Tiphiidae wasps of Madagascar ( Hymenoptera , Tiphiidae )

The tiphiid wasp fauna of Madagascar consists of 28 endemic species, including 12 species of Anthobosca (Anthoboscinae), nine species of Methocha (Methochinae), two species of Tiphia (Tiphiinae) and in the Myzininae three species of Meria, eight species of Mesa and two species of Myzinella. Seven species of Methocha, arcuata, depressa, flavipalpus, impunctata, nasiformis, robusta and strigosa, and one species of Myzinella, minima, are described as new. All of the genera are shared with mainland Africa.


Introduction
Madagascar has a relatively diverse, though seemingly recent tiphiid fauna.The least derived tiphiid subfamily Anthoboscinae is particularly species-rich on Madagascar, with nearly as many species as are found on continental Africa (Kimsey 2009).Three other tiphid subfamilies, Methochinae, Myzininae and Tiphiinae, are present as well, although they are less diverse relative to the mainland fauna than are the anthobos-cines.None of the Malagasy tiphiid genera are endemic to the island, but all of the tiphiid species in Madagascar are endemic.There are seven myzinine genera in the Afrotropical Region including Braunsomeria Turner, Meria Illiger, Mesa Saussure, Myzinella Guiglia, Parameria Guérin de Menevile, Poecilotiphia Cameron and Pseudomeria Saunders.Only Meria, Mesa and Myzinella are recorded from Madagascar.The subfamilies Methochinae and Tiphiinae are represented by species in the genera Methocha and Tiphia respectively.
The modern Malagassy tiphiid fauna clearly derives from mainland Africa.These wasps are strong fliers but many species have wingless or brachypterous females.Tiphiid wasps are, where known, parasites of beetle larvae, particularly in the Scarabaeidae and Cicindellidae, Their host larvae are largely found in the soil and the tiphiid females spend considerable time on the ground, which makes rafting on plant debris unlikely but not impossible.Thus, dispersal of tiphiid wasps across the oceanic channel to Madagascar from mainland Africa seems unlikely.However, over a period of tens of millions of years even a small number of successful dispersals might be sufficient to establish populations on the island (Tattersall 2008).Studies of other animals with relatively poor dispersal capabilities, such as non-flying vertebrates (Vences 2004) and ants (Fisher 1996) show a similar pattern of origin from Africa.
The California Academy of Sciences' Arthropods of Madagascar Project funded by the U.S. National Science Foundation revealed a more diverse tiphiid fauna than previously recorded, including seven new species of Methocha and one species of Myzinella that are described below.

Materials and methods
Specimens in this study were obtained from the California Academy of Sciences Arthropods of Madagascar Project and the Bohart Museum of Entomology.Holotypes and paratypes are deposited in the California Academy of Sciences, San Francisco (SAN FRANCISCO) and the Bohart Museum of Entomology, University of California, Davis (DAVIS).Other type repositories are indicated by the city names in parentheses in the species lists: BERLIN = Museum für Naturkunde, Berlin, Germany; BOLOGNA = Bologna Zoological Museum, Bologna, Italy; GENEVA = Museum of Natural History, Geneva, Switzerland; ITHACA = Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands; LONDON = Department of Entomology, the Natural History Museum, London; PARIS = Muséum National d'Histoire Naturelle, Laboratoire d'Entomologie, Paris, France; VIENNA = Naturhistorisches Museum, Vienna, Austria, and WASHINGTON = U.S. National Museum, Washington, D. C., USA.
The systematics of members of the family Tiphiidae is largely based on males.Females are less commonly collected and association of the sexes is problematic at best.This situation is also true in Madagascar, as a result the species level keys below are only to males where indicated as most of the females are unknown or unassociated.similar to the face in Tiphiinae.There are fourteen Anthobosca species described from mainland Africa and ten from Madagascar (Kimsey 2009).Hosts are unknown for the subfamily.Discussion.The genus Mesa is found from the Afrotropical Region, including Madagascar, north and east into Iran and Myanmar.Mesa males are characterized by having metasomal segment I more than three times as long as broad and the terga without a transverse subbasal carina or ridge.Hosts are unknown for the genus.

Key to species of
Although Bartalucci (2005) removed Mesa hova (Turner) from synonymy under nodosa (Guérin de Meneville 1837) he did not clearly describe the male and it is unclear, which male is associated with the female type of hova.Discussion.Bartalucci (2005) published the first record of Myzinella from Madagascar, although the genus is known from mainland Africa, India and Iran.A second, new species, minima, was present in the materials examined for this review from the California Academy of Sciences Madagascar project and is described below.

Key to species of
The most distinctive features of male Myzinella are the abruptly up curved uncus, short, nodose metasomal tergum I, metasomal terga III-VII with an abruptly elevated transverse ridge or flange separating the smooth anterior surface that slides beneath the preceding tergum from the tergal disk and the subapical metasomal sternum is broadly truncate apically.
Hosts: unknown for the genus.Diagnosis.Myzinella minima can be readily distinguished from Myzinella festiva by the black metasomal segment I, evenly convex and ovoid tegula and apically thickened antenna.Myzinella minima is also smaller than festiva, which ranges from 10-12 mm long.

