Corresponding author: Tomasz W. Pyrcz (
Academic editor: C. Peña
Updated data on the distribution, ecology and taxonomy of
Recent years have witnessed a rising interest in advanced studies focusing on species-rich African brush-footed butterflies (
The subgenus
Adults of
Sierra Leone
1 ♂: Sierra Leone, Guma, 01.03.1982, prep. genit. 10/09.05.2012, J. Lorenc; 1 ♀: same data, prep. genit. 11/09.05.2012, J. Lorenc; 1 ♀: Sierra Leone, Guma Valley, 400 m, 11.1991; 1 ♀: Sierra Leone, no locality, no date; 1 ♂: Ivory Coast, Abidjan (erroneous locality), 1966, S. Collins leg.; 1 ♀: same data (erroneous locality), ABRI; 6 ♂ and 2 ♀: Sierra Leone, Guma, 01.III.1982, H. Warren-Gash leg.; 1 ♂: Guinea, Diecke, 07.2000, HWG; 1 ♂ and 5 ♀: Sierra Leone, SMTD; 4 ♂: Guinée, Guinée forestière, Province de Yomou, Forêt classée de Diecke, 1-26.III.2003, leg. Ph. Leonard and E. Vingerhoedt; 3 ♂ and 1 ♀: same data but VI.2003; 2 ♂: same data but III.2005, PhO; 3 ♂ and 1 ♀: Guinée, Forêt classée de Diecké, GF; 2 ♂ and 2 ♀: Liberia, Mount Swa, Sz. Sáfián leg, MZUJ; 1 ♂: same data, ABRI.
Upperside colour black and apple green with a delicate yellow sheen, somewhat more prominent on the HWD median area.
Adults, males:
The known range of this subspecies extends from Sierra Leone, Liberia to SE Guinea. The pair in ABRI labeled as coming from Abidjan represent a conundrum. However, given the extensive research and collecting by one of the authors (HWG) in the Abidjan area without encountering any other specimens, we have concluded that they must be mislabeled.
Tai Forest (Forêt de Taï), Ivory Coast
12 ♂: Ivory Coast, Tai, 13–14.05.2000, S. Collins leg.; 1 ♂: same data, prep. genit. 07/09.05.2012, J. Lorenc; 1 ♀: same data; 1 ♀: same data, prep. genit. 08/09.05.2012, J. Lorenc; 1 ♂: same data but XII.1998, K. Adams leg., ABRI; 2 ♂: Ivory Coast, Tai, 12.I.1999, H. Warren-Gash leg.; 1 ♂: Ivory Coast, Tai, 13.I.1999, H. Warren-Gash leg.; 1 ♂ and 1 ♀: Ivory Coast, Tai, XII.1998, K. Adams; 2 ♂ and 1 ♀: Ivory Coast, Tai, 02.I.2000, H. Warren-Gash leg.; 1 ♂ and 1 ♀: Ivory Coast, Tai, 13.V.2000, H. Warren-Gash leg., HWG.
Upperside colour black and pine green.
The known distribution is limited to the Tai National Park in western Ivory Coast. The same pine green colour can be found in
Mount Nimba, Guinea
Upperside black and golden green, similar to
The subspecific epithet of this taxon derives from its type locality, the massif of Mount Nimba (1750 m) on the Ivory Coast – Guinea border.
The range is apparently disjunct with most specimens coming from the lower slopes of Mont Nimba and the surrounding area, and two collected in the Fouta Djalon range in northern Guinea, the latter record needs confirmation.
N. Awasso, Bibiani, Western Region, Ghana
Upperside black and golden green with a yellow subapical patch strongly suffused with green, compared to the slightly suffused patch in
Adults, males:
Adults, females:
The epithet of this taxon is an allusion to its resemblance to the next subspecies,
A widely distributed subspecies found in coastal forests in the Ivory Coast and Ghana, and also in inland forests in eastern Ivory Coast and Ghana.
