''Charaxes saturnus'', the foxy charaxes or koppie charaxes, is a

butterfly Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

that flies through most of the

Savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...

of the Eastern and Southern

Afrotropical realm The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the island of Madagascar, and the islands of the western Indian Ocean. It was formerly known as the Ethiopi ...

, and also occurs in suitable forest habitat locations including the forest belt of west-central Africa.

Description

''Charaxes saturnus'' tat.rev.2005/small> is a medium to large butterfly, with forewing length of 40–44 mm in the male, and 46–50 mm in the female.van Someren (1963), Revisional Notes on African ''Charaxes'' (Lepidoptera:Nymphalidae) Part 1. Bull. Br. Mus. nat. Hist. (Ent.) 13,(7), 205 The background colour of all four wings is a very dark blackish brown, with a broad orange postdiscal band traversing both pairs of wings from the leading edge of the forewings almost to the anal margin of the hindwings. The forewing outer margins are orange traversed by black-scaled veins. The hindwing has six orange marginal lunules, the lower three are white-edged to completely white. There are typically three or four blue submarginal spots towards the anal angle. The body and the basal area of all four wings is a tawny orange-brown. The underside exhibits a very characteristic mosaic appearance broadly similar to a number of closely related species, traversed by a jumble of bands, and of reddish, brown, and greyish patches, all edged with a filigree of white. Beyond the inner mosaic, a white complete postdiscal band bridges across fore- to hindwings. The outer orange marginal coloration is present on the underside as narrow white-edged lunules, bounded by a grey submarginal band. The female resembles the male but is larger. Both sexes of ''Charaxes saturnus'' are similar to females of '' Charaxes achaemenes''. and '' Charaxes guderiana''.

Hybridisation

Where ''C. saturnus'' is

sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...

with '' Charaxes epijasius'' tat.rev.2005/small>, over an extensive zone of overlap, the two species hybridize regularly, producing highly variable transitional specimens(''C. saturnus'' x ''C. epijasius''). The distribution range of hybrid forms extends from

Ethiopia Ethiopia, officially the Federal Democratic Republic of Ethiopia, is a landlocked country located in the Horn of Africa region of East Africa. It shares borders with Eritrea to the north, Djibouti to the northeast, Somalia to the east, Ken ...

to Western Kenya & Northern Tanzania. The variable hybrid forms have historically been named as ''harrisoni'', ''saturnalis'', pl. 4, f. 20-24, pl. 5, f. 25-28; TL= "Kenya, South Kavirondo, Suna" and ''pagenstecheri'' Observation of hybrid forms in Tanzania, as an example, beyond the recognised range of ''C. epijasius'', strongly indicates that the hybrid forms may exist as fertile hybrids, at a lower prevalence than the relatively more stable phenotype of ''C. saturnus''. More detailed phylogenetic research and breeding studies are required to elucidate further the relationships between the two species and their intermediate hybrid phenotypes, and the degree of fertility exhibited amongst their highly variable intermediate forms.

Subspecies

*''C. saturnus saturnus'', Butler, 1866. (TL="Interior of Africa") In the western areas of its range, it is more variable which has led to some earlier authors to suggest a western subspecies (e.g. Poulton, 1926, TL= N. Angola). Recent detailed re-evaluation found no western locality-depended areas with consistent population phenotypes which could merit infraspecific definition, including in Angola.L. F. Mendes, A. Bivar-de-Sousa, S. Vasconcelos & C. Van-Dúnem Santos (2017). Description of two new subspecies and notes on ''Charaxes'' Ochsenheimer, 1816 (Lepidoptera: Nymphalidae) of Angola. SHILAP Revta. lepid., 45 (178) 2017: 299-315. See p.303-4 for discussion 312,313 Figs. 7-18 onlin
hosted by Researchgate here
/ref> The ''Charaxini'' taxa ''harrisoni'', ''saturnalis'', and ''pagenstecheri'' are now thought to be at least partially fertile hybrid variable phenotypes of (''C. saturnus'' x '' C. epijasius''). This would suggest that evolutionary divergence to fully stable species separation between ''C. epijasius'' and ''C. saturnus'' is not complete. Further

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

research is required to clarify the current position further.

Related species

Historical attempts to assemble a cluster of presumably related species into a "''Charaxes jasius'' Group" have not been wholly convincing. More recent taxonomic revision, corroborated by phylogenetic research, allow a more rational grouping congruent with cladistic relationships. Within a well-populated

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

of 27 related species sharing a common ancestor approximately 16 mya during the

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

, 26 are now considered together as The ''jasius'' Group. One of the two lineages forms a robust clade of seven species sharing a common ancestor approximately 2-3 mya, i.e. during the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58"Out of Africa again: A phylogenetic hypothesis of the genus ''Charaxes (Lepidoptera: Nymphalidae)'' based on five gene regions"
. Aduse-Poku, Vingerhoedt, Wahlberg. Molecular Phylogenetics and Evolution (2009) 53;463–478 and are considered as the ''jasius'' subgroup. The ''jasius'' Group (26 Species) Clade 1: ''jasius'' subgroup (7 species): *''

Charaxes jasius ''Charaxes jasius'', the two-tailed pasha, is a butterfly in the family Nymphalidae. It is the only European species of the genus ''Charaxes''. Divergence of the Mediterranean species ''C. jasius'' from the last common ancestor with its closest r ...

'' *'' Charaxes epijasius'' tat.rev.2005ref name="Turlin 2005"/> *'' Charaxes legeri'' *''Charaxes saturnus'' tat.rev.2005ref name="Turlin 2005"/> *''

Charaxes pelias ''Charaxes pelias'', the protea emperor or protea charaxes, is a butterfly of the family Nymphalidae, and is endemic to the Cape Provinces in South Africa.Victor Gurney Logan Van Someren, 1963 Revisional notes on African ''Charaxes'' (Lepidopte ...

