Acraea (genus)


Acraea (genus)

Taxobox
name = "Acraea"



image_width = 240px
image_caption = The Tawny Coaster has been identified as the mysterious "Papilio terpsicore". Formerly known as "Acraea violae", it is now "A. terpsicore".
regnum = Animalia
phylum = Arthropoda
classis = Insecta
ordo = Lepidoptera
familia = Nymphalidae
subfamilia = Heliconiinae
tribus = Acraeini
genus = "Acraea"
genus_authority = Fabricius, 1807
type_species = "Papilio horta"
type_species_authority = Linnaeus, 1764
subdivision_ranks = Species
subdivision =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
See references in Haaramo (2007)]

"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. Most species assembled here are restricted to the African region, but some are found in India, Southeast Asia, and Australia.Silva-Brandão "et al." (2008)]

The foodplants of their 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.

ystematics 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 "Acrena" might actually be closer to other genera in the tribe Acraeini.Pitkin & Jenkins (2004), Silva-Brandão "et al." (2008). See also references in Haaramo (2007)]

With increasing availability of DNA sequence data, it is confirmed that "Acraea" as loosely defined does not constitute a monophyletic group. Even before the attepts to split up "Acraena" in earnest had begun, Jacob Hübner in 1819 suggested to separate species around "Acraena 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 "Acrena", 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 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 Acraena – the type species –, and hence which of the any further subdivisions will get to bear the name "Acraena", 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.

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.Silva-Brandão "et al." (2008); see also references in Haaramo (2007)]

pecies

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

"Acraea acrita" species group
* "Acraea acrita"
* "Acraea chaeribula"
* "Acraea eltringhamiana"
* "Acraea guluensis"
* "Acraea lualabae"
* "Acraea manca"
* "Acraea pudorina"
* "Acraea utangulensis""Acraea andromacha" species group (close to part of "A. serena" group?)
* "Acraea andromacha" "Acraea anemosa" species group
* "Acraea anemosa"
* "Acraea pseudolycia"
* "Acraea turna""Acraea aureola" species group
* "Acraea aureola"

"Acraea bonasia" species group (close to "A. oberthuri" and "A. rahira" groups? Paraphyletic?)
* "Acraea acerata"
* "Acraea alicia"
* "Acraea bonasia"
* "Acraea burgessi"
* "Acraea cabira"
* "Acraea eponina"
* "Acraea excelsior"
* "Acraea goetzei"
* "Acraea karschi"
* "Acraea jordani"
* "Acraea lumiri"
* "Acraea mirabilis"
* "Acraea miranda"
* "Acraea rangatana"
* "Acraea sotikensis"
* "Acraea uvui"
* "Acraea ventura"

"Acraea caecilia" species group (close to "A. cepheus" and "A. egina" groups?)
* "Acraea aglaonice"
* "Acraea asboloplintha"
* "Acraea atergatis"
* "Acraea axina"
* "Acraea braesia"
* "Acraea caecilia" (Type species of "Stephenia")
* "Acraea caldarena"
* "Acraea doubledayi"
* "Acraea ella"
* "Acraea equatorialis"
* "Acraea leucopyga"
* "Acraea lygus"
* "Acraea marnois"
* "Acraea natalica"
* "Acraea oncaea"
* "Acraea pseudegina"
* "Acraea pudorella"
* "Acraea rhodesiana"
* "Acraea stenobea"
* "Acraea sykesi"

"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 chambezi"
* "Acraea diogenes"
* "Acraea guillemei"
* "Acraea lofua"
* "Acraea mansya"
* "Acraea nohara"
* "Acraea onerata"
* "Acraea periphanes"
* "Acraea petraea"
* "Acraea punctellata"
* "Acraea rohlfsi"
* "Acraea violarum"

"Acraea circeis" species group (close to "A. masamba" and "A. pentapolis" groups?)
* "Acraea buschbecki"
* "Acraea circeis" (Type species of "Gnesia")
* "Acraea conradti"
* "Acraea grosvenori"
* "Acraea kuekenthali"
* "Acraea mairessei"
* "Acraea melanoxantha"
* "Acraea newtoni"
* "Acraea ntebiae"
* "Acraea oreas"
* "Acraea orina"
* "Acraea orinata"
* "Acraea parrhasia"
* "Acraea pelopeia"
* "Acraea peneleos"
* "Acraea penelope"
* "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" (Type species of "Rubraea")
* "Acraea medina"

"Acraea encedon" species group (close to "A. jodutta" and "A. pharsalus" groups?)
* "Acraea encedana"
* "Acraea encedon" – Common Acraea (Type species of "Hyalites")
* "Acraea neocoda"

"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"
* "Acraea insularis"
* "Acraea jodutta"
* "Acraea johnstoni"
* "Acraea lycoa" (Type species of "Planema")
* "Acraea masaris"
* "Acraea niobe"
* "Acraea safie"

