014 Genus Graphium Scopoli

14th edition (2015).

Genus Graphium Scopoli, 1777

Introductio ad historium naturalem sistens genera lapidum, plantarum et animalium 433 (506 + [34] pp.). Pragae. Type-species: Papilio sarpedon Linnaeus, by subsequent designation (Hemming, 1933. Entomologist 66: 199 (196-200).) [extralimital].

An Old World genus of 97 species, 42 of which are found in the Afrotropical Region. Generic review of the Afrotropical taxa by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 503-719).

Swordtails are large butterflies and are closely related to the true Swallowtails. Characteristically they possess a long, sword-like tail on each hindwing, for example the Large Striped Swordtail. Some species, such as the Angola White-lady Swordtail and the Veined Swordtail, are, however, tailless. Swordtail species are variable in both colour and patterning but the sexes of a particular species are very similar. They are strong and rapid fliers, usually keeping from one to three metres above the ground. The 10 species found in southern Africa occur mostly in coastal bush, and in frost-free savanna in the more inland areas. They are therefore confined to the more easterly and northerly portions of the Subregion. They fly during the summer months, often with a big hatch of adults in spring (October-November). They are not often seen feeding from flowers but the males of some species occasionally congregate in large numbers on damp sand or on mud. They rarely engage in hilltopping behaviour. Whereas the Swallowtails utilize plants belonging mainly to the Citrus family (Rutaceae) as larval foodplants, the Swordtails appear to breed mainly on members of the Custard-apple family (Annonaceae). The early stages are considered under Swallowtails.

Relevant literature: Libert, 2007b [Notes]. Hancock, 2006 [Notes]. Makita et al., 2003 [Molecular phylogeny].

Subgenus Arisbe Hübner, [1819]

In Hübner, [1816-[1826]]. Verzeichniss bekannter Schmettlinge 89 (432 + 72 pp.). Augsburg. Type-species: Papilio leonidas Fabricius, by action of the I.C.Z.N. (Opinion 823, 1967. Bulletin of Zoological Nomenclature 24: 218 (218-219).). [Arisbe placed on the Official List of Generic Names in Zoology;Opinion 678, 1963. (Bulletin of Zoological Nomenclature 20: 339-342.).]

= Zelima Fabricius, 1807. In: Illiger, K., Magazin für Insektenkunde 6: 279 (277-289). Type-species: Papilio pylades Fabricius, by original designation. [Placed on the Official List of Rejected and Invalid Names in Zoology; Opinion 678, 1963. (Bulletin of Zoological Nomenclature 20: 339-342.).

= Ailus Billberg, 1820. Enumeratio Insectorum. In Museo Gust. Joh. Billberg 81 (138 pp.). Holmiae. 1

[Replacement name for Zelima Fabricius.]

The subgenus Arisbe is confined to the Afrotropical Region and contains 42 species. Smith & Vane- Wright (2001) divided them into the following groupings: (summarized at htpp://www.nhm.ac.uk/entomology/graphium/)

Swordtailed species The antheus clade (antheus, evombar) The policenes clade (policenes, liponesco, biokoensis, policenoides) A group of paraphyletic species (kirbyi, junodi, polistratus, colonna, illyri, gudenusi, porthaon)

Short tailed and tailless species The angolanus clade (angolanus, endochus, morania, taboranus, schaffgotschi, ridleyanus ) The leonidas group (leonidas, levassori, cyrnus) The tynderaeus clade (tynderaeus, latreillianus) An unplaced species (philonoe) The adamastor group (adamastor, agamedes, schubotzi, olbrechtsi, abri, almansor, auriger, fulleri, ucalegonides, rileyi, poggianus, kigoma, hachei, aurivilliusi, ucalegon, simonsi )

Swordtailed species. Consists of two well-supported clades, the antheus clade with 2 species, and the policenes clade with 4 very similar species.

The antheus clade

Afrotropical members of this clade appear to be more closely related to Oriental members of the genus than they are to the policenes clade (Smith & Vane-Wright, 2001; Makita et al., 2003).

*Graphium (Arisbe) antheus (Cramer, [1779])# Large Striped Swordtail

Large Striped Swordtails (Graphium antheus). Left – male on wet sand, Malawi (image courtesy Raimund Schutte). Right – Female resting, Harold Johnson Nature Reserve, KwaZulu-Natal (image courtesy Steve Woodhall).

Papilio antheus Cramer, 1779 in Cramer, [1779-80]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en 2

America 3: 71 (176 pp.). Amsteldam & Utrecht. Papilio antheus Cramer. Trimen, 1862c. Papilio antheus Cramer, 1782. Trimen & Bowker, 1889. Papilio mercutius Smith & Kirby, 1894. Aurivillius, 1899. [synonym of antheus] Papilio antheus Cramer, 1779. Aurivillius, 1899. Papilio antheus Cramer. Swanepoel, 1953a. Papilio antheus Cramer, 1779. Dickson & Kroon, 1978. Graphium (Arisbe) antheus (Cramer, 1779). Pringle et al., 1994: 306.

Graphium antheus. Male (Wingspan 68 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal, South Africa. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium antheus. Female (Wingspan 73 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal, South Africa. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Type locality: [Africa]: “Eiland Amboina”. [False locality.]. Larsen (2005a) thinks that the type locality is Sierra Leone. Distribution: Sub-Saharan Africa, including Senegal, Guinea, Guinea-Bissau (Bivar de Sousa et al., 2007), Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2001), Nigeria (south), Cameroon, Equatorial Guinea, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Sudan, Uganda, Rwanda, Burundi, Ethiopia, Kenya, Tanzania, Malawi, Zambia,

Mozambique, Zimbabwe, Botswana, Namibia (north), South Africa (Limpopo Province, Mpumalanga, North West Province, Gauteng, KwaZulu-Natal, Eastern Cape Province), Swaziland. Specific localities: Senegal – Basse Casamance (Larsen, 2005a). Ghana – Kakum National Park (Larsen, 2006d); Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Togo – Mishahöhe (Strand, 1914); Sokadé (Strand, 1914); Basari (Strand, 1914). Benin – Noyau Central, Lama Forest (Fermon et al., 2001). Cameroon – Korup (Larsen, 2005a). Gabon – Troughout (van de Weghe, 2010). Central African Republic – Bangui (Basquin & Turlin, 1986); Dzanga (Noss, 1998). Democratic Republic of Congo – Sankishia (Dufrane, 1946). Uganda – Semuliki N.P. (Davenport & Howard, 1996); Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Kenya – Sabaki River (Hampson, 1891); Kibwezi (Strand, 1909); coast (Larsen, 1991c). Tanzania – Kunusi (Strand, 1914); Tabora (Strand, 1914); Ukaranga (Strand, 1914); Langenburg (Strand, 1914); practically throughout, including Pemba Island (Kielland, 1990d); Semdoe Forest Reserve (Doggart et al., 2001). Mozambique – Maputo (Grose-Smith & Kirby, 1894); Beira (Van Son, 1949); Rikatla (Van Son, 1949). Zimbabwe – Changadze River, near Birchenough Bridge (Van Son, 1949); Gwanda (Van Son, 1949); Bulawayo (Van Son, 1949); Matetsi (Van Son, 1949); Victoria Falls (Van Son, 1949); Harare (Van Son, 1949); Mutare (Van Son, 1949); Melsetter (Van Son, 1949); Mafussi (Van Son, 1949); Sanyati Valley (Van Son, 1949). Botswana – Chobe (Larsen, 1991l); Nata (N. Ashby vide Larsen, 1991l); Mpandama-Tenga (R. Plowes vide Larsen, 1991l); Sashe (G. Bailey vide Larsen, 1991l); Tswapong Hills (Larsen, 1991l); Radisele (Larsen, 1991l). Limpopo Province – Woodbush (Van Son, 1949); Lekgalameetse Nature Reserve (“Malta Forest”) (Swanepoel, 1953); Tubex (Swanepoel, 1953); Duiwelskloof (Swanepoel, 1953); Sibasa (Swanepoel, 1953); Entabeni Forest (Swanepoel, 1953); Wyliespoort (Swanepoel, 1953); Saltpan (Swanepoel, 1953); Messina (Swanepoel, 1953); Blouberg (Swanepoel, 1953); Doorndraai Dam Nature Reserve (Warren, 1990); Percy Fyfe Nature Reserve (Warren, 1990); Highlands Wilderness (Bode & Bode, unpublished checklist); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Komatipoort (Swanepoel, 1953); Barberton (Swanepoel, 1953); White River (Swanepoel, 1953); Graskop (Swanepoel, 1953); Acornhoek (Swanepoel, 1953); Mariepskop (Swanepoel, 1953). North West Province – Utopia Resort (C. Dobson, 2006). KwaZulu-Natal – Durban (Van Son, 1949); Oribi Gorge (Swanepoel, 1953); Eshowe (Swanepoel, 1953); Pietermaritzburg (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Empangeni (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); Kosi Bay Nature Reserve (Pringle & Kyle, 2002); Tembe Nature Reserve (Pringle & Kyle, 2002); Hluhluwe (male illustrated above). Eastern Cape Province – Port St. Johns (Swanepoel, 1953); Cebe, about 10 km south-west of Mazeppa Bay (Quickelberge). Swaziland – Throughout bushveld areas (Swanepoel, 1953); Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz).. Habitat: Frost-free savanna and forest margins. In Tanzania at altitudes from sea-level to 1 900 m (Kielland, 1990d). Specimens will fly far into true forest, presumably along river courses (Larsen, 2005a). Habits: The flight is very fast and direct, two to three metres above the ground, and with frequent changes of direction and tempo to investigate things that catch its attention. Flowers are often visited, red and yellow ones appearing to be favoured. Males mudpuddle, sometimes in very large assemblages, especially in September and October in southern Africa. Males do not appear to be territorial and both sexes fly randomly in the bush (Pringle et al., 1994). It is occasionally recorded migrating (Larsen, 1991c). Flight period: September to April in southern Africa. Early stages:

Monteiro, 1891: 219 4

Clark, in Van Son, 1949: 44 [as Papilio (Graphium) antheus]. “Egg like that of P. policenes Cr. and laid in the same way on the same plant. Larva. First instar practically the same as in P. policenes; second instar of the same shape as in policenes, but dark chocolate with yellow collar on first segment and some yellow on last segment; third and fourth instars of the same size and shape as in policenes, but chocolate with yellow collar and some yellow on the forked final segment above; fifth (final) instar: head light brown, body velvety- brown above spiracular line, below it and ventrally as in policenes; yellow collar with a black spine on each side; forked posterior segment mostly black, with a yellow basal patch extending over the dorsum only; body with traces of bands in some larvae; size slightly larger than in policenes (35 mm). Pupa. Rather cylindrical in shape, not widened angularly nor depressed as in policenes; cephalic projections flattened in front and more diverging apically; dorsal thoracic projection short, only slightly elevated towards tip and very little produced forward, hardly reaching the level of the cephalic projections, and connected with the middle of the head by a low, but sharply defined, vertical keel ending in a small, bisinuate transverse carina between the cephalic projections; the colour is greyish-brown, darker on posterior thoracic segments above and on all abdominal segments according to the only specimen at hand; length 23 mm.”

Clark, in Pringle et al., 1994: p.416; plate 39 [as Graphium (Arisbe) antheus]. “The eggs are laid singly on the tips of young shoots. They are 1 mm in diameter and 0,95 mm high and are pale, watery green in colour. The egg stage lasts about four days. The larva at first eats small holes in the leaves but later eats from the edge. The larva rests along the midrib of the leaf. It becomes green during the pre-pupal stage. The larval stage lasts about two weeks. The pupa is held upright by a silken girdle and is attached by the cremastral hooks to a silken pad. The pupal stage lasts from a week to several months, depending on the individual.”

Henning, Henning, Joannou, & Woodhall, 1997: 243 & 244 (photograph of early instar larva, final instar larva (two colour forms) and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Egg and third instar larva of Graphium antheus. Images courtesy Allison Sharp (left) and Raimund Schutte (right).

Graphium antheus final instar larvae. Green form left and brown form right. Images courtesy Raimund Schutte (left) and Allison Sharp (right).

Graphium antheus pupae. Images courtesy Raimund Schutte (left) and Allison Sharp (right).

Larval food: Annona reticulata L. (Annonaceae) (exotic) [Platt (1921) cited by Van Son, 1949: 44]. Annona senegalensis Pers. (Annonaceae) [Platt (1921) cited by Pringle et al., 1994: 306]. Artabotrys brachypetalus Benth. (Annonaceae) [Pringle et al., 1994: 306]. Artabotrys monteiroae Oliv. (Annonaceae) [Van Son, 1949: 44]. Cleistochlamys kirkii (Benth.) Oliv. (Annonaceae) [Mullin, in Pringle et al., 1994: 306]. Hexalobus monopetalus (A. Rich.) Engl. & Diels (Annonaceae) [Williams, in Pringle et al., 1994: 306]. Landolphia buchananii (Hallier f.) Stapf (Apocynaceae) [Van Someren, 1974: 316; as Landolphia ugandensis Staph.]. Monanthotaxis caffra (Sond.) Verdc. (Annonaceae) [Williams, 1996: 134; Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Monodora junodi Engl. & Diels (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Uvaria caffra E.Mey. ex Sond. (Annonaceae) [Van Son, 1949: 44]. Uvaria kirkii Hook.f. (Annonaceae) [Heath et al., 2002: 26]. Uvariodendron kirkii Verdc. (Annonaceae) [Congdon et al., 2009: 46].

antharis Godart, 1819 in Latreille & Godart, [1819], [1824] (as sp. of Papilio). Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 52 (1-328 [1819], 329-828 [1824]. Paris. [Africa]: “L’Amérique septentrionale”. [False locality.]

nyassae Butler, 1877 (as sp. of Papilio). Annals and Magazine of Natural History (4) 19: 459 (458-462). “Lake Nyassa”.

lurlinus Butler, 1883 (as sp. of Papilio). Annals and Magazine of Natural History (5) 12: 106 (101-107). “Victoria Nyanza”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 533 (503-719))].

evombaroides Eimer, 1889 (as ssp. of Papilio antheus). Die Artbildung und Verwandtschaft bei den Schmetterlingen [1]: 228 (243 pp.). Ghana: “Goldküste”.

utuba Hampson, 1891 (as sp. of Papilio). Annals and Magazine of Natural History (6) 7: 182 (179-184). Kenya: “East Africa, Sabaki River district”. [Lectotype designated by Smith & Vane-Wright, 2001 ( Bulletin of the Natural History Museum Entomology Series 70 (2): 533 (503-719))].

mercutius Grose-Smith & Kirby, 1894 in Grose-Smith & Kirby, 1892-7 (as sp. of Papilio). Rhopalocera exotica, being illustrations of new, rare and unfigured species of butterflies 2: 33 (261 pp.). London. Mozambique: “Morakwen, Delagoa Bay”. 6

hollandi Grose-Smith & Kirby, 1894 in Grose-Smith & Kirby, 1892-7 (as sp. of Papilio). Rhopalocera exotica, being illustrations of new, rare and unfigured species of butterflies 2: 33 (261 pp.). London. [Invalid; junior primary homonym of Papilio hollandi Edwards, 1892 [Papilionidae]. Name introduced on plate, but replaced by mercutius in description.]

scheffleri Strand, 1909 (as ab. of Papilio antheus). Archiv für Naturgeschichte 75 (1.3.): 367 (367-386). Kenya: “British Ost Afrika, Kibwezi”.

latepictulus Strand, 1914 (as ssp. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 4 (3-4, 10). Tanzania: “D.O. Afrika, Kongoë, Kunusi”.

combinata Strand, 1914 (as ab. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Tanzania: “Tabora in D.O. Afrika”.

micrevombaroides Strand, 1914 (as f. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Togo: “Mishahöhe; Sokadé, Basari”.

misanus Strand, 1914 (as ab. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Togo: “Misahöhe”.

hoesemanni Strand, 1914 (as ab. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Tanzania: “N.O. Tanganyika, Ukaranga; N. Nyassa See, Langenburg”.

rubrimacula Strand, 1914 (as ab. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Tanzania: “Langenburg am N. Nyassa See”.

comma Strand, 1914 (as ab. of Papilio antheus). Internationale Entomologische Zeitschrift 8: 10 (3-4, 10). Tanzania: “Langenburg am N. Nyassa See”.

mathieui Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) antheus antheus). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 120 (101-122). Democratic Republic of Congo: “Congo, Sankishia”.

paradoxa Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) antheus antheus ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 120 (101-122). Democratic Republic of Congo: “Congo Republic Belge”.

atrantheus Basquin & Turlin, 1986 (as f. of Papilio antheus). Bulletin de la Société Scientifique Naturelles (51): 1 ((49): 22; (51): 1.). Central African Republic: “Bangui, R.C.A.”.

anthaeus auct. (as sp. of Papilio). Vári et al., 2002: 129.

*Graphium (Arisbe) evombar (Boisduval, 1836)

Papilio evombar Boisduval, 1836. In: [Roret, Suites à Buffon] Histoire naturelle des Insectes. Species général des Lépidopteres 1: 254 (690 pp.). Paris. Papilio evombar Boisd., 1836. Aurivillius, 1899.

Type locality: Madagascar. Distribution: Madagascar, Comoro Islands. Habitat: Forest (Lees et al., 2003). Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) evombar evombar (Boisduval, 1836)

Type locality: Madagascar. Distribution: Madagascar.

Graphium (Arisbe) evombar viossati Collins, 1997

Graphium evombar viossati Collins, 1997. In: d’Abrera, 1997 Butterflies of the Afrotropical region. Part 1: 56.

Type locality: Comoro Islands: Anjouan, Forest of Moya. Distribution: Comoro Islands. Specific localities: Comoro Islands – Moya Forest, Anjouan (TL).

The policenes clade Consists of 4 very similar species. G. liponesco has distinctive male genitalia. Both G. liponesco and G. biokoensis are very difficult to differentiate from G. policenes.

*Graphium (Arisbe) policenes (Cramer, [1775])# Small Striped Swordtail

Left: Small Striped Swordtail (Graphium policenes) female, resting. Harold Johnson Nature Reserve, KwaZulu-Natal. Image courtesy Steve Woodhall. Right: Small Striped Swordtail (Graphium policenes) male, mudpuddling. Kibale Forest, Uganda. Image courtesy Raimund Schutte.

Papilio policenes Cramer, 1775 in Cramer., [1775-6]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 1: 61 (155 pp.). Amsteldam and Utrecht. Papilio policenes Cramer. Trimen, 1862c. Papilio policenes Cramer, 1779. Trimen & Bowker, 1889. Papilio policenes Cramer, 1775. Aurivillius, 1899. Papilio policenes Cramer. Swanepoel, 1953a. Graphium (Arisbe) policenes (Cramer, 1775). Pringle et al., 1994: 306.

Graphium policenes policenes. Male (Wingspan 58 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium policenes policenes. Female (Wingspan 63 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Alternative common name: Common Striped Swordtail. Type locality: [Africa]: “Surinamen”. [False locality.] Diagnosis: G. antheus differs in that three of the blue-green bars in the cell on the forewing upperside are S- shaped. In G. polistratus the bands are narrower, less green in colour, and the underside is much darker (Larsen, 1991c). In G. liponesco and G. biokoensis these bars are much narrower than in G. policenes; in addition these two species have narrower wings, lack a red tornal spot on the hindwing underside, and have proportionately longer tails (Larsen, 2005a). Distribution: Sub-Saharan Africa, including Guinea, Guinea-Bissau (Bivar de Sousa et al., 2007),Mali, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2001), Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Sudan, Uganda, Burundi, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, South Africa, Swaziland (Duke et al., 1999). Habitat: Coastal forest and other warm forests, including riparian forest (not in montane forest). In Tanzania from sea-level to 2 000 m (rarely at the higher elevations) (Kielland, 1990d). It may also be seen in quite open agricultural areas (Larsen, 2005a). Habits: This is the commonest of the long-tailed swordtails. Flies rapidly and erratically, often deviating from its course to investigate other butterflies or possible food sources. Males may be found feeding in large aggregations 9 at mud puddles and on damp sand. Both sexes feed from flowers. Both sexes fly randomly, from one to three, or more, metres above the ground. Migrations have been recorded (Larsen, 1968; 1991). In Kakum, Ghana, Larsen (2005a) noted red-headed agamid lizards (Agama agama) eating numbers of mudpuddling specimens. Flight period: End of August to April (Pringle et al., 1994). Early stages:

Saunders, 1857: (Saunders, W.W., Trans. Ent. Soc. Lond. 1857, 2nd Series, iv. pl. 13. figs a,b,c.). Trimen & Bowker, 1889 (footnote on p. 204) aver that these are the early stages of “another Papilio - either P. Demoleus [P. demodocus] or a near ally”).

Trimen & Bowker, 1889: 203 [as Papilio Policenes Cramer; Durban, KwaZulu-Natal]. “Larva. Back ferruginous-red, transversely striped with black. Head sandy-yellow. On each segment, except second, third, and fourth (thoracic), a median transverse streak of greyish-blue, black-edged both anteriorly and posteriorly; on thoracic segments no blue streak, but the transverse black edging streaks strongly marked; second segment bright yellow; on each thoracic segment, situate dorso-laterally, a pair of forward- and outward-pointing short acute black spines. Lower part of sides (separated from dorsal ferruginous by a lateral black stripe) pale-bluish, fading into whitish on the under surface. On anal segment a pair of short, acute, backward- and outward-pointing spines, superiorly yellow at base and mesially bluish, but at tip and inferiorly black. I received in 1878 four living larvae, collected near D’Urban by Colonel Bowker, and was led to think that they probably were those of Policenes because one of them became an imperfect pupa, agreeing in the main with some from which I reared this species, and also because they closely resembled a pencil-drawing by Mr. W.D. Gooch of a larva taken in the same locality, and referred by him doubtfully to Policenes. One of the four examples sent by Colonel Bowker was much smaller than the others, and evidently in a much earlier moult, but it differed in no particular except in the proportionally larger thoracic spines. Pupa. Flattened dorso-abdominally; thoracic dorsal projection much prolonged, pointing forward and upward, and extending as far forward as, or even a little farther than, the head; cephalic prominences short, acute, widely apart; lateral margin with two small acute projections, one behind the other; a little beyond these, abdominal margin considerably expanded laterally (almost foliated). Bright pale-green; roughened lateral edge of dorsal projection down to lateral abdominal expansion, median frontal line of that projection, and cephalic and thoracic lateral margin, all creamy- ferruginous, which tint forms an irregular lateral patch where edges of base of dorsal projection and those of thorax meet; two spots of the same colour on back of abdomen (third segment); spiracles superior, fuscous. (Plate II, fig. 4). I obtained twelve Policenes (six of each sex) from pupae sent by Colonel Bowker from D’Urban, Natal. They reached Cape Town on 30th November 1878, and the butterflies emerged during the succeeding fortnight, with the exception of one ♂, which did not make its appearance until 12 th February 1879. Seven of these had the date of their becoming pupae attached, and I was thus able to record that (with the exception just noted) the chrysalis state lasted from seven to fifteen days. All these pupae were suspended to leaves, in a position near the mid-rib.”

