Subtribe lachnocnemina




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FAMILY LYCAENIDAE

Leach, 1815


SUBFAMILY MILETINAE
Reuter, 1896



Relevant literature:

Ackery, 1990


TRIBE MILETINI

Reuter, 1896


SUBTRIBE LACHNOCNEMINA

Clench, 1955




Genus Lachnocnema Trimen & Bowker, 1887
South-African butterflies: a monograph of the extra-tropical species 2 Erycinidae and Lycaenidae: 233 (242 pp.). London.

Type-species: Hesperia bibulus Fabricius, by subsequent designation (Hemming, 1960. Annotationes lepidopterologicae (Part 1): 11 (7-11).).


A purely Afrotropical genus containing 36 species.

[Generic review by Libert, 1996 (in three parts).]



Lachnocnema bibulus group

Lachnocnema bibulus (Fabricius, 1793)
Hesperia bibulus Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 307 (488 pp.).

Lachnocnema bibulus. Male. Left – upperside; right – underside. Wingspan: 25mm. Estcourt, Natal. 3.10.1973. K.M. Pennington. (Transvaal Museum - TM3768).

Lachnocnema bibulus. Female. Left – upperside; right – underside. Wingspan: 26mm. Umkomaas, Natal. 9.IV.44. K.M. Pennington. (Transvaal Museum - TM3768).

Lachnocnema bibulus. Female. Left – upperside; right – underside. Wingspan: 29mm. Cintsa East, Eastern Cape Province, South Africa; 20-31 December, 2001; M.C. Williams (Williams collection).
Type locality: [West Africa]: “In Indiis”. [False locality.] Lectotype designated by Libert, 1996 - “Vallé du Pungoue, Mozambique, 1906 (G. Vasse), M.N.H.N., Paris.”

Distribution: Uganda (south), Burundi, Democratic Republic of Congo (south-east and east), Kenya (south), Tanzania, Malawi, Zambia (throughout), Angola, Mozambique, Zimbabwe, Botswana, Namibia (north), South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal, Eastern Cape Province), Swaziland (Duke, et al., 1999). Also in Madagascar (Stempffer, 1954; Lees et al., 2003).

Specific localities:

Kenya – Kisumu (Larsen, 1991); Nairobi (Larsen, 1991); Meru (Larsen, 1991); Rabai (Larsen, 1991); Teita Hills (Larsen, 1991); Kibwezi (Larsen, 1991).

Zambia – Victoria Falls (Heath, et al., 2002); Ndola (Heath, et al., 2002); Mufulira (Heath, et al., 2002).

Limpopo Province – Waterberg (Swanepoel, 1953); Potgietersrus (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Zoutpansberg (Swanepoel, 1953); Legalameetse Nature Reserve (“Malta Forest”).

Mpumalanga – Lydenburg (Swanepoel, 1953).

North West Province – Kgaswane Mountain Reserve (Williams).

Gauteng – Pretoria (Swanepoel, 1953); Witwatersrand Botanical Gardens (J. Dobson, unpublished checklist, 2001).

KwaZulu-Natal – Umkomaas (Swanepoel, 1953); Durban (Swanepoel, 1953) Hluhluwe (Swanepoel, 1953); Pietermaritzburg (Swanepoel, 1953); Greytown (Swanepoel, 1953); Muden (Swanepoel, 1953).

Eastern Cape Province – Grahamstown (Swanepoel, 1953); Peddie (Swanepoel, 1953); Bathurst (Swanepoel, 1953); King William’s Town (Swanepoel, 1953); Keiskamma (Swanepoel, 1953); Alicedale (Swanepoel, 1953); East London (Swanepoel, 1953); Butterworth (Swanepoel, 1953); Tsomo River (Swanepoel, 1953); Bashee River (Swanepoel, 1953); west of Port Elizabeth (Pringle, et al., 1994).

Madagascar – Ampandrandava in southern Madagascar (André Seyrig, April 1931 – specimen in MNHN, Paris; see also Libert, 1996: 496); near Betroka, not far east of Ampandrandava (Diehl, 1955).

Common name: Common woolly legs.

Habitat: Savanna and open forest. In Madagascar in the subarid bioclimatic zone (Lees et al., 2003).

