Lunaceps falcinellus Timmermann, 1954

Gustafsson, Daniel R. & Olsson, Urban, 2012, 3377, Zootaxa 3377, pp. 1-85 : 24-28

Lunaceps falcinellus Timmermann, 1954

( Fig 10a–d; Table 1)

Nirmus actophilus Kellogg and Chapman ; Waterston, 1914: 283 (partim)

Lunaceps falcinellus Timmermann, 1954a: 627 (partim)

Lunaceps incoenis (Kellogg and Chapman) ; Timmermann, 1954a: 623 (partim)

Lunaceps cabanisi Timmermann, 1954a: 625 (partim)

Lunaceps husainii Ansari, 1956: 49 New synonymy

Lunaceps husainii Ansari, 1958: 98

Lunaceps holophaeus incoenis (Kellogg and Chapman) ; Bechet, 1968: 126

Lunaceps holophaeus timmermanni Bechet, 1968: 126

Lunaceps falcinellus Timmermann ; Fedorenko, 1968: 73

Lunaceps incoenis (Kellogg and Chapman) ; Fedorenko, 1968: 73 (partim)

Lunaceps sp. II Fedorenko, 1968: 73

Lunaceps holophaeus timmermanni Bechet ; Eichler and Hackman, 1973: 92 New synonymy

Lunaceps sp. “ab Limicola f. falcinellus ” Eichler and Hackman, 1973: 92

Lunaceps sp. Rékási and Kiss, 1977: 105

Lunaceps cabanisi Timmermann ; Ledger, 1980: 169

Lunaceps sp. Rékási and Kiss, 1980: 76

Lunaceps incoenis (Kellogg and Chapman) s. l.; Pilgrim and Palma, 1982: 21

Lunaceps holophaeus (Burmeister) ; Literák, 1983: 743

Lunaceps sp. [ex Calidris ruficollis ] Palma, 1999: 380

Lunaceps falcinellus Timmermann ; Price et al., 2003: 196 (partim)

Lunaceps cabanisi Timmermann ; Price et al., 2003: 196 (partim)

Lunaceps incoenis (Kellogg and Chapman) ; Price et al., 2003: 196 (partim)

Lunaceps timmermanni Bechet ; Price et al., 2003: 196

Lunaceps incoenis (Kellogg and Chapman) ; Rékási, 2006: 183

Lunaceps timmermanni Bechet ; Rékási, 2006: 183

Lunaceps timmermanni Bechet ; Adam, 2007a: 137

Lunaceps drosti Timmermann ; Dik et al., 2010: 869 (identified in images)

Lunaceps drosti Timmermann ; Inci et al., 2010: 215

Lunaceps incoenis (Kellogg and Chapman) ; Dik et al., 2010: 869

Lunaceps incoenis (Kellogg and Chapman) ; Inci et al., 2010: 215

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

Lunaceps incoenis (Kellogg and Chapman) (“Clade 1”); Gustafsson and Olsson, 2012: 94 ( Figs. 1, 2) (in partim)

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

Lunaceps sp. ex Calidris ruficollis (“Clade 1”); Gustafsson and Olsson, 2012: 94 ( Figs. 1, 2).

Type host: Limicola falcinellus falcinellus ( Pontoppidan, 1763)

Other hosts: Limicola falcinellus sibirica Dresser, 1876 , Calidris ferruginea ( Pontoppidan, 1763) , Calidris ruficollis ( Pallas, 1776) , Calidris minuta (Leisner, 1812) .

Diagnosis: Head elongated ovoid ( Fig. 10a). Posterior margin flat or nearly flat. Marginal carina clearly defined, but not very heavily pigmented, especially anterior to the dorsal preantennal suture. Ventral anterior plate triangular. No lateral nodus, and dark preantennal band very vague. Dorsal preantennal suture broad, enveloping the apertures of the AVS in some individuals, and interrupted medially by a narrow bridge. AVS3 aligned with, or slightly posterior to, AVS2. Very rarely is AVS3 anterior to AVS2. In all cases, both AVS3 and AVS2 are markedly posterior to the ADS.

