(Neuroptera: Myrmeleontidae). Introduction and Genus Bankisus Navas

J. ent. Soc. sth. Afr. 1985 Vol. 48, No. r,pf. 189-212

The ant-lions of southern Africa (Neuroptera: Myrmeleontidae). Introduction and genus Bankisus Navas

M. W. MANSELL

Sational Collection of Insects, Plant Protection Research Institute, Private Bag X134, Pretoria, 0001.

This is the first in a series of papers revising the systematics of the southern African Myrmeleontidae (Neuroptera). It introduces the series, reviews litera- ture on Myrmeleontidae from the subregion, discusses zoogeography, biology and higher classification, provides a preliminary key to local genera and revises the genus Bankisus Navis. Bankisus comprises five species, B. oculatus Navis, B. carinifns (Esben-Petersen), B. elegantulus (Esben-Petersen), B. trisuttatus Navas and B. maculosus Holzel. The first two are recorded from southern Africa and are redescribed; the last three are known only by their holotypes, elegantulur and triguttatus from Zaire and maculosus from Oman. No new species are introduced and B. kristenseni (Esben-Petersen) is synonomised with B. oculatus.

INTRODUCTION Southern Africa has a rich fauna of Myrmeleontidae comprising about 140 species in 40 genera. It is the largest of 12 families of Neuroptera inhabiting the subregion and also includes the most striking representatives of the Order. The family has hitherto not been comprehensively studied, and accounts of local myrmeleontids are limited mainly to old scattered taxonomic papers. Existing descriptions arr generally superficial, relying on wing venation and external morphology; none provide details of genitalia or immature stages, many are based on unique specimens and illustrations are meagre. This has led to inadequately defined taxa, complicating the intrrpretation ofgenera, species and their relationships, and resulting in numerous synonomies. A study of the family is now being undertaken, aimed at providing a revision of the southern African species, and this is the first in a series of papers reporting on the work. It provides an introduction to the series by way of a litrrature review, a dis- cussion of zoogeography, biology and higher classification and a preliminary key to the local genera, followed by a revision of the genus Bankisus Navis. Each subsequent con- tribution will treat one or more genera, local taxa will be redescribed to incorporate new data, to ensure uniformity and to provide a convenient reference source. Many southern African genera and species are widespread and were originally described from arras to the north of the subregion. So, although dealing pri- ,J. enl. Soc. sth. flfr. Val. 48, No. 1, 1.98.5 marily with thr fauna south of the Zambezi and Curiene rivers (approximately 16"S to 18" S), the revision will also include relevant information on ant-lions from elsewhere in the Afrotropical region. Terminology follows that of Aspiick et al. (1980), who also provides a diagno- sis of the family, including morphological details of wing venation and genitalia. Trch- niques used in the study are similar to those described by them. Venational terminology and abbreviations are indicated in Figs I to 9, and that for genital structures ill Figs 12 to 29. Under rnaterial examined, irlformation in brackcts is supplementary data not reflected on the swecimen labels. Thr revision is based on the author's collrction of approximately 5000 specimens, in the National Collection of Insects, Pretoria, and on matrrial in thc following institutions, which will br referred to throughout the series by the accompanying abbreviations. BMNH-British Mlisrum (Natural History), London IRSN -Institute Koval des Scierices Naturcllrs. Brussels LS -Lilinaean Society, London MG -Musi?un~ dlHistoire Naturellr, Gcncva MP -Museum National d'Histoire Naturelle, Paris MRAC-Musee Royal dr I'Afrique Crritrale, Tervuren MZR -Muse0 dr Zoologia, Barcrlo~la NCI -National C:ollection of Insrcts, Pretoria NMZ -National Museum of Zimbabwe, Rulawayo SAM -South African Musrum, Cape Town UMO -University Museum, Hope Department of Entomology, Oxtbrd TM -'Transvaal Museum, Pretoria ZMC -Zoological Museum, Copenhagen University, Copenhagen ZML -Zoological Museum, I.und University, Lu~ld.

PREVIOUS \VORK ON SOUTHERN AFRICAN MYRMELEONTIDAE Publicatioris prrtinerit to the local taxa rover a 218 year period from 1758 to 1976, and all except three of these accounts are taxonomic. Apart from the studies of Pkringuey (1910, 1911) and Yuuthrd & Morari (Iy69a, 1969b, I 96yc), all research on Afrotropical Myrmeleontidae has been corlductrd t)y taxonomists outside Africa. The Lirst account of a southern African myrmeleontid appcarrd in I 758 when Linnaeus describrd I1emerohiu.c .\peciosum (Palpares specio~w).111 I 767 1,innaeus erected the genus Myrmeleon, an cxtant name, but considerably restricted compared to its original sense. Thunberg (I784), Olivier (1809, 18r r ) and Burmeister ( I 839) followed Linnarus and between them added eight species to the documerited fauna. Rambur (1842) described three new species from southern Akica arrd established the genus Palpares. Walker (18y3, 1860) described r I species from the subregion, but took the rrtrogressive step of igrloring all estal-dished genera except ~gyrmeleon.'l'he German entomologist and bibliographer, Herman Hagen, correlated intbrmation on Myrme- leorltidae in two public.ations (1860, 1866), thr latter being of hndamental importance in any study of Nruroptera. In threr other papers (Hagen 1853, 1862, 1887), he described and discussrd several southern African species arid erected the genera Tomatures and Pamexis. Gerstaec-ker (I863, 1885, 1888, I 894) workrd exterlsivrly on rast and

J. enl. Soc. sth. Afr. Vol. 48, No. 1, 1985

ZOOGEOGRAPHY

Aspock et al. (1980) estimate the existence of approximately 2000 species, dis- tributed on all the continents and especially in the arid regions of Africa, Asia and, to a lesser extent, Australia and the Americas. Myrmeleontidae also occur on many islands, for example, Hawaii (Zimmermann 1957), Galapagos (Stange 1969), Europa (Fraser 1951) and Mauritius. Some species extend as far north as British Columbia and Ontario in Canada, and Finland in Europe (Wheeler 1g3o), and as far south as New Zealand (Tillyard I 926). In southern Africa, Myrmeleontidae reveal two main distributional trends. There is a rich fauna, characterized by a high degree of endemism, inhabiting the more arid western parts of the subregion, and an easterly fauna, influenced by elements extending along the tropical corridor into south east Africa from east and central Africa. A few species are very restricted in distribution, especially in the south western Cape, whilst several are cosmopolitan. Approximately 34 species (25%) are known only from the western areas of the subregion, about 42 (31%) occur in the eastern parts, some 15 (II%) are restricted in distribution whilst the remaining 33% are either cosmopolitan or too inadequately known to categorize at present.