Clark, in Van Son, 1949: 46 [as Papilio (Graphium) policenes]. “Egg. Almost spherical in shape, flattened below, pale watery greenish-yellow, 1 mm high and of the same diameter. Laid singly on young shoots of the food-plant. The egg-stage lasts four days. Larva. There are five instars, the larval stage being 12 days. First instar: 2.5 mm long, dark brownish-grey, slightly thickened anteriorly, with pairs of spiny dorso-lateral appendages on the thoracic segments, the anterior pair being the largest, and bearing tubercles beset with distally forked spines; anal segment with spreading processes bearing about a dozen thin, black spines; abdominal segments with a dorso-lateral tubercle bearing apically a long, forked spine and at the base with two forked spines about one-third the length of the apical spine; a similar lateral tubercle is present above the spiracle bearing three forked spines of the same size as the basal spines of the dorso-lateral tubercle; a third tubercle bearing a single spine is situated between the two tubercles mentioned above, but slightly posterior in position; a fourth two-spined tubercle is present on the pleura below the spiracle, and a fifth, smaller one, with a single spine, is found a little posteriorly to it; there are two spines on the outer sclerites of the proleg. The day after hatching the larva reaches a length of 4.25 mm and prepares to moult, which event takes place on the next day. Second instar: yellow, with black-edged transverse stripes above and a lateral white stripe; the thoracic and anal processes have lost their spines, and the abdominal segments and sides of prolegs have now laterally a number of very short bristles, the tubercles of the first instar having completely disappeared; at the end of this stage the length reaches 7.5 mm, and moulting takes place two days after the final moult. Third instar: differs from the second in the lateral stripe which is now alternately white and yellow, more yellow before the anal processes, and in the absence of bristles; it prepares to moult on the following day and is then 10 mm long; moulting takes place a day after this. Fourth instar: This instar only differs from the third in the lateral stripe being pale blue, and the length reaches 20.5 mm. The larva prepares to moult on the following day, and a day later the moulting takes place. Fifth instar: 32 mm long when full grown. Head sandy-yellow; thoracic spines black, anterior pair very short, middle and posterior pairs long, acute, spreading horizontally. Dorsal surface ferruginous- yellow, with broad black edgings to second and third segments, the latter with a pale blue transverse narrow stripe enclosed by the black colour; abdominal segments each with a transverse light blue stripe at middle, narrowly edged with black both in 10 front and behind; intersegmental grooves brownish. A black lateral line. Lower half of sides light blue, fading ventrally into the whitish underside. Last segment light bluish-white, tinged with yellowish at base, with a pair of acute, diverging spines, which are light bluish-white edged with black above, the tips and underside being black. The larva stops feeding on the fourth day after the last moult, and pupates two days later; just before pupation the larva turns green and is very difficult to distinguish from the larva of P. morania.” “The pupal period varies greatly, even under the same conditions.” “It may be as short as a week, but may last for several months…”.

Carcasson, 1981 [larva and pupa].

Clark, in Pringle et al., 1994: p. 418; plate 40 [as Graphium (Arisbe) policenes laurentia]. “The eggs are laid singly on young shoots of the foodplant and are a pale, watery, greenish yellow. They are almost spherical, with a diameter of 1 mm. The egg stage lasts four days. There are five larval instars lasting 12 days. In the pre- pupal stage the larvae turn green. The pupa is attached by the cremastral hooks and is held upright by a silken girdle. The pupal stage is variable and may last from a week to several months.”

Henning, Henning, Joannou, & Woodhall, 1997: 245 & 246 (photograph of early instar larva, final instar larva (two colour forms) and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Graphium policenes policenes larva on Monodora mystica. Kibale Forest, Uganda. Image courtesy Raimund Schutte.

Larval foods: Annona reticulata (Annonaceae) [Larsen, 1991c: 118]. Annona senegalensis (Annonaceae) [Nichols, 1995]. Annona squamosa (Annonaceae) [Nichols, 1995]. Artabotrys monteiroae Oliv. (Annonaceae) [Van Son, 1949: 49]. Artabotrys species (Annonaceae) [Bowker, in Trimen & Bowker, 1889: 203; as Artabotrys, n. sp.; Durban, KwaZulu-Natal and Delagoa Bay, Mozambique]. Landolphia buchannani Engler (Apocynaceae) [Van Someren, 1974: 316; Sevastopulo, 1981]. Landolphia ugandensis Staph. (Apocynaceae) [Van Someren, 1974: 316]. Monanthotaxis caffra (Annonaceae) [Nichols, 1995]. Polyalthia species (Annonaceae) [Fontaine, 1985 (Democratic Republic of Congo)]. Uvaria bukobensis Engler (Annonaceae) [Van Someren, 1974: 316]. Uvaria caffra E. Mey. ex Sond. (Annonaceae) [Dickson & Kroon, 1978]. Uvaria chamae Beauvais (Annonaceae) [Van Someren, 1974: 316].

Note: Henning, Henning, Joannou & Woodhall (1997: 244) treat the populations of G. policenes found in the eastern and southern parts of the Afrotropical region as subspecies laurentia (Le Cerf) and those in the western portions as the nominate subspecies.

Graphium (Arisbe) policenes policenes (Cramer, [1775])#

Papilio policenes Cramer, 1775 in Cramer., [1775-6]. Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en

America 1: 61 (155 pp.). Amsteldam and Utrecht. Papilio policenes Cramer, 1775. Aurivillius, 1899. Papilio policenes laurentia le Cerf, 1924. Dickson & Kroon, 1978. [synonym of policenes] Graphium (Arisbe) policenes laurentia (Le Cerf, 1924). Pringle et al., 1994: 306. [synonym of policenes]

Type locality: [Africa]: “Surinamen”. [False locality.] Distribution: Sub-Saharan Africa, including Guinea, Guinea-Bissau (Bivar de Sousa et al., 2007), Mali, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2001), Nigeria (south), Cameroon, Equatorial Guinea (Bioko), Gabon, Congo, Democratic Republic of Congo, Angola (Bivar de Sousa & Fernandes, 1966), Sudan, Uganda, Burundi, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe (eastern border), South Africa (KwaZulu-Natal – coast, Eastern Cape Province – north-east), Swaziland (Duke et al., 1999). Specific localities: Guinea – Mamou (Dufrane, 1946); Parc National du Haut Niger (Larsen, 2005a). Ghana – Shai Hills (Larsen, 2005a); Kakum National Park (Larsen, 2006d); Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Benin – Noyau Central, Lama Forest (Fermon et al., 2001).

Nigeria – Bauchi (Larsen, 2005a); Jos (Larsen, 2005a); Alantika Mountains (Larsen, 2005a) Cameroon – Korup (Larsen, 2005a). Gabon – Troughout (van de Weghe, 2010). Democratic Republic of Congo – Dekese (Berger, 1960). Uganda – Semuliki N.P. (Davenport & Howard, 1996); Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Kenya – Arabuko-Sokoke Forest (Larsen, 1991c). Tanzania – most forested areas (scarce in northern and very common in western forests) (Kielland, 1990d). Malawi – Mt Mulanje (Congdon et al., 2010); Zomba Mountain (Congdon et al., 2010). Zambia – Choma (Heath et al., 2002); Ikelenge (Heath et al., 2002); Mwinilunga District (Heath et al., 2002); the Copperbelt (Heath et al., 2002); Lunzua Falls (Heath et al., 2002); Kalambo Falls (Heath et al., 2002); Mafinga Mountains (Heath et al., 2002); Nyika (Heath et al., 2002). Mozambique – Mt Chiperone (Timberlake et al., 2007); Mt Namuli (Congdon et al., 2010); Mt Mabu (Congdon et al., 2010). Zimbabwe – Vumba Mountains (Van Son, 1949); Chirinda Forest (Van Son, 1949). KwaZulu-Natal – Durban (Le Cerf, 1924); Oribi Gorge (Swanepoel, 1953); Port Shepstone (Swanepoel, 1953); Umkomaas (Swanepoel, 1953); Eshowe (Swanepoel, 1953); Empangeni (Swanepoel, 1953); Mtubatuba (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); False Bay (Swanepoel, 1953); Kosi Bay Nature Reserve (Swanepoel, 1953); Fanies Island (male illustrated above); Tembe Nature Reserve (Pringle & Kyle, 2002). Eastern Cape Province – Port St Johns (Van Son, 1949); Ngqeleni (Van Son, 1949); Bashee River (Swanepoel, 1953).

pompilius Herbst, 1788 (as sp. of Papilio). Natursystem aller bekannten in- und ausländischen Insekten. Der Schmetterlinge 3: 205 (232 pp.). Berlin. [Africa]: “Surinam”. [False locality.]

agapenor Fabricius, 1793 (as sp. of Papilio). Entomologia Systematica emendata et aucta 3 (1): 26 (488 pp.). Africa.

scipio Palisot de Beauvois, 1805 in Palisot de Beauvois, [1805-21] (as sp. of Papilio). Insectes receueillis en Afrique et en Amérique 70 (276 pp.). Paris. Nigeria: “Les déserts derrière les royaumes d’Oware et de Galbar ou Kalabar”.

polixenus Godart, 1819 in Latreille & Godart, [1819], [1824] (as sp. of Papilio). Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 52 (1-328 [1819], 329-828 [1824]). Paris. [Africa]: “L’Amérique septentrionale”.

laurentia Le Cerf, 1924 (as ssp. of Papilio policenes). Bulletin of the Hill Museum, Witley 1: 398 (369-399, 576, 578, 580, 582, 584). South Africa: “Natal, Durban”.

guineensis Dufrane, 1946 (as f. of Papilio (Cosmodesmus) policenes policenes). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 121 (101-122). Guinea: “Guinée, vallée de Mamou”.

coussementi Berger, 1960 (as f. of Graphium policenes). Lambillionea 60: 71 (71-72). Democratic Republic of Congo: “Congo, Kasai, Dekese”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

Graphium (Arisbe) policenes telloi Hecq, 1999

Graphium policenes telloi Hecq, 1999. Lambillionea 99: 517 (517-518).

Type locality: Central African Republic: “RCA, Région Nord, P.N., Brendja 12.XII.1998, leg J. Tello.” Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Nigeria (north), Cameroon (north), Central African Republic, Sudan (south). Specific localities: Nigeria – Mapeo (Schultze, 1916); Jos Plateau (Larsen, 2005a). 13

Central African Republic – Brendja (TL); Dzanga (Noss, 1998). Sudan – Tamboura (Larsen, 2005a).

Note: Larsen (2005a) notes that sudanicus (Schultze, 1916), described as a variety of G. policenes from northern Nigeria, is similar to specimens with light coloration and broad bands from the Jos Plateau, Tamboura in Sudan, and to ssp. telloi. Larsen and Libert are therefore of the opinion that telloi is a synonym of sudanicus. Larsen speculates that this may represent clinal variation, or that sudanicus may represent a valid subspecies or even species. He does not, however, make any formal taxonomic changes.

sudanicus Schultze, 1916 (as var. of Papilio policenes). Archiv für Naturgeschichte 81 (A.11.): 110 (110-112). Nigeria: “Mapéo im Alantika Gebirge, Mittel Adamaua”.

*Graphium (Arisbe) liponesco (Suffert, 1904) Long-tailed Striped Swordtail

Papilio policenes liponesco Suffert, 1904. Deutsche Entomologische Zeitschrift, Iris 17: 107 (12-107). Graphium liponesco (Suffert, 1904). Larsen, 1994: 152.

Published images: Larsen, 2005a (male upperside). Type locality: Togo: “Hinterlande von Togo”. Diagnosis: For distinguishing features from G. policenes, see this species. Distribution: Guinea, Sierra Leone, Ivory Coast, Ghana, Togo, Nigeria (west). Specific localities: Guinea – Boola (Strand, 1909). Ghana – Kakum National Park (Larsen, 2005a); Bobiri Butterfly Sanctuary (Larsen et al., 2007). Nigeria – Agege near Lagos (Larsen, 2005a). Habitat: Wet and moist forests in good condition (Larsen, 2005a). Habits: Though widely distributed this is a scarce species. The flight is slower and more gliding than in G. policenes (Larsen, 2005a). Males mud-puddle, usually together with the much more numerous G. policenes (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

boolae Strand, 1909 (as sp. of Papilio). Archiv für Naturgeschichte 75 (1.3.): 307 (303-311). Guinea: “Boola”. [Considered to be a junior synonym of G. liponesco by Larsen, 1994: 149.].

*Graphium (Arisbe) biokoensis (Gauthier, 1984) Gauthier’s Striped Swordtail

Papilio policenes biokoensis Gauthier, 1984. Entomologische Zeitschrift. Frankfurt. a.M. 94: 318 (314-320). Graphium policenes biokoensis (Gauthier, 1984). Ackery et al., 1995: 165. Synonym of Graphium policenoides Holland. Hancock, 1993: 568. Synonym of Graphium liponesco (Suffert). Larsen, 1994: 152. Graphium biokoensis (Gauthier, 1984). Smith & Vane-Wright, 2001: 566.

Type locality: Equatorial Guinea: “Insel Bioko”. Diagnosis: Very similar to G. liponesco but is slightly larger; light grey scales at base of space 2 are absent (Larsen, 2005a). Distribution: Nigeria (east), Equatorial Guinea (including Bioko), Cameroon, Gabon, Congo, Democratic Republic of Congo (east), Burundi. Specific localities: Nigeria – Okwangwo (Larsen, 2005a). Cameroon – Nyong River, near Makak (Birket-Smith, 1960). 14

Habitat: Forest. Habits: Flies together with the much commoner G. policenes (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

intermedia Birket-Smith, 1960 (as ab. of Papilio nigrescens). Bulletin de l’Institut Français d’Afrique Noire (A) 22: 540 (521-554, 924-983, 1259-1284). Cameroon: “French Cameroons, on the bank of the river Nyong just south of the town of Makak”.

*Graphium (Arisbe) policenoides (Holland, 1892)

Papilio policenoides Holland, 1892. Annals and Magazine of Natural History (6) 10: 287 (284-294). Papilio nigrescens Eimer, 1889. Aurivillius, 1899. [synonym of policenoides]

Type locality: Gabon: “Talaguga, upon the Upper Ogové”. Distribution: Cameroon, Gabon, Congo, Democratic Republic of Congo (Bas-Fleuve, Equateur) (Larsen, 1994: 151). Specific localities: Gabon – Talaguga on the Upper Ogove River (TL); Mouila (van de Weghe, 2010); Lake Gazengel (van de Weghe, 2010); ; Otoumbi (van de Weghe, 2010). Habitat: Wet forest in good condition (Larsen, 2005a). Habits: The species is extraordinarily rare: no more than about 25 specimens are known (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

nigrescens Eimer, 1889 (as ssp. of Papilio policines (sic)). Die Artbildung und Verwandtschaft bei den Schmetterlingen [1]: 223 (243 pp.). Cameroon: “Kamerun”. [Treated as an invalid junior primary homonym of Papilio philolaus nigrescens Eimer by Larsen, 1994 (Lambillionea 94 (2) (Tome I): 151, and in Ackery et al., 1995: 165.].

A group of 7 paraphyletic species

*Graphium (Arisbe) kirbyi (Hewitson, 1872) Kirby’s Swordtail

Papilio kirbyi Hewitson, 1872. Entomologist’s Monthly Magazine 9: 146 (146-147). Papilio kirbyi Hew., 1872. Aurivillius, 1899.

Type locality: “East Africa: Lagos”. [False locality]. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 540 (503-719))]. Diagnosis: As in Graphium polistratus, the light markings at emergence from the pupa are pale greenish but they soon fade to creamy white (Congdon et al., 2009). Distribution: Kenya (coast), Tanzania (coast; inland to Morogoro). Records from Nigeria (Boorman & Roche, 1957; Villiers, 1957) are erroneous (Larsen, 2005a). Specific localities: Kenya – coastal forests (Larsen, 1991c); Shimba Hills (Larsen, 1991c). Tanzania – Usambara (Aurivillius, 1899); Sanje, below the Uzungwa scarp (Kielland, 1990d); Kimboza Forest (Kielland, 1990d); Turiani (Kielland, 1990d); Kanga Mountain (Kielland, 1990d); Nguu Mountains (Kielland, 1990d); East Usambara (Kielland, 1990d); Msangazi Forest, western boundary of Mkwaja Ranch, coast (Congdon et al., 2009). 15

Habitat: Coastal forest. Closed forest in coastal forest-savanna mosaic (Congdon et al., 2009). In Tanzania it occurs from 200 to 500 m altitude (Kielland, 1990d). Habits: Both sexes visit flowers and males also mudpuddle (Larsen, 1991c). The flight is not quite as fast as most of the other tailed swordtails (Larsen, 1991c). Males are often encountered flying along forest roads. Females are more rarely seen but may be found in the earlier parts of the day while feeding from the flowers of forest shrubs (Kielland, 1990d). Flight period: September to November (Kielland, 1990d). Early stages:

Congdon et al., 2009 [photographs of larva and pupa]. It pupates head down, probably on the underside of a leaf.

Larval food: Annona senegalensis Pers. (Annonaceae) [Van Someren, 1974: 316]. Uvaria kirkii Hook f. (Annonaceae) [Congdon et al., 2009; soft leaves in captivity]. Uvaria species (Annonaceae) [Larsen, 1991c: 117]. Uvariodendron kirkii Verdc. (Annonaceae) [Congdon et al., 2009].

ottonis Aurivillius, 1899 in Aurivillius, 1898-9 (as var. of Papilio kirbyi). Kungliga Svenska Vetenskapakademiens Handlingar 31 (5): 492 (1-561). Tanzania: “Deutsch Ost-Afrika, Usambara”.

*Graphium (Arisbe) junodi (Trimen, 1893) Junod’s Swordtail

Papilio junodi Trimen, 1893. Transactions of the Entomological Society of London 1893: 138 (123-143). Papilio Junodi Trimen, 1893. Aurivillius, 1899. Papilio junodi Trimen, 1893. Dickson & Kroon, 1978. Graphium (Arisbe) junodi (Trimen, 1893). Pringle et al., 1994: 306.

Type locality: [Mozambique]: “Morakwen, Delagoa Bay”. Distribution: Angola, Democratic Republic of Congo, Uganda, Tanzania, Mozambique, Zimbabwe (eastern border). Specific localities: Mozambique – Maputo (40 km north of) (TL); Beira (Van Son, 1949); Amatongas (Chitty); Dondo Forest (Pennington); Inhamitango Forest (Pinhey); Maronga Forest (Pringle et al., 1994); Musapa Forest, east of the Chimanimani Mountains (Pinhey); Vila Fontes [Distrito de Caia] (Pinhey); Xiluvo Hills (D. Cookson). Zimbabwe – Chirinda Forest (McDermott, 2000: 51); Harare (Cox; single record); Witchwood Valley, below the Vumba (Barnes). Habitat: Warm forest. Habits: The flight is fast (Pringle et al., 1994). Flight period: Has been recorded from July to September and from January to April (Pringle et al., 1994). Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) polistratus (Grose-Smith, 1889) Dancing Swordtail

Papilio polistratus Grose-Smith, 1889. Annals and Magazine of Natural History (6) 3: 121 (121-137). Papilio sisenna Mab., 1890. Aurivillius, 1899. [synonym of polistratus] Papilio polistratus Smith, 1889. Aurivillius, 1899. Papilio polistratus Grose-Smith, 1889. Dickson & Kroon, 1978. Graphium (Arisbe) polistratus (Grose-Smith, 1889). Pringle et al., 1994: 306. 16

Type locality: [Kenya]: “East Coast of Africa, in the neighbourhood of Mombasa”. Diagnosis: Compared to G. policenes the green bars in the forewing cell are slightly wavy, narrower and paler (Kielland, 1990d). Distribution: Democratic Republic of Congo?, Kenya (coast), Tanzania, Malawi (north), Mozambique. Recorded, in error, from South Africa by Swanepoel (1953a) and from Kasane, Botswana by Larsen (Congdon et al., 2009). The Kasane record was for Graphium porthaon. Specific localities: Kenya – Mombasa (TL); coastal forest (Larsen, 1991c); Shimba Hills (Larsen, 1991c). Tanzania – Tanga (Weymer, 1892); East and West Usambara Mountains (Kielland, 1990d); Kimboza (Kielland, 1990d); Uluguru Mountains (Kielland, 1990d); Turiani (Kielland, 1990d); foot of Kanga Mountain (Kielland, 1990d); lower parts of Mwanihana Forest (Kilombero) (Kielland, 1990d); Pugu Hills (Kielland, 1990d); Masagati Forest south of Ifakara (Kielland, 1990d); Kahansi Gorge at base of Udzungwa Escarpment, 400 m (Congdon et al., 2009). Mozambique – Dondo Forest (Pinhey); on the banks of the Komati River, at Marracuene (Pennington). Habitat: Warm forest, including coastal forest. Also in heavy woodland (Kielland, 1990d). In Tanzania at altitudes from near sea-level to 900 m; also taken at 2 140 on Bondwa Mountain in the Ulugurus (Kielland, 1990d). Habits: Males are avid mudpuddlers (Larsen, 1991c) and both sexes feed from flowers (Kielland, 1990d). Males are readily attracted by decoys placed on the ground (Kielland, 1990d). Both sexes appear to keep to the shady undergrowth of forests (Gifford, 1965). Females are rarely seen as they tend to keep to the dark interior of the forests that the species inhabits (Larsen, 1991c). Flight period: Probably all year (Pringle et al., 1994). Larsen (1991) states that it is strongly seasonal. Early stages:

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: Annona senegalensis Pers. (Annonaceae) [Van Someren, 1974: 316]. Uvaria angolensis Oliv. (Annonaceae) [Congdon et al., 2009]. Uvaria species (Annonaceae) [Larsen, 1991c: 117].

sisenna Mabille, 1890 (as sp. of Papilio). Annales de la Société Entomologique de France (6) 10: 29 (17-51). Mozambique: “Côte de Mozambique”.

richelmanni Weymer, 1892 (as sp. of Papilio). Stettiner Entomologische Zeitung 53: 98 (79-125). Tanzania: “Ostafrika, Tanga”.

*Graphium (Arisbe) colonna (Ward, 1873)# Mamba Swordtail

Mamba Swordtail (Graphium colonna) male resting, Tembe Elephant Park, KwaZulu-Natal. Image courtesy SteveWoodhall.

Papilio colonna Ward, 1873. Entomologist’s Monthly Magazine 10: 151 (59-60, 1510152). Papilio colonna Ward, 1873. Trimen & Bowker, 1889. Papilio colonna Ward, 1873. Aurivillius, 1899. Papilio colonna Ward. Swanepoel, 1953a. Papilio colonna Ward, 1873. Dickson & Kroon, 1978. Graphium (Arisbe) colonna (Ward, 1873). Pringle et al., 1994: 307.

Graphium colonna. Male (Wingspan 58 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium colonna. Female. Left – upperside; right – underside. Dukuduku Forest, KwaZulu-Natal, South Africa. 26 February, 1977. P. Stobbia. Images M.C. Williams ex Henning Collection.

