Philopteroides novaezelandiae Mey, 2004 (by original designation).

Two species-groups:

beckeri species-group: two species

mitsusui species-group: ten species

Passeriformes, suborder Acanthisitti (Acanthisittidae), and suborder Passeri (Acanthizidae, Meliphagidae, Monarchidae Nectariniidae, Petroicidae, Platysteiridae, Pycnonotidae, Rhipiduridae).

Africa (Senegal, Uganda), Asia (India, Vietnam, Taiwan), Oceania (Micronesia, New Zealand).

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.

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) (Figs 11–12, 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.

Docophorus lineatus Giebel, 1874 was included in Philopteroides by Mey (2004: 174), when he described it as a new genus. Considering that Giebel’s types of this species were lost during the Second World War, and that we have not been able to examine any material from the type host, we follow Mey (2004) in including Docophorus lineatus in Philopteroides, but we are unable to place it in any of the two species-groups which we define below. A neotype for Docophorus lineatus is urgently needed to clearly define this species.

Arachnothera longirostra (Latham, 1790) – Little spiderhunter (Nectariniidae). See note below.

Unknown. The original description does not include a type locality. There are 13 subspecies of Arachnothera longirostra distributed throughout the Indo-Malayan region (Dickinson, 2003).

Considering current louse species descriptions, Giebel’s (1874) description of Docophorus lineatus is superficial and without any illustration. However, from Giebel’s original comparison of this species with Docophorus communis (= Philopterus citrinellae (Schrank, 1776) ), and in agreement with Harrison (1916), Hopkins and Clay (1952) and Price et al. (2003), we believe, without doubt, that Docophorus lineatus belongs to the Philopterus-complex.

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 Docophorus 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 Docophorus 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 Docophorus 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 Docophorus lineatus and (2) Philopteroides may be the correct genus for Docophorus lineatus. Furthermore, there is no species of Philopteroides recorded from any member of the Certhiidae.

Platysteira cyanea nyansae Neumann, 1905 – Brown-throated wattle-eye. (Platysteiridae)

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, 13–16, 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.

All species of the genus Tyranniphilopterus Mey, 2004 have the following characters, which are lacking in “Tyranniphilopterus” beckeri: (1) hyaline margin arises at a level below the as3; (2) conspicuous antero-lateral projections on anterior dorsal head plate; (3) prothoracic dorsal setae located on the posterior-lateral angles of the segment, not on its posterior margin.; (4) pleuro-tergal plates II–III with well developed postero-lateral projections. Instead, “Tyranniphilopterus” beckeri has the features which define Philopteroides. These morphological features justify placing it in the latter genus, as Philopteroides beckeri (Mey, 2004) new combination.

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.

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.

Holotype ♂ (NHMR #4322.c) and 1♀ paratype (NHMR #4322.b), ex Platysteira cyanea nyansae; UGANDA: Paraa, Murchison Falls, National park, 22.X.1998, P. Becker col.

1 nymph II (NHMR #4322.b), mounted on the same slide as the female paratype.

Gerygone igata igata (Quoy & Gaimard, 1830) – Grey warbler (Acanthizidae)

New Zealand.