Paratergal plates narrow, narrowing further posteriorly ( Fig. 10b). Tergal heads long, blunt, and re-entrant, with a broad tergal bar. In females, the tergal heads may be very broad, in some individuals they are approximately as broad as long. Paratergal plates and tergal heads less heavily pigmented in segment III than in more posterior segments. Tergal head on segment VII squat but pointed, and is usually transparent in females. Tergite III partially divided medially. Females pale. Many males with an extra submedian seta on ventral side of segment VII.

Female genital lobes oval with 3–5 posterior marginal setae, 3 sub-marginal setae, and 6 median margin setae ( Fig. 10c). Mesomere particularly short and stocky ( Fig. 10d). Dorsal mesomeral fingers very slender. Lower endomere narrow, with more or less parallel anterior and posterior margins. Apodemal bridge diffuse anteriorly.

Discussion: Gustafsson and Olsson (2012) found that Lunaceps populations from C. minuta , C. ferruginea , L. falcinellus , and C. ruficollis were genetically identical. The oldest available name for this clade is L. falcinellus . For a discussion on L. incoenis , the identity of its type specimen, and the assignment of host associations, see the entry for L. incoenis below.

Lunaceps falcinellus is almost indistinguishable from L. incoenis , but the tergal heads of abdominal segment VII are broader and blunter in L. falcinellus than in L. incoenis . In L. falcinellus , many males have two ventral setae on abdominal segment VII but, because they are sometimes missing, it is not a reliable character. Apart from L. incoenis , the most similar species to L. falcinellus is L. superciliosus sp. nov. Females of L. falcinellus and L. superciliosus sp. nov. differ in the number of genital setae, but this character is somewhat unreliable, and should not be used exclusively because the posterior marginal setae in particular have a tendency to fall off or perhaps never develop fully. Most males of L. superciliosus sp. nov. have a small suture medio-posterior to the antennae which readily separates it from L. falcinellus . However, this character can be hard to see in some individuals, and is missing entirely in female L. superciliosus .

It should be noted that Timmermann (1954a) included several individuals of an unidentified Quadraceps species among his paratypes for L. falcinellus , but both the type and the allotype are Lunaceps .

Ansari’s (1956) description of L. husainii mentions that this species could be separated from all other species of the genus by the shape of the head and body, and by the male genitalia. This is reiterated in his second description of the species ( Ansari, 1958); however this is accompanied by illustrations matching L. falcinellus as defined here. Price et al. (2003) listed L. husainii as a junior synonym of L. incoenis , following Ledger (1980: 168). Therefore, ours is a new synonymy for L. husainii under L. falcinellus .

Bechet (1968) mentioned “cut-off” paramere extremities in his description of L. timmermanni , but nothing resembling this has been found in any specimen from C. ferruginea studied by us. In the photo included with the original description it is not clear what that feature would look like, and the possibility that this is due to Bechet’s material being damaged cannot be excluded. The material studied by us cannot be morphologically separated from those collected from other hosts of L. falcinellus , and therefore we regard L. holophaeus timmermanni as a junior synonym of L. falcinellus .

Etymology: After the species name of the type host, Limicola falcinellus . The name means “little sickle”, and likely refers to the shape of the bill.

Material examined:

Ex Limicola falcinellus falcinellus

Holotype: ♂ 1, Pakistan?: Mekran Coast: Aisnee, 16 February 1872, J. Waterston Collection 1930-232 ( NHML). [ Note : The Mekran Coast lies in Iran and Pakistan, but the locality is given as “ India ” on slide. We have been unable to locate “ Aisnee ”.]

Allotype: ♀ 1, Pakistan: Karachi [slide has the locality as “ India ”], 8 February, 1872, J. Waterston Collection 1930-232 ( NHML).