Habits: Usually found in fairly well-defined colonies. Males defend territories from a perch on a bush or tree; the flight is rapid but usually not long sustained. Male territories may be on the flats or on hill-tops (Larsen, 1991). Small groups of adults are often found imbibing the honeydew secreted by sap-sucking homoptera (Henning, G., 1984). Females fly weakly, just above the ground, frequently settling on low vegetation (Pringle, et al., 1994).

Flight period: All year.

Early stages:
Jackson, 1937: 210 [Mt. Elgon, Kenya].

Larvae are found on many species of tree or shrub, provided that membracids or jassids (Homoptera) are present. The larva feeds on the secretions of the homopterans. “The fore-legs are much modified, being long, crab-like and capable of extended movement. Sitting back on its abdominal segments, the forepart of the body is raised over the anal tube of the membracid, the fore-legs are then vibrated rapidly over the insect, just in the same way as an ant strokes certain larvae with its antennae, when a drop of the secretion promptly appears and is greedily devoured. I have never seen a membracid actually eaten. Egg. The eggs are orange when freshly laid, small, circular, flattened, and strongly indented centrally above. They are laid among colonies of membracids or jassids but not in my experience among coccids, the female often pausing with abdomen raised away from the ants to feed on the secretion left on the stems or leaves. Larva. The larva is first red with black head, later brown. The dorsum is curved evenly from the anal extremity upwards and then down again, the head centrally very broad. Both collar and lateral carapace are present, the latter having scalloped edges. Dorsally there are rows of warts carrying long hair. Head black and small. Parasites. A tachinid was bred.”


Cripps and Jackson, 1940: 449 (Trans Roy Ent. Soc. Lond. 90: 449-453).

Cripps found eggs on a species of Vernonia (Asteraceae), not among the membracids that were also present on the plant. Females have also been seen, by Jackson, ovipositing on a low-growing species of Combretum, among ants; membracids were not present. The ants, in this case, were perforating the main veins on the undersides of the leaves, and feeding on the resultant flow of sap; the eggs were always placed close to these incisions. The young larvae, and both sexes of the adult, were also feeding on this sap. There were no signs of any but very young larvae, and no pupae were to be found. Cripps noted that “eggs are laid at about 14:00 to 15:00, on Vernonia, a shrub common on the edges of old native clearings. The young terminal shoots are often covered with membracids and jassids, and these are attended by masses of ants. I have seen the female ovipositing on several occasions, and she appears to be particular about the selection of a suitable leaf, those chosen being the old leaves about half-way down the stem. I have not seen a female ovipositing on the young shoots among ants, jassids, etc., nor have I found eggs or larvae there”. Eggs are often laid among a minute orange fungus on the leaves, which the eggs closely resemble [T.J.]. “Soon after the larva has hatched an ant finds it, and investigates it with its antennae for a few seconds. The ant then seizes the larva in its mandibles, from the side, and gently pulls it free of the leaf. The larva, stiffening and curling slightly, is lifted and turned upon its back; it is now easily and quickly carried down the branch to the base of the shrub. It is there dropped, and then finds its way to a resting place among the ants. Sometimes several ants approach the larva on its leaf, each taking hold of it, but always from the side, when a tug-of-war ensues until one eventually wins and carries off the prize. Sometimes the ant will drop the larva when a foot or so from the ground. I collected and placed full-grown larvae on the top of the shrub; upon each occasion an ant came and inspected it for a moment, then seized it, and took it down the stem to the nest. The ants are extremely strong; I saw one lift a full-grown larva, with another ant attached, and walk off with them down the shrub. On one occasion, an ant was descending the stem with a small larva I had placed at the top of a branch, when, about half-way down, they came to a hole in the bark on the underside of the stem; the ant popped the larva into the hole. Next day, I examined the hole and found the larva still there, and it remains there as I write, attended by ants all the time, who fight with each other over guarding and tickling it. I examined several other branches with holes in them; each hole contained a larva, and ants were at the entrance. The holes are always on the undersides of the stems, protected from the weather, and the larvae were always very small.”

“I opened an ant-run at the base of a mimosa thorn (Acacia stenocarpa Hochst.), the roots of which were within a foot of a Vernonia shrub. Larvae, in various stages, were found, lying quietly, five or six to a run, and several pupae were attached to the roots roofing the runs. Larvae and pupae were found also in the runs among the Vernonia roots. The pupae had no puparium whatsoever; they hung, head down, from the root, with the hollow of the run below them, attached by the anal extremity. There was no protection of any kind.” The Pupa case is hard and highly polished [T.J.]

