A revision of the montane Athetis aurogrisea and A. perplexa species groups (Lepidoptera, Noctuidae, Amphipyrinae)

M. Krüger

Lepidoptera Department, Transvaal Museum, N.F.I., South Africa

E-mail:

Krüger, M., 2001. A revision of the montane Athetis aurogrisea and A. perplexa species groups (Lepidoptera, Noctuidae, Amphipyrinae). Annals of the Transvaal Museum **: 000-000.

The predominantly montane Athetis aurogrisea and A. perplexa species groups are revised. With one exception (A. sordida from the eastern highlands of Zimbabwe) members of these species groups are endemic to the eastern part of South Africa, occurring from the Tsitsikama area in the easternmost Western Cape Province to the Soutpansberg Range in the Northern Province. Most species appear to occupy a narrow range, and many are known from the type locality only. The A. aurogrisea group comprises 7 species, 5 of which are described as new (A. auronitens Krüger, 2000, A. rhigognostis spec. nov., A. intricata spec. nov., A. aeneolineata spec. nov., A. aurogrisea Krüger, 2000, A. fulgens spec. nov. and A. aeneodeixis spec. nov.. The A. perplexa group comprises 10 species, 9 of which are described as new (A. assecta spec. nov., A. consocia spec. nov., A. alpina spec. nov., A. euxoa spec. nov., A. gracilis spec. nov., A. splendidula spec. nov., A. sordida spec. nov., A. collicola spec. nov., A. perplexa (Janse, 1938), and A. instrata spec. nov.).

Based on their distribution and biogeography it is suggested that these species groups represented a phylogenetically old lineage within the genus Athetis.

Keywords: Lepidoptera, Noctuidae, Amphipyrinae, Athetis, montane areas, endemics, Biogeography, southern Africa, Taxonomy.

INTRODUCTION

The genus Athetis Hübner, [1821] is one of the largest genera of amphipyrine noctuids in southern Africa, with 61 species presently recognized. In addition to a distinct savanna element in the subregion, which comprises mostly widespread species, there exists a specialized element particular to the fynbos and grassland biomes. These Athetis species occur near sea-level along the Cape coast, and at increasing altitudes further north, but are not well represented south of the barrier formed by the Limpopo.

In a recent paper, Krüger (2000) revised the Athetis species known from the kingdom of Lesotho, and drew attention to the large proportion of endemic species in this area. The present contribution forms a continuation of this study and is concerned with the predominantly montane Athetis aurogrisea and A. perplexa species groups. Members of these groups have not been recorded west of 23° E and attain their highest diversity in the central high-mountain part of their range, between Golden Gate National Park, the northern part of the Maloti Mountains of Lesotho and the Mont-aux-Sources area of the Natal Drakensberg (Maps 1, 2).

Although the A. aurogrisea and A. perplexa groups are almost entirely absent from the Western Cape, their combined distribution corresponds well to a pattern characterized by Stuckenberg (1962) as ‘montane-paleogenic’, suggesting that these groups represent ancient lineages within the genus Athetis.

MATERIALS AND METHODS

This revision is based on 332 adult specimens from the collection of the Transvaal Museum, Pretoria, South Africa (TMSA). All primary types were examined.

Genitalia dissections were made following Robinson (1976). Specimens were stained in a weak solution of ethanol and mercurochrome and slide-mounted in Euparal.

Label data of type specimens are quoted verbatim. However, the month of capture of the adult is given in Roman numerals and the name of the collector appears in brackets throughout.

On grounds of the strong overall similarity of the male and female genitalia in this group, this paper deviates from its predecessor (Krüger, 2000) in that only characters of taxonomic importance are dealt with in the descriptive part. For a general characterization of these structures I refer to the abovementioned article.

Because of the diagnostic importance of the configuration of cornuti on the vesica, the aedeagus is shown at a higher magnification than the genital capsule in Figs 28-43.

RANGE AND ALTITUDINAL DISTRIBUTION

The range shared by the A. aurogrisea and A. perplexa species groups extends in a broad band along the eastern part of South Africa, from the Tsitsikama area near the border between the Western and Eastern Cape Provinces at about 34° S to the Blouberg in Northern Province, an isolated western extension of the Soutpansberg range, situated at approximately 23°10'S. With the exception of A. sordida from the eastern highlands of Zimbabwe, a species which occupies an isolated position within the perplexa group, no representatives of either group are known to occur north of the Limpopo.

The centres of distribution and species richness of both groups are coincident, comprising the northern part of the Maloti range in Lesotho, the Golden Gate Highlands National Park in the Free State, adjoining to the north, and the Mont-aux-Sources area of the Natal Drakensberg, where several species occur sympatrically. As speciation is mostly seen as an accidental phenomenon, occurring in peripheral isolates with small gene pools, it is postulated that both species groups share a recent common ancestor and began to undergo their respective radiations in this area before extending their ranges to the north, south, and east. The hypothesis of an alpine origin of both groups is consistent with the observation that all constituent species today are confined to cold to moderately cold habitats that receive a substantial amount of rainfall.

In this central area, records are from an altitudinal range of 2,800 to 3,100 m. Towards the periphery of the species groups’ range, the recorded altitude drops to around 1,500 m in the montane grasslands of Gauteng (Suikerbosrand Nature Reserve), KwaZulu-Natal (Karkloof), the Eastern Cape (Amatola Range) and Mpumalanga (Mariepskop), and to near sea-level in the Tsitsikama area (Western Cape). With decreasing latitude and increasing temperature, the range is confined to the highest peaks in the former Transvaal, at around 2,000 m: Athetis intricata, a species endemic to the Blouberg, has not been recorded from other, more low-lying parts of the Soutpansberg massive. This phenomenon has been noted for several other, non-phytophagous groups of invertebrates identified as ‘palaeogenic elements’ by Stuckenberg (1962), and is known inter alia from other groups of geometrid moths (Krüger, 1998, 1999).

Most members of these species groups appear to be narrowly endemic and have been recorded from the type locality only. An exception to this is provided by Athetis perplexa (Janse), which enjoys a comparatively wide distribution encompassing parts of the Norhern Province, Mpumalanga, KwaZulu-Natal, the Eastern Cape Province and Swaziland.

SYSTEMATICS

Adults can be referred to the respective groups using external appearance alone. Within groups, however, differences between species are often very small, and in most cases both morphological and genital characters need to be consulted for reliable identification. This is especially necessary in cases where several species share the same number of apical cornuti on the vesica.

KEY TO THE SPECIES OF THE ATHETIS AUROGRISEA AND A. PERPLEXA GROUPS, BASED ON THE MALE GENITALIA

¹ Adults of these groups are of characteristic habitus and generally similar. Within the southern African fauna, confusion is only possible with Athetis interlata (Walker), which, however, is easily recognized (Pl. 29, Fig. 12 in Janse, 1940). The genitalia structure of this species clearly places it outside the groups dealt with in this paper.

² With one exception all members of the aurogrisea and perplexa groups are based on male holotypes. The exception is 17. A. instrata spec. nov. , whose striking genitalia are illustrated in Fig. 54. Although the female genitalia of three members of the perplexa group remain unknown, the habitus of the moth as well as distributional data make it unlikely that A. instrata is referable to any of them.

1. Moths with forewings relatively paler and longitudinal streaks clearer than below; median streak encompassing discal spots not distinct (Figs 1-11)¹. Male genitalia with vesica bearing between 1 and 4 apical cornuti (Figs 28-34). Female genitalia, where known, as in Figs 44-47

..... aurogrisea group, 2

- Moths with forewings relatively darker and indistinct longitudinal streaks; median streak encompassing discal spots distinct (Figs 12-27). Male genitalia with vesica bearing between 3 and 6 apical cornuti (Figs 35-43). Female genitalia, where known, as in Figs 48-54, structurally not different from those of aurogrisea group

..... perplexa group, 8

2.(1) Forewings broad, pale golden, with indistinct longitudinal streaks (Fig. 1). Male genitalia with a single, unusually small apical cornutus only, much smaller than thorn-like cornutus (Fig. 28). Female genitalia unknown. Lesotho (Maloti Mountains)

.....1. A. auronitens Krüger, 2000

- Forewings slightly to distinctly narrower, pale bronze, distinctly streaked longitudinally (Figs 2-11). Male genitalia with between 1 and 4 apical cornuti, similar in size to thorn-like cornutus (Figs 29-34). Female genitalia, where known, as in Figs 44-47. Eastern part of South Africa, one species in Lesotho

..... 3

3.(2) Adults as in Figs 2-3. Male genitalia (Fig. 29) with a single, large apical cornutus. Female genitalia (Fig. 44) with moderately deep antrum. Free State (Golden Gate National Park)

.....2. A. rhigognostis spec. nov.