Paratypes: ♂ 1, Pakistan?: Mekran Coast: Aisnee , 16 February 1872, J. Waterston Collection 1930-232 ( NHML) . ♀ 1, ♂ 2, Pakistan: Karachi [slide has the locality as “ India ”], 8 February, 1872, J. Waterston Collection 1930-232 ( NHML) . ♀ 1, India: Calcutta , 28 December 1872, J. Waterston Collection 1930-232 ( NHML) . ♀ 1, Pakistan?: Mekran Coast: Aisnee , 16 February 1872, J. Waterston Collection 1930-232 ( NHML) . ♀ 2, Karrachee [= Pakistan: Karachi?], 1 February 1872. J. Waterston Collection 1930-232 ( NHML) . ♂ 3, Norway: Finnmark, 22 November 1883, J. Waterston Collection 1930-232 ( NHML) . ♀ 1, Karalla Klang [= Malaysia: Selangor: Karala: Klang?], 15 January, 1880, J. Waterston Collection, 1930-232 ( NHML) .

Non-types: ♂ 1, Namibia [ Slide has the locality as “ South West Africa ”]: Sandwich Harbour, 26 January, 1973, BM 1974-33 ( NHML) . ♂ 2, India ( NHML) . ♂ 1, Poland: Piaseczno : [Konstancin-] Jeziorna, 27 August 1954, Yurezyk, 42/ac/1-1 ( MFN) . ♀ 3, ♂ 2, [ Collection data not given on slide], 20070g, m, o, p, r ( MFN) . ♂ 1, Sweden: Öland , Ottenby Bird Observatory, 25 May 2007, ID: 103, Band number: 3531454, D. Gustafsson ( SMNH) . ♀ 1, Sweden: Öland : Ottenby Bird Observatory, 25 May 2007, ID:106, Band number: 3531453, D. Gustafsson ( SMNH) . ♀ 1, Sweden: Öland : Ottenby Bird Observatory, 29 May 2007, ID: 134, Band number: ob29312, D. Gustafsson ( SMNH) . ♂ 1, Sweden: Öland , Ottenby Bird Observatory, 25 May 2007, ID: 103, Band number: 3531454, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland : Ottenby Bird Observatory, 25 May 2007, ID:106, Band number: 3531453, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland : Ottenby Bird Observatory, 29 May 2007, ID: 134, Band number: OB29312, D. Gustafsson ( GNM) .

Ex Limicola falcinellus sibirica

♂ 1, Australia: Western Australia: Broome Area: Richard’s Point , 11 November 2008, ID: 830, Band Number: 042-43332, D. Gustafsson ( GNM) . ♀ 1, Australia: Western Australia: Broome Area: Richard’s Point , 11 November 2008, ID: 829, Band Number: 042-43330, D. Gustafsson ( SMNH) . ♀ 1, Australia: Western Australia: 80 Mile Beach, Near Anna Plain Cattle Station , 17 November 2008, ID: 892, Band Number: 042-43482, YIO-P-01156, D. Gustafsson ( YIO) .

Ex Calidris ferruginea (synonyms: Erolia ferruginea or Erolia testacea )