"Acraea issoria" species group (a rather distinct lineage?)
* "Acraea issoria" – Yellow Coster (Type species of "Pareba") "Acraea masamba" species group (close to "A. circeis" and "A. pentapolis" groups?)
* "Acraea aubyni"
* "Acraea cinerea"
* "Acraea fornax"
* "Acraea igola"
* "Acraea masamba"
* "Acraea orestia"
* "Acraea quirinalis"
* "Acraea sambavae"
* "Acraea siliana"
* "Acraea simulata"
* "Acraea strattipocles""Acraea oberthuri" species group (close to "A. bonasia" group?)
* "Acraea althoffi"
* "Acraea bergeri"
* "Acraea oberthuri"Verify source|date=August 2008
* "Acraea pseudepaea""Acraea pentapolis" species group (close to "A. circeis" and "A. masamba" groups?)
* "Acraea pentapolis"
* "Acraea vesperalis" "Acraea pharsalus" species group (close to "A. encedon" and "A. jodhutta" groups?)
* "Acraea pharsalus"
* "Acraea vuilloti"

"Acraea rahira" species group (close to "A. bonasia" group?)
* "Acraea anacreon" (Type species of "Auracraea")
* "Acraea anacreontica"
* "Acraea bomba"
* "Acraea calida"
* "Acraea guichardi"
* "Acraea lusinga"
* "Acraea mirifica"
* "Acraea rahira"
* "Acraea speciosa"
* "Acraea wigginsi"
* "Acraea zitja"

"Acraea satis" species group
* "Acraea rabbaiae"
* "Acraea satis"
* "Acraea zonata"

"Acraea serena" species group (formerly "A. terpsicore" group, polyphyletic?)
* "Acraea admatha"
* "Acraea boopis"
* "Acraea burni"
* "Acraea camaena" (close to part of "A. andromacha" group?)
* "Acraea cerasa" (a rather distinct lineage?)
* "Acraea cuva"
* "Acraea dammii"
* "Acraea eltringhami"
* "Acraea eugenia"
* "Acraea hamata"
* "Acraea horta" (Type species of "Acraea")
* "Acraea hova"
* "Acraea humilis"
* "Acraea igati" (Type species of "Solenites")
* "Acraea insignis"
* "Acraea iturina" (close to "A. circeis" group?)
* "Acraea iturinoides"
* "Acraea kalinzu"
* "Acraea lia"
* "Acraea kraka"
* "Acraea leucographa"
* "Acraea machequena"
* "Acraea mahela"
* "Acraea matuapa"
* "Acraea neobule" (close to "A. zetes" group?)
* "Acraea obeira" (close to "A. andromacha" group?)
* "Acraea quirina" (close to "A. andromacha" group?)
* "Acraea ranavalona" (Type species of "Phanopeltis")
* "Acraea rileyi"
* "Acraea rogersi" (a rather distinct lineage?)
* "Acraea serena" (formerly "A. eponina", "A. terpsicore". Type species of "Telchinia".)

"Acraea terpsicore" species group (formerly "A. violae" group)
* "Acraea terpsicore" – Tawny Coaster (formerly "A. violae")

"Acraea zetes" species group (might include part of "A. serena" group; a very distinct lineage?)
* "Acraea acara"
* "Acraea barberi"
* "Acraea chilo"
* "Acraea hypoleuca"
* "Acraea magnifica"
* "Acraea trimeni"
* "Acraea zetes"

Species group undetermined
* "Acraea acutipennis"
* "Acraea adrasta"
* "Acraea aganice"
* "Acraea alalonga"
* "Acraea alcinoe"
* "Acraea alticola"
* "Acraea annonae"
* "Acraea atatis"
* "Acraea bellona"
* "Acraea bergeriana"
* "Acraea brainei"
* "Acraea comor"
* "Acraea consanguinea"
* "Acraea dondoensis"
* "Acraea elongata"
* "Acraea encoda"
* "Acraea endoscota" (close to "A. andromacha" group?)
* "Acraea epaea"
* "Acraea epiprotea"
* "Acraea excisa"
* "Acraea formosa"
* "Acraea hecqui"
* "Acraea indentata"
* "Acraea induna"
* "Acraea intermedia"
* "Acraea intermediodes"
* "Acraea kappa"
* "Acraea kia"
* "Acraea kinduana"
* "Acraea lapidorum"
* "Acraea leopoldina"
* "Acraea loranae"
* "Acraea lyci"
* "Acraea macaria"
* "Acraea macarista"
* "Acraea mima"
* "Acraea obliqua"
* "Acraea odzalae"
* "Acraea oscari"
* "Acraea overlaeti"
* "Acraea parei"
* "Acraea peetersi"
* "Acraea persanguinea"
* "Acraea pierrei"
* "Acraea polis" (close to "A. circeis", "A. masamba" and "A. pentapolis" groups?)
* "Acraea pseudatolmis"
* "Acraea pseuderytra"
* "Acraea pullula"
* "Acraea quadricolor"
* "Acraea rupicola"
* "Acraea scalivittata"
* "Acraea silia"
* "Acraea simulator"
* "Acraea tellus"
* "Acraea toruna"
* "Acraea turlini"
* "Acraea umbra"
* "Acraea vestalis"
* "Acraea zambesina"
* "Acraea zoumi"

Footnotes

References

* (2004): [http://www.nhm.ac.uk/jdsml/research-curation/projects/butmoth/index.dsml 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 – [http://www.funet.fi/pub/sci/bio/life/insecta/lepidoptera/ditrysia/papilionoidea/nymphalidae/heliconiinae/acraea/index.html "Acraea"] . Version of 2007-JUN-05. Retrieved 2008-AUG-15.
* (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 (HTML abstract)

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