The great Swiss botanist Augustin Pyramus de Candolle (1778–1841) described several species of Anemone with ovoid hairy carpels, having an involucre and single flowers with 5–15 sepals, as his sect. Anemonanthea DC., now regarded as sect. Anemone (Candolle 1818). He subdivided the section into an unnamed group with sessile involucres and tuberous roots, in which he included A. coronaria, A. pusilla DC. non Gaertn., A. palmata, A. pavonina and A. stellata along with another five globally distributed species (Candolle 1818). Candolle included A. hortensis incorrectly as a synonym of A. stellata, which he stated had 10–12 oblong, more or less obtuse sepals. He described his concept of A. pavonina as having 10–12 acute, lanceolate sepals, adding that he had rarely seen single-flowered plants, mostly those in gardens with double flowers (see comments on A. pavonina under Lamarck below).

Candolle continued with his treatment of Anemone sect. Anemonanthea in the first volume of his Prodromus (Candolle 1824). Under his unnamed grouping of species with sessile involucres and tuberous roots, he maintained A. coronaria and A. pusilla as he had previously (Candolle 1824). He added two new varieties: A. palmata var. florepleno DC. and A. pavonina var. fulgens DC. (see full discussion on this below). Candolle once again also included A. hortensis as a synonym of A. stellata and added the significant statement pertaining to this (translated from Latin): “It is more rarely cultivated in gardens and is more often provided with a single flower than the Anemone pavonina, with which it has long been confused” (Candolle 1824).

The German botanist and physician Kurt Polycarp Joachim Sprengel (1766–1833), in his treatment of the genus Anemone a year later, divided it into four unranked subdivisions (Sprengel 1825). His second division, Anemone [unranked] Stephanomata Spreng., was described as having divided leaves, solitary flowers and villose fruits (Sprengel 1825). He further subdivided this unranked group into those plants with tuberous roots (Group 1), as opposed to those with roots that were either woody (Group 2) or fibrous (Group 3). In the former group he included A. coronaria, A. formosa Clark (see synonymy below), A. palmata, A. pavonina and A. stellata and another six species from various parts of the world (Sprengel 1825). He followed Candolle by incorrectly including A. hortensis as a synonym of A. stellata and distinguished A. pavonina from A. stellata and A. formosa by having lanceolate, acute sepals, as opposed to sepals with apices somewhat blunt in A. stellata and sepals broadly ovate in A. formosa (Sprengel 1825).

In his Anemonarum revisio, the German botanist Georg August Pritzel (1815–1874) included A. coronaria, A. formosa, A. fulgens, A. hortensis and A. palmata in an unnamed subsection characterised by woolly fruits, tuberous roots and sessile involucral leaves, within Anemone sect. Anemonanthea DC. (Pritzel 1841). His description of A. fulgens included sepals 8–15, oblong with obtuse apices and included a var. β in polynomial form, comprising numerous extremely narrow sepals with most acute apices, and cited for that variety A. pavonina Lam. in synonymy (Pritzel 1841). Pritzel described A. hortensis as having sepals 8–14, more or less broadly oblong-lanceolate and with somewhat obtuse apices. He also included A. stellata Lam. in synonymy (Pritzel 1841). Pritzel referred to Sprengel (1825) for his description of A. formosa as having larger, broadly ovate sepals (Pritzel 1841).

Oskar Eberhard Ulbrich (1879–1952), in his revision of Anemone, placed what he considered to be Anemone pavonina next to the Spanish species A. palmata in Anemone sect. Eriocephalus Ulbr. subsect. Longistylae Ulbr. series Oriba (Adans.) Ulbr. (Ulbrich 1905). He also included A. hortensis and A. coronaria in series Oriba and considered A. fulgens to be synonymous with A. pavonina (Ulbrich 1905: 202, 247). He did not provide any diagnostic characters in order to distinguish A. pavonina from its closest relatives, merely stating that it was a very variable species that occurs from southern France to Greece and Turkey (Ulbrich 1905). Ulbrich regarded A. pavonina var. regina (Risso) Rouy et Fouc. (albeit an invalid name; see discussion on Rouy and Foucaud under Synonymy below) as being found only in the western part of Europe, limited to southern France, northern Italy and Corsica, but again provided no morphologically diagnostic evidence for this statement. He likewise referred A. pavonina var. purpureo-violacea (Boiss.) Halacsy to eastern Europe, with a distribution from Thrace and Macedonia through Greece, including the islands of Melos, Chios and Kos, but again without providing any distinguishable characters for doing so (Ulbrich 1905).