Alternative common name: Black Swordtail. Type locality: [Kenya]: “East Africa, Ribé”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 548 (503-719))]. Distribution: Liberia, Sierra Leone, Ghana, Ivory Coast, Togo, Equatorial Guinea, Gabon, Congo, Democratic Republic of Congo, Central African Republic, Sudan, Ethiopia (south-west), Somalia (south), Uganda, Kenya,

Tanzania, Malawi, Mozambique, Zimbabwe (east), South Africa (KwaZulu-Natal – north), Swaziland (Duke et al., 1999), Madagascar? Specific localities: Ethiopia – Lower Omo River. Kenya – Ribe (TL); Arabuko-Sokoke Forest (Larsen, 1991c); Shimba Hills (Larsen, 1991c); Thika (Larsen, 1991c); Meru (Larsen, 1991c); Mount Kulal (Larsen, 1991c). Tanzania – Mikindani (Suffert, 1904); Lindi (Suffert, 1904); coastal areas (Kielland, 1990d); Pemba Island (Kielland, 1990d); Usambara (Kielland, 1990d); Uluguru (Kielland, 1990d); Turiani (Kielland, 1990d); Sanje in the Kilombero Valley (Kielland, 1990d); Semdoe Forest Reserve (Doggart et al., 2001). Malawi – Mt Mulanje (single record). Mozambique – Maputo (Van Son, 1949). KwaZulu-Natal – Empangeni (Van Son, 1949); Ubombo (Van Son, 1949); Mtubatuba (Swanepoel, 1953); Greater St Lucia Wetland Park (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953); Kosi Bay Nature Reserve (Swanepoel, 1953); Dukuduku Forest (Pringle et al., 1994); Emanguzi Forest (Pringle et al., 1994); Makatini Flats (Pringle et al., 1994); Richards Bay (Pringle et al., 1994); Tembe Nature Reserve (Pringle et al., 1994). Habitat: Coastal forest and heavy woodland. In Tanzania it is found at altitudes from sea-level to 600 m (Kielland, 1990d). Habits: The flight is a little slower than other tailed swordtails but nevertheless still fast. They also fly slightly lower down (less than two metres above the ground) than other swordtails. Both sexes tend to fly in denser vegetation than other species, often weaving their way through thick, shady vegetation. The dark colouration and white tips of the tails are then very noticeable (Larsen, 1991c). Both sexes feed from flowers and males mudpuddle. Males do not appear to be territorial. Females are rarely seen (Larsen, 1991c). Flight period: October to April in southern Africa. The main flight period seems to be from December to February (Pringle et al., 1994). Early stages:

Trimen & Bowker, 1889: 211 [as Papilio Colonna Ward]. “Larva. Dull yellowish-green, with an interrupted brownish longitudinal streak along each side of the back. Sides of posterior half dark-brown; in one place (apparently on ninth segment) the strongly sinuated dorsal margins of this dark- brown closely approximate. Head brownish-sandy. Each of the three thoracic segments bearing dorso-laterally a pair of short forward- and laterally-pointing tubercular projections, - the second and third pairs clustered with small spines. Pupa. Pale bluish-green. Margin expanded laterally about wing-covers, but constricted in two places towards head, so that the outline viewed dorsally or ventrally is sinuated. Dorsal thoracic projection and upper part of edge of wing-covers inferiorly edged conspicuously with white. These descriptions of the earlier stages were made from drawings by Mrs. Monteiro, which she kindly lent to me in the year 1883.”

Monteiro, 1891: 217 [and plate facing p. 214]. Larva. “… the light green and brown caterpillar of the lovely Papilio colonna… is even more like a fish than this last (Papilio morania), as the tail is more strongly marked and the yellow-brown spines on the hump are branched, and consequently thicker and more like fins” Larva illustrated on the plate opposite page 214.

Larsen, 1991c: 118 [final instar larva]. “The larva is a magnificent creature, in strong green with a deep brown saddle and additional ornamentation.”

Henning, Henning, Joannou & Woodhall, 1997: 250 [material ex northern KwaZulu-Natal]. “Egg: unrecorded. Larva: third instar mid-brown ventrally, becoming dark chocolate-brown to black dorsally on second and third segments and dorso-laterally on others; head brown; dorsum of first and fourth to thirteenth segments yellow-green to bright green in a distinctive “saddle” pattern on fourth to ninth segments; brown to black coloration spreads dorsally on tenth and eleventh segments; first to third segments and bifid tail carry well-developed pairs of brown-black dorso-lateral projections carrying whitish spiny setae. It attains a length of 10 mm. Fourth instar larva similar in appearance but setose projections smaller in proportion to body and green becoming darker and more leaf-like. It attains a length of 18 mm. Fifth (final) instar still with characteristic saddle marking but thoracic segments invaded by bright leaf green so that dorsal brown is reduced to a series of spots forming broken transverse bands, dark coloration of ninth and tenth segments is maintained albeit much lighter; and the strongly bifid tail and thoracic projections still carry relatively large (compared to other Graphium larvae) orange-brown setose processes, the setae now being black. The green may be invaded by yellow to 19 give a mottled effect. Final instar attains a length of 25 mm. Pupa: pale bluish-green stippled with pale brown or yellow; dorso-ventrally flattened with wing-cases broadened and sinuate, the extremities of which bear silver metal marks. This unusual feature has not been recorded in any other member of the family. The above description is based on specimens collected in KwaZulu-Natal.”

The saddle marking and large setose processes make the larva of this species somewhat similar to that of the well known European notodontid Cerula vinula, the puss moth. It is easily distinguished from all its Southern African congeners and with its distinctive pupa hints that the taxonomy of this species may require further investigation. The pupa is attached by the cremaster to a silken pad and held upright by a silken girdle around the middle. The silken girdle is apparently weak in this species and often breaks, leaving the pupa hanging downwards. This unusual trait coupled with the unique silver metal mark make this pupa appear similar to that of some of the nymphalids that inhabit the same forests, such as Euphaedra neophron (Hopffer). The pupal stage lasts about 15 days.

Henning, Henning, Joannou, & Woodhall, 1997: 251 (photograph of early instar larva, final instar larva (two colour forms) and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: Annona species (Annonaceae) [Van Someren, 1974: 316]. Artabotrys monteiroae Oliv. (Annonaceae) [Dickson & Kroon, 1978]. Artabotrys species (Annonaceae) [Van Son, 1949: 55]. Uvaria caffra E.Mey. ex Sond. (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Uvaria kirkii Hook f. (Annonaceae) [Congdon et al., 2009; coastal Tanzania]. Uvaria species (Annonaceae) [Larsen, 1991c: 118; Pringle, 1995: 5].

tragicus Butler, 1876 (as sp. of Papilio). Entomologist’s Monthly Magazine 13: 56 (56-57). [Mozambique]: “Zambesi”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 548 (503-719))].

loncona Suffert, 1904 (as ssp. of Papilio colonna). Deutsche Entomologische Zeitschrift, Iris 17: 107 (12-107). Tanzania: “Deutsch Ost Africa, Mikindani und Lindi”.

*Graphium (Arisbe) illyris (Hewitson, 1873) Cream-banded Swordtail

Papilio illyris Hewitson, 1873. Entomologist’s Monthly Magazine 9: 232 (232-233). Papilio illyris Hew., 1873. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside). Type locality: [Ghana]: “Gold Coast”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 552 (503-719))]. Distribution: Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Congo, Gabon, Equatorial Guinea, Central African Republic, Democratic Republic of Congo, Tanzania. Habitat: Forest. Habits: This is an uncommon to rare butterfly and is seasonal in West Africa (mainly February and March) (Larsen, 2005a). Most of the time specimens keep to the interior of the forest but also fly along forest roads (Larsen, 2005a). Males mud-puddle, and are attracted to rotten fish as well as perspiration on humans (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) illyris illyris (Hewitson, 1873)

Papilio illyris Hewitson, 1873. Entomologist’s Monthly Magazine 9: 232 (232-233). Papilio illyris Hew., 1873. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside). Type locality: [Ghana]: “Gold Coast”. Distribution: Sierra Leone, Liberia, Ivory Coast, Ghana (central). Specific localities: Ghana – Friapere Forest, Coomassie (Le Cerf, 1924); Atewa Range (Larsen, 2005a); Kakum National Park (Larsen, 2005a).

stictica Le Cerf, 1924 (as f. of Papilio illyris). Bulletin of the Hill Museum, Witley 1: 399 (369-399, 576, 578, 580, 582, 584). Ghana: “Friapere Forest, Coomassie”.

Graphium (Arisbe) illyris flavisparsus (Fruhstorfer, 1903)

Papilio illyris flavisparsus Fruhstorfer, 1903. Stettiner Entomologische Zeitung 64: 359 (359-361). Synonym of Graphium illyris illyris (Hewitson, 1873). Ackery et al., 1995: 163. Graphium flavisparsus (Fruhstorfer, 1903). Canu, 1994: 313. Graphium flavisparsus (Fruhstorfer, 1903). d’Abrera, 1997: 54. Graphium illyris flavisparsus (Fruhstorfer, 1903). Smith & Vane-Wright, 2001: 554 stat. rev.

Type locality: Equatorial Guinea: “Fernando Po”. Distribution: Equatorial Guinea (Island of Bioko).

Graphium (Arisbe) illyris girardeaui Guilbot & Plantrou, 1978

Graphium illyris girardeaui Guilbot & Plantrou, 1978. Bulletin de la Société Entomologique de France 83: 70 (68-73).

Type locality: Central African Republic: “Empire centrafricain, Bangui”. Distribution: Gabon (van de Weghe, 2010), Central African Republic. Specific localities: Gabon – Kinguele (van de Weghe, 2010); Ipassa (van de Weghe, 2010); Waka (van de Weghe, 2010); Abeilles (van de Weghe, 2010); Lambarene (van de Weghe, 2010). Central African Republic – Bangui (TL); Dzanga (Noss, 1998).

Note: Larsen (2005a) considers this to be a synonym of hamatus but does not formally sink it.

Graphium (Arisbe) illyris hamatus (Joicey & Talbot, 1918)

Papilio illyris hamatus Joicey & Talbot, 1918. Proceedings of the Zoological Society of London 1917: 271 (271-272).

Type locality: [Tanzania]: “German East Africa”. Distribution: Nigeria, Cameroon, Equatorial Guinea, Congo, Democratic Republic of Congo, Tanzania. Specific localities: Nigeria – Sapoba (Larsen, 2005a); Oban Hills (Larsen, 2005a). Democratic Republic of Congo – Salonga River, near Watsikengo (Dufrane, 1946).

addenda Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) illyris). Bulletin et Annales de la Société Royale

Entomologique de Belgique 82: 121 (101-122). Democratic Republic of Congo: “Congo, le long de la rivière Salonga, environs de Watsikengo”.

*Graphium (Arisbe) gudenusi (Rebel, 1911)

Papilio gudenusi Rebel, 1911. Annalen des (K.K.) Naturhistorischen Museums. Wien 24: 409 (409-414).

Type locality: [Democratic Republic of Congo]: “Urwalde am Nordwestufer de Tanganika Sees, 1900-2100 m.”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 557 (503-719))]. Distribution: Democratic Republic of Congo (Kivu), Uganda (south-west – Kigezi district), Rwanda, Burundi. Specific localities: Democratic Republic of Congo – Kitembo, Kivu (Le Cerf, 1931). Uganda – Bwindi Forest (Davenport, 2002). Habitat: Forest (Davenport, 2002). Early stages: Nothing published. Larval food: Nothing published.

babaulti Le Cerf, 1931 (as sp. of Papilio). Bulletin de la Société Entomologique de France 1931: 275 (275-278). Democratic Republic of Congo: “Congo Belge oriental, Kitembo, région du lac Kivu”.

*Graphium (Arisbe) porthaon (Hewitson, [1865])# Cream Striped Swordtail

Papilio porthaon Hewitson, 1865 in Hewitson, 1862-6. Illustrations of new species of exotic butterflies 3: 4 [124 pp.]. London. Papilio porthaon Hewitson, 1865. Trimen & Bowker, 1889. Papilio porthaon Hew., 1865. Aurivillius, 1899. Papilio porthaon Hewitson. Swanepoel, 1953a. Papilio porthaon Hewitson, 1865. Dickson & Kroon, 1978. Graphium (Arisbe) porthaon (Hewitson, 1865). Pringle et al., 1994: 306.

Graphium porthaon porthaon. Male (Wingspan 58 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium porthaon porthaon. Female. Left – upperside; right – underside. Mount Selinda, Zimbabwe. 28 December, 1971. J. Tsalavoutos. Images M.C. Williams ex Henning Collection.

Alternative common names: Dark Swordtail; Coastal Swordtail. Type locality: [Mozambique]: “Zambesi”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 573 (503-719))]. Distribution: Kenya, Tanzania, Democratic Republic of Congo, Angola, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia, South Africa, Swaziland (Duke et al., 1999). Habitat: Frost-free savanna, coastal bush and warm forest. In Tanzania the nominate subspecies occurs from near sea-level to 1 900 m; ssp. tanganyikae is found from 780 to 1 000 m (Kielland, 1990d). Habits: Males fly rapidly and erratically, from one to three metres above the ground. Females fly a little more slowly. Both sexes feed from flowers and males mudpuddle (Kielland, 1990d). Larsen (CD-ROM) found the species mudpuddling in large numbers on roads parallel to the Chobe River in northern Botswana in January, 1992. Both sexes appear to fly randomly within their habitats. Flight period: September to April, with peak emergence usually in December and January (Pringle et al., 1994). Subspecies mackiei, from Meru, Kenya, flies only in November and December (Larsen, 1991c). Early stages:

Trimen & Bowker, 1889: 208 [as Papilio Porthaon Hewitson; pupa only]. “Pupa. Form almost as in Policenes, but dorso-thoracic anterior projection shorter, and marginal abdominal expansion less prominent. Pale-brownish, finely speckled with fuscous; on upper side, two blackish spots on head, one at apex of thoracic projection, and a row on each side of abdomen, about half-way between middle and lateral margin. Described from two specimens (which yielded a ♂ and a ♀ of Porthaon), brought from Delagoa Bay by Mrs Monteiro in July 1886. The butterflies did not emerge till the following December, and were both dwarfs. Other pupae of this species and of P. Colonna, Ward, which were taken on to England by Mrs. Monteiro, appear to have come out very satisfactorily in the Insect House at the Zoological Society’s Gardens.”

Monteiro, 1891: 219 [“Delagoa Bay”]. “First instar like a bit of shining steel.”

Aurivillius, in Seitz, 1925: 26 [a brief description of the final instar larva]. “The adult larva is velvety-green, with a variegated transverse band on each segment, composed of a white central line and on each side of this a black-brown, a green-yellow, and again a black-brown transverse line;..”.

Henning, Henning, Joannou & Woodhall, 1997: 247 [ex northern KwaZulu-Natal]. “Egg: whitish-yellow when first laid, darkening to yellow-green; almost spherical with a flattened base, diameter 0.8 mm. Larva: first instar blackish-purple, lightening to mauve-grey as it grows, fading to whitish ventrally; head black, tail bifid; first and second segments become paler as larva grows; thoracic segments with paired dorso-lateral spiny processes. It grows from 3 mm to 6 mm in six days. Second instar grey-white with fine transverse dark grey lines close together dorsally (four per segment) which make it appear mauve-grey from a distance; head brown-yellow and first and last segment orange- 23 yellow; segments each with a pair of dorso-lateral spined processes; tail bifid with a setose process on each half. It grows from 6 mm to 10 mm in four days. Third instar resembles second but has a yellow head. It grows from 10 mm to 17 mm in seven days. Fourth instar pale green ventrally, dorsum finely striped transversely from front with stripes of black, then yellow, black again, then white-blue, black again, yellow again and finally black once more, on each segment; thoracic segments with posterior yellow stripes more pronounced; head, first segment and tail orange; bifid tail tipped with black; same setose processes as in earlier instars but smaller in relation to size of larva. It grows from 17 mm to 28 mm in five to six days. Fifth (final) instar larva has two colour morphs. Dark morphs resemble fourth instar but head is brown-yellow, first segment and tail sulphur-yellow and black stripes broadened to give the larva a velvety black appearance when viewed dorsally; dorsal yellow and blue-white stripes broadened laterally to give an appearance of a pale lateral band along spiracles; thoracic segments carry dorso-lateral pairs of small black spikes. Green or yellow morph with same shape and processes, and head and ventral surfaces are the same colour as dark morph; differs dorsally in being bright leaf green or yellow with only dark shadows indicating where the previous instar had black transverse stripes; just above spiracles is a pale yellow lateral stripe and a green line along them; thoracic spikes black with yellow bases. Both morphs have a pale blue-green osmeterium. Pupa: green to pale brown with yellow striations and suture lines. Dorsal thoracic projection shorter than G. policenes and abdominal expansion less prominent. The above description is based on specimens collected in northern KwaZulu-Natal.”

Eggs are laid singly on leaf buds or on the underside of young leaves. Eggs take from three to four days to hatch, the larva becoming visible through the egg shell prior to hatching. The egg shell is not eaten. The young larvae hide in the flower bracts or along the eaten edge of young shoots. Older ones rest on the upperside of a leaf along the midrib, where they are nontheless hidden by other leaves. The pupa is leaf-like or bark-like and attached by the cremaster to a pad of silk spun on the stem of the food plant. It is held upright by a silken girdle around the middle.

Henning, Henning, Joannou, & Woodhall, 1997: 249 (photograph of early instar larva, final instar larva (three colour forms) and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: Annona species (Annonaceae) [Van Someren, 1974: 316]. Artabotrys monteiroae Oliv. (Annonaceae) [Dickson & Kroon, 1978]. Cleistochlamys kirkii (Benth.) Oliv. (Annonaceae) [Mullin, in Pringle et al., 1994: 307]. Friesodielsia obovata (Benth.) Verdc. (Annonaceae) [Paré, in Pringle et al., 1994: 307]. Monanthotaxis caffra (Sond.) Verdc. (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Monodora junodi Engl. & Diels (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Uvaria caffra E.Mey. ex Sond. (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Uvaria kirkii Hook.f. (Annonaceae) [Heath et al., 2002: 26]. Uvaria species (Annonaceae) [Kielland, 1990d: 48].

Graphium (Arisbe) porthaon porthaon (Hewitson, [1865])#

Graphium porthaon porthaon. Male (Wingspan 58 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium porthaon porthaon. Female. Left – upperside; right – underside. Mount Selinda, Zimbabwe. 28 December, 1971. J. Tsalavoutos. Images M.C. Williams ex Henning Collection.

Type locality: [Mozambique]: “Zambesi”. Distribution: Kenya (coast), Tanzania, Democratic Republic of Congo (east; single record, according to Larsen, 1991c: 119), Angola, Malawi, Zambia, Mozambique, Zimbabwe, Botswana (north), Namibia (Caprivi), South Africa (Limpopo Province – north-east, Mpumalanga – east, KwaZulu-Natal – north), Swaziland (Duke et al., 1999). Specific localities: Kenya – Arabuko-Sokoke Forest (Larsen, 1991c); Shimba Hills (Larsen, 1991c); Jilore (Larsen, 1991c). Tanzania – Langenburg (Thurau, 1903); Usambaras (Kielland, 1990d); Irindi on the lower slopes of Image Mountain (Kielland, 1990d); Kimboza (Kielland, 1990d); Uluguru Mountains (Kielland, 1990d); Turiani with the Nguru and Kanga Mountains (Kielland, 1990d); below the Uzungwa scarp (Kielland, 1990d); Pugu Hills (Kielland, 1990d). Malawi – Mt Mulanje (Congdon et al., 2010). Zambia – Lake Mweru (Heath et al., 2002); Lake Tanganyika (Heath et al., 2002); Luapula Valley (Heath et al., 2002); Luangwa River (Heath et al., 2002); Zambezi Valley (Heath et al., 2002). Mozambique – Maputo (Van Son, 1949); Bopira, Busi District (Van Son, 1949). Zimbabwe – Changadze River (Van Son, 1949); Lupani (Van Son, 1949); Gwaai River Bridge (Van Son, 1949); 25

Wankie (Van Son, 1949); Matetsi (Van Son, 1949); Victoria Falls (Van Son, 1949); Sinoia (Van Son, 1949); Sabi Valley (Van Son, 1949); Chipinga (Van Son, 1949). Botswana – Kabulabula, Chobe River (Van Son, 1936); Kasane (Larsen, 1991l); Kazungulu (Larsen, 1991l). Namibia – Katima Mulilo (Pringle et al., 1994). Limpopo Province – Lekgalameetse Nature Reserve (“Malta Forest”) (Van Son, 1949); Wyliespoort (Swanepoel, 1953); Saltpan (Swanepoel, 1953); Sibasa (Swanepoel, 1953); Mpaphuli Nature Reserve. KwaZulu-Natal – Mtubatuba (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); False Bay (Swanepoel, 1953); Magut (Swanepoel, 1953); Kosi Bay Nature Reserve (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953; male illustrated above); Muden (Pringle et al., 1994); Tugela River Valley (Pringle et al., 1994); south of Umhlanga Rocks (Dickson); Tembe Nature Reserve (Pringle & Kyle, 2002).

Note: The taxonomic status of vernayi van Son, 1936 is discussed below.

adjectus Thurau, 1903 (as ab. of Papilio porthaon). Berliner Entomologische Zeitschrift 48: 143 (117-143). Tanzania: “Langenburg”.

vernayi van Son, 1936 (as ssp. of Papilio porthaon). Annals of the Transvaal Museum 17: 138 (121-140). Botswana: “Kabulabula, Chobe River”. Note: Long series from Chobe are consistent and argue for considering vernayi a valid subspecies. Larsen writes in his CD-ROM on Botswana butterflies: “Ssp. vernayi was described from the Chobe area and later by van Son downgraded to an individual form; material I have seen supports maintaining it as a distinct subspecies. The forewing bands are broader and better fused than in coastal populations.”

Graphium (Arisbe) porthaon mackiei Collins & Larsen, 1991

Graphium porthaon mackiei Collins & Larsen, 1991. In: Larsen, 1991. The butterflies of Kenya and their natural history 119, 438 (490 pp.). Oxford.

Type locality: Kenya: “Meru Forest, xi 1988 (S.C. Collins, leg.)”. Holotype (male) in the National Museum, Nairobi. Description: “There are three main characteristics: (1) the discal band of the forewings (and some of the hindwing markings) are considerably broader, often twice as broad in space 1a; (2) on the forewing the four large discal spots from space 1a to 3 are wholly fused; those in 2 and 3 are free in the nominate subspecies; (3) the ground colour of the bands is a much deeper and clearer cream. In the very large series of both sexes studied there is no single specimen which is in any way intermediate.” “Interestingly, the Meru subspecies shows some similarity to that named ssp. vernayi van Son, 1936 from northeastern Botswana.” Etymology: Named for Mr. Leo Mackie of Pembroke House School, “through whose tutelage many young lepidopterists have passed.” Distribution: Kenya (lower Meru Forest on the eastern slopes of Mt Kenya). Specific localities: Kenya – Meru Forest (TL).

Graphium (Arisbe) porthaon tanganyikae Kielland, 1978

Graphium porthaon tanganyikae Kielland, 1978. Tijdschrift voor Entomologie 121: 161 (147-237).