Lokoja, Nigeria
1 ♂: Nigeria, Butatong, Calabar, 14.XI.1995, S. Collins leg.; 1 ♀: same data but 14–18.XI.1995; 2 ♂: Nigeria, east, T. Davey leg.; 1 ♀: same data; 1 ♂: Nigeria, Lagos, IX.1957, J. Boorman leg.; 1 ♀: same data but X.1962; 3 ♂: Nigeria, Mamu Forest, IV.1966, S. Collins leg.; 1 ♀: same data; 8 ♂: Nigeria, Obudu, 09–12.XI.1995, S. Collins leg.; 1 ♂: same data, prep. genit. 12/09.05.2012, J. Lorenc; 2 ♀: same data; 1 ♀: same data, prep. genit. 13/09.05.2012, J. Lorenc; 1 ♂: Nigeria, Okrasa, Calabar, S. Collins leg.; 1 ♀: Nigeria, Old Ekuri Calabar, 24–28.XI.1995, S. Collins leg.; 2 ♀: Nigeria, Olokomeji, IX.1957, J. Boorman leg.; 2 ♂: Cameroon, Ebogo, IV.1983, S. Collins leg., prep. genit. 16/09.05.2012, J. Lorenc, ABRI; 3 ♀: Nigeria, Isheri F., XII.1989, II-III.1990, A. A. Knoop Lindeboom leg; 2 ♂: Nigeria, Oban XI.1995, T. B. Larsen leg.; 1 ♂: Nigeria, Anambra F. R., VII.1962, St. Leger leg.; 6 ♂: Nigeria, Isheri F., IV.1988, II.1989, IV.1989, XII.1989, II.1990, A. A. Knoop Lindeboom leg.; 1 ♂: Nigeria, Isheri F., II.1991, D. P. Knoop leg. DK; 2 ♂: Nigeria, Isheri Forest, 29.III.1989, A. Knoop leg.; 3 ♂: same data but 09.V.1990; 1 ♂: same data but 23.I.1990; 1 ♂: same data but 14.II.1990; 1 ♂: same data but 09.IV.1990; 2 ♀: same data but I.1990; 1 ♀: same data but 10.III.1990; 1 ♀: same data but 05.IV.1989; 1 ♀: same data but 18.II.1990; 1 ♂: Nigeria, Olokomeji, 16.V.1988, A. Knoop leg.; 1 ♂: same data but 20.IV.1988; 1 ♀: same data; 1♂: Nigeria, Anambra State, Nsukka, 28.IX.1982, J. Wojtusiak leg.; 1 ♂: same data but 01.II.1983; 2 ♂: no data, MZUJ; 1 ♂: Nigeria, Olokomeji, 13.V.1989, leg. A. Knoop; 1 ♀: same locality, 20.IV.1988, leg. A. Knoop, PhO; 1 ♀: Gabon, Mondah, VIII.1993, leg. G. Faravel, GF.
Upperside colour black and mint green with a golden yellow subapical patch, usually with little or no green overcast.
Adults, females:
Found in the rain forests throughout southern Nigeria, western Cameroon and Gabon. An individual (a female) collected by Faravel in Mondah, Gabon, is tentatively associated with this subspecies although it is lighter orange on the FW and HWD, and is therefore similar to
Mongoumba, Central African Republic
Upperside colour black and olive drab, same as in the nominate
The epithet of this taxon is an allusion to its resemblance to the nominotypical subspecies.
On present knowledge, the range of this subspecies is limited to the CAR and eastern Cameroon, though its distribution pattern indicates it also occurs in adjacent areas of the DRC. The individual collected on an offshore platform in the Gulf of Guinea is an extraordinary finding. It is also a proof that
Ghana “Gold Coast”
1 ♂: Western Ghana, Bibiani, N. Awaso, 250–400 m, 09.IV.2007, ABRI; 1 ♂: same data, prep. genit. 04/20.06.2012, J. Lorenc; 1 ♀: Nsukka, Anambra State, Nigeria, 04.XII.1982, leg. J. Wojtusiak, prep. genit. 02/20.06.2012, J. Lorenc; 1 ♀: Banco N. P., Ivory Coast, 18.IX.1999, H. Warren-Gash leg., prep. genit. 