Lunaceps holophaeus (Burmeister, 1838)

Gustafsson, Daniel R. & Olsson, Urban, 2012, 3377, Zootaxa 3377, pp. 1-85 : 28-31

Lunaceps holophaeus (Burmeister, 1838)

( Fig. 11a–d; Table 1)

Nirmus holophaeus Burmeister, 1838: 427

Nirmus bicolor Piaget, 1880: 175

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

Degeeriella holophaea (Burmeister) ; Harrison, 1916: 115

Degeeriella bicolor (Piaget) ; Harrison, 1916: 109

Degeeriella bicolor (Piaget) ; Séguy, 1944: 279

Degeeriella holophaea (Nitzsch) ; Séguy, 1944: 285

Quadraceps holophaea (Nitzsch) ; Ansari, 1947: 279

Degeeriella holophaea (Burmeister) ; Blagoveshtchensky, 1948: 280

Lunaceps holophaeus (Burmeister) ; Hopkins and Clay, 1952: 201

Lunaceps bicolor (Piaget) ; Hopkins and Clay, 1952: 201

Lunaceps holophaeus (Burmeister) ; Timmermann, 1954a: 625

Lunaceps holophaeus (Burmeister) ; Ansari, 1956: 49

Lunaceps holophaeus (Burmeister) ; Ansari, 1958: 98

Lunaceps holophaeus (Burmeister) ; Malcomson, 1960: 190

Lunaceps holophaeus (Burmeister) ; Malcomson, 1960: 191

Lunaceps bicolor (Piaget) ; Touleshkov, 1964: 132

Lunaceps holophaeus (Burmeister) ; Touleshkov, 1964: 132

Lunaceps holophaeus (Burmeister) ; Keirans, 1967: 113

Lunaceps holophaeus holophaeus (Burmeister) ; Bechet, 1968: 125

Lunaceps holophaeus holophaeus (Burmeister) ; Emerson, 1972: 93

Lunaceps holophaeus holophaeus (Burmeister) ; Eichler and Hackman, 1973: 92

Lunaceps holophaeus (Burmeister) ; Rékási and Kiss, 1977: 105

Lunaceps holophaeus (Burmeister) ; Ledger, 1980: 169

Lunaceps holophaeus (Burmeister) ; Rékási and Kiss, 1980: 76

Lunaceps bicolor (Piaget) ; Rékási and Kiss, 1980 b: 104

Lunaceps holophaeus (Burmeister) ; Literák, 1983: 741

Lunaceps holophaeus (Burmeister) ; Price et al., 2003: 196

Lunaceps holophaeus (Burmeister) ; Martín Mateo, 2006: 26

Lunaceps holophaeus (Burmeister) ; Dik et al., 2010: 869

Lunaceps holophaeus (Burmeister) ; Inci et al., 2010: 215

Lunaceps holophaeus (Burmeister) (“Clade 2”); Gustafsson and Olsson, 2012: 94 ( Figs. 1, 2)

Type host: Philomachus pugnax ( Linnaeus, 1758)

Other host: Vanellus vanellus ( Linnaeus, 1758) (possibly straggler)

Diagnosis: Head elongated ovoid ( Fig. 11a). Marginal carina thick and continuous around the anterior margin of the head. Dorsal preantennal suture does not reach the lateral margins of the head. In some individuals, the dorsal preantennal suture is further subdivided into a median section, and two small kidney-shaped sections anterior-lateral to the ADS. In many, but not all, individuals the dorsal preantennal suture bulges posteriorly to envelope the apertures of the ADS. Ventral anterior plate triangular. Ventral carina, ventral lateral plates and ventral anterior plate ill defined. AVS3 and AVS2 more or less aligned. PCS very small, and cannot be seen in all individuals. Coni very broad and rounded. Postantennal area square-shaped, but posterior margin of head slightly concave. MTS4-5 thorn-like.

Paratergal plates narrow and dark. Re-entrant tergal heads present in segments III–IV ( Fig. 11b), but small and transparent, and hard to see in some individuals. On more posterior segments, tergal heads are still transparent, but blunt and not re-entrant. Many females are missing the re-entrant tergal heads entirely.

Female genital lobes with 4 posterior marginal setae, 3 sub-marginal setae, and 6 median marginal setae ( Fig. 11c). Notably, the median marginal setae are shorter than the sub-marginal setae. Parameres slender with small heads ( Fig. 11d). Mesomere almost triangular with straight lateral sides. Lower endomere crescent-shaped. Apodemal bridge indistinct.

Discussion: Overall, L. holophaeus is morphologically similar to L. falcinellus (which, according to molecular data, it is closely related to; Gustafsson and Olsson, 2012) and L. incoenis , particularly in the male genitalia and in preantennal characters, but its paratergal plates and tergal heads are more similar to L. actophilus , L. drosti , L. mintoni , and L. schismatus . Lunaceps holophaeus is the only Lunaceps species for which the dorsal preantennal suture does not reach the lateral edges of the head.