The Italian botanist Gina Luzzatto (1904–c. 1987) undertook a very comprehensive morphological examination of A. fulgens, A. hortensis, A. pavonina and A. stellata, which included several infraspecific names. Luzzatto surveyed the literature associated with the names; however, while examining many collections in herbaria and living material from the various territories where the taxa occur, she inferred relationships based on her own observations and from the descriptions of earlier authors without reference to type material (Luzzatto 1932a, b, 1933, 1934a, b). In a field study of the area around Nice, where Luzzatto examined material of what she referred to as Anemone hortensis L. var. pavonina Lam., near Levens, she stated that the plants agreed with Lamarck’s description “in cui il fiore è spesso doppio, con un gran numero di sepali lanceolato-lineari acuti, d’un rosso scarlatto, salvo gli esterni che sono verdi” [in which the flower is often double, with a large number of acute lanceolate-linear sepals, of a scarlet red, except for the external ones which are green] (Luzzatto 1932b). Luzzatto maintained her concept of Lamarck’s double-flowered plants in her later treatment, in which she described A. pavonina as having (translated from Italian) “double flowers with many oblong narrow pointed sepals” (Luzzatto 1934a).

The horticulturalist Edward Augustus Bowles (1865–1954) wrote a historical assessment of what he considered to be A. hortensis and A. pavonina but did not mention Lamarck’s type material (Bowles 1948). A bibliographical summary in Bowles’s account by the botanist William Thomas Stearn (1911–2001), who was then librarian at the Royal Horticultural Society, did, however, include the statement: “the type of A. pavonina is the monstrous double form listed below as var. pavonina” (Stearn in Bowles 1948). It is in that paper where the first mention of Candolle’s A. pavonina var. fulgens is considered to be of hybrid origin as A. × fulgens. Bowles asserted that this was between what was stated to be the native A. hortensis and the introduced A. pavonina, which he concluded must have been introduced from Greece, Macedonia and western Asia (Bowles 1948).

A more recent revision of the genus Anemone based on examination of comparative morphology (Ziman et al. 2008) concluded that Anemone subgen. Anemone sect. Anemone subsect. Anemone comprised the species A. coronaria, A. hortensis and A. palmata, while subsect. Somaliense Ziman, Bulakh and Kadota comprised the single species A. somaliensis Hepper (Ziman et al. 2008). The three species in subsect. Anemone are distinguished from A. somaliensis by the basal parts of the leaf petioles having an expanded sheath and deciduous tepals, vs no basally inflated leaf sheath and persistent tepals (Ziman et al. 2008). Moreover, A. somaliensis differs from A. hortensis by having much larger involucral leaves (similar to the basal leaves) and a smaller perianth (Ehrendorfer et al. 2009). Neither A. fulgens nor A. pavonina was mentioned in the treatment by Ziman et al. (2008).

Maïa and Venard (1976) undertook cytotaxonomic and phytochemical studies on A. coronaria, A. hortensis, A. palmata, A. pavonina (from southern France) and A. variata Jord. (see synonymy below). These authors followed the assumption, instigated by Bowles (1948), that A. hortensis is restricted in its origin to southwest Europe, occurring in Albania, Italy and France, whereas A. pavonina occurs in Greece and southwest Asia (Maïa and Venard 1976). Their morphological diagnosis of “A. hortensis (= A. stellata Lamarck)” was that it had 12 to 18 sepals, whereas A. pavonina (including the statement “with four varieties: typica, ocellata, purpureo-violacea and pavonina”) had nine sepals. They also followed Bowles’s classification, stating that A. × fulgens was a hybrid between A. hortensis and A. pavonina, adding that A. pavonina var. ocellata, with large red tepals exhibiting a basal zone of white, had been introduced into southern France from Greece (Maïa and Venard 1976).

Two samples of what they considered to be A. pavonina showed different chromosome lengths. One collection (sample a) was from a population of A. pavonina var. ocellata gathered near the village of Valbonne near Grasse in Provence, and the other (sample b) was from a mixed horticultural strain cultivated in the Station d’Amélioration des Plantes at Fréjus, Var, also in Provence. This difference in chromosome lengths they regarded as intraspecific variation, i.e. variation within the same species (Maïa and Venard 1976).

