Neuroptera: Myrmeleontidae: Acanthaclisini)

Zootaxa 4497 (3): 346–380 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2018 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4497.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:F847993D-11E7-41A9-8E57-08C64450D1A3

Antlions of southern Africa: Syngenes Kolbe, 1897, with descriptions of two new species and comments on extra-limital taxa (Neuroptera: Myrmeleontidae: Acanthaclisini)

MERVYN W. MANSELL Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa. E-mail: [email protected]

Abstract

The southern African species of Syngenes Kolbe, 1897, are revised and compared with their Afrotropical congeners. Three species occur in the region: S. longicornis (Rambur, 1842) and two new species, S. medialis and S. scholtzi, described and illustrated below. A key to the Afrotropical taxa is provided. The larva of S. longicornis is illustrated. The genus is widespread in the Afrotropical Region and extends to the Middle East and Oriental Region. Syngenes species are typically broad-winged antlions, with irregular bifurcate and biaereolate cells between the costa and subcostal veins, which distinguish them from other members of the tribe Acanthaclisini.

Key words: Neuroptera, Myrmeleontidae, Acanthaclisini, Syngenes, new species, larva, southern Africa, Afrotropical Re- gion

Introduction

The antlion genus Syngenes Kobe, 1897, (Neuroptera, Myrmeleontidae, Myrmeleontinae, Acanthaclisini) includes ten available names. Syngenes longicornis (Rambur, 1842), S. inquinatus (Gerstaecker, 1885), S. debilis (Gerstaecker, 1888) and S. carfii Insom & Terzani, 2017 are recorded from Afrotropical Africa. Syngenes maritimus (Needham, 1913), S. dolichocercus Navás, 1914 and S. alluaudi (van der Weele, 1909) were described from Aldabra Island and Madagascar, and S. arabicus Kimmins, 1943 from Saudi Arabia. Syngenes horridus (Walker, 1853) and S. palpalis Banks, 1931 are known from the Oriental Region (India, Indochina, Pakistan) (Stange 2004). Syngenes species are readily distinguished from other Acanthaclisini by the irregular biaereolate cells and bifurcate veins in the costal area (between the costa and subcostal veins) of the forewings, and distinctly broader wings than other Afrotropical genera in the tribe: Centroclisis Navás, 1909, Jaya Navás, 1912, Fadrina Navás, 1912 and Phanoclisis Banks, 1913. Accurate identification of the Afrotropical species has remained complex owing to infraspecific variation in wing markings and venation, lack of sufficient representative specimens for analysis and especially, uncertainty regarding the correct identity of the first species to be described, S. longicornis (as Acanthaclisis) by Rambur (1842). A study of the southern African taxa has revealed three distinctive species from south of the Kunene and Zambezi Rivers: S. longicornis and two undescribed taxa documented here. The revision has also established the correct identity and provenance of S. longicornis, and consequently, an accurate interpretation of one of the ancient names in Myrmeleontidae. These three taxa are the focus of this paper, while comments are provided on the remaining six Afrotropical species names. Of these, S. maritimus is reinstated from synonymy with S. longicornis (Ohm & Hölzel 1995), S. alluaudi is transferred to Jaya and S. carfii is synonymized with S. longicornis. Five species, S. inquinatus, S. debilis, S. maritimus S. dolichocercus and a new species from southern and central Africa are morphologically similar and may form a complex that extends from West Africa to the Indian Ocean Islands and Madagascar. It is consequently proposed that the two species from West Africa (S. inquinatus S.

346 Accepted by A. Letardi: 23 Aug. 2018; published: 10 Oct. 2018 debilis), the central species described below, and the Indian Ocean Islands species (S. maritimus, S. dolichocercus) be regarded as separate entities to avoid any incorrect association, until further morphological and molecular analyses are conducted to determine their status. This can only be achieved once sufficient material has been procured for morphological and molecular analyses. The current revision is consequently limited, primarily to the southern African taxa where sufficient material is available. Whereas, for two of the species, S. inquinatus, and S. debilis, only the type specimens are definitive, while for S. maritimus, two syntypes, the two type specimens of S. dolichocercus, and six other poorly preserved specimens from Madagascar were studied. Stange & Miller (1985) described the larva of S. longicornis from South Africa, and it is illustrated below (Fig. 12). This is the ninth in a series of papers revising the genera of southern African Myrmeleontidae (see Mansell 2018) and the first to treat the tribe Acanthaclisisini.

Material and methods

Material examined is in the following institutions: Natural History Museum, London (BMNH); Ernst-Moritz-Arndt Universität, Greifswald (EMAU); Institut royal des Sciences naturelles de Belgique, Bruxelles (IRSN); South African National Collection of Insects, Pretoria, South Africa (SANC); Transvaal Museum, Pretoria, (now Ditsong Museum) (TMSA), Muséum national d'Histoire naturelle, Paris (NMHN), Musée royal de l’Afrique centrale, Tervuren, Belgium (MRAC); Zoological Institute, Lund, Sweden (ZILS). Accession numbers, as recorded in the Palpares Relational Database (Mansell & Kenyon 2002), are provided for material examined. The first four letters of each unique accession code reflect the institutional acronym as provided above. All specimens recorded below are in SANC except where otherwise indicated. Numbers cited in bold after references refer to the universal numbering system of the Lacewing Digital Library (Oswald 2016). Terminology and abbreviations for wing venation are provided in Figure 3, and for genital structures in Figures 8, 13–16 and 17–19, with the following additional terms used in the text: A1–A3, anal veins; T1–T9, abdominal tergites; Ta1–Ta5, tarsomeres 1–5; S1–S9 abdominal sternites. Terminalia were dissected after maceration (5–6 hours) in a cold 10% KOH solution, and photographed using a Leica® Z16 APOA camera. Images were edited using Leica® Application Suite software version 4.4.0 (Leica Microsystems) and Irfan View. A Canon® EOS550D camera coupled with a 100mm Canon® macro lens was used to image adults, using Helicon Focus Auto montage software (Helicon Soft Ltd).

Syngenes Kolbe, 1897

Syngenes Kolbe, 1897: 15. 3434. Type species: Acanthaclisis debilis Gerstaecker, 1888: 100, by monotypy. 2558. Onclus Navás, 1912: 166. 549. Stange & Miller 1985: 38 (synonymy). 5823.

Diagnosis. Characterized by broad forewings with irregular biaereolate cells and bifurcate veins in the costal area (between C and Sc). Forefemur with two femoral sense hairs, one on middle femur, absent from hind femur. Body coloration yellow and black. Males with long ectoprocts. Thorax and abdomen with sparse pilosity. Labial palps very small with short slit-shaped palpimacula. The known larva (Fig. 12) has a distinct median lobe on the clypeal- labrum and is white with sparse black markings (Stange 2004).

Key to species of Afrotropical Syngenes

1. Taxa from Africa ...... 2 - Taxa from Madagascar and Indian Ocean Islands...... 6 2. Tibial spurs right-angular, with prominent internal flange (Figs 4, 5); biaereolate cells present in costal area of forewing before Rs (Fig. 3); abdomen with chevron-like pattern (Fig. 11). Eastern African coastal belt, South Africa to Somalia ...... S. longicornis (Rambur).

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 347 - Tibial spurs not sharply bent at a right angle, more evenly curved and lacking prominent triangular flanges...... 3 3. Biaereolate cells absent from costal area of forewing before Rs and rarely present before Rs2 (Fig. 27); unbroken series of biaereolate costal cells only commencing beyond Rs2; tibial spurs slender, strongly curved (but not bent at right angle), with slight flange in forelegs (Figs 28, 29); Central, South and Eastern Africa ...... S. medialis sp. nov. - Biaereolate cells present in costal area of forewing before Rs or commencing at Rs with three or more before Rs2 ...... 4 4. Wings narrow relative to other Syngenes species (Figs 44, 45); body black with bright yellow markings (Figs 46, 51). Namibia and Northern Cape Province, South Africa...... S. scholtzi sp. nov. - Wings broad; unbroken series of biaereolate cells present in forewing costal area from before Rs2 to pterostigma. Central and West Africa...... 5 5. Costal area with 1–2 biaereolate cell before Rs in each forewing (based only on the Holotype from Congo) (Figs 73, 74). Wings heavily marked...... S. inquinatus (Gerstaecker). - Costal area with variable number of biaereolate cells (1–4) before Rs (based only on one syntype from Lagos, Nigeria), slender species, wings not heavily marked, spurs curved, slender (Figs 76, 77) ...... S. debilis (Gerstaecker). 6. Forewing costal area with many (> 8) biaereolate cells before Rs ...... S. dolichocercus Navás - Forewing costal area with fewer (< 6) biaereolate cells before Rs ...... S. maritimus (Needham)

Syngenes longicornis (Rambur, 1842) Figs 1–24, 66, 69, 72.

Acanthaclisis longicornis Rambur, 1842: 382. 5314. Myrmeleon longicornis (Rambur): Walker 1853: 320. 6194. Syngenes longicornis (Rambur): van der Weele 1907: 266. 406. Syngenes carfii Insom & Terzani, 2017: 55. 16257. Syn. nov.

Redescription. Based on Holotype ♀, 10 ♂ and 27 ♀. Diagnosis. Medium-sized yellow and black antlions. Antennae long, clavate, narrowly annulated with black and yellow. Abdomen with characteristic chevron-like pattern (Fig. 11). Wings broad, with posterior margins of forewings slightly undulating, alternating pale and light brown markings on wing veins, usually with a faint curved mark in distal region of forewing, occasionally with one or two large spots at posterior margin of forewing (Fig. 1). Forewing with characteristic irregular biaereolate cells or bifurcate crossveins in costal area, biaereolate cells extending from before Rs to pterostigma (Fig. 3). Distinguished from other Syngenes species by the right-angular tibial spurs bearing a prominent triangular flange on inner surface (Figs 4, 5), and by the characteristic abdominal pattern.

