Lunaceps numenii numenii ( Denny, 1842 )

Gustafsson, Daniel R. & Olsson, Urban, 2012, 3377, Zootaxa 3377, pp. 1-85 : 51-55

Lunaceps numenii numenii ( Denny, 1842)

( Fig. 21a–d; Table 1)

Nirmus numenii Denny, 1842: 53

Nirmus inaequalis Piaget, 1880: 176

Nirmus phaeopi Denny ; Waterston, 1915: 33 (partim)

Degeeriella numenii (Denny) ; Harrison, 1916: 119

Degeeriella numenii (Denny) ; Séguy, 1944: 289

Degeeriella numenii (Denny) ; Blagoveshtchensky, 1948: 279

Lunaceps numenii (Denny) ; Hopkins and Clay, 1952: 201

Lunaceps numenii (Denny) ; Timmermann, 1954a: 631

Lunaceps haematopi Timmermann, 1954a: 635 . New synonymy

Lunaceps haematopi Timmermann ; Malcomson, 1960: 190

Lunaceps numenii (Denny) ; Malcomson, 1960: 190

Lunaceps numenii (Denny) ; Touleshkov, 1964: 132

Lunaceps numenii (Denny) ; Fedorenko, 1968: 73

Lunaceps phaeopi (Denny) ; Brander, 1969: 7 [Ref: Eichler and Hackman, 1973: 92]

Lunaceps numenii numenii (Denny) ; Emerson, 1972: 94 (partim)

Lunaceps haematopi Timmermann ; Emerson, 1972: 93

Lunaceps numenii (Denny) ; Eichler and Hackman, 1973: 92

Lunaceps numenii (Denny) ; Rékási and Kiss, 1977: 104

Lunaceps numenii (Denny) ; Ledger, 1980: 169

Lunaceps haematopi Timmermann ; Ledger, 1980: 169

Lunaceps numenii (Denny) ; Rékási and Kiss, 1980: 75

Lunaceps numenii numenii (Denny) ; Mayberry et al., 2000: 60

Lunaceps numenii numenii (Denny) ; Price et al., 2003: 196 (partim)

Lunaceps haematopi Timmermann ; Price et al., 2003: 196

Lunaceps numenii numenii (Denny) ; Palma and Jensen, 2005: 56

Lunaceps numenii (Denny) (“Clade 9”); Gustafsson and Olsson (2012): 94 ( Figs. 1, 2)

Type host: Numenius arquata arquata ( Linnaeus, 1758)

Other hosts: Numenius arquata orientalis C.L. Brehm, 1831 , Haematopus ostralegus Linnaeus, 1758 , Haematopus bachmani Audubon, 1838 , Dromas ardeola Paykull, 1805

Note: The two species of Haematopus are listed by Price et al. (2003: 196, 289) as hosts for L. haematopi (= L. numenii ), one as the type host and the other taken from Emerson (1972: 93), who merely stated that H. bachmani and H. palliatus Temminck, 1820 , were likely hosts. No other record of L. haematopi being found on any of the North American Haematopus species is known to us.

Diagnosis: Hyaline margin comparatively broad ( Fig. 21a). Marginal carina interrupted or weakened by dorsal preantennal suture. Anterior to dorsal preantennal suture most clearly visible as an “island” near the AS 1 and AS 2, however there is a colourless connection between this and the medio-anterior bulge of the marginal carina, which forms a prominent extension (commonly pointed in males, rounded in females) that augments the ventral anterior plate. Dorsal preantennal suture interrupted medially, bulging to envelop the ADS. Preantennal setae quite long and thick. Lateral nodus small but distinct. Dark preantennal band narrow, and almost continuous across the head. AVS2 and AVS3 aligned, in males and most females both are aligned with ADS. MTS1 and OS slender, but long, particularly in females where they approach the size of the MTS3.

Pterothorax with rounded posterior margin.

Abdomen relatively broad ( Fig. 21b). Only in segment II is the anterior margin of the tergite indented medially. Paratergal plates broad. Antero-lateral ends of tergites III–IV broadly pointed, but without tergal heads. The type of Lunaceps inaequalis studied has perhaps more pointed antero-lateral ends of tergites than is standard in L. numenii numenii , but is otherwise similar. Sublateral indentation narrow and long in all segments. Lateral setae of segment III not thorn-like on either sex.

Female genital lobes with 7–8 (9 in Haematopus material) posterior marginal setae, 5–6 (6–7 in Haematopus material, 8 in Lunaceps numenii inaequalis ) sub-marginal setae, and 7–9 median marginal setae ( Fig. 21c). Parameres gently curved, with a ventral aperture approximately one fifth from distal tip 8 Fig. 21d). Mesomere roughly triangular but elongated posteriorly, with straight or nearly straight lateral sides which are interrupted in distal half by two small apertures, only rarely with visible setae (though these are usually visible in material from H. ostralegus ). Dorsal fingers convergent and extending anteriorly to ventral blades, with median groove broad and shallow. Ventral blades thickened medially. Lower endomere semi-circular, with posterior margin flat dorsally and concave ventrally, and connected to apodemal bridge by vaguely defined triangle.