Female. Habitus as in Fig. 3 and head as in Fig. 10. Anterior setae 3 (as3) rigid, 0.04–0.05 long, anterior dorsal head setae (ads) minute, 0.02 long. Hyaline membrane with shallow concavity and thin sclerotization; anterior dorsal plate slightly concave and ventral head plate deeply concave (Fig. 32). Pterothorax with 6–8 posteromarginal setae on each side. Tergo-pleural plate II with a reduced postero-lateral projection overlapping segment III, tergo-pleural plate III without projection. Tergocentral setae: II, 5–6 (plus 2 anterior setae); III, 6; IV, 5–6; V, 5–6; VI, 4–6; VII, 5–6; VIII, 5. Tergites VII–IX+X as in Fig. 17. Abdominal sternal setae: segment II, LS-SL; III, LS-SL; IV, SS-SS (rarely LS on one side only); V, SS-SS; VI, LL-LL (one female with one S on one side only). Paratergal setae (all long): II–III, 0; IV–V, 2; VI–VIII, 3. Tergites VI–VIII with an innermost seta (included in the paratergal count), lateral to postspiracular seta. Sternites III–VI well-developed as large, rectangular plates. Vulva with 3 medium long setae each side. Subgenital plate, inner genital sclerite, subvulval sclerites, and vulvar margin as in Fig. 18, and intraspecific variation of two additional females as in Figs 18a and 18b. Measurements (n = 8): HL 0.39–0.43, ANW 0.13–0.15, POL 0.16–0.17, POW 0.28–0.31, ADPL 0.21–0.22, ADPLL 0.14, ADPW 0.13–0.16, TRL 0.09–0.10, TRW 0.03–0.04, TW 0.35–0.41, PW 0.22–0.28, PTL 0.13–0.14, PTW 0.29–0.35, TPVL 0.19–0.25, AW 0.44–0.54, EWG 0.06–0.08; IWG 0.03–0.04, TL 1.35–1.65.

Male. Similar to female, except in dimensions and morphology of terminalia (Figs 4, 19, 20). Pterothorax with 5–6 posteromarginal setae on each side. Tergocentral setae: II, 6 (plus 2 anterior setae); III, 6; IV, 6; V, 6; VI, 6; VII, 6; VIII, 4. Dorsal terminalia as in Fig. 19. Sternal setae as for female. Paratergal setae (all long): II–III, 0; IV–V, 2; VI–VIII, 3. Sternal plates well developed and entire on segments III–VI; subgenital plate with 4 long setae as in Fig. 20, the anterior pair on the plate, and the posterior on the plate margin. Genitalia: length of parameres 0.40 (Fig. 33). Measurements (n = 1): HL 0.38, ANW 0.14, POL 0.15, POW 0.28, ADPL 0.18, ADPLL 0.13, ADPW 0.14, TRL 0.09, TRW 0.03, TW 0.36, PW 0.21, PTL 0.15, PTW 0.31, TPVL 0.22, AW 0.47, GL 0.19, GW 0.08, TL 1.30.

Holotype ♀ (MONZ AI. 030137), ex Gerygone igata igata; NEW ZEALAND, no other data. Paratypes: 1♂, 2♀ (MONZ AI.017303) 1♀ (MZUSP #2885), same data as holotype; 2♀ (MONZ AI.017299), same host species, NEW ZEALAND: Orongorongo Valley, 20.V.1976, B.M. Fitzgerald col.; 1♀ (MONZ AI.017301), same host species, NEW ZEALAND: Kowhai Bush, Kaikoura, 15.XI.1976, B. Gill col.; 1♀ (MONZ AI.017302), same host species, NEW ZEALAND: Orongorongo Valley, 12.V.1977, B.M. Fitzgerald col..

This species is named in memory of the late Professor Robert L.C. Pilgrim (1921–2010), for his outstanding contribution to knowledge of ectoparasitic insects, and for his long friendship with RLP (Palma 2011).

Morphologically close to Philopteroides 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 Philopteroides pilgrimi can be distinguished by (1) the presence of spine-like setae on sternite V (absent in Philopteroides beckeri); (2) female tergites IX+X with long innermost setae situated on the tergal plate (Fig. 17) (on soft tegument in Philopteroides beckeri, Fig. 13); (3) female subgenital plate without medial setae (Fig. 18) (against three pairs of setae on each side, as in Philopteroides beckeri, Fig. 14); (4) shape of sub-vulval sclerites (compare Figs 14 and 18); (5) male subgenital plates (compare Figs 16 and 20); and (6) male genitalia (compare Figs 31 and 33).

Rhipidura fuliginosa placabilis Bangs, 1921 – New Zealand fantail (Rhipiduridae)

New Zealand.

