Charaxes

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Charaxes
Charaxes solon at MNP.jpg
C. solon at Mahim Nature Park, Mumbai
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Nymphalidae
Subfamily: Charaxinae
Tribe: Charaxini
Genus: Charaxes
Ochsenheimer, 1816
Species

Many, see text

Synonyms
  • PaphiaFabricius, 1807
  • EriboeaHübner, [1819]
  • JasiaSwainson, 1832
  • MonuraMabille, 1877
  • HaridraMoore, [1880]
  • ZinghaHemming, 1939
  • HadrodontesStoneham, 1964
  • StonehamiaCowan, 1968

The rajah and pasha butterflies, also known as emperors in Africa and Australia, [1] [2] (genus Charaxes) make up the huge type genus of the brush-footed butterfly subfamily Charaxinae, or leafwing butterflies. They belong to the tribe Charaxini, which also includes the nawab butterflies ( Polyura ). Charaxes are tropical Old World butterflies, with by far the highest diversity in sub-Saharan Africa, a smaller number from South Asia to Melanesia and Australia, and a single species ( C. jasius ) in Europe. They are generally strong flyers and very popular among butterfly collectors.

Contents

Etymology

Charaxes means "to sharpen" or "to make pointed", referring to the pointed 'tails' on the hind wing. Charaxes may also be related to charax, meaning 'a sharp stake', or charaxis, a 'notch' or 'incision', which are also features of the hind wing. [3]

Biology

Charaxes frequent sunny forest openings and glades where they rest with open or partly open wings sunning themselves. When alerted, they close the wings exposing the cryptic underside. Certain favoured perches are selected and intruders are chased and driven off, Charaxes feed in part at oozes from tree wounds infested with beetle or other larvae and on rotting fruit (they come to hanging traps baited with fermenting banana). They lay their eggs on small understorey or marginal trees. Tender shoots are usually selected for egg laying but as the larvae grow they move to older leaves. Larvae rest on the upper surface of a leaf on which they have spun a protecting silk pad.

Photograph from Schultze, 1917 Die Charaxiden und Apaturiden der Kolonie Kamerun, illustrating aspects of Charaxes biology SchultzeKameroonCharaxesHabitatPlate.jpg
Photograph from Schultze, 1917 Die Charaxiden und Apaturiden der Kolonie Kamerun, illustrating aspects of Charaxes biology

The most striking features in the habits of Charaxes are the powerful rapid flight, the partiality to putrid matter and the constancy with which a specimen returns to the same spot. Few species are found in the open country ( C. pelias pelias , C. jasius and C. fabius ), where there are only bushes and rarely trees; most species inhabit the more wooded country and some are found only in and near larger forests. The males come often in some numbers to water pools on roads where they mud puddle; both sexes are fond of the juice of trees, decaying fruits, dung of animals and putrid meat and can successfully be entrapped by the use of such baits; one is known to come to flowers ( C. zoolina ). Some species locate mates by hill-topping.

The larvae feed variously on Rhamnaceae, Leguminosae, Sapindaceae, Melianthaceae, Euphorbiaceae, Gramineae, Ochnaceae, Lauraceae, Tiliaceae, Meliaceae, etc. [4]

Charaxes acraeoides is part of a mimetic ring with Pseudacraea and Acraea . The closely related Charaxes fournierae mimics species of Euphaedra .

Life history

"We have examined the eggs of a number of species, taken from the bodies of dry specimens. Apart from size, there is no apparent difference in the eggs of the various species. The micropyle end is flattened, slightly concave, with the centre raised; from the centre radiate slight ribs, which gradually disappear at the sides and do not reach the base or underside; the longitudinal ribs are connected by extremely feeble transverse ridges, and bear, as Doherty correctly points out, small projecting points, which are easily visible under a good lens.

