Melibrueelia novaeseelandiae Valim & Palma, new species

Melibrueelia novaeseelandiae Valim & Palma new species

(Figs 1–3, 6 A–C, 7A,B)

Brueelia sp. nov.; Watt 1971: 235, 244. Brueelia sp.; Pilgrim & Palma 1982: 27. Brueelia sp.; Murray et al. 2001: 1263. Brueelia sp.; Palma 2010: 408.

Type host. Prosthemadera novaeseelandiae novaeseelandiae (Gmelin, 1788) —tui (Meliphagidae).

Type locality. Reefton, South Island, New Zealand.

Other hosts. Anthornis melanura melanura (Sparrman, 1786) —bellbird; Anthornis melanura obscura Falla, 1948 —Three Kings bellbird; Anthornis melanura oneho Bartle & Sagar, 1987 —Poor Knights bellbird (Meliphagidae).

Diagnosis. Melibrueelia novaeseelandiae is morphologically close to some Corvonirmus species from members of the Corvidae and Brueelia species from members of the Timaliidae. In particular, it is close to Corvonirmus argulus (Burmeister, 1838), C. leucocephalus (Nitzsch [in Giebel], 1866), C. quadrangularis (Rudow, 1869) and C. afzali (Ansari, 1957a) from several Corvus species, by having the same general body chaetotaxy, swollen antennal scapus in males, and similar pattern of male genitalia (Ansari 1957a). However, they can be distinguished by the generic characters discussed above (see Table 1). Also, M. novaeseelandiae is close to four Brueelia species from Timaliidae belonging to the pengya -group (sensu Ansari 1956b: 135), here regarded as the “' longisternus -subgroup” (i.e. Brueelia longisternus Ansari, 1956b, B. sternotransversa Ansari, 1956b, B. sternotypicus Ansari, 1956b, and B. ventratum Ansari, 1956b) by having similar: chaetotaxy in both sexes, mesosomal sclerite in the male genitalia, width of the "cross piece" (sensu Ansari 1956b), and reticulated pattern in the female subgenital plate (see Table 1). However, both sexes of M. novaeseelandiae can be distinguished from species of the longisternus -subgroup by having: a complete medial interruption of the marginal carina; a slight interruption of the lateral side of marginal carina (entire in the longisternus -subgroup); no pleural setae on segment III (present in the longisternus -subgroup); a well-developed anterior dorsal head plate (absent in the pengya - group); and sternites III–VI without lateral incrassations (“well developed antero-lateral regions” of Ansari, 1956b; present in the longisternus -subgroup). Furthermore, males of M. novaeseelandiae differ from those of the longisternus -subgroup by having the scapus and the last antennal flagellomeres distinctly swollen, while the scapus is only slightly swollen in the longisternus -subgroup. Also, females of M. novaeseelandiae differ from those of the longisternus -subgroup by having tergal plate XI fused with IX+X (see Table 1).

Description. Male . Habitus as in Figs 1 A, 3A. Body pigmentation light-brown, except for the head marginal carina and the antennal pedicel which are slightly darker (Fig. 3 A). Flagellomeres colourless dorsally.

Head as in Figs 1 A, 2A, 3A. Preantennal region triangular, occipital and preantennal margins almost straight, and marginal temporal margins slightly convex. Small and almost straight hyaline margin between tips of the premarginal carina each side (Fig. 2 A); anterior dorsal head plate with posterior margin straight, lateral margins convex, deeply concave anteriorly, and completely surrounded by the dorsal preantennal suture (Fig. 2 A). Preantennal margin with internal incrassation almost straight (Fig. 2 A). Frontoclypeal suture indistinct, its nodal area irregular in shape and well sclerotized. Gular plate roughly pentagonal, uniformly pigmented. Temples forming an almost square angle at mts3 level; marginal temporal carina darker pigmented and thin, with its inner margin straight and narrowing from mts3 level to prothorax level (Fig. 2 A).

Thorax: As in Figs 1 A. Pterothorax with 6 marginal setae on each side (rarely 7 in one or both sides); pterothoracic apodeme well developed and with internal end narrow, reaching the lateral margins of the segment. Meso- and metasternal plates not fused, the former slightly longer than wide, and the latter square-shaped and slightly more pigmented, both plates bearing a pair of long setae each.