- Adults as in Figs 4-11. Male genitalia (Figs 30-34) with 2-4 slightly smaller apical cornuti. Female genitalia (Figs 45-47) with antrum of varying depth. Eastern part of South Africa, including a further, more robust species from the Golden Gate area; one species in Lesotho

..... 4

4.(3) Small moths (Figs 4, 5). Male genitalia (Fig. 30) with 2 apical cornuti. Female genitalia (Fig. 45) with shallow antrum. Soutpansberg Range in northern South Africa, known from summit of Blouberg only; a single doubtful record from the Kruger Park (Nyandu Bush)

.....3. A. intricata spec. nov.

- Larger moths (Figs 6-11); if of similar small size, then vesica bearing 4 instead of 2 apical cornuti. Male genitalia (Figs 31-34) with 3 or 4 apical cornuti on vesica. Female genitalia (Figs 46, 47), where known, with deeper antrum. Distribution more southerly (Golden Gate, Maloti Mtns, Natal Drakensberg, Amatola range)

..... 5

5.(4) Adult male as in Fig. 6, comparatively robust for the group. Male genitalia (Fig. 31) with 3 apical cornuti. Female unknown. Natal Drakensberg, known from Mont-aux-Sources only

.....4. A. aeneolineata spec. nov.

- Adults as in Figs 7-11. Male genitalia (Figs 32-34) with 4 apical cornuti. Female genitalia, where known, as in Figs 46, 47. Free State (Golden Gate), Lesotho and Eastern Cape Province (Amatola range), not recorded from Natal Drakensberg

..... 6

6.(5) Adults as in Figs 7, 8. Male genitalia (Fig. 32) with the two most posterior microcornuti of the ventral band strongly enlarged. Female genitalia (Fig. 46) having moderately shallow antrum with convex rim. Lesotho, Maloti Mountains

.....5. A. aurogrisea Krüger, 2000

- Adults as in Figs 9-11, either more (Fig. 11) or less robust than A. aurogrisea (Figs 9, 10). Male genitalia (Figs 33, 34) with four apical cornuti; most posterior microcornuti of the ventral band not enlarged. Female genitalia, where known, with markedly deeper antrum (Fig. 47)

..... 7

7.(6) Smaller moths (Figs. 9, 10), forewing pattern distinct. Male genitalia (Fig. 33) with valvae widening distinctly apically; female genitalia with deep antrum and slender corpus bursae (Fig 47). Eastern Cape Province, Amatola range (Hogsback area)

.....6. A. fulgens spec. nov.

- Larger moths (Fig.11), forewing pattern indistinct. Male genitalia (Fig. 34) with valvae hardly widening towards apex. Female unknown. Free State, Golden Gate National Park

.....7. A. aeneodeixis spec. nov.

8.(1) Adults as in Figs 12, 13. Male genitalia (Fig. 35) with 3 apical cornuti. Female genitalia (Fig. 48) with deep antrum. Free State, Golden Gate National Park

.....8. A. assecta spec. nov.

- Adults as in Figs 14-27. Male genitalia (Figs 36-43) with 4-6 apical cornuti. Female genitalia, where known, with mostly shallower antrum (Figs 49-54). Variously distributed in eastern part of southern Africa, including Golden Gate area

..... 9

9.(8) Adult male as in Fig. 14, small. Male genitalia (Fig. 36) with 4 apical cornuti; genital capsule narrow and dorsal band of microcornuti long (upside down in Fig. 36). Female unknown. Free State, Golden Gate National Park

.....9. A. consocia spec. nov.

- Adults as in Figs 15-27, larger. Male genitalia (Figs 37-43) with 4-6 apical cornuti; genital capsule less narrow and more or less drop-shaped; dorsal band of microcornuti shorter. Female genitalia, where known, as in Figs 49-54. Variously distributed in southern Africa, but not recorded from Golden Gate National Park

..... 10

10.(9) Adults as in Figs 15, 16. Male genitalia (Fig. 37) with 4 apical cornuti; aedeagus comparatively stout, with posterior cornuti in dorsal band large and cornuti in ventral band indistinct. Female genitalia (Fig. 49) with large bursa copulatrix and moderately deep antrum. Natal Drakensberg, known from Mont-aux-Sources area only

..... 10. A. alpina spec. nov.

- Adults as in Figs 17-27. Male genitalia (Figs 38-43) with 4-6 apical cornuti, aedeagus less stout and dorsal and ventral band of microcornuti as illustrated. Female genitalia, where known, as in Figs 50-54. Variously distributed in southern Africa

..... 11

11.(10) Adults as in Figs 17, 18. Male genitalia (Fig. 38): aedeagus large, with 4 prominent apical cornuti; microcornuti of ventral band relatively large (upside down in Fig. 38). Female genitalia (Fig. 50) with deep antrum and comparatively large corpus bursae. Mpumalanga, known from Mariepskop mountain only

.....11. A. euxoa spec. nov.

- Adults as in Figs 19-27. Male genitalia (Figs 39-43) with 4-6 apical cornuti; if aedeagus of similar size and vesica bearing 4 apical cornuti, then the latter smaller. Female genitalia, where known, with shallower antrum and comparatively smaller corpus bursae (Figs 51-54). Variously distributed in eastern part of southern Africa

..... 12

12.(11) Adults as in Figs 19-21. Male genitalia (Figs 39, 40) with 4 apical cornuti. Females, where known, as in Fig. 51. Eastern Cape Province (Tsitsikama area, 1 species) and Gauteng (Suikerbosrand, 1 species)

..... 13

- Adults as in Figs 24-26. Male genitalia (Figs 42, 43) with 5 or 6 apical cornuti. Females, where known, as in Fig. 53. Variously distributed in southern Africa but not recorded from the above areas

..... 14

13.(12) Adults as in Figs 19, 20. Male genitalia (Fig. 39) with aedeagus short and tips of valvae rounded. Female genitalia (Fig. 51) with moderately deep antrum and narrow ductus bursae. Eastern Cape Province (Tsitsikama area)

..... 12. A. splendidula spec. nov.

- Adult male as in Fig. 21. Male genitalia (Fig. 40) with aedeagus longer and tips of valvae less rounded. Female unknown. Gauteng (Suikerbosrand)

.....13. A. gracilis spec. nov.

14.(12) Adults as in Figs 22-24. Male genitalia (Figs 41, 42) with 5 apical cornuti. Female genitalia, where known, as in Fig. 52

..... 15

- Adults as in Figs 25, 26. Male genitalia (Fig. 43) with 6 apical cornuti; microcornuti of dorsal band very numerous, those in ventral band few but distinct, as illustrated. Female genitalia (Fig. 53) with moderately deep antrum and slender corpus bursae. Widely distributed in eastern part of South Africa (Northern Province to Transkei)

.....16. A. perplexa (Janse)

15.(14) Adult male as in Fig. 24, small, wing pattern resembling that of other members of the group. Male genitalia (Fig. 42) with genital capsule pear-shaped, microcornuti in ventral band arranged in a single band as in other members of the group. Female unknown. KwaZulu-Natal (Karkloof)

.....15. A. collicola spec. nov.

- Adults as in Figs 22, 23, small, forewings much darker than in other members of the group. Male genitalia (Fig. 41) with genital capsule spindle-shaped, microcornuti in ventral band arranged in two arcs. Female genitalia (Fig. 52) with strongly sclerotized corpus bursae; bursal wall spinose. Eastern highlands of Zimbabwe (Vumba Mts.)

.....14. A. sordida spec. nov.

CHECKLIST

Genus Athetis Hübner, [1821]

aurogrisea group

auronitens Krüger, 2000

rhigognostis spec. nov.

intricata spec. nov.

aeneolineata spec. nov.

aurogrisea Krüger, 2000

fulgens spec. nov.

aeneodeixis spec. nov.

perplexa group

assecta spec. nov.

consocia spec. nov.

alpina spec. nov.

euxoa spec. nov.

splendidula spec. nov.

gracilis spec. nov.

sordida spec. nov.

collicola spec. nov.

perplexa (Janse, 1938) Proxenus

instrata spec. nov.

SPECIES ACCOUNTS

Genus Athetis Hübner, [1821] 1816: 209. Type species: Noctua dasychira Hübner, [1817], by subsequent designation by Hampson, 1909, but cited as ‘furvula Hübner, [1808]’. Type locality: Europe (Nye, 1975).

