Cryptic genera, cryptic species: phylogeny of the genus Philopteroides Mey, 2004, sensu lato, with descriptions of two new genera and one new species Author Ren, Mengjiao 0000-0002-0108-6104 Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Library of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, 105 Xingang West Road, Haizhu District, Guangzhou, 510260, Guangdong Province, China Author Tian, Chunpo 0000-0001-8681-4475 College of Life Sciences, Shaanxi Normal University, 620 West Chang’an Street, Chang’an District, Xi’an City, 710119, Shaanxi Province, China Author Grossi, Alexandra A. 0000-0002-1397-4221 Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Library of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, 105 Xingang West Road, Haizhu District, Guangzhou, 510260, Guangdong Province, China Author Zou, Fasheng 0000-0002-8913-5651 Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Library of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, 105 Xingang West Road, Haizhu District, Guangzhou, 510260, Guangdong Province, China Author Gustafsson, Daniel R. 0000-0001-8868-8310 Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Library of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, 105 Xingang West Road, Haizhu District, Guangzhou, 510260, Guangdong Province, China text Arthropod Systematics & amp; Phylogeny 2024 2024-08-06 82 585 605 journal article 10.3897/asp.82.e114351 8070F22C-9721-418D-B82F-9333A0076455 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 ). Classification of the species previously placed in Philopteroides , along with host information. Type species of each genus are denoted with an asterisk (*).
Louse taxon Type host Host family Notes
Philopteroides Mey, 2004 - - -
Philopteroides fuliginosus Valim & Palma, 2013 Rhipidura fuliginosa placabilis Bangs, 1921 Rhipiduridae -
Philopteroides gigas Najer et al., 2016 Paramythia montium De Vis, 1892 Paramythiidae 1
Philopteroides lineatus ( Giebel, 1874 ) Arachnothera longirostra (Latham, 1790) Nectariniidae -
Philopteroides macrocephalus Valim & Palma, 2013 Petroica macrocephala macrocephala (Gmelin, 1789) Petroicidae -
Philopteroides mitsusui ( Uchida, 1948 ) Myzomela rubrata dichromata Wetmore, 1919 Meliphagidae 2
Philopteroides novaezelandiae Mey, 2004 * Acanthisitta chloris chloris (Sparrman, 1787) Acanthisittidae -
Philopteroides pilgrimi Valim & Palma, 2013 Gerygone igata igata (Quoy & Gaimard, 1830) Acanthizidae 3
Philopteroides sclerotifrons ( Tandan, 1955 ) Cinnyris asiaticus asiaticus (Latham, 1790) Nectariniidae -
Philopteroides sinancorellus Najer et al., 2016 Oreocharis arfaki (Meyer, 1874) Paramythiidae 1
Philopteroides xenicus Mey, 2004 Xenicus longipes longipes (Gmelin, 1789) Acanthisittidae -
- - - -
Coronedax new genus - - -
Coronedax longiceps new species * Terpsiphone incei (Gould, 1852) Monarchidae -
Coronedax terpsiphoni (Najer & Sychra [in Najer et al.], 2012 b ) new combination Terpsiphone viridis (Müller, 1776) Monarchidae -
- - - -
Stasiasticopterus new genus - - -
Stasiasticopterus cucphuongensis ( Mey, 2004 ) new combination Pycnonotus finlaysoni eous Riley, 1940 Pycnonotidae -
Stasiasticopterus flavala (Najer & Sychra [in Najer et al.], 2012 a ) new combination * Hemixos flavala Blyth, 1845 Pycnonotidae -
Stasiasticopterus haerixos ( Gustafsson et al. 2022 b ) new combination Ixos mcclellandii (Swinhoe, 1861) Pycnonotidae -
Stasiasticopterus holosternus ( Gustafsson et al., 2022 b ) new combination Pycnonotus goiavier goiavier (Scopoli, 1786) Pycnonotidae -
Stasiasticopterus kayanobori ( Uchida, 1948 ) new combination Spizixos semitorques cinereicapillus Swinhoe, 1871 Pycnonotidae 4
Stasiasticopterus longiclypeatus ( Gustafsson et al. 2022 b ) new combination Hypsipetes everetti samarensis Rand & Rabor, 1959 Pycnonotidae -
- - - -
I ncerta sedis - - -
Philopteroides beckeri ( Mey, 2004 ) Platysteira cyanea nyansae Neumann, 1905 Platysteiridae 5
1 These two species, from mountain endemics of New Guinea, are difficult to place in the present classification. They appear to be similar to both Philopteroides and Coronedax , but the parameres are much reduced or absent, and the dorsal preantennal suture appears to reach the lateral margin of the head. We here retain them in Philopteroides , but note that as more species of this genus are described, this may need to be reevaluated. 2 This species cannot be satisfactorily placed based on the original description and illustrations of Uchida (1948) and is in need of redescription. Notably, other species known from honeyeaters all fall within Philopteroides as defined here, but as many of Uchida’s specimens appear to be contaminations or stragglers, this cannot be assumed. 3 The female vulval chaetotaxy and the shape of the male genitalia (Fig. 9 ) is similar to that of Philopteroides , and the sternal chaetotaxy of Ph. pilgrimi is similar to that of Coronedax . However, no specimens were examined, and the species has never been fully illustrated. Notably, the original illustration does not show any long apical setae of the parameres ( Valim & Palma 2013 ; fig. 33). Possibly, Ph. pilgrimi is close to the unidentified species from Eugerygone rubra that was placed as a sister to Coronedax in our phylogeny. 4 Placed in Stasiasticopterus based on host associations and the fact that specimens from the same host from the Chinese subspecies Spizixos semitorques semitorques Swinhoe, 1861 belong to this genus. However, the species cannot be identified satisfactorily based on Uchida’s description and illustration ( Gustafsson et al. 2022 b ) and is in need of redescription. 5 The placement of this species is uncertain. The male genitalia, as illustrated by Valim & Palma (2013 ; fig. 31), suggests that it may be close to Coronedax , but the sternal abdominal chaetotaxy is dissimilar to that of other Coronedax [cf. Mey (2004 ; fig. 29) and Figs 18 , 19 ]. A reexamination of the type material is necessary before this species can be placed in the present classification.
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.