IMMATURE STAGES AND BIOLOGY Larvae possess important morphological and biological attributes which can be used in determing genera and species, in supra-generic classification and in resolv- ing phylogenetic questions. Stange (I97ob) listed the early publications dealing with myrmeleontid larvae, and recent papers include those by him (Stange 197ob, 1980). Steffan (1964, 1968, 1971, 1975) and Willmann (1977). Apart from these few contributions, which deal with non-Afrotropical forms, larvae have been largely ignored, as many species are difficult to find and rear. Larvae of Afrotropical species have hitherto not been described, and the vast majority are still unknown. Of the estimated 140 species in southern Africa, the larvae of 22 have been correlated with the adults. Of these, 13 species belong to Myrmeleon, Cueta and Hagenomyia, which are the only three southern African genera whose larvae are known to construct pits. Cymothales larvae in- habit tree-holes, those of Tricholeon and some Neuraleon species occur in caves or under rock overhangs, whilst the majority are probably freeliving in sand. Significant morphological characters include: shape of mandibles and tooth number, form of the eyes, shape of the thoracic spiracle, development of thoracic and abdominal scoli, form of the fossorial structures on sternite 8 and the structure of the setae. Tooth number in the southern African species varies from three to six, with three teeth the most prevalent. In most species the eyes are borne on prominent tubercles. but in the pit-building species these have become reduced, and the eyes are often sessile on the head. The prothoracic spiracle is sometimes situated on a protuberance of the pleurites. Lateral processes of the thorax and abdomen are well developed in some cave-dwelling forms (eg. Tricholeon), whilst the sand-inhabiting larvae are usually devoid of such processes. The fossorial setae of most species are stout bristles, but in Cueta the bristles are fused into small toothed plates, whilst in Palpares, Auia and Fadrina therr are paired tringular digging appendages. One of the objectives of this study is to discover and correlate larvae of the southern African species and to use the biological and morphological features in the systematic revision of the family. Mansell: ant lions of the genus Bankisus

REVIEW OF HIGHER CLASSIFICATION The supra-generic classification of the Myrmeleontidae is largely urlresolved (Stange 1g7oa, Aspock et al. 1980). Several authors, whose main contributions are reviewed below, have attempted to arrange the family into subfamilies, tribes and sub- tribes, but overall consensus on these categories has not been achieved. Tribal classification on a world basis is particularly controversial and complex because of insufficient knowledge of many genera, especially the immature stages. In all, some 7 subfamilies and 33 tribes have been proposed, of which about 4 subfamilies and 23 tribes are still taxonomically available. One of the earliest classifications was by Banks (18gg),who distinguished two groups, Myrmeleoni and Dendroleoni, in the North American fauna. Later, in his study on the African Myrmeleontidae (Ranks 191I), he elevated these to subfamilies Myrmeleoninae and Dendroleoninae, each comprising two tribes, Palparini and Myrmeleonini, and Dendroleoni and Nemoleonini respectively. Navis (19I 2b, I 9 12d, 19I 3d, I 914e, I g I 4f, I 926~)added eight more tribes, Gymnocnemini, Acanthaclisisini, Creagrini, Megistopini, Neuroleini, Formicaleonini, Dimarini, Porrerini and Pignatelli- ni, but did not consider subfamilies. Tillyard (1916) recognized Bank's two subfamilies and added two more tribes, Distoleonini and Protoplectrini, in studies on the Austra- lian fauna. Esben-Petersen (1918) split the Myrmeleontidae into two groups, Archaemyr- meleonidae and Neomyrmeleonidae, the former containing the Palparini and related forms, the latter comprising the Myrmeleontinae and Dendroleontinae, with four and six tribes respectively. He erected three new tribes, Macronemurini, 1,opezini and Myr- mecaelurini. Esben-Petersen considered the Archaemyrmeleonidae to be the more primitive ant-lions, a view generally supported by later workers (eg. Holzel 1972:74), but not shared by the present author. The Palparini are regarded as an advanced group for several reasons, the most important beirig distribution and larval morphology. The subfamily is not represented in Australia or South America, indicating that the group probably evolved after Africa, Australia and South America had separated. It is unlikely that this large and successful Afrotropical subfamily could have disappeared without trace from the latter two continents. In the Palpaririae, larval tooth number varies between three and six, and an increased number of teeth can be considered apo- morphic relative to the other Neuroptera. The more primitive forms should be sought amongst the Dendroleontini and not Palparini as generally thought. Banks (1927) raised the Palparini and Macronemurini to subfamily status, bringing the number to four, and added the tribes Brachynemurini and Glenurini for the Neartic fauna. In 1938 he mentioned the tribe Gamini, and in 1943 elevated the Acanthaclisisini to subfamily level. In 1954, Markl classified the myrmeleontid genera of the world into 23 tribes, adding 10 new tribes in the process. He synonymized Neuroleini with Formicaleonini, 1,opezini with Myrmecaelurini and Creoleonini was established as a replacement name for Creagrini. Markl apprared to overlook the existing tribes Macronemurini, Distoleo- nini Gamini and Pignatellini, and erected the following new tribes: Psrudimarini, Pal- paridiini, Echthromyrmicini, Maulini, Acanthoplectrini, Gepini, Nesoleonini, Obini, Nyutini, and Dimarellini. He did not attempt to arrange the tribes into subfamilies. Stange ( I 96 I) mentioned three subfamilies, Acanthaclisinae, Macronemuri- nae and Dendroleontinae, from the New World, but did not deal with the Myrmeleon- tinae or Palparinac. In 1967 and agairl in 197oa, Stange referred to four of the subfami- 194