(For generic synonymies see Poole, 1989 and Krüger, 2000.)

I. The Athetis aurogrisea group

Diagnosis. Athetis species with glossy wings, forewing pattern formed by longitudinal streaks. Moths paler than species of the A. aurogrisea group. Females usually markedly smaller than males (see Figs 1-11).

Description: Adult (Figs 1-11). Slender-bodied, delicate moths; females, where known, smaller than males but not brachypterous. Eyes large; labial palpi recurved, somewhat rough-scaled, dark leather- to blackish-brown, 1.5 to 2 times diameter of eyes. Antennae serrated, with serrations more prominent in males, and bearing fine cilia. Forewings pale golden or bronze, intensely streaked with dark grey along venation. Reniform and claviform stigma reduced, orbicular (discal spot) white, small, enclosed in dark median streak.

Genitalia. Male (Figs 28-34). Uniform, corresponding to diagnosis given in Krüger, 2000: 141, and apparently without species group-specific characteristics. Aedeagus with numerous cornuti that may conveniently be classified in four groups: a single curved and stout cornutus near apex (the so-called thorn-like cornutus); immediately anterior of this 1-4 large, nail-like and strongly sclerotized cornuti (the so-called apical cornuti); finally, numerous microcornuti arranged in a dorsal and a ventral band. The number of apical cornuti is often diagnostic. Female (Figs 44-47). Variable in size but answering to diagnosis given in Krüger, 2000: 141-142. Bursa copulatrix pipe-shaped; ductus bursae separated into a more or less bowl-shaped antrum and a long, cylindrical anterior section, the ductus proper; both strongly sclerotized but joined by a membranous section. Corpus bursae approximately tear-shaped; bursal wall tough, finely spinose.

Biology and early stages. Unknown.

Distribution (Map 1). Entirely confined to high altitudes unlike members of the A. perplexa group, which descend to near sea level in the Tsitsikama area. However, members of both groups have a common centre of distribution encompassing the Golden Gate National Park (Free State), the northern part of the Maloti Mountains in Lesotho and the Natal Drakensberg. Outside this area isolated records are from the Northern Province (Blouberg) and Eastern Cape Province (Hogsback).

Map 1

1. Athetis auronitens Krüger, 2000, Figs 1, 28

Athetis auronitens Krüger, 2000: 148. Holotype %, LESOTHO, Maluti Ski Chalet, Themeda-Festuca alpine veld, 3000 m; 28°47'S 28°12'E, 15.-16.I.1996 (Krüger, Dombrowsky), to U.V. light (TMSA) [examined].

DIAGNOSIS. Easily recognized on account of the pale golden forewings (see Fig. 1). Within the A. aurogrisea group, this species is characterized by its derived, paler wing markings. In the male genitalia, the vesica bears a single apical cornutus, as in A. rhigognostis below, which, however, differs markedly in habitus. In the other members of the A. aurogrisea group 2 to 4 apical cornuti are present.

DESCRIPTION. Adult male (Fig. 1). Of medium size, forewing length 13-15 mm (n = 11). Forewings broad, with pale golden ground colour. Dark suffusion of interneural areas weakly developed. For details refer to Krüger, 2000: 148.

Male genitalia (Fig. 28). Valvae slender, with rounded tips and short, basally thickened harpe. Aedeagus large relative to size of genital capsule; thorn-like cornutus prominent. A single apical cornutus present, hardly larger than posterior cornuti of ventral band. Microcornuti in both dorsal and ventral band small and few in number.

PHENOLOGY. Adults have been collected in January. The female is unknown and may be partly brachypterous as in other Athetis species occurring at high altitudes.

DISTRIBUTION. Lesotho (Maloti Mountains).

MATERIAL EXAMINED. 16 % (1 dissected, TM Lep. Heter. Genitalia slide No. 13708). 16 Pretoria (TMSA).

Localities. LESOTHO: Maluti Ski Chalet (16).

2. Athetis rhigognostis spec. nov., Figs 2, 3, 29, 44

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Free State]: Golden Gate G[olden] G[ate] H[ighland] N[ational] P[ark] Survey, 28.I.-3.II.1971 (Potgieter & Snyman); TM Lep. Heter. Genitalia slide No. 13737.- (TMSA).

Paratypes (3%, 4&). [SOUTH AFRICA, Free State]: 2&, ibidem, dated 16.-25.I.1968 (Potgieter & Jones) (2&); 3%, 2&, ibidem, dated 1978-1979 (L. Vári) (1& dissected, TM Lep. Heter. Genitalia slide No. 13796).- (TMSA).

DIAGNOSIS. Moths intermediate in size between the larger Athetis aurogrisea from Lesotho and the sympatric A. aeneodeixis, and the smaller A. intricata and A. fulgens, and most closely approaching A. aeneolineata in stature (compare Figs 2, 3 and 6).-

Very similar externally to most other members of the A. aurogrisea group, and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, A. rhigognostis is characterized by the presence of a single apical cornutus only; in other, similar species their number varies between two and four. A single apical cornutus is also present in A. auronitens from Lesotho; however, this species is readily recognized by its pale golden forewing colouration (compare Figs 1-3).

DESCRIPTION. Adult (Figs 2, 3). Small, forewing length 12-14 mm (%) (n = 31)and 11-13 mm (&) (n = 33). Antennae of male serrated, with long cilia; antennae of female very finely serrated, almost filiform. Moths slightly sexually dimorphic, females having a relatively reduced wing area. Both pairs of wings glossy, pale golden suffused with grey. Forewings intensely streaked with grey along venation; a narrow, blackish streak enclosing a small, whitish discal spot extending medially across wings. Hind wings greyish with a metallic sheen; discal spots absent. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 29). Valvae elongated, narrow and somewhat angular, exceeding genital capsule in length; harpe well developed, curved. Aedeagus stout; vesica with a single large apical cornutus and a thorn-like cornutus of approximately equal size, plus a dorsal and ventral band of smaller cornuti as illustrated.

Female genitalia (Fig. 44). Antrum wide-mouthed, with convex rim. Bursa copulatrix pipe-shaped, slender, with relatively short ‘stem’. Wall of corpus bursae and base of ductus with wrinkled or crenulated texture, very finely spinose.

PHENOLOGY. Adults have been collected in January and February.

DISTRIBUTION. Golden Gate National Park, Free State, South Africa.

ADDITIONAL MATERIAL. 27% and 29&. 56 Pretoria (TMSA).

Localities. South Africa, Free State: Golden Gate National Park.

ETYMOLOGY. From Greek ÕÃ(@H, cold, and (<fFJ0H, one who knows: referring to the frequently cold conditions at the type locality, even during the flight period of this species in mid-summer.

3. Athetis intricata spec. nov., Figs 4, 5, 30, 45

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Northern Province]: Blouberg, T[rans]v[aa]l, Motlakeng, 5-6,000 ft, 6.-15.I.1955, Transv[aal] Mus[eum] Exp[edition]; TM Lep. Heter. Genitalia slide No. 13740.- (TMSA).

Paratypes (3%, 3&). [SOUTH AFRICA, Northern Province]: 3%, 2&, same data as holotype; 1&, Nyandu Bush, K[ruger] N[ational] P[ark] Survey, 3.XII.1964 (Vári & Potgieter) (dissected, TM Lep. Heter. Genitalia slide No. 13742).- (TMSA).

DIAGNOSIS. Athetis intricata is the smallest member of the A. aurogrisea group. Apart from its small size, the species is similar externally to most other members of the A. aurogrisea group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, A. intricata is unique in possessing two prominent apical cornuti. The species has an isolated distribution in the north of the former Transvaal.

DESCRIPTION. Adult (Figs 4, 5). Small, forewing length 10-12 mm (%) (n = 7) and 10-11 mm (&) (n = 6). Antennae of both sexes finely serrated, with sparse cilia; serrations and cilia very fine in females. Moths slightly sexually dimorphic, females having a relatively reduced wing area. Both pairs of wings faintly glossy, pale golden suffused with grey. Forewings delicately streaked with grey along venation; a narrow, blackish streak enclosing a small, whitish discal spot extending medially across wings. Hind wings greyish with a metallic sheen; discal spots indicated by a weak grey shade. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, pale grey, densely suffused with darker grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 30). Ring formed by tegumen and vinculum narrow, guttiform, with tegumen narrower than vinculum. Valvae long, slender and somewhat angular, equalling genital capsule in length; harpe well developed, slightly curved. Juxta large, cleft posteriorly. Aedeagus stout; vesica with two large apical cornuti and a somewhat smaller thorn-like cornutus; ventral microcornuti concentrated in a patch rather than a band (upside down in illustration).

