Felicia is a genus of small shrubs, perennial or annual herbaceous plants, with 85 known species, that is assigned to the daisy family (Compositae or Asteraceae). Like in almost all Asteraceae, the individual flowers are 5-merous, small, and clustered in typical heads, which are surrounded by an involucre of between two and four whorls of bracts. In Felicia, the centre of the head is taken by yellow, seldom whitish or blackish blue disc florets, and is almost always surrounded by one single whorl of mostly purple, sometimes blue, pink, white, or yellow ligulate florets, and rarely, ligulate florets are absent. These florets sit on a common base (or receptacle) and are not individually subtended by a bract (or palea). Most species occur in the Cape Floristic Region, which is probably the area where the genus originates and had most of its development. Some species can be found in the eastern half of Africa, up to Sudan and the south-western Arabian Peninsula, while on the west coast, species can be found from the Cape to Angola, with one species having outposts on the Cameroon-Nigeria border and central Nigeria. Some species of Felicia are cultivated as ornamentals, and several hybrids have been developed for that purpose.
Taxonomy Aster tenellus, discovered by the early English botanist Leonard Plukenet in 1692, was the first species recorded that is now included in the genus Felicia (as F. tenella). This was followed in 1700 by Aster fruticosus, again by Plukenet, repeated by Caspar Commelin in 1701, a species now called Felicia fruticosa. Such names published before 1753, the year that was chosen as a starting point for the binomial nomenclature proposed by Carl Linnaeus, are not valid. In 1761, John Hill suggested the genus Coelestina, widely considered a later synonym of Ageratum, but the plant that was illustrated is most likely the same species as F. amelloides. In 1763, Carl Linnaeus described Cineraria amelloides (F. amelloides), the first valid name for a species assigned today to Felicia. In 1763, French botanist Michel Adanson described a new genus, Detris, without mentioning a species, but apparently having F. amelloides in mind. Peter Jonas Bergius, a botanist from Sweden, described Aster hyssopifolius in 1767, now known as F. hyssopifolia. A year later, Dutch botanist Pieter Burman the Younger added Aster aethiopicus (now F. aethiopica). This was followed by Aster cymbalariae (William Aiton, 1789), Leysseria ovata and Pteronia echinata (Carl Peter Thunberg, 1800 and 1823), Aster filifolius (Étienne Pierre Ventenat, 1804), and Cineraria bergeriana (Kurt Polycarp Joachim Sprengel, 1826), now F. cymbalariae, F. ovata, F. echinata, F. filifolius and F. bergeriana.
In 1815, French Asteraceae specialist Henri Cassini begun his work to rearrange the genera of the Asteraceae, splitting off species that had been assigned to Aster up till that moment but are now accepted in Felicia, starting with Agatacha. This name was probably a typographical error, since he replaced it with Agathaea in the following year. In 1817 he created Charieis, and followed up with Felicia in 1818. He created Munychia in 1825 to accommodate Aster cymbalariae. Of the species he assigned to his new genus, Diplopappus (1819), none are currently included in Felicia. At that moment, 17 species currently included in Felicia were known to science.
Because of the extensive collecting in the Cape Region by Drège, Ecklon and Zeyher, during the next decade, the number of known species expanded rapidly. In 1832, Christian Friedrich Lessing moved most of the species that Cassini distributed over his new genera back to Aster, although he assigned Cineraria bergerana to Elphegea, and created the new combinations Diplopappus fruticosa and D. filifolia. A year later, Nees von Esenbeck considered the assignment of current Felicia species. He thought Charieis was synonymous to the older genus Kaulfussia, agreed mostly with the split created by Cassini, includes the species of the section Lignofelicia in Diplostephium (Kunth, 1820), and created the genus Detridium for Cineraria bergerana. Augustin Pyramus de Candolle in 1836 recognised Agathaea, Diplopappus, Felicia and Munychia, altogether 49 species, most of which are assigned to Felicia today. Consecutive authors can roughly be divided into lumpers, who brought together many species in Aster, and splitters, who proposed narrow taxa; both groups eventually converged on Felicia in its current circumscription. Jürke Grau revised the genus in 1973 and described seventeen new species (F. alba, F. caespitosa, F. canaliculata, F. clavipilosa, F. comptonii, F. joubertinae, F. merxmuellerii, F. microcephala, F. monocephala, F. nigrescens, F. nordenstamii, F. oleosa, F. stenophylla and F. tsitsikamae), alongside eight new subspecies. Only four species have been described since: F. flava by Henk Jaap Beentje (1999), F. martinsiana (2007) by Santiago Ortiz, F. josephinae (2002) by John Manning and Peter Goldblatt, and F. douglasii in 2018 by Manning, Anthony Maggee and James Boatwright. Currently, 85 species have been assigned to Felicia.
Although Felicia has been commonly used for a long time, it is not the oldest name. In principle, Detris, as the earliest name, should have priority, but it was suppressed in favour of the widely used name Felicia. In 2010, it was proposed to also conserve Felicia over Agathaea, Charieis and Coelestina, because these names also precede Felicia and should get priority otherwise.
Naming The genus is presumed to be named in honor of Fortunato Bartolomeo de Felice, an Italian-Swiss scientist.
Subdivision The genus Felicia is divided into six groups called sections. These are Anhebecarpaea, Dracontium, Felicia, Lignofelicia, Longistylis and Neodetris. F. douglasii has not yet been assigned to any section.
