Spania Meigen, 1830

Kerr, Peter H., 2010, 2592, Zootaxa 2592, pp. 1-133 : 97-98

Genus Spania Meigen

Figs. 19, 60, 81, 104, 148, 163.

Spania Meigen 1830: 335 . Type species Spania nigra Meigen 1830 , by monotypy.

Archicera Szilády 1934a: 264 . Type species Archicera avavorum Szilády 1934a , by monotypy.

Diagnosis. I consider the reduced form of the female first segment of the cercus (approximately half the length or less of the fairly elongate second segment) and the form of female sternite 8, which lacks a medial invagination along its posterior margin, as autapomorphic character states that define Spania . Since the closest relatives of Spania are mandibulate ( Spaniopsis and Symphoromyia ), the loss of mandibles may represent an additional autapomorphy.

Spania species are small (2.1 to 3.0 mm), dark brown to black in color, and slight in build. Wings are lightly infuscate, especially along costal vein; male holoptic, eyes separated in female; antenna with stylus arising ventrally or terminally from enlarged flagellomere base, laterally compressed; mandibles absent; laterotergite bare; M 3 completely or incompletely present; tibial spur formula 0:2:0; hind tibia without macrochaetae; tergite 9 with ventrolateral arms, extending posteriorly, surrounding and fusing to sternite 9 laterally; female spermathecal ducts with accessory glands arising near base of spermathecae. Spania is most likely to be confused with Ptiolina or Bolbomyia . It may be distinguished most easily from Ptiolina species by the absence of hind tibial spurs and its small size. Spania is approximately the same size as Bolbomyia , however, it has a stylate antenna and M 3 at least incompletely present, and lacks fore and hind tibial spurs. Spania was once synonymized with Spaniopsis ( Paramonov 1962) , however Spania may be distinguished from this genus by its small size, its delicate build, scape clearly smaller than the pedicel, and mandibles absent. Spania is restricted to the Holarctic Region, with a distribution that includes North America, Europe, and Japan.

Description. Head. Clypeus bulbous. Scape approximately same size as pedicel. First flagellomere of antenna enlarged, laterally compressed, bearing ventro-apical stylus. Eyes in male ommatidia split into dorsal and ventral areas and smaller ventrally. Eyes inconspicuously setulose; in female, dichoptic; in male, holoptic. Parafacials in male not swollen. Labella with pseudotracheae, longer than palpus. Theca short and stout, with medial suture. Palpus one-segmented. Lacinia present, but very reduced. Lacinia apex not serrated. Mandibles absent. Cibarial pump short, as wide as long or wider. Cornu shorter than cibarial pump. Pharyngeal pump approximately same length as cibarial pump (including cornu).

Thorax. Mesonotum lacking vittae, black or brown, without dorsocentral bristles. Anepisternum bare. Postspiracular sclerite smooth, bare. Proscutellum narrowly present or absent. Subscutellum slightly swollen or not. Laterotergite bare. Wing hyaline, without markings. Lower calypter reduced. Upper calypter well developed, with broad curvature, lobe-like, width twice length or less. Costa extends past wing tip. Humeral crossvein well developed. Sc-r crossvein absent or weakly developed, positioned distal to h by approximate length of h. Dorsal side of R 1 setulose, ventral side bare. All other wing veins bare. R 1 and R 2+3 separated at wing margin. R 2+3 sinuous, apical third ultimately bends anteriorly slightly, toward leading edge of wing margin. Length of R 2+3 about as long as R 5. Base of R 4 –R 5 fork proximal or directly above distal end of cell dm. R 4 mostly straight apically. R 5 ending at wing tip or anterior to wing tip, clearly longer than R 4+5 (r-m to R 4 origin). R-m crossvein at proximal one-third to near halfway of discal cell. M 3 wing vein incompletely present (not reaching margin) or complete, reaching wing margin. Origin of CuA 1 at discal cell or at crossvein separating discal and basal medial cells; m-cu crossvein absent. Length of CuA 2 v. posterior vein of cell bm greater than 1/2 length, less than 2/3 length of posterior vein of cell bm. Alula with broad curvature that is slightly shifted distally. Anal lobe well developed. Cell cu p open or closed. Halter knob approximately 1/2 length of stem. Tibial spur formula 0:2:0. Hind coxal tubercle absent. Hind tibial macrochaetae absent. Postmetacoxal bridge absent.