Female genitalia (Fig. 45). Antrum wide-mouthed but shallow, not tapering anteriorly. The membranous connection to ductus bursae proper long. Bursa copulatrix pipe-shaped, slender, with long ‘stem’ and elongated corpus bursae. Wall of ductus bursae with wrinkled texture in anterior half, otherwise smooth; wall of corpus crenulated, very finely spinose.

PHENOLOGY. Adults have been collected in December and January.

DISTRIBUTION. Blouberg, Northern Province of South Africa, with a single record from the north of the Kruger National Park.

ADDITIONAL MATERIAL. 3% and 3& (1 dissected, TM Lep. Heter. Genitalia slide No. 13741). 6 Pretoria (TMSA).

Localities. SOUTH AFRICA, Northern Province: Blouberg, Motlakeng.

ETYMOLOGY. From Latin intricatus, tangled, intricate: the delicate forewing pattern peculiar to this group is particularly clearly developed in this species.

4. Athetis aeneolineata spec. nov., Figs 6, 31

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, KwaZulu-Natal]: Mont-aux-Sources, Natal National Park, 1.-3.II.1961 (A.L. Acutt); TM Lep. Heter. Genitalia slide No. 13789.- (TMSA).

DIAGNOSIS. Based on the single specimen known, Athetis aeneolineata cannot be distinguished externally from A. aurogrisea from the Maloti Mountains of Lesotho. It is characterized, however, by the presence of three apical cornuti on the vesica, whereas there are four in A. aurogrisea (compare Figs 31 and 32).

DESCRIPTION. Adult male (Fig. 6). Indistinguishable externally from A. aurogrisea Krüger, 2000, below. Forewing length 14 mm (n = 1).

Male genitalia (Fig. 31). Ring formed by tegumen and vinculum guttiform, with tegumen tapering posteriorly. Valvae elongated, surpassing genital capsule in length; harpe well developed, only slightly curved. Juxta large and rounded, cleft posteriorly. Aedeagus stout; vesica with three large apical cornuti, larger than thorn-like cornutus; ventral band of microcornuti strongly developed.

PHENOLOGY. The only specimen known was collected at the beginning of February.

DISTRIBUTION. Mont-aux-Sources area in the Natal Drakensberg, KwaZulu-Natal province of South Africa.

ETYMOLOGY. From Latin aëneus, brazen and linea, a line, referring to the forewing pattern.

5. Athetis aurogrisea Krüger, 2000, Figs 7, 8, 32, 46

Athetis aurogrisea Krüger, 2000: 150. Holotype %, LESOTHO, Maluti Ski Chalet, Themeda-Festuca alpine veld, 3000 m; 28°47'S 28°12'E, 15.I.1996 (Krüger, Dombrowsky), to U.V. light (TMSA) [examined].

DIAGNOSIS. The male is indistinguishable from that of A. aeneolineata from the Natal Drakensberg. In the male genitalia, the present species is characterized by the presence of four apical cornuti, whereas three are found in A. aeneolineata. Furthermore, the genital capsule is more broadly elliptical in A. aurogrisea (compare Figs 31 and 32).

DESCRIPTION. Adult (Figs 7, 8). Because of the extreme similarity to A. aeneolineata, the original description is cited here in full: ‘Small, forewing length 12-15 mm (%) and 11-13 mm (&). Antennae serrated, ringed with pale golden and grey. Labial palpi upcurved, slightly less than twice diameter of eyes. Abdomen clearly extending beyond hind margin in males, for a short distance only in females. Both pairs of wings glossy, broad, but forewings slightly narrower than in preceding species [i.e., A. auronitens]. Forewings pale golden, intensely streaked with dark grey along venation. A narrow, dark streak enclosing a small discal spot extending from wing base across discal cell to termen, Hind wings greyish; discal spots not visible. Cilia mixed light grey and gold. Underside of forewings dark grey, of hind wings, lighter grey, with small, grey discal spots. Vestiture of body generally grey, mixed with pale golden scales; thorax dorsally pale golden with two grey longitudinal stripes.

Male genitalia (Fig. 32). Uncus small. Ring formed by tegumen and vinculum delicate, with vinculum longer and broader than tegumen; vinculum extended to form a short, stout saccus. Transtilla well developed. Juxta large, rounded, exhibiting a deep median cleft. Valvae elongated, with well-rounded, densely setose tips; harpe with a long and curved upper process; clavus with a long series of setae. Aedeagus large relative to rest of genitalia, vesica bearing a long band of small, needle-like cornuti, preceded by a group of larger cornuti.

Female genitalia (Fig. 46). Papillae anales large. Both pairs of apophyses rod-like, moderately robust; a. anteriores slightly shorter than a. posteriores. Bursa copulatrix pipe-shaped: ductus bursae long and well sclerotized, posteriorly forming a wide, bowl-shaped antrum and joining corpus bursae near base. Corpus bursae tear-shaped, slightly longer than ductus. Bursal wall spinose, with small spicula.

PHENOLOGY. Adults have been collected in January.

DISTRIBUTION. Lesotho (Maloti Mountains).

MATERIAL EXAMINED. 52% (2 dissected, TM Lep. Heter. Genitalia slide Nos. 13709, 13801) and 6&(1 dissected, TM Lep. Heter. Genitalia slide No. 13710). 58 Pretoria (TMSA).

Localities. LESOTHO: Maluti Ski Chalet (46); Oxbow (4); Pass of Guns (3); Mahlasela Hill Pass (2); 6 km below Monteng Pass (3).

6. Athetis fulgens spec. nov., Figs 9, 10, 33, 47

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Eastern Cape Province]: Hogsback, Cape, 12.II.[19]86 (N.J. Duke).- (TMSA).

Paratypes (2%, 1&). [SOUTH AFRICA, Eastern Cape Province]: 1%, 1&, Alice, Hogsback, o.c. 2.1942 (dissected, TM Lepidoptera Heterocera Genitalia slide Nos. 13803, 13804); 1 %, Hogsback, C[ape] P[rovince], 24.-25.II.1978 (M.J. Scoble).- (TMSA).

DIAGNOSIS. Of the three members of the A. aurogrisea group with four apical cornuti on the vesica, A. fulgens is the smallest (compare Figs 7-11), and is similar in size to A. intricata (2 apical cornuti) and A. rhigognostis (1 apical cornutus). The male genitalia are further characterized by the valvae being broad apically and bearing a well-developed harpe.

DESCRIPTION. Adult (Figs 9, 10). Small, forewing length 12-13 mm (%) (n = 4) and 10 mm (&) (n = 1). Wing pattern as in other members of the group (for a general description, see Krüger, 2000: 150). Basal pale streak on forewings prominent.

Male genitalia (Fig. 33). Ring formed by tegumen plus vinculum broadly guttiform, with bell-shaped tegumen. Valvae elongated, surpassing genital capsule in length, with broad, angular tips; harpe well developed, curved. Juxta square, with poorly defined contours and an indistinct posterior cleft. Aedeagus stout; vesica with four apical cornuti in a larger and a smaller pair; microcornuti in dorsal and ventral band comparatively large and well-defined.

Female genitalia (Fig. 47). Delicate in keeping with the size of the moths. Antrum parallel-sided and comparatively deep, separated by a wide, thinly membranous area from the narrow ductus bursae. Bursal wall finely and evenly spinose.

PHENOLOGY. Adults have been collected in February.

DISTRIBUTION. Only known from Alice/Hogsback area, Eastern Cape Province of South Africa.

FURTHER MATERIAL. 1%. 1 Pretoria (TMSA).

Localities. SOUTH AFRICA, Eastern Cape Province: Hogsback.

ETYMOLOGY. From Latin fulgens, shining, referring to the glossy wings.

7. Athetis aeneodeixis spec. nov., Figs 11, 34

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Free State]: Golden Gate, G[olden] G[ate] H[ighland] N[ational] P[ark] Survey, 1978-1979 (L. Vári) (dissected, TM Lep. Heter. Genitalia slide No. 13797).- (TMSA).

Paratypes (2%). [SOUTH AFRICA, Free State]: 1%, same data as holotype; 1%, ibidem, dated 16.-25.I.1968 (Potgieter & Jones) (dissected, TM Lep. Heter. Genitalia slide No. 13798) .- (TMSA).