Type locality: Tanzania: “Kigoma, Kasoge”. Diagnosis: Differs from the nominate subspecies in the following respects: ground colour darker; hindwing upperside with reduced submarginal spots; underside darker; red and black markings better developed (Kielland, 1990d). Distribution: Tanzania (Kigoma District). Specific localities:

Tanzania – Kasoge Forest at Mahale Mountain (TL); Kemfu Forest (Kielland, 1990d); Mihumu (Kielland, 1990d); Mukuyu (Kielland, 1990d).

Short tailed and tailless species

The angolanus clade A well-supported clade of six species, all with similar genitalia.

*Graphium (Arisbe) angolanus (Goeze, 1779)# Angola White Lady

Two male Angola White-lady Swordtails (Graphium angolanus) mudpuddling with a Brown-veined White (Belenois aurota) in Zambia. Image courtesy Raimund Schutte.

Papilio eques achivus angolanus Goeze, 1779. Entomologische Beyträge zu des Ritter Linné zwölften Ausgabe des Natursystems 3 (1): 87 (390 pp.). Leipzig. Papilio pylades Fabricius. Trimen, 1862c. [synonym of Graphium angolanus baronis] Papilio corinneus Bertoloni, 1849. Trimen & Bowker, 1889. [synonym of Graphium angolanus angolanus] Papilio pylades Fabr., 1793. Aurivillius, 1899. [synonym of Graphium angolanus baronis] Papilio pylades angolanus Goetz [sic]. Swanepoel, 1953. [synonym of Graphium angolanus baronis] Papilio angolanus Goeze, 1779. Dickson & Kroon, 1978. Graphium (Arisbe) angolanus Goeze, 1779. Pringle et al., 1994: 305.

Graphium angolanus angolanus. Male (Wingspan 68 mm). Left – upperside; right – underside. Nibela Peninsula, KwaZulu-Natal. November, 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium angolanus angolanus. Female (Wingspan 73 mm). Left – upperside; right – underside. Manguzi Forest, KwaZulu-Natal. 25 February 2010. M. Williams. Images M.C. Williams ex Williams Collection.

Alternative common names: Angola White-lady Swordtail; White Lady; Angolan White Lady. Type locality: Angola: “Angolae”. Distribution: Senegal, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Ivory Coast, Mali, Burkina Faso, Ghana, Togo, Benin (Fermon et al., 2001), Niger, Nigeria, Cameroon, Equatorial Guinea, Sao Tome & Principe, Gabon, Congo, Democratic Republic of Congo, Central African Republic, Chad, Sudan, Uganda, Rwanda, Burundi, Ethiopia, Kenya, Tanzania, Angola (Bivar de Sousa & Fernandes, 1966), Malawi, Zambia, Mozambique, Zimbabwe, Botswana, Namibia, South Africa, Swaziland, Comoro Islands. Habitat: Frost-free savanna (Pringle et al., 1994). In Tanzania it is found from sea-level to 2 200 m (Kielland, 1990d). It readily colonizes degraded forest habitats (Larsen, 2005a). In West Africa it is a savannah butterfly par excellence and flies well into the driest zones, only occasionally occurring in the forest zone (Larsen, 2005a). Habits: The flight is very fast, random, and from one to three metres above the ground. Males do not hilltop, as do those of G. morania, instead they have been noted flying up the spine of ridges in Lekgalameetse Nature Reserve, a behaviour that may be termed “ridging” (Williams, unpublished). Both sexes feed from flowers, such as those of Tridax (Larsen, 2005a) and males mud-puddle, often in large aggregations (Schultze, 1916; Pringle et al., 1994). Sometimes migrates, either on its own or as part of mixed migrations (Larsen, 1991c). Larsen (2005a) recounts that in 1968 his father saw a huge single-species migration of G. angolanus in the Guinea savanna of Nigeria, near the present Abuja. Larsen (1978) noted the species as part of a mixed migration north-east of Natitingou in Benin in April, 1978.

Flight period: September to April in southern Africa. Early stages:

Coleridge, 1916: 91 (Coleridge, P.L., 1916. J. of the E. Afr. and Uganda Nat. Hist. Soc. 5: 91-93).

Seth-Smith, 1938 [Ghana].

Clark, in Van Son, 1949: 36 [as Papilio (Graphium) pylades angolanus]. “Egg of the usual Papilionid shape, greenish-white, laid singly on the upper surface of a young leaf of the foodplant. Larva. First instar chocolate-coloured, with white dorsal markings on segments 6-10, pairs of spiny processes on thoracic segments (Coleridge says on segments 2, 4 and 5, which is probably incorrect) and white posterior end; second instar with the dorsal markings taking on a chess-board appearance; third instar with the dorsal markings in the shape of transverse bands, and with yellow longitudinal bands; fourth instar at first similar to the third, but before the final moult with greenish-yellow head stained with faint crimson; dorsal region green bordered at sides by a yellow spiracular line; a dorsal yellow line on segments 5-11; the yellow dorso-lateral processes of second segment are joined by a black transverse line, and the segment is margined posteriorly with black; third segment with two fine maroon transverse lines; fourth and fifth segments with the two dorso-lateral blue-black spines slightly wavy, joined by two maroon transverse bands, with the interspace pale blue; segments 6-11 marginal maroon, with two transverse maroon lines separated by a blue stripe overlaying the yellow dorsal line; the two posterior segments are slightly shaded with maroon; posterior tubercles yellow; lateral and ventral sides greenish stained with crimson, legs and prolegs green, the latter slightly hairy; final instar nearly two inches long, green with a pale yellow supra-spiracular line connecting the processes of the second segment with the posterior tubercles, and which carries on each segment two brighter chrome-yellow spots; second segment narrowly margined with black in front and behind; processes yellow tipped with black; segments 4 and 5 each with two simple blue-black dorso- lateral spines sprouting from orange tubercles; posterior tubercles yellow; osmeterium green; before pupation the larva turns a very pale, clear green. Larval period about two weeks. Pupa. Head with two fairly sharp points, thorax keeled and produced forward into a fairly long horn; body tapering gradually and evenly to the sharp posterior end; imitates the colour of the leaf of the food-plant, each segment being regularly veined; a pale green spiracular stripe from tip of horn to the posterior end; two longitudinal subdorsal slightly divergent stripes of similar colour from base of horn to posterior end where they unite. Pupal stage from eleven to twelve days.”

Darlow, 1949b.

Henning, S.F., 1984: 33.

Clark, in Pringle et al., 1994: p. 410; plate 36 [as Graphium (Arisbe) angolanus angolanus]. “The eggs are laid singly on the upper surface of the leaves of the foodplant. They are 0,95 mm in diameter and 0,8 mm high and are pure white when first laid, changing later to a pale yellowish brown and developing pale, salmon-pink maculae. The young larvae eat their way out near the top of the egg and consume the eggshell. The larva feeds on the surface of the leaves, at first eating small holes but later devouring the entire leaf. It moults where it is feeding and just before pupation turns pale green. The larval stage lasts about two weeks. The pupa is held upright by a silken girdle and is attached by the cremastral hooks to a silken pad. The pupal stage lasts about 12 days but up to two years has been recorded.”

Henning, Henning, Joannou, & Woodhall, 1997: 239 (photograph of early instar larva, final instar larva and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: Annona senegalensis Pers. (Annonaceae) [Platt (1921) cited by Van Son, 1949: 36; Seth-Smith, 1938 (Ghana)]. Friesodielsia obovata (Benth.) Verdc. (Annonaceae) [De Freina, 2009: 169; genus spelt as Frisodielsia]. Hexalobus monopetalus (A.Rich.) Engl & Diels (Annonaceae) [Congdon et al., 2009; eastern Zambia]. Landolphia buchannanii (Hallier f.) Stapf (Apocynaceae) [Van Someren, 1974: 316]. Landolphia buchannanii (Hallier f.) Stapf (Apocynaceae) [Van Someren, 1974: 316; as Landolphia ugandensis Staph.]. Sphedamnocarpus pruriens (A.Juss.) Szyszyl. (Malpighiaceae) [Platt (1921) cited by Van Son, 1949: 36]. Uvaria species (Annonaceae) [Kielland, 1990d: 46].

Graphium (Arisbe) angolanus angolanus (Goeze, 1779)#

Papilio eques achivus angolanus Goeze, 1779. Entomologische Beyträge zu des Ritter Linné zwölften Ausgabe des Natursystems 3 (1): 87 (390 pp.). Leipzig. Papilio pylades Fabricius. Trimen, 1862c. [Synonym of Graphium angolanus baronis] Papilio corinneus Bertoloni, 1849. Trimen & Bowker, 1889. [Synonym of Graphium angolanus angolanus] Papilio pylades Fabr., 1793. Aurivillius, 1899. [synonym of Graphium angolanus baronis] Papilio pylades angolanus Goetz [sic]. Swanepoel, 1953. Papilio angolanus Goeze, 1779. Dickson & Kroon, 1978. Graphium (Arisbe) angolanus angolanus Goeze, 1779. Pringle et al., 1994: 305.

Type locality: Angola: “Angolae”. Distribution: Angola (Bivar de Sousa & Fernandes, 1966), Democratic Republic of Congo (south), Kenya (coast), Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana (north), Namibia (north), South Africa (Limpopo Province, Mpumalanga, KwaZulu-Natal – north), Swaziland, Comoro Islands. Specific localities: Democratic Republic of Congo – Kimuenza (Schultze, 1913); Luebo (Dufrane, 1946); Banana (Dufrane, 1946); 30

Leopoldtville (Dufrane, 1946); Mulundu River (Dufrane, 1946). Tanzania – Throughout, including Pemba Island (Kielland, 1990d); Kilossa (Suffert, 1904); Msamwia (Strand, 1911); Kitungulu (Strand, 1911); Mtembwa River (Strand, 1911); Ngezi Forest, Pemba (Turlin & Lequeux, 1992); Jozani Forest, Zanzibar (Turlin & Lequeux, 1992) ; Katavi National Park (Fitzherbert et al., 2006). Malawi – Mt Mulanje (Congdon et al., 2010); Zomba Mountain (Congdon et al., 2010). Mozambique – Mt Chiperone (Timberlake et al., 2007); Mt Namuli (Congdon et al., 2010); Mt Mabu (Congdon et al., 2010); Mt Mecula [-12.0772 37.6297] (Congdon & Bayliss, 2013). Zimbabwe – Plumtree (Van Son, 1949); Marula (Van Son, 1949); Syringa (Van Son, 1949); Westacre (Van Son, 1949); Mount Selinda (male illustrated above). Botswana – Shakawe (Larsen, 1991l); Tsodilo Hills (Larsen, 1991l); Kachekawbe (Larsen, 1991l); Kasane (Larsen, 1991l); Kazungulu (Larsen, 1991l). Limpopo Province – Throughout bushveld areas (Swanepoel, 1953); Percy Fyfe Nature Reserve (Warren, 1990); Lekgalameetse Nature Reserve (“Malta Forest”); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015). Mpumalanga – Throughout bushveld areas (Swanepoel, 1953); Buffelspoort Nature Reserve (Williams). North West Province – Throughout bushveld areas (Swanepoel, 1953). KwaZulu-Natal – Durban (Swanepoel, 1953); Verulam (Swanepoel, 1953); Eshowe (Swanepoel, 1953); Empangeni (Swanepoel, 1953); Hluhluwe (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); Kosi Bay Nature Reserve (Swanepoel, 1953); Weenen (Swanepoel, 1953); Greytown (Swanepoel, 1953); Tembe Nature Reserve (Pringle & Kyle, 2002). Swaziland – Throughout bushveld areas (Swanepoel, 1953); Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz).

corinneus Bertoloni, 1850 (as sp. of Papilio). Memorie della Reale Accademia del Scienze dell’Instituto di Bologna. Class de Scienze Fisiche. Bologna (1) 2: 173 (165-188). Mozambique: “Mozambico”.

anthemenes Wallengren, 1857 (as sp. of Papilio). Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar. Stockholm annis 1838-1845. Collecta (n.s.) 2 (4): 6 (55 pp.). South Africa: “Caffraria”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/a). [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 579 (503-719))].

lapydes Suffert, 1904 (as ssp. of Papilio pylades). Deutsche Entomologische Zeitschrift, Iris 17: 103 (12-107). Tanzania: “Deutsch Ost Africa, Kilossa”.

kitungulua Strand, 1911 (as ab. of Papilio pylades angolanus ). Mitteilungen aus dem Zoologischen Museum in Berlin 5: 294 (275-304). Tanzania: “Msamwia; Kitungulu, Mtembwa Fluss”.

spoliatus Schultze, 1913 (as ab. of Papilio angolanus). Entomologische Rundschau 30: 50 (49-50). Democratic Republic of Congo: “Belgisch Kongo, Kimuenza am Stanleypool”.

dawanti Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Congo belge, Luebo”.

wansoni Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Bas Congo, Banana”.

jottrandi Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Congo, Léopoldville”.

deficiens Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Léopoldville”.

blariauxi Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Kasaï, rivière Mulundu, affluent de Longatschimo, 85 km. de Tshikapa”.

addenda Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades angolanus ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo: “Congo belge”.

howelli Turlin & Lequeux, 1992 (as ssp. of Graphium angolanus). Lambillionea 92 (4): 312 (311-321). Tanzania: “Ngezi forest, extrémité nord ouest de l’île Pemba, Tanzania, Juin 1988.” In M.N.H.N., Paris. Synonymized with Graphium angolanus (Goeze, 1779) by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 579 (503-719)). Note: Congdon et al., 2009 suggest that this taxon may be a valid subspecies of Graphium angolanus and that the matter requires investigation.

jozani Turlin & Lequeux, 1992 (as f. of Graphium angolanus howelli). Lambillionea 92 (4): 313 (311-321). Tanzania: “forêt de Jozani, sud de Zanzibar”. [Synonymized with G. angolanus (Goeze, 1779) by Smith & Vane- Wright, 2001. (Bulletin of the Natural History Museum Entomology Series 70 (2): 579 (503-719))].

corineus auct. (as sp. of Papilio). Vári et al., 2002: 129.

Graphium (Arisbe) angolanus baronis (Ungemach, 1932)

Papilio pylades baronis Ungemach, 1932. Mémoires de la Société des Sciences Naturelles (et Physiques) du Maroc 32: 21 (1-122). Graphium angolanus baronis (Ungemach, 1932). Smith & Vane-Wright, 2001.

Published images: Larsen, 2005a (male upperside and underside). Type locality: [Ethiopia]: “Baro”. Distribution: Senegal, Gambia, Guinea-Bissau, Guinea, Sierra Leone, Ivory Coast, Mali, Burkina Faso, Ghana, Togo, Benin (Fermon et al., 2001), Niger, Nigeria, Sao Tome & Principe, Cameroon, Gabon, Congo, Central African Republic, Democratic Republic of Congo (north – Ubangi, Mongala, Uele, Ituri), Chad, Sudan, Uganda, Rwanda, Burundi, Ethiopia, Kenya (west), Tanzania. Specific localities: Senegal – Dakar (Larsen, 2005a). Guinea – Mamou (Dufrane, 1946). Ghana – Kakum National Park (Larsen, 2006d); Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Togo – Klouto [6°57'15.07"N 0°34'54.40"E] (Safian et al., 2009). Benin – Noyau Central, Lama Forest (Fermon et al., 2001). Gabon – Bateke Plateau (van de Weghe, 2010); Franceville (van de Weghe, 2010); Lastoursville (van de Weghe, 2010); Mayumba (van de Weghe, 2010). Congo – Diélé (Boullet & Le Cerf, 1912); Brazzaville (Boullet & Le Cerf, 1912). Uganda – Semuliki N.P. (Davenport & Howard, 1996). Ethiopia – Baro (TL).

pylades Fabricius, 1793 (as sp. of Papilio). Entomologia Systematica emendata et aucta 3 (1): 34 (488 pp.). “Africa”. [Invalid; junior primary homonym of Papilio pylades Stoll, [1782]: [Castniidae].] [Synonymized with G. angolanus baronis (Ungemach, 1932) by Smith & Vane-Wright, 2001. (Bulletin of the Natural History Museum Entomology Series 70 (2): 582 (503-719))].

hypochroa Boullet & Le Cerf, 1912 (as ab. of Papilio pylades-angolanus). Bulletin de la Société Entomologique de France 1912: 246 (246-247). Congo: “Congo français, Diélé; Brazzaville”.

houzeaui Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) pylades pylades). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Guinea: “Guinée, vallée de Mamou”.

calabar Hancock, 1985 (as sp. of Graphium). Papilio International 2: 97 (97-103). “Africa”. [Synonymized with G. angolanus baronis (Ungemach, 1932) by Smith & Vane-Wright, 2001. (Bulletin of the Natural History Museum Entomology Series 70 (2): 581 (503-719))].

*Graphium (Arisbe) endochus (Boisduval, 1836)

Papilio endochus Boisduval, 1836. In: [Roret, Suites à Buffon] Histoire naturelle des Insectes. Species général des Lépidopteres 1: 243 (690 pp.). Paris. Papilio endochus Boisd., 1836. Aurivillius, 1899.

Type locality: Madagascar. Distribution: Madagascar (north and east). Habitat: Forest (Lees et al., 2003). Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) morania (Angas, 1849)# White Lady

Small White Lady Swordtails (Graphium morania). Images courtesy Steve Woodhall.

Papilio morania Angas, 1849. The Kafirs illustrated in a series of drawings taken among the Amazulu, Amaponda, and Amakosa tribes; [etc.]: [52] ([52] pp.). London. Papilio morania Angas, 1849. Trimen & Bowker, 1889. Papilio morania Angas, 1849. Aurivillius, 1899. Papilio morania Angas. Swanepoel, 1953a. Papilio morania Angas, 1849. Dickson & Kroon, 1978. Graphium (Arisbe) morania (Angas, 1849). Pringle et al., 1994: 305.

Graphium morania. Male (Wingspan 53 mm). Left – upperside; right – underside. Tembe, KwaZulu-Natal, South Africa. 19 April 2003. J. Dobson. Images M.C. Williams ex Dobson Collection.

Graphium morania. Female. Left – upperside; right – underside. Mount Selinda, Zimbabwe. 28 December, 1971. W. Teare. Images M.C. Williams ex Henning Collection.

Alternative common name: Small White-lady Swordtail. Type locality: [South Africa]: “Natal and the Zulu Country”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 585 (503-719))]. Distribution: Malawi (? MCW), Mozambique (south), Zimbabwe (south-east), Botswana (east), South Africa (Limpopo Province, Mpumalanga, Gauteng, KwaZulu-Natal), Swaziland. Specific localities: Botswana – Tswapong Hills (Larsen, 1991l). Limpopo Province – Waterberg (Van Son, 1949); Waterpoort (Van Son, 1949); Great Salt Pan (Van Son, 1949); Griffin Mine (Van Son, 1949), Pietersburg District (Van Son, 1949); Sibasa (Swanepoel, 1953); Duiwelskloof (Swanepoel, 1953); Gravelotte (Swanepoel, 1953); Nylsvley (Grei, 1986); Doorndraai Dam Nature Reserve (Warren, 1990); Lekgalameetse Nature Reserve (“Malta Forest”) (Williams) Highlands Wilderness (Bode & Bode, unpublished checklist); Bateleur Nature Reserve (Williams); Soetdoring Farm [-24.561 28.233] (A. Mayer, pers comm. 2015); Bateleur Nature Reserve (Williams & Dobson, unpub., 2015). Mpumalanga – Komatipoort (Swanepoel, 1953). Gauteng – Magaliesberg Mountains, Pretoria (Williams). KwaZulu-Natal – Durban (Van Son, 1949); Eshowe (Van Son, 1949); Umkomaas (Van Son, 1949); Oribi Gorge (Swanepoel, 1953); Scottburgh (Swanepoel, 1953); Isipingo (Swanepoel, 1953); Verulam (Swanepoel, 1953); Empangeni (Swanepoel, 1953); Mtubatuba (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); Nongoma (Swanepoel, 1953); Kosi Bay Nature Reserve (Swanepoel, 1953); Umzimkulu River (Pringle et al., 1994); Tembe Nature Reserve (Pringle & Kyle, 2002). Swaziland – Throughout (Swanepoel, 1953); Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz).. Habitat: Frost-free savanna (Pringle et al., 1994). Habits: In this endemic southern African species the flight is fast, erratic and from one to three metres above the ground. Both sexes feed from flowers. Males are often found mudpuddling, sometimes in enormous assemblages. In October, 1973 hundreds of males were seen mudpuddling on the banks of the Sand River at Waterpoort in Limpopo Province (Williams, unpublished). Males are somewhat territorial and are known to defend territories on hill-tops as well as on slopes and flat ground (Williams, unpublished). Flight period: September to May (Pringle et al., 1994). Early stages:

Trimen & Bowker, 1889: 221 [as Papilio Morania]. “Larva. Dull-green with a yellowish tinge; two parallel darker transverse lines across back of each segment; incisions of segments pale bluish-grey. First thoracic segment and lateral stripe bright-yellow, the latter deepening to orange on last segment, and edged inferiorly throughout by dark-green. Lower portion of sides pale bluish-green, with a bluish- white stripe immediately above legs. Head pale-green. Each thoracic segment with a pair of short, pointed, laterally projecting spines, of which the first pair is shortest and mainly black, but sandy-yellow at base, - the second ferruginous 34 basally, but thence black, - and the third whitish tinged with blue; between the bases of the second and third pairs a transverse ferruginous-brown streak (incomplete on second thoracic segment). Anal segment bearing dorsally a pair of terminal, short, acute, yellow spines. A small example in an earlier moult is very different, having three dorsal longitudinal blackish lines, of which the middle one is thinnest, the transverse darker dorsal lines more developed, and a blackish transverse bar on the penultimate segment. The lower sides and the under side are fuscous, the head is sandy-yellow, the thoracic spines are proportionally larger, and the transverse bars between the second and third pairs broader and much darker. Pupa. Stout, rounded, but tapering posteriorly to rather a long point; not depressed or widened dorsally, as in Brasidas [= leonidas]. Not nearly so blunt and rounded anteriorly as in Brasidas, the cephalic prominences, though short, being acute and widely divergent. Dorso-thoracic peak not so long as in Brasidas, and more ascendant, so as not to project so far in front of the head. About one inch in length. Bright-green; a well-defined, raised, pale-yellow line curving each side from apex of dorso-thoracic peak, and meeting at anal extremity. On back a similar thinner line, starting from same peak (but near its base), almost immediately branches into two gently divergent lines, which from third abdominal segment gradually converge, to meet again at anal extremity. Between these lateral and dorsal lines, on each side a series of very thin V-like yellow marks (with the angles directed backward), and all truncated by the segmental incisions. On third abdominal (seventh) segment, a small ferruginous spot in a dull-creamy ring adjoins each dorsal line on its inner side about the middle. About midway between the lateral line and the cephalic prominence on each side, a somewhat sinuous, less distinct, pale- yellow line, which on wing-cover marks position of median nervure, and emits thin indications of positions of nervules. In addition to these principal pale-yellow markings, there are numerous minor reticulations and dottings of the same colour distributed over the surface. Frontal line of dorso-thoracic peak, from its apex to head, minutely roughened, and coloured dull-ferruginous, with a few creamy specks. The larva is described from four Natalian specimens (one of an early moult) forwarded by Colonel Bowker in November 1878. I did not rear the perfect insect from them, but two of them became pupae quite like those of Morania; and coloured drawings (accompanied by the wings of that species) lent me by Mrs. Monteiro in 1883 (made from Delagoa Bay larvae), agreed closely with my description above given. Of the pupa, I received from Colonel Bowker thirteen living specimens, and obtained the imago from each one, ten butterflies appearing between 24 th November and 16th December 1878. Of the remaining three, two did not make their appearance before a year afterwards (14th November and 9th December 1879), and the last not until 14 th March 1880. Colonel Bowker noted the date of pupation in five instances, so that I can record the normal duration of the chrysalis state in those few cases as varying from twelve to sixteen days. It is noticeable that in outline this pupa does not simulate a complete leaf so well as that of P. Brasidas [= Graphium leonidas], but the anterior extremity, with the concave, roughened, and ferruginous-tinted line from the tip of the thoracic peak to the head, gives precisely the effect of the irregular edge of a leaf gnawed and partly eaten away by insects. The under side of the mid-rib of a leaf is usually the situation of the suspended pupa; in two instances only, among the specimens sent by Colonel Bowker, was the insect attached to the twigs of the foodplant. As regards the surface of the pupa, the yellow lines, V-streaks, and minuter markings admirably represent the aspect of the leaves, and effectually conceal the insect among the foliage.”