The larvae in the runs were seen in any position, facing the entrance or the reverse, and at various depths, sometimes several together, sometimes alone. Ants were constantly in attendance, stroking the larvae with their antennae, sometimes as many as six or eight at the same time. At the same spot underground, I found, at about 8 a.m. on a very dull, cloudy day, two female butterflies with wings fully spread. They walked slowly out, attended all the time by ants who stroked them with their antennae (C.C.). Jackson speculates that the hairs on the legs protect the butterfly when it emerges in and then leaves the ant nest. Cripps noted trophallaxis between a mature larva and ants in captivity. An ant would approach the larva and antennate it in the craniodorsal region. After some time the larva raised its anterior portion and rapidly vibrated its legs against the head of the ant. The ant then vibrated its antennae for a second then spread them wide apart and kept them still. The larva placed its mouth-parts within the mandibles of the ant, which were now wide open. A small drop of clear fluid was regurgitated [by the ant] and seized by the larva which then drew away, the drop slowly disappearing into the larva. The larva then dropped to the ground and walked away. This behaviour was noted on several occasions but a drop of fluid was not always produced. The whole interaction lasts about two minutes. Sometimes a larva, upon being antennated, will shake the anterior segments violently and turn its head away; this does not appear to alarm the ant. Although there is a well developed gland on the abdominal segments this does not appear to be utilized by the ants. Jackson believes that the larvae are ‘honey-dew’ feeders and that only exceptionally (e.g. when starved) will they actually feed on homoptera [but see Clark & Dickson, 1971].

Pupa. Placed below ground in the ant-runs, and attached head downwards, to the roots of the tree or shrub, by the anal extremity. Shape squat, broad, much shortened, angulate at the head-case and shoulders; abdominal segments broad, evenly excurved from anal extremity and incurved to a pronounced constriction or ‘waist’ at a point about the centre of the wing-cases. A lateral view shows that this ‘waist’ extends round and above the pupa, the abdominal and thoracic segments being raised and domed above it. Colour pale dirty brown, wing-cases lighter; surface highly polished. Length 11 mm; width 8 mm at the broadest part.”

Larval gland (described by H. Eltringham). “An examination of the dorsal half of a longitudinal section showed, on the inner surface of the eighth segment, a pale patch lying between the hypoderm and the cuticle. Transverse sections were made of this eighth segment. These sections show a flattened gland of bifid structure, the division being in the long axis of the body, i.e. along the dorsal centre line. There is a containing membrane, and glandular lobes cut in various planes. Some of the lobes show comparatively large cells, and others have a more fibrous appearance. No very obvious ducts are apparent. Careful examination of the whole series fails to show any special opening to the exterior, nor is any such opening visible on examination of the dorsal cuticle from the outside. The active appearance of the gland in section suggests that it is functional, and if its secretion does in fact make its way to the exterior, it must do so by some form of diffusion or osmosis. There would be nothing very unusual about this as I have found glands in Lepidoptera which are very obviuosly active and functional but which show no evident external orifice. The chitinous cuticle of this larva shows the usual pore canals, and there seems no reason why the excretion, if any, should not occur through them. As usual in such larvae, there are numerous subcuticular muscle fibres and the contraction of these would cause pressure on the glands with a tendency to expel the contents.”
Clark, 1940: 50.
Owen, 1971 (Owen, D.F., 1971. Tropical butterflies. Clarendon Press, Oxford).
Clark and Dickson, 1971: 248 [as Lachnocnema bibulus; near Port Elizabeth, Eastern Cape].

Egg: 0.5 mm diam. x 0.25 mm high. Laid singly, as a rule, but two or three eggs may be laid in close proximity. They are white when laid, gradually changing to pinkish. Eggs hatch after 9-12 days. Larva: 1st instar 0.95 mm, growing to 3 mm in 3 days; 2nd instar growing to 4.5 in 4 days; 3rd instar growing to 8 mm in 6 days; 4th instar growing to 13 mm in 9 days. Although they remain among ant-attended Jassids, larvae are not molested in any way by the ants. In the early instars they feed on the young Jassids and on their droppings, and in the final instar larvae rear themselves up over a Jassid and suddenly pounce on it or crawl up behind it and seize the wing tips. The claws on segments 1-3 are abnormally long and are thus adapted to this procedure. There are many broods but diapause may take place in very cold weather. The characteristic features of the larvae in all the instars are shown clearly in the accompanying plate, while the larva was described in some detail in an earlier publication. Pupa: 7 mm. Secured by the cremastral hooks and by a silken girdle to twigs or dead leaves. The imago emerges after some 20 days. Recorded from eggs and larvae from near Port Elizabeth.”