DIAGNOSIS. Next to the quite differently marked A. auronitens from Lesotho, A. aeneodeixis is the most robust member of the group. Similar externally and in size to A. aurogrisea but wing pattern coarser, reminiscent of members of the perplexa group (compare Figs 7-8 and 11).

DESCRIPTION. Adult male (Fig. 11). Large for the group, forewing length 13-14 mm (%) (n = 5). Antennae serrated, ciliate. Wing pattern as in other members of the group, but streaks rather widely spaced, resulting in a comparatively coarse pattern. Median streak enclosing discal spot fairly prominent.

Male genitalia (Fig. 34). Ring formed by tegumen plus vinculum elliptical. Valvae elongated as in other members of the group, gradually widening towards tips. Juxta broadly triangular. Aedeagus short and stout, vesica with three of the four subapical conuti large and of approximately equal size, the fourth smaller, of the size of the thorn-like cornutus (the latter hidden in illustration). Microcornuti in the shorter ventral band longer and fewer in number in comparison to the dorsal band.

PHENOLOGY. Adults have been collected in January and February.

DISTRIBUTION. Golden Gate National Park, Free State, South Africa.

ADDITIONAL MATERIAL. 2%. [SOUTH AFRICA, Free State]: Golden Gate (2). Abdomen missing.

ETYMOLOGY. From Latin aeneus, brazen, and Greek *,Ã>4H, a display, with reference to the colouration.

II. The Athetis perplexa group

Diagnosis. Moths darker than members of the A. aurogrisea group as a result of more intense dark irroration. Wing pattern similar but appearing less ‘tidy’. Females, where known, hardly smaller than males (Figs 12-27).

Description: Adult (Figs 12-27). Moderately robust moths; females of approximately the same size as males. Eyes large; labial palpi recurved, somewhat rough-scaled, dark leather- to blackish-brown, 1.5 to 2 times diameter of eyes. Antennae on average slightly more robust than in members of the aurogrisea group, serrated, with serrations more prominent in males, and bearing fine cilia. Forewings dark bronze, incompletely streaked with dark grey along venation. Reniform and claviform stigma reduced, orbicular (discal spot) white, small, enclosed in dark median streak.

Genitalia. Male (Figs 35-43) Uniform, corresponding to diagnosis given in Krüger, 2000: 141, and apparently without species group-specific characteristics. Aedeagus as in the previous group with numerous cornuti that may conveniently be classified in four groups: a single curved and stout cornutus near apex (the so-called thorn-like cornutus); immediately anterior of this 3-6 large, nail-like and strongly sclerotized cornuti (the so-called apical cornuti); finally, numerous microcornuti arranged in a dorsal and a ventral band. The number of apical cornuti is often diagnostic. Female (Figs 48-54). Variable in size but corresponding to diagnosis given in Krüger, 2000: 141-142. Bursa copulatrix pipe-shaped; ductus bursae separated into a more or less bowl-shaped antrum and a long, cylindrical anterior section, the ductus proper; both strongly sclerotized but joined by a membranous section. Corpus bursae approximately tear-shaped; bursal wall tough, finely spinose. Athetis instrata is characterized by the mostly strongly spinose bursal wall of any member of the A. aurogrisea and A. perplexa species group.

Biology and early stages. Unknown.

Distribution (Map 2). Although members of the perplexa group share most high-altitude habitats (2800-3100m, Golden Gate, Mont-aux-Sources, Maloti Mountains) with species belonging to the aurogrisea group, others descend to elevations around 1500 m in neighbouring provinces, with one species each occurring near sea level (A. splendidula, Tsitsikama area) and north of the Limpopo (A. sordida, eastern highlands of Zimbabwe).

8. Athetis assecta spec. nov., Figs 12, 13, 35, 48

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Free State]: Golden Gate G[olden] G[ate] H[ighland] N[ational] P[ark] Survey, 16.-25.I.1968 (Potgieter & Jones).- (TMSA).

Paratypes (3%, 4&). [SOUTH AFRICA, Free State]: 2%, 2&, same data as holotype; 1%, ibidem, dated 1978-1979 (L. Vári) (dissected, TM Lep. Heter. Genitalia slide No. 13787); 2&, ibidem, dated 28.I.-3.II.1971 (Potgieter & Snyman) (both dissected, TM Lep. Heter. Genitalia slide Nos. 13738, 13990).- (TMSA).

Map 2

DIAGNOSIS. Very similar externally to most other members of the A. perplexa group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. assecta bears three apical cornuti - in other members of the group this number varies between four and six. The female genitalia are characterized by a massive antrum and relatively distinctly spinose wall of the bursa copulatrix.

DESCRIPTION. Adult (Figs 12, 13). Forewing length 12-13 mm (both sexes, %: n = 15, &: n = 20). Antennae of male faintly serrated, with short cilia; antennae of female filiform. Females not appreciably smaller than males. Both pairs of wings glossy, pale bronze, and strongly suffused with grey in fresh specimens; worn examples appearing paler. Forewings intensely streaked with grey along venation; the blackish median streak enclosing the white discal spot well developed. Hind wings pale bronze; discal spots absent or indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 35). Tegumen bell-shaped, vinculum broadly rounded. Valvae with tips rounded; harpe almost straight. Aedeagus with vesica bearing three apical cornuti; microcornuti of dorsal and ventral band as illustrated.

Female genitalia (Fig. 48). Antrum conspicuous: wide-mouthed, deep, and hardly tapering anteriorly. Bursa copulatrix pipe-shaped as in other members of the group; wall of ductus bursae with wrinkled texture near transition to corpus, otherwise smooth. Wall of corpus bursae with wrinkled texture, very faintly but distinctly spinose.

PHENOLOGY. Adults have been collected in January and February.

DISTRIBUTION. Golden Gate National Park, Free State, South Africa. Single records are from the Monteng Pass area in Lesotho and Clove (spelt Clova on the specimen label) in KwaZulu-Natal.

ADDITIONAL MATERIAL. 11% (1 dissected, TM Lep. Heter. Genitalia slide No. 13805) and 16& (1 dissected, TM Lep. Heter. Genitalia slide No. 13800). 27 Pretoria (TMSA).

Localities. SOUTH AFRICA, Free State: Golden Gate (24). KwaZulu-Natal: Clova (2). LESOTHO: 6 km below Monteng Pass (1).

ETYMOLOGY. From Latin assector, to accompany, to follow. The present species is one of four members of the A. aureogrisea and A. perplexa groups occurring in the Golden Gate area.

9. Athetis consocia spec. nov., Figs 14, 36

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Free State]: Golden Gate, G[olden] G[ate] H[ighland] N[ational] P[ark] Survey, 16.-25.I.1968 (Potgieter & Jones); TM Lep. Heter. Genitalia slide No. 13799.- (TMSA).

DIAGNOSIS. Although comparatively small, the holotype is very similar externally to most other members of the A. perplexa group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A consocia. bears four apical cornuti, a condition shared with A. alpina, A. euxoa, A. gracilis and A. splendidula, but is distinguished by the narrow genital capsule, the tegumen : vinculum length ratio of 2:1 and rather angular valvae (compare Figs 36-40).

DESCRIPTION. Adult male (Fig. 14). Forewing length 11 mm (holotype). Antennae faintly serrated, bearing short and dense cilia. Both pairs of wings glossy, pale bronze, and suffused with grey. Forewings streaked with grey along venation; the blackish median streak enclosing the white discal spot moderately well developed. Hind wings pale bronze; discal spots indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings but very small, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 36). Genital capsule elongate. Tegumen and vinculum U-shaped, vinculum just over half length of tegumen. Valvae exceeding genital capsule in length, with broad, truncated tips; harpe curved. Juxta medium-sized, with indistinctly notched posterior margin. Aedeagus with vesica bearing four apical cornuti; cornuti of dorsal band large, those of ventral band quite small and very numerous.

PHENOLOGY. The holotype was collected in January.

DISTRIBUTION. Golden Gate National Park, Free State, South Africa.

ETYMOLOGY. From Latin consocius, a companion: one of several species occurring sympatrically in the Golden Gate area.

10. Athetis alpina spec. nov., Figs 15, 16, 37, 49

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, KwaZulu-Natal]: Mont-aux-Sources, Natal National Park, 6.-9.II.1961 (A.L. Acutt); TM Lep. Heter. Genitalia slide No. 13788.- (TMSA).