'' *'' Charaxes castor'' *'' Charaxes hansali'' Clade 2: contains the well-populated three additional subgroups (19 species) of the jasius Group, called the ''brutus'', ''pollux'', and ''eudoxus'' subgroups. Further exploration of the phylogenetic relationships amongst existing ''Charaxes'' taxa is required to improve clarity.

Distribution

Where appropriate habitat occurs, from Ethiopia, Somalia, Kenya, Uganda, Tanzania, across to the west in Central African Republic, Gabon, Democratic Republic of Congo, Angola, and all of southern Africa down to East London (Eastern Cape Province), but not the Mediterranean climate of the Weastern Cape Province, where it is replaced by ''

''. It is not known to occur in Equatorial Guinea, Rwanda or Burundi.Global Biodiversity Information Facility
''Charaxes saturnus''
/ref>

Habitat

Savannah and open woodland, from sea level – 2200 m; lower slopes of Kilimanjaro; and similar habitats found across the South African

Veldt Veld ( or , Afrikaans language, Afrikaans and Dutch language, Dutch: ''veld'', field), also spelled veldt, is a type of wide-open, rural landscape in Southern Africa. Particularly, it is a flat area covered in grass or low scrubland, scrub, ...

including

Koppie An inselberg or monadnock ( ) is an isolated rock hill, knob, ridge, or small mountain that rises abruptly from a gently sloping or virtually level surrounding plain. In Southern Africa, a similar formation of granite is known as a koppie, an ...

habitats. In Angola and the central African forest belt, it occurs in open and closed-canopy woodland and secondary forest areas also.

Natural history

''C. saturnus'' is continuously brooded and flies throughout the year. File:Foxy Emperor (Charaxes jasius saturnus) and Fruit Chafers (Chondrorrhina trivittata) and others (6029321985).jpg, alt=Foxy Emperor (Charaxes saturnus) & Fruit Chafers, Kruger, South Africa, Foxy emperor (''Charaxes saturnus''), Kruger, South Africa

Life cycle

The early stages from egg to pupa are described in detail by Dr. V. G. L. van Someren & Rev. K. St. A. Rogers hosted by Biodiversity Heritage Library (BHL)

Larva

After hatching, the larva eats the remaining eggshell, rests for 12hrs, and commences nocturnal habit of devouring leaves on the host plant, non-selective as to age and size of leaf. The larva spins a silken refuge on a selected leaf to rest during the day. The larva is initially yellowish-olive with a blackish-brown head bearing short tubercles. The body is finely papillated, and two pale tails arise from the anal segment. After the first moult, the larva is green, with a green face.

Larval food plants

Larvae

polyphagous Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...

on a wide range of hosts including ''

Brachystegia ''Brachystegia'' is a genus of tree of the subfamily Detarioideae that is native to tropical Africa. Trees of the genus are commonly known as miombo, and are dominant in the miombo woodlands of central and southern tropical Africa. The Zambezia ...

'' , ''

Hibiscus ''Hibiscus'' is a genus of flowering plants in the Malva, mallow family, Malvaceae. The genus is quite large, comprising List of Hibiscus species, several hundred species that are Native plant, native to warm temperate, Subtropics, subtropical ...

'', '' Croton sp.'', ''

Afzelia quanzensis ''Afzelia quanzensis'' (pod mahogany, , , ) is a species of tree native to sub-Saharan Africa. It ranges from Somalia and Democratic Republic of the Congo to South Africa. It is a protected tree in South Africa. It belongs to the subfamily Caesal ...

'', '' Bauhinia galpinii'', ''

Burkea africana ''Burkea africana'', the wild syringa (), is a deciduous, medium-sized, spreading, flat-topped tree which grows in the woodlands and savannas of sub-Saharan Africa. It is the sole species in genus ''Burkea'', which belongs to the subfamily Caesalp ...

'', ''

Schotia brachypetala ''Schotia brachypetala'', the weeping boer-bean, is a leguminous flowering tree in the family Fabaceae (bean family/pod-bearing family/legumes) and the subfamily Detarioideae. The woodland tree is native to Africa south of the Zambezi River, ...

'', the red spike-thorn shrub and tree '' Gymnosporia senegalensis'', ''

Colophospermum mopane ''Colophospermum mopane'', commonly called mopane, mopani, butterfly tree, turpentine tree, or balsam tree, is a tree in the legume family (Fabaceae), that grows in hot, dry, low-lying areas, in elevation, in parts of Southern Africa. The tree ...

'',
Xanthocerces zambesiaca
', ''

Julbernardia globiflora ''Julbernardia globiflora'' is a tropical African tree widespread at moderate altitudes in Miombo woodland to the south and east of the equatorial forest region of the Congo Basin. Its common name is mnondo. It is ecologically important over wid ...

'',
Xeroderris stuhlmannii
', '' Guibourtia conjugata'', ''

Catha edulis Khat (''Catha edulis''), also known as Bushman's tea, especially in South Africa, is a flowering plant native to eastern and southeastern Africa. It has a history of cultivation originating in the Harari Region, Harar area (present day easter ...

''.

References

External links

*Images and public data o
''C. saturnus''
on Bold Systems 4 at

Consortium for the Barcode of Life The Consortium for the Barcode of Life (CBOL) was an international initiative dedicated to supporting the development of DNA barcoding as a global standard for species identification. CBOL's Secretariat Office is hosted by the National Museum of ...

. {{Taxonbar, from=Q15801208 saturnus IUCN Red List least concern species Butterflies described in 1865 Butterflies of Africa Taxa named by Arthur Gardiner Butler