Their isolation of the floral pigments in A. hortensis, A. pavonina var. ocellata and A. pavonina var. purpureo-violacea revealed that all three samples they examined contained the pigments kaempferol (yellow) and quercitol (off-white), and that both their samples of A. hortensis and A. pavonina var. purpureo-violacea contained cyanidol (or cyanidin, blue or red), which was lacking in A. pavonina var. ocellata, where it was substituted by pelargonidol (or pelargonidin, orange, red or red-blue). The floral pigments, however, in A. coronaria included all five pigments (Maïa and Venard 1976).

Maïa and Venard concluded that the chromosome morphology of A. palmata was very different from the group comprising A. coronaria, A. hortensis and the two samples of A. pavonina. They stated that the chromosomes from A. hortensis and sample (a) of A. pavonina var. ocellata did not differ from each other and that the chromosome lengths of A. hortensis and both samples of A. pavonina also coincided in length. They also ascertained that within the second group, the chromosome morphology of A. coronaria was quite distinct from the other three samples (Maïa and Venard 1976).

Hoot et al. (1994) undertook phylogenetic analyses based on combined data from cpDNA restriction sites, ribosomal DNA restriction sites and morphology but did not include data for A. palmata. Their results confirmed the sister relationship between A. coronaria with pantoporate pollen and the spiraperturate pollen of A. fulgens sensu Hoot et al., A. hortensis and A. pavonina sensu Hoot et al. (Hoot et al. 1994). Their results, obtained from the phylogenetic analyses of restriction site data, showed that a clade with A. coronaria on a branch with 13 sites was sister to a polytomy comprising A. hortensis, A. fulgens and A. pavonina, each with zero restriction site variation (Hoot et al. 1994). The phylogenetic tree from the combined analysis of morphological data, cpDNA and restriction site variation revealed the same topology for the four sampled taxa, sister to a clade with four species occurring in the southwest USA. The clade with A. coronaria sister to a branch with A. hortensis, A. fulgens and A. pavonina was referred to as part of the “Coronaria Group” within Anemone sect. Anemone (Hoot et al. 1994).

Ehrendorfer and Samuel (2001) sequenced the plastid atpB/rbcL intergenic spacers of 21 taxa of Anemone, obtaining a single most parsimonious tree. They included samples of A. coronaria, A. hortensis, A. palmata and a Greek sample from Olympia, Peloponnese, of their understanding of A. pavonina (Ehrendorfer and Samuel 2001). Their results showed moderate support (68% bootstrap) for what they termed the “Coronaria” clade, comprising A. coronaria and two samples of A. palmata sister to A. hortensis and A. pavonina on a branch with 85% bootstrap support (Ehrendorfer and Samuel 2001). In their discussion they concluded that “A. hortensis and A. pavonina are close and possibly only elements of one polymorphic Mediterranean species, whereas the frequently sympatric A. coronaria appears well separated” (Ehrendorfer and Samuel 2001).

The results from the study by Ehrendorfer and Samuel (2001) were corroborated by results from cytogenetic data by Mlinarec et al. (2006) from Adriatic populations of A. hortensis, including samples of what they referred to as A. pavonina. Their results revealed that A. coronaria, A. hortensis and A. pavonina all had three acrocentric and five sub-metacentric chromosomes and all had similar heterochromatin banding patterns (Mlinarec et al. 2006). They concurred with the results from Hoot et al. (1994) and Ehrendorfer and Samuel (2001), stating that “A. hortensis and A. pavonina are the most closely related species within the Coronaria group” (Mlinarec et al. 2006).

The revised molecular and morphological study undertaken by Ehrendorfer et al. (2009) has enormously clarified the species relationships and ascertained the wild distribution of the taxa in Anemone sect. Anemone. It has also gone a long way towards resolving the extremely muddled taxonomic history of both A. coronaria and A. hortensis (Ehrendorfer et al. 2009). The classification in the study was based on sequence data from the cpDNA intergenic region atpB-rbcL previously undertaken by Ehrendorfer and Samuel (2001). This study has broadly expanded on the phylogenetic analyses of Hoot et al. (1994); however, these authors admitted that the variation within A. hortensis had not been sufficiently studied. They treated material classified as A. fulgens, A. pavonina and A. stellata, and other infraspecific named taxa, as variations within a polymorphic A. hortensis (Ehrendorfer et al. 2009).

