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Acraea
	Adult male fiery acraea (A. acrita)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Nymphalidae
Tribe:	Acraeini
Genus:	Acraea; Fabricius, 1807
Type species
	Papilio horta; (Linnaeus, 1764)
Species
	Presently about 220, see text
Synonyms
	; Alacria Henning, 1992; Aphanopeltis Mabille, 1887; Auracraea Henning, 1993; Aurora Henning, 1992 (non Ragonot 1887: preoccupied); Gnesia Doubleday, 1848; Hyalites Doubleday, 1848; Pareba Doubleday, 1848; Phanopeltis Mabille, 1887; Planema Doubleday, 1848; Rubraea Henning, 1992; Solenites Mabille, 1887; Stephenia Henning, 1992; Telchinia Hübner, 1819; (but see text)

Acraea is a genus of brush-footed butterflies (family Nymphalidae) of the subfamily Heliconiinae. It seems to be highly paraphyletic and has long been used as a "wastebin taxon" to unite about 220 species of anatomically conservative Acraeini. Some phylogenetic studies show that the genus Acraea is monophyletic if Bematistes and Neotropical Actinote are included (see Pierre & Bernaud, 2009). Most species assembled here are restricted to the Afrotropical realm, but some are found in India, Southeast Asia, and Australia.^[2]

Biology

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 caterpillars are usually Urticaceae or, like in most Heliconiinae, Passifloraceae. Some feed on other plants, such as Fabaceae, "Flacourtiaceae", or Violaceae. Their preferred species contain cyanogenic glycosides, which make the larvae and adults poisonous to predators. The aposematic 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.^[2] Their flight is slow and flapping.

Systematics and taxonomy

That all these species were properly placed in Acraea has never been generally accepted. In 1807, Johan Christian Fabricius established the genus for the garden acraea, described as Papilio horta by Carl Linnaeus 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 Acraeini.^[3]

With increasing availability of DNA sequence data, it is confirmed that Acraea as loosely defined does not constitute a monophyletic group. Even before the attempts to split up Acraea in earnest had begun, Jacob Hübner in 1819 suggested to separate species around Acraea serena as Telchinia. This name has been applied to a generally African group whose members usually feed on Urticaceae, and they had already been noted to bear some uncanny resemblances to the American Actinote in anatomical 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.^[3]

But while several informal species groups 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—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, do not feed on Urticaceae, it may be that the separation of Telchinia is unwarranted and other proposed genera might be resurrected instead.^[1]

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 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 Indian 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's De uitlandsche Kapellen, had been used all the time.^[4] Another instance of the confusion rife in this genus is exemplified by Boisduval's Acraea manjaca.

These developments^[which?] come from two papers written by J. Pierre and D. Bernaud.^[5]^[6]

Species

Since the proposed phylogenetic sequence of the species groups is almost certainly incorrect for a large part, the groups are simply listed alphabetically.^[7]

J. Pierre & D. Bernaud have published a complete systematic and synonymic list.^[8]

^[4]

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)

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

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?)

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 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 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 – 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

^ ^a ^b See references in Haaramo (2007)
^ ^a ^b Silva-Brandão et al. (2008)
^ ^a ^b Pitkin & Jenkins (2004), Silva-Brandão et al. (2008). See also references in Haaramo (2007).
^ ^a ^b Silva-Brandão et al. (2008); see also references in Haaramo (2007)
^ "Acraea terpiscore" (PDF). Acraea.com. Retrieved 15 March 2022.
^ "Acraea serena" (PDF). Acraea.com. Retrieved 15 March 2022.
^ Woodhall (2005)
^ "CLASSIFICATION ET LISTE SYNONYMIQUE DES TAXONS DU GENRE ACRAEA" (PDF). Acraea.com. Retrieved 15 March 2022.

References

Pitkin, B. & Jenkins, P. (2004): Butterflies and Moths of the World: Generic Names and their Type-species. Version of 2004-NOV-05. Retrieved 2008-AUG-15.
Savela, Markku (2007): Markku Savela's Lepidoptera and some other life forms – Acraea. 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)
Silva-Brandão, Karina Lucas; Wahlberg, Niklas;, Francini, Ronaldo Bastos; Azeredo-Espin, Ana Maria L.; Brown, Keith S. Jr.; Paluch, Márlon; Lees, David C. & Freitas, André V.L. (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. doi:10.1016/j.ympev.2007.11.024 PDF fulltext
Woodhall (2005): Field Guide to Butterflies of South Africa. Cape Town: Struik Publishers.