Willman (1977) mentioned two subfamilies, Palparinae and Myrmeleontinae, including five tribes in the latter. He synonymized Isoleonini Holzel with Gepini Markl, stating that the latter name had priority. Aspock et al. 1980 divided the family into two sister groups, Palparinae and Myrmeleontinae, but did not undertake a tribal classification of the European Myrmeleontidae. Most recently, New (1982) reappraised the status of the family Stilbopterygidae, referring the genus Albardia to the Ascalaphi- dae, and Stilbopteryx and Aeropteryx to the Myrmeleontidae, where they constitute the subfamily Stilbopteryginae. Whilst some agreement on subfamilies seems to be emerging, three (Palpari- nae, Acanthaclisinae, Myrmeleontinae) according to Stange (1976) and two (Palpari- nae and Myrmeleontinae) recognised by Aspock et al. (198o), plus Stilbopteryginae (New 1982), the tribes are still subjectively defined.

In the present study, southern African genera will be arranged in existing tribes as far as possible, where these are well defined. The situation will be reviewed again when more information is available. 'I'he genus Bankisus, now being reviseti, to- gether with Cymothales and Tricholeon belongs to the Dendroleontini (Stange, 1976). Other genera could be accommodated as follows: Pa1pare~-,Pamexis, Tomatares, Crambo- morphus, Golafrus, and Lachlathetes in Palparini; Palparidius in Dimarini; Avia, Centroclisis, Syngenes and Fadrina in Acanthaclisisini; Mynneleon and Hagenomyia in Myrmeleontini; Banyutu.r, Distoleon, Creoleon, Brachyplectron, Macronemurus, Nannoleon, Nemoleon and Neuro- leon in Nemoleontini; Cueta, Furgella and Nesoleon in Gepini; Capophanes, Obus and Mochus in Obini; Maula and Isonemurus in Maulini, whilst Exaetoleon is uncertain. These tribes have been defined by Markl (I954).

Figs 1-9. Key figures. I. Forewing of Palpares sp.; 2. Forewing of Cueta sp.; 3. Forewing of Crambo- morphus sp. showing double costal series; 4. Forewing of Palpares sp. with single costal series; 5. Forewing of Palparidius sp.; 6. Hind wing of Palpares sp.; 7. Hind wing of Tri- choleon sp.; 8. Hind wing of Myrmeleon sp.; g. Hind wing of Creoleon sp.; Abbreviations:

rA - first anal vein; C - costa; Cua - antrrior cubitus; Cup - posterior cubitus; Cua~- upper branch of cubital fork; Cuar - lower branch of cubital fork; Ma - anterior median; Mpr - upper branch of posterior median; Mpz - lower branch of posterior

median (oblique vein); Ps - presectorial veins; R - radius; Rs - radial sector; Rv - recurrent vein; Sc - subcosta. Mansell: ant lions of the genus Bankisus J. ent. Soc. sth. Afr. Vol. 48, h70. 1, 198.5

PROVISIONAL KEY TO THE GENERA OF SOUTHERN AFRICAN MYRMELEONTIDAE

I Veins <:up and rA distinct, not fused (Fig. I); large forms with heavily markrd, usually broadwings ...... 2 Veiris Cup and 1.4fusrd closc to wing has? (Fig. 2); smaller forrns with narrower, often unmarkedwings ...... 8 2 Costal arca of forcwirlgs with'somc cells douhle (Fig. 3); wings vcry narrow, with sinuatc hindmargins ...... Crambomorphus MacLachlan Costal arca of forewings with singlc row of cells, except occasionally near pterostigma (Fig.4) ...... 3 3 Wings narrow, each with a prqjrcting lobe on hindmargin; hirid lcgs long, slender, at least one half as long as ahdomcn ...... Golafrus Navis Wings broad, hindmargins not lobed; llirld legs shortcr than oric halfthe abdomen lcrigth .... 4 4 Antennae short, clubbed, distinctly shortrr than combined Icngth of lhrc tibia and tarsus; smallisti forms with bright yellow, heavily marked wings ...... ,i Antcnna as long or distinctly longer than combincd lerigth of fore tibia and tarsus; wings may be yellowish, but not bright yellow ...... 6 5 Basal half of hind wings yellow, unmarked ...... Tomatares Hagen Basal half of hind wings with dark brown markings ...... Pamexis Hagen 6 In hind wings, Cu,, slightly curved at junction with posterior fork of M,,, (Fig. 5); male cctoprocts very long, at least I 2 mm in length ...... Palparidius Ptringucy Hind wings with Cu,, bent sharply at junction with posterior fork of M,, to form recurrent vein (Fig. 6); ectoprocts of male short, not approaching I 2 mm ...... 7 7 Wings vcry broad, posterior margins sinuate; antennae long, cylindrical without distinct club ...... Lachlathetes Navas Posterior margiris ofwings not distinctly sinuate; antennae clavate ...... Palpares Ilambur 8 One or two prrsectorial veins in hind wings (Fig. 7) ...... 18 Thrcr or more cesectorial veins in hind wings (Fig. 8) ...... (1 y Labial palps very long, at least one half as long as aritcrlria ...... Maula Navis Labial palps normal, not one halfas long as ariterina ...... ro ro Tibial spurs stout, curved through go0 or right angled, longcr than Iirst tarsomcre (which is short) in all three pairs of legs; combined Icngth of tarsomcrcs I to 4 less th;m that of 5. Usually robust dull grey hairy sprcirs ...... I I Tibial spurs slender, straight or slightly curved, shorter than first tarsomere, at least in hind legs legs; combined length of tarsomeres I to 4 usually grrater than that oTg ...... 14 I I Costal veins of forewings with double row of cclls or costal vcins mostly forked (see Fig. 3) 12 Costal area with single row ol'crlls, except near ptcrostignia ...... 13 12 Costal veins forked or formirig irregular cells; tibia1 spurs cvenly curved .... Syngenes Kolhe Costal cells in two even rows; spurs strongly right angled ...... Fadrina NavLs 13 Tibial spurs cvcrily curved ...... Avia Navis Tibial spurs sharply right angled ...... Centroclisis Navas 14 In forewings, Rs arises slightly before or approximately on the same level as Cu fork (Cuar andCua2) ...... HagenomyiaBanks In forewings Rs arises well bryond Cu fork ...... 15 15 First tarsomere in fore- and middlr lcgs short only slightly longcr than srcond; tibial spurs curved longrr than first and second tarsomercs on forr- and middle legs ..... Furgella Markl First tarsomere long (in all three pairs of Icgs), about twice the lrngth of the second tar- somerc, spurs straight equal or shorter than first tarsomere in all legs ...... 16 16 Yrllow species, with ycllow and brow11 striped thorax; cisht to twelve prcscctorial veins in hindwings ...... CuetaNavis Not riniformly ycllow or thorax not striped with brown arld yellow; usually eight or fewer prescctorial veins in hand wings ...... I 7 Mansell: ant lions of the genus Bankisus I97

