Lukeniana Lehmann, Zahiri & Husemann, 2023

Lehmann, Ingo, Zahiri, Reza & Husemann, Martin, 2023, Revision of the Metarbelodes Strand, 1909 genus-group (Lepidoptera: Cossoidea: Metarbelidae) with descriptions of two new genera and 33 new species from high elevations of eastern and southern Africa, Zootaxa 5267 (1), pp. 1-106 : 11-14

Lukeniana Lehmann, Zahiri & Husemann gen. nov.

urn:lsid:zoobank.org:act:19013856-B5BB-4F78-8591-16D3F33027CA

Type Species : Metarbela obliqualinea Bethune-Baker, 1909 . The Annals and Magazine of Natural History (8)3(17): 425, by present designation .

Diagnosis. Lukeniana obliqualinea is designated herein as the type species to avoid further confusion with Metarbelodes umtaliana . This group comprises species that share two characters: CuP vein fold in the forewing is weak, not continuous, and not tubular; and a strongly sclerotized, hollow, upward bent, thorn-like process representing a ventro-distal extension of the sacculus. One species from the southern Ethiopian Plateau ( L. rajaeii sp. nov.) and one species from various localities in north-central Namibia ( L. raymondrevellii sp. nov.) have a less pronounced (smaller) thorn-like process.

Description. Head ( Fig 3a ; Fig. 2c ): Rough-scaled; long hair-like scales on fronto-clypeus; a pair of pits absent from lower fronto-clypeus, but a pair of small conical projections present on lower fronto-clypeus in both sexes (more pronounced in females); projections separated, but close together and in between two large pits behind labial palpi ( Fig. 1c ); labial palpi long, almost as long as eye diameter, rarely longer (e.g., L. mbalaensis sp. nov.), consisting of three segments; 2 nd segment longest, up to 1.5–2.3 longer than 1 st (basal) segment; apical palpomere shortest, 0.5 length of basal segment in both sexes. Labial palpi in females sometimes narrower than in males ( L. friederikebauerae sp. nov.). Male antennae bipectinate; female antennae unipectinate, sometimes slightly bipectinate towards tip. Dorsal and lateral sides of branches and flagellum scaled in both sexes (branches appearing less scaled in females). In males, tip of branches broader, sometimes slightly spoon-shaped, always densely scaled, sometimes extremely densely scaled (eg. L. mikerobertsi sp. nov.).

Thorax: Densely covered with hair-like scales (slightly finer scales in females); short scale crest on metathorax. Fore and mid legs ochre with long dense hair-like structures. Epiphyses absent in both sexes. Hindlegs with one pair of narrow tibial spurs in both sexes (shorter inner spur at least 0.5 mm long); only holotype and paratype of L. hausmanni sp. nov. with two narrow pairs of tibial spurs; upper pair smaller and shorter (inner spur 0.2 and 0.3 mm long, outer spur 0.6 and 0.9 mm). Forewing upperside essentially without scale pattern, except CuA 2 is white and brown or brown and cream-coloured in both sexes; usually not shiny, but rarely shiny towards edge of wing and termen in both sexes. Fringe scales hair-like and very long in both sexes (up to 2.1 mm). Forewing venation similar in both sexes ( Figs 9b; 9e ; 10a; 10b ) with 1A+2A forked at base; CuP obsolete, with few exceptions ( L. mbalaensis ); CuA 2 originating from hind margin of posterior cell in both sexes; CuA 1, M 3 and M 2 separate and originating from apical angle of posterior cell in both sexes; M 1 originating from distal margin of median cell, sometimes close to base of a small areole in both sexes; R 1 originating from anterior margin of median cell in both sexes; R 2 originating from anterior angle of areole and is separated or short-stalked with R 3 +R 4: (i) R 2 originating from anterior angle of areole; short-stalked with R 3 +R 4 ( L. georgeadamsoni sp. nov., L. bergsteni sp. nov., L. lenzi sp. nov.), or (ii) R 2 originating from anterior angle of areole and sharing the same basal point with R 3 +R 4 ( L. bergsteni , L. utaheidenreichae sp. nov., L. stueningi ), or (iii) R 3 +R 4 well separated from R 2 ( L. friederikebauerae , L. rajaeii ); R 5 originating from or close to posterior angle of areole in both sexes; Sc more or less parallel to R 1. In hindwing CuA 2 originating from hind margin of posterior cell; CuA 1, M 3 and M 2 originating from apical angle of posterior cell, separated; M 1 and Rs originating from apical angle of anterior cell, usually stalked in both sexes; with or without a bar from Rs to Sc+R 1; with a small vein in discocellular cell on both fore- and hindwing; fringe with very long cilia in both sexes, 1.1–2.1 mm in length. Retinaculum and frenulum absent.