TABLE 1. Measurements (mm) of Syngenes longicornis. Figure above is the mean, size range below. mm Antenna Head width Forewing Forewing width Hind wing length Hind wing width Body length length length Holotype ca. 36.0 ca. 9.5 ca. 29.0 ca. 9.0 Female Males 9.3 4.1 41.1 11.1 37.1 9.9 36.0 n = 10 8.4–10.3 3.9–4.4 36.5–45.0 10.0–12.0 32.0–40.7 9.0–11.0 32.3–38.6 Females 9.8 4.2 43.0 11.5 39.3 10.1 35.4 n = 27 8.1–11.4 3.6–4.7 35.6–46.3 10.0–12.2 29.2–42.4 9.0–11.0 31.3–37.5

Head: wider than prothorax, vertex slightly raised, rounded. Frons and vertex black with two large laterally elongated embossed orange patches on vertex, two smaller spots in between, vertex also with row of small, embossed yellow marks, occiput with three large embossed yellow patches. Frons and vertex with short white recumbent setae. Face below antennae, clypeus, genae, labrum uniformly yellow, with long, sparse, pale setae. Antennae long, clavate, longer than twice head width, toruli less than scape diameter apart, scape almost touching eyes, scape and pedicel uniformly yellow, with sparse long white setae, flagellomeres short, annulated with black and yellow, covered in short black setae. Eyes large, greater than hemispherical, with sparse ocular setae usually present. Maxillary and labial palps very small, much less than head width, yellow, terminal labial palpomere spindle-shaped with long acute apex, palpimacula short, slit-shaped (Fig. 6).

348 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL FIGURES 1–2. Syngenes longicornis (Rambur). 1, male habitus NEUR11837 (forewing length 42 mm); 2, female habitus NEUR11833 (forewing length 42 mm), showing wing pattern variation.

Thorax: yellow with black markings, sparsely pilose. Pronotum longer than wide, tapering anteriorly. Anterior margin with two curved black lines laterally merging anteriorly, sides of pronotum with two diffuse black streaks, another two oblique streaks on each side of midline at posterior margin, central pronotum with distinct dots. Sparse long white and black forwardly curved setae situated laterally, short black bristles present along anterior raised margin, posterior margin with long curved black setae. Mesothorax: mesoprescutum yellow, brown anteriorly, a large diffuse black mark on either side of midline, anterior margin with long curved black setae; mesoscutum yellow with four longitudinal black lines on either side above wing bases and sparse black setae; mesoscutellum yellow with two black central stripes and two larger diffuse black spots posterior to central stripes. Metathorax: metaprescutum yellow with two large black marks on either side, metanotum with two velvety spots on each side, two longitudinal stripes laterally above wing bases, long curved white setae laterally; metascutum yellow with a large black mark on either side of midline, devoid of setae. Pleurites below wings yellow with irregular brown marks, sternites yellow, dense long white setae present on pleurites and sternites. Wings: broad, forewings longer, broader than hind wings, apices sub-acute, membrane hyaline, veins with alternating sections of yellow and black, bearing very short sparse black and yellow setae on correspondingly coloured areas. Forewings broad with slightly undulating posterior margin, apices rounded with sub-acute tip, usually with a faint curved mark in distal part, occasionally with one or two spots at posterior margin. Axillary protuberance at base of forewing yellow with long dense white and black setae. Costa alternating yellow and black with tuft of short black bristles at base, costal area narrow at base, broadening distally; costal veins irregular,

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 349 usually 7 cells (but variable) before Rs, some veins bifurcate, with biaereolate cells in unbroken series extending distally from Rs. Pterostigma pale yellow, indistinct. Apical margin of wing with densely arranged narrowly-forked veins. Hypostigmatic cell elongate. Sc, R and proximal region of CuA with yellow and black chevron-like pattern. Rs arises well beyond CuA fork; area between Sc and R mostly pale with black cross-markings; 8–10 presectoral veins present. Mp2 (oblique crossvein) arising beyond CuA fork, CuP arises just before basal crossvein, fused with A1, A2 very short (sometimes not manifest, merged with A1 from base), curved before merging with A1. Hind wings narrower than forewings, lanceolate, always unmarked; axillary protuberance at base yellow with long dense white setae, C mainly yellow with tuft of long white recumbent setae at base, costal area narrow at base, broadening then narrowing, uniareolate, veins unbranched. Pterostigma yellow, hardly discernible. Hypostigmatic cell long. 5–6 occasionally 7 presectoral crossveins, Rs arises beyond Mp2 fork, Cu fused with posterior branch of Mp2 fork; anterior banksian line visible in distal portion of wing. Pilula axillaris in males conspicuous with densely packed short brown recumbent setae, females with long white setae in this position.

FIGURES 3–6. Syngenes longicornis. 3, base of forewing showing main veins and biaereolate cells in subcostal area; 4, foreleg tibial spurs with flange and facial brush; 5, hind leg tibial spurs with flange; 6, labial palp with palpimacula. Abbreviations: A1 = First anal vein, C = Costal vein, CuA = Anterior cubital vein, CuP = Posterior cubital vein, Cc = Costal cell, Mp2 = branch of posterior medial vein (oblique vein), R = radial vein, Rs = Radial sector, Rs2 = second radial sector, Sc = Subcostal vein.

Legs: forelegs strongly developed with short stout spines. Coxa, trochanter yellow with long white pubescence. Femur yellow with faint diffuse brown marks, very stout, thickened at base tapering distally, two femoral sense hairs near base, ventral surface with two rows of strong short black spines accommodating tibia on closure, long

350 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL white setae with occasional black setae present. Tibia slender, yellow with four black stripes at femoral articulation and two black annulations proximally, one distally; lateral surfaces with long white setae, black setae dorsally, with dense pad of short black setae ventrally fitting between the two rows of black bristles on femur. A dense facial brush of short golden pubescence present on interior apical surface (Fig. 4). Tibial spurs (Figs 4, 5) characteristically bent at a right angle with prominent triangular flange. Tarsus with Ta5 longer than Ta1–Ta4 combined, Ta1–Ta4 yellow with black annulation on each, covered in short black bristles, Ta5 long, yellow, black apically, covered with short black bristles; preapical claws dark reddish-brown, stout, strongly curved. Middle legs shorter, more slender than forelegs, entirely yellow; coxa, trochanter shorter than in foreleg, with long white pubescence. Femur yellow with one sensory seta proximally, two rows of black bristles ventrally as in foreleg, covered with long white setae and sparse patch of long black bristles dorso-apically. Tibia, tarsus as in forelegs, but without facial brush. Hind legs long slender, yellow, lacking femoral sense hair; coxa, trochanter as in middle leg, femur sometimes with small black dots at setal bases, dorsal surface with short black setae, laterally with long black curved setae, long white setae proximally, ventral surface with two rows of short black bristles; tibia, tarsus as in middle leg.

FIGURES 7–9. Syngenes longicornis 7, male terminalia lateral; 8, dorsal; 9, ventral. Abbreviation: T9, Tergite 9.

Abdomen: yellow with characteristic black markings imparting a chevron-like pattern (Fig. 11), tergites with sparse short black setae, sternites yellow with sparse short white setae but sternites 1–2 with long soft dense white pubescence. T1 yellow, black centrally, T2 yellow laterally black centrally, T3 yellow with black mark on either side of midline tapering to two parallel stripes anteriorly. Male (Figs 7–10, 13–16): T9 yellow, divided dorsally,

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 351 two black streaks along posterior margins. IX brown, short, rounded with pale central stripe and fringe of long black posteriorly directed setae. Ectoprocts yellow, very long, anterior dorsal margin with 2 black streaks and long anteriorly-curved white and black setae becoming much longer posteriorly, base of ectoprocts swollen bearing a dense tuft of short black setae. Gonarcus (Figs 13–16) bears a prominent beak-shaped Mu; Pa (Figs 13–16) sclerotized, articulating with Gs, terminating in upwardly curved sharply pointed extremities. Female (Figs 17–20): T9 divided dorsally, Ga digitiform with long white setae, Gl clavate bearing long stout, slightly curved pale spines, Epr rounded, bearing slender, slightly curved setae. Spermatheca (Fig. 20) broad proximally becoming slender and coiled, tapering distally, with fine setae along distal extremity.

FIGURES 10–12. Syngenes longicornis. 10, abdomen dorsal with hair pencil details; 11, abdomen dorsal showing pattern; 12, third instar larva, photo: H. de Klerk©.