Abdomen: As in Fig. 1 A. Tergites uniformly pigmented, except for a narrow area around spiracles (Fig. 3 A). Tergal chaetotaxy: postspiracular long on V–VII; accessory setae long on III–VII; and one medium long sutural seta on II–VII. Porotaxy: present on tergites II–V and sternites II–VI. Tergite VIII: trichoid lateral setae thin and medium long, and two setae sub-equal in length to trichoid setae. Tergite IX+X medially divided, with 2 medium long and 2–3 short setae. Paratergal chaetotaxy: II–III 0; IV–V 2; VI–VIII 3. Sternal plates II–VI ribbon-shaped and well pigmented, each with one pair of long setae. Subgenital plate sub-triangular, wide anteriorly, tapering towards its distal end and uniformly pigmented (Figs 1 A, 3A).

Genitalia: As in Figs 1 C, 6A–C. Basal plate wide, with thickened and sub-parallel lateral margins; base of parameres curved and forming a well-defined head, then slightly curved and tapering distally, each bearing one subapical small seta and one sensillum at the tip; mesosomal complex reaches distally the mid-length of the parameres, with small dorsal reticulations, bearing 2 pairs of sensilla each side and a gonopore in the centre, which joins the endomeres through a long sclerotized tube (Fig. 1 C).

Female. Habitus as in Figs 1 B, 3B. Pigmentation darker than male, especially the tergopleural plates. Abdomen much longer than in male and with almost parallel sides; terminalia and tergal chaetotaxy distinct. Pterothorax with 5 marginal setae on each side (rarely 6, less often 4 in one side). Tergites II–VIII divided medially, with internal end of each half plate almost square; tergites IX–X–XI fused and uniformly pigmented (Fig. 7 A). Tergal chaetotaxy: postspiracular long on V–VII, and one medium long sutural seta on II–VII. Porotaxy: present on tergites II–V and sternites II–VII. Tergite VIII: each side with one thin trichoid lateral seta, one postero-medial seta and one sutural seta. Paratergal chaetotaxy: II–III 0; IV–V 2; VI–VIII 3. Sternal plates rectangular and well pigmented, with same chaetotaxy as in males (Figs 1 B, 3B).

Subgenital plate with strong reticulations medially, a wide vulvar margin with small medio-posterior notch, and 3–6 small setae on each side (Fig. 7 B). Gonapophysis with 6–10 medium long setae (Fig. 1 B). Vulva with 14–22 short spiniform setae and 8–12 long thin setae on its distal margin (Figs 1 B, 7B).

Measurements as in Table 2.

Etymology. The species epithet novaeseelandiae (Latin) = New Zealand, refers to both the name of the country of origin and the species name of the type host of this louse.

Abbreviations fοr characters measured are explained in Materials and methods

Characters Host: Prosthemadera novaeseelandiae Host: Anthornis melanura

♂♂ n = 30 ♀♀ n = 42 ♂♂ n = 10 ♀♀ n = 15

Range Mean ± SD Range Mean ± SD Range Mean ± SD Range Mean ± SD

0.33-0.37 0.34 ± 0.010 0.38-0.45 0.41 ± 0.014 0.32-0.36 0.33 ± 0.009 0.37-0.40 0.39 ± 0.010 POW 0.27-0.35 0.30 ± 0.017 0.31-0.35 0.33 ± 0.010 0.26-0.28 0.27 ± 0.008 0.27-0.31 0.30 ± 0.011 TW 0.36-0.41 0.38 ± 0.012 0.40-0.46 0.42 ± 0.012 0.34-0.36 0.35 ± 0.006 0.36-0.40 0.39 ± 0.010 ADHPL 0.06-0.07 0.06 ± 0.002 0.06-0.10 0.08 ± 0.006 0.06-0.07 0.06 ± 0.004 0.07-0.08 0.08 ± 0.003 ADHPW 0.08-0.09 0.09 ± 0.004 0.09-0.11 0.10 ± 0.005 0.07-0.08 0.08 ± 0.002 0.08-0.10 0.09 ± 0.005 PW 0.20-0.24 0.22 ± 0.011 0.23-0.27 0.24 ± 0.010 0.20-0.23 0.21 ± 0.009 0.21-0.24 0.22 ± 0.007 PTW 0.34-0.41 0.37 ± 0.016 0.36-0.43 0.39 ± 0.015 0.32-0.35 0.34 ± 0.008 0.34-0.37 0.35 ± 0.011 AW-v 0.45-0.55 0.49 ± 0.020 0.44-0.57 0.52 ± 0.029 0.44-0.48 0.46 ± 0.012 0.44-0.52 0.47 ± 0.023