Female. Habitus as in Fig. 5 and head as in Fig. 11. Anterior setae 3 (as3) rigid, 0.04–0.05 long; anterior dorsal head setae (ads) peg-like, 0.02 long. Hyaline membrane with deep concavity and thick sclerotization of its margin, 0.03–0.04 long; anterior margin of the anterior dorsal plate deeply concave and ventral head plate more concave than the dorsal one (Fig. 34). Pterothorax with 8 posteromarginal setae on each side. Tergo-pleural plate II with a reduced postero-lateral projection overlapping segment III, tergo-pleural plate III without any projection. Tergocentral setae: II, 12–13 (plus 2 anterior setae); III, 15–16; IV, 13–18; V, 16–19; VI, 16–17; VII, 12–14; VIII, 6–12. Tergites VII–IX+X as in Fig. 21. Sternal setae (intraspecific variation in parentheses): II, SL-SL (plus 1–2 long setae laterad to sternal plate); III, LL-SLL (LSL-SLL); IV, LLSLL-LLSLL (LLSLL-LSLLL); V, LLSLL-LLSLL (LLSLL-LSLLL); VI, LLLL-LLLL (LLLLL-LLLL). Segments IV-VII with 1–4 long additional setae situated on soft tegument, between the sternal plate and the pleural setae. Paratergal setae (all long): II–III, 0; IV–V, 3; VI, 4 (rarely 3 on one side only); VII, 3–4; VIII, 1–2. Segments VI–VIII with an innermost dorsal seta on each tergite (included in the paratergal count), besides the postspiracular seta. Sternites III–VI with well-developed, large plates, roughly rectangular. Vulva with 7–9 medium long setae each side, and 4–6 small setae on the middle of the vulvar margin. Subgenital plate, inner genital sclerite, subvulval sclerites, and vulvar margin as in Fig. 22. Measurements (n = 8): HL 0.45–0.48, ANW 0.09–0.10, POL 0.20–0.22, POW 0.31–0.32, ADPL 0.18–0.20, ADPLL 0.15–0.16, ADPW 0.13–0.14, TRL 0.09–0.10, TRW 0.03–0.04, TW 0.43–0.46, PW 0.25–0.26, PTL 0.16–0.18, PTW 0.35–0.37, TPVL 0.21–0.23, AW 0.53–0.60, EWG 0.05–0.06; IWG 0.03–0.04, TL 1.37–1.49.

Male. Similar to female, except in dimensions and morphology of terminalia (Fig. 6). Pterothorax with 7–9 posteromarginal setae on each side. Tergocentral setae: II, 11–14 (plus 2 anterior setae); III, 10–18; IV, 10–17; V, 9–19; VI, 10–18; VII, 11–13; VIII, 7–8. Tergites VII–IX+X as in Fig. 23. Paratergal setae (all long): II–III, 0; IV–V, 2–4; VI–VII, 3–4; VIII, 2. Sternal plates well developed and entire on segments III–VI; subgenital plate with 4 proximal long setae and 2 distal postero-lateral long setae (Fig. 24). Genitalia: length of parameres 0.50 (Fig. 35). Measurements (n = 8): HL 0.42–0.46, ANW 0.09–0.10, POL 0.20–0.21, POW 0.28–0.30, ADPL 0.18–0.19, ADPLL 0.14–0.15, ADPW 0.11–0.12, TRL 0.09–0.10, TRW 0.03–0.05, TW 0.40–0.42, PW 0.24–0.25, PTL 0.14–0.16, PTW 0.32–0.35, TPVL 0.20–0.21, AW 0.47–0.53, GL 0.21–0.23, GW 0.06–0.07, TL 1.12–1.24.

