''Acraea'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of brush-footed butterflies (

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Nymphalidae) of the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

Heliconiinae The Heliconiinae, commonly called heliconians or longwings, are a subfamily of the brush-footed butterflies (family (biology), family Nymphalidae). They can be divided into 45–50 genera and were sometimes treated as a separate family Heliconi ...

. It seems to be highly

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

and has long been used as a " wastebin taxon" to unite about 220

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

anatomical Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...

ly conservative

Acraeini The Acraeini are a tribe (biology), tribe of butterflies of the subfamily Heliconiinae in the family Nymphalidae. Genera The recognized genera are: Tree of Life * ''Abananote'' Potts, 1943 * ''Acraea (genus), Acraea'' Fabricius, 1807 – acr ...

. Some phylogenetic studies show that the genus ''Acraea'' is monophyletic if '' Bematistes'' and

Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...

'' Actinote'' are included (see Pierre & Bernaud, 2009). Most species assembled here are restricted to the

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 ...

, but some are found in

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...

, and

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

.Silva-Brandão et al. (2008)

Biology

Acraea zetes caterpillar to pupae to butterfly metamorphosis by Nick Hobgood

The eggs are laid in masses; the larvae are rather short, of almost equal thickness throughout, and possessing branched spines on each segment, young larvae group together on a protecting mass of silk; the pupa is slender, with a long abdomen, rather wide and angulated about the insertion of the wings, and suspended by the tail only. '' A. horta'', '' A. cabira'', and '' A. terpsicore'' illustrate typical life histories. The food plants of ''Acraea''

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...

s are usually

Urticaceae The Urticaceae are a family, the nettle family, of flowering plants. The family name comes from the genus ''Urtica''. The Urticaceae include a number of well-known and useful plants, including nettles in the genus ''Urtica'', ramie (''Boehmeria ...

or, like in most Heliconiinae, Passifloraceae. Some feed on other plants, such as

Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...

, "

Flacourtiaceae The Flacourtiaceae is a defunct family of flowering plants whose former members have been scattered to various families, mostly to the Achariaceae and Salicaceae. It was so vaguely defined that hardly anything seemed out of place there and it bec ...

", or

Violaceae Violaceae is a family of flowering plants established in 1802, consisting of about 1000 species in about 25 genera. It takes its name from the genus '' Viola'', the violets and pansies. Older classifications such as the Cronquist system plac ...

. Their preferred species contain

cyanogenic glycoside In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycosides. ...

s, which make the larvae and adults poisonous to

predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...

s. The

aposematic Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...

coloration of the adults announces this, and some species are mimicked by less noxious butterflies. At least some ''"Acraea"'' are able to produce the toxins themselves. Their flight is slow and flapping. Eltringham1912TransEntSocLondPlate6

Systematics and taxonomy

That all these species were properly placed in ''Acraea'' has never been generally accepted. In 1807,

Johan Christian Fabricius Johann Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is co ...

established the genus for the garden acraea, described as ''Papilio horta'' by

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

in 1764, and its relatives. By and by, an increasing number of species were placed here. As early as 1848, and again in 1887 and the early 1990s, it was attempted to divide the genus into groups of closest relatives, as it was suspected that some ''"Acraea"'' might actually be closer to other genera in the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

.Pitkin & Jenkins (2004), Silva-Brandão et al. (2008). See also references in Haaramo (2007). With increasing availability of

DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...

data, it is confirmed that ''Acraea'' as loosely defined does not constitute a

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

group. Even before the attempts to split up ''Acraea'' in earnest had begun,

Jacob Hübner Jacob Hübner (20 June 1761 – 13 September 1826, in Augsburg) was a German entomologist. He was the author of ''Sammlung Europäischer Schmetterlinge'' (1796–1805), a founding work of entomology. Scientific career Hübner was the author of '' ...

in 1819 suggested to separate species around '' Acraea serena'' as ''Telchinia''. This name has been applied to a generally