Abdomen: With dense hair-like scales and abdominal tuft, usually short, not longer than one-third of abdomen, very rarely up to 40% of abdomen ( L. georgeadamsoni ). Male genitalia ( Fig. 13 ; Fig. 14e ; Fig. 11e ; Fig. 15b–d ) with tegumen and vinculum fused, forming firm, narrow ring. Uncus large, broad, one uncus lobe ca. 0.6–0.8 length of valva (lateral view), rarely less than 0.5 ( L. hausmanni ), setose ventrally and occasionally dorsally. Basal edge of uncus well developed (strongly, slightly or not bent at center). Gnathos with gnathos arms covered by many short tooth-like processes at distal ends, resembling a hand. Gnathos arms not connected by a sclerotized band basally (as in Shimonia Lehmann & Rajaei 2013 ), but rarely connected by a thin membranous structure basally ( L. rajaeii ); usually not bent towards uncus, but sometimes strongly bent towards uncus ( L. timdavenporti sp. nov.). Valva ventroposteriorly with one thorn-like process, well developed, always hollow inside and strongly sclerotized, usually bent upward, rarely setose, sometimes with a smaller thorn-like process at its base ( L. bergsteni , L. jankiellandi sp. nov.), extending from near base of valva and along costal margin with a large, setose and weakly-sclerotized projection (so-called “inner valva”). Usually with rounded, rarely ovoid emargination below. Broad transverse semi - transtilla leaf-like, attached to costa of valva and opposite of vinculum, usually without setae. Sacculus sometimes absent. Juxta usually broad and elongate, rarely small and narrow ( L. rajaeii , L. mikerobertsi ), usually with two acuminate lobes with a flag-like process at each apex and a deep posterior emargination. Broad vinculum sometimes opposite saccus, forming a plate as broad as length of saccus ( L. stueningi , L. mbalaensis , L. utaheidenreichae ). Vinculum with broadened lower half. Saccus broad, narrowly triangular or digitate, sometimes smaller than or much larger than juxta; usually rounded at tip. Phallus simple, tube-like, long to very long (1.0–1.5 length of valva), straight, or strongly bent at middle; vesica without cornuti. Female postabdominal structure and genitalia ( Fig. 11d 1-2 ) with papillae anales broad; dorsal part obliquely 8-shaped or elliptic in posterior view, covered with short and long setae. Segment 8 setose along its posterior margin and with long anterodorsal setae, not emarginate. Ventro-lateral surface of segment 8 with two large strongly sclerotized bands, partly separate or sometimes entirely fused along the posterior margin of the bands; always broadest at middle. Ductus bursae and corpus bursae broad, membranous, extremely thin, corpus bursae often pear-shaped, rarely rounded, large or very large (> size of segment 8 in lateral view); ductus bursae or corpus bursae lacking any processes.

Species richness. Currently, 29 species are included in this new genus of which 28 species are described as new to science.

Distribution. Species of Lukeniana occur in southern and eastern Africa, ranging from the Southern Ethiopian Plateau, southward via eastern Uganda, western and central Kenya to north-central and western Tanzania into the southeastern Democratic Republic of the Congo ( DRC), eastern and central Zambia, all of Malawi, and further south, but north of the Limpopo River, to western Mozambique and via Zimbabwe westward into Botswana and north-central Namibia ( Figs 19–20). Mount Elgon (on the frontier of Uganda and Kenya and ca. 120 km northeast of Lake Victoria) is currently the most northwestern distribution limit and the Otjozondjupa Region in north-central Namibia (ca. 2,800 km southwest of Lake Victoria) the most southwestern limit; the area around Gidole (southern Ethiopian Plateau) is currently the northeastern limit and Chipinge (south-eastern Zimbabwe) the southeastern limit. Within this vast area, the majority of Lukeniana species have scattered distributions (defined here as less than 50,000 km 2). They are found at elevations greater than 600 m on mountain ranges, dormant volcanoes, highland and/or plateau areas (including plateau inselbergs), and on high plains associated with the “East African Rift System (EARS)” sensu McConnell (1972), to the “Southern African Plateau” sensu Nyblade & Sleep (2003) and Said et al. (2015), and to the adjacent Chimanimani-Bvumba-Inyanga mountainland. Species of Lukeniana occur from 600– 2,419 m in a range of climate types, e.g., warm temperate upland, subtropical humid highland climate and tropical dry climate. The average annual rainfall in these habitats varies from at least 325 mm in the Otjozondjupa Region, 1594 mm at Lubumbashi to 1956 mm at Kakamega Forest (western Kenya). As regular element of the Afromontane Region ( White 1981) north of the Limpopo River, species of Lukeniana are also likely to occur in relict forests or woodlands in submontane and/or montane areas from Cameroon through Nigeria westward to Sierra Leone (West Africa).

