Halipeurus spadix subclavus Timmermann, 1961 —new status

(Figs 9, 30, 45, 64, 66–67)

Halipeurus subclavus Timmermann, 1961: 411 . Type host: Puffinus lherminieri (? persicus Hume, 1873). Holotype 3 in NHML.

Halipeurus (Halipeurus) subclavus; Timmermann, 1965: 144.

Halipeurus (Halipeurus) subclavus; Price et al., 2003: 188.

DIAGNOSIS: Male: habitus as in Fig. 66; clypeal signature as in Fig. 9; terminalia (ventral view) as in Fig. 30; genitalia as in Fig. 64. Female: habitus as in Fig. 67; clypeal signature as for male; terminalia (ventral view) as in Fig. 45.

Measurements of both sexes as in Table 2.

MATERIAL EXAMINED

Types

Ex Puffinus lherminieri (? persicus): Holotype 3, allotype Ƥ, N. Male Atoll, Maldive Is, Indian Ocean, 23 Jan. 1957, W.W.A. Phillips (NHML 1957-283).

Non-types

Ex Puffinus lherminieri bailloni (Bonaparte, 1857): 323, 24Ƥ, Réunion I., Indian Ocean, Jan. 1996, V. Bretagnolle (MONZ). New host record.

Ex Puffinus lherminieri nicolae Jouanin, 1971: 23, Cousine I., Seychelles Is, Indian Ocean, 12 Jan. 1996, V. Bretagnolle (MONZ). New host record.

Ex Puffinus gavia (Forster, 1844): 2Ƥ, New Zealand, 21 May 1934, #23 (CMNZ); 13, The Brothers Is, Cook Strait, N.Z., 12–18 Jun. 1956, G.W. Ramsay (NZAC); 1Ƥ, The Brothers Is, Cook Strait, N.Z., 29 May 1962, A. Wright, AW97 (MONZ); 33, 1Ƥ, Australia, Mar. 1968, D. Sawyer (ANIC); 23, 1Ƥ, Great I., Three Kings Is, N.Z., Nov. 1970, J.C. Watt (NZAC); 23, Motuara I., Queen Charlotte Sound, N.Z., 23 Jan. 1971, J.R. Jackson (RLCP); 23, Bay of Plenty, N.Z., May 1971, O.M.196 (RLCP); 23, Botany Bay, N.S.W., Australia, 27 Aug. 1972, B. Jones (ANIC); 13, 2Ƥ, Karewa I., Bay of Plenty, N.Z., 9 Nov. 1972, L. Moran (NZAC); 13, 3Ƥ, Ikamaru Bay, Wellington, N.Z., 27 Jul. 1975, J.R. Jackson (RLCP); 13, 1Ƥ, Opau Bay, Wellington, N.Z., 31 Aug. 1975, J.R. Jackson (MONZ); 13, 1Ƥ, Farewell Spit, N.Z., 1977, K. Owen (MONZ); 13, Petone Beach, Wellington, N.Z., 31 Jul. 1978, S. Cotter (MONZ); 33, 3Ƥ, Lyall Bay, Wellington, N.Z., 1 Aug. 1978, P. Roberts (MONZ); 33, 3Ƥ, Petone Beach, Wellington, N.Z., 31 Mar. 1979, S. Cotter (MONZ); 73, 7Ƥ, White Beach, Nubeena, S.E. Tasmania, Australia, Feb. 1980, H.D. Barker (TMTA; RLCP); 13, 2Ƥ, Blumine I., Queen Charlotte Sound, N.Z., 28 Aug. 1983, G. Wragg (MONZ); 23, 1Ƥ, Moturipa I., Anaura Bay, N.Z., 2 Sep. 1983, G. Wragg (RLCP); 143, 18Ƥ, Petone Beach, Wellington, N.Z., 5 Feb. 1988, S. Cotter (MONZ); 33, 2Ƥ, Hot Water Beach, Coromandel, N.Z., 15 Dec. 1992, R.L. Palma (MONZ). New host record.

* If parameres are asymmetrical, measurement given corresponds to longer paramere.

DISCUSSION: Timmermann’s (1961) description of Halipeurus subclavus is very brief and without illustrations, but includes measurements and a comparison with H. spadix (as H. intestatus), which he regarded as the closest species. Besides the holotype and allotype, no additional specimens of H. subclavus have been reported in the literature. My examination of the types of H. subclavus (Figs 66–67) shows that they are extremely similar to H. spadix; hence, the logical conclusion would be to demote H. subclavus to a junior synonym of H. spadix . However, considering that the dimensions of H. subclavus —including many specimens from three other hosts—are consistently smaller in both mean values and ranges than those of H. spadix from five hosts (see Table 2), I believe that regarding both taxa as subspecies of H. spadix is the best option to show that, despite being morphologically very close, they are still different and identifiable as separate entities. There are a few specimens of H. spadix spadix from Puffinus huttoni that slightly overlap in head dimensions with specimens of H. spadix subclavus from P. l h e rminieri bailloni and P. g a v i a. However, they can be clearly separated by their total length in both sexes as well as by paramere length in males (Table 2). Timmermann (1961: 412) commented that studies of larger series than those available to him might show that his three new species ( H. spadix, H. intestatus and H. subclavus) would have to be considered as conspecific.

Based on molecular data, Austin et al. (2004: 858) proposed that Puffinus lherminieri persicus and P. l h e r m i n - ieri bailloni should be regarded as subspecies of P. bailloni; the host distribution of Halipeurus spadix subclavus is congruent with that arrangement. Austin et al. (2004: 858) also concluded that Puffinus lherminieri nicolae should be synonymised with Puffinus lherminieri dichrous (as Puffinus bailloni dichrous). However, this latter synonymy does not agree with the distribution of Halipeurus lice on those hosts because a very different species, H. forficulatus, is the regular Halipeurus parasitising P. lherminieri dichrous in the Pacific Ocean (28 males and 28 females examined—including the holotype, the allotype and two paratypes of H. forficulatus —from nine hosts collected in eight localities). If the synonymy between P. l. nicolae and P. l. dichrous is correct, the Indian Ocean population of P. l. dichrous (known as P. l. nicolae) must have had its ancestral population of H. forficulatus replaced by H. spadix subclavus after a host switch from one of the other Indian Ocean subspecies of P. lherminieri . On the other hand, the presence of H. spadix subclavus on Indian Ocean birds, regarded as P. bailloni dichrous by Austin et al. (2004: 858), may in fact indicate that P. l. nicolae should not be synonymised with P. l. dichrous .

Two other subspecies of Puffinus lherminieri have been described from the Indian Ocean: P. l. temptator Louette & Herremans, 1985 from the Comoro Islands, and P. l. colstoni Shirihai & Christie, 1996 from Aldabra Island (Dickinson 2003: 76). However, no lice have been recorded from those shearwaters yet.