0.24-0.29 0.26 ± 0.013 - - 0.25-0.27 0.26 ± 0.006 - -

. 0.07-0.09 0.08 ± 0.006 - - 0.07-0.08 0.08 ± 0.002 - -

1.10-1.35 1.20 ± 0.048 1.54-2.02 1.74 ± 0.089 1.07-1.23 1.14 ± 0.048 1.52-1.78 1.66 ± 0.067 ANT-Sca.L 0.08-0.09 0.08 ± 0.004 0.04-0.05 0.04 ± 0.003 0.06-0.09 0.08 ± 0.006 0.04-0.05 0.04 ± 0.003 ANT-Sca. W 0.05-0.06 0.05 ± 0.003 0.03-0.04 0.03 ± 0.002 0.05-0.06 0.05 ± 0.003 0.03-0.04 0.03 ± 0.003 ANT-Ped.L 0.04-0.06 0.05 ± 0.003 0.04-0.05 0.04 ± 0.003 0.04-0.05 0.05 ± 0.004 0.04-0.05 0.04 ± 0.003 ANT-Ped. W 0.03-0.04 0.03 ± 0.002 0.02-0.04 0.03 ± 0.003 0.03-0.04 0.03 ± 0.002 0.02-0.03 0.02 ± 0.002 ANT-FlaIL 0.03-0.05 0.04 ± 0.003 0.02-0.03 0.03 ± 0.002 0.03-0.04 0.04 ± 0.004 0.02-0.03 0.02 ± 0.002 ANT-FlaIW 0.03-0.04 0.03 ± 0.002 0.02-0.03 0.02 ± 0.002 0.03-0.04 0.03 ± 0.002 0.02-0.03 0.02 ± 0.002 ANT-FlaIIL 0.03-0.04 0.04 ± 0.004 0.02-0.03 0.03 ± 0.002 0.03-0.04 0.03 ± 0.004 0.02-0.03 0.02 ± 0.002 ANT-FlaIIW 0.04-0.04 0.04 ± 0.002 0.02-0.03 0.03 ± 0.002 0.03-0.04 0.04 ± 0.004 0.02-0.03 0.02 ± 0.002 ANT-FlaIIIL 0.02-0.04 0.03 ± 0.006 0.03-0.05 0.04 ± 0.004 0.02-0.05 0.03 ± 0.007 0.03-0.04 0.04 ± 0.003 ANT-FlaIIIW 0.03-0.04 0.04 ± 0.002 0.02-0.03 0.03 ± 0.002 0.03-0.04 0.04 ± 0.002 0.02-0.03 0.02 ± 0.001

Type material. Ex Prosthemadera novaeseelandiae novaeseelandiae: Holotype ♂, Rahu Saddle Reefton, Buller, N.Z., 21 Nov. 1977, P. Grant (MONZ AI. 032893). Paratypes: 8♂, 15♀, same data as for the holotype (MONZ AI.015776); 2♂, 2♀, same data as for the holotype (MZUSP #5984–#5985).

Additional material examined (non-types). Ex Prosthemadera novaeseelandiae novaeseelandiae: 1♀, Mahinapua, Westland, South Island, N.Z., 1 Jan. 1965, J.R. Jackson (MONZ AI.015766); 1♂, Low flat, Raoul Island, Kermadec Is, N.Z., 24 Dec. 1966, D.S.I.R., D22333 (NZAC); 3♀, Raoul Island, Kermadec Is, N.Z., 25 Dec. 1966, D.S.I.R (NZAC); 1♂, 4♀, Otira, N.Z., 29 Sep. 1970, R.L. Pilgrim (MONZ AI.015770); 1♂, 1♀, Otira, N.Z., 29 Nov. 1970, J.R. Jackson (MONZ AI.015769); 7♀, Rotorua, N.Z., 14 Aug. 1972, P.A. Oppenheim (MONZ AI. 020612); 3♂, 3♀, Ranui Cove, Auckland Island, N.Z., 5 Feb. 1973, D.S. Horning (MONZ AI.015771); 1♂, 2♀ Raoul Island, Kermadec Is, N.Z., 23 May 1973, J. Ireland (NZAC); 3♂, 3♀, Orongorongo Valley, N.Z., 18 Feb. 1976, B.M. Fitzgerald, A. I-2030 (MONZ AI.015773); 33♂, 31♀, Lower Hutt, Wellington, N.Z., 10 Jul. 1977, M. Bar-Brown (MONZ AI.015774); 1♀, Western Springs, Auckland, N.Z., 23 Oct. 1977, A. Nordhof (NZAC); 1♂, 5♀, Little Barrier Island, CL, N.Z., 1 Feb. 1979, C.R. Veitch (MONZ AI.015777); 1♂, 5♀, Karamea, Westland, N.Z., no date, T.P. Fisher (MONZ AI.015778).