Clark, in Van Son, 1949: 38 [as Papilio (Graphium) morania]. “Egg. 1 mm in diameter by 0.95 mm high. Spheroidal with flattened base, pale watery greenish-yellow when laid, with a very faint irregular hexagonal reticulation. Laid singly on young shoot of the food-plant. Egg-stage about 4 days. Larva. There are five larval instars lasting only about 15 days; first instar 2.5 mm long when hatched, with brownish- yellow thoracic and last three abdominal segments, lighter elsewhere, with three dorsal dark brown lines widened at middle of the segments; sides whitish, with blackish longitudinal lines; there are three pairs of blunt dorso-lateral processes on thoracic segments and a pair of posterior processes; all these processes are densely covered with peculiar distally-forked setae which are also present on the dorsal surface of the thoracic segments; second instar lighter, most of the ground colour being white, with the lines blackish; the setose processes are yellow, and the setae are simple; a double dorsal row of yellow spots on 4th to 11th segments, bordering the dorsal line; third and fourth instars rather similar, yellow with black transverse band on posterior sides of second and third thoracic segments, and with narrow black longitudinal lines; lower half of body brownish; final instar green; two parallel darker transverse lines or rows of dots across the middle, and a third similar row at the anterior edge of each abdominal segment; each thoracic segment with a pair of acute laterally projecting spines, of which the pair on the first segment is shortest and mainly black, but sandy-yellow at base, the second ferruginous basally, but thence black and the third whitish, tinged with blue (Mr. Clark’s painting shows the first pair black, and both the second and third pairs black with crimson bases); between the bases of the second and third pair a transverse ferruginous-brown streak (incomplete on second segment) (Mr. Clark’s painting shows a reticulated brown marking on third thoracic and first abdominal segment; presumably there is a variation in this respect); last segment with a pair of terminal, short, acute diverging yellow spines.”

Clark, in Pringle et al., 1994: p. 412; plate 37 [as Graphium (Arisbe) morania]. “The eggs are laid singly on young leaves or shoots. They are 1 mm in diameter and 0,95 mm high and are a pale, watery green. The egg stage lasts about four days. There are five larval instars lasting approximately 15 days. The pupa is 35 attached by the cremastral hooks to a silken pad and supported by a silken girdle. The pupal stage can last for a couple of weeks or even up to two years.”

Henning, Henning, Joannou, & Woodhall, 1997: 241 (photograph of early instar larva, final instar larva and pupa).

Larval food: Annona senegalensis Pers. (Annonaceae) [Pringle et al., 1994: 305]. Artabotrys brachypetalus Benth. (Annonaceae) [Pringle et al., 1994: 305]. Artabotrys monteiroae Oliv. (Annonaceae) [Pringle et al., 1994: 305]. Artabotrys species (Annonaceae) [Bowker, in Trimen & Bowker, 1889: 221; as Artabotrys n. sp.; Durban, KwaZulu-Natal, and Delagoa Bay, Mozambique]. Hexalobus monopetalus (A.Rich.) Engl. & Diels (Annonaceae) [Williams, in Pringle et al., 1994: 305]. Uvaria caffra E.Mey. ex Sond. (Annonaceae) [Dickson & Kroon, 1978].

vansoniana Storace, 1953 (as f. of Graphium morania). Annali del Museo Civico di Storia Naturale (di Genova) Giacomo Doria 66: 253 (252-264). South Africa: “Natal, Unkomaas”.

holoplaga Grei, 1986 (as var. of Graphium morania). Fauna and Flora. Pretoria No. 44: 11, 10 (10-12). South Africa: “Transvaal, Boekenhout, Nylsvley”.

*Graphium (Arisbe) taboranus (Oberthür, 1886)

Papilio taboranus Oberthür, 1886. Bulletin de la Société Entomologique de France (6) 6: cxiv (114-115).

Graphium taboranus. Male. Left – upperside; right – underside. Wingspan: 53mm. Lumbumbashi, Katanga. Nov 71. Dr Y. Allard. (Transvaal Museum – TM2956).

Type locality: [Tanzania]: “Tabora, dans l’Ounyanyambé”. Distribution: Angola, Democratic Republic of Congo (Shaba), Tanzania (central, west and south), Malawi, Zambia (north). Recorded, in error, from Southern Africa by Dickson & Kroon, 1978. Specific localities: Democratic Republic of Congo – Lumbumbashi (male illustrated above). Tanzania – Ounyanyambe, Tabora (TL); southern, central and western Tanzania, in the Mpanda District (Kielland, 1990d). Malawi – Mt Mulanje (Congdon et al., 2010).

Zambia – Ikelenge (Heath et al., 2002); the Copperbelt (Heath et al., 2002); Mumbwa (Heath et al., 2002); Mansa (Heath et al., 2002); Chambeshi River (Heath et al., 2002); Lofu (Lufubu) River (Heath et al., 2002); Mporokoso (Heath et al., 2002); Kawambwa (Heath et al., 2002). Habitat: Woodland, including Brachystegia woodland. In Tanzania at altitudes of 1 000 to 1 600 m (Kielland, 1990d). Early stages:

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: Annona species (Annonaceae) [Kielland, 1990d: 49]. ?Annona senegalensis Pers. (Annonaceae) [Congdon et al., 2009]. ?Hexalobus monopetalus (A.Rich.) Engl & Diels (Annonaceae) [Congdon et al., 2009; eastern Zambia].

nivinox Butler, 1894 (as sp. of Papilio). Proceedings of the Zoological Society of London 1893: 667 (643-684). “Lake Mweru”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 587 (503-719))].

*Graphium (Arisbe) schaffgotschi (Niepelt, 1927) Schaffgotsch’s Swordtail

Papilio schaffgotschi Niepelt, 1927. Internationale Entomologische Zeitschrift 21: 53 (53). Graphium (Arisbe) schaffgotschi (Niepelt, 1927). Pringle et al., 1994: 305.

Graphium schaffgotschi. Male. Left – upperside; right – underside. Wingspan: 56mm. Zambia, N.W. Province, Kasangeji, 11° 40'S; 24° 25'E. 1500 m. 25.X.1999. F.M. Kayombo. (Newport Collection).

Graphium schaffgotschi. Female. Left – upperside; right – underside. Wingspan: 63mm. Zambia, N.W. Province, Kasangeji, 11° 40'S; 24° 25'E. 1500 m. 22.I.1997. F.M. Kayombo. (Newport Collection).

Type locality: [Namibia]: “Deutsch Sudw. Africa”. [False locality.] Diagnosis: Similar to G. taboranus, from which it differs as follows: larger; white areas on upperside of wings more extensive; on the forewing upperside the dark band between the white discal area and postdiscal spots is narrower (Pringle et al., 1994). Distribution: Angola (Bivar de Sousa & Fernandes, 1966), Democratic Republic of Congo (south), Zambia (north-west). Recorded, in error, from Southern Africa by Dickson & Kroon, 1978. Specific localities: Democratic Republic of Congo – Zilo, Katanga (Pringle et al., 1994). Zambia – Mwinilunga (Heath et al., 2002); Ikelenge (Heath et al., 2002); Mongu (Heath et al., 2002); Kasangezhi (male and female illustrated above). Habitat: Savanna. Flight period: All year (Pringle et al., 1994). Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) ridleyanus (White, 1843) Acraea Swordtail

Papilio ridleyanus White, 1843. Annals and Magazine of Natural History 12 (1): 262 (262-268). Papilio ridleyanus White, 1843. Aurivillius, 1899.

Graphium ridleyanus. Male. Left – upperside; right – underside. Wingspan: 66mm. Bangui, R. C. A. 78.12.03. R.P. Godart. (Curle Trust Collection – 8).

Graphium ridleyanus. Female. Left – upperside; right – underside. Wingspan: 77mm. Bangui, R. C. A. 1978.01.07. R.P. Godart. (Curle Trust Collection – 9).

Type locality: [Democratic Republic of Congo]: “oras fluminis Zaire”. [Lectotype designated by Smith & Vane- Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 590 (503-719))]. Distribution: Nigeria (east), Cameroon, Equatorial Guinea, Sao Tome & Principe, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Chad, Sudan (south), Uganda, Rwanda, Burundi, Tanzania (west), Zambia. Records for Sierra Leone (Berger, 1981; Mathot, 1990) and Benin (Smith & Vane-Wright, 2001) are regarded as erroneous (Larsen, 2005a). Specific localities: Nigeria – Okwangwo (Larsen, 2005a); Gashaka-Gumpti (Larsen, 2005a). Cameroon – Kulembembe (Schultze, 1913). Gabon – Lambarene (van de Weghe, 2010); Waka (van de Weghe, 2010); Mouila (van de Weghe, 2010); Tchibanga (van de Weghe, 2010); Oyem (van de Weghe, 2010); Nouna (van de Weghe, 2010); Ipassa (van de Weghe, 2010); Langoue (van de Weghe, 2010); Bakouaka (van de Weghe, 2010); Franceville (van de Weghe, 2010); Mpassa, Bateke Plateau (van de Weghe, 2010). Central African Republic – Dzanga (Noss, 1998); Mangoumba (Schultze, 1930); Bangui (specimens illustrated above). Democratic Republic of Congo – Kassai River (Niepelt, 1915); Micozi River (Dufrane, 1946). Sudan – Yambio (Holland, 1920); Bahrel Ghazal (Holland, 1920); Njam Njam (Röber, 1928). Uganda – Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Tanzania – Upper Ruwubu River (Le Cerf, 1924); from the Mahale Mountains and Leugele River to just south of Kigoma (Kielland, 1990d). Zambia – Kalene Hill (Ikelenge) (Heath et al., 2002). Habitat: A forest species which will visit flowering trees some distance from forest. Larsen (2005a) regards it as a forest/savanna transition zone species. In Tanzania at altitudes from 780 to 1 200 m (Kielland, 1990d). Habits: This is quite a rare species but in certain localities and at certain times it may be more numerous (Larsen, 2005a). A mimic of large Acraea species, particulary A. egina (Kielland, 1990d) and A. perenna (Birket-Smith, 1960; Larsen, 2005a). As noted by Larsen (2005a), Schultze (1917) pointed out that the species rarely flies in the same places as A. egina. Some females have the red areas obfuscated, then resembling A. zetes and A. pharsalus (Larsen, 2005a). Males are known to mud-puddle (Kielland, 1990d). Early stages: Nothing published. Larval food: Monanthotaxis laurentii (De Wild.) Verdc. (Annonaceae) [Fontaine, 1985 (Democratic Republic of Congo); as Popowia congoensis].

infuscatus Schultze, 1913 (as ab. of Papilio ridleyanus). Archiv für Naturgeschichte 79 (A.7.): 5 (4-5). Cameroon: “Süd-Kamerun, Kulembembe”.

fumatus Niepelt, 1915 (as var. of Papilio ridleyanus). Internationale Entomologische Zeitschrift 9: 58 (58). Democratic Republic of Congo: “Kassai-Fluss, Kongostaat”.

fumosus Holland, 1920 (as var. of Papilio ridleyanus). Bulletin of the American Museum of Natural History 43: 247 (109-369). Sudan: “S. Sudan, Yambio, Bahrel Ghazal”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

rosa Le Cerf, 1924 (as f. (? an subsp.) of Papilio ridleyanus). Bulletin of the Hill Museum, Witley 1: 394 (369-399, 576, 578, 580, 582, 584). Tanzania: “E. Tanganyika, Upper Ruwubu River, Urindi Distr.”.

njami Röber, 1928 (as ssp. of Papilio ridleyanus). Internationale Entomologische Zeitschrift 22: 67 (67-72). Sudan: “Njam Njam”.

semivitreus Schultze, 1930 (as ab. of Papilio ridleyanus). Deutsche Entomologische Zeitschrift, Iris 44: 176 (176- 188). Central African Republic: “Mongoumba”.

hecqueti Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) ridleyanus). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 118 (101-122). Democratic Republic of Congo: “Sud Kivu, rivière Micozi, zône des mines M.G.L.”.

The leonidas group In the analyses of Smith & Vane-Wright (2001) this group of three species, usually placed together, were found to be paraphyletic.

*Graphium (Arisbe) leonidas (Fabricius, 1793)# Veined Swordtail

Veined Swordtails (Graphium leonidas). Left – perching, hilltopping male (image courtesy Steve Woodhall). Centre – mudpuddling male, Zambia (image courtesy Raimund Schutte). Right – Female (image courtesy Steve Woodhall).

Papilio leonidas Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 35 (488 pp.). Hafniae. Papilio leonidas Fabricius. Trimen, 1862c. Papilio leonidas Fabricius, 1793. Trimen & Bowker, 1889. Papilio brasidas Felder, 1864. Trimen & Bowker, 1889. [Synonym of Graphium leonidas leonidas] Papilio leonidas Fabr., 1793. Aurivillius, 1899. Papilio leonidas Fabricius. Swanepoel, 1953a. Papilio leonidas Fabricius, 1793. Dickson & Kroon, 1978. Graphium (Arisbe) leonidas (Fabricius, 1793). Pringle et al., 1994: 305.

Graphium leonidas leonidas. Male (Wingspan 78 mm). Left – upperside; right – underside. Lekgalameetse N.R., Limpopo Province, South Africa. 6 December, 2012. M. Williams. Images M.C. Williams ex Williams Collection.

Graphium leonidas leonidas. Male form brasidas (Wingspan 78 mm). Left – upperside; right – underside. Umtamvuna River, KwaZulu-Natal, South Africa. 22 December, 2009. J. Dobson. Images M.C. Williams ex Dobson Collection.

Graphium leonidas leonidas. Female. Left – upperside; right – underside. Malta Forest, Limpopo Province, South Africa. December, 1957. W. Teare. Images M.C. Williams ex Henning Collection.

Graphium leonidas leonidas. Female form brasidas (Wingspan 80 mm). Left – upperside; right – underside. Umtamvuna River, KwaZulu-Natal, South Africa. 20 December, 2008. J. Dobson. Images M.C. Williams ex Dobson Collection.

Type locality: “Africa”. Distribution: Sub-Saharan Africa, including Senegal, Gambia, Guinea-Bissau, Guinea, Mali, Burkina Faso, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2001), Nigeria, Cameroon, Sao Tome & Principe, Equatorial Guinea, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Chad, Sudan, Ethiopia, Uganda, Rwanda, Burundi, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana, South Africa, Swaziland. Habitat: Frost-free savanna and the ecotone between savanna and forest. In Tanzania from sea-level to 2 200 m. Very common in the Brachystegia woodlands of western Tanzania (Kielland, 1990d). Habits: The flight pattern of this common, widespread species is fast and erratic. It is both highly vagile and migratory (Larsen, 1991c; Larsen, 2005a). Both sexes regularly visit flowers and males are often found mudpuddling, sometimes in large species assemblages. When roosting they often hang from a twig with folded wings, thereby enhancing their resemblance to the distasteful danaine Tirumala petiverana (Doubleday). Sevastopulo (1973) maintains that leonidas itself may be toxic and thus be a Müllerian rather than a Batesian mimic. This seems improbable, given the larval host-plants. Males are strongly territorial, establishing and defending these on hilltops and ridges or in clearings in the bush. Within these territories males select a perch from two to five metres above the ground. Territories are usually about 10 to 20 metres in diameter and are patrolled with a circling, gliding flight, some metres above the ground. Flight period: All year, but more plentiful in the midsummer months. Early stages:

Trimen & Bowker, 1889: 213 and 215 [as Papilio Leonidas Fabricius (p. 213) and as Papilio Brasidas Felder (p. 215)]. Page 213: “A pupa [of Leonidas] received from Mrs. Monteiro resulted in a ♂ on the 9th December 1886. I could not discover that this Delagoa Bay specimen differed at all from the chrysalides of Brasidas from Natal, which are described below.” Page 215: “Larva (second moult). Yellowish-green above, paler on the sides, pale-green beneath; back of segments 3, 4, and 5 occupied by a lozenge-shaped space of rich dark-green. A median dorsal narrow stripe, whitish on thoracic segments, but beyond that dull pinkish-yellow, edged on both sides with dark-green. Second segment reddish-brown superiorly, with a short but acute tubercular process on each side; third and fourth segments each bearing dorso-laterally a pair of short acute yellow tubercular processes, edged inwardly and basally by an ovate black spot, and connected by a transverse reddish-brown band; a blackish line runs longitudinally from each ovate black spot on third segment to the reddish-brown second segment. On each side, from sixth to last segment, a series of seven oblique slender dark-green lines; also a longitudinal (spiracular) dull-whitish stripe, commencing superiorly on third segment, curving downward on fifth, and thence running straight to extremity. Anal segment bearing superiorly a pair of rather long acute divergent processes. Retractile forked scent organ of second segment deep indigo-blue. (Adult). Paler and brighter green above, and of a bluer tint beneath. Dorsal markings, except reddish-brown of second segment and black spots of third and fourth segments, apparently very much fainter and partly obsolete; and the dorsal and lateral stripes thinner dark-green. Segmental incisions marked, shining.” “Pupa. Semi-transparent. Bright yellowish-green. On each side of back, from apex of dorso-thoracic projection to anal extremity, a narrow ochrey-yellow marginal ridged or elevated narrow stripe; on summit of back a pair of similar ridged

42 stripes, commencing abruptly and considerably apart near base of dorso-thoracic projection, and converging till they meet at anal extremity; a slender dark-green dorsal madian line. On each side, a series of oblique thin streaks, rather darker than ground-colour, extending from near dorsal median line to marginal yellow ridge (and beyond it on cephalic area). Very thick and robust, especially towards head, which is remarkably blunt and sub-rotund. Dorsally much concaved and widened about middle; dorso-thoracic peak prolonged upward and forward a good deal beyond front of head, and slightly curved; anal extremity elongate and attenuated. Attached to mid-rib or to foot-stalk and mid-rib of under side of leaves; usually head downward. The above description of the larva is made from drawings and notes of Natalian examples furnished by Captain H.C. Harford in 1869 and 1872; that of the pupa is from living Natalian specimens received from Colonel Bowker in 1878. With the latter arrived two larvae, one already suspended for pupation, the other in the stage described as the “second moult” by Captain Harford. Only the former of these larvae produced the perfect insect, but I obtained ten butterflies from the pupae received. Of these, the dates of pupation of nine being on record, I am able to note that the duration of the chrysalis state in November and December varied from twelve to eighteen days. These eleven pupae were (with the exception of two on the twigs of the food-plant, and one – from the larva that pupated after arrival – attached to the side of the box) suspended on the under side of the leaves, the tail being fixed at the base of the mid-rib, or, in a few cases, just free of the beginning of the leaf, to the short foot-stalk. In this position, the difficulty of distinguishing the pupa from the adjacent leaves is considerable, as the concavity and depression of the dorsal aspect, from the attenuated tail to the prominent and curved thoracic peak (which intercepts the view of the blunted head), and the outline of this aspect, the resemblance of the four raised dorsal stripes converging at the tail to a mid-rib or short foot-stalk, and the slightly darker oblique streaks and minute flecks representing the venation, all combine to assimilate the pupa to the surrounding foliage in the most deceptive manner.”

Clark, in Van Son, 1949: 41 [as Papilio (Graphium) leonidas]. “Egg. Pale watery-yellow, deepening slightly in colour; almost spherical, laid singly on young shoot of the foodplant. Egg-stage lasts about 4 days. Larva. There are five larval instars lasting altogether about 25 days. First instar: 2.5 mm long on hatching, pale yellow with light brown diagonal stripes and darker brown dorsal and subdorsal lines; the body of the larva thickens from the head to the third segment, then tapers to the forked last segment; paired spiny processes are present on first three segments, and numerous forked setae on all body segments. Second instar: darker than first, and forked setae replaced by short single ones. Third instar: yellowish-green above, paler on sides, pale green beneath; back of segments 3, 4 and 5 occupied by a lozenge-shaped space of rich dark green; a median dorsal narrow stripe, whitish on thoracic segments, but beyond that dull pinkish-yellow, edged on both sides with dark green; first thoracic segment reddish- brown above, with short acute lateral processes; second and third segments with the lateral processes yellow, edged with black and connected by a transverse reddish-brown band; a blackish line runs longitudinally from each black edging to the reddish-brown first segment; lateral diagonal lines dark green; there is a lateral (spiracular) whitish stripe commencing superiorly on second segment, curving downwards on fourth and thence running straight to the posterior end; caudal processes acute and divergent; osmeterium deep indigo blue. Fourth instar: deep brown, with darker diagonal stripes and lighter longitudinal lines: a double dorsal line on thoracic and first abdominal segments, thence continued as a single line to posterior end, and two lines on each side of the thoracic and first two abdominal segments, the outer one along the bases of the spiny processes; traces of a third line on second to fourth abdominal segments, between the dorsal and the first subdorsal lines. Fifth instar: brown after moulting, but gradually changing to dull green, with a light brown dorsal, broad longitudinal stripe on thoracic and first abdominal segments except the posterior part of the first segment, widened on anterior part of first segment and crossed by transverse bands of the same colour along posterior edges of second and third segments; this marking is edged by dark green forming a lozenge-shaped patch narrowly edged with yellowish-white; intersegmental incisions marked and shining; length up to 45 mm. Pupa. Fully described by Trimen” (see above). “The unusual head-downward position of the pupa observed by Trimen is due to the extreme thinness of the silken girdle by means of which the pupa is originally fastened in an upright position (as is usual in Papilios), and which often breaks, causing the pupa to hang freely (as in the Nymphalidae).”

Carcasson, 1981 [larva and pupa].

Clark, in Pringle et al., 1994: p. 414; plate 38 [as Graphium (Arisbe) leonidas leonidas]. “The eggs are laid singly on the young shoots of the foodplant. They are pale, watery yellow when first laid, deepening slightly in colour with age. The young larva eats its way out of the side of the egg near the top and generally devours the eggshell. The egg stage lasts about four days. The larva rests on a midrib of a young leaf and crawls away to feed. At first it eats small holes in the leaves but later it feeds on the edge of the leaves. There are five larval instars lasting about 25 days. The pupa is very blunt-headed compared with others of the genus. This stage lasts from a couple of weeks to several months. The pupa is attached by the cremastral hooks to a silken pad and is supported by a silken girdle.”