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

n group whose members usually feed on

, and they had already been noted to bear some uncanny resemblances to the American '' Actinote'' in

details. Indeed, they seem to be closer relatives of these than of the other butterflies placed in ''Acraea'', which usually feed on Passifloraceae and are at least in part quite close relatives of the African genus '' Bematistes''. Those closest to that genus might warrant separation as ''Rubraea'' or ''Stephenia''. But while several informal

species group In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

s have been established, it is not clear which of these are monophyletic and how to split the apparently still paraphyletic genus further. The placement of the garden acraea—the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

—and hence which of the any further subdivisions will get to bear the name ''Acraea'', remains unresolved. As it is traditionally included in the former ''A. terpsicore'' group (now ''A. serena'' group) and its caterpillars, while

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 ...

, do not feed on Urticaceae, it may be that the separation of ''Telchinia'' is unwarranted and other proposed genera might be resurrected instead. There was one major misidentification which still causes confusion today. ''Acraea terpsicore'', described as ''Papilio terpsicore'' by Linnaeus in 1758, was held to be the

senior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

of ''A. serena'', described by Fabricius as ''Papilio serena'' in 1775. Hence, the former name was commonly used for that African species. But as it turned out, Linnaeus had actually described an

n species—the well-known tawny coaster. Fabricius in 1793 believed it was new to science and described it again, as ''Papilio violae''. Consequently, it had been long known as ''A. violae''. It was also recognized that Fabricius' little-studied ''P. serena'' was none other than the orange acraea. For this, the name ''A. eponina'', from the ''Papilio eponina'' established in the 1780 issue of

Pieter Cramer Pieter Cramer (21 May 1721 (baptized) – 28 September 1776) was a wealthy Netherlands, Dutch merchant in linen and Spanish wool, remembered as an entomologist. Cramer was the director of the Zealand Society, a scientific society located in Flushin ...

's ''De uitlandsche Kapellen'', had been used all the time.Silva-Brandão et al. (2008); see also references in Haaramo (2007) Another instance of the confusion rife in this genus is exemplified by Boisduval's '' Acraea manjaca''. These developments come from two papers written by J. Pierre and D. Bernaud.

Species

Since the proposed

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 ...

sequence of the species groups is almost certainly incorrect for a large part, the groups are simply listed alphabetically. J. Pierre & D. Bernaud have published a complete systematic and synonymic list. Acraea andromacha 2

''Acraea acrita'' species group * '' Acraea acrita'' – fiery acraea * '' Acraea annonae'' * '' Acraea bellona'' * '' Acraea chaeribula'' * '' Acraea eltringhamiana'' * '' Acraea guluensis'' * '' Acraea loranae'' * '' Acraea lualabae'' * '' Acraea manca'' * '' Acraea pudorina'' – Kenyan fiery acraea * '' Acraea utengulensis'' – Tanzanian fiery acraea ''Acraea andromacha'' species group (close to part of ''A. terpsicore'' group?) * '' Acraea andromacha'' – small greasy, glasswing ''Acraea anemosa'' species group * '' Acraea anemosa'' – broad-bordered acraea * '' Acraea pseudolycia'' * '' Acraea turna'' ''Acraea aureola'' species group * '' Acraea aureola'' (= onerata) (cepheus group) Yellow-banded Acraea 19 07 2010

''Acraea bonasia'' species group (close to ''A. oberthuri'' and ''A. rahira'' groups?

Paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

?) * '' Acraea acerata'' – small yellow-banded legionnaire, falls acraea * '' Acraea alicia'' * '' Acraea bonasia'' * '' Acraea burgessi'' * '' Acraea cabira'' – yellow-banded acraea * '' Acraea serena'' – dancing acraea * '' Acraea eponina'' – orange acraea or small orange acraea * '' Acraea excelsior'' * '' Acraea goetzei'' * '' Acraea karschi'' * '' Acraea lumiri'' * '' Acraea rangatana'' * '' Acraea hecqui'' * '' Acraea pierrei'' * '' Acraea sotikensis'' – Sotik acraea, Sotika legionnaire * '' Acraea uvui'' – tiny acraea * '' Acraea ventura'' ''Acraea caecilia'' species group (close to ''A. cepheus'' and ''A. egina'' groups?) Natal legionnaires (''A. natalica'') * '' Acraea aglaonice'' – clear-spotted acraea * '' Acraea mirabilis'' – marvelous acraea * '' Acraea miranda'' – desert acraea, Somali acraea * '' Acraea asboloplintha'' – black-winged acraea, black-winged legionnaire * '' Acraea atergatis'' * '' Acraea atatis'' * '' Acraea axina'' – little acraea * '' Acraea braesia'' * '' Acraea caecilia'' – pink legionnaire (type species of ''Stephenia'') * '' Acraea caldarena'' – black-tipped acraea, black tip acraea * '' Acraea doubledayi'' * '' Acraea ella'' * '' Acraea equatorialis'' * '' Acraea intermediodes'' * '' Acraea leucopyga'' * '' Acraea lygus'' – lygus acraea * '' Acraea marnois'' (= '' Acraea caecilia'') * '' Acraea natalica'' – Natal acraea, Natal legionnaire * '' Acraea oncaea'' – window acraea, window legionnaire * '' Acraea pseudegina'' * '' Acraea pudorella'' * '' Acraea rhodesiana'' * '' Acraea stenobea'' – suffused acraea * '' Acraea lyci'' * '' Acraea sykesi'' – Sykes' acraea ''Acraea cepheus'' species group (close to ''A. caecilia'' and ''A. egina'' groups?) * '' Acraea abdera'' * '' Acraea asema'' * '' Acraea atolmis'' * '' Acraea bailundensis'' * '' Acraea buettneri'' * '' Acraea cepheus'' * '' Acraea bergeriana'' * '' Acraea chambezi'' * '' Acraea diogenes'' * '' Acraea guillemei'' * '' Acraea lapidorum'' * '' Acraea lofua'' * '' Acraea mansya'' * '' Acraea nohara'' – light red acraea * '' Acraea dondoensis'' * '' Acraea onerata'' * '' Acraea periphanes'' * '' Acraea petraea'' – blood-red acraea, blood acraea * '' Acraea punctellata'' * '' Acraea rohlfsi'' * '' Acraea violarum'' – speckled red acraea ''Acraea circeis'' species group (close to ''A. masamba'') * '' Acraea buschbecki'' * '' Acraea circeis'' – white legionnaire (type species of ''Gnesia'') * '' Acraea conradti'' * '' Acraea grosvenori'' * '' Acraea kuekenthali'' * '' Acraea melanoxantha'' * '' Acraea newtoni'' * '' Acraea ntebiae'' * '' Acraea oreas'' * '' Acraea orina'' – Orina legionnaire * '' Acraea orinata'' * '' Acraea parrhasia'' – yellow-veined legionnaire * '' Acraea pelopeia'' * '' Acraea peneleos'' * '' Acraea penelope'' – Penelope's acraea, Penelope's legionnaire * '' Acraea perenna'' * '' Acraea semivitrea'' (type species of ''Alacria'') * '' Acraea translucida'' * '' Acraea ungemachi'' * '' Acraea vumbui'' Acraea_encedon_HotMarina_2012-01-21

