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FAMILY PAPILIONIDAE

Subfamily Papilioninae

A comprehensive and recognised stable phylogeny for the genus Papilio has been an ongoing problem and debated from a number of studies over the last 40 years. The most recent research has created a time-calibrated species-level Papilio molecular phylogeny producing a reference framework (Condamine et al., 2023).

 

From a total of approximately 600 species within this subfamily, there is a global total of 235 Papilio species of which 65 occur within the Afrotropical region. Afrotropical species have been split into four subgenera, 22 species being recorded in Uganda. There is a global total of 100 Graphium species, of which 41 are found within the Afrotropical region and 12 within Uganda. Within the park there have been 19 species from the two genera recorded at the park; Papilio with 14 species and Graphium with five. Being nectar feeders they were never found within traps and none of the species recorded within the park could be described as common and generally would be found within the forest fringes.

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Subgenus Arisbe

There are a number of subgenera within the genus, with only one occurring in the Afrotropical region - Subgenus Arisbe. This subgenus is divided into seven species groups. 

Genus Graphium 

There are 41 recognised Graphium species within the Afrotropical region from around 100 species globally. The five species so far recorded at Semuliki are Graphium policenes, G. leonidas, G. antheusG. angolanus and G. ucalegon. A further eightspecies have been recorded from the DRC Ituri Forest and Semliki Valley. Graphium ucalegon is the only new species record additional to the Forbes (2018) species list.

​Antheus species group. A small group with two described species, one occurring at Semuliki: Graphium antheus.

Graphium antheus (Cramer, 1779)

Graphium antheus.jpeg
Graphium antheus.jpeg

Graphium antheus (right forewing, upper and underside, sexes similar)

Policenes species group. Containing 11 species with one recorded from Semuliki, Graphium policenes.

Graphium policenes policenes (Cramer, 1775)

Graphium policenes.jpeg
Graphium policenes.jpeg

Graphium policenes (upper and underside, sexes similar)

Leonidas species group. A small group containing three species with one recorded from Semuliki, Graphium leonidas.

Graphium leonidas leonidas (Fabricius, 1793)

G. leonidasA.jpg

Graphium leonidas (underside, sexes similar)

Image by Timothy Lloyd (original posted on inaturalist)

Adamastor species group. The largest group containing 16 species with one recorded from Semuliki, Graphium ucalegon.

Graphium ucalegon schoutedini (Berger, 1950)

Graphium ucalegon.jpg

Graphium ucalegon (male, upperside)

Image by Sven Bontenbal (Semuliki National Park Collection - www.observation.org)

Angolanus species group. A group containing six species with one recorded from Semuliki, Graphium angolanus.

Graphium angolanus baronis (Ungemach, 1932)

Graphium angolanus (waiting on field images)

Genus Papilio
The 65 Afrotropical species are divided into four subgenera: PapilioPrinceps, Druryia and Nireopapilio. The 14 species recorded at Semuliki are within three of the subgenera. Each subgenus has been subdivided into species groups.

Subgenus Nireopapilio
This has been split into three groups: zalmoxis, hesperus and nireus. The seven Papilio species within this subgenus from Semuliki are divided within two of these groups: hesperus and nireus. The zalmoxis group is represented by only one species, Papilio zalmoxis which has been recorded from the DRC Ituri Forest. 

Nireus species group. A group comprising 21 species, with six being recorded from Semuliki: Papilio nireus, P. mackinnoni, P. nobilis, P. sosia, P. charopus and P. chrapkowskoides.

Papilio nireus nireus (Linnaeus, 1758) 

Papilio nireus.jpeg
Papilio nireus.jpeg

Papilio nireus (male, upper and underside)

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Papilio nireus.jpeg

Papilio nireus (female, upper and underside)

Papilio chrapkowskoides chrapkowskoides (Storace, 1952)

Papilio chrapkowskoides.jpeg
Papilio chrapkowskoides.jpeg

Papilio chrapkowskoides (male, upper and underside, sexes similar)

Papilio sosia pulchra (Berger, 1950)

Papilio sosia (waiting on field images)

Papilio nobilis crippsianus (Stoneham, 1936)

Papilio nobilis (waiting on field images)

Papilio charopus montuosus (Joicey and Talbot, 1927)

Papilio charopus (waiting on field images)

Papilio mackinnoni mackinnoni (Sharpe, 1891)

Papilio mackinnoni (waiting on field images)

Hesperus species group. A group comprising four species, with one being recorded from Semuliki: Papilio hesperus.

Papilio hesperus hesperus (Westwood, 1843)

Papilio hesperus (waiting on field images)

Subgenus Druryia
This subgenus has been split into seven groups, included an undefined group (incertae sedis). The six Papilio species within this subgenus recorded from Semuliki are divided into four of these groups: cynorta, dardanus, zenobia and menestheus

Cynorta species group. A group comprising three species, with one being recorded from Semuliki: Papilio cynorta.