Henning, Henning, Joannou, & Woodhall, 1997: 236 (photograph of early instar larva, final instar larva and pupa).

Congdon et al., 2009 [photographs of larva and pupa].

Graphium leonidas egg (left) and 1st instar larva (right). Images courtesy Raimund Schutte (left) and Steve Woodhall (right).

Graphium leonidas 4th instar larva (left) and final instar larva (right). Images courtesy Raimund Schutte (left) and Steve Woodhall (right).

Graphium leonidas pupa. Image courtesy Steve Woodhall.

Larval food: Annickia chlorantha (Oliv.) Setten & Maas (Annonaceae) [Lees, 1989 (Korup, Cameroon)]. Annona senegalensis Pers. (Annonaceae) [Van Someren, 1974: 316]. Artabotrys brachypetalus Benth. (Annonaceae) [Woodhall, 2005a]. Artabotrys cinerea (Annonaceae) [Henning, Henning, Joannou & Woodhall, 1997: 237]. Friesodielsia obovata (Benth.) Verdc. (Annonaceae) [Larsen, 1991l (Kasane and Shakawe, Botswana); Paré, in Pringle et al., 1994: 306 (Zimbabwe)]. Landolphia buchannani (Hallier f.) Stapf (Apocynaceae) [Van Someren, 1974: 316]. Landolphia buchannani (Hallier f.) Stapf (Apocynaceae) [Van Someren, 1974: 316; as Landolphia ugandensis Staph.]. Monanthotaxis caffra (Sond.) Verdc. (Annonaceae) [Harford, in Trimen & Bowker, 1889: 215; as Popowia Caffra; KwaZulu-Natal].

Monanthotaxis fornicate (Baill.) Verduc. (Annonaceae) [Congdon, et al., 2009]. Uvaria acuminata Oliv. (Annonaceae) [Heath et al., 2002: 26]. Uvaria caffra E.Mey. ex Sond. (Annonaceae) [Williams, 1996: 134; near Tembe Elephant Reserve and Makatini Flats, KwaZulu-Natal]. Uvaria kirkii Hook.f. (Annonaceae) [Heath et al., 2002: 26]. Uvaria leptocladen Oliv. (Annonaceae) [Van Someren, 1974: 316].

Note: Congdon et al. (2009) state that populations of leonidas on Bioko Island may not be synonymous with the nominate subspecies on the mainland and should be investigated with respect to their taxonomy.

Graphium (Arisbe) leonidas leonidas (Fabricius, 1793)#

Graphium leonidas leonidas. Male form brasidas (Wingspan 78 mm). Left – upperside; right – underside. Umtamvuna River, KwaZulu-Natal, South Africa. 22 December, 2009. J. Dobson. 45

Images M.C. Williams ex Dobson Collection.

Graphium leonidas leonidas. Female. Left – upperside; right – underside. Malta Forest, Limpopo Province, South Africa. December, 1957. W. Teare. Images M.C. Williams ex Henning Collection.

Type locality: “Africa”. Diagnosis: There are two distinct forms (the nominate and f. brasidas) (Pringle et al., 1994). Distribution: Sub-Saharan Africa, including Senegal (south), Gambia, Guinea-Bissau, Guinea, Mali, Burkina Faso, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin (Fermon et al., 2001), Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Chad, Sudan, Ethiopia, Uganda, Rwanda, Burundi, Kenya, Tanzania, Malawi, Zambia, Mozambique, Zimbabwe, Botswana (north-east), South Africa (Limpopo Province, Mpumalanga, KwaZulu-Natal, Eastern Cape Province), Swaziland. Specific localities: Ghana – Ankasa (Larsen, 2005a); Kyabobo N.P. (Larsen, 2005a); Kakum National Park (Larsen, 2006d); Bobiri Butterfly Sanctuary (Larsen et al., 2007); Boabeng-Fiema Monkey Sanctuary (Larsen et al., 2009). Togo – Klouto [6°57'15.07"N 0°34'54.40"E] (Safian et al., 2009). Benin – Noyau Central, Lama Forest (Fermon et al., 2001); Lokoli (Tchibozo et al., 2008). Cameroon – clear-felled forest in Korup (Larsen, 2005a). Gabon – Almost throughout (van de Weghe, 2010). Central African Republic – Chari (Berger, 1950); Djema (Berger, 1950); Dzanga (Noss, 1998). Democratic Republic of Congo – Luebo (Dufrane, 1946); Camayenne (Dufrane, 1946); Sankisishia (Dufrane, 1946); Stanleyville (Berger, 1950); Lukombe (Berger, 1950). Sudan – Between Dalba to the Uma River (Rothschild & Jordan, 1905); Konta (Rothschild & Jordan, 1905). 46

Uganda – Mount Elgon (Bryk, 1928); Busia (Bryk, 1928); Semuliki N.P. (Davenport & Howard, 1996); Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Tanzania – Muanza (Suffert, 1904); Katavi National Park (Fitzherbert et al., 2006); Mkwaja Ranch (Congdon et al., 2009). Malawi – Mt Mulanje (Congdon et al., 2010); Zomba Mountain (Congdon et al., 2010). Mozambique – Maputo (Van Son, 1949). Zimbabwe – Victoria Falls (Storace, 1953); Chirinda Forest (Van Son, 1949). Botswana – Kasane (Larsen, 1991l); Shakawe (Larsen, 1991l); Shashe (G. Bailey vide Larsen, 1991l); Nata (N. Ashby vide Larsen, 1991l); Serowe (Larsen, 1991l); Kazungulu (Larsen, 1991l). Limpopo Province – Throughout bushveld areas (Swanepoel, 1953); Lekgalameetse Nature Reserve (“Malta Forest”) (Van Son, 1949); Shilouvane (Van Son, 1949). Mpumalanga – Throughout bushveld areas (Swanepoel, 1953); Barberton (Van Son, 1949); Marieps Kop (Van Son, 1949); Buffelspoort Nature Reserve (Williams). North West Province – Throughout bushveld areas (Swanepoel, 1953). KwaZulu-Natal – Durban (Felder & Felder, 1864); Malvern (Le Cerf, 1924); Shangwana (Van Son, 1949); Eshowe (Van Son, 1949); Oribi Gorge (Swanepoel, 1953); Port Shepstone (Swanepoel, 1953); Ixopo (Swanepoel, 1953); Umkomaas (Swanepoel, 1953); Pietermaritzburg (Swanepoel, 1953); Estcourt (Swanepoel, 1953); Empangeni (Swanepoel, 1953); Mtubatuba (Swanepoel, 1953); St. Lucia Bay (Swanepoel, 1953); Magut (Swanepoel, 1953); Kosi Bay Nature Reserve (Pringle & Kyle, 2002); Tembe Nature Reserve (Pringle & Kyle, 2002); Ngome Forest (male illustrated above). Eastern Cape Province – Ngqeleni (Van Son, 1949); Port St Johns (Van Son, 1949); East London (Swanepoel, 1953); King William’s Town (Swanepoel, 1953); Bashee River (Swanepoel, 1953); Umtata (Swanepoel, 1953); Cintsa East (the Dobsons and Williams); Haga Haga (the Dobsons and Williams); Cwebe-Dwesa Nature Reserve (E. Pringle, Jan., 2015). Swaziland – Throughout bushveld areas (Swanepoel, 1953); Mlawula N. R. (www.sntc.org.sz); Malolotja N. R. (www.sntc.org.sz).

similis Cramer, 1775 in Cramer, [1775-6] (as sp. of Papilio). Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America 1: 14 (155 pp.). Amsteldam and Utrecht. Africa: “China; Java”. [False locality.] [Invalid; junior primary homonym of Papilio similis Linnaeus, 1758 [Danainae].]

brasidas Felder & Felder, 1864 (as sp. of Papilio). Verhandlungen der Zoologische-Botanischen Gesellschaft in Wien 14: 307 (289-378). South Africa: “Zulu, Port Natal”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 596 (503-719))].

interniplaga Aurivillius, 1899 in Aurivillius, 1898-9 (as ab. (var.?) of Papilio leonidas). Kungliga Svenska Vetenskapakademiens Handlingar 31 (5): 487 (1-561). Tanzania: “Deutsch Ost-Afrika”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/l).

onidale Suffert, 1904 (as sp. of Papilio). Deutsche Entomologische Zeitschrift, Iris 17: 106 (12-107). Tanzania: “Muanza am Südufer des Victoria Nyanza”.

umanus Rothschild & Jordan, 1905 (as ssp. of Papilio similis). Novitates Zoologicae 12: 191 (175-191). Sudan: “Between Dalba to the Uma River, Konta”.

plagifera Le Cerf, 1924 (as f. of Papilio leonidas leonidas). Bulletin du Muséum National d’Histoire Naturelle. Paris 30: 138 (137-139). “Port Lakkah”.

melusina Le Cerf, 1924 (as f. of Papilio leonidas brasidas). Bulletin of the Hill Museum, Witley 1: 397 (369-399, 576, 578, 580, 582, 584). South Africa: “Malvern, Natal”.

petiveranoides Bryk, 1928 (as f. of Papilio leonidas). Societas Entomologica 43: 15 (13-16). Uganda: “Mount Elgon-Steppe; Busia”.

obliterata Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) leonidas leonidas). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo: “Congo Belge, Luebo”.

subobliterata Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) leonidas leonidas). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo?: “Camayenne”.

vreuricki Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) leonidas leonidas ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo: “Congo Belge”.

mathieui Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) leonidas leonidas ). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo?: “Sankisishia Congo”.

vrydaghi Berger, 1950 (as ab. of Graphium leonidas leonidas). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 91 (102 pp.). Democratic Republic of Congo: “Stanleyville”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

leucosina Berger, 1950 (as ab. of Graphium leonidas leonidas). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 91 (102 pp.). Democratic Republic of Congo: “Lac Leopold II, Lukombe”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

djema Berger, 1950 (as ab. of Graphium leonidas leonidas). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 91 (102 pp.). Central African Republic: “Afrique Equat. Franç., Chari, Djema”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

cyrnoides Storace, 1953 (as ab. of Graphium leonidas). Annali del Museo Civico di Storia Naturale (di Genova) Giacomo Doria 66: 260 (252-264). Zambia/Zimbabwe: “Zambesi, Cascate Victoria”.

Graphium (Arisbe) leonidas pelopidas (Oberthür, 1879)

Papilio leonidas var? pelopidas Oberthür, 1879. Études d’Entomologie 4: 55 (1-117).

Type locality: Tanzania: “Zanguebar, Tchouacka”. Tchouacka is probably Chwaka today and is on the island of Pemba (Congdon et al., 2009). Diagnosis: Differs from the nominate subspecies in the following respects: forewing upperside with much larger white spots in the cell and in spaces 3 and 4; hindwing upperside with more extensive median white patch and submarginal spots reduced to mere points (Kielland, 1990d). Distribution: Tanzania (Pemba Island, Zanzibar Archipelago). Specific localities: Tanzania – Tchouacka (= Chwaka), Pemba Island, Zanzibar Archipelago (TL).

Note: Congdon et al. (2009) suggest that pelopidas may deserve specific rather than subspecific rank.

Graphium (Arisbe) leonidas zanzibaricus Kielland, 1990

Graphium leonidas zanzibaricus Kielland, 1990. Butterflies of Tanzania 47 (363 pp.). Melbourne. Synonym of Graphiium leonidas pelopidas Oberthür, 1879. Ackery et al., 1995. Graphium leonidas zanzibaricus Kielland, 1990. Congdon et al., 2009. stat. rev.

Type locality: Tanzania: “Zanzibar”. [Actually Unguja Island (Congdon et al., 2009)]. Diagnosis: The individuals in the population on Unguja Island, the larger island in the Zanzibar Archipelago, are smaller and brighter than mainland individuals (Congdon et al., 2009). Distribution: Tanzania (Unguja Island, Zanzibar Archipelago). Specific localities: Tanzania – [Unguja Island], Zanzibar [Archipelago] (TL).

Graphium (Arisbe) leonidas santamarthae (Joicey & Talbot, 1927)

Papilio leonidas santa-marthae Joicey & Talbot, 1927. Encyclopédie Entomologique (B.3. Lepidoptera) 2: 12 (1-14).

Type locality: Sao Tome & Principe: “St. Principe, 1500-2000 ft.”. Distribution: Sao Tome & Principe (Island of Principe).

Graphium (Arisbe) leonidas thomasius (Le Cerf, 1924)

Papilio leonidas thomasius Le Cerf, 1924. Bulletin du Muséum National d’Histoire Naturelle. Paris 30: 137 (137-139).

Type locality: Sao Tome & Principe: “San Thome”. Distribution: Sao Tome & Principe (Island of Sao Tome).

*Graphium (Arisbe) levassori (Oberthür, 1886) Comore Swordtail

Papilio levassori Oberthür, 1886. Études d’Entomologie 13: 10 (9-15). Papilio Levassori Oberthür, 1890. Aurivillius, 1899. [date of authorship erroneous]

Type locality: Comoro Islands: “Grand Comore”. Distribution: Comoro Islands. Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) cyrnus (Boisduval, 1836)

Papilio cyrnus Boisduval, 1836. In: [Roret, Suites à Buffon] Histoire naturelle des Insectes. Species général des Lépidopteres 1: 239 (690 pp.). Paris. Papilio cyrnus Boisd., 1836. Aurivillius, 1899.

Type locality: Madagascar. Distribution: Madagascar. Specific localities: Madagascar – Mojange (Suffert, 1904). Habitat: Forest (Lees et al., 2003). Early stages: Nothing published. Larval food: Nothing published.

nuscyrus Suffert, 1904 (as ssp. of Papilio cyrnus). Deutsche Entomologische Zeitschrift, Iris 17: 104 (12-107). Madagascar: “Mojange”. Synonymized with Graphium cyrnus by Lees et al., 2003.

similis Storace, 1953 (as f. of Graphium cyrnus). Annali del Museo Civico di Storia Naturale (di Genova) Giacomo Doria 66: 260 (252-264). Madagascar.

adiecta Storace, 1953 (as f. of Graphium cyrnus). Annali del Museo Civico di Storia Naturale (di Genova) Giacomo Doria 66: 260 (252-264). Madagascar.

The tynderaeus clade

*Graphium (Arisbe) tynderaeus (Fabricius, 1793) Electric Green Swordtail

Papilio tynderaeus Fabricius, 1793. Entomologia Systematica emendata et aucta 3 (1): 35 (488 pp.). Papilio tynderaeus Fabr., 1793. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside). Type locality: No locality given. Probably Sierra Leone (Larsen, 2005a). Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo (Ituri Forest and north Shaba), Tanzania (west). Specific localities: Ghana – Kakum National Park (Larsen, 2006d). Cameroon – Edea (Gaede, 1916); Korup (Larsen, 2005a). Gabon – Libreville (van de Weghe, 2010); Crystal Mountains (van de Weghe, 2010); Oyem (van de Weghe, 2010); Bitam (van de Weghe, 2010); Lope (van de Weghe, 2010); Langoue (van de Weghe, 2010); Ipassa (van de Weghe, 2010); Kongou (van de Weghe, 2010); Lamberene (van de Weghe, 2010); Ogooue (van de Weghe, 2010); Waka (van de Weghe, 2010); Fernan Vaz (van de Weghe, 2010); Rabi (van de Weghe, 2010); Gamba (van de Weghe, 2010); Mouila (van de Weghe, 2010); Tchibanga (van de Weghe, 2010); Lastoursville (van de Weghe, 2010); Bakouaka (van de Weghe, 2010); Nouna (van de Weghe, 2010); Franceville (van de Weghe, 2010); Bateke Plateau (van de Weghe, 2010). Central African Republic – Dzanga (Noss, 1998). Democratic Republic of Congo – Lopori Beach (Capronnier, 1889); Djamba (Dufrane, 1936) Sankuru Berger, 1950); Kondue (Berger, 1950). Tanzania – near Mkulya River, Kigoma District at 1 450 m (Kielland; single record in 1954). Habitat: Primary lowland forest. Habits: An uncommon and very beautiful butterfly (Larsen, 2005a). Schultze (1917) regards it to be the papilionid with the fastest flight. It hurtles down tracks and along rivers at a furious rate (Larsen, 2005a). Males sometimes mud-puddle but are extremely wary if approached (Larsen, 2005a). Larsen (2005a) recounts an instance of a specimen perching on half a boiled egg laid out as bait. Early stages: Nothing published. Larval food: Nothing published.

nausinous Godart, 1819 in Latreille & Godart, [1819], [1824] (as sp. of Papilio). Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 45 (1-328 [1819], 329-828 [1824]). Paris. “Afrique”.

ochrea Capronnier, 1889 (as var. of Papilio tynderaeus). Bulletin de la Société Entomologique de Belgique 1889: cxix (118-127). Democratic Republic of Congo: “Lopori Strand”.

fraudatus Gaede, 1916 (as ab. of Papilio tynderaeus). Internationale Entomologische Zeitschrift 9: 105 (105-106, 109-112, 125-126). Cameroon: “Süd Kamerun, Edea”.

incompleta Dufrane, 1936 (as ab. of Papilio tynderaeus). Lambillionea 36: 42 (40-42). Democratic Republic of Congo: “Congo Belge, Djamba”.

confluens Berger, 1950 (as ab. of Graphium tynderaeus). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 67 (102 pp.). Democratic Republic of Congo: “Sankuru, Kondue”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

*Graphium (Arisbe) latreillianus (Godart, [1819]) 50

Coppery Swordtail

Papilio latreillianus Godart, 1819 in Latreille & Godart, [1819], [1824]. Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 44 (1-328 [1819], 329-828 [1824]). Paris. Papilio latreillianus God., 1819. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside and underside). Type locality: “Afrique”. Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Sao Tome & Principe, Equatorial Guinea, Gabon, Congo, Angola, Central African Republic, Democratic Republic of Congo, Chad, Uganda, Tanzania. Habitat: Primary forest (Larsen, 2005a). Habits: May be quite common at certain times, such as at the begining of the rainy season (March in Ghana) (Larsen, 2005a). Males mud-puddle and are also attracted to urine-soaked sand, human pespiration and camp rubbish tips (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) latreillianus latreillianus (Godart, [1819])

Papilio latreillianus Godart, 1819 in Latreille & Godart, [1819], [1824]. Encyclopédie Méthodique. Histoire Naturelle [Zoologie] 9 Entomologie: 44 (1-328 [1819], 329-828 [1824]). Paris.

Published images: Larsen, 2005a (male upperside and underside). Type locality: “Afrique”. Distribution: Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana. Specific localities: Guinea – Fouta Djalon (Larsen, 2005a); Conakry (Larsen, 2005a). Ghana – Ankasa (Larsen, 2005a); Kakum National Park (Larsen, 2006d).

Graphium (Arisbe) latreillianus theorini (Aurivillius, 1881)

Papilio latreillanus [sic] var. theorini Aurivillius, 1881. Entomologisk Tidskrift 2: 45 (38-47). Type locality: Gabon: “Gaboon”. Holotype in the Swedish Natural History Museum (images available at www2.nrm.se/en/lep_nrm/l). Diagnosis: Differs from the nominate subspecies in that the green markings on the hindwing and in spaces 1a and 1b of the forewing, are narrower, and it is a larger insect (Larsen, 2005a). Distribution: Nigeria, Cameroon, Sao Tome & Principe, Equatorial Guinea, Gabon, Congo, Democratic Republic of Congo, Angola, Central African Republic, Chad, Uganda (west), Tanzania. Specific localities: Cameroon – Duala (Strand, 1914); Lolodorf (Avinoff, 1926); Nyong River, south of Makak (Birket-Smith, 1960); Korup (Larsen, 2005a). Gabon – Throughout (van de Weghe, 2010). Democratic Republic of Congo – Tsibobe, Katanga (Berger, 1950). Central African Republic – Dzanga (Noss, 1998).

potamonianus Ehrmann, 1909 (as sp. of Papilio). Canadian Entomologist 41: 86 (85-87). Democratic Republic of Congo: “Upper Congo, West Africa”.

infrapicta Strand, 1914 (as ab. of Papilio latreillianus theorini ). Archiv für Naturgeschichte 80 (A.2.): 159 (139- 161). Cameroon: “Duala”.

aurivillianus Avinoff, 1926 (as sp. of Papilio). Annals of the Carnegie Museum 16: 355 (355-374). Cameroon: “Lolodorf”.

xerophila Berger, 1950 (as f. of Graphium tynderaeus theorini ). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 69 (102 pp.). Democratic Republic of Congo: “Katanga, Tsibobe”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

lutea Birket-Smith, 1960 (as ab. of Papilio latreillanus [sic] theorini). Bulletin de l’Institut Français d’Afrique Noire (A) 22: 533 (521-554, 924-983, 1259-1284). Cameroon: “French Cameroons, River Nyong, south of Makak station”.

An unplaced species In the phylogenetic analyses of Smith & Vane-Wright (2001) G. philonoe emerged as part of a polytomous terminal bush with the adamastor group, despite the distinctive appearance and genitalia.

*Graphium (Arisbe) philonoe (Ward, 1873) Eastern White-lady Swordtail

Papilio philonoë Ward, 1873. Entomologist’s Monthly Magazine 10: 152 (59-60, 151-152). Papilio philonoe Ward, 1873. Aurivillius, 1899.

Alternative common names: Eastern White Lady; White-dappled Swallowtail. Type locality: [Kenya]: “Ribé”. Diagnosis: Similar to G. angolanus and G. taboranus, from which it differs in the cell spots, which consist of an upper, wide and nearly even white band with three white dots below it (Kielland, 1990d). Distribution: Sudan, Ethiopia, Somalia, Uganda, Kenya, Democratic Republic of Congo, Tanzania, Malawi, Mozambique. Habitat: Coastal and riparian forest. Prefers the forest-savanna ecotone (Larsen, 1991c). Habits: The flight is not as powerful as most of the other species of the genus Graphium. Both sexes are fond of flowers; Larsen (1991) records large numbers feeding from the flowers of a species of Maerua in Brachystegia- dominated portions of the Arabuko-Sokoke Forest on the Kenyan coast. It is only infrequently seen mudpuddling. Early stages:

Congdon et al., 2009 [photographs of larva and pupa of both subspecies]. Ovipositing females insert the egg into a tightly-furled leaf bud, making them difficult to find even if oviposition is observed.

Larval food: Annona species (Annonaceae) [Larsen, 1991c: 117]. Uvaria acuminata Oliv. (Annonaceae) [Congdon et al., 2009; subspecies philonoe]. Uvaria chamae P.Beauv. (Annonaceae) [Van Someren, 1974: 316]. Uvaria leptocladen Oliv. (Annonaceae) [Van Someren, 1974: 316; Congdon et al., 2009 (for subspecies whalleyi)].

Graphium (Arisbe) philonoe philonoe (Ward, 1873)

Papilio philonoë Ward, 1873. Entomologist’s Monthly Magazine 10: 152 (59-60, 151-152). Papilio philonoe Ward, 1873. Aurivillius, 1899.