Cottrell, 1984.
Larval food:

(Probably) Sorghum sp. (Poaceae) [Bowker, in Trimen & Bowker, 1889: 418] Note: this is erroneous, in the light of our knowledge today.



Ossana bicolor Distant (Homoptera) [Lamborn, 1914: 470 (Nigeria)].

(Secretions of) membracids (Homoptera: Membracidae) and jassids [now psyllids] (Homoptera: Psyllidae) [Jackson, 1937: 210; Mt. Elgon, Kenya].

Ant regurgitations [Cripps and Jackson, 1940].

Plant juice exuding from the mid-rib of leaves of a species of Combretum that had been punctured by ants [Cripps & Jackson, 1940: 449].

Nymphs and adults of an unidentified homopteron (Homoptera) [Clark, 1940: 50].

Psyllids (Homoptera: Psyllidae) [Clark and Dickson, 1971: 248; as jassids; near Port Elizabeth, Eastern Cape].

Juices exuding from a damaged plant stem [Cottrell, 1984].

Associated ant:

Camponotus akwapimensis Mayr. var. poultoni For. (Formicidae) [Lamborn, 1914: 470 (Nigeria)].
delegorguei Boisduval, 1847 (as sp. of Lycaena). In: Delegorgue, A., Voyage dans l’Afrique australe 2: 588 (585-602). South Africa/Zimbabwe: “Massilicatzi” [on Limpopo River, apparently near Messina, Limpopo Province].

Lachnocnema laches (Fabricius, 1793)
Hesperia laches Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 317 (488 pp.).

Synonym of Lachnocnema bibulus (Fabricius). Ackery et al., 1995: 542.



Lachnocnema laches (Fabricius, 1793). Libert, 1996.

Type locality: [West Africa]: “In America”. [False locality.] Lectotype designated by Libert, 1996, as: Burundi: “Usumbura (700 m), Urundi, 28 VIII 1961 (M. Fontaine), Coll. Stempffer, M.N.H.N., Paris.”

Distribution: Angola, Democratic Republic of Congo, Rwanda, Burundi, Tanzania (west and Dar es Salaam), South Africa (Limpopo, North-West Province, KwaZulu-Natal).

Specific localities:

Limpopo Province – Lapalala (Woodhall); Mphupuli (Woodhall); Levubu (Woodhall).

North-West Province – Brits (Woodhall).

KwaZulu-Natal – Mkuze (Collins); Manguzi (Woodhall).

Habitat: Savanna.

Flight period: ?

Early stages: Nothing published.

Larval food: Nothing published.

Lachnocnema pseudobibulus Libert, 1996
Lachnocnema pseudobibulus Libert, 1996. Lambillionea 96 (1) (Tome II): 196 (185-202).

Type locality: Tanzania: “Lagoda, Mufindi, Tanzanie, VI 1984 (S.C. Collins), N.H.M., Londres.”

Distribution: Uganda, Kenya, Tanzania, Democratic Republic of Congo, Malawi, Zimbabwe.

Habitat:

Early stages: Nothing published.

Larval food: Nothing published.

Lachnocnema sosia Libert, 1996
Lachnocnema sosia Libert, 1996. Lambillionea 96 (1) (Tome II): 197 (185-202).

Type locality: Uganda: “Katera, Sango Bay, Ouganda, IV. 1956 (T.H.E. Jackson), Coll. Stempffer, M.N.H.N., Paris.”

Distribution: Uganda, Kenya, Democratic Republic of Congo, Tanzania (north).

Specific localities:

Tanzania – Mount Meru (single record) (Congdon and Collins, 1998).

Habitat:

Early stages: Nothing published.

Larval food: Nothing published.

Lachnocnema riftensis Libert, 1996
Lachnocnema riftensis Libert, 1996. Lambillionea 96 (1) (Tome II): 197 (185-202).

Type locality: Kenya: “Foot of Kikuyu Escarpment, nr. Naivasha, 7500 ft, Kenya, 3. III.1911 (S.A. Neave), N.H.M., Londres.”

Distribution: Kenya, Burundi.

Habitat:

Early stages: Nothing published.

Larval food: Nothing published.

