Halipeurus confusus Palma

Palma, Ricardo L., 2011, New taxa, new synonymies and new host records in the louse genus Halipeurus (Insecta: Phthiraptera: Philopteridae) parasitic on petrels (Aves: Procellariiformes), Zootaxa 3017, pp. 1-45 : 3-6

Halipeurus confusus Palma , new species

( Figs 1–3 , 28 , 43 , 57 )

Halipeurus (Halipeurus) accentor Edwards, 1961: 151 , figs 3R–7R (in part Halipeurus confusus ; in part H. leucophryna Timmermann, 1960 ; in part H. theresae Timmermann, 1969 ).

“ Halipeurus accentor ” Nelson, 1969: 199 (not Halipeurus accentor Edwards, 1961 ).

Halipeurus theresae ; Amerson & Emerson, 1971: 4 (in part Halipeurus confusus ; in part H. theresae Timmermann, 1969 ).

Halipeurus leucophryna ; Amerson & Emerson, 1971: 5 (in part Halipeurus confusus ; in part H. leucophryna Timmermann, 1960 ).

“ Halipeurus pelagicus ” Amerson & Emerson, 1971: 5 (not Lipeurus pelagicus Denny, 1842 ).

“ Halipeurus leucophryna ” Watt, 1971: 236, 242 (not Halipeurus leucophryna Timmermann, 1960 ).

Halipeurus (Halipeurus) sp.; Pilgrim & Palma, 1982: 9, 30.

TYPE HOST: Pterodroma nigripennis (Rothschild, 1893) .

TYPE LOCALITY: South East Island, Chatham Islands, New Zealand.

HOLOTYPE: 3 in MONZ.

DIAGNOSIS: Male: habitus as in Fig. 1 ; clypeal signature as in Fig. 3 ; terminalia (ventral view) as in Fig. 28 ; genitalia as in Fig. 57 . Female: habitus as in Fig 2 ; clypeal signature as for male; terminalia (ventral view) as in Fig. 43 . Measurements of both sexes as in Table 1.

ETYMOLOGY: The species epithet confusus is from Latin, meaning confused, mingled, and is used here as an adjective in the nominative singular.

MATERIAL EXAMINED

Types

Ex Pterodroma nigripennis : Holotype 3 (MONZ, AI.023491), allotype Ƥ (MONZ, AI.023492) and 23, 2Ƥ paratypes, South East I., Chatham Is, New Zealand, 8 Jan. 1973, D.E. Crockett (MONZ). Other 274 paratypes as follows: 1Ƥ, Chatham Is, N.Z., Aug. 1900 (MONZ); 73, 9Ƥ, Herald I., Kermadec Is, N.Z., 11 Nov. 1925, R.H. Beck (MONZ; RLCP); 1Ƥ, Meyer I., Kermadec Is, N.Z., 3 Jan. 1963, C.M. Clark (BPBM); 23, N. Terraces, Raoul I., Kermadec Is, N.Z., 16 Jan. 1963, C.M. Clark (BPBM); 83, 3Ƥ, Raoul I., Kermadec Is, N.Z., 3 Mar. 1963, C.M. Clark (BPBM); 1Ƥ, Pacific Ocean, 13 Sep. 1963 (USNM); 23, 3Ƥ, Pacific Ocean, 4 Nov. 1963, P. Gould (USNM); 13, 2Ƥ, Pacific Ocean, 4–15 Jun. 1964 (USNM); 13, 2Ƥ, Pacific Ocean, 8 Oct. 1964 (USNM); 13, 43 miles S off Raoul I., Kermadec Is, N.Z., 23 Nov. 1964, F.C. Kinsky (MONZ); 13, 1Ƥ, Pacific Ocean, 25 Nov. 1964 (USNM); 2Ƥ, Pacific Ocean, 6o04’N- 154o56’W, 11 Jun. 1965, POBSP 4032-496099 (USNM); 13, 1Ƥ, Pacific Ocean, 7o18’N- 159o30’W, 12 Jun. 1965, POBSP 4024 (USNM); 13, 1Ƥ, Pacific Ocean, 15o38’N- 169o55’W, 17 Jun. 1965, POBSP 4003 (USNM); 53, 4Ƥ, at sea, 13o46’N- 172o56’W, 18 Jun. 1965, POBSP 4269-495656 (AMNH; KCEM; USNM); 1Ƥ, at sea, 9o00’N- 155o W, 5 Jul. 1965, POBSP 4210-495272 (USNM); 13, Meyer I., Kermadec Is, N.Z., 22 Dec. 1966, J.C. Watt & C.R. Veitch (NZAC); 203, 22Ƥ, Meyer I., Kermadec Is, N.Z., 31 Dec. 1966, D.E. Crockett & J.C. Watt (NZAC; MONZ; RLCP); 23, 1Ƥ, Norfolk I., Australia, 26 Nov. 1968, H.J. Disney (AMSA); 33, Norfolk I., Australia, 21 Mar. 1969, H.J. Disney (AMSA); 23, 2Ƥ, Sumner, Canterbury, N.Z., 4 Apr. 1970, J. Warham (RLCP); 63, 4Ƥ, Raoul I., Kermadec Is, N.Z., 29 Nov. 1972, J. Ireland (NZAC; RLCP); 53, 2Ƥ, Raoul I., Kermadec Is, N.Z., N.Z., 4 Dec. 1972, J. Ireland (NZAC; RLCP); 113, 11Ƥ, Raoul I., Kermadec Is, N.Z., 27 Jan. 1973, J. Ireland (NZAC; RLCP); 23, Waipu, Ruakaka, N.Z., 7 Apr. 1974, M. O’Reilley (MONZ); 13, 1Ƥ, Suva, Viti Levu I., Fiji, 4 Apr. 1975, W.N. Beckon (MONZ); 33, 3Ƥ, Kermadec Is, N.Z., 28 Jan. 1979 (MONZ); 63, 1Ƥ, Muriwai Beach, Auckland, N.Z., 10 Feb. 1979, S.M. Reed (MONZ); 23, 1Ƥ, Dargaville Beach, Northland, N.Z., 17 Feb. 1979, D.E. Crockett (MONZ); 73, 7Ƥ, Matthew I., New Caledonia, 18 Dec. 1979, R. de Naurois (MONZ); 23, 1Ƥ, South East I., Chatham Is, N.Z., 11 Feb. 1980, A.C.G. Heath (MONZ); 33, 3Ƥ, Ruatahuna, Urewera National Park, N.Z., winter 1980, C. Whiting (MONZ); 33, 3Ƥ, Macauley I., Kermadec Is, N.Z., 21 Nov. 1980, D.M. Cunningham (MONZ); 43, 4Ƥ, Raoul I., Kermadec Is, N.Z., 2 Dec. 1982, M. Frazer (MONZ); 53, 4Ƥ, Starkeys I., Chatham Is, N.Z., 8 Dec. 1982, S. Cotter (MONZ); 23, 4Ƥ, South East I., Chatham Is, N.Z., 10 Feb. 1986, M.J. Imber (MONZ); 83, 6Ƥ, Muri, Rarotonga I., Cook Is, Feb. 1986, G. McCormack (MONZ); 63, 5Ƥ, Macauley I., Kermadec Is, N.Z., 29 Nov. 1988, A.J.D. Tennyson (MONZ); 53, 5Ƥ, Curtis I., Kermadec Is, N.Z., 8 Nov. 1989, A.J.D. Tennyson (MONZ); 43, 4Ƥ, Islets in lagoon, S. New Caledonia, 1–10 Mar. 1995, V. Bretagnolle (MONZ); 13, 2Ƥ, Herald I., Kermadec Is, N.Z., no date (AMNH); 13, 1Ƥ, Kermadec Is, N.Z., no date (NHML, Thompson Collection 1980-40. These specimens are also paratypes of Halipeurus accentor Edwards, 1961 ).

