folded under the mesosome and attached to the basal apodeme much farther anterior than in most other species in the morpho-group (reproduced in Fig. 10 ). Overall, the male genitalia of this species resemble those of Coronedax (Figs 11 , 12 ) more than those of any other species of Philopteroides (Figs 2–8 ). However, sternal chaetotaxy, head shape, and the lack of elongated pst 2 in Ph. beckeri separate it from Coronedax . Possibly this species represents either the sister clade to Coronedax or the sister clade to Stasiasticopterus in our tree (Fig. 1 ). As this mosaic of characters complicates any assessment of the placement of this species within the Philopteroides morpho-group, we presently consider Philopteroides beckeri to be incerta sedis within Philopteroides s. lat.

The two species described by Najer et al. (2016) from New Guinean berrypeckers are difficult to place presently, not least because the two species are so different from each other. The male genitalia appear to lack or have much reduced parameres (Figs 6 , 7 ), similar to species of Philopteroides from sunbirds (Fig. 8 ), but in overall shape of the male genitalia and in the broad heads they are more similar to those of the beckeri - species group, in which they were originally placed. Presumably, as more species of the Philopteroides morpho-group are described, the relationships of these two species with the rest of the morpho-group may be clarified. No genetic data are available for either species. They are here considered to belong to Philopteroides until more is known about this group.

Figures

Figure 1. Phylogeny of the Philopterus - complex based on the mitochondrial COI and nuclear EF- 1 α genes generated in BEAST v 1.10. 4. Nodes with posterior probabilities of 1.0 are marked with asterisks (*), whereas nodes with posterior probabilities > 0.95 are marked with circlets (˚); all unmarked clades received posterior probabilities of <0.95. Louse voucher numbers are reduced compared to Table 1 for simplicity; voucher numbers are given only when disambiguation is necessary. For information on the collection locality of all specimens, see Table 1. The three genera of the Philopteroides morpho-group are named to the right, along with numbered clades discussed in the text. Within Clades II – III, specimens derived from African hosts are marked with gray circles after the name.

Figures 2–17. Comparison of the male genitalia of some species of Philopteroides Mey, 2004, Coronedax gen. n., and Stasiasticopterus gen. n. All figures redrawn from their original descriptions (Tandan 1955; Mey 2004; Valim & Palma 2013), unless otherwise noted. Illustrations have been rescaled to be roughly the same size. Illustrations are in ventral view, unless otherwise noted. No illustrations of the genitalia have ever been published for Philopteroides lineatus (Giebel, 1874), Philopteroides mitsusui (Uchida, 1948), or Stasiasticopterus kayanobori (Uchida, 1948). 2 Philopteroides novaezelandiae Mey, 2004. 3 Philopteroides xenicus Mey, 2004. 4 Philopteroides fuliginosus Valim & Palma, 2013. 5 Philopteroides macrocephalus Valim & Palma, 2013. 6 Philopteroides gigas Najer et al., 2016. 7 Philopteroides sinancorellus Najer et al., 2016. 8 Philopteroides sclerotifrons (Tandan, 1955); no scale in original. 9 Philopteroides pilgrimi Valim & Palma, 2013. 10 Philopteroides beckeri (Mey, 2004) (redrawn from Valim & Palma 2013). 11 Coronedax terpsiphoni (Najer & Sychra [in Najer et al.], 2012 a). 12 Coronedax longiceps sp. n. 13 Stasiasticopterus longiclypeatus (Gustafsson et al., 2022 b). 14 Stasiasticopterus holosternus (Gustafsson et al., 2022 b). 15 Stasiasticopterus haerixos (Gustafsson et al., 2022 b). 16 Stasiasticopterus flavala (Najer & Sychra [in Najer et al.], 2012 a. 17 Stasiasticopterus cucphuongensis (Mey, 2004) (redrawn from Gustafsson et al. 2022 b). Bold grey lines signify the generic divisions used here; narrow grey lines signify groups that are morphologically different from the type species of the respective genera, but where there is insufficient data to propose addition (e. g., subgeneric) limits. The two species here considered incerta sedis are placed in separate groups, pending further investigations. — Abbreviations used: BA = basal apodeme; GP = gonopore; MS = mesosome; lpmes = lateral posterios mesosomal setae; PM = parameres; pst 1–2 = parameral setae 1–2 (2 distal to 1).

Figures 18, 19. Coronedax longiceps sp. n. 18 male habitus, dorsal and ventral views. 19 female habitus, dorsal and ventral views. — Abbreviations used: mms = marginal mesometanotal setae; ppss = pronotal post-spiracular seta; ps = paratergal setae; psps = principal post-spiracular setae; ss = sutural setae; sts = sternal setae; tps = tergal posterior setae.