Tonnoiriella veronikae sp. nov.

(Figs 19–35)

Type material. Holotype ♂: Madagascar, Toamasina prov., Analamazaotra 1.4 km SSW Andasibe vill. (Périnet), cca 940 m a.s.l., 48° 24' 48'' E, 18° 56' 09'' S, S on the first place (see N. nyahururuensis) 20.i.2000, P. Chvojka leg. Slide with a dissected specimen, Cat. No. 34699, Inv. No. 22010 (NMPC).

Paratypes 2 ♂ (slides, NMPC): The same locality, date and collector, Cat. No. 34700–34701, Inv. No. 22011– 22012 (NMPC). All material P. Chvojka leg. (by sweep netting).

Type locality. Madagascar: Toamasina prov., Analamazaotra 1.4 km SSW Andasibe vill. (Périnet).

Etymology. The new species name (adjective) is based on the first name of the granddaughter of the first author: Veronika Tomková.

Bionomics. Unknown. Adults were collected in secondary forest habitat, specifically a swampy area with a nearby rill.

Distribution. Currently known only from Madagascar.

Description. Male. Head 0.95 times as long as broad (Fig. 19). Vertex inflated dorsally, frons with numerous setae alveoli in a stripe, divided and divergent both near vertex and near frontal suture, sometimes hardly divided in the vertical axis, sparse on both sides of head margins (Fig. 19). Eye bridge is moderately developed, of three complete facet rows (Fig. 31), upper apices of eyes tapered medially. Eyes separated by about three facet diameters, connected by arched interocular suture, a little strengthened laterally, sometimes interrupted medially (Fig. 31), with a few postocular bristles (4–6). Setae alveoli of frontoclypeus arranged in a strangulated triangular central scar patch near basis of antennae tapering to a tongue-shaped area below frontal suture. Ratio of distances of apices of eyes (tangential points) to minimum width of frons approximately 2.5:1. Antennae (Figs 25, 32) with 16 articles. Scape slender basally, pedicel almost ball-shaped, 1.5 times as long as scape. Scales of pedicel broadly lanced in contrast to stiletto-shaped scales of fagellomeres. Flagellomeres mostly spindle-shaped, however the last three flagellomeres a little reduced in size (cask-shaped or ovoid). Antennomere 14 with a hemispherical axial apiculus. Sensory filaments (ascoids) nedle-shaped, bent, paired, 2.8 times shorter than the flagellomeres bearing them. Mouthparts extending beyond basal palp segments. Length ratio of maxillary palps 1.0:2.0:2.1:3.8, the last segment is annulate, connected somewhat subapically with preceding one (Fig. 20). For the terminal lobes of labium, as shown in Fig. 33, the lines of spines between both lobes are not developed. Ratio of maximum length of cibarium (Figs 19, 26) to length of epipharynx 1.7:1, labrum with microsetae.

Thorax with mesothoracic allurement protuberance almost flattened, with a circular thoracic spiracle anteriorly, anepisternum with a triangular setose patch (Fig. 21). Wings (Fig. 34) 1.6 mm long - holotype, both paratypes 1.5 mm, well sclerotised, lancet-shaped, not enlarged in anal and humeral areas, with rounded top, clear, wing membrane bare. Basal costal nodes distinct, Sc straight, uninterrupted. Medial fork incomplete in contrast to radial one. R5 ends beyond wing apex. Bases of M3, CuA1 and CuA2 are not connected. The ratio of maximum wing length to maximum of width 2.7:1. Strengthened veins: Sc (thickened basally), whole R2 and R5, all veins distally, however, R4 and M1 stout also in basal cell, CuA1 as well distally and basally with a declined turn. CuA2 strongly thickened and arched at basis, gradually tapering distally. Halteres (Fig. 27) have a prolonged stem and ovate knob with a row of minute hairs laterally. Ratio of maximum length of halteres to their maximum width approximately 2.6:1. Length ratios of femora, tibiae and first tarsal segments: P1 1.9:2.0:1.0, P2 1.9:2.5:1.2, P3 2.0:2.8:1.2. Claws of P1 bulbose at base, bent, haired in two basal thirds, as shown in Fig. 28, twice pointed, subapical tooth is blunt in contrast to the sharp prolonged terminal spike.