The topologies for these taxa in the studies by Hoot et al. (1994) and Ehrendorfer and Samuel (2001) are broadly congruent with the results from analyses of chloroplast atpB-rbcL spacer and nuclear ITS regions undertaken on 55 species of Anemone by Hoot et al. (2012). The results by Hoot et al. (2012) indicated a Bayesian probability value (BI) of 1 and 82% maximum parsimony bootstrap support for the clade comprising A. coronaria, A. somaliensis, A. hortensis and A. pavonina (Hoot et al. 2012). One branch on this clade comprised A. coronaria and A. somaliensis (BI 1; 78% bootstrap), sister to a branch with A. hortensis and A. pavonina (BI 1; 100% bootstrap). The study by Hoot et al. (2012) recognised A. coronaria, A. hortensis, A. palmata, A. pavonina and A. somaliensis as forming Anemone subgen. Anemone sect. Anemone subsect. Anemone ser. Anemone (Hoot et al. 2012). These authors stated that although A. hortensis and their understanding of A. pavonina sampled from Greece are very well supported as sisters to each other, the branch lengths indicated a number of unique substitutions, suggesting that species status might be correct for A. pavonina, but they did not include any indications as to potential differences in morphology (Hoot et al. 2012).

Examination of relict satellite DNA sequences of the AhTR1 group in 22 species from four sections of Anemone also showed that A. hortensis from the island of Hvar, Croatia, and their sampling of A. pavonina from Bogdanci, Paljurci, North Macedonia, shared a very similar species-specific satDNA profile. Their results support the treatment by Ehrendorfer et al. (2009) that A. hortensis and A. pavonina are a single species (Besendorfer and Mlinarec 2013).

The topologies from all previous molecular analyses are congruent with results from sequence data from the nuclear 5S rDNA gene and spacer region (Mlinarec et al. 2016). The latter’s research showed that four samples of A. hortensis from Croatia and four samples of their understanding of A. pavonina from North Macedonia formed a clade with Bayesian inference of 1 but were unresolved with respect to a strongly supported clade of three samples of A. palmata, also with BI of 1, and a clade of eight samples of A. coronaria, also with BI of 1. These taxa formed a discrete clade with moderate support of 0.62 BI, which was referred to incorrectly as “sect. Coronaria” (Mlinarec et al. 2016). Branch lengths on the phylogram for the samples on each of the three branches on the clade representing A. hortensis and A. pavonina, A. palmata and A. coronaria all showed differing lengths. The lengths shown by the three samples of A. pavonina and four samples of A. hortensis, albeit well supported, all showed greater differences in their branch lengths than did the eight samples of A. coronaria (Mlinarec et al. 2016). Branch lengths between the three samples of A. pavonina and four of A. hortensis were, however, comparable to the differences in branch lengths of most other species of Anemone in that study.

The most recent phylogenetic study using data from cpDNA (atpB-rbcL, trnL-F, rbcL) and nrDNA ITS sequences on 159 individuals representing 146 species of Anemone was conducted using maximum likelihood (ML) and Bayesian inference (BI) (Xiang et al. 2024). Their results also concurred with the topologies of those from previous molecular analyses. A large clade of 50 species was fully supported (100% bootstrap, BI 1), which was referred to as sect. Anemone (Xiang et al. 2024). Within that large clade, A. coronaria, A. hortensis, A. palmata, A. pavonina and A. somaliensis all formed a moderately supported clade (58% bootstrap; BI 1) comprising two subclades: A. palmata, A. hortensis and A. pavonina sister to A. coronaria and A. somaliensis (Xiang et al. 2024). The subclade with A. palmata sister to A. hortensis and A. pavonina was also moderately supported (67% bootstrap, BI 1), while the branch with A. hortensis and A. pavonina was well supported (83% bootstrap, BI 1).

Linnaeus validated A. hortensis in his “Species plantarum” with the following protologue:

Anemone hortensis L. Sp. pl. 1: 540 (1753)

9. Anemone foliis digitatis

Pulsatilla foliis digitatis Hort. cliff. 224.

Anemone geranii rotundo folio, purpurascens. Bauh. pin. 173.

Anemone hortensis latifolia 3. Clus. hist. 1. p. 249.