05/20.06.2012, J. Lorenc, MZUJ; 1 ♂: Ghana, Oda Big Tree, XII.2010, Sáfián, Sz. leg., SZS; 1 ♂: Ghana, Eastern Region, Asuom Amanfrom, Amanfrom Forest, Kade District, 20–24.III.2005, Sáfián, Sz., Csontos, G. and Kormos, B. leg., SZS; 1 ♂: Ivory Coast, Alepe-Yaya, 06.II.1999, H. Warren-Gash leg.; 1 ♂: Ivory Coast, Alepe-Yaya, 12.XII.1999, H. Warren-Gash leg. 1 ♂: Ivory Coast, Lamto, 05.XI.2000, H. Warren-Gash leg.; 2 ♂ and 2 ♀: Ivory Coast, Alepe, 21.I.2000, H. Warren-Gash leg.; 1 ♀: Ivory Coast, Alepe, 07.I.2001, H. Warren-Gash leg., HWG; 1 ♂ and 1 ♀: Ivory Coast, Teapleu, V.1977; 10 ♂ and 3 ♀: Guinée, Guinée forestière, Province de Yomou, Forêt classée de Diecke, VI.2003, Eric Vingerhoedt leg.; 1 ♂: Cameroon, Douala, IX.1996, leg. P. Prouvost; 1 ♂: Cameroon, no exact locality, II.1997, E. Vingerhoedt, 1 ♂: Cameroon, Ebogo, I.1994, T. Garnier leg., PhO; 40 ♂ and 28 ♀: Ghana; 2 ♂ and 2 ♀: Ivory Coast; 1 ♂: S. Nigeria, Ewohimi, 29.XI.55, J. Boorman; 1 ♂: no label; 2 ♂: Cameroon, Ebogo, Mbalmayo, X.2001; 1 ♂: Cameroon, Ebogo, II.1994, S. Collins leg., ABRI; 1 ♂: Ghana, Kibi Atewa Mts., II.2007., J. Boersma; 1 male ♂: Nigeria, Ewohimi, I.1955, J. St. Leger leg., DK; 2 ♂: Nigeria, Sapoba, III.1967, J. Riley leg., ex Cornes and Riley coll., DK; 2 ♂ and 2 ♀: Liberia, Mount Swa, Sz. Sáfián leg, MZUJ; 1 ♂: same data, ABRI; 1 ♂: same data, SSS.
Upperside ground colour predominantly golden green, with the exception of the light yellow subapical patch.
Adults, males:
The nominate subspecies is widely distributed from central Liberia across Ivory Coast, Ghana and Nigeria and into Cameroon and northern Gabon, in both in perhumid coastal forests and drier forests in the interior. It is phenotypically quite stable throughout with the notable exception of the Mount Swa population in Liberia where all green individuals were detected flying alongside typically patterned ones. Such an individual variation may be explained by phenotype selection to a heterogenous environment of patchy rain forest and savanna. Nevertheless, it remains to be confirmed that the green and typical morphs are indeed syntopic and we cannot exclude the possibility that the green morph represents a separate parapatric subspecies.
Tai Forest, Ivory Coast
1 ♂: Ivory Coast, Tai, 12.I.1999, H. Warren-Gash leg.; 3 ♂: Ivory Coast, Tai, 02.I.2000, H. Warren-Gash leg.; 1 ♂: Ivory Coast, Tai, XII.1998, K. Adams leg.; 1 ♀: Ivory Coast, Tai, 01–10.IV.2001, H. Warren-Gash leg., HWG; 1 ♂: Ivory Coast, Moyen Cavally, Tai Forest, IV.2001, prep. genit. 03/09.11.2012, J. Lorenc, HWG; 1 ♂: same data but 02.I.2000, MZUJ; 1 ♂: Tai, Ivory Coast, 13–14.VIII.2001, S. Collins; 1 ♂: same data but 13.IX.2001; 1 ♂: same data but 13–14.V.2000 (paratype
Upperside ground colour black and pine green.
Adults, males:
Adults, females:
Known distribution is limited to the Tai National Park in western Ivory Coast.
Mount Nimba, Guinea
Upperside ground colour golden yellow, same as in
The subspecific epithet of this taxon derives from iron in Latin,
Distribution limited to the Mount Nimba area. Considered that southern Ivory Coast was extensively sampled for several years, also by one of the authors (HWG), we believe that one paratype was labeled as from “Azagny” by error.
Ketta Forest, Ouesso, Moyen Congo
Upperside ground colour black and apple green.
This subspecies was described from Ouesso in Congo, and apart from the holotype there are only a couple of other specimens known.
Ititye Camp, Mihuno, 25 miles east of Kigoma, Western Province, Tanzania.