''Acraea egina'' species group (close to ''A. caecilia'' and ''A. cepheus'' groups?) * '' Acraea egina'' – elegant legionnaire (type species of ''Rubraea'') * '' Acraea medea'' ''Acraea encedon'' species group (close to ''A. jodutta'' and ''A. pharsalus'' groups?) – common acraea or white-barred acraea or encedon acraea * '' Acraea encedana'' – Pierre's acraea * '' Acraea encedon'' – common acraea, white-barred legionnaire (type species of ''Hyalites'') * '' Acraea necoda'' * '' Acraea encoda'' ''Acraea issoria'' species group (a rather distinct lineage?) * '' Acraea issoria'' – yellow coster (type species of ''Pareba'') ''Acraea jodutta'' species group (close to ''A. encedon'' and ''A. pharsalus'' groups?) * '' Acraea actinotina'' * '' Acraea acuta'' * '' Acraea alciope'' * '' Acraea alciopoides'' * '' Acraea amicitiae'' * '' Acraea ansorgei'' * '' Acraea aurivillii'' * '' Acraea baxteri'' * '' Acraea disjuncta'' * '' Acraea esebria'' – dusky acraea * '' Acraea comor'' * '' Acraea insularis'' * '' Acraea jodutta'' – jodutta legionnaire * '' Acraea johnstoni'' * '' Acraea lycoa'' – lycoa legionnaire (type species of ''Planema'') * '' Acraea masaris'' * '' Acraea niobe'' * '' Acraea safie''

''Acraea masamba'' species group (close to ''A. circeis''?) * '' Acraea aubyni'' – Aubyn Rogers' acraea * '' Acraea cinerea'' – grey acraea * '' Acraea fornax'' * '' Acraea igola'' – dusky-veined acraea * '' Acraea masamba'' * '' Acraea quirinalis'' * '' Acraea sambavae'' * '' Acraea siliana'' * '' Acraea simulata'' (tentatively placed here; '' Bematistes''?) * '' Acraea strattipocles'' ''Acraea oberthuri'' species group (close to ''A. bonasia'' group?) * '' Acraea althoffi'' – Althoff's acraea * '' Acraea bergeri'' * '' Acraea oberthueri'' * '' Acraea pseudepaea'' ''Acraea pentapolis'' species group (close to ''A. circeis'' and ''A. masamba'' groups?) * '' Acraea pentapolis'' – musanga legionnaire * '' Acraea vesperalis'' * '' Acraea obeira'' * '' Acraea burni'' – pale-yellow acraea (close to ''A. andromacha'' group?) * '' Acraea orestia'' * '' Acraea lia'' ''Acraea pharsalus'' species group (close to ''A. encedon'' and ''A. jodhutta'' groups?) * '' Acraea pharsalus'' – familiar legionnaire * '' Acraea vuilloti'' ''Acraea rahira'' species group (close to ''A. bonasia'' group?) * '' Acraea anacreon'' – large orange acraea, orange acraea (type species of ''Auracraea'') * '' Acraea anacreontica'' * '' Acraea bomba'' * '' Acraea calida'' * '' Acraea guichardi'' * '' Acraea lusinga'' * '' Acraea induna'' – induna acraea (included in ''"Telchinia"'') * '' Acraea kaduna'' * '' Acraea parei'' * '' Acraea alalonga'' – long-winged orange acraea (included in ''"Telchinia"'') * '' Acraea mirifica'' * '' Acraea rahira'' – marsh acraea * '' Acraea speciosa'' * '' Acraea wigginsi'' * '' Acraea zitja'' ''Acraea satis'' species group * '' Acraea rabbaiae'' – clear-wing acraea, clearwing legionnaire * '' Acraea satis'' – east coast acraea * '' Acraea zonata'' ''A. terpsicore'' group * '' Acraea admatha'' – Hewitson's glassy legionnaire * '' Acraea boopis'' – rainforest acraea * '' Acraea endoscota'' (close to ''A. andromacha'' group?) * '' Acraea kappa'' * '' Acraea kia'' * '' Acraea kinduana'' * '' Acraea camaena'' (close to part of ''A. andromacha'' group?) * '' Acraea cuva'' * '' Acraea dammii'' * '' Acraea eltringhami'' * '' Acraea eugenia'' * '' Acraea hamata'' * ''

Acraea horta ''Acraea horta'' or the garden acraea is a butterfly of the family Nymphalidae. It is found in South Africa and Zimbabwe. Description ''A. horta'' L. (53 e, f) is one of the longest known and commonest Acraeids of the Cape, but it is also nati ...