Paratypes (2&). [SOUTH AFRICA, KwaZulu-Natal]: 1&, same data as holotype (dissected, TM Lep. Heter. Genitalia slide No. 13000); 1&, ibidem, dated 14.-30.III.1961 (A.L. Acutt).- (TMSA).

DIAGNOSIS. Very similar externally to most other members of the A. perplexa group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. alpina bears four apical cornuti - a condition shared with A. consocia, A. euxoa, A. gracilis and A. splendidula. Athetis alpina may be separated from these species by the shape of the genital capsule, the stout and curved cornutus and the size and configuration of the microcornuti in the dorsal and ventral band (compare Figs 36-40). The female genitalia have the antrum comparatively shallow and the corpus bursae quite broad.

DESCRIPTION. Adult (Figs 15, 16). Forewing length 13 mm (%) (n = 1), 11-12 mm (&) (n = 2). Antennae of male holotype mostly lost but apparently faintly serrated, with short cilia; antennae of female so weakly serrated as to appear filiform. Labial palpi, especially of male , long and extending beyond base of antennae. Both pairs of wings glossy, bronze, and strongly suffused with grey. Forewings intensely streaked with grey along venation; the blackish median streak enclosing the white discal spot well developed; in addition, a slightly narrower streak extending from wing base to tornus. Hind wings pale bronze; discal spots indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots absent on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 37). Genital capsule relatively short; tegumen bell-shaped, vinculum broadly rounded. Valvae barely exceeding genital capsule in length, with well-rounded tips; harpe gently curved. Juxta equally bell-shaped. Aedeagus stout and curved; vesica bearing four apical cornuti; microcornuti of dorsal and ventral band as illustrated.

Female genitalia (Fig. 49). Antrum shallow, tapering anteriorly, the membranous connection to ductus bursae proper wide. Bursa copulatrix pipe-shaped with broad corpus; wall of ductus bursae with wrinkled texture near transition to corpus, otherwise smooth. Wall of corpus bursae spinose, especially posteriorly

PHENOLOGY. Adults have been collected later than for most other members of this group, with records from early February to late March.

DISTRIBUTION. Natal Drakensberg in the South African province of KwaZulu-Natal, only known from the Mont-aux-Sources area.

ETYMOLOGY. From Latin alpinus, Alpine, montane, referring to the habitat.

11. Athetis euxoa spec. nov., Figs 17, 18, 38, 50

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Mpumalanga]: Mariepskop, 15.-24.III.1965 (Potgieter & E. v. Son); TM Lep. Heter. Genitalia slide No. 13988.- (TMSA).

Paratypes (2%, 3&). [SOUTH AFRICA, Mpumalanga]: same data as holotype (1& dissected, TM Lep. Heter. Genitalia slide Nos. 13989).- (TMSA).

DIAGNOSIS. Very similar externally to most other members of the A. perplexa group, especially A. perplexa itself, and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. euxoa bears four apical cornuti - a condition shared with A. consocia, A. alpina, A. gracilis and A. splendidula, but is distinguished by the shape of the genital capsule, well-developed and curved harpe and the large and comparatively straight aedeagus (compare Figs 36-40). The female genitalia resemble those of other members of this group, especially A. assecta and A. alpina, but have the sides of the antrum more strongly convex.

DESCRIPTION. Adult (Figs 17, 18). Forewing length 12-13 mm (%) (n = 49), 11-14 mm (&) (n = 37). Antennae of male slender, hardly thicker than those of female; faintly serrated, with short cilia; antennae of female almost filiform. Females exhibiting some variation in size. Both pairs of wings glossy, bronze, and strongly suffused with dark grey in fresh specimens; worn examples appearing paler. Forewings streaked with grey along venation; the blackish median streak enclosing the white discal spot fairly well developed, basal streak faint. On hind wings, discal spots absent or indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots absent on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 38). Genital capsule broadly spindle-shaped; tegumen with slightly concave sides, vinculum broadly rounded, its base truncated. Valvae barely surpassing genital capsule in length, tips between rounded and truncated in shape; harpe prominent, curved. Aedeagus large, less curved than in other members of the group, with vesica bearing four apical cornuti; cornuti of dorsal and ventral band as illustrated.

Female genitalia (Fig. 50). Antrum large, with convex sides and rim. Bursa copulatrix pipe-shaped as in other members of the group; wall of ductus bursae with wrinkled texture near transition to corpus, otherwise smooth. Corpus bursae well developed, bursal wall very faintly but distinctly spinose.

PHENOLOGY. Adults have been collected in March.

DISTRIBUTION. Mariepskop Mountain in Mpumalanga Province, South Africa. It is not known at which altitude or in which vegetation type the species occurs.

ADDITIONAL MATERIAL. 46% and 34&. 80 Pretoria (TMSA).

Localities. SOUTH AFRICA, Mpumalanga: Mariepskop (80).

ETYMOLOGY. From Greek §L>@@H, well-polished, referring to the glossy wings.

12. Athetis splendidula spec. nov., Figs 19, 20, 39, 51

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Western Cape Province]: Tsitsikama, Goesabos Forestry, 1.-3.III.1980 (Scoble & Kroon).- (TMSA).

Paratypes (2%, 1&). [SOUTH AFRICA, Western Cape Province]: 1%, same data as holotype (dissected, TM Lep. Heter. Genitalia slide No. 13993); 1%, 1&, ibidem, dated 13.-22.III.1979 (Potgieter & Scoble) (dissected, TM Lep. Heter. Genitalia slide No. 13994) (&) and 17.III.1979 (Potgieter & Scoble) (%).- (TMSA).

DIAGNOSIS. Moths relatively robust for the group, most similar externally perhaps to A. euxoa and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. gracilis bears four apical cornuti as in A. consocia, A. alpina, A. euxoa and A. gracilis. The species is most similar to A. gracilis and separated from it in the male sex by the shorter aedeagus and more rounded tips to the valvae (compare Figs 39 and 40). In addition, the abdomen of the male is more slender in A. gracilis below, with different ventral sclerotizations. The female genitalia resemble those of A. alpina, although the bursa copulatrix is markedly smaller (compare Figs 49 and 51).

DESCRIPTION. Adult (Figs 19, 20). Forewing length 12-13 mm (%: n = 4; &: n = 1), the single female showing no tendency towards wing reduction. Antennae of male faintly serrated, with short cilia; antennae of female so weakly serrated as to appear filiform. Both pairs of wings glossy, pale bronze, and strongly suffused with greyish-brown in fresh specimens. Forewings streaked with grey along venation; the blackish median streak enclosing the white discal spot well developed. Hind wings pale bronze; discal spots absent or indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, pale cream, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 39). Tegumen bell-shaped, vinculum broadly rounded. Valvae with well-rounded tips; harpe weakly curved. Aedeagus short, vesica bearing four apical cornuti; microcornuti of dorsal and ventral band as indicated, large.

Female genitalia (Fig. 51). Antrum broad and comparatively shallow, rim slightly convex. Bursa copulatrix pipe-shaped, relatively small and narrow. Wall of corpus bursae very faintly spinose only.

PHENOLOGY. Adults have been collected in March.

DISTRIBUTION. Only known from the Tsitsikama area in the easternmost Western Cape Province, South Africa.

ADDITIONAL MATERIAL. 1%. 1 Pretoria (TMSA).

Localities. SOUTH AFRICA, Western Cape Province: Tsitsikama, Goesabos Forestry (1).

ETYMOLOGY. From Latin splendidulus, diminutive formed from splendidus, shining, referring to the slightly glossy wings.

13. Athetis gracilis spec. nov., Figs 21, 40

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, Gauteng]: Suikerbosrand Natuurreservaat, 15.-16.XI.1980 (D.M. Kroon).- (TMSA).

Paratypes (3%). [SOUTH AFRICA, Gauteng]: same data as holotype (1% dissected, TM Lep. Heter. Genitalia slide No. 13802).- (TMSA).

DIAGNOSIS. Very similar externally to most other members of the A. perplexa group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. gracilis bears four apical cornuti as in A. consocia, A. alpina, A. euxoa, and A. splendidula. Athetis gracilis is most similar to the latter species but separated from it by the relatively longer aedeagus and less rounded tips to the valvae (compare Figs 39 and 40). In this species the two most posterior microcornuti of the dorsal band are much enlarged and could possibly be interpreted as forming part of the apical group.