17 Smallish species with roundecl wingtips, wirigs heavily marked or blotched with brown or distinct black mark over ptcrostigma; seven or eight presectorial veins in hind wings ...... Nesoleon Uariks Wingtips usually acute, wings mostly hyalirie, generally three to five (occasionally three to eight) prescctorial veins in hand wings ...... Myrmeleon Linnarus 18 Cubital forks in forewirlgs are parallel for most ofthcir length (Fig. 9) ...... 19 Cubital forks in forewings diverge (see Fig. 8) ...... 20 19 Median forks in hind wings diverge; tibial spurs well developed, longer than first tarsonlrre ...... Cre~leon~illyard Median forks in hind wings parallel; tibial spurs weakly developed, shortcr than first tarsomere or absent ...... Obus Navas 20 Tibialspursabsent ...... 21 'Tibialspurspresent ...... 23 21 Costal area offorewings with double row ofcells (see Fig. 3) ...... Capophanes Banks Costal area of forewings with single row of cells ...... 22 22 Radial sector arises before cubital fork in forcwings, wings hyaline with brown marks; Vertex raised ...... Bankisus NavLs Rs arises beyond (:u Sork in forewings, wings pink with brown spots; vertex rounded ...... ExaetoleonKimmins 23 Hind wings strongly falcate with acute apex; both pairs of wings prominently marked ...... CymothalesGerstaecker Hind wings not falcate, if slightly falcate then not marked ...... 24 24 Labial palps Long, at least one half as long as antenna ...... Isonemurus Esben-Pctersen Labial palps normal, not one half as lorig as antenna ...... 25 25 In forewings Ks arises well before Cu fork; males with pilulae ...... 26 In forewings Rs arises well beyond Cu fork or at approximately the Sam? level as the Cu fork; malrs lack pilulac ...... 27 26 Legs long slender, hind legs reaching beyond fourth abdominal tergite ...... Tricholeon Esben-Petersen Legs short, hind legs not extending beyond third abdominal tergite ...... NannoleonEsbcn-Petersen 27 Tarsi longcr than tibiae in forr- and middle legs; with long narrow wings and long abdomen ...... NemoleonNavas Tarsi shorter or same length as tibiae ...... 28 28 Tibial spurs less than or equal to the length of the first tarsomere in hind legs ...... 29 Tibial spurs lo~lgerthan the first tarsomerr in hind legs ...... 29 Tibial spurs as long or longer tharl half the first tarsomere in fore- and middle legs ...... NeuroleonNavis .libial . spurs usually shortcr than halfof first tarsomere in all legs ...... Brachyplectron Esben-Petersen 30 Antennar long cylindcrical, longer than thorax, not distinctly clavatc; rorewing shortcr thanhindwing ...... BanyutusNavas Antennae distinctly clavatc, shorter or equal to length of thorax; Ibre- and hind wings of equallength ...... 31 31 Spurs on Sore tibiae longer or equal to the combined Icngth of tarsomeres one to Sour ...... DistoleonBanks Spurs less than the combined lengtt~of tarsomerrs one to four ...... Macronemurus Costa

The status of the following genera is uncertain, some are undoubtedly synonynls, so they have been omitted li-om the key: Campestretus Navis, Bordus Navis, Ladrus Navis, Mochus Navas, Gandulus Navis, Suca Navis, Mironus Navis, Nosa Navks and Negretus Navis. .J. ent. Soc. sth. Afr. Vol. 48, No. 1, 1985

Genus Bankisus Naviis Bankisus Navis, rgI2a: 45; 1g26a: 105; Banks, 1938: 126; Markl, 1954: 221; Stange, 1976: 262, 287; Holzel, 1983: 211. Navasius Esben-Petersen, 1936: 202; Markl, 1954: 22 1. Type-species: Bankisus oculatus Navis, by original designation.

REDESCRIPTION.Small species, characterized by hyaline wings with brown markings, long slender legs and lack of tibial spurs.