Larva (Fig. 12). Described by Stange & Miller (1985), who provided notes on the habitat and behavior. It is a beautiful black and white larva that is fast moving and free-living in coastal dunes on the fringes of vegetation along the eastern seaboard of Africa. Distribution (Fig. 72). Extending along the eastern coastal belt from the Western Cape Province, South Africa to Mozambique and into Tanzania and Somalia. Comments. The first Syngenes species to be described, S. longicornis (as Acanthaclisis) (Rambur 1842) has been misinterpreted by most previous authors: van der Weele (1907, 1908), Esben-Petersen (1916, 1920), Banks (1920), Navás (1924, 1935), Fraser (1951), Handschin (1963), Ohm & Hölzel (1995), Insom & Terzani (2017). The correct identification of this taxon was consequently crucial to determining the subsequent species described from

352 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL the Afrotropical Region. Rambur (1842) specifically stated that the country of origin was unknown, but there is now a label attached to the type specimen indicating “Senegal” (Fig. 23). This is clearly an erroneous subsequent addition (Prost 1998), who also stated that S. longicornis is a southern African taxon that does not occur in West Africa, an assumption verified here. This name however, still required association with a known population of Syngenes to understand its characteristics and extent. This relationship has now been clearly associated with a species that occurs in coastal forests and verges of dune vegetation along the east coast of Africa. It does not extend far inland, unless ideal sandy conditions exist (e.g. Tembe Elephant Reserve, South Africa). This species, with its characteristic white larvae (Fig. 12), has been uncritically, but correctly, identified as S. longicornis by recent authors: Stange & Miller (1985), Gess & Gess (1998). A characteristic of S. longicornis is that the tibial spurs are sharply bent at a right angle and there is a prominent triangular flange along the ventral surface (Figs 4, 5). This is also the typical spur structure of the genus Centroclisis, but is not as pronounced in other species of Syngenes. Furthermore, biaereolate cells are present in the forewing costal area from before Rs (Fig. 3). There are other features, including a chevron-like (inverted V-shape) abdominal pattern (Fig. 11) that characterize this species. Examination of the female holotype of Acanthaclisis longicornis in IRSN, and from photographs provided by Jerome Constant (IRSN) (Figs 21–24), has confirmed the tibial spur structure (Fig. 22), and features of the wings and abdomen closely match that of the southern and eastern African population, which is now confirmed as S. longicornis. It consequently does not occur in the other countries listed by Stange (2004): Benin, Madagascar, Senegal, Seychelles and Togo.

FIGURES 13–16. Syngenes longicornis. Male gonarcus and parameres complex: 13, lateral; 14, dorsal; 15, caudal; 16, ventral. Abbreviations: Gs = Gonarcus, Mu = Mediuncus, Pa = Paramere.

The specimen illustrated by van der Weele (1907, Pl. 9, Fig. 9) is not S. longicornis, as already stated by Navás (1912), but is clearly S. maritimus. Syngenes carfii Insom & Terzani, 2017, shares all the main characters for distinguishing S. longicornis: tibial spurs right-angled with internal flange, chevron-like abdominal pattern and the forewing costal area with biaereolate cells before Rs. It is consequently relegated to synonymy with S. longicornis.

FIGURES 17–20. Syngenes longicornis. Female Terminalia: 17, lateral; 18, ventral; 19, dorsal; 20, spermatheca. Abbreviations: Ga = anterior gonapophyses, Gl = lateral gonapophyses, Epr = Ectoproct, T9 = Tergite 9.

Additional material examined. SOUTH AFRICA, Western Cape Province: 1♀, Wilderness, 33°59’S 22°35’E, 24.ii.1965, J.S.Taylor, ZILS00114 (ZILS); 1♀, Still Bay 34°22’17”S 21°24’32”E, 40m, 15.ii.1971, D.J.van Wyk, NEUR11835. Eastern Cape Province: 1♂ 1♀, Kenton on Sea, 33°40’S 26°40’E, 50m, 15.ii.1971, R.A.Jubb, Malaise trap, NEUR11829; 1♀, Kleinemonde, 33°32’29”S 27°02’54”E, 20m, 17.ii.1971, M.W.Mansell, NEUR11836; 1♂, St Francis Bay, 34°10’S 24°50’E, 20m, 12.iii.1972, M.W.Mansell, NEUR09458; 4♂ 2♀, Port Alfred, 33°36’S 26°54’E, 20m, 15.xii.1988, E.Grobbelaar, NEUR11842. KwaZulu-Natal: 1♀, Lake Sibaya, 27°22’04”S 32°42’56”E, 50m, 7.i.1968, M.W.Mansell, NEUR11834; 1♀, same data but 23.i.1970, NEUR11833; 1♀, Cape Vidal, 28°07’24”S 32°33’23”E, 30m, 10.xii.1973, L.R.Minter, NEUR11156; 1♂, Umlalazi Nature Reserve, 28°57’S 31°45’E, 10m, 28.i.1988, J.G.H.Londt, NEUR11837; 3♀, Sodwana Bay, 27°32’S 32°41’E, 30m, 3.xii.1988, H.Terblanche, NEUR01956; 2♀, same locality, 8.ii.1997, R.G.Oberprieler, NEUR10145; 2♀, Kosi Bay Nature Reserve, 26°59’S 3°48’E, 10m, 8.ii.1990, E.Grobbelaar, NEUR01584; 1♀, Kuleni Farm, 27°54’43”S 32°21’52”E, 115m, 12.ii.1990, E.Grobbelaar, NEUR01612; 1♀, Sihangwana, Tembe Elephant Park, 27°02’30”S 32°25’20”E, 85m, 1.ii.1996, R.Stals, NEUR11828; 1♀, same data but 6.iv.1996, NEUR05699; 1♂ 1♀, same locality, 1.ii.1996, E.Grobbelaar, NEUR05595; 4♀, same data but 25.i.2006, NEUR09438; 1♀, St Lucia Lake,

354 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL 28°23’S 32°24’E, 20m, 11.ii.1997, R.G.Oberprieler, NEUR10144; 1♀, Ndumu Game Reserve, 26°51’60”S 32°15’00”E, 45m, 27.i.2006, E.Grobbelaar, NEUR09433; 1♀, Maphelana, 28°23’53”S 32°24’59”E, 29.i.1993, 6m, J.deG.Harrison, TMSA00885 (TMSA). MOZAMBIQUE: 1♂, Inhaca Island, 26°02’19”S 32°54’14”E, 10m, 23.i.1971, M.W.Mansell, NEUR11830; 1♀, Beira 20 km N, 19°45’22”S 35°01’28”E, 5m, 3.iv.2000, R.D.Stephen, NEUR05587; 1♂ 1♀, Chidenguele, 24°56’03”S 34°11’09”E, 18m, 27.ii.2016, A.J.Gardiner, NEUR12539.

FIGURES 21–24. Acanthaclisis longicornis Rambur, (Syngenes longicornis). Holotype female (IRSN): 21, habitus; 22, tibial spur; 23, associated labels; 24, forewing showing details of subcostal cells. Photographs: J. Constant (IRSN).

Syngenes medialis sp. nov. Figs 25–43, 68, 70, 72.

Etymology. Derived from its geographic distribution between the populations of S. scholtzi n. sp. in the west and S. longicornis in the east of southern Africa and in the middle of the distribution of Syngenes species extending from West Africa to Madagascar. Description. Based on holotype male, 14 male and 61 female paratypes. Diagnosis. Distinguished from other Syngenes species by the absence of biaereolate cells in the costal area before Rs and usually beyond Rs2 (Figs 27, 70). Wings broad, variously marked, sometimes with distinct large spots at hind margin of forewing. Abdomen banded, no chevron pattern. Tibial spurs in forelegs slender, strongly bent but not right-angular with very narrow flange, not expanding apically as in S. longicornis, hind tibial spurs evenly curved, without flange, TA 1–4 uniformly black, T5 pale. Labial palp with short slit-shaped palpimacula (Fig. 41).

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 355 TABLE 2. Measurements (mm) of Syngenes medialis. Figure above is the mean, size range below. mm Antenna length Head width Forewing length Forewing Hind wing Hind wing width Body length width length Holotype 9.7 4.6 46.7 12.5 41.7 10.5 44 Males 10.3 4.5 45.7 12.5 40.6 10.8 41.6 n = 15 9.1–11.6 4.0–5.0 41.1–49.0 11.0–12.8 36.2–43.8 9.5–12.0 36.0–46.5 Females 10.4 4.6 46.9 12.3 41.9 10.9 37.3 n = 60 8.6–11.8 3.8–5.1 44.3–50,3 11.2–14.0 36.0–44.8 9.0–12.3 31.0–42.6

FIGURES 25–26. Syngenes medialis n.sp. 25, Holotype male, NEUR09562 (Forewing length 47mm); 26, Paratype female NEUR11845 (Forewing length 47mm), showing variability in wing markings.

Head: wider than prothorax, vertex slightly raised, rounded. Frons and vertex black with short white recumbent setae; three large laterally-elongated embossed yellow marks present on upper frons, vertex with small irregular yellow spots, occiput with three distinct embossed orange patches. Face below antennae, clypeus, genae and labrum uniformly yellow, with long, sparse, pale setae. Antennae long, clavate, longer than twice head width, toruli less than scape diameter apart, scape almost touching eyes, scape and pedicel uniformly yellow, with sparse long white setae, flagellomeres short, annulated with black and yellow, covered in short black setae. Eyes large, greater than hemispherical, sparse ocular setae usually present. Maxillary and labial palps short, much less than head width, yellow, terminal labial palpomere spindle shaped with long acute apex, palpimacula short, slit shaped (Fig. 41).

356 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL FIGURES 27–30. Syngenes medialis. 27, Holotype base of forewing, NEUR09562, showing absence of biaereolate cells to beyond Rs2; 28, foreleg tibial spurs and facial brush; 29, hind leg tibial spurs; 30, abdominal pattern, NEUR11845.

