Three new species of the genus Philopteroides Mey, 2004 (Phthiraptera, Ischnocera, Philopteridae) from New Zealand

Abstract We describe and illustrate three new species of chewing lice in the genus Philopteroides parasitic on passerines (Order Passeriformes, families Acanthizidae, Rhipiduridae and Petroicidae) from New Zealand. They are: Philopteroides pilgrimi sp. n. from Gerygone igata igata; Philopteroides fuliginosus sp. n. from Rhipidura fuliginosa placabilis and Rhipidura fuliginosa fuliginosa; and Philopteroides macrocephalus sp. n. from Petroica macrocephala macrocephala and Petroica macrocephala dannefaerdi. The identity of Docophorus lineatus Giebel, 1874 is discussed based on its morphology and host association. We also transfer Tyranniphilopterus beckeri to the genus Philopteroides, and provide a key to identify adults of 12 of the 13 species now included in Philopteroides.


Introduction
The genus Philopteroides was erected by Mey (2004) to include seven species, three of which he described as new and four of which he transferred from other genera. Najer and Sychra (2012a, b) described two additional species, and we herewith describe three further new species from New Zealand, as well as transfer a species described in the genus Tyranniphilopterus by Mey (2004) to Philopteroides, bringing the total number of species to 13. We also give additional diagnostic characters to distinguish Philopteroides from closely related genera included in the Philopterus-complex (sensu Mey 2004).
Species of the Philopterus-complex are relatively sedentary lice belonging to the docophorid ecotype (Mey 2004) and highly adapted to live on feathers of the host's head & neck, on which they spend their entire life cycle. The hosts of the 13 species of Philopteroides belong to several families within the avian order Passeriformes, covering a wide geographical distribution over Africa, Asia and Oceania (see below).

Methods
Specimens examined from New Zealand hosts belong to the Museum of New Zealand Te Papa Tongarewa, Wellington, N.Z. (MONZ), except for some paratypes deposited in the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP). The remaining material examined is held in the Naturhistorischen Museum, Rudolstadt, Thüringen, Germany (NHMR). The lice were treated and mounted on slides following the Canada balsam technique (Palma 1978).
All measurements are in millimeters, taken from digitalized images from slidemounted specimens using the software Leica Application Suite (LAS), and identified by the following abbreviations: as3, length of the anterior setae 3; ADPL, anterior dorsal plate length (taken at middle line); ADPLL, anterior dorsal plate lateral length (taken from the base of anterior dorsal setae -ads, to lateral apices of the plate); ADPW, anterior dorsal plate width (taken at its widest point); ANW, width of anterior notch (taken between the bases of as3); AW, maximum width of the abdomen (taken at level of segment V); EWG, external width of genital chamber; GL, male genitalia length; GW, male genitalia width (taken at the basal plate); HL, head length (excluding hyaline membrane); IWG, internal width of genital chamber; POL, preantennal length (taken from the base of the conus to the bases of as3, obliquely to the head axis); POW, preantennal width (taken between bases of the coni); PTW, pterothorax width; PTL, pterothorax length; PW, prothorax width; TL, total length; TPVL, tergal plate V length; TRL, trabecula length; TRW, trabecula width; TW, temporal width.
The sternal abdominal setae are given in sequence from the left to the right side and named by the letters: "L" meaning long and flexible, and "S" meaning short and spine-like. Thus, the first letter indicates the outermost seta on the left side and the last letter the outermost on the right side. For example: LSLS-SLSL if the pattern is symmetrical, or LSLS-L-SLSL if asymmetrical.
The minute spine-like and the long trichoid setae present on each side of the pterothorax are not included in the number of pterothoracic setae. The chaetotaxy of the abdominal tergocentral setae does not include the postspiracular setae, except for tergite II where postspiracular setae, if present, can not be distinguished from the remaining setae. We regard as pleural setae those situated on the lateral sides of the tergo-pleurites, and as sternal setae those next to the sternites. Some species have ventral setae between the innermost pleural and the outermost sternal setae, which we regard as additional setae.
The nomenclature of head features and setae follows Clay (1951), as amended by Mey (1994). Scientific nomenclature, vernacular names, and the classification of hosts follow those in Dickinson (2003).
In addition to those characters mentioned by Mey (2004) in his original description of the genus, we add further diagnostic characters to support the generic position of Philopteroides within the Philopterus-complex.
Diagnosis. Member of the Philopterus-complex by presence of well-developed trabeculae. Anterior dorsal head plate with posterior median projection well developed and sclerotized, but without antero-lateral projections. Hyaline membrane and anterior head plates deeply concave forming an "osculum" (mitsusui group) 34,36); some species with wide frons (beckeri group) (Figs 9-10). Hyaline membrane deeply or slightly concave, arising from the level of the tips of the marginal carinae or above the anterior setae 3 (as3), with a conspicuous median sclerotization and without additional setae. Marginal carina not interrupted laterally, but with a conspicuous lateral suture on the dorsal surface, at the level of the posterior dorsal sub-medial setae (d.sm.s.). Conus ranging from much reduced to well developed. Marginal temporal setae 2 (m.t.s.2) and pre-ocular setae (p.o.s.) median to short. Prothoracic dorsal setae close to the middle of the segment, and to its posterior margin. Pleuro-tergal plates II-IV without postero-lateral projections ('posterior heads'), but few species with at most a slightly pronounced angle on segment II, but not on III or IV. Spine-like setae present on some of the sternites II-VI.