In contrast, species of Mountelgonia are only found at higher elevations in eastern Africa ( Rwanda, Burundi, Uganda, Kenya, Tanzania and extreme northeastern Zambia). In Kenya, and probably in other parts of eastern Africa, species of Mountelgonia are locally sympatric with species of Lukeniana , e.g. on Mount Elgon ( Lukeniana timdavenporti / Mountelgonia percivali Lehmann, 2013 ), around Thika ( L. kammeri sp. nov. / Mountelgonia thikaensis Lehmann, 2013 ), on the Game Ranch of Dr. David Hopcraft on the Athi River-Kapiti Plains and adjacent to the Lukenia Hills ( L. georgeadamsoni / Mountelgonia arcifera Hampson, 1909 ) and on the Lolldaiga Hills Ranch ( L. mikerobertsi / Mountelgonia sp. nov. 9 / Fig 133 in Lehmann 2019b).

Habitat. See Appendix 1.

Biological traits. The biology of all species of Lukeniana is unknown at present.

Etymology. The genus is named after the “Lukenia Hills” (1,635 –1,840 m, ca. 126′01′′S–129′27′′S and 3705′14′′E–3704′03′′E), located ca. 25 km southeast of Nairobi ( Kenya). The areas in and around Nairobi support high species richness of this genus, including the type species, L. obliqualinea . The spelling represented here for the “Lukenia Hills” follows the Geographical Section of the Naval Intelligence Division in the Handbook of Kenya Colony and Protectorate (1920). The same area is also known as “Lukenya”, e.g. on the map published by the Survey of Kenya in 1965, or as “Mount Lukenya” or “Lukenya Hill” (D. Hopcraft pers. com. to I.L. in 1995). The gender of the new genus is feminine.

Figures

FIGURE 1. a. Zambezia jennyhuntae, holotype, male, Zimbabwe, Manicaland Province, Mutare (formerly Umtali); b. Z. durrellbarnesi, holotype, female, Zimbabwe, Manicaland Province, Bvumba Mountains; c. Z. darrelplowesi, holotype, female, Zimbabwe, Manicaland Province, Mutare (formerly Umtali), Hospital (today Kopje House); d. Metarbelodes umtaliana, holotype, male, Zimbabwe, Manicaland Province, Old Umtali or Umtali (today Mutare).

FIGURE 2. a. Lukeniana butleri, paratype, male, Zimbabwe, Harare Province, Harare, “Fawlty Towers”; b. L. butleri, paratype, female, Zimbabwe, Harare Province, Harare, “Fawlty Towers”; c. L. kollhorsti, holotype, male, Zimbabwe, Manicaland Province, Busi Farm; d. Zambezia diredaouaensis, holotype, male, Ethiopia, Harar Plateau, Dire Daua, today: chartered city of Dire Dawa; e. Z. madambae, holotype, male, Botswana, Chobe District Council, Kasane, Chobe River, Chobe Rapids; f. Z. madambae, paratype, female, Botswana, Chobe District Council, Kasane, Chobe River, Chobe Rapids.

FIGURE 3. a. Lukeniana enaiposha, holotype, male, Kenya, Rift Valley Province, near Lake Naivasha; b. L. obliqualinea, male, Kenya, County Nairobi, Nairobi; c. L. obliqualinea, female, Kenya, County Nairobi, Loresho Ridge; d. L. raymondrevellii, holotype, male, Namibia, Otjozondjupa Region, Ozombanda; e. L. rajaeii, paratype, male, Ethiopia, Southern Nations, Nationalities and Peoples’Region, Gidole; f. L. tubiraensis, holotype, male, Tanzania, Kigoma Region, Tubira Forest.