Thorax: yellow with black markings, sparsely pilose. Prothorax slightly longer than wide, tapering anteriorly, lateral margins with two curved black streaks that merge anteriorly and posteriorly, central pronotum with single streak at anterior margin and two short parallel streaks posteriorly, with distinct black dots at setal bases. Sparse long white and black forwardly curved setae present laterally, short black bristles along anterior raised margin, posterior margin with long curved black setae. Mesothorax: mesoprescutum yellowish-brown, black anteriorly and laterally, with two yellow spots laterally, anterior margin with long curved black setae; mesoscutum yellowish- brown with three diffuse longitudinal black lines on either side above wing bases, with sparse black setae; mesoscutellum yellowish-brown with two curved black marks centrally, anterior and lateral margins narrowly black. Metathorax: metaprescutum black with pale midline, metanotum yellow with black V-shaped mark centrally, a black streak present laterally above wing bases, velvety spots not discernible, long curved white setae present laterally; metascutum black with three yellow spots, one centrally, two laterally. Pleurites below wings yellow with black streaks, sternites yellow, dense long white setae present on pleurites and sternites. Wings: broad, forewings longer, broader than hind wings, apices sub-acute, forewings more rounded than hind wings, membrane largely hyaline, veins with alternating sections of yellow and black, bearing very short sparse black and yellow setae on correspondingly coloured areas. Forewings broad with slightly undulating posterior margin, a distinct curved mark present in distal third, frequently with one or two large spots at posterior margins (22 of 75 specimens). Axillary protuberance at base of forewing dark yellow with long dense white and black setae.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 357 Costal vein alternating yellow and black with tuft of short black bristles at base, costal area narrow at base, broadening then narrowing slightly towards pterostigma Proximal costal area veins bifurcate but mainly unbranched, with no biaereolate cells; cells only becoming irregularly biareolate beyond Rs and usually beyond Rs2 (Fig. 27). Pterostigma white. Apical margin of wing with densely arranged narrowly forked veins. Hypostigmatic cell elongate. Sc, R and proximal region of CuA with yellow and black chevron-like pattern. Membrane between Sc and R heavily marked with alternating yellow and black bands. Rs arises well beyond CuA fork, with Rs2 at or just beyond first fork of Rs, 8–10 presectoral veins present. Mp2 (oblique vein) arising beyond CuA fork, CuP arises at basal crossvein, bifurcating after short free base with upper branch linking with CuA and lower uniting with A1, A2 not parallel to A1, evenly curved. Membrane at posterior base with long soft white setae. Hind wings narrower than forewings, lanceolate, with faint marks usually discernible in distal half, axillary protuberance at base yellow with long dense white setae, C yellow and black, costal area uniareolate, veins unbranched until pterostigma, costal area narrow at base expanding into a bulge then narrowing and parallel to Sc. Pterostigma white, hardly discernible. Hypostigmatic cell long. Usually 5–6, presectoral crossveins, Rs arises beyond Mp2 fork, Cu fused with posterior branch of Mp2 fork, anterior banksian line visible in distal portion of wing. Pilula axillaris in males conspicuous, bearing densely packed short brown recumbent setae. Females with long white setae in this position.

FIGURES 31–33. Syngenes medialis. Male terminalia. 31, lateral; 32, dorsal with hair pencils; 33, ventral.

Legs: forelegs strongly developed with short stout spines. Coxa, trochanter yellow with long white pubescence. Femur yellow with diffuse brown marks dorsally, very stout, thickened at base tapering distally, two femoral sense

358 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL hairs present near base, ventral surface with two rows of strong short black bristles accommodating the tibia on closure, long white setae occur laterally. Tibia slender, yellow with four black stripes at femoral articulation and two black annulations proximally; lateral surfaces with long white setae, black setae dorsally, with densely arranged short black setae ventrally that fit between two rows of black bristles on femur. A dense facial brush of short golden pubescence present on interior and ventral apical surfaces. Tibial spurs (Figs 28, 29) slender, strongly bent but not right-angular with slightly raised flange. Tarsus with Ta5 longer than Ta1–Ta4 combined, Ta1–Ta4 shiny black, covered in short black bristles, Ta5 long, yellow, black apically, covered with short black bristles, preapical claws dark reddish-brown, stout, strongly curved. Middle legs shorter, more slender than forelegs, entirely yellow. Coxa, trochanter shorter than in foreleg, yellow with long white pubescence. Femur yellow with one sensory seta proximally, two rows of black bristles ventrally as in foreleg, covered with long white setae and sparse patch of long black bristles dorso-apically. Tibia, tarsus as in forelegs, but spurs more evenly curved lacking raised flange. Hind legs long slender, yellow, lacking femoral sense hair. Coxa, trochanter as in middle leg. Femur with faint black diffuse black stripe laterally, dorsal surface with short black setae, laterally with long black curved setae, long white setae proximally, ventral surface with two rows of short black bristles, tibia yellow, slender with short coarse black setae, tibial spurs slender, gently curved.

FIGURES 34–37. Syngenes medialis. Male gonarcus and parameres complex: 34, lateral; 35, dorsal; 36, ventral; 37, caudal.

Abdomen: yellow with medially divided black bands at distal half of each segment (Fig. 30), tergites with sparse short black setae, sternites yellow with sparse short white setae but sternites 1–2 with long soft dense white pubescence, sternite 8 black proximally, sternite 9 black. Males with black hair pencils between segments 6 and 7 (Fig. 32). Male (Figs 31–37): T9 yellow divided dorsally; S9 black, short, rounded with pale central stripe and long black posteriorly directed setae (Fig. 33). Ectoprocts yellow with long postventral lobe, dorsal margin with black stripe (Figs 31, 32), postventral lobes with long anteriorly curved white and black setae becoming longer

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 359 posteriorly (Figs 31–33). Gonarcus (Figs 34–37 bears a prominent beak-shaped Mu; Pa (Figs 34–37) sclerotized, articulating with Gs, terminating in upwardly curved sharply pointed extremities. Female (Figs 38–40, 42): T9 divided dorsally, Ga digitiform with long pale setae; Gl clavate bearing very long stout, slightly curved brown spines, Epr rounded, bearing slender, slightly curved setae. Spermatheca (Fig. 42) very long relative to other taxa, sclerotized, broad proximally becoming slender, with long convoluted, coiled distal portions bearing fine setae along distal extremity.

FIGURES 38–40. Syngenes medialis. Female terminalia: 38, lateral; 39, ventral; 40 dorsal.

Larva. Very similar to that of S. longicornis, being white with the characteristic black markings. Unfortunately, the four specimens received in the larval stage pupated before photographs or measurements could be obtained. They were collected in the sand of a dry riverbed. Distribution (Fig. 72). This species is widespread in the eastern and northern provinces of South Africa: Limpopo, Mpumalanga, KwaZulu-Natal, extending south to the Eastern Cape. Also northwards into Botswana, Mozambique, Zimbabwe, northern Namibia, Malawi, Congo and Tanzania. Comments. Syngenes medialis resembles both S. inquinatus from West Africa and S. maritimus from Aldabra Island and Madagascar, and it was uncertain as to which of these two taxa this common and widespread species should be assigned. Examination of 76 specimens, extending over a wide area from Brazzaville (Congo) to South