Philopteroides lineatus
Distribution. Unknown. The original description does not include a type locality. There are 13 subspecies of A. longirostra distributed throughout the Indo-Malayan region (Dickinson, 2003).
In his original description, Giebel (1874) clearly states "... Schläfenecke drei sehr lange über den Prothorax hinausragende" (= "... marginal temporal carina with three very long macrochaetae projecting beyond the prothorax"). This feature is present in species of Philopterus Nitzsch, 1818 sensu stricto, Clayiella Eichler, 1940, and in a few species of Mayriphilopterus Mey, 2004, but not in species of Philopteroides which have, at most, two long marginal temporal setae (mts), with the mts2 always short (Mey 2004). Hence, it would appear that D. lineatus is not a Philopteroides sensu stricto, unless Giebel (1874) included the ocular seta -which is medium to long in Philopteroides -in his three long marginal temporal setae, but the ocular seta does not project beyond the pronotum. Therefore, until material from the type host becomes available to allow for a neotype designation, it is not possible to establish with certainty the correct generic position of D. lineatus within the Philopterus-complex.
Regarding the host-louse association of Docophorus lineatus, Giebel (1874) gave the type host as "Arachnothera (Certhia) longirostris". Hopkins and Clay (1952: 285) interpreted that host species as being conspecific with "Certhia b. brachydactyla Brehm", according to the bird nomenclature of the time. However, Dickinson (2003: 714) listed Arachnothera longirostra (Latham, 1790) as a valid species in the family Nectariniidae, as well as Certhia brachydactyla Brehm, 1820 as a valid species in the family Certhiidae (Dickinson 2003: 648). In our opinion, the name of the species of the type host given by Giebel (1874) is the most important piece of information to establish the correct type host, regardless of the genus or subgenus associated with that species. Therefore, in agreement with Mey (2004: 174, footnote), we consider that the type host of D. lineatus is Arachnothera longirostra (Latham, 1790). The fact that another species of Philopteroides has a type host in the family Nectariniidae (see Philopterus sclerotifrons Tandan, 1955 below) is further evidence that (1) Arachnothera longirostra is the correct type host for D. lineatus and (2) Philopteroides may be the correct genus for D. lineatus. Furthermore, there is no species of Philopteroides recorded from any member of the Certhiidae.

beckeri species-group
The trapezoidal shape of the head is a distinctive character in the two species of this group. The preantennal region is short (POL 0.15-0.18) and broad (ANW 0.13-0.15), with a hyaline margin shallowly concave at midline. Conus very reduced. Distribution. Uganda. This species was recently described in detail and, therefore, it is not necessary to redescribed it again. We only include habitus images of the holotype male and one paratype female (Figs 1-2), not figured in the original description by Mey (2004). We also include illustrations to support our change of the original generic combination of this species, and have re-drawn only those characters (Figs 9,(13)(14)(15)(16)30) useful to distinguish it from the second species described below. In addition, the second nymphal stage is described from a single specimen (Fig. 29) mounted together with the female paratype, and a re-interpretation of the male genitalia is presented in Fig. 31.
Furthermore, the hosts of all species of Tyranniphilopterus -except for "Tyranniphilopterus" beckeri -belong to the passerine suborders Tyranni and Passeri, and are confined to the Americas, while the hosts of all species of Philopteroides belong to the passerine suborders Passeri and Acanthisitti distributed in Africa, Asia and Oceania. Hence, the geographical distribution of its host is further evidence that placing "Tyranniphilopterus" beckeri in the genus Philopteroides is correct.
Description of second nymphal stage. Head, thorax and abdomen as in Fig. 29. Head sub-conical, marginal carina entire laterally, with well developed anterior dorsal and ventral head plates. Anterior setae 3 (as3) rigid, 0.04 in length. Dorsal head plate with convex lateral margins and almost straight posterior margin, bearing rigid anterior dorsal head setae (ads), as the adults. Anterior ventral plate cordiform. Marginal temporal setae 3 (m.t.s.3) very long, other temporal setae short to minute. Each of the pair of posterior setae on pronotum located half way between the middle of the segment and its lateroposterior angle. Only four long pterothoracic setae, as in the second nymphal stage of most species of Philopteridae (Mey 1994). Abdomen mostly membranous, except for the tergo-pleural plate of second segment (first visible), and the pleural plates of segments III-V. Segments VI and VII with small patches of light sclerotization. Abdominal chaetotaxy as in Fig. 29, with two long dorsal setae plus the postspiracular seta on each side of segments III-VIII. Sternites II-III with one spine-like setae among three long setae on each side, IV-V with a pair of long setae each side, and VI-VIII with only one long setae on each side. One long pleural seta on III-VIII. Measurements (n=1): HL 0.37, ANW 0.11, POL 0.14, POW 0.27, ADPL 0.12, ADPW 0.14, TRL 0.08, TRW 0.04, TW 0.36, PW 0.21, PTW 0.29, AW 0.43, TL 1.06.
Remarks. Morphologically close to Ph. beckeri, especially by features of the head. In addition to the key characters mentioned below, the habitus of both species is distinct (compare Figs 1 and 3; 2 and 4, respectively). Both sexes of Ph. pilgrimi can be distinguished by (1) the presence of spine-like setae on sternite V (absent in Ph. beckeri); (2) female tergites IX+X with long innermost setae situated on the tergal plate ( Fig. 17)

mitsusui species-group
The approximately triangular shape of the head is the distinctive character in the ten species included in this group. The preantennal region is longer (POL 0.22-0.29) and narrower (ANW 0.10-0.12) than in the beckeri species-group, and the hyaline margin is deeply concave at midline. Conus well-developed.