The slug-shaped caterpillar is widest in or before the middle, narrowed behind, and has a finely granulated skin. The head is flat, prognathous, and bears four processes which point backwards and are rough with tubercles, as is the hinder edge of the head between the processes; cheeks also tuberculated; the anal segment bears dorsally two more or less prominent processes, which are longer in the young larva than in the full-grown one. The colour of the larvae is generally green, often yellowish, the head bears, on each side, a light line which runs along the outer horn, and there is a spot on one or more abdominal segments, the colour of these latter markings, which are mostly more or less halfmoon-shaped, is as a rule buffish, the spots having often a darker (reddish or bluish) border. The caterpillar is a very slow creature, which does not voluntarily leave the twig on a leaf of which the egg was deposited.

The thick chrysalis is bright green as a rule, very smooth and shining, dorsally very convex; head bluntly bipartite; end of abdomen with two rounded tubercles ventrally." [5] :282

Description of the imago

Adult Charaxes have a robust thorax and abdomen. Their wingspan is around 8 to 10 cm. The ground colours of the wing uppersides vary from tawny to black to pale. Wing markings may be spots, bars or bands of white, orange or blue. Some species have metallic blue spots and others have an overall iridescent, metallic gloss. Charaxes zingha is partially red and Charaxes eupale is light green. The males of the Charaxes etheocles complex are largely black and are known as the "black Charaxes".

Various colours and shapes of Charaxes
NovaraExpZoologischeTheilLepidopteraAtlasTaf59.jpg
Lepidoptera2Butler.jpg
Illustrations of new species of exotic butterflies Charaxes V.jpg
NovitatesZoologicae4Plate5.jpg
Illustrations of new species of exotic butterflies Nymphalis I.jpg

The outer margin of the forewing varies from being nearly straight to deeply concave. The hindwing is abdominally always longer than costally, and in most species presents a triangular shape. The teeth on the distal (or outer) margin of both wings vary much according to species and groups of species, and is also not entirely constant within a species. The anal angle (wing corner) is in many species more pronounced than the outer margin between the tails, best seen in Charaxes zingha .

The pattern of the upperside is often very different in the sexes and in the various species. The females can be arranged according to the pattern of the upperside of the forewing into four groups:

The underside is sometimes very colourful with orange, grey and auburn silver areas.

Sexual differences

Sexual dimorphism is among Charaxes a much commoner phenomenon than similarity of the sexes in colour and shape. The wings of the female are always broader than those of the male, and the hindwing is, as a rule, less triangular; the tails are broader, often widened at the end; in some species the male has one, the female two tails, in other the tails of the male are obliterated (absent), while the female possesses such an appendage.

Taxonomy

Charaxes wing
Charaxesvenation2.jpg
Venation terminology
Charaxesvenation.jpg
Wing mark terminology

The taxonomy is complex with very many ranks (subspecies, forms and variants) and Charaxes demonstrates allopatric , sympatric speciation, parallel evolution and complex mimicry. There have been many rank and placement changes. Splitters, possibly with commercial as well as entomological interests, have named very many forms – an example is Georges Rousseau-Decelle. Many species show geographic and climatic (including altitudinal) clines. An additional problem is the publication of new taxa in journals which are not peer reviewed.

"No group of African butterflies arouses stronger emotions than Charaxes. Gaining an understanding of their phylogenetic relationships will add a new chapter to their convoluted literature". [6]

Significant monographs on the taxonomy of Charaxes include:

Van Someren studied long series of museum specimens, drawing attention, as had his predecessors, to the variability of Charaxes species over their often vast range, identifying and describing subspecies and isolating new species. This clinal variation is complicated by variation associated with climate (season), altitude and mimicry. Van Someren recognizes species groups, the analysis is in part subjective and the number of taxa remains uncertain.

The type species of the genus Charaxes is Charaxes jasius . [7]

Species groups

Cladogram of the basal subdivision of the genus Charaxes and its nearest relatives Charaxes basal cladogram.jpg
Cladogram of the basal subdivision of the genus Charaxes and its nearest relatives

Defining species groups is a convenient way of subdividing well-defined genera with a large number of recognized species. Charaxes species are so arranged in assemblages called "species groups" (not superspecies, but an informal phenetic arrangement). These may or may not be clades. As molecular phylogenetic studies continue, lineages distinct enough to warrant some formal degree of recognition become evident and new groupings are suggested, but consistent ranking remains a problem.

