313

KEYS Tú THE ETHIOPIAJ.'f -III MACQUARTIINAE BY The late F. 1. VAN ENIDEN Commonwealth Institute of Entomology (Accepted 12th May, 1959) (With 11 figures in the text) CüNTENTS , Page Introduction 313 Abhreviations 314 Deñnition of the subfamily Macquartiinae 315 Biology of tha Ethiopian Macquartiinae 315 Key ta the tribes of Ethiopian Macquartiinae 317 Key ta tbe sub-tribes of Echinornyini 320 l. :Macqaartiini 320 2. Wagneriini 332 3. Campyiochaetini 350 4. Helocerini 353 5. Nemoraeini 358 6. Acemyini 368 7. Gennariini 371 8. Thelairini 372 9. Minthoini 377 10. 384 11. Echinornyini 402 a. Ernestiina 402 b. Linnaernyina. 407 c. Echinomyina 467 References 486

[The present report was completed by DI' van Ernden with the exception of Olle descrip­ tion, of the figures (to which, however, he had made reference in the appropriate places) and of the section dealing with the biology of the group. The biological data have kindly been summarised by DI' S. V. Peris, of the Instituto de Edafología, Madrid, and the bibliographical references at the end of the paper were provided to support this parto It had been van Emden's intention to inelude a paragraph of thanks in the íntro• duction. This was not completed, but acknowledgments of help received are given at varíous places in the text, and on his behalí thanks are bere tendered to the authoritie" of the British Museum for the facilities which they have so kindly afforded. The present annotator is responsible fol' the deseription of J.l1arshallornyia and its single , for the text figures and for following the original orthography of the generie names Acemya, and Echinornya. As both forms of the Jatter part of each of these names (myia, mya) are given in the lexicon and as the original orthography corresponds with oue of thero, there is no evidence of positive error. None of the subsequent emendations has been placed on the Officia1 List of Generic Names and a11 must be held to be invalid.-R, G. Fennah.} INTRODUCTION The Macquartiinae, the third subfamily of<¡:achinidae in this series, is very much larger than the two subfamilies studi"d hitherto (Phasiinae, 1944, Proc. zool. Soco Lond. 114 ; 389-436, and Dexiinae, 1947, I.c. 116, 627-674). Most of the introductory remarks to the first paper of,the series also apply to this papero In its present scope the subfamily was first defined by :vJ:esnil

P.Z.S.L.-134 21 314 F. l. VA.~ ElVIDEN

(1939) under the name Larvaevorinae, but a similar group had previously been outlined in its main contents and in some of its characters by Villeneuve (1924, 1932) under the name Protachininae. The latter name is not derived from a " type genus " and is therefore not available under the "Rules", and the name Larvaevara :Vleigen 1800 appears likely to be suppressed. Actually, Larvaevara ~eigen 1800 does not belong to the present group, although most authors have accepted Hendel's (1908) identification of Larvaevara :Vleigen 1800 with Taehina :Vleigen 1803 and the alleged absence of Exorista ~Ieigen 1803 from the 1800 paper. In fact, the descriptions of Larvaevara and Exarista are practically identical (antennae with two segments, the "second" pris­ matic, etc.) whereas that of Taehina is very clifferent (antennae with three segments, the second cylindrical, the third subcircuJar). The ouJy species mentioned under Exarista is larvarum F., to which the description of the antennae of Larvaevara applies, and the generic name of 1800, like the specific name of 1803, refers to the typical parasitism of caterpillars by this species. Larvaev ara is thus synonymous with Exorista :Vleigen 1803, Eutachina B.B.1890, and Taehina )Ieigen 1838 and does not fati in the present subfamily. The oldest available name for an aggregate containing one of the genera now in this group appears to be ~acquartünae (" Macquartidae " Robineau-Desvoidy 1830, p. 203). In the genera composing this subfamily 113 species have been described before, but twenty of them have to be relegated to synonymy and one to varietal status. Seven genera and fifty-eight species are described as new. Literature and synonymy are not cited so far as they are found in Town­ send's " Manual ofMyiology " and Becker's "Kat. pal. Dipt.", or, so far as biolo­ gical literature is concerned, in Cuthbertson and Munro's "Sorne records of tachinid parasites and their hosts in southern Africa" (1941, Trans. Rhodesia sei. Ass. 38: 88-118). Species marked with an asterisk (*) have been incorporated from the literature, as specimens are not available. The types of the new species are in the British Museum (Natural History). The first two connate abdominal segments are counted as one, so that the last entirely free tergite is the fourth. Abbreviations Abbreviations used in describing the legs :--{l;, anterior; ad, antero-dorsal ; av, antero-ventral ; d, dorsal; p, posterior; pd, postero-dorsal; pv, postero­ ventral; v, ventral. In using these expressions one shouJd imagine the leg extended in a straight line at right angles to the longitudinal axis of the body. Other abbreviations aer, acrostichal bristles (hairs) ia, intra-alar bristles B.:YI., British ~useum ()fatural History) m, fourth vein 0, costal cel! (20, second costal cell) m-m, post&ior cross vein c, costa npl. notopleural setae de, dorsocentral bristles ph, posthumeral seta h, humeral cross-vein post, postsutural KEYS TÚ THE ETHIÚPB..N TACHINIDAE-MACQUARTIINAE 3]5 pra, prealar brjstle T 4+5, third vein prsc, prescutel1ar rs, radial sector prst, presutural r-m, small cross vein R margjnal cel! sa, supra-alar brjstle