Nymphs: Ventral carina very angular. Dorsal preantennal suture long and broad, bending posteriorly and almost reaching the base of antennae.

Discussion: Lunaceps numenii is easily distinguished by the narrow and almost complete dark preantennal band, which is only barely interrupted medially. Overall, the head is heavily sclerotized, reminiscent of L. actophilus . The ventral anterior plate is quite large, but similar to that of L. numenii phaeopi and L. kukri sp. nov., and differs somewhat between the sexes.

Most similar to L. kukri sp. nov., with which it was previously considered conspecific, but males of these two species differ in the genitalia, particularly in the shape of the mesomeres and parameres. Females identical to L. kukri sp. nov. apart from the number of genital setae.

Lunacepsnumenii numenii is also similar to L. numenii phaeopi , but they differ in size— L. numenii numenii and L. kukri sp. nov. are the only Lunaceps species measuring above 2 mm —and in the shape of the male genitalia. In general, L. numenii numenii is broader in the abdomen than L. numenii phaeopi , and the paratergal plates are narrower, with a dark border marking, which is missing in L. numenii phaeopi . The ventral sublateral setae of abdominal segment III is long and slender in both sexes, which immediately sets it apart from L. numenii phaeopi .

No significant consistent differences were found between material from the two subspecies of Numenius arquata .

The type of Lunaceps inaequalis differs only very slightly from Lunaceps numenii , in the number of submarginal setae of the female genitalia and perhaps in the shape of the antero-lateral ends of tergites. These differences are small, and do not in our opinion warrant recognition of Lunaceps inaequalis as a separate species. We therefore accept the synonymisation of Lunaceps inaequalis with Lunaceps numenii ( Hopkins and Clay, 1952; Price et al., 2003).

There is a small series of Lunaceps specimens collected from Haematopus ostralegus and Dromas albeola at the NHML, which Timmermann (1954a) described as Lunaceps haematopi . Even in the original description, Timmermann seems to have had some doubts on the validity of the species, as so few individuals were available despite the type host ( H. ostralegus ) having been frequently sampled. However, in his review of ischnoceran lice from the Charadriiformes, Timmermann (1957: 81) appears more certain about the validity of L. haematopi , as he wrote: “Obwohl keine Veranlassung besteht, ihr normales Vorkommen bei Haematopus und Dromas anzuzweifeln, dürfte es sich doch empfehlen, mit einem abschliessenden Urteil über die Art zu warten, bis sie besser bekanntgeworden ist, da L. haematopi auffallenderweise auf beiden Wirten ziemlich selten zu sein scheint.” (“Although there is no reason to doubt its natural occurence on Haematopus and Dromas , it might be advisable to wait with a final judgement about this species until it is better known, as L. haematopi seems to be quite rare on both hosts.”; our translation)

Both Dromas and Haematopus would be atypical hosts for Lunaceps , as Haematopus appears to be more closely related to avocets and stilts (e.g., Paton et al., 2003; Thomas et al., 2004b; Fain and Houde, 2007), and Dromas more closely related to gulls and terns ( Thomas et al., 2004a) or pratincoles and coursers ( Pereira and Baker, 2010). It is possible that some populations of Dromas and/or Haematopus are hosts to Lunaceps . However, we feel it is best to include this material in L. numenii numenii until additional specimens have been collected. The Lunaceps from these two host species are similar in size, and both the male genitalia and the preantennal area of the lice from these two host species are virtually identical. The lateral setae of the mesomere appear to be longer and easier to see in material from H. ostralegus than in that of N. arquata , and there are some differences in the female genital setae. However, there is some overlap in setal counts, particularly in the sub-marginal setae. There are, above all, no clear characters to separate them.

The specimens from Dromas ardeola differs slightly from that of Numenius arquata and Haematopus ostralegus in head setal characters, male and female genital characters, as well as some small differences in abdominal setal arrangement. The material from D. ardeola comes from two samples with a small total number of individuals (see below). If this collection proves to represent a natural population on Dromas ardeola , it will be necessary to study them further to establish if they deserve species recognition but, at present, we prefer to include it in L. numenii numenii until more material becomes available.

Etymology: From numeni = curlew, the new moon. The name refers to the host genus, which in turn is named after the second meaning of numenii , as their beaks can be likened to thin moon crescents.