Species

In older literature, some of the nawab butterflies, such as Polyura dolon , may be included under Charaxes.

Species not found in the Afrotropical realm

Species found in the Afrotropical realm

Listed alphabetically. [8] [9] Recently updated taxonomy is referenced within the list.

Related Research Articles

<i>Charaxes jasius</i> Species of butterfly

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 it shared with its closest related species still flying in the Afrotropical realm most probably occurred around 2 mya, i.e. during the Pliocene.

<i>Charaxes brutus</i> Species of butterfly

Charaxes brutus, the white-barred emperor or white-barred Charaxes, is a butterfly of the family Nymphalidae. It is found in Africa.

<i>Charaxes pelias</i> Species of butterfly

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.

<i>Charaxes castor</i> Species of butterfly

Charaxes castor, the giant emperor or giant charaxes, is a butterfly of the family Nymphalidae. It is found throughout the Afrotropical realm below the Sahel.

<i>Charaxes druceanus</i> Species of butterfly

Charaxes druceanus, the silver-barred emperor or silver-barred charaxes, is a butterfly of the family Nymphalidae. It is found throughout tropical Africa.

<i>Charaxes smaragdalis</i> Species of butterfly

Charaxes smaragdalis, the western blue charaxes, is a butterfly of the family Nymphalidae. It is found from Senegal to Somalia, from Angola to Kenya and from Sudan to Egypt.

Charaxes schiltzei is a butterfly in the family Nymphalidae. It is found in the Democratic Republic of the Congo, Uganda, Rwanda and Burundi.

<i>Charaxes hansali</i> Species of butterfly

Charaxes hansali, the cream-banded charaxes, is a butterfly in the family Nymphalidae. It is found in Tanzania, Rwanda, Kenya, Uganda, Sudan, Ethiopia, Somalia, Saudi Arabia, Yemen and Oman.

Charaxes junius is a butterfly in the family Nymphalidae. It is found in Ethiopia and Sudan. The habitat consists of forests and woodland savanna.

<i>Charaxes ansorgei</i> Species of butterfly

Charaxes ansorgei is a butterfly in the family Nymphalidae. It is found in Sudan, the Democratic Republic of Congo, Uganda, Rwanda, Burundi, Kenya, Tanzania, Malawi and Zambia. The habitat consists of montane forest on altitudes between 2,000 and 3,000 meters. The larvae feed on Bersama abyssinica abyssinica, Bersama abyssinica englerana and Bersama paullinoides.It was once considered to be a very rare species but it is relatively common in dense and inaccessible montane forests

<i>Charaxes pollux</i> Species of butterfly

Charaxes pollux, the black-bordered charaxes, is a butterfly in the family Nymphalidae.

<i>Charaxes phraortes</i> Species of butterfly

Charaxes phraortes is a butterfly in the family Nymphalidae. It is found in north-eastern Madagascar. The habitat consists of wet coastal forests.

<i>Charaxes eudoxus</i> Species of butterfly

Charaxes eudoxus, the Eudoxus charaxes, is a butterfly in the family Nymphalidae. It is found in Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea, Gabon, the Republic of the Congo, Angola, the Democratic Republic of the Congo, the Central African Republic, Sudan, Uganda, Rwanda, Burundi, Kenya, Tanzania, Cameroon and Zambia.

Charaxes lucyae is a butterfly in the family Nymphalidae. It is found in Tanzania.

Charaxes richelmanni is a butterfly in the family Nymphalidae. It is found in Cameroon, Gabon, the Republic of Congo, the Central African Republic, the Democratic Republic of Congo and Tanzania. The habitat consists of lowland evergreen forests.

<i>Charaxes subornatus</i> Species of butterfly

Charaxes subornatus, the ornate green charaxes, is a butterfly in the family Nymphalidae. It is found in Ivory Coast, Ghana, Togo, Nigeria, Cameroon, the Central African Republic, the Republic of the Congo, Gabon, the Democratic Republic of the Congo, Uganda and Kenya. The habitat consists of evergreen forests.