Lunaceps holophaeus has been reported from a variety of atypical hosts, including Limosa limosa ( Blagoveshtchensky, 1948; Golikova, 1959) [Ref: Fedorenko, 1968]), Numenius arquata ( Blagoveshtchensky, 1948; Rékási, 1993), Calidris ruficollis (Belopolskaya, 1959 [Ref: Fedorenko, 1968]), Xenus cinereus ( Blagoveshtchensky, 1948) , Calidris temminckii ( Blagoveshtchensky, 1948) , and Tringa totanus ( Rékási, 1993) , and two non-charadriiform hosts Asio flammeus ( Blagoveshtchensky, 1948) , Somateria mollissima ( Rékási, 1993) . Some of these may be stragglers or misidentifications.

Timmermann (1954a) synonymised Lunaceps bicolor , originally described from Vanellus vanellus , with L. holophaeus , based on size and shape, and under the assumption that it was a straggler on V. vanellus . The measurements of the L. bicolor and L. holophaeus specimens studied by Timmermann (1954a) match each other. However, Timmermann (1954a) does not mention any morphological characters, and thus this synonymisation may be unjustified. Eichler (1959) mentions having in his collection a single individual of L. bicolor , which supposedly has male genitalia similar to those of Abumarkub koenigi (= Neophilopterus platyclypeatus ; see Tandan and Hajela, 1962) in their general structure. Eichler’s specimen (located at the MFN) has its genitalia covered with a blotch of Canada balsam on the cover slide, but the details that can be see are typical for a Lunaceps , and unlike that of Eichler’s (1959) illustration of the male genitalia of Abumarkub koenigi . The specimen is a Lunaceps schismatus sp. nov. The only other collections of L. bicolor are cited by Touleshkov (1964) and Rékási and Kiss (1980 b).

Timmermann (1971) suggested that the lice parasitic on Vanellus vanellus are more closely related to the lice parasitic on other boreal shorebirds than to the lice from other Vanellus species. Quadraceps junceus from Vanellus vanellus appear to be more closely related to Quadraceps charadrii from Pluvialis apricaria and Quadraceps hospes from Pluvialis squatarola than to Quadraceps renschi from the Australian Vanellus miles (Gustafsson and Olsson, in prep.). No Lunaceps have been found on Vanellus vanellus examined in Sweden (n = 2)

Etymology: From holo = whole, and phae = dusky. Probably a reference to the even brown colour that characterises most members of the genus.

Material examined: Ex Philomachus pugnax (synonym: Machetes pugnax )

Neotype: ♂ 1, United Kingdom: England: Suffolk, August 1926, Meinertzhagen Collection 4921 ( NHML).

Neoallotype: ♀ 1, United Kingdom: England: Suffolk, August 1926, Meinertzhagen Collection 4921 ( NHML) .

Neoparatypes: ♀ 11, ♂ 9, United Kingdom: England: Suffolk, August, 1926, Meinertzhagen Collection 4921 ( NHML) (one of these slides contains several divided individuals, all mixed together). ♀ 7, ♂ 3, United Kingdom: Scotland: Hebrides: N. Uist, August 1941, Meinertzhagen Collection 14531 ( NHML). ♀ 4, ♂ 1, Palestine, April 1953, Meinertzhagen Collection BM 1953-225 ( NHML). ♀ 28, ♂ 25, Estonia, August 1934, Meinertzhagen Collection 1540 ( NHML) (one of these slides also contain an unidentified Quadraceps , a female L. schismatus and what may be a male L. limosella , but the individual is pale and some characters are difficult to see).

Non-types: ♀ 20, ♂ 18, Uganda: Bunyoro: Butiaba , 10 May 1935, Hopkins Collection ( NHML) . ♂ 2, United Kingdom: England: Kent: Stoke Lagoon , 22 August 1964, BM 1966-165 ( NHML) . ♂ 1, Wells, 26 August , 1892, J. Waterston Collection BM 1930-232 ( NHML) . ♀ 1, ♂ 2, n#1, India: Uttar Pradesh: Etawah , 19 September 1865 ( NHML) . ♀ 1, ♂ 1, Israel: Kfar Masaryk , 27 August 1958, 488, BM1959-56 ( NHML) . ♀ 1, ♂ 1, Israel: Kfar-Rupin , 29 March 1960, 1046/7 ( NHML) . ♂ 1, Sri Lanka [as “Ceylon”]: Leawaya: Palatopana , 27 November 1955, BM1956-133 ( NHML) . ♀ 1, ♂ 1, United Kingdom: Shetlands: Lerwick , 7 September 1957, BM 1956-724 ( NHML) . ♀ 3, ♂ 2, South Africa: Guderstepoort , 18 September 1930, G.B. Thompson Collection, BM1980-40 ( NHML) . ♀ 1, ♂ 2, Canada: Saskatchewan: Regina , 8 May 1965 (PIPeR) . ♀ 1, ♂ 1, Poland: Near Gdansk: Mikoszewo , 31 July 1963, Zuige, 42/y/18 and 42/y/12 ( MFN) . ♂ 1, [Collection data not given on slide], 20064a ( MFN). ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 7 August 2007, ID: 321, Band number: 6130114, D. Gustafsson ( SMNH) .