Ex Anthornis melanura melanura: 3♂, 4♀, Turiwhati, Westland, N.Z., 13 Oct. 1970, J.R. Jackson (MONZ AI.015779); 2♀, 1N, Ranui Cove, Auckland Island, N.Z., 30 Jan. 1973, D.S. Horning (MONZ AI.015781); 1♂, 1♀, Whataroa River, West Coast, N.Z., 14 Jan. 1975, D.J. Greenwood (MONZ AI.015782); 1♂, 4♀, Orongorongo Valley, N.Z., 18 May 1976, B.M. Fitzgerald, A.3.1020 (MONZ AI.015783); 3♂, 9♀, Auckland Islands, N.Z., no date, E. Jennings, Otago Museum A.03.118 (MONZ AI.015788).

(MONZ AI.015784); 1♀, Tawhiti Rahi, Poor Knights Islands, N.Z., 5 Dec. 1980, R.H. Kleinpaste (MONZ AI.015785); 1♀, same locality, 6 Dec. 1980, R.H. Kleinpaste (MONZ AI.015786); 1♂, same locality, 10 Dec. 1980, R.H. Kleinpaste (MONZ AI.015787).

Remarks. No significant qualitative differences were found among all the populations of Melibrueelia novaeseelandiae (80♂, 117♀) examined from the two species (four subspecies) of regular hosts listed above. A comparison of measurements between the populations from the two host genera shows that there is a complete overlap of the ranges (Table 2). Although mean values of lice collected from Anthornis melanura are somewhat smaller than those from the type host (Table 2), we do not regard those differences as significant to justify the division of Melibrueelia novaeseelandiae into two taxa; instead, they could represent another example of Harrison's Rule (Johnson et al. 2005), considering that P. novaeseelandiae is larger and heavier (length 27–32 cm; weight 85 to 120 g) than A. melanura (length 17–20 cm; weight 25 to 32.5 g) (Marks 2001: 1173, 1191).

The family Meliphagidae comprises 175–180 species distributed over Australasia, Indonesia and other Pacific Islands (Joseph et al. 2014). Several records of “ Brueelia sp.” have been published from Australian meliphagid hosts (Murray et al. 2001: 1262) but, to the best of our knowledge, none has been described and named yet. Hence, a comparison of Melibrueelia novaeseelandiae with species which are likely to be the closest relatives is not possible at present.

We can only compare Melibrueelia novaeseelandiae with described species which share morphological similarities with it, but those similarities are more likely to be the result of convergent evolution than an indication of a close phylogenetic relationship. For example, the chaetotaxy and general morphology of M. novaeseelandiae is similar to those of some species of Brueelia parasitic on Corvidae and Timaliidae (Table 1) (Ansari 1956a, b, 1957). Also, the genitalia show a similar pattern to those of some species from Timaliidae (see genus diagnosis above). However, Gardner et al. (2010: 1093) concluded that four other avian families—not Corvidae or Timaliidae—were the closest relations to the Meliphagidae, placing them all in the superfamily Meliphagoidea. Considering that there are no Brueelia species known from those four families (Acanthizidae, Dasyornithidae, Maluridae and Pardalotidae; see Price et al. 2003, Mey & Barker 2014), again, it is not possible to even speculate about which louse could be the closest relative of M. novaeseelandiae. Future genetic studies of M. novaeseelandiae and its similar species from Corvidae and Timaliidae, in addition to samples of “ Brueelia ” from members of Meliphagoidea, may reveal their true relationships.

Although there is no molecular data of the louse populations living on Anthornis and Prosthemadera to assess their age and relationship, the New Zealand meliphagid lineage appears to have diverged from the Australian sister taxon (genus Pycnopygius Salvadori, 1880) about 12 million years ago (Joseph et al. 2014: fig. 1). Judging from the position of the node where the New Zealand genera diverged from each other and the length of the branches, Anthornis and Prosthemadera would at least be about 6 million years old (Joseph et al. 2014: fig. 1). Therefore, some morphological variation between the two Melibrueelia populations would have been expected if they had remained completely isolated from each other. However, the lack of such morphological variation would imply that there has been continuous gene flow between the Melibrueelia lice living on tuis and bellbirds, preventing their speciation. Such gene flow may have been facilitated by dispersal of lice by parasitic louse-flies which move from one host to another (phoresy) regardless of the host relationships, as Johnson et al. (2002: 245) have shown for lice of the genus Brueelia.