Key to species of
Description.Male (Fig. 6).Body length.5-8 mm.Face (Fig. 7).Oral cavity unmodified; genal bridge opaque, barely covering the tongue base; vertex impunctate between hindocellus and eye, except for single row of punctures along eye margin; flagellomere I twice as long as broad; flagellomeres II and III 1.7× as long as broad; flagellomere XI 1.4× as long as broad, apical flagellar segments twice the breadth of basal segments.
Thorax.Pronotum more than twice as broad as long; metasomal dorsum and mesopleuron with punctures separated by 0.5-2.0puncture diameters; tegula ovoid, evenly convex; forewing apical cells separated from wing apex by more than length of discoidal cell.
Metasoma.Tergum I highly polished, punctures 2-4 or more puncture diameters apart; terga I-VII polished with punctures 2-4 puncture diameters apart; tergum VII apically acute on either side of apicomedial notch; sternum I with punctures 1-2 puncture diameters apart; sterna II-V with transverse medial and apical bands of punctures; sternum VI impunctate, except for medial patch of punctures.
Color.Black, with pale yellow markings, mandibles pale yellow, pronotum with yellow transverse band, tegula pale yellow, femoral apices, tibiae partly and tarsi completely yellow, metasomal terga II-VI with medial and lateral yellow spots, tergum I with short transverse yellow band along posterior margin; wing veins whitish basally, black apically and medially, wing membrane whitish.
Vestiture.Long white, largely decumbent.Diagnosis.The most distinctive features of arcuata are the flattened clypeus, welldeveloped antennal lobes and propodeal sculpturing consisting of fine transverse to dorsally arcuate ridges.This species most closely resembles arnoldi but can be distinguished by the longer flagellomeres and the finely ridged propodeum.
Metasoma.Terga highly polished, punctures tiny and separated by 4 or more puncture diameters; sterna impunctate except for basal band of punctures 1 puncture diameter apart.Genitalia (Fig. 22).Paramere slender, with small medial angle, marked by transparent lobe, breadth at medial angle 2.0-2.3×breadth at halfway point between apex and medial angle, paramere length versus breadth at submedial angle is 6×.
Color.Black, with pale yellow mandibles and fore and midcoxae, fore and mid legs may be yellowish brown; palpi whitish; wing membrane faintly brown-tinted apically.
Vestiture.Sparse erect, silvery.Diagnosis.The most distinctive features of depressa are the coarsely areolate propodeum, the shape of the frontal projection above and between the antennal lobes and the sublaterally impressed pronotal disk.This species does not closely resemble any of the other Malagasy Methocha species, although the depressed pronotum resembles that of robusta, which differs in having the pronotal depression interrupted medially.
Vestiture.Sparse erect, silvery.Diagnosis.Methocha flavipalpus resembles nasiformis and strigosa based on the strong medial clypeal projection.It can be distinguished from those species by the coloration of the parategula, which has a transparent margin and the yellowish mandibles, palpi and legs; all black to dark or reddish brown in nasiformis and strigosa.
Color.Black; mandibles and palpi whitish to yellow; fore and midcoxae may be pale brown, fore and mid legs brown with paler joints and tarsi; parategula bicolored, brown and whitish with transparent margin; wing membrane untinted, except posterior margin slightly brown-tinted in some specimens.
Vestiture.Sparse erect, silvery.Diagnosis.The highly polished, largely impunctate and unsculptured body is the most distinctive feature of impunctata.The unsculptured propodeum will immediately separate it from arcuata, which shares the largest number of characteristics with impunctata.
Color.Black, with dark red mandibles, scapal carina yellow legs black to dark brown, trochanters sometimes paler; palpi reddish; wing membrane brown-tinted.
Vestiture.Sparse erect, silvery.Diagnosis.Methocha robusta is one of two species with a medial depressed pronotal disk.It can be distinguished from the other species, depressa, by the frons concave between the antennal lobes and the strongly convex clypeus.In addition, the pronotal depression lacks the longitudinal medial elevation seen in depressa.

Meria species are sexually dimorphic, as are members of the genera Mesa and Myzinella. Males are elongate and slender with straight, cylindrical antennae. Females are heavy-bodied, with coiled antennae. Males also have the distinctive hook- like apical metasomal sternum (uncus) found in other myzinines. Meria males can be
distinguished from Mesa and Myzinella by the short, broad metasomal sternum I and petiolate forewing submarginal cell.Hosts are unknown.
Methocha nasiformis is one of three species with an acute or even digitate medial clypeal projection.It can be distinguished from flavipalpus by the black, concolorous parategula, dark legs and nearly straight apical clypeal margin (features shared with strigosa).It can be distinguished from strigosa by the coarsely punctate, not finely ridged propodeum.