'' – garden acraea (type species of ''Acraea'') * '' Acraea hova'' * '' Acraea igati'' (type species of ''Solenites'') * '' Acraea insignis'' * '' Acraea leucographa'' * '' Acraea machequena'' – machequena acraea * '' Acraea mahela'' * '' Acraea matuapa'' * '' Acraea neobule'' – wandering donkey acraea, wandering donkey (close to ''A. zetes'' group?) * '' Acraea brainei'' * '' Acraea quirina'' – common glassy legionnaire (close to ''A. andromacha'' group?) * '' Acraea ranavalona'' (type species of ''Phanopeltis'') * '' Acraea rileyi'' * '' Acraea rogersi'' – Rogers's large legionnaire (a rather distinct lineage?) * '' Acraea serena'' – dancing acraea, small orange legionnaire (formerly misidentified as ''A. eponina'', ''A. terpsicore''; type species of ''Telchinia'') ''Acraea terpsicore'' species group (formerly ''A. violae'' group) * '' Acraea terpsicore'' – tawny coster (formerly ''A. violae'') ''Acraea zetes'' species group (might include part of ''A. serena'' group; a very distinct lineage?) * '' Acraea acara'' – acara acraea * '' Acraea barberi'' – Barber's acraea * '' Acraea chilo'' * '' Acraea hypoleuca'' * '' Acraea magnifica'' * '' Acraea trimeni'' – Trimen's acraea * '' Acraea zetes'' Species group undetermined * '' Acraea acutipennis'' * '' Acraea alticola'' * '' Acraea cerasa'' – tree top acraea * '' Acraea ducarmei'' ("''epaea'' group") * '' Acraea humilis'' * '' Acraea kalinzu'' * '' Acraea kraka'' – kraka glassy acraea * '' Acraea macarista'' * '' Acraea mima'' * '' Acraea odzalae'' * '' Acraea omrora'' * '' Acraea oscari'' * '' Acraea overlaeti'' * '' Acraea peetersi'' * '' Acraea polis'' (close to ''A. circeis'', ''A. masamba'' and ''A. pentapolis'' groups?) * '' Acraea pseudatolmis'' * '' Acraea pullula'' * '' Acraea punctimarginea'' * '' Acraea silia'' * '' Acraea simulator'' * '' Acraea iturina'' – Ituri glassy acraea * '' Acraea toruna'' * '' Acraea turlini'' * '' Acraea viviana'' * '' Acraea zoumi'' Some other species formerly in ''Acraea'' have now been definitely assigned to other genera, e.g. '' Bematistes''.

Footnotes

References

* (2004)
Butterflies and Moths of the World: Generic Names and their Type-species
Version of 2004-NOV-05. Retrieved 2008-AUG-15. * (2007): Markku Savela's Lepidoptera and some other life forms �

Version of 2007-JUN-05. Retrieved 2008-AUG-15.
Seitz, A. ''Die Gross-Schmetterlinge der Erde'' 13: Die Afrikanischen Tagfalter. Plate XIII 53 et seq.

Seitz, A. ''Die Gross-Schmetterlinge der Erde'' 13: Die Afrikanischen Tagfalter. Text (in German)
* (2008): Phylogenetic relationships of butterflies of the tribe Acraeini (Lepidoptera, Nymphalidae, Heliconiinae) and the evolution of host plant use. '' Mol. Phylogenet. Evol.'' 46(2): 515–531. PDF fulltext
* (2005): ''Field Guide to Butterflies of South Africa''. Cape Town: Struik Publishers.

External links

Le Site des Acraea de Dominique Bernaud

Systematic and synonymic list (Pierre & Bernaud)

Alphabetic list (Pierre & Bernaud)

African description references (Pierre & Bernaud)

''Acraea terpsicore'' rehabilitation paper

''Acraea serena'' rehabilitation paper

''Acraea epaea'' group (ex ''Bematistes'') revision paper

''Acraea'' at EOL

African Butterfly Database
especially bibliography

image overview {{Taxonbar, from=Q420732 Acraeini Nymphalidae genera Taxa named by Johan Christian Fabricius