360 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL Africa has however, revealed a consistent feature in which the costal veins of S. medialis are either simple or forked, without any biaereolate cells before the origin of Rs, and usually also beyond Rs2. By contrast, S. inquinatus and S. maritimus have a variable number of crossveins before Rs and biaereolate cells after Rs before Rs2. It was consequently decided to allocate separate specific status to avoid an incorrect association with either. It is clear that these three morphologically similar taxa constitute a species complex, with S. inquinatus occurring in West Africa, S.maritimus inhabiting Madagascar and Aldabra Island while S. medialis occupies a large area in between, in southern, eastern and central Africa. A specimen in MRAC labeled as Cotype of S. dolichocercus by Navás (1925) is S. medialis. Type material examined. SOUTH AFRICA, Limpopo Province: Holotype ♂, Makhutsi Camp, Lekgalameetse Nature Reserve, 24°11’57”S 30°20’22”E, 1200m, 9.i.1997, hand-netted, M.W.Mansell, NEUR09562. Paratypes: 2♂ 3♀, same data as holotype; 1♀, Nylstroom, 24°42’S 28°20’E, 15.iii.1978, S.J.van Tonder, NEUR11818; 1♀, 10 km N Wyllies Poort, 22°53’S 29°52’E, 30.xii.1978, L.R.Minter, NEUR09075; 1♂ 1♀, Mogol Nature Reserve, 23°58’S 27°45’E, 20.i.1983, M.W.Mansell, NEUR00118; 1♀, D’Nyala Nature Reserve, 23°45’03”S 27°49’32”E, 893m, 8.xii.1989, M.W.Mansell, H.&U.Aspöck, NEUR01314; 1♀, same locality, 13.i.1991, E.Grobbelaar, NEUR10482; 1♀, Lapalala Nature Reserve School, 23°54’06”S 28°17’31”E, 1158m, 18.ii.1994, R.G.Oberprieler, NEUR11868; 1♀, Natuurpraal Resort, Melkrivier, 24°00’S 28°25’E, 1100m, 20.ii.1995, M.W.Mansell, H.&U.Aspöck, NEUR02038; 1 ♀, Messina, 22°19’41”S 30°01’37”E, 525m, 22.xi.1996, C.Peyper, NEUR10178; 1♀, Ben Lavin Nature Reserve, 23°09’S 29°57’E, 19.i.1998, M.D.Picker, NEUR09583; 1♀, Alldays, 22°40’S 29°06’E, 8.iii.2000, R.D.Stephen, NEUR05580; 1♀, Mapungubwe, 22°12’09”S 29°21’50”E, 518m, 22.xii.2009, No collector name, NEUR11867; 1 ♂, Dongola Ranch, 22°13’57”S 29° 41’19”E, 630m, 23.xii.2009, No collector name, NEUR11866; 1♀, Northwich Game Farm, 22°53’21”S 29°52’25”E, 836m, 19.i.2014, D.K.Bakkes, NEUR11922; 1♀, Bievack 14 Farm, 22°28’20”S 28°55’14”E, 636m, 22.iii.2014, D.Kamffer, NEUR11900; 1♀, Zuurfontein Farm, 23°48’29”S 27°46’27”E, 835m, 28.iii.2014, N.J.Parry, NEUR11885; 1♂, Gnu Ranch, 24°01’44”S 27°29’13”E, 998m, 19.iv.2014, R.W.Mansell, NEUR11878. Mpumalanga Province: 1♀, Louws Creek, 25°37’S 31°16’E, 15.iii.1920, A.Rob, TMSA00889, (TMSA); 1♂ 3♀, Ngweti River, 25°31’S 31°42’E, 8.xi.2001, G.Nelson, NEUR05752. KwaZulu-Natal: 1♀, Lake Sibaya, 27°22’04”S 32°42’56”E, 50m, 15.xii.1968, J.L.Minshull, NEUR11854; 1♀, Empangeni, 28°44’S 31°53’E, 22.xi.1989, P.E.Reavell, NEUR11901. Eastern Cape Province: 1♀, Kei Bridge, 32°30’23”S 27°58’45”E, 3.iv.1990, N.J.Duke, NEUR11832. Kruger National Park: 1♀, Satara Camp, 24°23’40”S 31°46’40”E, 269m, 1.iii.1968, A.C.Kemp, NEUR11852; 1♀, same locality and collector, 9.ii.1969, NEUR11851; 1♀, same locality and collector, 15.ii.1969, NEUR11850; 1♀, same locality and collector, 11.iii.1969, NEUR11849; 1♀, same locality, 26.iv.1969, J.H.Potgieter, A.Strydom, TMSA00890 (TMSA); 1♀, same locality, 1.ii.1987, A.J.van Rensburg, NEUR11872; 1♀, same locality, 5.ii.1988, M.D.Picker, NEUR11831; 1♀, Crocodile Bridge, 25°21’S 31°54’E, 12.iv.1969, A.Braack, NEUR11848; 1♀, Skukuza, 24°59’41”S 31°36’49”E, 17.xii.1970, 300m, A.Braack, NEUR11846; 1♀, same locality and collector, 7.i.1972, NEUR11855; 1♀, same locality and collector, 9.iii.1972, NEUR11856; 1♀, same locality, 14.i.1985, M.W.Mansell, NEUR11817; 1♀, same locality, 4.iv.1990, L.E.O.Braack, NEUR05636; 1♀, same locality, 26.xi.1993, M.Krüger, TMSA00891 (TMSA); 1♀, same locality, 7.iv.1999, H.Stapelberg, NEUR11871; 5♂ 3♀, Sabi-Sand Rivers Junction, 24°57’28”S 31°42’30”E, 238m, 20.i.1984, hand-netted, M.W.Mansell, NEUR00409; 1♀, Shirombe Pan, 22°44’S 31°24’E, 19.xii.1970, A.&H.Braack, NEUR11859; 1♀, Pumbe Picket, 22°37’S 31°16’E, 7.iv.1972, A.Braack, NEUR11857; 1♀, Pafuri, 22°25’52”S 31°12’45”E, 216m, 20.xi.1973, J.van Reenen, TMSA00888 (TMSA); 1♀, same locality, 15.i.1982, R.M.Miller, NEUR11838; 1♂, same locality, 27.xii.1983, L.E.O.Braack, NEUR00320; 1♀, same locality and collector, 25.iv.1984, NEUR00327; 1♀, Nyandu Sandveld, 22°39’S 31°21’E, 25.xi.1973, A.&H. Braack, NEUR11845; 1♀, Bobomene, 22°25’45”S 31°12’31”E, 124m, 17.iii.1974, A.Braack, NEUR11847; 1♀, same locality, 29.i.1984, M.W.Mansell, NEUR00267; 1♀, Magaludzo, 22°30’45”S 31°05’09”E, 317m, 25.xi.1969, A.Braack, NEUR11858; 1♀, same locality, 1.i.1970, A.&H.Braack, NEUR11844; 1♀, Tambotie Camp, 24°27’16”S 31°24’21”E, 426m, 20.iv.2017, H. de Klerk, NEUR12444. BOTSWANA: 1♀, Chobe Safari Lodge, 17°48’21”S 25°08’46”E, 940m, 1.iv.1988, A.Kvist, NEUR11843; 1♂ 1♀, Moremi Gorge, 22°37’S 27°26’E, 23.xii.2000, A.J.Gardiner, NEUR08976. CONGO: 1♀, Brazzaville, 04°16’S 15°17’E, 15.iii.1965, L.Detaille, NEUR11903. NAMIBIA: 1♀, Buffalo Camp, 18°07’06”S 21°40’43”E, 29.ii.1992, R.G.Oberprieler, NEUR11819; 1♂, Katima Mulilo, 17°30’13”S 24°16’30”E, 950m, 20.xii.2002, H.Bodenstein, NEUR06068.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 361 TANZANIA: 1♀, Moba, 07°02’22”S 29°45’58”E, 782m, 15.iv.1954, H.Bomans, NEUR11902. ZIMBABWE: 1♀, Kariba Campsite, 16°31’S 28°50’E, 27.xii.1975, M.W.Mansell, NEUR11853; 1♀, Tashinga Camp, 16°48’48”S 28°26’26”E, 8.iv.1988, A.J.Gardiner, NEUR01298; 1♀, Matsheumhlope, Bulawayo, 20°10’S 28°43’E, 11.ii.2000, A.J.Gardiner, NEUR05775. (All in SANC except where otherwise indicated).

FIGURES 41–43. Syngenes medialis. 41, labial palp with palpimacula; 42, spermatheca NEUR11818; 43, living female specimen NEUR12444. Photo: H. de Klerk©.

Syngenes scholtzi sp. nov. Figs 44–65, 67, 71, 72.

Etymology. Named for Clarke H. Scholtz, Professor and Head of the Department of Entomology, University of Pretoria, for years of support and encouragement. Also, in recognition of his significant contribution to Entomology as a teacher, mentor and prolific scientific writer in the field of Scarabaeoidea taxonomy and biology, and especially for his contributions to the Lacewing project. Description. Based on male holotype, 2 male and 17 female paratypes. Diagnosis. Medium-sized black antlions with yellow markings (Figs 44–47). Antennae long, clavate, narrowly annulated with black and yellow. Prothorax with black lateral margins. Abdomen mainly black, with a chevron-like pattern on T2 and T3 (Fig. 51). Wings narrow relative to other Syngenes species, with posterior margin of forewings slightly undulating; alternating pale and light brown markings on wing veins, with a distinct curved mark in distal region of forewing, often with one large spot at posterior margin of forewing (11 of 20 specimens). Forewing with characteristic irregular biaereolate cells or bifurcate veins in the costal area, biaereolate cells extending from before Rs to pterostigma. Legs with relatively slender strongly curved tibial spurs bearing a very slight flange on inner surface (Figs 49, 50).

362 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL TABLE 3. Measurements (mm) of Syngenes scholtzi. Figure above is the mean, size range below. mm Antenna length Head width Forewing Forewing Hind wing length Hind wing width Body length length width Holotype 9.1 4.2 42.4 10.7 37.9 9.3 41.6 Males 8.9 4.1 43.6 11.0 39.1 9.4 39.6 n= 2 8.7–9.1 3.7–4.4 41.3–47.2 10.2–12.0 36.8–42.6 9.0–10.0 37.0–41.6 Females 10.2 4.3 44.7 11.3 40.4 9.9 37.4 n =17 9.1–10.9 3.9–4.8 40.3–49.8 9.9–12.3 36.4–42.6 8.7–10.6 32.0–42.5

FIGURES 44–45. Syngenes scholtzi n.sp. 44, Holotype male, NEUR12250 (forewing length 43mm); 45, female paratype, NEUR11899 (forewing length 48mm).

Head: wider than prothorax, vertex slightly raised, rounded. Antennae long, clavate, longer than twice head width, toruli less than scape diameter apart, scape almost touching eyes, scape, pedicel yellow with a brown spot on dorsal surface, sparse long white setae ventrally. Flagellomeres short, annulated with black and yellow, but yellow ventrally in proximal region, covered in short black setae. Eyes large, greater than hemispherical, sparse ocular setae usually present. Maxillary and labial palps short, much less than head width, yellow, terminal labial palpomere spindle-shaped with long acute apex, palpimacula slit-shaped (Fig. 57). Frons and vertex black with short white recumbent setae, vertex with two large laterally elongated embossed orange spots, often two smaller spots in between, also with irregular black and yellow markings, occiput with three distinct embossed orange spots. Face below antennae, clypeus, genae and labrum uniformly yellow, with long, sparse, pale setae.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 363 FIGURES 46–47. Syngenes scholtzi. Holotype male, NEUR12250, living specimen, Verwater Pass, Tswalu Kalahari Reserve. Photos: H. de Klerk©.

