Genus Bicyclus | Semuliki National Park

Genus Bicyclus: There are around 100 species of the Afrotropical genus Bicyclus so far described and presently the genus is undergoing a major revision. There are 26 - including B. mesogenina which was not included in the published Semuliki checklist or possibly 27 (depending on the alboplaga/xenoides taxonomic status) of the current described species to have been recorded at Semuliki. A number of species display seasonal polymorphism and B. vulgaris and B. buea were two of the most common assemblage species sampled in the fruit traps. Some females from a number of species are difficult to differentiate (eg. buea, smithi and sanaos or vulgaris and sandace) and in this instance I will still include a female specimen for reference although the specific species can't be verified. The website follows the species groups categories detailed in the 'Systematic checklist (with complete synonymic listing) of currently known Bicyclus species' by Aduse-Poku et al., (2016).

The evadne species-group and the alboplaga-complex

Although listed as 2 separate species (B. alboplaga and B. xeneoides) in 'The butterflies (Lepidoptera: Papilionoidia) of Semuliki National Park, western Uganda' the two species show little genetic differences and are morphologically variable over their range. The field images below show the specimens (females and males) of the two species trapped at Semuliki. The two species were not common.

Bicyclus alboplaga/xenoides (males left)

The ignobilis species-group

There is only one species: B. ignobilis recorded at Semuliki from this group of six described species. Only one female specimen was ever trapped and female upper and underside is much lighter. The species-group was revised by Brattström in 2015.

Bicyclus ignobilis (males above)

The sciathis species-group

This species-group was revised by Brattström et al., in 2016. Of the 13 described species in this group, three are recorded at Semuliki: hyperanthus, uniformis and procora.

Bicyclus hyperanthus (males above)

Bicyclus uniformis (males above)

Bicyclus procora (males above)

The hewitsoni species-group

This species-group has 12 described species. Only two species are recorded at Semuliki: graueri which is a common species and sebetus which was rarely trapped.

Bicyclus sebetus (males above)

Bicyclus graueri (males left)

The safitza species-group

This species-group has 3 described species. Only one species has been recorded at Semuliki: safitza which is one of the more regularly trapped Bicyclus species. This species shows seasonal polyphenism in wing pattern between dry and wet season forms with respect to eyespot diameter size.

Bicyclus safitza (males above)

The funebris species-group

There is only one species described in this group and like B. safitza, B. funebris displays seasonal plasticity in wing pattern, principally eyespot diameter.

Bicyclus funebris (males above, dry season form)

The rhacotis species-group

There are seven described species in this group with at least two officially recorded at Semuliki but most likely three: B sambulos, B mesogena and B. mesogenina. Both B. mesogena and B.mesogenina are sympatric over the Congo Basin region (and in Kibale National Park, a submontane forest around 40km south-east of Semuliki in Uganda) with differences in both forewing and andocronial brush shape and also genitalia.. B. mesogena or mesogenina was one of the more common Bicyclus species and was regularly trapped.

Bicyclus sambulos (males above)

Bicyclus mesogena (males left)

The angulosa species-group

There are 12 described species in this group with five species recorded at Semuliki: B collinsi, B. auricruda, B. campa, B. mollitia and B. angulosa. In comparison to the abundant B. buea and B. vulgaris, none of these species can be regarded as common.

Bicyclus collinsi (males above)

Bicyclus campa (males above)

Bicyclus auricruda (males left)

Bicyclus mollitia (males left)

Bicyclus collinsi and auricruda, males comparison

Bicyclus angulosa (males above)

The martius species-group

From the 14 described species in this group, five species have been recorded at Semuliki: B buea, B. sanaos, B. smithi, B. golo and B. sophrosyne. Females of some of these species are particularly difficult to identify in the field (buea and sanaos: smithi, golo and sophrosyne). In this instance an image of a female that could either be for example Sanaos or Buea, even if it may be ambiguous will be displayed. This is one of the groups identified by the work of Aduse-Poku et al., (2017) that requires a revision.