3 ♀: Kasye Forest, Kigoma, Tanzania, III.1990, leg. local dealer, ex coll. P. Boyer, MZUJ; 3 ♂: R.C.A., Bangui, C.A.R., VIII.2000, S. Collins leg.; 1 ♂: R.C.A., Bangui, IX.2000, S. Collins leg., 2 ♂ and 1 ♀: R.C.A., Bangui, IV.2004, S. Collins leg.; 1 ♂: R.C.A., Bomoloto, R. Longamp, VII.1985, S. Collins leg.; 2 ♀: R.C.A., Bangui, VIII.2000, S. Collins leg.; 1 ♀: R.C.A., I.2004, S. Collins leg.; 2 ♂: R.D.C. (NW), Lukolela, Congo River, V.2012; 11 ♂ and 10 ♀: Tanzania, ABRI; 1 ♂: Congo, Province de l’Equateur, Mbandaka, 12.II.1998, prep. genit. 01/04.12.2012, J. Lorenc; 1 ♂: same data, prep. genit. 05/04.1.2012, J. Lorenc; 1 ♀: same data but XII.1997, prep. genit. 08/04.12.2012, J. Lorenc; 1 ♂: Congo, Province de l’Equateur, Environ de Mbandaka – Kuluboku, 27.IV.1995, prep. genit. 06/04.1.2012, J. Lorenc; 1 ♀: same data but 15.XI.1995, prep. genit. 07/04.12.2012, J. Lorenc; 1 ♂: Congo, Province de l’Equateur, Lukolela, 26.X.1993, prep. genit. 03/04.12.2012, J. Lorenc; 1 ♂: same data but 27.X.1993, prep. genit. 04/04.12.2012, J. Lorenc; 1 ♂: same data but XI.1993, PhO; 1 ♂: Sandoa, Katanga, R.D.C., Sandoa, 16.I.1991; 1 ♂: same data but 30.X.1920; 1 ♂: Kafakumba, Katanga, R.D.C., VII.1939; F.G. Overlaet leg.; 2 ♂: same data but II.1939; 1 ♂: same data but IX.1933; 1 ♂: Kinda, Katanga, R.D.C., 31.III.1916; 1 ♂: same data but 31.VIII.1916; 1 ♂: Montamba, 24.V.1916; 1 ♂: Kondué, East Kasai, R.D.C., no date, E. Luja leg.; 1 ♂: Tshilolo, Sankuru, East Kasai, 10.III.1951, Dr. Vallard leg.; 1 ♀: Kafakumba, Katanga, R.D.C., II.1939, F. G. Overlaet leg.; 1 ♀: same data but III.1939; 1 ♀: same data but IV.1939; 1 ♀: same data but IX.1933; 1 ♀: Kinda, 27.II.1916, F. G. Overlaet leg., MRAC.
Upperside ground colour golden yellow, same as in
Adults, females:
Male genitalia (in lateral view, aedeagus extracted):
Female genitalia (in lateral view):
Male genitalia (in lateral view, aedeagus extracted):
Female genitalia (in lateral view):
Distribution map:
This subspecies was described from the now largely destroyed Kasye forest in western Tanzania. It is however widely distributed across the Congo Basin south to Katanga (Shaba). The individuals coming from the area of Mbandaka in the western part of the Congo Basin are attributable to this subspecies. They present however an unusual, apparently individual variation in the ground colour of the upperside, ranging from orange-yellow, through typically golden yellow to yellow with a strong green overcast. Such an apparent individual variation, quite common in some species of
In terms of geographical range, the two species are largely sympatric, though, as noted above,
The apparent gaps in the range of
In terms of habitat, the ecological preferences of
It is interesting to observe that where the two species fly sympatrically, as they do through much of their range, the wing colour markings in one species are closely matched by the other. There are two plausible explanations for that. Either, it is a parallel evolution reflecting an adaptation to a particular forest light structure, a common background. In fact, all green populations are found in rain forest (
Apart from the slender body and elongated wings, we did not find any other outstanding morphological characters in
The most interesting trait of
In the light of this study, previously published articles and the ongoing research on the taxonomy of
The authors would like to thank Steve C. Collins (Nairobi) for hosting T. Pyrcz and Sz. Sáfian in Nairobi and for the unrestricted access to the ABRI collections, Teresa di Micco de Santo (Nairobi) for her assistance in photography and data basing in ABRI; Torben B. Larsen (Copenhagen) for cross checking the identifications of