<i>Charaxes dilutus</i> Species of butterfly

Charaxes dilutus is a butterfly in the family Nymphalidae. It is found in Uganda, Tanzania, Malawi, the Democratic Republic of the Congo, Zambia and Angola. The habitat consists of tropical evergreen forests.

<i>Charaxes montis</i> Species of butterfly

Charaxes montis is a butterfly in the family Nymphalidae. It is found in the Democratic Republic of the Congo, south-western Uganda, Rwanda and Burundi. The habitat consists of montane forests.

<i>Charaxes epijasius</i> Species of butterfly

Charaxes epijasius, the cream-bordered charaxes or Sahel charaxes, is a butterfly in the family Nymphalidae. It flies through most of the Savannah of the Afrotropical realm except southern Africa.

<i>Charaxes saturnus</i>

Charaxes saturnus, the foxy charaxes or koppie charaxes, is a butterfly that flies through most of the Savannah of the Eastern and Southern Afrotropical realm, and also occurs in suitable forest habitat locations including the forest belt of west-central Africa.

References

  1. Woodhall, Steve (2005). Field Guide to Butterflies of South Africa. Cape Town, South Africa: Struik. ISBN   978-1-86872-724-7.
  2. Braby, Michael F. (2004). Complete Field Guide to Butterflies of Australia. CSIRO Publishing. pp. 174–175. ISBN   0643090274.
  3. Otto, Herbert. "Butterflies of the KNP". krugerpark.co.za. Retrieved 19 January 2019.
  4. "Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni & L. M. Hernández (2010). HOSTS - A Database of the World's Lepidopteran Hostplants. Natural History Museum, London".
  5. 1 2 Rothschild, W.; Jordan, K. (1900). "A monograph of Charaxes and the allied prionopterous genera". Novitates Zoologicae . 7: 281–524.
  6. Ackery P. R., Smith C. R., and Vane-Wright R. I. (eds.) (1995). Carcasson's African butterflies-An Annotated Catalogue of the Papilionidea and Hesperioidea of the Afrotropical Region. Canberra: CSIRO. A collated edited and amplified version of Carcasson's manuscripts. Included are species, genus group, species group and infra-subspecific names of Afrotropical butterflies and text on host plants, general reviews of bionomics, biology, biogeography and bibliography.
  7. Rydon, AHB (1971). "The Systematics of the Charaxidae (Lep. : Nymphaloidea)". The Entomologist's Record and Journal of Variation. London. 83: 219–233. Retrieved 15 March 2018. Pg. 220 "Recognising this fact, Ochsenheimer (1816, Schmett. Europa, vol. 4) removed Papilio jasius from Paphia, and placed it instead in his own genus Charaxes, thus making P. jasius the type-species of the latter by monotypy."
  8. "Charaxes Ochsenheimer, 1816" at Markku Savela's Lepidoptera and Some Other Life Forms
  9. "Genus Charaxes (2019 Rev.)" Ed. M.C.Willams (2019) at Metamorphosis.org.za
  10. [Sp. nov.] Vingerhoedt, E. & Faravel, G. 2009. Lambillionea, 2009:109;389-94
  11. [Sp. nov.] Basquin, P. Entomologia Africana, 2012;2:51-54
  12. [Sp. nov.] Faravel, G. & Bouyer, Th. Entomologia Africana 2022; 27: 17 – 27
  13. [Sp. nov.] Collins, Congdon & Bampton. Entomologia Africana, 2017; 22: 19 – 30
  14. [Sp. Nov.] Vingerhoedt & Vande Weghe. Lépodoptères - Revue des Lépidoptères de France, 2011; 48: 2-6
  15. Bouyer, Zakharov, Rougerie, Vingerhoedt (2008); Les Charaxes du "groupe eupale:description d'un nouveau genre, revision et approche genetique (Lepidoptera, Nymphalidae, Charaxinae). Entomologia Africana, Hors Serie no.3, 2008: 1-32. p5. Comment: Stat.Rev. from [schultzei syn=latimargo] to schultzei bona species

Further reading