DESCRIPTION. Adult male (Fig. 21). Forewing length 11-13 mm (n = 6). Antennae of male faintly serrated, with short cilia. Both pairs of wings glossy, pale bronze, and strongly suffused with grey in fresh specimens; worn examples appearing paler. Forewings intensely streaked with grey along venation; the blackish median streak enclosing the white discal spot well developed. A pale and indistinct postmedian line of ground colour present. Hind wings pale bronze; discal spots absent. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa of hind wings. Discal spots whitish on forewings, dark grey on hind wings. Vestiture of body concolorous with wings.

Male genitalia (Fig. 40). Genital capsule pear-shaped, sides of tegumen faintly concave. Juxta slightly narrower and harpe slightly more curved than in A. gracilis above. Aedeagus longer, with vesica bearing four apical cornuti; cornuti of dorsal and ventral band as indicated.

PHENOLOGY. Adults have been collected in November, the earliest recorded time for any member of this group.

DISTRIBUTION. Suikerbosrand Nature Reserve near Heidelberg, Gauteng Province, South Africa.

ADDITIONAL MATERIAL. 2%. 2 Pretoria (TMSA).

Localities. SOUTH AFRICA, Gauteng: Suikerbosrand Nature Reserve (2).

ETYMOLOGY. From Latin gracilis, slender; the reference is to the comparatively delicate adult habitus.

14. Athetis sordida spec. nov., Figs 22, 23, 41, 52

TYPE MATERIAL. Holotype %, ZIMBABWE: Cloudlands, 9.III.[19]95 (N.J. Duke); TM Lep. Heter. Genitalia slide No. 13996.- (TMSA).

Paratypes (3&). ZIMBABWE: 1&, Aberfoyle, Honde Valley, 20.-21.IV.[19]96 (N.J. Duke); 2&, Laurenceville, Vumba, S[outhern] Rh[odesia], 6.-12.III.1964 (Vári & van Son) (1 dissected, TM Lep. Heter. Genitalia slide No. 14056).- (TMSA).

DIAGNOSIS. Together with A. consocia the smallest member of the A. perplexa group, and also the darkest (compare Figs 14 and 22, 23). In the male genitalia, the vesica of A. sordida bears five apical cornuti. The same number is present in A. collicola , but adults of that species are larger, with paler wings. Athetis sordida is also the only member of the A. perplexa group known to occur in Zimbabwe.

DESCRIPTION. Adult (Figs 22, 23). Small, forewing length 11-12 mm (both sexes, %: n = 1; &: n = 3). Antennae of male weakly serrated, with short cilia; antennae of female filiform. Both pairs of wings slightly glossy, whitish-grey, and strongly and evenly suffused with dark greyish-brown on forewings; females darker than males. Forewings weakly streaked with blackish-grey along venation in male; in females such streaking present as a series of short streaks in postmedian area only. A black median streak enclosing the white discal spot present but poorly developed. Hind wings pale whitish, lightly suffused with grey; discal spots indicated by a faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, suffused with blackish-grey over forewings, but suffusion lighter along inner margin, and in a broad band along costa of hind wings, extending along termen for a short distance. Discal spots minute on forewings, slightly better developed on hind wings, which also display an indistinct postmedian line, well developed near costa but usually not reaching anal angle. Vestiture of head and thorax dark greyish-brown, of abdomen, grey.

Male genitalia (Fig. 41). Genital capsule relatively narrow; tegumen bell-shaped, vinculum extended into a broad point. Valvae somewhat angular, only moderately widening towards apex; harpe long, distinctly curved. Juxta approximately hexagonal, indistinctly cleft posteriorly. Aedeagus with vesica bearing five apical cornuti with gradual transition in size to cornuti of dorsal band.

Female genitalia (Fig. 52). Antrum hardly tapering anteriorly, i.e. parallel-sided, with straight rim. Bursa copulatrix pipe-shaped as in other members of the group; wall of ductus bursae locally ribbed near transition to corpus, otherwise smooth. Wall of corpus bursae tough, with somewhat wrinkled texture, evenly spinose.

PHENOLOGY. Adults have been collected in March and April.

DISTRIBUTION. Vumba Mountains in the eastern highlands of Zimbabwe.

ETYMOLOGY. From Latin sordidus, dirty, referring to the blackish wings.

15. Athetis collicola spec. nov., Figs 24, 42

TYPE MATERIAL. Holotype %, [SOUTH AFRICA, KwaZulu-Natal]: Karkloof, Natal, 28.Jan[uary 19]17 (A.J.T. Janse); TM Lep. Heter. Genitalia slide No. 13995 (TMSA).

DIAGNOSIS. Very similar externally to most other members of the A. perplexa group and distinguishable with certainty only by means of genitalia dissection. In the male genitalia, the vesica of A. collicola bears five apical cornuti - the same number is present in A. sordida, adults of which are of similar size but immediately recognizable by their much darker forewings. The female is unknown.

DESCRIPTION. Adult male (Fig. 24). Forewing length 12 mm (n = 1). Antennae weakly serrated, with short cilia. The holotype, the only specimen known, is rather worn, but is very similar in habitus to other members of this group except A. sordida. For a description see for example under A. consocia and A. alpina .

Male genitalia (Fig. 42). Genital capsule distinctly pyriform; tegumen bell-shaped, vinculum much shorter, with broadly rounded base. Valvae sole-shaped, hardly widening towards apex; harpe curved. Juxta trapezoidal, posterior margin indistinctly cleft. Aedeagus with vesica bearing five apical cornuti; microcornuti of both dorsal and ventral bands well developed and distinct.

PHENOLOGY. The holotype was collected in January at an altitude of approximately 1,500 m.

DISTRIBUTION. Only known from Karkloof in the Natal Midlands.

ETYMOLOGY. From Latin collis, a hill, and colo, to inhabit: the species occurs at lower altitudes than its close relatives.

16. Athetis perplexa (Janse, 1938), Figs 25, 26, 43, 53

Proxenus perplexa Janse, 1938: 267. Holotype %, [SOUTH AFRICA, Northern Province]: Haenertsburg, I.1923 (Swinb[urne]/Stevens); Athetis perplexa Janse % type; G[enitalia preparation] 3183; Proxenus perplexa Janse Type No. 1922 (TMSA) [examined].

Paratypes (1& examined). [SOUTH AFRICA, Mpumalanga]: Kastrol Nek, I.[19]22 (A. Rob[erts]/G. v.D[a]m); Athetis perplexa Janse &type; Athetis perplexa Janse Type No. 2877 (TMSA) [examined].

DIAGNOSIS. Comparatively large and broad-winged for the group and most closely approaching A. euxoa in habitus. Athetis perplexa is the only member of its group in which the vesica of the aedeagus bears six apical cornuti - in the other species of the group this number varies between three and five. The female genitalia are characterized by having the antrum distinctly wider than the remainder of the ductus bursae.

DESCRIPTION. Adult (Figs 25, 26). Forewing length 11-13 mm (%: n = 16, &: n = 8). Both sexes variable in size, but females on average hardly smaller than males: the specimen illustrated in Fig. 26 therefore is quite small. Antennae of male weakly serrated, with short cilia; antennae of female filiform. Moths in habitus not distinguishable from A. euxoa above. For a description, see under that species and also in Janse, 1938: 267.

Male genitalia (Fig. 43). Tegumen bell-shaped with slightly concave sides, vinculum broadly rounded. Valvae sole-shaped, with rounded apex; harpe hardly curved. Juxta rhomboid, indistinctly cleft posteriorly; transtilla normal. Aedeagus with vesica bearing six apical cornuti; transition in size between the latter and the microcornuti of the dorsal band distinct.

Female genitalia (Fig. 53). Antrum strongly developed, wider than anterior section of ductus bursae, and with convex sides and rim. Ductus bursae gently tapering anteriorly, with a distinctly grooved area near transition to corpus, otherwise smooth. Wall of the tear-shaped corpus bursae finely but distinctly spinose.

PHENOLOGY. Adults have been collected from January to March in a variety of habitats including montane grassland (Malolotja, Kastrol Nek) and subtropical bush/forest (Mbabane, Mbona).

DISTRIBUTION. Athetis perplexa appears to be the most widely distributed member of the A. perplexa group and has been recorded from the Northern Province, Mpumalanga, KwaZulu-Natal and the Eastern Cape Province of South Africa, as well as from Swaziland.

MATERIAL EXAMINED. 16% (4 dissected, TM Lep. Heter. Genitalia slide Nos. 13790, 13791, 13807, 13808) and 8&(2 dissected, TM Lep. Heter. Genitalia slide Nos. 13739, 13792). 24 Pretoria (TMSA).