Holotype ♀ (MONZ AI. 030138), ex Rhipidura fuliginosa placabilis, NEW ZEALAND: Otaihanga, Paraparaumu, WN, 23.III.1996, N. Hyde col. Paratypes: 2♂ (MONZ AI.017297), 1♂, 1♀ (MZUSP #2886), same data as holotype; 2♂, 1♀ (MONZ AI.017295), same host species, NEW ZEALAND: Wallaceville Animal Research Centre, Upper Hutt, 4.IV.1974, D.M. Pearce col.; 1♂ (MONZ AI.017296), same host species, NEW ZEALAND: Little Barrier I., 1.IX.1977, C.R. Veitch col.; 1♂ (MONZ AI.017298), same host species, NEW ZEALAND: Days Bay, Wellington, 8.II.2003, E.W. Dawson col.

1♂ (MONZ AI.017290), ex Rhipidura fuliginosa fuliginosa (Sparrman, 1787), NEW ZEALAND: Jackson Bay, Westland, 6.VII.1969, W. Spiekman col.; 1♀ (MONZ AI.017291), NEW ZEALAND: Coutts I., Canterbury, 22.XII.1970, J.R. Jackson col.; 1♂, 1♀ (MONZ AI.017292), NEW ZEALAND: Nelson, 8.V.1972, B.A. Holloway col.; 2♂, 2♀ (MONZ AI.017293), NEW ZEALAND: Nelson, 22.V.1972, G. Kuschel col.; 1♀ (MONZ AI.017294), NEW ZEALAND: Franz Joseph Glacier, 1973–1974, P. Fletcher col.

The s pecies epithet is a noun in apposition derived from the species name of the host, and also refers to the dark colour of the lice.

Morphologically close to Philopteroides macrocephalus by having: anterior dorsal head setae (ads) peg-like; segment III without pleural setae; segment IV with 2 pleural setae; sternite VI without spine-like setae; setae of the male subgenital plate situated on the plate. However, both sexes of Philopteroides fuliginosus can be distinguished from those of Philopteroides macrocephalus by: (1) the hyaline margin with a shallower indentation (0.03–0.04mm) at its midline (compare Figs 11 and 12; 34 and 36); (2) the female tergal chaetotaxy and shape of tergites IX+X (Fig. 21); (3) the male subgenital plate (Fig. 24); and (4) the male genitalia (Fig. 34).

Petroica macrocephala macrocephala (Gmelin, 1789) – New Zealand tit or tomtit (Petroicidae)

New Zealand.

Female. Habitus as in Fig. 7, and head as in Fig. 12. Anterior setae 3 (as3) rigid, peg-shaped, 0.05–0.06 long; anterior dorsal head setae (ads) peg-like, 0.02 long. Hyaline membrane with deep concavity and thin sclerotization, 0.06–0.08 long; anterior dorsal plate slightly concave and ventral head plate deeply concave (Fig. 36). Pterothorax with 9 posteromarginal setae on each side. Tergo-pleural plate II with a much reduced postero-lateral projection overlapping segment III; tergo-pleural plate III without any projection. Tergocentral setae: II, 17–19 (plus 2 anterior setae); III, 23–25; IV, 25–27; V, 26–27; VI, 24–26; VII, 23–25; VIII, 12–14. Tergites VII–IX+X as in Fig. 25. Sternal setae: II, SL-LS; III, LLLLSL-LSLLLL; IV, LLLLSLL-LLSLLLL; V, LLLLLSLL-LLSLLLLL; VI, LLLLL-LLLLL (some specimens show slight variations from this symmetrical pattern). Segments II–VII with 3–5 additional long setae situated on the soft tegument, between the sternal plate and the pleural setae. Paratergal setae (all long): II–III, 0; IV, 5–6; V, 4–5; VI–VII, 4; VIII, 2. Tergites VI–VII with an innermost dorsal seta (included in the paratergal count), besides the postspiracular setae (not included in the count). Sternites III–VI with large, well-developed plates, roughly rectangular. Vulva with 9–11 medium to long setae each side, and 14–15 small setae in the middle of the vulvar margin. Subgenital plate, inner genital sclerite, subvulval sclerites, and vulvar margin as in Fig. 26, plus a variation of an additional female in Fig. 26a. Measurements (n = 7): HL 0.59–0.62, ANW 0.10–0.12, POL 0.28–0.30, POW 0.40–0.41, ADPL 0.24–0.25, ADPLL 0.20–0.21, ADPW 0.18–0.19, TRL 0.11–0.13, TRW 0.04–0.05, TW 0.58–0.64, PW 0.30–0.36, PTL 0.21–0.25, PTW 0.44–0.48, TPVL 0.25–0.27, AW 0.72–0.79, EWG 0.05–0.06, IWG 0.03–0.04, TL 1.85–2.13.