Head with square raised vertex. Maxillary palps five-segmented, labial palps-. three-segmented, sense organ on third palpomere with rounded afperture. Prothorax longer than broad with long setae. Wings hyaline, marked with brown, pterostigma of forewings prominent, maroon-coloured, pale in hind wings; forewings broader than hind wings. Forewings with 3 presectorial veins, Rs arises close to wing-base well before Cu fork, Mpn meets Cua~just beyond Cu fork, Cu forks widely divergent, rA fused with Cup, banksian lines discernible; hind wings with I crossvein between R and M before Rs, Rs arises before M fork, IA fused with Cup, banksian lines discernible, pilulae present in males. Legs long, slender, lacking tibial spurs; pre- tarsal claws long, straight, folding back against bristle-pad on fifth tarsomere, first tarsomere long, fifth slightly shorter, intermediate three shorter still; femoral sense hairs absent. Tibiae longeru than tarsi in all lets." Abdomen of males with tergite g divided, ectoprocts short; gonarcus, parameres distinct, parameres sclerotized, free from one another. Pleuritocavae absent. Fe- male with tergite g divided, ectoprocts short, rounded; two or three pairs of gonapophyses present, gonapophyseal plate not developed; fossorial setae well developed. Larvae: unknown The genus comprises five species, two of which, B. oculatus Navas and B. cari- nijions (Esben-Petersen), are recorded from southern Africa. The other three, B. elegantulus (Esben-Petersen), B. triguttatus Navis and B. maculosus Holzel, are known only by their respective holotypes, elegantulus and triguttatus from Zaire and maculosus from Oman. Navasius kristenseni (Esben-Petersen) originally included in Gymnocnemia Schnei- der, by Esben-Petersen (1915) is a synonym of B. oculatus NavBs. Markl (1954) synonymized Navasius Esben-Petersen with Bankisus NavLs and Stange (1976) transferred the species in Navasius to Bankisus. Bankisus is a widespread genus, ranging from southern Africa to Yemen and Oman on the Arabian peninsula.

Bankisus oculatus NavBs, Figs 10, I 2-20, 30. Bankisus oculatus Navas, 1912a: 46; 1g12b: 96; Banks, 1913a: I 54; Markl, 1954: 198; Stange, 1976: 288; Holzel, 1983: 211. Gymnocnemia kristenseni Esben-Petersen, 1915: 1 79; 1936: 206. Navasius kristrnscni (Esben-Petersen); Esben-Petersen, 1936: 206. Bankisus kristenseni (Esbcn-Petersen);Stange, 1976: 288. syn. nov.

REDESCRIPTION:based on the female holotype of B. oculatus NavBs, and other material examined. Sexes alike, characterized by long slender antennae and legs, long maculated wings, forewings broader than hind wings and whitish thoracic tergites. Mansell: ant lions of the genus Bankisus I99

Fig. 10. Bankisus oculatus Navis.

Size: mean measurements in mm for 38 specimens, ranges in brackets: length of body (excluding head) I7,4 (14,o-20,o); forewing 20,9 (17,o-23,o); hind wing 19,6 (16,o-22,o); antenna 5,4 (4,q-6,o). Head: frons, vertex, occiput dark shiny brown, darker towards toruli, inter- antenna1 mark shiny black, usually extending onto clypeus as a short vertical stripe; clypeus and labrum uniformly pale yellowish-brown or reddish. Maxillary palps uniformly pale yellowish, terminal segment spindle-shaped with acute apex, distal region of stripe with brown annulation, articulation of cardo and stripe brown; labial palps pale yellowish, terminal segment long, spindle-shaped; submentum black, bearing four long black setae. Antennae long, about one third the forewing length, slightly thickened apically; flagellum comprising 30 to 35 segments, pale yellowish white with two brown stripes on basal one third, covered with short black setae; scape and pedicel pale, each with a dark mark on inner surface. Pronotum with transverse furrow anteriorly; pale midline flanked by two broad blackish marks, with four dark marks on anterior raised portion, lateral margins brown medially, pale anteriorly; prominent long white setae project over the head from anterior margin of prothorax, with shorter white sctar on lateral margins and dorsal surface, dorsal surface also with some long black setae, all alveoli on pronotum black, imparting a stippled appearance. Cervical sclerites black, sternites pale. Mesothorax: mesoprescutum black anteriorly, pale posteriorly with long white setae set in black alveoli (remainder of pterothoracic tergites pale creamy-white except for a short narrow black stripe on mesopostnotum and metaprescutum); four prominent clumps of erect white setae occur on mesonotum, with mesoscutellum covcred in fine whit? setae. J. ent. Soc. slh. Afr. Vol. 48, No. I, 198.5

Metathorax: metaprescutum with long soft white setae, metanotum devoid of setar, metascutum with very line sparsr setae. Pterothoracic pleurites with a broad black band situated above coxae, but not extending to wing bases; long white setae present on pleurites. Sternites pale yellowish-white. Legs: forelegs; coxae brown on outer surfacrs, otherwise pale, trochanters with prominent black spot, femora pale with four narrow brown stripes and long white and shorter black setar interspersed with recumbent black setae, tibiae very long with two thin brown stripes and sparsr long white setae amongst recumbent black setar, tarsi with first tarsomere only slightly longer than other four, pale with short black setae; middle legs similar to forelegs, but slightly shorter, first tarsomcre distinctly longer than apical four; hind legs with coxae and trochanters similar to forelegs, femora completely brown dorsally, pale ventrally with recuniberlt black setar but lacking white sctae, tibiae pale with short black setae, first tarsomere distinctly longer than apical four. Wings: pointed apically, not falcate marked as depicted in Fig. 10. Rs with about 10 branches in forewing, 8 in hind wing, other venation as in genus. Abdomen: shorter than hind wing; pale, banded with black, sparsely covered with short hyaline setae and black setar on external terminalia. Male (Figs 12-17, 30) with ventral margins of tergite g just reaching sternite 9 which is broad and convex with rounded posterior margin; ectoprocts almost rectangular, posterioventral margin not projecting. Gonarcus arcuate with fairly broad proximal regions; mediuncus not discernible apparently membranous; parameres sclerotized, hollow, boat-shaped, widest in proximal region with vertical projections and fine longitudinal striations ventrally (Fig. 30), gonosetae present; gonarcus and parameres enveloped in eversible membranous sac; hypandrium difficult to discern, apparently membranous. Female