FIGURE 9. Wing venation of: a. Zambezia darrelplowesi, female, holotype (Zambezia = subgroup G1 in Lehmann 2019b); b. Lukeniana obliqualinea, female, Kenya, County Nairobi, Ngong (Lukeniana = subgroup G3 in Lehmann 2019b); c. Zambezia madambae, male, holotype; d. Z. diredaouaensis, male, paratype; e. Lukeniana tubiraensis, male, holotype; f. Metarbelodes umtaliana, male, holotype (Metarbelodes = subgroup G2 in Lehmann 2019b). The areole (blue arrows) is a possible plesiomorphy (in Lehmann 2019b, p. 24), the continuous CuP fold (red arrows), partly sclerotized, is a very rare apomorphy (in Lehmann 2019b, p. 25) among species of Metarbelidae. It has been found only in species close to or to the South of the Zambezi River. In the same areas an intact CuP on the forewing occurs only in basal species (as defined by Lehmann 2019b, pp. 24 and 25) of Metarbelidae (Lehmann 2019b). The complete loss of the CuP in the forewing is another apomorphy (Lehmann 2019b, p. 322) that occurs in the majority of species of Metarbelidae in theAfrotropical Region and Oriental Region (seeFigure 39).Drawings by I.L.

FIGURE 10. An intact CuP vein on the forewing is a plesiomorphy (cf. Fig. 9) and occurs only in basal Metarbelidae in northeastern parts of the Republic of South Africa, mainly on the Southern African Plateau, and on the plateau area of Mount Mulanje, only 90 km to the North of the Zambezi River (Lehmann 2019b). Figured are examples of apomorphies in regard to the CuP on forewing (red arrows): a. CuP absent, CuP fold is weak but not continuous in Lukeniana michaelgrzimeki, female, paratype; b. The CuP fold is almost entirely absent in L. rajaeii, male, paratype; c. Zambezia darrelplowesi, female, holotype and d. Z. madambae, male, paratype, both have a strong CuP fold, continuous and partly sclerotized. Images were taken by Dr. Hossein Rajaei (SMNS) in November 2015.

FIGURE 11. a. Zambezia madambae, holotype, male, Botswana, Chobe District Council, Kasane, Chobe River, Chobe Rapids; b. Z. jennyhuntae, holotype, male, Zimbabwe, Manicaland Province, Mutare (formerly Umtali); c. Metarbelodes umtaliana, holotype, male, Zimbabwe, Manicaland Province, Old Umtali or Umtali (today Mutare); d1. Lukeniana mzuzuensis, paratype, female, Malawi, Northern Region, Mzuzu, Nkho-rongo, female genitalia with ductus bursae and corpus bursae, both thinly membranous and without any structures; d2. Detail of female genitalia of L. mzuzuensis, paratype, with narrow bandson lateroventral surface of segment 8 (cf. red arrows); e. L. obliqualinea, holotype, male, Kenya, County Nairobi, Nairobi (picture taken by Kim Buckmaster, NHMUK, London (picture taken in 1996)..

FIGURE 13. a. Lukeniana enaiposha, holotype, male, Kenya, Rift Valley Province, near Lake Naivasha; b. L. obliqualinea, male, Kenya, County Nairobi, Nairobi; c. L. raymondrevellii, holotype, male, Namibia, Otjozondjupa Region, Ozombanda; d. L. rajaeii, paratype, male, Ethiopia, Southern Nations, Nationalities and Peoples’Region, Gidole; e. L. tubiraensis, holotype, male, Tanzania, Kigoma Region, Tubira Forest; f. L. lutztoepferi, holotype, male, Kenya, County Nairobi, Ololua Forest.

FIGURE 14. a. Lukeniana bergsteni, paratype, male, Tanzania, Kilimanjaro Region, Mount Kilimanjaro, Kibonoto; b. L. carolae, holotype, male, Malawi, Central Region, Mount Ntchisi, Ntchisi Forest Reserve; c. L. stevecollinsi, holotype, male, Malawi, Northern Region, Nyika Plateau, Nyika National Park, near Thazima Gate; d. L. butleri, paratype, male, Zimbabwe, Harare Province, Harare, “Fawlty Towers”; e. L. kollhorsti, holotype, male, Zimbabwe, Manicaland Province, Busi Farm; f. Zambezia diredaouaensis, holotype, male, Ethiopia, Harar Plateau, Dire Daua, today: chartered city of Dire Dawa.

FIGURE 15. Examples of male genitalia in lateral view: a. Zambezia madambae, paratype, with extremely small thorn-like process and small weakly-sclerotized projection (red arrows); b. Lukeniana timdavenporti, paratype, with strongly bent ventral edge of valva and well developed thorn-like process (red arrows); c. L. bergsteni, paratype, without strongly bent ventral edge of valva and two thorn-like processes (red arrows); d. L. butleri, paratype, with many long setae on valva that appear to be black in colour and one thorn-like process (red arrows).