Male genitalia with ejaculatory apodeme straight in dorsal view, almost parallel-sided distally with an ovoid chamber, widened proximally in convergent margins, terminating anteriorly in a sclerotized axial rim (Fig. 30), compressed dorso-ventrally and bent from lateral view (Fig. 24). Aedeagal complex asymmetrical, wrinkled basally (Figs 24, 30), distal part with a boomerang or sickle-shaped paramere, laterally with a cut side arch without a bulbous protuberance, superficial spines quite missing. Gonocoxites almost cylindrical (Figs 23, 30), with a stout seta on inner side basally and 2–3 small setae near distal part of the ejaculatory apodeme. Gonocoxal apodemes forming a pair of rectangular projections anteriorly (Fig. 30). Gonostyli conical, not gradually tapering, with a small swelling in the middle, hooked terminally. Length ratio of ejaculatory apodeme and gonostylus approximately 1.3:1. Epandrium (Figs 29, 35) almost bare, with two patches of scanty latero-caudal hair insertions. Basal paired apertures almost elliptical, not connected, developed in a proximal small dark area of epandrium. Remainders of ventral plate missing. Hypandrium (Figs 22, 30) narrow, triangular in the middle, conspicuously strangulated subapically, hemicircular terminally. Epiproct lobule-shaped, featureless (Fig. 35), hypoproct (Figs 29, 35) narrowed proximally, rounded distally, both parts haired. Epandrial clasping lobes slightly C-shaped (Fig. 35), almost cylindrical, only a little widened basally (Figs 29, 35), with five not frazzled tenacula subapically, as a fraction of surstylus (1.5:1).

Female. Unknown.

Differential diagnosis. The Afrotropical species of genus Tonnoiriella are morphologically rather uniform, except for the male genitalia (see Table 2). T. veronikae sp. nov. described here is very similar to T. drepanopenis (Duckhouse, 1975), which has relative lengths of three first antennomeres 1.4:1.7:1.0, relative lengths of palpal segments 1.0:1.1:1.2:2.5, hypandrium in the middle triangular, rounded terminally, caudal sclerite of aedeagal complex boomerang-shaped, distal margin of caudal sclerite arched horizontally, gonostyli gradually tapering and apex of gonostylus rounded. On the other hand T. veronikae sp. nov. differs by the relative length of antennomeres 1–3 1.2:1.3:1.0, relative length of palpal segments 1.0:1.7:1.8:3.2, hypandrium almost triangular in the middle, strangulated subapically, semicircular terminally, caudal sclerite of aedeagal complex almost boomerang-shaped, but with cut side arch, distal margin of caudal sclerite approximately straight horizontally, gonostyli not gradually tapering, with a small swelling in the middle, apex of gonostylus conspicuously hooky.

cracens drepanopenis fasciola stuckenbergi veronikae Comments. Wagner (1978, 1990, 1994) and Wagner & Salamanna (1984) included Neoarisemus in tribe Brunettiini; Vaillant (1982b, 1986, 1990), Wagner (1997a, b) and Quate & Brown (2004) placed it in Setomimini and Ježek (2004) in Mormiini (Brunettiina). Moreover, Duckhouse (1987) placed Neoarisemus in Maruinini, as followed by Kvifte (2012).

A tribal placement of Tonnoiriella was historically contraversial as well. Vaillant (1971, 1982a), Duckhouse (1975), Wagner & Salamanna (1984), Wagner (1979, 1987, 1990) and Ježek (1999) placed this genus in Pericomaini, Vaillant (1982b) and Wagner (1997a, b) in Setomimini and Duckhouse (1987), Wagner & Andersen (2007), Kvifte et al. (2011), Kvifte (2012) and Afzan & Belqat (2016) in Maruinini.

Duckhouse (1987) considered both Neoarisemus and Tonnoiriella in Maruinini by the anterior plate-like gonocoxal condyles and dorsoventral ejaculatory apodeme, radial wing fork before medial one, R2 is several times longer than R2+3, flagellomeres are barrell-shaped and narrower than scpe and pedicel. We follow here Duckhouse´s opinion in concordance with Kvifte (2012), however, Duckhouse (1991) additionally concluded that Maruinini are probably paraphyletic; ultimately, molecular evidence may be necessary to accurately resolve the placement of Neoarisemus and Tonnoiriella as well as future exact modern study.