11 Fig. I I. Banki,cus ca~inijons(Esben-Petersen). Mansell: ant lions of the genus Bankisus

Figs I 2-20. Bankisuc oculatus NavLs. I 2-1 7 malr grnitalia. r 2. Apex of abdomen, lateral. I 3. Apex oS abdomen, ventral. 14. Gonarcus and parameres, lateral. 15. Gonarcus and parameres, dorsal. 16. Gonarcus and paramcrcs, vcntral. 17. Gonarcus and parameres, caudal. 18-20 Scmale genitalia. 18.Apex of abdomen, lateral. 19. Apex of abdomen,

vcntral. 20. Sperrnatheca. Abbreviations: Epr - ectoproct; (;a - anterior gonapo-

physes; GI - latcral gonapophyscs; (;p - posterior gonapophyses; Gs - gonarcus; Pa -

paramerr; Prg - prcqcnital platc; I3 - t(.raitc 8; (1 - tcrgitc 9; ix - sternitc g. 202 J. ent. Soc. sth. Afr. Vol. 48, No. 1, 1985

(Figs 18-2o)with tergite 8 tapering to acute apices ventrally; each half of tergite 9 elongately oval; ectoprocts rounded with long erect black setae, merging with short stout upwardly curved fossorial setae. Pregenital plate small, tooth-like, lying medially at posterior margin of sternite 7. PosteEior go~apophysessmall, oval, with dense straight setae; anterior gonapophyses digitiform, articulating with apices of tergite 8, bearing a dorsal brush-like pad of long erect setae with sparse long setae ventrally; lateral gonapophyses separated from each other by narrow membranous area, bearing short stout fossorial setae posteriorly and shortish bristles ventrally above genital opening; a plate-like structure ventral to genital opening bearing short straight stout fossorial setae. Spermatheca (Fig. 20) sclerotized, darkly pigmented, C-shaped, broad at base, tapering slightly. The holotype of Navasius kristenseni (Esben-Petersen) is a small faded specimen, with slightly narrower wings than the typical forms of B. oculatus. Apart from these superficial differences, there are no features to distinguish it from B. oculatus. A comparison of the male genitalia of N. kristenseni with those of other specimens of B. oculatus showed no differences, so it was concluded that the two species are the same, with the latter having priority. This conclusion is further supported by a specimen identified by Esben-Petersen (in SAM) as Gymnocnemia kristenseni. It is a typical B. oculatus, with broad wings, but Esben-Petersen did not consider it different from his G. kristenseni, so identified it as this species. A female specimen from Yemen (in BMNH) is also identified as B. oculatus as no differences could be found to distinguish it, indicating that B. oculatus probably has a wide distribution in the Afrotropical Region. Specimens have been captured at light, and by sweeping dry twigs hanging from trees. At present nothing is known about the biology. Distribution: widespread in the Afrotropical Region; recorded from South Africa, Mocambique, Zimbabwe, Malawi, Tanzania, Ethiopia and Yemen.

MATERIALEXAMINED. Holotype? Bankisus oculatus NavAs, holotype 6 Navasiur kristenseni (Esben-Petersen), I I 6 and 26 P specimens. SOUTH AFRICA. Transuaal: I 9, National Collection of Insects accession number AcNE 258, Shirombe Pan (22.44s 31.24E), 1g.xii.1970, H & A. Braack, (NCI); I 9, AcNE 290, Punda Maria (22.41S 31.ozE), 27.iii.1973, A. Braack, (NCI); I 6 I 9,AcNe 226, Punda Maria, 22.41s 31.ozE, 25.xii.1983, M. W. Mansell, (NCI); I 6, AcNE 204, Skukuza, 24.593 31.55E, 9.i. 1984, M. W. Mansell, (NCI); all the above localities in the Kruger National Park; I 6 I 9,Pretoria North (25.40s 28. I IE), xi. 1955, G. van Son, (TM); I 9,Pre- toria North (25.40s 28.1 IE), ~o.xi.1g61,G. van Son, (TM); I 9,Percy Fyfe Nature Reserve, Potgietersrus District (24.03s zg.ogE), 18-zo.i.1971, R. Jones, (TM); I 9, AcNE 260, Nylsvley Nature Reserve, 24.393 28.42E, 10-11.xii.1979, C. D. Eardley, (NCI); r P, Rustenburg Nature Reserve (25.40s z7.12E), 3-6.xi.1975, Potgieter & Scoble, (TM); I 6, AcNE 225, Rustenburg Nature Reserve, 25.40s 27.12E, 7.xii.1983, M. W. Mansell, (NCI); I 6, AcNE 185, Soutpan, Prrtoria District, 25.24s 28.06E, 15.xi.1983, M. W. Mansell, (NCI); I ?, AcNE 340, Wylliespoort (22.58s 29.57E), 28.xi.1978, L. Minter, (NCI); Natal: 2 0, M'fongosi (28.40s 30.5oE), ii.1914, W. E. Jones, (SAM): MOCAMBIQUE: I 9,Vallet du Pungout, GuengCre, 1906, G. Vasse, (MP), (also recorded by NavAs, 1912b: 96): ZIMBABWE: holotype ? of B. oculatus, Mashonoland, Mazoe, 5000 ft, (17.30s 31.o3E), 26.xii.1905, G. A. K. Marshall, (BMNH); I 8,Arcturus, Salisbury (Harare) (17.438 31.05E), 1917,Dr Melle, (SAM); Mansell: ant lions of the genus Bankisus 203