Abdomen. Abdominal segments evenly tapered. In female, tergite 7 much wider than long; intersegmental membrane between segments 7 and 8 short, as throughout abdomen; sternite 8 sclerite entire, wider than long. Male terminalia with epandrium simple, not containing hypandrium ventrally. Epandrium wider than long, strongly notched anteriorly. Tergite 10 absent. Hypoproct elliptical, flattened, tomentose, without setae. Cercus directly adjacent to epandrium; widely displaced from one another, separation distance greater than three quarters width of cercus; held vertical in relation to rest of abdomen; in posterior view cupped, forming circular outline medially. Hypandrium fused entirely to gonocoxites. Gonocoxite with sinuous dorsal ridge, leading to gonocoxal apodemes. Gonocoxal apodemes short, not reaching anterior margin of hypandrium. Sperm sac slightly bulbous ventrally, produced into slightly developed paired lobes or smooth. Ejaculatory apodeme laterally compressed; long, reaching anterior margin of hypandrium. Lateral ejaculatory processes present, integrated into sperm sac membrane. Aedeagal tines absent. Endoaedeagal process absent. Female terminalia with tergite 9 entire, with narrow anteriorly-directed ventrolateral projections, enveloping sternite 9. Spermathecae three, spherical, lightly to moderately sclerotized. Spermathecal ducts no more than three times length of sternite 9, with swelling halfway between genital chamber and spermathecae. Spermathecal duct accessory glands arise at base of each spermatheca. Spermathecal ducts sclerotized and thickened into narrow ring near junction with common spermathecal duct, otherwise smooth, not enlarged, and unsclerotized. Common spermathecal duct thickened, short, shorter than longest diameter of genital chamber. Genital chamber teardrop shaped, small, occupying fraction of sternite 9 area. Accessory gland posterior to genital chamber present, inconspicuous, easily overlooked even after staining. Sternite 9 anterior end broadly paddle-shaped. Sternite 9 with broad posterolateral projections that are held at an angle. Tergite 10 present, entire, short (length less than half width). Sternite 10 present, entire, roughly rectangular. Cercus twosegmented. Basal cercal segment not elongated, without ventral process, separated from each other dorsally by approximate width of second cercal segment. Ventral lobes of first segment of cercusdo not curve ventrally towards one another to form ring. Second cercal segment narrow, elongated (~2.5x longer than wide or more), without apical sensory pits.

Larva. Undescribed, but see Biology below.

Biology. Although the larvae of Spania nigra Meigen have not been characterized, Mik (1896) reportedly found a Spania nigra larva mining in the thallus of Pellia neesiana (Bryophyta: Pelliaceae ), a liverwort species ( Nartshuk 1995).

Literature. Nagatomi & Saigusa (1982) give a key to the Japanese fauna that includes all species.

Notes. Szilády (1934: 264) distinguished Archicera from Ptiolina and Spania by the antennal flagellum, which he stated, had faintly visible divisions. This feature, as Nagatomi (1982a: 54) has noted, is within the morphological variation already present within Spania (see Nagatomi & Saigusa 1982). On account of the small size of Archicera avarorum and the description of the flagellomere as being lanceolate, this species is certainly placed among the Spania and has been treated as a synonym of Spania nigra ( Majer 1988) . The holotype of Archicera avarorum has been destroyed [ Hungary National History Museum, Budapest]. Therefore a neotype, preferrably a specimen from either Austria or Croatia, needs to be designated for Archicera avarorum so that its appropriate taxonomic status may be documented formally.