Material examined:

Ex Numenius arquata

Nontype material: ♀ 5, ♂ 5, United Kingdom: Scotland: Hebrides: South Uist , January 1934, Meinertzhagen Collection 73-110, 237, 273-425 ( NHML) . ♀ 1, ♂ 1, Denmark: Faroes Islands: Nólsoy , 29 January 1985 ( MONZ) . ♀ 9, ♂ 11, United Kingdom: England: Sheffield Park , 15 December 1975 ( MONZ) . ♀ 1, ♂ 3, United Kingdom: Wales: Radnorshire: Llanbrochllyn , 27 April 1933, BM 1980-40 ( NHML) . ♀ 1, United Kingdom: England: Lancashire: Langton: Preston , 23 January 1923, BM1980-40 ( NHML) . ♂ 3, Hungary, November 1936, Meinertzhagen Collection 8035 ( NHML) . ♀ 1, ♂ 1, [ Collection data not given on slide], 1913 ( NHML) (as Nirmus inaequalis ) . ♀ 2, ♂ 1, United Kingdom: Cheshire: Lace , 25 February 1933, Hopkins Collection ( NHML) . ♀ 1, Slovenia: Ljubljana: Iziea ?, 23 May 1950, BM 1959-232 ( NHML) . ♀ 13, ♂ 9, United Kingdom: Scotland: Hebrides: North Uist , August 1941, Meinertzhagen Collection 14563 ( NHML) . n#4, Europe , 2 December 1968, BM 1968-213 ( NHML) . ♀ 1, United Kingdom: Scotland: Dornoch Firth , December 1934, Meinertzhagen Collection 2244 ( NHML) . ♀ 5, ♂ 3, United Kingdom: Scotland: Ross-shire , October 1935, Meinertzhagen Collection 4536 ( NHML) . ♀ 2, ♂ 2, United Kingdom: England: Boston: Lines , 2 March 1938, 222 ( NHML) . ♀ 35, ♂ 40, United Kingdom: Orkney Islands, August-September 1938, Meinertzhagen Collection 11626 (one slide in Hopkins Collection), some marked BM 1951-171 ( NHML) . ♀ 3, [ Collection data not given on slide], marked “ENTOMOLOGY, Mallophaga , Louse/Parasite of Curlew”, BM 1980-40 ( NHML) . ♂ 1, [ Collection data not given on slide], unmarked ( NHML) . ♀ 17, ♂ 8, Glos [= United Kingdom: England: Gloucestershire?], November 1936, Meinertzhagen Collection 8025 and 8027 ( NHML) . ♀ 13, ♂ 6, United Kingdom: England: Suffolk, September 1935, Meinertzhagen Collection 4003 ( NHML) . ♀ 3, ♂ 3, United Kingdom: Scotland: Hebrides: South Uist , September 1953, Meinertzhagen Collection 20230, BM 1953-658 ( NHML) . ♀ 8, ♂ 3, United Kingdom: Scotland: Hebrides: South Uist , December 1933, Meinertzhagen Collection 73 ( NHML) . ♂ 2, Glos [= United Kingdom: England: Gloucestershire?], October 1936, Meinertzhagen Collection 8027 ( NHML) . ♀ 2, ♂ 2, United Kingdom: England: Northumberland: Monk’s House Bird Observatory , 21 May 1954, BM1980-40 (one of these slides also contains a female L. falcinellus ) ( NHML) . ♂ 5, United Kingdom: Scotland: Watherspoon, L. Harrison Collection ( NHML) . ♀ 3, ♂ 3, n#1, [ Collection data not given on slide], 2688 131 ( MFN) . ♀ 2, ♂ 2, [ Collection data not given on slide], 2824 vih and vgh ( MFN) . ♀ 2, ♂ 3, [ Collection data not given on slide], 20046e, f, h, i, k ( MFN) . ♀ 4, ♂ 4, [ Collection data not given on slide], 3047a–h ( MFN) .

Ex Numenius arquata orientalis (synonym: Numenius arquata lineata )

♀ 9, ♂ 3, Arabia , March 1948, Meinertzhagen Collection 17569 ( NHML) . ♀ 2, ♂ 2, n#1, Maldives: Gan-Addu Atoll , 6 Decmber 1958, BM 1959-560 ( NHML) .

Ex Haematopus ostralegus (possibly stragglers)

Holotypes, allotypes, and paratypes of L. haematopi : ♀ 34, ♂ 17, Ireland, September 1937, Meinertzhagen Collection 10568 ( NHML) .

Ex Dromas ardeola (possibly stragglers)

♀ 6, ♂ 3, Tanzania: Zanzibar, October 1912, Meinertzhagen Collection 3591 ( NHML) . ♂ 1, East Africa ( NHML) .

Ex Numenius sp.

♀ 1, ♂ 1, [Collection data not given in slide], 2259 fi and 2250 fh ( MFN). ♀ 1, [collection data not given on slide], 2259 [as Lunaceps inaequalis ] ( MFN).

Ex Limosa limosa (possibly stragglers) ♀ 1, ♂ 2, England, L. Harrison Collection ( NHML) .

Material of uncertain status due to condition of slide:

Ex Numenius arquata :

♂ 1, Asia, L. Harrison Collection ( NHML) . ♂ 1 (male genitalia only), United Kingdom: Shetlands : Ronas Voe, 23 March 1911, J. Waterston Collection BM1930-232 ( NHML) . ♀ 1, ♂ 1, [ Collection data not given on slide], 2824 w ae ( MFN) (as Lunaceps phaeopi numenii ) . ♀ 1, [ Collection data not given on slide], 716 ( MFN) (as Lunaceps phaeopii numenii ) .

Ex Circus aeruginosus (possibly straggler): ♀ 1, [Collection data not given on slide], 1758a ( MFN).