Male. Similar to female, except in dimensions and morphology of terminalia (Fig. 8). Pterothorax with 7–9 posteromarginal setae on each side. Tergocentral setae: II, 17–18 (plus 2 anterior setae); III, 17–19; IV, 22–24; V, 18–20; VI, 17–20; VII, 16–18; VIII, 10–12. Paratergal setae (all long): II–III, 0; IV–V, 4 (rarely 2 setae on one side only); VI–VII, 4–5 (rarely 3 setae on one side only); VIII, 2. Sternal plates III–VI well developed and entire; 4 long setae on the subgenital plate with only the anterior pair situated on the plate (Fig. 28). Genitalia: length of parameres 0.50 (Fig. 37). Measurements (n = 9): HL 0.55–0.57, ANW 0.10–0.12, POL 0.27–0.28, POW 0.37–0.39, ADPL 0.23–0.25, ADPLL 0.27–0.28, ADPW 0.16–0.17, TRL 0.10–0.12, TRW 0.04–0.05, TW 0.51–0.55, PW 0.28–0.30, PTL 0.17–0.20, PTW 0.42–0.44, TPVL 0.25–26, AW 0.62–0.65, GL 0.27–0.32, GW 0.07–0.08, TL 1.49–1.61.

Holotype ♀ (MONZ AI.030122), ex Petroica macrocephala macrocephala, NEW ZEALAND: Haast Pass, 30.IX.1969, C.N. Challies col. (University of Canterbury). Paratypes: 3♂, 3♀ (MONZ AI.017278 & AI.017279), 1♂, 1♀ (MZUSP #2887), same data as holotype.

2♀ (MONZ AI.017281), ex Petroica macrocephala dannefaerdi (Rothschild, 1894), NEW ZEALAND: Penguin Creek, Snares Island, 29.III.1972, D.S. Horning & C.J. Horning cols.; 1♂ (MONZ AI.017282), NEW ZEALAND: Boat Harbour, Snares Is, 22.II.1975, H.A. Best col.; 1♂ (MONZ AI.017283), NEW ZEALAND: Station Point, Snares Island, 25.II.1977, D.S. Horning col.; 1♂ (MONZ AI.017284), NEW ZEALAND: Tern Point, Snares Islands, 10.I.1987, A. Tennyson col.; 1♂ (MONZ AI.017280), NEW ZEALAND: Snares Is, no date, CMu Av. 4635. 1♂ (MONZ AI.017277), ex Petroica macrocephala, NEW ZEALAND: Springs Junction, Maruia, 20.VIII.1966, J.R. Jackson col.

The s pecies epithet is a noun in apposition derived from the species name of the host, and also refers to the larger head of this louse species.

Morphologically close to Philopteroides fuliginosus by the features mentioned under that species above. However, both sexes of Philopteroides macrocephalus can be distinguished from those of Philopteroides fuliginosus by (1) the hyaline margin with a deeper incision (0.06–0.08mm) at its midline (compare Figs 11 and 12, Figs 34 and 36); (2) the female tergal chaetotaxy and shape of tergites IX+X (compare Figs 21 and 25); (3) the male subgenital plate (compare Figs 24 and 28); and (4) the male genitalia (compare Figs 35 and 37).