R3", submarginal cell se, auxiliary vein R 5, first posterjor cel! Se, pterostigma (sub-costal cell) TI' first vein stpl, sternopleural bristles T 2+3 , second vein

DEFINITlüN üF THE SUBFMTILY J.1ACQU.ARTIINAE Tachinidae with the prosternum bare or the pv apical seta of the hind tibia well developed (usual!y both these charaeters present simultaneously), reclinate frontal setae absent. Normally 3 post ia, if only 2, the first is missing. J owls narrower than the combined length of the seeond and third antennal segments, or if not, with a wel!-developed occipital di!ation. Face without a keel between the antennae (except in Euthera). Arista not plumose (except in Thelairini). There can be no doubt that the division of the subfami!y into two main groups of tribes accorcling to the development of the basal dorsal excavation of the abdomen (Mesni!, 1939) or the relative position of the mouth margin and the vibrissae (van Emden, 1954) is a natural one. Similarly the classifi­ cation of the seeond of these groups containing the Echinomyiini and Leskiini is largely uncontroversial, the only problem, whieh is an interesting one, being posed by the -group. Mesnil, using the dorsal excavation as the decisive character, placed the Leskiini in what is here treated as the first group and the Aphriini (as Aphriina) in the second, mentioning that the Aphriine genus had the excavation of the first group. In studying the British fauna I brought the Leskiini into the second group, using the projecting mouth margin, because the type of the abdominal excavation is subject to a consider­ able number of exceptions, but left the Aphria group in Echinomyini. The present revision of the Ethiopian species has shown that it is desirable to go even further by uniting Aphriini and Leskiini in one tribe, the two groups being connected by several intermediate forms and in fact very closely related. Many more problems arise in the first group of trjbes, and the solution offered here is in many ways based more on expecliency than on scientific convictions. Nevertheless, it appears that Wagneriini and Campylochaetini are closely related, and the same applies to Thelairini and Minthoini. The position of Germariini and Nemoraeini close to the latter tribes can also be assumed to be correcto The definition and contents of the Macquartiini and Helocerini and the position of the Acemyini in the present subfamily remain wholly problematjc.

BIOLOGY OF THE ETHIOPIAN MACQUARTIINAE Little is known concerning the biology of Ethlopian species of this group of fiies. The followlng list of records is possibly incomplete, as several papers have not been available for examination. 21' 316 F. 1. VA..'f IDIDE~