Species Head width Head length Total length Paramere length * number & sex (at temples) (including hyaline margin) (including hyaline margin)

Halipeurus confusus

Holotype 3 0.37 0.74 3.78 0.41 30 3 0.356 (0.34–0.38) 0.719 (0.69–0.75) 3.670 (3.50–3.87) 0.406 (0.39–0.42) 30 Ƥ 0.391 (0.36–0.41) 0.725 (0.69–0.76) 3.913 (3.69–4.10) –

Halipeurus pricei

Holotype 3 0.32 0.66 3.07 0.30 25 3 0.309 (0.30–0.33) 0.635 (0.62–0.66) 2.960 (2.80–3.14) 0.302 (0.29–0.31) 25 Ƥ 0.359 (0.34–0.37) 0.686 (0.66–0.70) 3.561 (3.46–3.68) – * If parameres are asymmetrical, measurement given corresponds to longer paramere.

DISCUSSION: The type series of H. confusus comprises 148 males and 132 females divided into 25 samples from eight different host-breeding localities, and 18 samples from birds taken at sea or found dead onshore. This extensive series shows that H. confusus is a morphologically uniform species, closer to H. kermadecensis ( Johnston & Harrison, 1912) and H. turtur Edwards, 1961 than to all other species of Halipeurus . However, males of H. confusus differ from those two species by the genitalia having much longer and wider parameres, as well as a different configuration of the aedeagal sac ( Fig. 57 ), and by the shape and chaetotaxy of the ventral terminalia ( Fig. 28 ). In addition, the abdomen of male H. confusus is about 30% longer than that of H. turtur . Females of H. confusus can be separated from those of H. turtur by their greater total length and by fine details of the terminalia. Females of H. confusus and H. kermadecensis are very similar and can only be separated by a detailed comparison of their terminalia.