Localities. SOUTH AFRICA, Eastern Cape Province: Transkei, Langeni Forest (9). KwaZulu-Natal: Mbona (6); Yellowwoods/Balgowan (1). SWAZILAND: Malagwane Hill/Mbabane (4); Malolotja (4).

17. Athetis instrata spec. nov., Figs 27, 54

TYPE MATERIAL. Holotype &, [SOUTH AFRICA, Mpumalanga]: E[astern] Transvaal, Berlin, gorge-edge, 25.32 S - 30.44 E; 3.2.1987, E-Y: 2407, UV light collection, leg. Endrödy-Younga; TM Lep. Heter. Genitalia slide No. 13991- (TMSA).

Paratypes (1&). [SOUTH AFRICA, Mpumalanga]: same data as holotype.- (TMSA).

DIAGNOSIS. The male is unknown. Females are narrow-winged and characterized by dark greyish-brown forewings and are perhaps most similar externally to A. assecta (compare Figs 12, 13 and 27). In the female genitalia the ductus bursae is less heavily sclerotized than in any other member of the A. perplexa group, whereas the wall of the corpus bursae is more strongly spinose, perhaps suggesting an isolated position of this taxon within the group.

DESCRIPTION. Adult female (Fig. 27). Forewing length 11-12 mm (n = 2). Antennae very weakly serrated, appearing almost filiform. Both pairs of wings faintly glossy, pale bronze, and suffused with greyish-brown on forewings. Forewings in addition with indistinct darker streaking along venation; a blackish median streak enclosing the small white discal spot present but not conspicuous. Hind wings pale greyish-bronze, discal spots indicated by a very faint grey shade only. Cilia concolorous with respective pair of wings, i.e. paler on hind wings. Underside also glossy, whitish, densely suffused with grey over forewings and in a broad band along costa and termen of hind wings. Discal spots greyish on both pairs of wings, hind wings in addition with a faint grey postmedian line. Vestiture of head and thorax concolorous with forewings, of abdomen, concolorous with hind wings.

Female genitalia (Fig. 54). Antrum conspicuous: wide-mouthed, parallel-sided and hardly tapering anteriorly, markedly wider than anterior section of ductus bursae. The latter less heavily sclerotized than in other members of the group, widening towards the pear- rather than tear-shaped corpus bursae. Bursal wall strongly spinose.

PHENOLOGY. Adults have been collected in February.

DISTRIBUTION. Area of the Berlin gorge in Mpumalanga province, South Africa.

ETYMOLOGY. From Latin insterno (past participle instratus), to cover: the bursal wall of the female genitalia is densely covered with star-shaped sclerotizations.

ACKNOWLEDGEMENTS

I thank Mrs. L.M. Molony, University of Pretoria, and Dr D.M. Kroon, Sasolburg, for assistance with photographic work.

REFERENCES

JANSE, A. J. T., 1937-39. The Moths of South Africa, Vol. 3, Cymatophoridae, Callidulidae and Noctuidae (partim). E.P. & Commercial Printing Co., Durban [pp. 145-320 containing the genus Athetis (as Proxenus) published in 1938].

JANSE, A. J. T., 1940. The Moths of South Africa. Plates to Volumes III. John Bale, Sons & Staples, Ltd., London.

KRÜGER, M., 1998. Revision of the genus Hypotephrina Janse, 1932 (Lepidoptera: Geometridae: Ennominae). Annals of the Transvaal Museum 36(26): 351-372.

KRÜGER, M., 1999. Revision of Afrotropical Ennominae of the Drepanogynis group III: the genera Argyrophora Guenée, Pseudomaenas Prout and Microligia Warren (Lepidoptera: Geometridae: Ennominae). Annals of the Transvaal Museum 36(32): 427-496.

KRÜGER, M., 2000. The genus Athetis Hübner, [1821] in Lesotho (Lepidoptera, Noctuidae, Amphipyrinae). Annals of the Transvaal Museum 37: 139-161.

NYE, I. W. B., 1975. The generic names of moths of the world. Vol. 1, Noctuoidea (Part): Noctuidae, Agaristidae, and Nolidae. British Museum (Natural History), London. 568 pp.

POOLE, R. W., 1989. Lepidopterorum Catalogus (New Series), Fascicle 118: Noctuidae, Parts 1-3. E.J. Brill/Fauna & Flora Publications. 1134 pp.

ROBINSON, G.S., 1976. The preparation of slides of Lepidoptera genitalia with special reference to the Microlepidoptera. Entomologists’ Gazette 27: 127-132.

STUCKENBERG, B.R., 1962. The distribution of the montane palaeogenic element in the South African invertebrate fauna. Annals of the Cape Provincial Museums 2: 190-205.

GAZETTEER

South Africa

Western Cape Province

Tsitsikama, Goesabos Forestry 33°58'S 23°45'E

Eastern Cape Province

Hogsback 32°06'S 27°01'E

Langeni Forest, Transkei 31°29'S 28°28'E

Free State

Golden Gate National Park 28°31'S 28°30'E

KwaZulu-Natal

Balgowan 29°24'S 30°03'E

Clova (also Clove) 28°24'S 29°23'E

Karkloof 29°26'S 30°19'E

Mbona 30°18'S 30°22'E

Mont-aux-Sources 28°47'S 28°52'E

Yellowwoods: see Balgowan

Mpumalanga

Berlin Gorge 25°32'S 30°44'E

Kastrol Nek 25°17'S 30°19'E

Mariepskop 24°35'S 30°50'E

Gauteng

Suikerbosrand Nature Reserve 26°30'S 28°15'E

Northern Province

Blouberg, Motlakeng c. 23°10'S 29°00'E

Haenertsburg 23°56'S 29°57'E

Nyandu Bush 22°41'S 31°23'E

Lesotho

Mahlasela Hill Pass 28°50'S 28°45'E

Maluti Ski Chalet 28°47'S 28°12'E

Monteng (also Moteng) Pass 28°46'S 28°04'E

Oxbow 28°46'S 28°10'E

Pass of Guns 28°50'S 29°55'E

Swaziland

Malagwane Hill: see Mbabane

Malolotja 27°01'S 31°39'E

Mbabane 26°20'S 31°08'E

Zimbabwe

Aberfoyle, Honde Valley c. 18°30'S 32°50'E

Cloudlands - see Vumba Mountains

Laurenceville - see Vumba Mountains

Vumba Mountains c. 19°05'S 32°44'E

LEGENDS

Figs 1-8, adults. 1, Athetis auronitens %; 2-3, A. rhigognostis, %,&; 4-5, A. intricata, %,&; 6, A. aeneolineata, %; 7-8, Athetis aurogrisea, %,&. Scale bar in mm. Fig 1-4 & Fig 5-8

Figs 9-16, adults. 9-10, Athetis fulgens, %,&; 11, A. aeneodeixis, %; 12-13, A. assecta, %,&; 14, A. consocia, %; 15-16, A. alpina, %,&. Scale bar in mm. Fig 9-12 & Fig 13-16

Figs 17-24, adults. 17-18, Athetis euxoa, %,&; 19-20, A. splendidula, %,&; 21, A. gracilis, %; 22-23, A. sordida, %,&; 24, A. collicola, %. Scale bar in mm.Fig 17-20 & Fig 21-24

Figs 25-27, adults. 25-26, Athetis perplexa, %,&; 27, A. instrata, &. Scale bar in mm.Fig 25-27

Figs 28-33, male genitalia. 28, Athetis auronitens; 29, A. rhigognostis; 30; A. intricata; 31, A. aeneolineata; 32, A. aurogrisea; 33, A. fulgens. Scale bar = 0.3 mm. 

Figs 34-39, male genitalia. 34, Athetis aeneodeixis; 35, A. assecta; 36, A. consocia; 37, A. alpina; 38; A. euxoa; 39, A. splendidula. Scale bar = 0.3 mm. 

Figs 40-43, male genitalia; figs 44-46, female genitalia. 40, Athetis gracilis; 41, A. sordida; 42, A. collicola; 43, A. perplexa. Scale bar = 0.3 mm. 44, Athetis rhigognostis; 45, A. intricata; 46, A. aurogrisea. Scale bar = 0.5 mm.

Figs 47-54, female genitalia. 47, Athetis fulgens; 48, A. assecta; 49, A. alpina; 50, A. euxoa; 51, A. splendidula; 52, A. sordida; 53, A. perplexa; 54, A. instrata. Scale bar = 0.5 mm. 

Map 1. Recorded geographical distribution of A. aurogrisea group.

Map 2. Recorded geographical distribution of A. perplexa group.