♀ 10, ♂ 5, Morocco, October 1939, 11953 ( NHML) . ♀ 1, New Zealand: Lake Ellesmere , January 1928, E.F. Stead, R. C.L. Pilgrim Collection (Cmu Av L588) ( MONZ) . ♂ 1, Australia: Victoria: Western Port Bay: Stockyard Point , 13 January 1996, W. Steele ( MONZ) . ♀ 3, ♂ 10, Estonia, August 1934, Meinertzhagen Collection, 1444 ( NHML) . ♀ 1, ♂ 1, Sudan, May 1936, Meinertzhagen Collection 7946 ( NHML) . ♀ 6, ♂ 5, Papua New Guinea: Western District: near Wando , 14 October 1969, I.L., Owen, 691014/11b, BM 1970-381 ( NHML) . ♀ 1, ♂ 1, Angola: Port Alexander , 20 September 1912, 964, BM 1954-318 ( NHML) . ♀ 1, ♂ 4, Kenya, April 1916, Meinertzhagen Collection 3880 ( NHML) . ♀ 2, ♂ 6, Seychelles: Aldabra: South Island: Cing Cases , January 1968, B. Cogan and A. Hutson, Roy. Soc. Exp., BM 1968-333 ( NHML) . ♀ 1, United Kingdom: England: Weymouth , 8 October 1910, J. Waterston Collection BM 1930-232 ( NHML) . ♀ 1, Malawi: Lake Chilwa [slide has this as “Nyasaland”], 21 November, 1960 ( NHML) . ♀ 1, ♂ 1, Mauritius, 30 November 1973, BM 1974-4 ( NHML) . ♀ 1, ♂ 1, Kenya: Nakuru, January 1956, Meinertzhagen Collection 20420 ( NHML) . ♀ 1, ♂ 1, Sri Lanka [as “Ceylon”]: Northern Province: Mullaitivu , 25 March 1955, BM 1955-299 ( NHML) . ♀ 1, Cyprus: Akrotiri, 21 September 1967, K.C. Emerson Collection ( OSU) . ♀ 4, ♂ 5, India: Tamil Nadu: Point Calimere, June–September , 1969, K.C. Emerson Collection ( OSU) . ♀ 2, ♂ 2, Thailand: Samutprakan: Bang Pu , 29 September 1965, K.C. Emerson Collection ( OSU) . ♂ 1, Australia: Western Australia: Broome Area: Richard’s Point , 11 November 2008, ID: 842, Band Number: 042-43406, D. Gustafsson ( GNM) . ♂ 1, Australia, Western Australia, Broome Area: Richard’s Point , 11 November 2008, ID: 840, Band Number: 042-43364, D. Gustafsson ( SMNH) . ♂ 1, Australia, Western Australia, Broome Area: Richard’s Point , 11 November 2008, ID: 838, Band Number: 042-43353, YIO-P-01157, D. Gustafsson ( YIO) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 30 July 2007, ID: 253a-1, Band number: 3533877, D. Gustafsson ( YIO) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 23 July 2007, ID: 168a-1, Band number: 3532199, YIO-P-01158, D. Gustafsson ( YIO) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 30 July 2007, ID: 253a-2, Band number: 3533877, YIO-P-01159, D. Gustafsson ( YIO) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 23 July 2007, ID: 168a-2, Band number: 3532199, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 23 July 2007, ID: 167, Band number: 3532168, D. Gustafsson ( SMNH) . ♀ 1, Sweden, Öland: Ottenby Bird Observatory , 25 July 2007, ID: 196, Band number: 3532705, D. Gustafsson ( SMNH) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 30 July 2007, ID: 253a-1, Band number: 3533877, D. Gustafsson ( SMNH) . n#1, Sweden: Öland: Ottenby Bird Observatory , 26 July 2007, ID: 208, Band number: 3533046, D. Gustafsson ( SMNH) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 21 May 2007, ID: 72, Band number: 3531328, D. Gustafsson ( GNM) . ♀ 1, ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 24 July 2007, ID: 178-1, 178-2, Band number: 3532387, D. Gustafsson ( GNM) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 25 July 2007, ID: 196-s, Band number: 3532705, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 25 July 2007, ID: 199, Band number: 3532812, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 30 July 2007, ID: 250, Band number: 3533595, D. Gustafsson ( GNM) . ♀ 1, Sweden: Öland: Ottenby Bird Observatory , 2 August 2007, ID: 279, Band number: 3534673, D. Gustafsson ( GNM) .