Thorax: yellow, mostly covered with heavy black markings, sparsely pilose. Prothorax slightly longer than wide, tapering anteriorly, bearing sparse long white and black forwardly-curved setae laterally, short black bristles present along anterior raised margin, sides of pronotum uniformly black, two curved black streaks on each side of midline at posterior margin, central pronotum with distinct dots and circular black mark centrally, posterior margin with long curved black setae. Mesothorax: mesoprescutum yellow, black anteriorly, a large diffuse black mark on either side of midline, anterior margin with long curved black setae; mesoscutum largely black (not striped) on either side above wing bases and sparse black setae; mesoscutellum yellow with two black central stripes and two larger diffuse black spots posterior to central stripes. Metathorax: metaprescutum mostly black with pale central mark, metanotum with two velvety spots surrounded by yellow on each side, black laterally above wing bases, long curved white setae laterally; metascutum black with pale central mark, sparse long white setae laterally. Pleurites below wings largely black, sternites mainly black, dense long white setae present on pleurites and sternites. Wings: forewings longer, broader than hind wings, apices sub-acute, membrane hyaline, veins with alternating sections of pale and black, bearing short curved sparse black setae. Forewings broad with slightly undulating posterior margin, apices rounded with sub-acute tip, with distinct curved mark in distal part of forewings, usually with one spot at posterior margin (11 of 17); axillary protuberance at base of forewing orange with long white and black setae. Costa alternating pale and black with tuft of short black bristles at base, costal area narrow at base, broadening then narrowing slightly towards pterostigma. Veins in costal area irregularly biaereolate ranging from 0–9 cells and usually 2 (0–4) bifurcate veins before Rs, becoming irregularly biareolate distally beyond Rs. Pterostigma hardly discernible. Apical margins of wings with densely arranged narrowly forked veins Hypostigmatic cell elongate. Sc and R with yellow and black chevron-like pattern extending onto membrane

364 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL between Sc and R, Rs arises well beyond CuA fork, with Rs2 at or beyond first fork of Rs, 9–12 presectoral veins. Mp2 (oblique vein) arising just beyond CuA fork, CuP arises close beyond basal crossvein, curving downwards to unite with A1, A2 fused basally with A1, then curving sharply towards A3 (crossvein-like). Membrane at posterior base with long soft white setae, posterior margin with dense short white and black setae. Hind wings narrower than forewings, lanceolate, with faint black streak in apico-posterior region, axillary protuberance pale yellow with long dense white setae. Costa mainly yellow with tuft of long white recumbent setae at base, short dense curved black setae along length, costal area uniareolate, veins unbranched, Sc and R marked with black and white, membrane unmarked. Pterostigma pale, hardly discernible. Hypostigmatic cell long. Usually 7, rarely 6–8 presectoral crossveins. Rs arises beyond Mp2 fork, Cu fused with posterior branch of Mp2 fork; anterior banksian line visible in distal portion of wing. Pilula axillaris in males conspicuous with densely packed short brown recumbent setae, females with long white setae in this position.

FIGURES 48–51. Syngenes scholtzi. 48, base of forewing showing biaereolate cells before Rs; 49, foreleg tibial spurs and facial brush; 50, hind leg tibial spurs; 51, abdomen dorsal showing pattern.

Legs: forelegs strongly developed with short stout spines. Coxa, trochanter yellow with long white pubescence. Femur yellow with two brown marks separated by yellow dorsally, femur not as stout as in S. longicornis, two femoral sense hairs present near base, ventral surface with two rows of strong short black bristles accommodating the tibia on closure, long white setae laterally, black towards apex, with pad of dense brush-like bristles antero- laterally. Tibia slender, yellow with four black stripes at femoral articulation and two black annulations proximally; lateral surfaces with long white setae, black setae dorsally, with dense pad of short black setae ventrally that fits

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 365 between the two rows of black bristles on femur. A dense facial brush of short golden pubescence present on interior apical surface, extremity of tibia black. Tibial spurs (Figs 49, 50) curved with low flange, not as prominent as S. longicornis. Tarsus with TA5 longer than TA1–TA4 combined, TA1 pale, Ta2– TA4 black, covered in short black bristles, TA5 long, yellow, black apically, covered with short black bristles; preapical claws dark reddish- brown, slender, evenly curved. Middle legs shorter, more slender than forelegs, entirely yellow. Coxa, trochanter shorter than in foreleg, yellow with long white pubescence. Femur yellow with one sensory seta proximally, two rows of black bristles ventrally as in foreleg, with brown patch dorsally, covered with long white setae and sparse patch of long black bristles dorso-apically. Tibia, tarsus as in forelegs but lacking facial brush, preapical claws distinctly shorter stouter strongly curved. Hind legs long slender, yellow, lacking femoral sense hair. Coxa, trochanter as in middle leg; femur with brown lateral stripe and brown patch dorsally, dorsal surface with short black setae, laterally with long black curved setae, long white setae proximally, ventral surface with two rows of short black bristles. Tibia long, slender, with four black stripes at articulation, brown annulation distally, with row of black setae dorsally, tibial spurs slender evenly curved. Tarsi and preapical claws as in middle leg.

FIGURES 52–53. Syngenes scholtzi. Male terminalia. 52, dorsal; 53, ventral.

Abdomen: largely black with yellow markings imparting a chevron-like pattern on T3 and T4 (Fig. 51), tergites with very sparse short black setae, sternites yellow with sparse short white setae but sternites 1–2 with long soft dense white pubescence. T1 yellow, black centrally, T2 yellow laterally black centrally. Male (Figs 52–61) with black hair-pencils present between T6 and T7. T9 yellow divided dorsally, S9 black short rounded with pale central stripe and long black posteriorly directed setae Ectoprocts yellow with long postventral lobe, dorsal margin of

366 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL ectoprocts with two black stripes, postventral lobes with long anteriorly-curved white and black setae becoming longer posteriorly. Gonarcus (Figs 58–61) bears a prominent curved beak-shaped Mu; Pa (Figs 58–61) sclerotized, articulating with Gs, terminating in upwardly curved sharply pointed extremities. Female (Figs 62–65): T9 divided dorsally, Ga digitiform with long pale setae, Gl clavate bearing long stout, slightly curved black spines, Epr rounded, bearing slender, slightly curved setae. Spermatheca (Fig. 65) sclerotized proximally becoming slender, coiled, tapering distally, with fine setae along distal extremity.

FIGURES 54–57. Syngenes scholtzi. Male terminalia: 54, lateral showing hair pencils; 55, dorsal showing hair pencils; 56, hair pencil detail. 57, Labial palp with palpimacula.

Larva. Unknown. Distribution. Namibia and Northern Cape Province of South Africa. Comments. This species is distinguished from its congeners by the relatively narrower wings and predominantly black body coloration, prothorax with uniformly black sides and the characteristic markings on T2 and T3. It is confined to Namibia and the Northern Cape Province of South Africa, which are drier than other areas in which Syngenes species occur. Type material examined. SOUTH AFRICA, Northern Cape Province: Holotype ♂, Verwater Pass, Tswalu Kalahari Reserve, 27°19’41”S 22°31’22’E, 1285m, 10.i.2018, M.W.Mansell, J.B.Ball, H.de Klerk, NEUR12250 (SANC). Paratypes: SOUTH AFRICA, Northern Cape Province: 2♀, Groblershoop, 28°53’44”S 21°59’04”E, 868m, 1.iv.1980, C.G.E.Moolman, NEUR11821; 1♀, same locality and collector, 8.iv.1982, NEUR11822; 1♀, Molopo Motel, 26°55’54”S 20°39’44”E, 890m, 25.x.1990, J.H.Hoffmann, V.C.Moran, NEUR05727; 1♀,

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 367 Olifantshoek, 27°55’40”S 22°44’08”E, 1250m, 2.ii.1991, M.W.Mansell, NEUR11825; 1♀, Driekop Nature Reserve, Kenhardt, 29°24’23”S 21°06’18”E, 850m, 19.ii.1991, M.W.Mansell, NEUR11823.

FIGURES 58–61. Syngenes scholtzi. Male gonarcus and parameres complex: 58, lateral; 59, dorsal; 60, ventral; 61, caudal.

NAMIBIA, 1 ♂, Ameib Ranch, 21°48’S 15°39’E, 1000m, 7.xii.1988, R.B.Miller, NEUR02168; 1♀, same locality, 10.iii.1987, R.G.Oberprieler, NEUR00978; 1♀, same locality, 22.ii.1988, M.W.Mansell, H.Rausch, NEUR01061; 1♂, Ongombeanavita, 21°31’S 16°32’E, 14.i.1956, F.Gaerdes, NEUR11827; 1♀, Valencia Ranch, 23°08’38”S 16°28’42”E, 1627m, 15.ii.1970, A.C.Kemp, NEUR11824; 1♀, Helmeringhausen, 25°53’S 16°49’E, 1400m, 5.iii.1987, R.G.Oberprieler, NEUR11899; 1♀, same locality, 17.ii.1988, M.W.Mansell, NEUR01005; 1♀, Ugab River near Onverwag, 19.iv.1991, E.Holm, S.Gussmann, NEUR11826; 1♀, Usakos, 22°00’15”S 15°35’08”E, 870m, 21.ii.1988, M.W.Mansell, NEUR01039; 1♀, Satco Farm, 27°55’S 18°43’E, 900m, 25.ii.1988, M.W.Mansell, NEUR01114; 1♀, Uis Mine, 21°13’S 14°52’E, 850m, 1.iii.1995, P.Bayliss NEUR02013; 1♀, Namseb Ranch, Maltahöhe, 24°45’25”S 16°54’07”E, 1388m, 21.iii.2017, M.W.Mansell, C.H.Scholtz, H.de Klerk, NEUR12459. All in SANC.

Syngenes inquinatus (Gerstaecker, 1885) Figs 73–75

Acanthaclisis inquinata Gerstaecker, 1885: 11. 2556. Syngenes inquinatus (Gerstaecker): Navás 1912: 166. 549. Syngenes longicornis inquinatus (Gerstaecker): Handschin 1963: 224. 2832. Type Depository: EMAU. Holotype: female (badly damaged).