Type locality: Kenya: “Ribé”. 52

Distribution: Democratic Republic of Congo, Somalia (coast), Kenya (coast), Tanzania (east), Malawi, Mozambique. Specific localities: Kenya – Ribe (TL); Arabuko-Sokoke Forest (Stoneham, 1931); Mount Sagala (Larsen, 1991c). Tanzania – Amani in the Usambaras to Kimboza (Kielland, 1990d); Uluguru (Kielland, 1990d); Turiani (Kielland, 1990d); Sanje at the foot of the Uzungwa scarp (Kielland, 1990d); Pugu Hills (Kielland, 1990d); Rondo Plateau (Kielland, 1990d); Semdoe Forest Reserve (Doggart et al., 2001).

dulcinea Stoneham, 1931 (as f. of Papilio philonoë). Bulletin of the Stoneham Museum (2): [1] ([1 p.]). Kenya: “Protectorate of Kenya, Gede, Arubuka Forest”.

Graphium (Arisbe) philonoe whalleyi (Talbot, 1929)

Papilio philonoe whalleyi Talbot, 1929. Bulletin of the Hill Museum, Witley 3: 72 (72-77).

Type locality: Sudan: “S. Sudan, Imatong Mountains”. Distribution: Sudan (south), Ethiopia (south-west), Uganda (north), Kenya (extreme north-west). Specific localities: Sudan – Imatong Mountains (TL).

Note: Larsen (1991) suspects that whalleyi may be a species distinct from philonoe. This is supported by Congdon et al. (2009) who state that whalleyi is more closely tied to forest than is the nominate subspecies.

The adamastor group A group of 17 species which appear to mimic members of the danaine genus Amauris.

*Graphium (Arisbe) adamastor (Boisduval, 1836) Boisduval’s White Lady Swordtail

Papilio adamastor Boisduval, 1836. In: [Roret, Suites à Buffon] Histoire naturelle des Insectes. Species général des Lépidopteres 1: 371 (690 pp.). Paris. Papilio adamastor Boisd., 1836. Aurivillius, 1899.

Alternative common name: Boisduval’s White Lady. Published images: Larsen, 2005a (male upperside and female upperside). Type locality: “Cote de Guinée”. [= Coast of Guinea] Diagnosis: See G. almansor for differentiation from that species. It differs from G. agamedes as follows: clear white subapical markings on forewing (absent in agamedes); a much larger triangular white spot in the cell (Larsen, 2005a). Distribution: Guinea, Sierra Leone, Mali, Burkina Faso, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Congo, Central African Republic, Democratic Republic of Congo (Ubangi, Mongala). Specific localities: Guinea – Mamou (Dufrane, 1946); 30 km N of Mamou (Dufrane, 1946). Ghana – Boabeng-Fiema (Larsen, 2005a); Aburi (Larsen, 2005a). Togo – Klouto [6°57'15.07"N 0°34'54.40"E] (Safian et al., 2009). Cameroon – Akom, S. of Ebolowa (Libert, 2007); Yoko on the Adamaoua Plateau (Libert, 2007); Nkolkomou, a hill near Yaounde (Libert, 2007). Democratic Republic of Congo – Ubangi (Berger, 1950); Zongo-Mokoanghay (Berger, 1950). Habitat: Dry forest and the transition between forest and Guinea savanna (Larsen, 2005a). Habits: The flight is slow, especially females, which are good mimics of Amauris damocles. Like G. almansor it 53 has periodic population irruptions (Larsen, 2005a). Both sexes feed from flowers and Larsen (2005a) noted their partiality for those of Calodendrum splendens. They have also been noted ‘tree-topping’ in Aburi Botanical Gardens, Ghana but it is not known whether this behaviour is usual for the species (Larsen, 2005a). Early stages:

Larsen, 2005a When ovipositing, the female selects an unfurled shoot and forces it open with her abdomen. The egg that is laid is then covered by the leaves when the female removes her abdomen.

Larval food: Annonaceae [Larsen, 2005a; Boabeng-Fiema; oviposition only].

guineensis Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) charcedonius [sic]). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 120 (101-122). Guinea: “Vallée de Mamou”.

houzeaui Dufrane, 1946 (as ab. of Pailio (Cosmodesmus) charcedonius [sic]). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 120 (101-122). Guinea: “kil. 30 au N. de Mamou, route vers Labé”.

dimbroko Berger, 1950 (as ssp. of Graphium adamastor). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 89 (102 pp.). Sierra Leone. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. [Synonymized with G. adamastor by Berger (1974)].

zongo Berger, 1950 (as ssp. of Graphium adamastor). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 87 (102 pp.). Democratic Republic of Congo: “Congo- Ubangi, Zongo-Mokoanghay”. Paratype in the Royal Museum for Central Africa, Tervuren, Belgium. [Synonymized with G. adamastor by Berger (1974)].

*Graphium (Arisbe) agamedes (Westwood, 1842) Westwood’s White Lady Swordtail

Papilio agamedes Westwood, 1842. Annals and Magazine of Natural History 9: 38 (36-39). Papilio agamedes Westw., 1842. Aurivillius, 1899.

Alternative common name: Westwood’s White Lady. Published images: Larsen, 2005a (male upperside and underside). Type locality: [Ghana]: “Ashantee, tropical Africa”. Diagnosis: Similar to G. adamastor but white markings more regular; light subapical markings absent; triangular cell spot smaller and often divided in two (Larsen, 2005a). Distribution: Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon, Central African Republic, Democratic Republic of Congo (Mongala, Uele). Recorded, in error, from Sierra Leone by Strand, 1909 (Larsen, 2005a). Specific localities: Ivory Coast – Mount Tonkoui (Larsen, 2005a). Ghana – Ashanti (TL); Volta Region (Maessen, vide Larsen, 2005a). Habitat: Forest/savanna transition zone (Larsen, 2005a). Habits: This is a rare butterfly (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

medesaga Suffert, 1904 (as ssp. of Papilio agamedes). Deutsche Entomologische Zeitschrift, Iris 17: 106 (12-107). Togo: “Hinterland”.

*Graphium (Arisbe) schubotzi (Schultze, 1913)

Papilio [odin] var. schubotzi Schultze, 1913. Entomologische Rundschau 30: 73 (73-74). Graphium auriger schubotzi (Schultze, 1913). Graphium schubotzi (Schultze, 1913). Smith & Vane-Wright, 2001: 622 stat. rev.

Type locality: Democratic Republic of Congo: “Duma, Ubangi-Distrikt; Angu, Ubangi-Distrikt”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium (taxon name spelt as schoubotzi). Distribution: Cameroon, Congo, Central African Republic, Democratic Republic of Congo. Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) schubotzi schubotzi (Schultze, 1913)

Papilio [odin] var. schubotzi Schultze, 1913. Entomologische Rundschau 30: 73 (73-74). Graphium auriger schubotzi (Schultze, 1913). Ackery et al., 1995: 162. Graphium schubotzi (Schultze, 1913). Smith & Vane-Wright, 2001: 622 stat. rev.

Type locality: Democratic Republic of Congo: “Duma, Ubangi-Distrikt; Angu, Ubangi-Distrikt”. Distribution: Cameroon (east), Congo, Central African Republic, Democratic Republic of Congo (Ubangi). Specific localities: Cameroon – Duma (TL); Angu (Schultze, 1913); Düm-Mündung (Strand, 1910); Batouri district (Smith & Vane- Wright, 2001). Democratic Republic of Congo – Lemfu (Berger, 1950).

odin Strand, 1910 (as sp. of Papilio). Wiener Entomologische Zeitung 29: 34 (29-35). Cameroon: “Kamerun, Düm- Mündung”. [Invalid; junior primary homonym of Papilio odin Fabricius, 1793 [Riodinidae].]

eyeni Berger, 1950 (as ssp. of Graphium odin). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 84 (102 pp.). Democratic Republic of Congo: “Bas Congo, Lemfu”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. [Given as a subspecies of auriger (Butler) in Ackery et al., 1995: 162. Regarded as a synonym of aurivilliusi (Seeldrayers) by Hancock, 1993 (Arnoldia Zimbabwe 9 (40): 568 (557-583.)). Synonymized with Graphium schubotzi (Schultze, 1913) by Smith & Vane-Wright, 2001 (Bulletin of the British Museum of Natural History Entomology Series 70 (2): 623 (503-719))].

Graphium (Arisbe) schubotzi maculata Libert, 2007

Graphium schubotzi maculata Libert, 2007. Lambillionea 107 (1): 2 (1-11).

Type locality: Cameroon: “Nkolbisson, IV 1988 (M. Libert); Holotype (male) in M.N.H.N., Paris”. Description: “The distinctive character is the presence of marginal marks. On forewing, the visibility of these marks is very variable: they are hardly visible in some specimens, but they are present in all cells from R 4 to CuA2 in others; they are always paler than discal marks, and almost always separated in two by intervenosa. In cell CuA 1, the discal mark tends to be smaller than in the nominate subspecies for males (it is even very small for two of them), and it is missing for the three females (it is present, with a normal size, in all the nine females from Bangui examined). On hindwing, the marginal (and post-discal) marks have the same white color as discal marks. Marginal marks are present in all cells from R5 to CuA1; in each cell, the mark is divided in two small spots either side of the midline, the lower one being often less marked. Post-discal marks occur in cells M2 (small), M3 (large) and CuA1 (medium), the latter two slightly notched in the middle of their distal margin. The disposition of the post-discal marks is the same as in G. (A.) adamastor, but their shape is quite different: in this species, they are deeply notched, almost separated in two lines, like a U.” Distribution: Cameroon (central). Specific localities: Cameroon – Nkolbisson, near Yaounde (TL); Mts Akondoue, Febe, Kala, Ngoekele, Nkolkomou (Libert, 2007); 60 km NW of Afanessele (Libert, 2007); 50 km S of Zoetoupsi (Libert, 2007); Dja Forest (Libert, 2007); 55

Makouopsap near Foumban (Libert, 2007); Lena on the Admaoua Plateau (Libert, 2007).

*Graphium (Arisbe) olbrechtsi Berger, 1950

Graphium olbrechtsi Berger, 1950. Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 85 (102 pp.).

Type locality: Democratic Republic of Congo: “Sankuru, Mwene-Ditu”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Democratic Republic of Congo. Early stages: Nothing published. Larval food: Nothing published. Relevant literature: Berger, 1969 – Notes on.

Graphium (Arisbe) olbrechtsi olbrechtsi Berger, 1950

Graphium olbrechtsi Berger, 1950. Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 85 (102 pp.).

Type locality: Democratic Republic of Congo: “Sankuru, Mwene-Ditu”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Democratic Republic of Congo (Kabinda, Lomami, Lualaba). Specific localities: Democratic Republic of Congo – Sankuru, Mwene-Ditu (TL).

Graphium (Arisbe) olbrechtsi tongoni Berger, 1969

Graphium olbrechtsi tongoni Berger, 1969. Lambillionea 67: 84 (84-85).

Type locality: Democratic Republic of Congo: “Congo ex Belge, Maniema, Tongoni”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Democratic Republic of Congo (east). Specific localities: Democratic Republic of Congo – Maniema, Tongoni (TL).

*Graphium (Arisbe) abri Smith & Vane-Wright, 2001

Graphium (Arisbe) abri Smith & Vane-Wright, 2001. Bulletin of the Natural History Museum Entomology Series 70 (2): 628 (503-719).

Type locality: ? Distribution: Central African Republic. Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) almansor (Honrath, 1884) Almansor White Lady Swordtail

Papilio almansor Honrath, 1884. Berliner Entomologische Zeitschrift 28: 210 (203-212). Papilio almansor Honrath, 1884. Aurivillius, 1899.

Graphium almansor almansor. Male. Left – upperside; right – underside. Wingspan: 60mm. Kitwe, Zambia. 64.08.22. A.I. Curle. (Curle Trust Collection – 10).

Alternative common name: Honrath’s White Lady. Type locality: “Guinea, Ashanti”. [False locality.] Larsen (2005a) argues that the type locality is probably Angola. Diagnosis: G. almansor can be distinguished from the similar G. adamastor by the following: forewing dark brown not blackish grey; no, or very small, white postdiscal spot space 2 of forewing; hindwing cell entirely white (distal end brown in adamastor); the small submarginal spots on both wings less well developed (Larsen, 2005a). Distribution: Guinea, Ghana, Togo, Nigeria, Cameroon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo, Sudan, Ethiopia, Uganda, ?Rwanda, ?Burundi, Kenya, Tanzania, Zambia. Habitat: Forest, including riparian forest. Also in heavy Brachystegia woodland (Heath et al., 2002). In Tanzania the nominate subspecies occurs at altitudes from 780 to 1 500 m, but is commonest at lower altitudes (Kielland, 1990d). In West Africa it is found in hilly country at the transition between Guinea savanna and small patches of forest, between 700 and 1 100 m (Larsen, 2005a). In Cameroon it is found between 800 and 1 200 m, i.e. it is regarded as submontane (Libert, 2007). Habits: Both sexes are mimics of distasteful danaines belonging to the genus Amauris; in West Africa A. damocles and in East Africa the A. echeria-group (Larsen, 2005a). Males are greatly attracted to damp spots on river banks and both sexes feed from flowers. Flight period: In West Africa it is often common in late January (Larsen, 2005a). Early stages:

Congdon et al., in Libert, 2007 The pupa is similarly marked to that of Graphium poggianus but is much smaller.

Congdon et al., 2009 [photographs of larva and pupa]. Ovipositing females insert the egg into a tightly-furled leaf bud, making them difficult to find even if oviposition is observed. The pupae pupate head downwards on the underside of a leaf.

Larval food: Pseudospondias microcarpa (A.Rich.) Engl. (Anacardiaceae) [Van Someren, 1974: 316; this record is dubious (Larsen, 2005a). Congdon et al, in Libert, 2007: 10 state categorically that this record is incorrect.] A liana in the Annonaceae (Congdon et al., in Libert, 2007). Uvaria angolensis Oliv. (Annonaceae) [Congdon et al., 2009].

Graphium (Arisbe) almansor almansor (Honrath, 1884)

Papilio almansor Honrath, 1884. Berliner Entomologische Zeitschrift 28: 210 (203-212). Papilio almansor Honrath, 1884. Aurivillius, 1899.

Graphium almansor almansor. Male. Left – upperside; right – underside. Wingspan: 60mm. Kitwe, Zambia. 64.08.22. A.I. Curle. (Curle Trust Collection – 10).

Type locality: “Guinea, Ashanti”. [False locality.] Distribution: Angola (Bivar de Sousa & Fernandes, 1966), Democratic Republic of Congo (Lualaba, Lomami, Kasai, Sankuru, Tanganika), Zambia (north). Specific localities: Democratic Republic of Congo – Kapanga (Berger, 1950); Kinda (Berger, 1950). Zambia – 100 km south of Mwinilunga (Heath et al., 2002); Mwinilunga (Heath et al., 2002); Ikelenge (Heath et al., 2002); South Mutundu River (Heath et al., 2002); Ndola (Heath et al., 2002); upper Kafue River (Heath et al., 2002); Samfya (Heath et al., 2002); Nchelenge (Heath et al., 2002); Chambeshi River (Heath et al., 2002); Mbala (Heath et al., 2002); Kitwe (male illustrated above).

albescens Berger, 1950 (as f. of Graphium almansor almansor). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 81 (102 pp.). Democratic Republic of Congo: “Lualaba, Kapanga”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

depuncta Berger, 1950 (as f. of Graphium almansor almansor). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 81 (102 pp.). Democratic Republic of Congo: “Lualaba, Kinda”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

Graphium (Arisbe) almansor birbiri (Ungemach, 1932)

Papilio charcedonius [sic] birbiri Ungemach, 1932. Mémoires de la Société des Sciences Naturelles (et Physiques) du Maroc 32: 21 (1- 122).

Type locality: Ethiopia: “Baro”. Distribution: Ethiopia (south-west). Specific localities: Ethiopia – Baro (TL).

Graphium (Arisbe) almansor carchedonius (Karsch, 1895)

Papilio carchedonius Karsch, 1895. Entomologische Nachrichten. Berlin 21: 285 (275-286). Papilio carchedonius Karsch, 1895. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside). Type locality: Togo: “Bismarckburg, Misahöhe”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 633 (503-719))]. Diagnosis: Unlike the nominate subspecies and subspecies escherichi the large white spot in forewing cell is rounded and does not touch the large spot in space 3 broadly. Distribution: Ghana, Togo. Specific localities: Ghana – Kyabobo National Park (Larsen, 2005a). Togo – Bismarckburg (TL); Klouto [6°57'15.07"N 0°34'54.40"E] (Safian et al., 2009).

Graphium (Arisbe) almansor escherichi (Gaede, 1915)

Papilio escherichi Gaede, 1915. Internationale Entomologische Zeitschrift 9: 71 (38-40, 71-74).

Type locality: [Central African Republic] (Libert, 2007): “Neu-Kamerun, Carnot”. The holotype is missing but the three other males in the type series are in the Berlin Museum (Libert, 2007). Libert (2007: 4) designates one of these males, captured by Escherich during the Muni Expedition, as the lectotype (“Kamerun-Carnot, C.A.R. (S.G. Escherich); Z.M.H.U., Berlin”). Distribution: Nigeria (east), Cameroon, Congo, Central African Republic, Democratic Republic of Congo (Mongala, Uele), Sudan (south), Uganda. Specific localities: Nigeria – Boggo, Obudu Plateau (C. Olory, vide Larsen, 2005a); Mambilla Plateau (Larsen, 2005a); near Yola (Larsen, 2005a). Cameroon – Toro in N. (Libert, 2007); Akoafim (Tessmann, vide Libert, 2007; suspected to be erroneous); Msungli near Bamenda (Libert, 2007); Yoko on the Adamaoua Plateau (Libert, 2007); Kounden Plateau (Libert, 2007); Malap Forest (Libert, 2007); Mt Mbam (Libert, 2007); Kimbi (Libert, 2007); Makouopsap (Libert, 2007); Banyo (Libert, 2007); Gaoui (Libert, 2007); Pangar (Libert, 2007); Ngaoundaba (Libert, 2007); Wakaso (Libert, 2007). Central African Republic – Carnot (TL).

snowi Gabriel, 1945 (as ssp. of Graphium almansor). Entomologist 78: 151 (151-152). Sudan: “S. Sudan”.

Graphium (Arisbe) almansor uganda (Lathy, 1906)

Papilio uganda Lathy, 1906. Transactions of the Entomological Society of London 1906: 9 (1-10).

Type locality: Uganda: “Entebbe”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 636 (503-719))]. Diagnosis: Light markings on the upperside of the wings reduced in extent and yellowish rather than white (Kielland, 1990d). Distribution: Democratic Republic of Congo (Kivu), Uganda, Kenya (west), Tanzania (extreme north-west), Rwanda?, Burundi? Specific localities: Uganda – Entebbe (TL); Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Kenya – Kakamega Forest (Larsen, 1991c); Mara River (Larsen, 1991c); Nyanza (Larsen, 1991c). 59

Tanzania – western and north-western Tanzania, from Sibwesa in Mpanda District to Kigoma (Kielland, 1990d); Bukoba Region (Kielland, 1990d).

graueri Grünberg, 1908 (as sp. of Papilio). Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1908: 59 (50-62). Uganda.

Graphium (Arisbe) almansor dufranei Collins & Larsen, 2008

Graphium almansor dufranei Collins & Larsen, 2008. Metamorphosis 19 (2): 44 (42-113).

Published images: Larsen, 2005a (male upperside and underside). Type locality: Guinea: “Fouta Djalon, Labé (11°32'N 12°28'W), ii.2001 (ABRI leg. et coll.)” Diagnosis: “Male forewing 40 mm. In the male the white cell spot Etymology: Named for Abel Dufrane, who described a large number of taxa between 1929 and 1954 (Collins & Larsen, 2008). Distribution: Guinea. Specific localities: Guinea – Fouta Djalon (TL); Dubreka (Collins & Larsen, 2008).

*Graphium (Arisbe) auriger (Butler, 1876)

Papilio auriger Butler, 1876. Entomologist’s Monthly Magazine 13: 57 (56-57). Papilio auriger Butler, 1876. Aurivillius, 1899.

Type locality: Gabon: “Gaboon”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 638 (503-719))]. Distribution: Senegal, Cameroon, Equatorial Guinea, Gabon, Congo, Central African Republic, Democratic Republic of Congo. Specific localities: Cameroon – Kribi (Libert, 2007). Gabon – Bitam (van de Weghe, 2010); Oyem (van de Weghe, 2010); Mitzic (van de Weghe, 2010); Kinguele (van de Weghe, 2010); Tchimbele (van de Weghe, 2010); Kougouleu (van de Weghe, 2010); Kitomba (van de Weghe, 2010); Waka (van de Weghe, 2010); Fernan Vaz (van de Weghe, 2010); Abanga River (van de Weghe, 2010); Ogooue (van de Weghe, 2010); Koulamoutou (van de Weghe, 2010); Tchibanga (van de Weghe, 2010). Early stages: Nothing published. Larval food: Nothing published.

harpagon Grose-Smith, 1890 (as sp. of Papilio). Annals and Magazine of Natural History (6) 5: 224 (224). Gabon: “Gaboon”.

*Graphium (Arisbe) fulleri (Grose-Smith, 1883)

Papilio fulleri Grose-Smith, 1883. Entomologist’s Monthly Magazine 19: 234 (234-235). Papilio Fulleri Smith, 1883. Aurivillius, 1899.

Type locality: Cameroon: “Camaroons”. Distribution: Cameroon, Equatorial Guinea, Gabon, Congo, Angola, Democratic Republic of Congo, Central African Republic. Early stages: Nothing published. 60

Larval food: Nothing published.

Graphium (Arisbe) fulleri fulleri (Grose-Smith, 1883)

Papilio fulleri Grose-Smith, 1883. Entomologist’s Monthly Magazine 19: 234 (234-235). Papilio Fulleri Smith, 1883. Aurivillius, 1899.

Type locality: Cameroon: “Camaroons”. Distribution: Cameroon (north-west), Congo. Specific localities: Cameroon – Baliko (Libert, 2007); Loum (Libert, 2007); Mt Cameroon (Libert, 2007); Bafut, north of Bamenda (Libert, 2007); Dohon (Libert, 2007); Tanyon (Libert, 2007). Congo – San Benito (Le Cerf, 1912).

transiens Le Cerf, 1912 (as var. of Papilio boulleti). Bulletin de la Société Entomologique de France 1912: 383 (382-383). Congo: “Congo français, rivière San Benito”.

Graphium (Arisbe) fulleri boulleti (Le Cerf, 1912)

Papilio boulleti Le Cerf, 1912. Bulletin de la Société Entomologique de France 1912: 382 (382-383).