Ex Calidris minuta (synonym: Erolia minuta )

♀ 1, ♂ 1, Israel: Kfar Masaryk , 26 August 1958, BM 1959-56 ( NHML) . ♀ 2, ♂ 1, Kenya, March 1936, Meinertzhagen Collection (7264, 7268) ( NHML) . ♀ 4, ♂ 1, Uganda, April 1936, Meinertzhagen Collection (7712) ( NHML) . ♀ 1, ♂ 1, Yemen: Aden, December 1948, Meinertzhagen Collection (17948) ( NHML) . ♀ 1, Arabia, January , 1948, Meinertzhagen Collection (17123) ( NHML) . ♀ 1, Angola: Port Alexander , 23 October 1912, 1092, BM1954-318 ( NHML) . ♀ 2, ♂ 1, Uganda: Kayanja?: Tolo ?, 28 April 1944, Hopkins Collection ( NHML) . ♀ 1, ♂ 1, V. Dvår ?, 9 September 1950, Coll. Dr. F. Balát ( NHML) . ♀ 1, Iraq: Baghdad, L. Harrison ( NHML) . ♂ 1, Sri Lanka [as “Ceylon”]: Southern Province: Bundala , 24 September 1954, BM 1954-699 ( NHML) . ♂ 1, Sri Lanka [as “Ceylon”]: Hambantona District: Bundala lagoon, 6 March 1955, BM 1955-332 ( NHML) . ♀ 3, India: Tamil Nadu: Thanjavur: Point Calimere , 26 September 1969, K.C. Emerson Collection ( OSU) . ♂ 2, India: Rajasthan: Bharatpur , 15 February 1971, K.C. Emerson Collection ( OSU) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 21 May 2007, ID 71-1, Band number: 2kn50208, D. Gustafsson ( SMNH) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 21 May 2007, ID 71-2, Band number: 2kn50208, D. Gustafsson ( GNM) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 21 May 2007, ID 77-1, Band number: 2kn50207, D. Gustafsson ( SMNH) . ♂ 1, Sweden: Öland: Ottenby Bird Observatory , 21 May 2007, ID 77-2, Band number: 2kn50207, D. Gustafsson ( GNM) .

Ex Calidris ruficollis (synonyms: Erolia ruficollis , Ereunetes ruficollis , Ereunetes minuta ruficollis or Pisobia minuta ruficollis )

♀ 2, ♂ 2, Australia: Western Australia: Perth , August 1967, R. H. Stranger, BM 1969-48 ( NHML) . ♀ 3, ♂ 3, Australia: Tasmania: Ralphs Bay , 20 November 1981, K. Heiden ( MONZ) . ♂ 1, New Zealand: Rangitikei River Mouth, 28 February 1983, F. Sibley, NM 22743 ( MONZ) . ♀ 1, Japan: Tyototen ?, 31 August 1957, BM 1958-245 ( NHML) . ♀ 1, Papua New Guinea: Western District: Wando , 14 October 1969, I. L. Owen, 691014/3c, BM 1970- 381 ( NHML) . ♀ 1, Myanmar: Rangoon, April 1900, Meinertzhagen Collection 4297 ( NHML) . ♀ 3, ♂ 6, Philippines: Luzon: Batangas: Calatagan , September–December 1964, K.C. Emerson Collection ( OSU) . ♀ 1, ♂ 1, Thailand: Samutprakan: Chao Praya River , 23 September 1965, K.C. Emerson Collection ( OSU) . ♀ 1, Australia: Western Australia: Broome Area: Two Dog Hermit , 12 November 2008, ID: 844, Band number: 036-13725, D. Gustafsson ( SMNH) . ♂ 1, Australia: Western Australia: Broome Area: Coconut Wells , 9 November 2008, ID. 817, Band number: 036-13701, D. Gustafsson ( GNM) .

Ex Xenus cinereus (synonym: Tringa terek ) (possibly stragglers):

♂ 1, Sri Lanka [as “Ceylon”]: Hambantota: Bundala , 23 October 1955, W. W. A. Phillips, 1956-133 ( NHML) . ♂ 1, India: Tamil Nadu: Point Calimere , 7 July 1969, K.C. Emerson Collection ( OSU) .