Bicyclus buea (males above)

Bicyclus golo (males left)

Bicyclus sanaos (males left)

Bicyclus smithi (males above)

Bicyclus sophrosyne (males above)

The dorothea species-group

There are five described species in this group, three species have been recorded at Semuliki: B sandace, B. jefferyi and B. vulgaris. Included within this species-group is the dorothea-complex containing three morphologically and genetically similar species: B. jefferyi, B. moyses and B. dorothea where B. jefferyi has more of an eastern distribution, ranging into Kenya. B. jefferyi was most commonly trapped along with B. safitza within the grassland patch.

Bicyclus jefferyi (males above)

Bicyclus vulgaris (males left)

Bicyclus sandace (males above)

The ena species-group

There are four described species in this group, with only one occurring at Semuliki: B campina. This species is highly variable and the specimens included here can be compared with B. campa (images further above) with respect to the discal line of the forewing underside which in B. campus is usually angled more predominantly to 90° and this can be seen in the field images.

Bicyclus campina (males above)

Field Images: There are field images for all but one of the recorded species: B angulosa. This is a savannah species and although the grassland was sampled with traps this species was never trapped.

Bicyclus xenoides (male, under and upperside)

Bicyclus xenoides (female, underside)

Bicyclus xenoides (female, upperside)

Bicyclus ignobilis (female, under and upperside)

Bicyclus ignobilis (male and female)

Bicyclus hyperanthus (female, upperside)

Bicyclus hyperanthus (male, under and upperside)

Bicyclus hyperanthus (female, underside)

Bicyclus uniformis (male, under and upperside)

Bicyclus uniformis (female, underside)

Bicyclus uniformis (female, upperside)

Bicyclus procora (female, upperside)

Bicyclus procora (male, under and upperside)

Bicyclus procora (female, underside)

Bicyclus sebetus (male, upper and underside)

Bicyclus sebetus (male, upperside showing absence of subapical band)

Bicyclus sebetus (male, underside)

Bicyclus sebetus (female, upper and underside)

Bicyclus graueri (female, upper and underside)

Bicyclus graueri (male, upperside)

Bicyclus graueri (male, underside)

Bicyclus safitza (males and female, underside - dry season form)

Bicyclus safitza (males and female, uppersides)

Bicyclus safitza (male and female, underside - wet season form)

Bicyclus safitza (miniature male, trapped during dry season)

Bicyclus funebris (male and female, underside - dry season form)

Bicyclus funebris (male underside - wet season form)

Bicyclus funebris (female,underside - wet season form)

Bicyclus funebris (males, upperside - seasonal forms)

Bicyclus funebris (females, upperside - seasonal forms)

Bicyclus sambulos (male, under and upperside)

Bicyclus sambulos (female, underside)

Bicyclus sambulos (female, upperside)

Bicyclus mesogena (male, under and upperside)

Bicyclus mesogena (female, under and upperside)

Bicyclus collinsi (male, under and upperside)

Bicyclus collinsi (female, upperside)

Bicyclus auricruda (male, under and upperside)

Bicyclus auricruda (female, upperside)

Bicyclus collinsi (female, underside)

Bicyclus auricruda (female, underside)

Bicyclus campa (male, underside)

Bicyclus campa (female under and upperside)

Bicyclus mollitia (male, under and upperside)

Bicyclus angulosa (waiting on field image)

Bicyclus mollitia (female, underside)

Bicyclus mollitia (female, upperside)

Bicyclus buea (male, under and upperside)

Bicyclus buea (female, under and upperside)

Females of Bicyclus buea and B. sanaos (species below) are impossible to identify in the field as reliable identification cues have yet to be described (female examples of trapped individuals of either B. buea and B. sanaos)

Bicyclus sanaos (male, under and upperside)

Bicyclus sanaos (female, upper and underside)