I 9,Cross Koppie, Umtali (Mutare) (18.578 32.42E), xi.1963, Nat. Mus. 7. Rhodesia, (NMZ); I 9, Khami, Matebeleland (20.20s 28.30E), i. 1960, Nat. Mus. S. Rhodesia, (NMZ); I 6, Khami, Matebeleland (20.2oS 28.3oE), xi.1960, Nat. Mus. S. Rhodesia, (NMZ); 2 9,Turk Mine, Matebeleland (19.39s 28.54E), i.1961, Nat. Mus. S. Rhode- sia, (NMZ); I 9, Bulawayo, Hillside (20. IOS 28.43E), 26.k 197I, Mrs Donnelly, (NMZ); I 9,Bulawayo (20.10s 28.43E), xii.1970, Mrs Robinson, (NMZ); I 6, Bula- wayo, Matsheamhope (20. I~S28.43E), 17.i.1979, C. A. Carr, (NMZ); I 6,Bulawayo, Riverside (20.10s 28.43E), xii.1982, Mrs Hughes, (NMZ); I 9,Bulawayo, Riverside (20.1oS 28.43E), i.1984, Mrs Hughes, (NMZ); I 9,Bulawayo, Burnside (2o.1oS 28.43E), g.xii.1982, Mrs P. Lorber, (NMZ); I 6, Bulawayo, Burnside (20.10s 28.43E), 17.i.1g83, Mrs P. Lorber, (NMZ): MALAWI: I 9,Nyasaland, Bowa District (Dowa ?) (13.40s 33.55E), 1918, J. B. Davey, (BMNH); I 6, Livingstonia, N. Nyasaland (ro.yjS 34.1oE), 8.xii.1941, R. C. Wood, (NMZ); I 9,ACNE 259, Monkey Bay (14.04s 35.05E), 6.i.1978, B. Sharp, (NCI): TANZANIA: I 9,Maboya, German E. Africa, 1914, Dr E. J. Baxter, (BMNH); I 9,G. E. Africa, road to Kilosa, Usagara District, 1500-2500 ft (approx. 06.49s 37.ooE), 22-26.xii.1910, S. A. Neave, (BMNH): ETHIOPIA: holotype 6 of Navasius kristenseni, Abessinien, Kristensen leg., (ZMC): YEMEN: I 9,San'a, ca 7000 ft (15.25N 44.14E), vi.1938, Dr P. W. R. Petrie, (BMNH).

Bankisus carinifons (Esben-Petersen), Figs I I, 2 1-29, 31. Navasius carinifrons Esben-Petersen, 1936: 204. Bankisus carinifrons (Esben-Petersen); Stange, 1976: 288; Holzel, 1983: 21 I REDESCRIPTION:based on the female holotype of Navasius carinzyrom Esben- Petersen, and other material examined. Sexes alike, characterized by short broad maculated wings, whitish thoracic tergites, short clavate antennae and longish legs. Size: mean measurements in mm for I I specimens, ranges in brackets: length of body 13,4 (12,o-15,o); forewing 18,3 (16,o-20,o); hind wing 17,7 (15,5-19,0); antenna 2,g (2,q-3,6). Head: frons, vertex, occiput yellowish, three black marks between vertex and occiput, a broad black band extends from above toruli, between antennae onto clypeus; clypeus dark above pale distally, labrum pale. Maxillary and labial palps black, articu- lations pale, terminal labial palpomere spindle-shaped; submentum pale, bearing about eight long pale setae. Antennae short, less than one third the forewing length, distinctly clavate; flagellum comprising 20 or 21 flagellomeres, pale with brown annulations, a few completely pale flagellomeres precede the black club; scape pale with dark spot on inner surface, pedicel and proximal flagellomere brown. Pronotum with shallow anterior transverse furrow; three dark stripes extend posteriorly from furrow, with two lateral dark spots and diffuse markings usually discernible, markings may vary; setae on pronotum long, curved, pale on anterior and lateral margins, black centrally. Pleurites and sternites dark brown to black. Mesothorax: mesoprescutum black anteriorly, pale posteriorly, bearing long intermingled black and white setae in brown alveoli (remainder of pterothoracic tergites yellowish-white dorsally, except for diffuse marks on metatergites); sparse intermingled black and white setae occur on mesonotum and mesoscutellum, with long soft white setae along posterior margin of mesoscutellum. Metathorax: metanotum with sparse pale setae, metascutel- 204 J. enl. Soc. srh. Afr. Vol. 48, No. 1, 198.5

Figs 2 I -29. UankZSus carinfins (Esbcn-Petrrsen). 2 1-26 ~nalrgrnitalia. 2 I. Apex of abdomen. lateral. 22. Apex of abdomen, ventral. 23. Gonarcus and parameres, lateral. 24. Conarcus and paramcres, dorsal. 25. <;onarcus and paramew" ventral. 26. Gonar- cus and paramerrs, caudal. 27-29 femalc genitalia. 27. Apex of abdomen, lateral. 28.

Apex of abdomen, ventral. 29. Sprrmathrra. Abhreviations: Mrd - mrdiuncus; others as in Figs 12-20. lum with sparse black and whitr setae. Pterothoracic pleurites with broad black band above coxae, but not reaching wing bases; long white setae present on pleurites. Legs: Sorrlegs; coxar brown on outer surSaces, otherwise palr, trochanters pale, femora with threr brown stripes, and black and white srtae in black alveoli, tibiar pale with narrow annulation distally and bearing black and white sctae in black alvro- Mansell: ant lions of thegenus Bankisus 205 li, tarsi with first tarsomere long, remaining four shorter, all covered with short black sctac, first three tarsomeres pale, apical two black; middle legs similar to forelegs but with two femoral stripes, tibiae with three longtitudinal stripes; hind legs similar to forelegs but with femora almost completely dark brown, tibiae pale with faint apical stripes, no white setae present. Wings: short, broad, forewings broader than hind wings; marked as depicted in Fig. I I. Rs with about 7 or 8 branches in both wings, other venation as for genus. Abdomen: shorter than hind wings, with markings and pubescence similar to B. oculatus. Male (Figs 21-26, 31) with each half of tergite g reaching sternite g which is broad with rounded apex; ectoprocts with postrrioventral margin projecting. Gonarcus arcuate, mediuncus prominent, sclerotized, surface covered with delicate scale-like sculpturing (Fig. 31); parameres broad, sclerotized, almost shoe-shaped with inner poste~iormargins projecting upwards, ventral surfaces with tooth-like sculpturing (Fig. 31): gonosetae present; gonarcus, mediuncus, parameres enveloped in eversible membranous sac; hypandrium internum very delicate, difficult to discern. Female (Figs 27-29) with tergite 8 tapering ventrally; each half of tergite 9 elongate; ectoprocts oval, bcaring long black setae but no fossorial setae. Pregenital plate small, tooth-like, lying mcdially at posterior margin of sternite 7. Posterior gonapophyses absent; anterior gonapophyses large, digitiform, densely covered with black setae, membranously connected te apices of tergite 8; lateral gonapophyses separated from each other by narrow membranous area, bearing short stout curved fossorial setae posteriorly, and bristles ventrally above genital opening; a plate-like structure bearing setae occurs below the opening. Spermatheca sclerotized, darkly pigmented, shaped as in Fig. 29.