Diagnosis. The holotype (Figs 73–75) is badly damaged and discolored, most of the abdomen and antennae are missing. Forewings heavily marked with an apical streak and large spots at the posterior margins. Costal area with biaereolate cells from before origin of Rs, one in right wing, two in left forewing and a series of biaereolate cells from Rs to pterostigma.

FIGURES 62–64. Syngenes scholtzi. Female terminalia: 62, lateral; 63, ventral; 64, dorsal.

Type material examined. Holotype ♀, labelled “inquinatus Gerst.* Congo Stdgr. (Staudiger)”/ “Zool. Mus. Greifswald II 27416”. (EMAU). Distribution. Congo. Comments. Syngenes inquinatus is distinguished from S. debilis by its larger size, more heavily marked wings and fewer biaereolate cells before Rs, and from S. medialis by the biaereolate cells from before Rs and an unbroken series of biaereolate cells from Rs2 to pterostigma. Syngenes maritimus is very similar to this species, but is separated primarily by the disjunct distribution of the two populations.

FIGURES 66–68. Syngenes species. Comparative lateral gonarcus and parameres complex: 66, S. longicornis; 67, S. scholtzi; 68, S. medialis.

FIGURE 72. Recorded distribution of Syngenes species in southern Africa. Map: C. Deschodt.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 371 FIGURES 73–75. Acanthaclisis inquinatus Gerstaecker (Syngenes inquinatus). Female Holotype: 73, habitus 74, forewing details, 75, Associated labels. Photos: P. Michalik (EMAU).

Syngenes debilis (Gerstaecker, 1888) Figs 76–79

Acanthaclisis debilis: 100. 2558. Syngenes debilis (Gerstaecker): Kolbe 1897: 16. 3434. Type depository: EMAU. Holotype: male. Type locality: Lagos (06°27’N 03°28’E), Nigeria.

Diagnosis. A small species, forewings 38–40 mm, hind wing 33–35 mm (Gerstaecker 1888). Wings without spots, but forewing with faint curved streak in apical third. Costal area with biaereolate cells from near base in right forewing, with fewer biaereolate cells before Rs in left forewing, the biaereolate cells regular and uninterrupted in both wings from Rs to pterostigma. The abdominal pattern and the evenly curved tibial spurs “Schiensporen bogig gekrummt” (Gerstaecker, 1888) and its West African distribution distinguish this species from the East African S. longicornis. It is distinguished from S. inquinatus by its apparent smaller size and more numerous biaereolate cells before Rs, but this separation is also tentative, pending further specimens. Type material examined. Holotype ♂, labelled: “debilis Gerst* Lagos, Krchdf” [Kricheldorff]/ “Zool. Mus. Greifswald II 274150 ”. (EMAU).

372 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL Distribution: Benin, Côte d’Ivoire, Cape Verde islands, Ghana, Mali, Nigeria, Senegal and Togo (Prost 1998). Comments. According to details of the type specimens (Fig. 78), there are two male syntypes in EMAU. Although Gerstaecker (1888) did not state the number of specimens that he studied, it is clear from the description that measurements are only provided for one specimen, which is depicted in Fig.76, and this should be regarded as the holotype of Acanthaclisis debilis. The original label also bears an asterisk next to the name, which is indicative of Gerstaecker’s designation of a type specimen. In the original description, Gerstaecker (1888) did not refer to his earlier species, S. inquinatus or attempt to make any comparison with this species.

FIGURES 76–79. Acanthacisis debilis Gerstaecker (Syngenes debilis). Male Syntype: 76, habitus; 77, forewing details; 78 & 79, associated labels. Photos: P. Michalik (EMAU).

Syngenes maritimus (Needham, 1913) Figs 80–87.

Acanthaclisis maritimus Needham, 1913: 244. 4426. Ohm & Hölzel 1995: 8. 8376 (Incorrect synonymy with S. longicornis). Syngenes maritimus (Needham): Stange 2004: 360. 11168.

Diagnosis. Characterized by biaereolate cells from Rs to pterostigma, with the bottom row of cells (along Sc) being regular and evenly spaced. This sets it apart from its African congeners where the cells are more irregular and not as evenly spaced. There are a variable number of biaereolate cells (0–5) before Rs. This appears to be a variable character in S. maritimus. It can be distinguished from S. medialis by the number of biaereolate cells immediately after Rs, well before Rs2 that are usually lacking in S. medialis. The evenly curved spurs clearly distinguish it from S. longicornis with which it was synonymized by Ohm & Hölzel (1995), without substantiation.

FIGURES 80–81. Syngenes maritimus. 80, male Syntype; 81, female Syntype. Photos: B. Price (BMNH).

Distribution. Aldabra Island and Madagascar. Comments. Needham (1913) recorded four specimens in the original description, all from Aldabra Island, Seychelles, collected by J.C.F. Fryer in 1908. Only two of the specimens remain in BMNH. The other two, including the specimen photographed by Needham (1913, Fig. 2) have been on loan since 2001 and possibly lost.

374 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL Photographs of the remaining two, a male with terminalia detached in a vial with the specimen, and a female are available (Figs 80, 81). In the absence of the specimen photographed by Needham, the male depicted in Fig. 80, and labelled as below, is designated as lectotype of Syngenes maritimus Needham. Type material examined. SEYCHELLES, Aldabra Island. Photographs. 1♂ Syntype, Syngenes maritimus Needham (Fig. 80), labelled “Syn-type (round label with blue border) / Seychelle Islands, Percy Sladen Trust Expedition, 1913–170 / NHMUK010288440” + Barcode icon. Male terminalia in separate vial (BMNH); 1♀ Syntype Syngenes maritimus Needham (Fig. 81), labelled “Syn-type (round label with blue border) / Aldabra, 1908, J.C.Fryer / Seychelle Islands, Percy Sladen Trust Expedition (remainder obscured) / NHMUK010288439” + Barcode icon. (BMNH).

FIGURES 82, 82a, 83, 83a. Syngenes dolichocercus. 82, female Holotype; 82a, associated labels; 83, male Paratype; 83a, associated labels. Photos: A. Mantilleri (NMHN).

Additional material examined. MADAGASCAR. 2♀ Madagascar, Vadon, no further data (specimens in poor condition) (MRAC); 1♀ Madagascar, Sud, Schouteden, no further data (MRAC); 1♂ Fort Dauphin 5 km W, 25°02’56”S 46°57’39”E, 25m, 21.iv.1991, A.Freidberg, F.Kaplan, NEUR11839 (terminalia defective) (SANC).

Syngenes dolichocercus Navás, 1914: 91. 606. Syngenes dolichocerus (!): Ábrahám & Doboz 2011: 114 (incorrect spelling). 15015. Type Depository: MNHN. Holotype: female. Type locality: Diego Suarez (12°25’S 49°20’E), Madagascar.

Diagnosis. The female holotype (Fig. 82) has a curved streak in the apical third of the forewings and an indistinct mark at the anastomosis of CuA2 and A1 at the posterior margin. The costal area has numerous (>8) biaereolate cells from before the origin of Rs and an unbroken series of biaereolate cells from Rs to pterostigma. The abdomen is distinctly banded. Distribution. Madagascar.

FIGURES 84–87. Syngenes species. Comparative forewing wing venation of types: 84, S. maritimus, female Syntype; 85, S. maritimus, male Syntype; 86, Syngenes dolichocercus, female Holotype; 87, Syngenes dolichocercus, male Paratype. Photos: B. Price (BMNH) and A. Mantilleri (NMHN).

Comments: This species is very similar to S. maritimus but the holotype female has 9 biaereolate cells before Rs in each forewing, apparently distinguishing it from S. maritimus. Navás (1914) based the description of S. dolichocercus on two specimens: a female (Fig. 82) labeled: “Museum Paris Madagascar Diego-Saurez [12°25’S 49°20’E], G. Grandidier, 1910 / Syngenes dolichocercus ♀, Nav. Navás, S.J. det./ TYPE” (red label) (Fig. 82a); and a male (Fig. 83) labeled: “Museum Paris, Madagascar, A. Grandidier, 1867 / Syngenes dolichocercus Nav. P.

376 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL Navas, S.J. det.” (Fig. 83a). These two specimens are also labelled: LECTOTYPE (red label), Syngenes dolichocerus Navas, 1914, Lectotype, J. Legrand, Dec, 1992 (handwritten) (Fig. 82a) and LECTOTYPE (red label), Syngenes dolichocercus Navas, 1914, Lectotype, design. Prost 2001 (handwritten) (Fig. 83a), respectively. The female specimen that is labelled “TYPE” must be regarded as the holotype as intended by Navás (1914), also by his description “area radiali partim ante sectorum biaereolata” It is also the specimen specifically listed in the description of the species. The male specimen, mentioned in the original description, differs significantly from the female holotype by having fewer biaereolate cells before Rs and is clearly the same as specimens described by Needham (1911) from Aldabra Island as S. maritimus. If S. dolichocercus and S. maritimus are to be regarded as separate species then the two specimens in Navás’s (1914) type series are not conspecific. If further series of specimens reveal a continuum from fewer to more biaereolate cells in the costal areas, then S. dolichocercus would require synonymization with S. maritimus.

FIGURES 88–89. Jaya alluaudi. 88, habitus male; 89, female.

Furthermore, a specimen in MRAC labeled as a cotype of S. dolichocercus by Navás (1925) has no cotype status as it was designated by Navás only in 1925, eleven years after the original description S. dolichocercus. It is also from the African mainland (Kinda, DRC), and is the same as S. medialis.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 377 Type material examined. Holotype ♀, photograph, as documented above; paratype ♂, photograph, as documented above. Additional material examined. 1♀ Madagascar, Vadon, no further data (specimen in poor condition).