Bicyclus smithi (male, under and upperside)

Bicyclus smithi (female, under and upperside)

Females of Bicyclus smithi and B. golo (species below) are impossible to identify in the field as reliable identification cues have yet to be described (female examples of trapped individuals of either B. smithi and B. golo)

Bicyclus golo (male, under and upperside)

Bicyclus golo (female, underside)

Bicyclus golo (female, upperside)

Bicyclus sophrosyne (male underside)

Bicyclus sophrosyne (male upperside)

Bicyclus sophrosyne (female underside)

Bicyclus jefferyi (male, underside)

Bicyclus jefferyi (male, upperside)

Bicyclus jefferyi (female, underside)

Bicyclus jefferyi (female, upperside)

Females of Bicyclus jefferyi, B. vulgaris and B. sandace can be impossible to identify in the field as reliable identification cues have yet to be described (female examples of trapped individuals of possibly either B. jefferyi, B vulgaris and B. sandace)

Bicyclus vulgaris (male, underside)

Bicyclus vulgaris (male, upperside)

Bicyclus vulgaris (female, underside)

Bicyclus vulgaris (female, upperside)

Bicyclus sandace (male, under and upperside)

Bicyclus sandace (female, underside)

Bicyclus campina (male, underside)

Bicyclus campina (female, underside)

Specific genus site links, references and bibliography:

Aduse‐Poku, K., Brakefield, P.M., Wahlberg, N. & Brattström, O., (2016). Supplemental data S5: Systematic checklist (with complete synonymic listing) of currently known Bicyclus species.

Aduse‐Poku, K., Brakefield, P.M., Wahlberg, N. & Brattström, O., (2017). Expanded molecular phylogeny of the genus Bicyclus (Lepidoptera: Nymphalidae) shows the importance of increased sampling for detecting semi-cryptic species and highlights potentials for future studies. Systematics and Biodiversity, 15(2), 115-130.

Brakefield, P.M. & Reitsma, N. (1991). Phenotypic plasticity, seasonal climate and the population biology of Bicyclus butterflies (Satyridae) in Malawi. Ecological Entomology 16, 291-303.

The two papers below describe ongoing taxonomic revision of the genus.

Brattström, O., Aduse‐Poku, K., Collins, S.C. & Brakefield, P.M., (2015). Revision of the Bicyclus ignobilis species‐group (Lepidoptera: Nymphalidae: Satyrinae) with descriptions of two new species. Zootaxa 4018, 57‐79.

Brattström, O., Aduse‐Poku, K., Collins, S.C., Di Micco De Santo, T. & Brakefield, P.M., (2016). Revision of the Bicyclus sciathis species group (Lepidoptera: Nymphalidae) with descriptions of four new species and corrected distributional records. Systematic Entomology 41, 207‐228.

Condamin, M., (1973). Monographie du Genre Bicyclus (Lepidoptera, Satyridae). L’Institut Fondamental D’Afrique Noire, Dakar. To date the most comprehensive resource covering this genus, unfortunately, all images are in black and white.

Dongmo, M.A.K, Bonebrake, T.C., Hanna, R. & Fomena, A. (2018). Seasonal polyphenism in Bicyclus dorothea (Lepidoptera:
Nymphalidae) across different habitats in Cameroon. Environmental Entomology, 47(6), 1601–1608.

Nymphalidae.Net provides a detailed taxonomic overview of the Subtribe Mycalesina. Subtribe Mycalesina (Wahlberg, N).

Williams, M.C., (2020). Genus Bicyclus. A section of Afrotropical Butterflies (17th Edition). Publication is available from Lepidopterists' Society of Africa website https://www.lepsocafrica.org/?p=publications&s=atb.

Online resource. www.bicyclus.se. The research site of Oskar Brattström for up to date research and publications on all things Bicyclus.

BUTTERFLY Research, Conservation and Education Semuliki National Park, Uganda