Papilio cynorta (Fabricius, 1793)

Papilio cynorta.jpg
Papilio cynorta.jpg

Papilio cynorta (male, upper (white band) and underside)

Papilio cynorta.jpg
Papilio cynorta.jpg
Papilio cynorta.jpg
Papilio cynorta.jpg

Papilio cynorta (male, upper (cream band) and underside)

Papilio cynorta (female, upper and underside)

Dardanus species group. A group comprising four species, with two being recorded from Semuliki: Papilio dardanus and P. phorcas. Papilio dardanus is categorised into a number of subspecies (possibly around 13) from male traits. The two subspecies recorded at Semuliki are Papilio dardanus dardanus and P. dardanus meseres. Six forms (based on female traits) have been described within the subspecies, three of which are found at Semuliki f. hippocoonf. planemoides and f. trophonissa (waiting on field images).

Papilio dardanus dardanus (Brown, 1776) \ Papilio dardanus meseres (Carpenter, 1948) 

Papilio dardanus.jpg

Papilio dardanus (male, underside)

Papilio dardanus.jpeg
Papilio dardanus.jpeg

Papilio dardanus dardanus f. hippocoon (female, upper and underside)

Image by Sven Bontenbal (Semuliki National Park Collection - www.observation.org)

Papilio dardanus.jpeg
Papilio dardanus.jpeg
Papilio dardanus.jpeg

Papilio dardanus meseres f. mixtoides (female, upper and underside)

Papilio dardanus meseres f. mixtoides (female, upperside)

Papilio dardanus.jpeg

Papilio dardanus meseres f. mixtoides (female, underside)

Papilio dardanus.jpeg
Papilio dardanus.jpeg

Papilio dardanus dardanus f. planemoides (female, upper and underside)

Papilio phorcas congoanus (Rothschild, 1896)

Papilio phorcas.jpeg
Papilio phorcas.jpeg

Papilio phorcas (male upper and underside, sexes similar)

Papilio phorcas.jpeg

Papilio phorcas (male upperside, yellow form) 

Papilio phorcas.jpeg

Papilio phorcas (male underside, yellow form)

Zenobia species group. A group comprising ten species, with two being recorded from Semuliki: Papilio zenobia and P. mechowi. 

Papilio mechowi mechowi (Dewitz, 1881)

Papilio mechowi.jpeg
Papilio mechowi.jpeg

Papilio mechowi (male, upper and underside)

Papilio mechowi.jpeg

Papilio mechowi (worn male, faded black upperside )

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Papilio mechowi (male, underside)

Papilio zenobia (Fabricius, 1775)

Papilio zenobia.jpeg
Papilio zenobia.jpeg

Papilio zenobia (sexes similar, upper and underside)

Papilio zenobia.jpeg

Papilio zenobia (sexes similar, upperside)

Papilio zenobia.jpeg

Papilio zenobia (sexes similar, underside)

Menestheus species group. A group comprising three species, with one being recorded from Semuliki: Papilio lormieri. 

Papilio lormieri lormieri (Distant, 1874) / Papilio lormieri semlikana (Le Cerf, 1924)

Papilio lormieri (waiting on field images)

Subgenus Princeps
This subgenus comprises only one species group - demodocus. 

Demodocus species group. A group comprising four species, with one being recorded from Semuliki: Papilio demodocus. 

Papilio demodocus demodocus (Esper, 1798)

Papilio demodocus.jpeg
Papilio demodocus.jpeg

Papilio demodocus (sexes similar, upper and underside)

Specific site links, references and bibliography

​Bernardi, G,, Pierre, J. and Nguyen, T.H. (1985). Le polymorphisme et le mimétisme de Papilio dardanus Brown [Lep. Papilionidae]. Bulletin de la Société Entomologique de France, 90 (1-4).

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Clarke, C.A., and Sheppard, P.M. (1960). The genetics of Papilio dardanus Brown II. Races Dardanus, Polytrophus, Meseres and Tibullus. Genetics, 45 (4).

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Clarke, C.A. and Sheppard, P.M. (1960). The evolution of mimicry in the butterfly Papilio dardanus. Heredity, 14.

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Condamine, F.L., Allio, R., Reboud, E.l. et al. (2023). A comprehensive phylogeny and revised taxonomy illuminate the origin and diversification of the global radiation of Papilio (Lepidoptera:Papilionidae). Molecular phylogenetics and evolution 183.

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Eltringham, H., Cantab, M.A. and Oxon, M.A. (1910). African Mimetic Butterflies. Clarendon Press, Oxford.

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Rumbucher, K., (2023). Butterflies of Africa Pt 3. Papilionidae II, Princeps Hübner, (1807), Papilio dardanus species group 1. Oliver Schaffler, Munich.

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Thompson, M.J. and Timmermans, M.J.T.N. (2014). Characterising the phenotypic diversity of Papilio dardanus wing patterns using an extensive museum collection. PLOS One, https://doi.org/10.1371/journal.pone.0096815.

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