The following taxa are assigned to the respective sections: Anhebecarpaea: F. echinata (type), F. nordenstamii, F. westae. Dracontium: F. rosulata (type), F. wrightii, F. quinquenervia, F. monocephala. Felicia: F. tenella (type), F. abyssinica, F. australis, F. bechuanica, F. cana, F. caespitosa, F. clavipilosa, F. comptonii, F. drakensbergensis, F. dentata, F. deserti, F. erigeroides, F. fascicularis, F. ferulacea, F. hirta, F. hirsuta, F. hyssopifolia, F. linearis, F. microcephala, F. microsperma, F. mossamedensis, F. muricata, F. nigrescens, F. ovata, F. serrata, F. uliginosa, F. venusta, F. zeyheri. Lignofelicia: F. fruticosa (type), F. brevifolia, F. burkei, F. canaliculata, F. dregei, F. esterhuyseniae, F. filifolia, F. gunillae, F. lasiocarpa, F. macrorrhiza, F. odorata, F. oleosa, F. rogersii, F. scabrida, F. whitehillensis. Longistylus: F. welwitschii (type), F. alba, F. anthemidodes, F. boehmii, F. grantii, F. smaragdina. Neodetris: F. amelloides (type), F. aculeata, F. aethiopica, F. amoena, F. annectens, F. bellidioides, F. bergeriana, F. cymbalariae, F. cymbalarioides, F. denticulata, F. diffusa, F. dubia, F. ebracteata, F. elongata, F. flanaganii, F. flava, F. heterophylla, F. hispida, F. josephinae, F. joubertinae, F. linifolia, F. martinsiana, F. merxmuelleri, F. minima, F. namaquana, F. petiolata, F. puberula, F. stenophylla, F. tenera, F. tsitsikame.
Phylogeny According to recent DNA-analysis, Felicia belongs to the tribe Asterinae. It is probably polyphyletic.
Reassigned species The species that were originally described as, or moved to Felicia or one of its synonyms, but have since been reassigned, include the following:
Agathaea leiocarpa = Mairia sp. Elphegea reflexa = Polyarrhena reflexa Aster caffrorum (= F. caffrorum) = Microglossa caffrorum F. cameronii = Helicia sp. F. ciliaris = Gymnostephium ciliare F. ericifolia = Macowania ericifolia F. ficoidea = to be assigned to the Heliantheae F. kraussii = Gymnostephium ciliare F. imbricata = Polyarrhena imbricata F. lasiocarpa = Zyrphelis lasiocarpa F. maritima = to be assigned to the Heliantheae F. monticola = Zyrphelis monticola F. pinnatifida = Aster erucifolius F. reflexa = Polyarrhena reflexa F. retorta = Senecio retortus F. rigidula = Chrysocoma rigidula Fresenia pinnatilobata = Pegolettia pinnatilobata Fresenia foliosa = Pegolettia retrofracta
Description The base number of chromosomes of the earliest common ancestor of Felicia is probably nine, with most species diploid (2n=18), but some taxa in the section Felicia are tetraploid (4n=36), such as F. clavipilosa subsp. transvaalensis, F. fascicularis, and F. uliginosa. Divergent base chromosome numbers occur in the derived section Neodetris, with eight (2n=16) in F. amoena and F. elongata, six (2n=12) in F. minima and five (2n=10) in F. heterophylla, F. merxmuelleri and F. namaquana.
About 60% of the species currently assigned to Felicia are shrubs, high, very rarely up to , most of which belong to the sections Lignofelicia, Anhebecarpaea and Felicia. 20% are herbaceous perennials, which are often in the sections Dracontium and Neodetris. Another 20% consists of annuals, many of which are assigned to the section Neodetris, while Longistylis is entirely made up of annual species. All the woody, and most herbaceous, species are well-branched and bear many flowerheads, while the section Dracontium is rich in species, with a leaf rosette and stems topped by a single head. Stressed specimens of annual Felicia species can sometimes have only one flower head per plant. Two related species in the section Neodetris, growing on vertical rock faces, have a pendulous habit: F. petiolata and F. flanaganii.
Vegetative reproduction is rare, but rhizomes occur in F. tenella, F. wrightii, F. amoena and more or less in F. uliginosa. A special type of vegetative reproduction can be found in F. fascicularis that has branches, which produce roots when in contact with soil. Several species develop short and long shoots, the combination of which creates a particular, well-recognisable habit in many species belonging to the sections Lignofelicia and Felicia. All types of unbranched hairs may be found within the genus, including glandular hairs. Felicia has resin ducts.
Leaves Leaves are mostly alternately set along the stem, but opposite leaves do also occur, most prevalent in the section Neodetris, while in the section Longistylus, many species have opposite leaves in lower parts of the stem and alternately set leaves nearer to the top. Species in the sections Lignofelicia, Anhebecarpaea and Dracontium never have opposite leaves. Leaves are not compounded, and mostly have an entire margin, although in all sections but Anhebecarpaea, some species have leaves with up to moderate size teeth. Further incised leaves are restricted to F. brevifolia and its close relatives in section Lignofelicia. Leaves are generally seated, and their foot may be widened, particularly when these are succulent. Leaves in other species assigned to Lignofelicia may be digit-shaped, not having a recognisable margin. Species with stalked leaves are rare and restricted to the sections Felicia and Neodetris. Each leaf has between one and five primary veins, a number that is directly linked with the width of the leaf blade, and varies on the same plant. Leaves vary in consistency between species from soft to leathery. Resin canals are common in leaves of species of the sections Lignofelicia, Anhebecarpaea and Felicia, but do not occur in the other sections.
Most species have leaves with at least some hairs. Only F. dentata subsp. nubica, F. filifolia, F. fruticosa and F. whitehillensis, and some forms of F. muricata, are fully glabrous. On the other end of the spectrum, species in the section Neodetris are particularly hairy. Margins are often more hairy than the rest of the surface, which generally has a cartilageous seam or carries fine stiff bristle-like teeth.
Flowerhead