No other species of Halipeurus has suffered such an identity crisis as that of H. confusus . The confusion began with Edwards (1961: 151) when he described H. accentor based on a mixture of samples from four hosts, which he believed were two species of Pterodroma , each including two subspecies, as follows: “ Pterodroma leucoptera masafuerae ”, “ Pterodroma leucoptera hypoleuca ”, “ Pterodroma cookii nigripennis ” and “ Pterodroma cookii difilipianna ” (sic). These host taxa are now regarded, respectively, as the following four species: Pterodroma longirostris (Stejneger, 1893) , Pterodroma hypoleuca (Salvin, 1888) , Pterodroma nigripennis (Rothschild, 1893) and Pterodroma defilippiana (Giglioli & Salvadori, 1869) (see Jouanin & Mougin 1979: 76 and Dickinson 2003: 75). Two species of Halipeurus ( H. leucophryna Timmermann, 1960 and H. theresae Timmermann, 1969 ) are known from three of those petrels (see Price et al. 2003: 187, 371) and a third, H. confusus , parasitises the fourth host species. Edwards’s type series of H. accentor contains all three Halipeurus species.

I have examined the holotype male, the allotype female and one paratype male of H. accentor (all deposited in AMNH) from Pterodroma longirostris , the type host, and I have no doubt that they are conspecific with H. leucophryna . Further, I have examined a male-female pair of paratypes of H. accentor (deposited in MCZC) from Pterodroma defilippiana , which is also H. leucophryna . However, another male-female pair of paratypes of H. accentor (deposited in NHML) from Pterodroma nigripennis is clearly not H. leucophryna but H. confusus (see Material examined). Emerson (1972: 80) listed H. accentor as a junior synonym of H. leucophryna without any comment and without citing any material examined to justify his synonymy. Nevertheless, I agree with Emerson’s new synonymy, which left the Halipeurus from Pterodroma nigripennis unnamed.

Timmermann (1965: 148) realised that both H. accentor and H. leucophryna had the same type host but he was not convinced that they represented the same species, hence he listed the former species under H. leucophryna as “Syn.? H. accentor ”. Timmermann correctly pointed out the gross differences between the genitalia of those two species, as far as he could judge from comparing his H. leucophryna material against Edwards’s (1961: 140, fig. 5R) illustration of H. accentor genitalia, but he was cautious and did not confirm the synonymy because he had not compared the types. After examining the types of H. accentor , I can only conclude that the figure of the male genitalia depicted by Edwards for H. accentor (1961: 140, fig. 5R) was drawn from a paratype male from Pterodroma nigripennis , and is therefore H. confusus . The male genitalia of H. leucophryna is, as stated and illustrated by Timmermann (1965: 149, fig. 89), very slender, thus differing significantly from the genitalia of H. confusus ( Fig. 57 ).

Amerson & Emerson (1971) listed many records of lice collected by the Pacific Ocean Biological Survey Program (POBSP) carried out by the Smithsonian Institution in the Pacific Ocean from 1963 to 1969. I have been able to examine some of the Halipeurus collected by the POBSP (see Material examined) but I found them to be mostly misidentified. The problem is that not only the lice but also the petrel hosts were incorrectly identified (G.E. Watson pers. comm. 1982, J.A. Bartle pers. comm. 1982). Thus, records of H. theresae listed by Amerson & Emerson (1971: 4) under Pterodroma hypoleuca actually refer to two species: H. theresae from Pterodroma hypoleuca and H. confusus from Pterodroma nigripennis . Similarly, records of H. leucophryna listed under Pterodroma cooki cooki (sic) in Amerson & Emerson (1971: 5) are a mixture of H. leucophryna and H. confusus , but none of them were from Pterodroma cookii : the hosts involved were Pterodroma longirostris and Pterodroma nigripennis . Another incorrect host-louse record is that of H. pelagicus under Pterodroma cooki cooki (sic) in Amerson & Emerson (1971: 5): the lice are again H. confusus and the host Pterodroma nigripennis .

My examination of samples from Norfolk Island identified by Nelson (1969), those from the Kermadec Islands published by Watt (1971), and specimens collected by the POBSP reported by Amerson & Emerson (1971) (see Material examined), confirm that the entries I have listed above in the synonymy of H. confusus are correct.

Figures

FIGURES 1 – 2. Halipeurus confusus: 1, habitus of male holotype. 2, habitus of female allotype. Scale = 1 mm.

FIGURES 3 – 12. Male clypeal signature (hyaline margin omitted): 3, Halipeurus confusus. 4, Halipeurus pricei. 5, Halipeurus atlanticus. 6, Halipeurus pelagodromae. 7, Halipeurus pelagicus. 8, Halipeurus vincesmithi. 9, Halipeurus spadix subclavus. 10, Halipeurus raphanus. 11, Halipeurus fallacis. 12, Halipeurus nesofregettae.

FIGURES 28 – 33. Ventral view of male terminalia: 28, Halipeurus confusus. 29, Halipeurus pricei. 30, Halipeurus spadix subclavus. 31, Halipeurus atlanticus. 32, Halipeurus pelagodromae. 33, Halipeurus pelagicus.

FIGURES 43 – 48. Ventral view of female terminalia: 43, Halipeurus confusus. 44, Halipeurus pricei. 45, Halipeurus spadix subclavus. 46, Halipeurus atlanticus. 47, Halipeurus pelagodromae. 48, Halipeurus pelagicus.

FIGURES 56 – 60. Male genitalia, dorsal view: 56, Halipeurus atlanticus. 57, Halipeurus confusus. 58, Halipeurus vincesmithi. 59, Halipeurus nesofregettae. 60, Halipeurus raphanus.