Jaya alluaudi comb. nov. Figs 88–93

Acanthaclisis alluaudi van der Weele, 1909: 62. 423. Sogra alluaudi (van der Weele): Navás 1912: 73. 557. Centroclisis alluaudi (van der Weele): Stange 2004: 343. 11168. Syngenes alluaudi (van der Weele): Ábrahám & Dobosz 2011: 114. 15015.

FIGURES 90–93. Jaya alluaudi. 90, right forewing; 91, forewing details; 92, foreleg spurs; 93, hind leg spurs.

Comments. This species, from Madagascar, has now been associated with three of the five valid genera of

378 · Zootaxa 4497 (3) © 2018 Magnolia Press MANSELL Acanthaclisisini in the Afrotropical Region, after its original placement in Acanthaclisis Rambur. It differs from Acanthaclisis by the biareolate costal cells only being present after the origin of Rs, from Syngenes by the costal cells being uniaereolate and not bifurcated before Rs and by the regular pattern of these cells. It is separated from Centroclisis by the evenly curved tibial spurs (right-angular in Centroclisis); and from Fadrina by the biaereolate cells present only from Rs (from close to base in Fadrina). Ábrahám & Dobosz (2011) erroneously transferred this taxon to Syngenes without any discussion or justification. It clearly does not belong in Syngenes and is consequently transferred here to Jaya based on the above characters (Figs 88–93). This species will be more fully described and illustrated in an upcoming revision of the genus Jaya, but figures are provided here to show the major characters that relate it to Jaya.

Acknowledgements

Persons who provided the specimens listed above are thanked for their invaluable contribution to this study. H. de Klerk and C. Deschodt (University of Pretoria) kindly provided Figures 43, 46, 47 and 72 respectively. J. Constant (IRSN), B. Price (BMNH), A. Mantilleri (NMHN), P. Michalik (EMAU), took photographs of the type specimens of S. longicornis, S. maritimus, S. dolichocercus, and S. debilis and S. inquinatus respectively. The National Parks Board of South Africa is acknowledged for permits enabling lacewing research in the Kruger National Park. A., H. & L.E.O. Braack are thanked for their contributions of material while in the employ of the Park. The South African Biodiversity Information Facility (SABIF) and the Global Biodiversity Information Facility (GBIF) are acknowledged for supporting the initial databasing initiatives of the Afrotropical Lacewing project. Special thanks are due to the JRS Biodiversity Foundation for support, which has greatly facilitated and enhanced our ongoing lacewing studies and databasing of the lacewings of southern Africa, and particularly for facilitating the acquisition of the Leica Photomicroscope, associated software and computer.

References

Ábrahám, L. & Doboz, R. (2011) Contribution to the ant-lion and owl-fly fauna of Madagascar with description of new taxa (Neuroptera: Myrmeleontidae, Ascalaphidae). Natura Somogyiensis, 19, 109–138. [15015] Banks, N. (1913) Notes on African Myrmeleonidae. Journal of the New York Entomological Society, 21, 149–157. [57] Banks, N. (1920) Neuroptera, Panorpata, and Trichoptera collected by the American Museum Congo Expedition, with lists of the species known from the Belgian Congo. Bulletin of the American Museum of Natural History, 43, 21–33. [72] Banks, N. (1931) Some oriental neuropteroid insects. Psyche, 38, 56–70. [86] Esben–Petersen, P. (1916) Notes concerning African Myrmeleonidae I. Arkiv för Zoologi, 10 (15), 1–22. [0138] Esben–Petersen, P. (1920) Revision of some of the type-specimens of Myrmeleonidae, described by Navas and placed in the Vienna Museum. Annales de la Société Entomologique de Belgique, 60, 190–96. [146] Fraser, F.C. (1951) Notes on the Neuroptera of Madagascar. Part 2. The Myrmeleontidae and Ascalaphidae. Memoires de l'Institut Scientifique de Madagascar, 5, 347–357. [2392] Gerstaecker, A. (1885) Vier decaden von Neuropteren aus der Familie Megaloptera Burm. Mittheilungen aus dem Naturwissenschaftlichen Vereine von Neu-Vorpommern und Rügen in Greifswald, 16, 1–49. [2556] Gerstaecker, A. (1888) Weitere Beiträge zur artenkenntniss der Neuroptera Megaloptera. Mitteilungen des Naturwissenschaftlichen Vereins für Neu-Vorpommern u. Rugen in Greifswald, 19, 89–130. [2558] Gess, F.W. & Gess, S.K. (1998) Terrestrial invertebrates: Insects. In: Lubke, R.A. & de Moor, I.J. (Eds.), Field Guide to the Eastern and Southern Cape Coasts. Second Edition. University of Cape Town Press, pp. 344–354. Handschin, E. (1963) Beitrag zur Kenntnis der Neuropterenfauna von Madagascar. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 35, 211–226. [2832] Insom, E. & Terzani, F. (2017) A new species of Syngenes Kolbe, 1897 from Somalia (Neuroptera: Myrmeleontidae: Acanthaclisini). Onychium, 13, 55–62. [16257] Kimmins, D.E. (1943) New species of Arabian Myrmeleonidae. Annals and Magazine of Natural History, (11) 10, 145–156. [214] Kolbe, H.J. (1897) Die Thierwelt Ost-Afrikas und der Nachbargebiete. Wirbellose Thiere 4. Netzflügler. In: Mobius, K. (Ed.), Deutsch-Ost-Afrika. Wissenschaftliche Forschungsresultate über Land und Leute Ostafrikanischen Schutzgebietes und der Angrenzenden Länder. Dietrich Reimer, Berlin, pp. 1–42. [3434] Mansell, M.W. (2018) Antlions of southern Africa: genus Crambomorphus McLachlan, 1867, including extra-limital species (Neuroptera: Myrmeleontidae: Palparinae: Palparini). Zootaxa, 4382 (3), 465–500. https://doi.org/10.11646/zootaxa.4382.3.3.

REVISION OF SYNGENES Zootaxa 4497 (3) © 2018 Magnolia Press · 379 Mansell, M.W. & Kenyon, B. (2002) The Palpares relational database: an integrated model for lacewing research, In: Sziráki, G. (Ed.), Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology (6-9 August 2000, Budapest, Hungary). Acta Zoologica Academiae Scientarum Hungaricae, 48 (2), 185–195. Navás, L. (1909) Notas neuropterológicas. XI. Mirmeleónido nuevo de Madagascar. Butlletí de la Institució Catalana d'Història Natural, 9, 71–72. [498] Navás, L. (1912) Insectos Neurópteros nuevos o poco conocidos. Memorias de la Real Academia de Ciencias y Artes de Barcelona, 10, 135–202. [549] Navás, L. (1914) [Neuroptera nova africana.] II series. Memorie dell'Accademia Pontifica dei Nuovi Lincei, Rome (1) 32, 90– 99. [606] Navás, L. (1924) Algunos insectos del Museo de París. 2.a serie [IIa]. Brotéria (Zoológica), 21, 99–114. [775] Navás, L. (1925) Insectes du Congo Belge. Série I. Revue de Zoologie Africaines, Bruxelles, 13, 123–132. [795] Navás, L. (1935) Comunicaciones entomológicas. 18. Insectos de Madagascar. Segunda [II] serie. Revista de la [Real] Academia de Ciencias Exactas Fisico–Quimicas y Naturales de Zaragoza, 18, 42–74. [962] Needham, J.G. (1913) Neuroptera, Myrmeleonidae from the Indian Ocean. Transactions of the Linnean Society of London, Zoology, 16, 243–246. [4426] Ohm, P. & Hölzel, H. (1995) Die Neuropteren der Seychellen. Entomologisches Nachrichtenblatt, Wien (NF) 2, 3–12. 8376. Oswald, J.D. (2016) Bibliography of the Neuropterida. Version 11.0. Available from: http://lacewing.tamu.edu/Biblio/Main (Accessed 30 July 2018) Prost, A. (1998) Les Acanthaclisinae d'Afrique occidentale et centrale [Neuroptera, Myrmeleontidae]. Revue Française d'Entomologie, (N.S.) 20, 157–173. [9290] Rambur, M.P. (1842) Histoire Naturelle des Insectes, Névroptères. Librairie encyclopédique de Roret. Fain et Thunot, Paris, pp. 1–534. [5314] Stange, L.A. (2004) A systematic catalog, bibliography and classification of the world antlions (Insecta: Neuroptera: Myrmeleontidae). Memoirs of the American Entomological Institute, 74, 1–565. [11168] Stange, L.A. & Miller, R.B. (1985) A generic review of the Acanthaclisine antlions based on larvae (Neuroptera: Myrmeleontidae). Insecta Mundi, 1, 29–42. [5823] Van der Weele, H.W. (1907) Les Myrméléonides de Madagascar. Bulletin Scientifique de la France et de la Belgique, 41, 249– 278. [406] Van der Weele, H.W. (1908) Notizen über Gerstaecker's Myrmeleoniden. Notes from the Leyden Museum, 30, 57–62. [421] Van der Weele, H.W. (1909) Les Planipennia recueillis par le Prof. Voeltzkow a Madagascar et dans les iles environnantes. Bulletin Scientifique de la France et de la Belgique, 42, 61–68. [423] Walker, (1853) List of the specimens of neuropterous insects in the collection of the British Museum. Part II. (Sialides– Nemopterides). British Museum, London, pp. 193–476. [6194]