Wagneria carbonaria Panz. is known as a parasite of Euxoa segetum Schiff. (Noctuidae) in S. Aii:ica and also of an unidentified moth larva; in Europe this species also parasitises Noctuidae such as Hoplodrina alsinea Brahm, Agrochom circellaris Hufn. and Lacanobia w-mtinum Hufn. Yemoraea capensis R.D. has been recorded from S. Africa from Heliothis armigera Hbn. and Diparopsis castanea Hmps. (Noctuidae), and Nemoraea fortuna Curr. from Dasychira goodi Holl. (Lymantrüdae). Themira nigripes F. is a parasite of the arctiid, Diacri. sia fmca Wllgrn. and Themira palliventris Curr. of Diacrisia maculosa Stóll. Dejeania bombymns F. attacks Heiiothis armigera Hbn. in S. Africa and D. wollastoni Aust. was obtained from Lepidoptera pupae collected on the Ruwen­ zari Expedition. Paratachina obliqua Lw. makes yet another parasite of Heliothis armigera in S. Africa, while Guphocera haemorrhoidalis Macq. is recorded from Agrotis segetum Schiff. and Laphygma exempta Walk., and hova Villen. from Leucania loreyi Dup. Linnaemya is the genus with most African records. The following have been found, all in S. Africa if not otherwise stated : Linnaemya affinis Corti recorded from Heliothis armigera Hbn. and Leueania leucosticha Hmps.; L. agilis Curro from Diparopsis castanea Hmps. and Heliothis armigera Hbn. and also from the latter in Tanganyika. Linnaemya alboseutellata Speis. from an unidentified caterpillar (probably Heliothis armigera Hbn.) and L. angulicornis Speis from Leucania loreyi and Leucania leucosticha Hmps. No host record for Fal!. has been found for Africa, but in Europe the species parasitises Hyloicus pinastri L. (Sphingidae), Saturnia pavonia L. (Saturniidae), Ag·rotis typsilon Rott. and other species of Noctuidae of the genera Laphygma, Peridroma, etc. Linnaemya conformis Curr. is recorded from Phytometra sp. (C{octuidae), and laxiceps Villen. from Diacrisia lutescens Walk. (Arctiidae) in S. Africa. L. longirostris Macq. in S. Africa and Uganda from Helopeltis armigera Hbn. and Phytometra limbata Guen., L. pareesetosa Vill. parasitises Earias insulana Boisd. (Westermanniidae) in S. Africa and L. obscurior is recorded from the same species in Tanganyika. Finally L. variegata Wied. has been recorded in S. Africa from the limacodid Goenobasis amoena Feld. Apart from these records the following general remarks can be made. The genus J1!1acquartia is normally found on coleopterous hosts (Chrysomelidae) but there is a doubtful record from Lepidoptera (Pyraustidae) ; the fiies are said to be larviparous or oviparous. Wagneria para'litises mainly noctuid caterpillar<;. The hosts of Gampylochaeta itself are mainly Geometridae; ohere are, however, records of other Campylochaetini from Sphingidae, Noto­ dontidae, Lymantrüdae and Arctiidae. Members of the Helocerini are the most varied in their choice of hosts; they attack principally Lepidoptera but there are records of other non-Ethiopian genera from Dermaptera, Diptera (Syrphidae) and a myriapod. Sorne at least are larviparous. The Nemoraeini seem to be parasites of Lepidoptera (Noctuidae, Sphingidae, Arctiidae, Georoe­ tridae) whereas the Acemyini, as far as is known, are exclusively parasites of Acrididae (Anacridium), though there is an isolated' record of a species froro a noctuid. Of the biology of the Germariini no records have been found. The Tbelairini attack caterpillars of larger Heterocera (Sphingidae, Xotodontidae, Lasiocampidae, Noctuidae and specially Arctiidae), and seem to be oviparous KEYS TQ THE ETHIOPlAN TACHI~IDAE-?r1ACQUA..RTITIAE 317 on the hosts. Minthoini are recorded as being oviparous, and parasitise Pyrali­ dae and ~octuidae in Europe. Leskiini, also oviparous, parasitise Lepidoptera, principally _\egeriidae but also Tortricidae, Pyralidae, Arctiidae, Noctuidae and a species has been recorded from a nymphalid. In the tribe Echinomyiui the Ernestiina para~itise motlLs of the following groups ~otodontidae, Geometridae, Noctuidae, Lymantriidae, Lasiocam­ pidae, Arctiidae, Sphingidae, Saturniidae, Thyatiridae, as well as nymphalid butterfiies ; they belong to Pantel's 4th group in which the fernales are ovi­ parous or larviparous. The Linnaernyina have as hosts species of Noctuidae, Arctiidae, Aegeriidae, Sphingidae and Lymantriidae and the Echinomyina, Sphingidae, ~octuidae and Lymantriidae. In Europe Peletieria parasitises Papilio, Agrotis, Arctia and Mjmantria, and Echinomya Arctiidae, ~octuidae, Sphingidae, Lymantriidae and Lasiocampidae : both belong to the 4th group of Pante!. In general the :YIacquartiinae appear to be parasites of Lepidoptera, mainly of large Heterocera. Of the Ethiopian groups the only exceptions are A'Iacquar­ tia and the acernyin Ceracia. The affinities of this tribe are still obscure and its members show features of both the (Phorocerinae sensu Mesnil) and the Phasiinae. Sorne members of the Helocerini, at least, seem to be relatively polyphagous, while others seem to have found hosts among arthro­ pods outside the Insecta. The economic importance of the group may be great, as among their hosts figure most noxious Lepidoptera. of economic importance, ineluding eutworms and the bollworms (Diparopsis castanea Hmps. and Earias insulana Boisd.), both among the most important pests of cotton in Africa.

K ey to the Tribes 01 Ethiopian Macquartiinae

1 (l8). )louth-rnargin not projecting between vibrissae (except in Euthera where it projects, but much less than middle of clypeus). Posterior surface of hind coxa. bare. 2 (17). Fore tarsi not rnarkedly compressed laverally (in Allo­ thelaira slightly compressed. and dilated towards apices of the segments)_ Clypeus wi1;hout a median keel. 3 (10). Occiput with numerous black setulae behind upper part of occipital row (except in a few Wagneria). 4 (9). Scutellum with 2-3 strong margina.ls, tha distal ones rather clase together and decussate or diverging, between them no pair of smaller setae. Integument dark. 5 (6). Basal dorsal excavation of abdomen often not reaching hind margino Basal half of scutellum without erect discal setae (but sometimes ~he decumbent preapical setae shift.ed to middle), apical scutellars decussate, only 1-2 further pairs of marginal scutellars presento Facial ridges and parafacialia (in Ethiopian species) without stroog erec~ seta.e 00 iotermed.iate pan (but sometimes with small seta.e). Ocellars proclinatd or indistinct. r~H setulose only on and oeRr basal nade, bend of m without a.n