Lachnocnema kiellandi Libert, 1996
Lachnocnema kiellandi Libert, 1996. Lambillionea 96 (1) (Tome II): 198 (185-202).

Type locality: Tanzania: “Minziro for. (Bukoba), Tanzanie, X. 1993 (I. Kielland), N.H.M., Londres.”

Distribution: Tanzania.

Specific localities:

Tanzania – Minziro Forest; Nyakanazi, Biharamulo District (single male) (Congdon & Collins, 1998).

Habitat: Forest (Congdon and Collins, 1998).

Early stages: Nothing published.

Larval food: Nothing published.

Lachnocnema durbani group

Lachnocnema durbani Trimen & Bowker, 1887
Lachnocnema d’urbani Trimen & Bowker, 1887. South-African butterflies: a monograph of the extra-tropical species 2 Erycinidae and Lycaenidae: 236 (242 pp.). London.

Lachnocnema durbani. Male. Left – upperside; right – underside. Wingspan: 25mm. Rosslyn, Pta, Transvaal. 20.1.1962. Russell Badham. (Transvaal Museum - TM3769).

Lachnocnema durbani. Female. Left – upperside; right – underside. Wingspan: 30mm. Muden, Nat. 8.V.1952. H. Cookson. (Transvaal Museum - TM3769).
Type locality: South Africa: “Cape Colony. Eastern Districts: Grahamstown; New Year’s River, Albany District; between Kowie and Fish Rivers, Coast of Bathurst District; King William’s Town. Kaffraria Proper: Bashee River. Natal. Upper Districts: Estcourt; Ladysmith; Biggarsberg; Rorke’s Drift”.

Original description:

“Exp. al., male 11.5 lin. – 1 in. 2 lin.; female 10 lin. – 1 in. 2 lin. Very closely allied to L. Bibulus (Fab.). Male. Dull greyish-brown, much paler than in Bibulus; terminal disco-cellualar darker spot in fore-wing more apparent. Under side: very pale grey, with a faint yellowish tinge over hind-wing and narrow costal border of fore-wing; steely spots as in Bibulus, but less brilliant; the transverse band and discal row of spots much yellower. Fore-wing: base dusky (except on inner margin) as far as end of discoidal cell; quadrate mark on discoidal nervules much farther from extremity of cell. Hind-wing: central band much narrower at its beginning on costa, more irregular, being broken up into mostly separate unequal spots; discal row of spots also more irregular. Female. Very much paler and duller than in Bibulus, without dark costal borders and with only ill-defined dusky hind-marginal borders; no discal white patches, but a diffused very pale grey discal shade, inclining to whitish in hind-wing; bluish-grey basal suffusion obsolete. Fore-wing: terminal disco-cellular fuscous spot rather conspicuous, isolated, much smaller. Hind-wing: a more or less indistinct sub-lunulate terminal disco-cellular fuscous spot. Under side: as in male; much duller than in Bibulus, the markings less distinct. Fore-wing: the discal quadrate marking often (in four out of eight examples) expanded into a series of dusky spots extending from near costa to first median nervule.”



Distribution: Uganda, Rwanda, Kenya, Tanzania, Democratic Republic of Congo, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia (north), South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal, Eastern Cape Province), Swaziland (Duke, et al., 1999).

Recorded, in error, from Ghana by Berger, 1981 (Larsen, 2005a).



Specific localities:

Zambia – Victoria Falls (Heath, et al., 2002).

Limpopo Province – Warmbaths (Swanepoel, 1953); Potgietersrus (Swanepoel, 1953); Polokwane (Swanepoel, 1953); Munnik (Swanepoel, 1953); Zoutpansberg (Swanepoel, 1953); Legalameetse Nature Reserve (Williams, April 2006).

North West Province – Kgaswane Mountain Reserve (Williams).

Gauteng – Pretoria (Swanepoel, 1953); Onderstepoort (Williams).

KwaZulu-Natal – Hluhluwe (Swanepoel, 1953); Howick (Swanepoel, 1953); Tugela (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Ladysmith (Swanepoel, 1953); Biggarsberg (Swanepoel, 1953).

Eastern Cape Province – Groendal dam, to the west of Uitenhage (Dickson); Grahamstown (Swanepoel, 1953); Port Alfred (Swanepoel, 1953); Fish River (Swanepoel, 1953); Bathurst (Swanepoel, 1953); King William’s Town (Swanepoel, 1953); East London (Swanepoel, 1953).
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