Figs 30-31. SEM of ventral surfaces of I)arameres. 30. Bankisus oculatus Navas. 31 Bankzsus carini- from (Esben-Petersen). 206 J. ent. Soc. sth. Afr. Vol. 48, No. 1, 1985

This species is easily distinguished from B. oculatus by its shorter broader wings, short clavate antennae and by features of the male and female genitalia, especially the form of the parameres. All the locally collected specimens were captured at light. Nothing is known about the biology. Distribution: widespread in the Afrotropical Region: recorded from South Africa, Zimbabwe and Zaire.

MATERIALEXAMINED. Holotype ? Navasius carinifom Esben-Petersen, 2 6 and 9 ? specimens. SOUTH AFRICA. Transvaal: I 9,ACNE 255, Satara (24.23s 31.47E), i.1969, A. C. Kemp, (NCI); I 9,AcNE 256, Skukuza (24.598 31.55E), 14.i.1971, A. Braack, (NCI); I 9, AcNE 257, Skukuza (24.59s 31.55E), 18.ii.1g73, A. Braack, (NCI); 1 ?, ACNE 289, Skukuza (24.59s 31.55E), 14.i.1969, H. & A. Braack, (NCI);2 d I 9,ACNE 207, Skukuza, 24.593 31.55E, 18-21 .i.1984, M. W. Mansell, (NCI); I 9, AcNE 208, Renoster Koppie, 25.07s 31.36E, 2o.i.1984, M. W. Mansell, (NCI); I 9, Lower Sabie (25.08s 31.55E), 26.iii.1952, Janse & Vari, (TM); I 9,Lower Sabie (25.08s 3 I .55E), 4-5.i~.1952, L. Vari, (TM); all these localities in the Kruger National Park. ZIMBABWE: I 9, Dotts Drift, Chisumbanji, Lower Sabi, Rhodesia (20.40s 32.16E), i.1969, Nat. Mus. S. Rhodesia, (NMZ): ZAIRE: holotype ? of Navasius carinifom, Ht. Katanga, Chinkolobwe (Shinkolobwe) (I1.1oS 26.4oE), 1g.x.rg30, J. Ro- mieux, (MG).

Bankisus elegantulus (Esben-Petersen). Navasius elegantulus Esben-Petersen, 1936:203. Bankisus elegantulus (Esben-Petersen); Stange, 1976: 288; Holzel, 1983: 21I. This species, known only from the holotype, is distinctive. The description by Esben-Petersen (1936) is accurate and cannot be eleborated without additional material. The holotype is a female, and not a male as stated by Esben-Petersen: the abdomen has been repaired, and the attached terminalia are those of a female. If an incor- rect abdomen has been attached, lack of pilulae indicate that the specimen is a female.

Bankisus elegantulus can be distinguished from B. carinzjirons, which it closely resembles, by the features outlined by Esben-Petersen (1936: 205)) but especially by the reddish-brown thoracic tergites of B. elegantulus.

MATERIALEXAMINED. ZAIRE: holotype 9 Navasius elegantulus Esben-Petersen, Ht. Katanga, Chinkolobwe (Shinkolobwe) (II.I~S26.40E), 28.ix.1930, J. Romieux, (MG). Bankisus triguttatus NavSs. Bankisus triguttatus Navas, 1gz6a:104; 1926b: 88; Stange, 1976:288; Hiilzel, 1983: 21 1. This species, also known only from the holotype is characterized by the excep- tionally broad forewings and long narrow hind wings, which are markedly iridescent. The wings are less heavily maculated than the preceding three Bankisus species. Banki- sus triguttatu~ can be distinguished from B. oculatus and B. carinzfrons by its reddish- brown thoracic tergites, and from B. elegantulu~by the much broader forewings, fewer Mansell: ant lions of the genus Bankisus 207 markings and by absence of a row of spots along the raised vertex. The description by Navas (1926a) is accurate and must suffice in the absence of additional specimens.

MATERIALEXAMINED. ZAIRE: Holotype ? Bankisus triguttatus NavBs, Tshela (04.57s 12.57E), 1o.xi.1920, Dr H. Schouteden, (MRAC).

Bankisus maculosus Holzel.

Bankisus maculosus Hiilzel, I 983: 2 I o. This species resembles B. oculatus Navis (Holzel, 1983), and is also known only from the holotype. 'The specimen has not been examined, so no comment can be made upon its status at present. It is the first Bankisus species to be described from outside Africa, and it was recorded from Oman on the Arabian Peninsula. ACKNOWLEDGEMENTS The following persons and institutions are thanked for the loan of material: S. J. Brooks (BMNH), B. Hauser (MG), H. M. Andre (MRAC), D. I,. Hancock (NMZ), V. B. Whitehead (SAM), R. B. Toms (TM), N. P. Kristensen (ZMC). The National Parks Board of South Africa and the Transvaal Department of Nature Con- servation are acknowledged for permission to collect on their reserves. I am grateful to H. Braack, A. Braack and L. E. 0. Braack for collecting material in the Kruger Nat- ional Park for this study. I thank G. L. Prinsloo (NCI) for reading and commenting on the manuscript; S. Schwartz (NCI) drew Figs 10 and I I, and H. van Tonder, Plant Protection Research Institute, Electron Microscope Unit, provided Figs 30 and 3 I.

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Accepted 10 December 1984 Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 4012

Reference Citation: Mansell, M. W. 1985 [1985.??.??]. The ant-lions of southern Africa (Neuroptera: Myrmeleontidae). Introduction and genus Bankisus Navás. Journal of the Entomological Society of Southern Africa 48:189-212.

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