Two New Labenopimpline Ichneumonids (Hymenoptera: Ichneumonidae) from the Upper Cretaceous of Southern Africa

ABSTRACT Two new species of Labenopimplinae, Labenopimpla orapa sp. n. and Rugopimpla botswana sp. n., are described from the Upper Cretaceous (Turanian) deposits of Orapa in Botswana. These are the first and only known Cretaceous Ichneumonidae from the southern hemisphere. The new species seem to be very similar to their relatives from the Cenomanian of the Russian Far East. This indicates a wide distribution of Ichneumonidae by the early Late Cretaceous at least.


INTRODUCTION
The parasitoid-wasp family Ichneumonidae consists of approximately 60 000 recent species worldwide (Wahl & Sharkey 1993). In spite of the great diversity and consistage of ichneumonid history has remained almost unexplored until recently. Until now only a few dozen Cretaceous ichneumonids have been described, and these are entirely from Asia.
Two subfamilies are known from the Early Cretaceous. The Tanychorinae Rasnitsyn, Rasnitsyn 1975, 1980Zhang 1991;Zhang & Rasnitsyn 2003;Kopylov 2010a). The Palaeoichneumoninae Kopylov, 2009 (three genera and 12 species) were less numerous during the Neocomian, but nearly completely supplanted the Tanychorinae in the later Early Cretaceous (Kopylov 2009). Both subfamilies failed to cross the Early/Late Cretaceous boundary. Instead of the archaic Tanychorinae and Palaeoichneumoninae, features of the modern Labeninae, Pimplinae and Tryphoninae, entered the fossil record in the early Late Cretaceous (Kopylov 2010b and present contribution). This subfamily was previously known only from the Cenomanian Obeshchayushchiy locality in the Russian Far East; the two new representatives of Labenopimplinae described below, found ichneumonids from Africa and the southern hemisphere. Although Brothers and Rasnitsyn (2003) listed four ichneumonids amongst the 108 specimens of Hymenoptera from Orapa, the specimens described below are the only ones complete enough for adequate treatment.

MATERIAL AND METHODS
The material studied came from the Orapa locality in north-eastern Botswana. The age of the deposits seems to be Turonian (Brothers & Rasnitsyn 2003;Gernon et al. http://www.africaninvertebrates.org.za 2009; and references therein). The fossils are stored in the Bernard Price Institute of Palaeontology (BPI), University of the Witwatersrand, Johannesburg, South Africa. The the organic remnants and the matrix, or using oblique incident light from the top left to emphasise sculpture. The drawings were prepared electronically by superimposition on the photographs and checked against the specimens, specially for the addition of sculptural details. The terminology for wing venation (see Fig. 2) is from Kopylov (2009 Type genus: Labenopimpla Kopylov, 2010 (by original designation).
The prime morphological characteristic differentiating this subfamily from the Palaeoichneumoninae is the presence of parallel (rather than posteriorly converging) notauli. Unfortunately, the notauli are not visible in either of the specimens described here. Nevertheless, they are assigned to the Labenopimplinae on the basis of their characteristically shaped areolet: Labenopimpla has the areolet pentagonal with a relatively long 4M and subvertical r-m, and Rugopimpla has a quadrangular areolet, but all Palaeoichneumoninae have the areolet pentagonal with oblique r-m. Thus, these the basis of the generic characters listed by Kopylov (2010b).
Turonian. Head and mesosoma poorly preserved, metasoma well preserved, ovipositor probably incomplete (apparent apex highly irregular); legs and antenna incomplete; fore and hindwings very well preserved.
Remarks: The description is based primarily on examination of counterpart specimen 25980 (Figs 3, 4). When recently sought, specimen 25240-1 was not found. It may currently be in the National Museum of Botswana, Gaborone, but this has not been ( Fig. 1) and sketches by A.P. Rasnitsyn.

Rugopimpla botswana sp. n.
Figs 5-7 Etymology: After the country of origin, Botswana; noun in apposition. Description: Entire body, wing veins and ovipositor very dark; antenna and legs less gellomeres 3× as long as wide, apical ones 1.5× so. Forewing with 1Rs&1M arched; r-rs nearly straight; 2Rs half as long as 2+3M and twice as long as 4M; r-m as long as 2+3M, with two bullae; 1m-cu 0.67× as long as 2Rs+M; 1m-cu&2Rs+M arched; ramulus twice as long as wide. Hindwing probably with 1Rs 2.5× as long as r-m; 1Cu and cu-a probably subequal in length, junction apparently weakly angled (venation unclear and confused because of superimposition of wings). Measurements (lengths) in mm: Antenna at least 5.2, head and mesosoma 2.7, metasoma 3.5, ovipositor at least 1.2 (visible free section), forewing 3.9. Comparison: Differs from R. vulgaris, R. fallax and R. matrona in 1Cu:cu-a and probably 1Rs:r-m ratios in the hindwing; also differs from R. vulgaris in having r-rs nearly straight, and from R. matrona in presence of r-m. Differs from R. angusticella in having areolet wider with 4M longer, and in presence of r-m and ramulus. Differs from R. macra in having 1Rs&1M arched, r-rs nearly straight, and longer ramulus.

DISCUSSION
Labenopimpla and Rugopimpla were recently described from the Upper Cretaceous (Cenomanian) deposits at the Obeshchayushchiy locality in the Russian Far East (Kopylov 2010b). The present two species from southern Africa are very similar to their congeners despite their origins almost from opposite ends of the Earth. Indeed, their localities are now separated by some 14 000 km, and were separated by the Tethys Ocean during the Cretaceous period. The source localities, Orapa and Obeshchayushchiy, are of nearly the same age (Rasnitsyn 2002;Brothers & Rasnitsyn 2003). This indicates an early expansion of distribution, the more so since even the mid-Early Cretaceous (Aptian) fauna demonstrates only the most basal ichneumonid subfamilies, Tanychorinae and Palaeoichneumoninae, which are apparently absent from Orapa and Obeshchayushchiy (Kopylov 2010b). All known Cretaceous ichneumonidbearing localities, other than Orapa, are concentrated in Central, Northern and Eastern Asia (Russian Transbaikalia, Taimyr and Far East, Mongolia, eastern China) and North America (New Jersey and Canadian ambers); nothing is known from North Africa and western Eurasia, the regions which could probably clarify the scenario of early ichneumonid origins and dispersal.

ACKNOWLEDGMENTS
The authors are grateful to M.B. Mostovski for help in preparation of this paper. The helpful comments of a reviewer, A.I. Khalaim (Zoological Institute, Russian Academy of Sciences, St Petersburg), are much appreciated. For DSK and APR, the study was supported by the Program of the Presidium of the Russian Academy of Sciences "Origin