Type locality: [Gabon]: “Congo français, chutes de Doumé (Ogooué)”. Distribution: Cameroon (central and south), Equatorial Guinea, Gabon, Congo (north), Democratic Republic of Congo, Central African Republic. Specific localities: Cameroon – Nzimulana, Ngoko Sanga (Strand, 1913); Moloundou (Strand, 1913); Lolodorf (Holland, 1917); Assobam (Berger, 1950); Mount Kala (Darge, 1995); Ngat II (Darge, 1995); Meukonong (Darge, 1995); Ebodenkou (Darge, 1995); Ekelemba (Libert, 2007); Afanessele (Libert, 2007). Gabon – Doumé (TL); Nouna (van de Weghe, 2010). Democratic Republic of Congo – Yakoma (Libert, 2007). Central African Republic – Bangui (Libert, 2007).

foersterius Strand, 1913 (as sp. of Papilio). Archiv für Naturgeschichte 79 (A.3.): 17 (17-18). Cameroon: “S. Kamerun, Nzimulana, Ngoko Sanga, 400-500 m.”.

stetteni Strand, 1913 (as sp. of Papilio). Archiv für Naturgeschichte 79 (A.7.): 140 (138-151). Cameroon: “Süd- Kamerun Molundu”.

sanganus Strand, 1913 (as sp. of Papilio). Archiv für Naturgeschichte 79 (A.3.): 17 (17-18). Cameroon: “Kamerun”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 642 (503-719))].

sanganoides Strand, 1913 (as sp. of Papilio). Archiv für Naturgeschichte 79 (A.3.): 18 (17-18). Cameroon: “Kamerun”.

divisimacula Strand, 1913 (as ab. of Papilio sanganoides). Archiv für Naturgeschichte 79 (A.3.): 18 (17-18). Cameroon: “Kamerun”.

weberi Holland, 1917 (as sp. of Papilio). Annales of the Carnegie Museum 11: 14 (14-18). Cameroon: “in the vicinity of Lolodorf”.

gabrieli Berger, 1950 (as f. of Graphium ucalegonides ucalegonides ). Annales du Musée Royal du Congo Belge. C- 61

Zoologie (3) 8 (1): 78 (102 pp.). Cameroon: “S. Cameroun, Assobam”.

beloni Darge, 1995 (as ssp. of Graphium ucalegonides). Lambillionea 95 (2) (Tome II): 284 (281-288). Cameroon: “Cameroun: Mont Kala, 900/1150m., 20 km. W. de Yaoundé, 15.III.1975 (Ph. Darge).” Place of deposition of holotype not stated. [Synonymized with G. fulleri boulleti (Le Cerf, 1912) by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomological Series 70 (2): 643 (503-719))].

*Graphium (Arisbe) ucalegonides (Staudinger, [1884])

Papilio ucalegonides Staudinger, 1884 in Staudinger & Schatz, 1884-8. Exotischer Schmetterlinge 1: 10 (333 pp.). Bayern. Papilio ucalegonides Staud., 1884. Aurivillius, 1899. Papilio phrynon Druce, 1895. Aurivillius, 1899. [synonym of ucalegonides] Graphium fulleri ucalegonides (Staudinger, 1884). Smith & Vane-Wright, 2001. Graphium ucalegonides (Staudinger, 1884). Larsen, 2005a. stat. rev.

Type locality: [Angola, Quango] (Larsen, 2005a): “Innern Ost afrikas”. Distribution: Cameroon, Congo, Angola (north), Central African Republic, Democratic Republic of Congo (except Shaba), Chad. Specific localities: Cameroon – Moloundou (Smith & Vane-Wright, 2001); Assamba [probably on the road between Moloundou and Yokaduma – Libert, 2007 (Smith & Vane-Wright, 2001); M’peum (Libert, 2007); Yokaduma (Libert, 2007). Angola – Quango (TL). Central African Republic – Dzanga (Noss, 1998; as fulleri ucalegonides). Democratic Republic of Congo – Punia (Dufrane, 1946); Sankuru, km. 50 de la route Lousambo-Batempa (Berger, 1950). Early stages: Nothing published. Larval food: Nothing published.

phrynon Druce, 1895 (as sp. of Papilio). Annals and Magazine of Natural History (6) 15: 332 (332-333). Democratic Republic of Congo: “Upper Congo”. Treated as a valid species by Aurivillius (1899).

cuvelieri Dufrane, 1946 (as ab. of Papilio (Cosmodesmus) ucalegonides). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 120 (101-122). Democratic Republic of Congo: “Punia Kivu”.

addenda Berger, 1950 (as f. of Graphium ucalegonides ucalegonides ). Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 78 (102 pp.). Democratic Republic of Congo: “Sankuru, km. 50 de la route Lousambo- Batempa”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

*Graphium (Arisbe) rileyi Berger, 1950 Riley’s Swordtail

Graphium ucalegonides rileyi Berger, 1950. Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 78 (102 pp.). Graphium fulleri rileyi Berger, 1950. Smith & Vane-Wright, 2001. Graphium rileyi Berger, 1950. Larsen, 2005a: 82, stat. rev.

Alternative common name: Riley’s Graphium. Published images: Larsen, 2005a (male upperside). Type locality: [Ghana]: “Gold Coast, West Africa, Begoro”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium. Distribution: Ivory Coast, Ghana (central). Specific localities: Ivory Coast – Abengourou area (Larsen, 2005a); Tiassale (Larsen, 2005a).

Ghana – Begoro (TL); Atewa Range (Maessen, vide Larsen, 2005a). Habitat: Semi-deciduous and upland forest (Larsen, 2005a). Habits: Apparently commonest at the beginning of the rainy season (February and March) (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) poggianus (Honrath, 1884)

Papilio poggianus Honrath, 1884. Berliner Entomologische Zeitschrift 28: 210 (203-212). Papilio poggianus Honrath, 1884. Aurivillius, 1899.

Graphium poggianus. Male. Left – upperside; right – underside. Wingspan: 77mm. Hillwood Nchila, Mwinilunga dist., Zambia. 6/X/2000. A.J. Gardiner. (Gardiner Collection).

Graphium poggianus. Female. Left – upperside; right – underside. Wingspan: 84mm. Female. Hillwood Nchila, Mwinilunga dist., Zambia. 6/X/2000. A.J. Gardiner. (Gardiner Collection).

Type locality: “Guinea”. [False locality.] Diagnosis: Differs from Graphium almansor in that it is larger; on the forewing upperside the white marking in the cell is triangular in shape (Kielland, 1990d). Distribution: Angola (north), Democratic Republic of Congo (south), Zambia (north-west). Specific localities: Zambia – Ikelenge (Heath et al., 2002); Zambezi Bridge (Libert, 2007); Hillwood (specimens illustrated above). 63

Habitat: Moist forest at about 1 450 m elevation (Congdon et al., in Libert, 2007). Habits: Males fly randomly through the forest and come to mud (Congdon et al., in Libert, 2007). The males are fast-flighted (Congdon et al., 2009). Females are scarcer and may be seen ovipositing at any height above the ground (the host plant is a liana, which grows into the canopy of the forest) (Congdon et al., 2009). Early stages:

Congdon, Bampton & Collins, in Libert, 2007 Females were seen ovipositing from ground level up into the canopy. The female inserts the egg into a furled leaf bud, so that it is invisible unless the bud is opened. The foodplant is a liana in the family Annonaceae. Artabotrys is present but is not used by poggianus. The larva feeds on the soft leaves at the growing point of a shoot. The final instar larva and the pupa are figured in the publication. The pupa has a pair of dark spots on the dorsal surface of the thorax. The long axis of the spots is head to tail. Occasionally there is a second pair of much smaller spots anterior to the large pair. Some pupae have a pair of thin dark lines running the length of the pupa and passing through the spots.

Congdon et al., 2009 [photographs of larva and pupa].

Larval food: A liana in the Annonaceae (Congdon et al., in Libert, 2007). Uvaria near angolensis Oliv. (Annonaceae) [Congdon et al., 2009; western Zambia].

deliae Libert & Collins, 2007 (as sp. of Graphium). Lambillionea 107 (1): 24 (19-29). Type locality: Zambia: “Chiwoma f., 100 km S. Mwinilunga, NW Zambia, 14 X 2002 (T.C. Congdon, S.C. Collins)”. Holotype (male) in ABRI, Nairobi. Described from 58 males and 2 females. Downgraded to a form by Gardiner, 2010b: 147 stat. nov. [Diagnosis: Male genitalia similar to those of G. poggianus, from which deliae differs in the greater extension of the white markings on the forewing (wider bar across the cell and a mark in CuA 1). The two butterflies [deliae and poggianus] differ not only in size but also in wing shape. In deliae both forewing and hindwing are narrower than in poggianus (Congdon et al., in Libert, 2007). Description: “Length of forewing: males, 38 to 41 mm, female, 43 mm. The new species is slightly smaller than poggianus. The ground colour of the upperside is dark brown, almost black on forewing; on hindwing, the dark region is restricted to the margin, but distinctly chocolate brown. Light markings are creamy, not pure white as in poggianus, while dark parts are lighter, which results in a less contrasted upperside. Opposite cells M 3 and M2, the forewing cell is crossed by a wide band (particularly wide in the holotype, but always wider than in poggianus), tapering distally in several males; two males have a faint apical spot in the cell. There is a small, but variable, spot in R 3, sometimes divided by intervenosa, and sometimes a very small one at the base of R4; the spot in R5 is much larger (always more than half the cell), and distinctly notched by intervenosa. In M1, along R5, the female has a narrow mark (about 6 mm), but only two males have a faint spot; more generally, apical markings have greater extension in the female. In M 2, the post-discal mark fills most of the posterior half; along the discocellular, its base is wider, extending in a line along M2 in the female. In M3, CuA1, CuA2 and 1A, the postdiscal marks fill most of the cells, with only a narrow dark margin (larger along nerves). The postdiscal mark in CuA 1 is the most variable; for half the males, it fills all the space between the nerves, but the dark margin extends more or less along CuA 1 in the others, and in two males CuA1 and CuA2 are completely separated. In poggianus, there is usually no mark in CuA1, but there are two notable exceptions, the type and one male from Ikelenge (see plate), in which there is a mark that is only slightly smaller than in the male of deliae with the smallest mark. On hindwings, clear markings are not significantly different from those in poggianus. The main difference is in the shade of the discal part of the wing (between the clear markings of the base and the dark margin), which is usually gold in males, with conspicuous intervenosa. The hindwings of the female are similar, but lighter, with the discal region more yellow. In cells M 2, M3 and CuA1, there is a wide streak on each side of intervenosa (these streaks are much fainter, hardly apparent, in males); in M1 there is only one streak, above intervenosa, and the streaks are not distinct from basal markings in R 5 and R1. As in poggianus, the underside is similar to the upperside, but paler; the basal tuft of white scales which is characteristic of the adamastor group is also present. The only difference is the presence of a black round spot at the base of the hindwing costal cell. Also on hindwing, the female and some males have also a much smaller dark spot at the base of R 1 (always missing in poggianus). Male genitalia. No difference could be found between one male of G. (A.) deliae and one male of poggianus; the horizontal blade of the ventral harp was not bifid. Both differ considerably from the genitalia of a male of G. (A.) a. almansor (from Chiwoma forest) by a much larger dorsal terminal process.” Distribution: Zambia (west). Specific localities: Zambia – Chiwoma Forest (TL); Mavunda Forest (Libert, 2007). Etymology: This species is named for Delia Curle, late wife of Alf Curle (Libert, 2007). Habitat: Dry Cryptosepalum exfoliatum forest on Kalahari sands at an altitude of 1 250 m elevation (Congdon et al., in Libert, 2007). Habits: Males appear to be territorial, patrolling along roads through the forest. The flight is fast and about

one to three metres above the ground. It is sympatric with Graphium almansor, both species using the same larval foodplant (Libert, 2007). The males of deliae, however, are much stronger and faster than those of almansor (Congdon et al., 2009). Early stages: Congdon et al., in Libert, 2007 The final instar larva is illustrated. It was found about a metre above the ground, feeding on the soft leaves at the tip of a shoot. The pupa is more heavily marked than that of poggianus or almansor. There are three pairs of spots, diminishing evenly in size towards the head. The largest pair is appreciably larger than those of poggianus, and the spots are of a different shape, being wider than they are long. The pupa is smaller than that of poggianus, reflecting the generally smaller size of the butterfly. This size difference appears to be unrelated to the food supply. “It is a consistent difference between the two taxa.” Congdon et al., 2009 [photographs of larva and pupa]. Larvae feed on the soft leaves at the tips of shoots. The pupae pupate head downwards on the underside of a leaf. Larval food: A liana species of Annonaceae (Congdon et al., in Libert, 2007). Uvaria angolensis Oliv. (Annonaceae) [Congdon et al., 2009; Chiwoma Forest].]

*Graphium (Arisbe) kigoma Carcasson, 1964

Graphium (Arisbe) almansor kigoma Carcasson, 1964. Journal of the East Africa Natural History Society & Coryndon Museum 24 (4): 66 (62-67). Graphium poggianus kigoma Carcasson, 1964. Graphium kigoma Carcasson, 1964. Smith & Vane-Wright, 2001: 652 stat. rev.

Type locality: [Tanzania]: “Tanganika, Makuyu, Kigoma”. Diagnosis: Differs from G. poggianus in the greenish-yellow suffusion in the pale areas, and on the hindwing upperside, which has a smaller pale discal area (Kielland, 1990d). Distribution: Tanzania (west – Kigoma district). Specific localities: Tanzania – Makuyu, Kigoma (TL); Gombe Stream National Park (Kielland, 1978); Mahale Mountains in Kigoma (Kielland, 1990d); Kampisa in Mpanda (Kielland, 1990d); Malagarasi River south of Kigoma town (Kielland, 1990d). Habitat: Forest, including riparian forest; also heavy woodland (Kielland, 1990d). Flight period: Commonest from December to April (the rainy season); rarely seen during the dry season (Kielland, 1990d). Early stages: Nothing published. Larval food: Nothing published.

wranghami Kielland, 1978 (as ssp. of Graphium almansor). Tijdschrift voor Entomologie 121: 160 (147-237). Tanzania: “Kigoma, Gombe Stream Nat. Park”. [Synonymized with G. kigoma Carcasson, 1964 by Smith & Vane- Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 652 (503-719))].

*Graphium (Arisbe) hachei (Dewitz, 1881)

Papilio hachei Dewitz, 1881. Berliner Entomologische Zeitschrift 25: 286 (286). Papilio Hachei Dewitz, 1881. Aurivillius, 1899.

Type locality: [Angola]: “West afrika, Quango”. [Lectotype designated by Smith & Vane-Wright, 2001 ( Bulletin of the Natural History Museum Entomology Series 70 (2): 655 (503-719))]. Distribution: Cameroon, Gabon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Central African Republic, Democratic Republic of Congo. Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) hachei hachei (Dewitz, 1881)

Papilio hachei Dewitz, 1881. Berliner Entomologische Zeitschrift 25: 286 (286). Papilio Hachei Dewitz, 1881. Aurivillius, 1899.

Type locality: [Angola]: “West afrika, Quango”. Distribution: Cameroon, Congo, Angola (Bivar de Sousa & Fernandes, 1966), Democratic Republic of Congo (west). Specific localities: Cameroon – Bitje, Ja River (Le Cerf, 1924). Angola – Quango (TL).

camerunicus Le Cerf, 1924 (as ssp. of Papilio hachei-möbii). Bulletin of the Hill Museum, Witley 1: 395 (369-399, 576, 578, 580, 582, 584). Cameroon: “Bitje, Ja River”.

Graphium (Arisbe) hachei moebii (Suffert, 1904)

Papilio moebii Suffert, 1904. Deutsche Entomologische Zeitschrift, Iris 17: 104 (12-107).

Type locality: [Democratic Republic of Congo]: “Central-Africa, 6°S. Breite, 25-26 ° östl.”. Distribution: Cameroon, Gabon, Congo, Democratic Republic of Congo (north and east), Central African Republic. Specific localities: Cameroon – Meukonong (Darge, 1995); Akoafim (Smith & Vane-Wright, 2001); Bitje (Smith & Vane-Wright, 2001); Mount Kala (Smith & Vane-Wright, 2001); Dja (Libert, 2007); Assobam (Libert, 2007); Yokaduma (Libert, 2007). Gabon – Waka (van de Weghe, 2010); Ipassa (van de Weghe, 2010); Belinga (van de Weghe, 2010). Central African Republic – Dzanga (Noss, 1998).

martensi Dufrane, 1946 (as sp. of Papilio (Cosmodesmus)). Bulletin et Annales de la Société Royale Entomologique de Belgique 82: 119 (101-122). Democratic Republic of Congo: “Congo Belge, Equateur”.

*Graphium (Arisbe) aurivilliusi (Seeldrayers, 1896)

Papilio aurivilliusi Seeldrayers, 1896. Annales de la Société Entomologique de Belgique 40: 499 (499-505). Papilio Aurivilliusi Seeldr., 1896. Aurivillius, 1899. Graphium aurivilliusi (Seeldrayers, 1896). Ackery et al., 1995: 162. Graphium auriger aurivilliusi (Seeldrayers, 1896). Hancock, 1993: 568. Graphium aurivilliusi (Seeldrayers, 1896). Smith & Vane-Wright, 2001.

Type locality: “Congo”. Distribution: Democratic Republic of Congo, Congo?. Early stages: Nothing published. Larval food: Nothing published.

*Graphium (Arisbe) ucalegon (Hewitson, [1865]) Creamy Swordtail

Papilio ucalegon Hewitson, 1865 in Hewitson, 1862-6. Illustrations of new species of exotic butterflies 3: [3] ([124 pp.]. London. Papilio ucalegon Hew., 1865. Aurivillius, 1899.

Alternative common name: Creamy Graphium. Published images: Larsen, 2005a (male upperside and underside). 66

Type locality: Nigeria: “Old Calabar”. [Lectotype designated by Smith & Vane-Wright, 2001 (Bulletin of the Natural History Museum Entomology Series 70 (2): 659 (503-719))]. Diagnosis: Distinguished from other species of the adamastor species group by the two basal black spots on the underside of the hindwing (Larsen, 2005a). Distribution: Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Angola, Central African Republic, Democratic Republic of Congo, Uganda, Tanzania. Recorded, in error, from Ghana (Larsen, 2005a). Habitat: Primary forest. Habits: Strongly seasonal, occurring mainly just before the beginning of the rains (Larsen, 2005a). The flight appears to be lazy but, in fact, is quite rapid (Larsen, 2005a). Males mud-puddle and also feed from rotten fish (Larsen, 2005a). They are usually seen flying along forest roads or river banks (Larsen, 2005a). Early stages: Nothing published. Larval food: Nothing published.

Graphium (Arisbe) ucalegon ucalegon (Hewitson, [1865])

Papilio ucalegon Hewitson, 1865 in Hewitson, 1862-6. Illustrations of new species of exotic butterflies 3: [3] ([124 pp.]. London. Papilio ucalegon Hew., 1865. Aurivillius, 1899.

Published images: Larsen, 2005a (male upperside and underside). Type locality: Nigeria: “Old Calabar”. Distribution: Nigeria (west), Cameroon, Equatorial Guinea (including Bioko), Gabon, Congo, Angola (north- west), Central African Republic, Democratic Republic of Congo (Mayumbe). Specific localities: Nigeria – Old Calabar (TL); Ondo (Larsen, 2005a); Abakaliki (Larsen, 2005a); Okwangwo (Larsen, 2005a); Cross River Loop (Larsen, 2005a). Cameroon – Bombe (Strand, 1912); Nlohe (Strand, 1912); Bipindi (Strand, 1913); Bipindi (Strand, 1913); Kribi (Strand, 1913); Mount Kala (Darge, 1995); Korup (Larsen, 2005a); Lomie (Libert, 2007); Ebogo (Libert, 2007); Yaounde (Libert, 2007). Equatorial Guinea – Alén Benitogebt (Strand, 1913). Gabon – Mondah (van de Weghe, 2010); Crystal Mountains (van de Weghe, 2010); Bitam (van de Weghe, 2010); Waka (van de Weghe, 2010); Inkoghou (Fernan Vaz) (van de Weghe, 2010); Akaka (van de Weghe, 2010); Tchibanga (van de Weghe, 2010); Mayumba (van de Weghe, 2010); Mouila (van de Weghe, 2010); Langoue (van de Weghe, 2010); Ipassa (van de Weghe, 2010); Nouna (van de Weghe, 2010); Ogooue (van de Weghe, 2010).

legonuca Suffert, 1904 (as ssp. of Papilio ucalegon). Deutsche Entomologische Zeitschrift, Iris 17: 106 (12-107). Cameroon: “Süd-Camerun”.

superfluosa Strand, 1912 (as ab. of Papilio ucalegon). Archiv für Naturgeschichte 78 (A.2.): 140 (136-141). Cameroon: “Bombe; Nlohe”.

addenda Strand, 1913 (as ab. of Papilio ucalegon). Archiv für Naturgeschichte 79 (A.2.): 25 (10-26). Cameroon: “Kamerun, Bipindi; Bipindi Kribi”; Equatorial Guinea: “Spanish Guinea, Alén Benitogebt”.

Graphium (Arisbe) ucalegon fonteinei Berger, 1981

Graphium ucalegon fonteinei Berger, 1981. Les Papillons du Zaire 51 (543 pp.). Bruxelles.

Type locality: Democratic Republic of Congo: “Sankuru, km. 43, route de Lusambo à Batempa”. Distribution: Democratic Republic of Congo (Sankuru, Kabinda, Lomami, Lualaba). Specific localities: 67

Democratic Republic of Congo – Sankuru, km. 43, route de Lusambo à Batempa (TL).

fontainei Berger, 1950 (as f. of Graphium ucalegon schoutedeni). Annales du Musée Royal du Congo Belge. C- Zoologie (3) 8 (1): 74 (102 pp.). Democratic Republic of Congo: “Sankuru, km. 43, route de Lusambo à Batempa”. Holotype in the Royal Museum for Central Africa, Tervuren, Belgium.

Graphium (Arisbe) ucalegon schoutedeni Berger, 1950

Graphium ucalegon schoutedeni Berger, 1950. Annales du Musée Royal du Congo Belge. C-Zoologie (3) 8 (1): 73 (102 pp.).

Type locality: Democratic Republic of Congo: “Uele, Bambesa”. Holotype and allotype in the Royal Museum for Central Africa, Tervuren, Belgium. Diagnosis: Compared to the nominate subspecies the forewing pale band is narrower and the hindwing band is indistinct (Kielland, 1990d). Distribution: Democratic Republic of Congo (east), Uganda (west), Tanzania (north-west). Specific localities: Democratic Republic of Congo – Bambesa, Uele (TL). Uganda – Maramogambo Forest, Queen Elizabeth N.P. (Tumuhimbise et al., 2001). Tanzania – Bukoba Region (Kielland, 1990d).

*Graphium (Arisbe) simoni (Aurivillius, [1899])

Papilio ucalegon var. simoni Aurivillius, 1899 in Aurivillius, 1898-9. Kungliga Svenska Vetenskapakademiens Handlingar 31 (5): 485 (1- 561).

Type locality: [Angola]: “Congogebiet, Bangala”. Distribution: Nigeria, Cameroon, Gabon, Congo, Angola, Democratic Republic of Congo, Central African Republic. Specific localities: Gabon – Oyem (van de Weghe, 2010). Angola – Bangala (TL). Early stages: Nothing published. Larval food: Nothing published.