Anemone 1 Dod. pempt. 434. (in fact 430/431)

Habitat ad Rhenum & in Italia ♃ (= perennial sign)

Due to their apparent morphological similarity, Linnaeus was initially uncertain as to how he could distinguish between Anemone coronaria and A. hortensis. He described both species under the group heading of “Anemones caule folioso, seminibus caudatis” [Anemones with stems bearing leaves and seeds with tails] (Linnaeus 1753). His validating description of A. coronaria was “7. Anemone foliis radicalibus ternato-decompositis, involucro folioso” [Anemone with much-divided ternate basal leaves, involucre leaf-like]. He also included a variety of A. coronaria with red flowers and multiple tepals, thereby recognising that A. coronaria with its narrow leaves also exhibited doubling of the floral parts “β Anemone tenuifolia multiplex rubra”. He stated that the origin for both varieties of A. coronaria was from “Oriente, Constantinople allata”, i.e. brought to me from Mediterranean eastern Europe and Constantinople (Linnaeus 1753).

Prior to the publication of his protologue, Linnaeus had alluded to the variation in colour and the number of tepals of A. hortensis “variat florum plenitudine et colore” on what he had then called “Pulsatilla foliis digitatis” (Linnaeus 1738 see Reference 1 below). His initial distinction between A. coronaria and A. hortensis was, however, largely based on leaf dissection – the former had much more dissected ternate leaves, whereas the leaves of A. hortensis were digitate. He indicated that A. hortensis was to be found in Germany along the river Rhine and in Italy (see References 1 and 3 below).

Linnaeus maintained his brief description for A. hortensis in the second volume of the tenth edition of “Systema naturae” (Linnaeus 1759). In that work he did not provide any additional information for the species, maintaining the simple phrase which he had used in his protologue, “9. A. foliis digitatis [Anemone with digitate leaves]” (Linnaeus 1759). The description for A. hortensis was repeated verbatim in “Species plantarum” ed. 2, but for A. coronaria, which again included his β variety, he added another variety, “γ Anemone angustifolia multiplex, mutata florum facie quotannis nova” [Anemone with multiple narrow leaves and flowers which alter their appearance every year] (Linnaeus 1762).

In the “Systema naturae” ed. 12 vol. 2, the entry for A. coronaria is the same as that in “Systema naturae” ed. 10, but for Anemone hortensis, it is enlarged to “9. A. foliis digitatis. Anemone fol. digitatis: lobis incisis, petalis lanceolatis numerosis. Ger. prov. 380 [Anemone with digitate leaves with incised lobes, petals lanceolate, numerous]” (Linnaeus 1767). The reference to “Ger. prov.” is to the French physician Louis Gérard (1733–1819), who wrote “Flora gallo-provincialis” (Gérard 1761). Gérard’s description of the anemone follows below.

The description of A. hortensis in the 12^th edition is repeated verbatim in “Systema naturae” ed. “13” (Linnaeus 1770) and again in the “Systema vegetabilium”, with the addition only of “seminibus lanatis” [woolly seeds] (Linnaeus 1774).

Linnaeus’s most comprehensive description of Anemone hortensis, however, was in his Mantissa altera 2, which was in effect an appendix to the 12^th edition of “Systema naturae”: “Anemone hortensis. Involucrum foliolis lanceolatis. Petala 9, lineata, extus pubescentia” [Anemone hortensis. Involucre of lanceolate leaflets. With 9 petals, each with lines and pubescent on the outside] (Linnaeus 1771).

Louis Gérard, who was cited in “Systema naturae” ed. 12, used the phrase name “5. Anemone foliis digitatis, lobis incisis, petalis lanceolatis numerosis” (Gérard 1761). Gérard added the following synonyms:

Anemone foliis digitatis. Linn. Spec. 540.

Anemone geranii rotundo folio, purpurascens. C. B. Pin. 173. Tour. 176.

Anemone italica, latiusculis spinosis foliis, tertia clusii. J. B. 3. p. 402

Anemone 1. Dod. pempt. 434. Dalech. p. 845.

Anemone 2. Camer. epit.

“Provenit in sterilibus abbatiae” du Thoronet. Perennis.

Gérard described this species from waste ground around the Abbey of Thoronet, c. 20 km from his home in Cotignac, Provence, and referred to Linnaeus’s protologue but without mentioning the name A. hortensis. Gérard referred to an illustration in Daléchamps (1587). This illustration was the same, reversed, as that already used in Clusius (1576) and in L’Obel (1576, 1581) and again in Dodoens (1583). Gérard included a reference to Tournefort (1700), who, in turn, cited Caspar Bauhin (C. Bauhin 1623). Gérard also referred to an engraving in Camerarius (1586). All the illustrations cited by Gérard depict A. hortensis L.

Jean-Baptiste-Pierre-Antoine de Monet, chevalier de Lamarck (1744–1829), described Anemone pavonina in his “Encyclopédie méthodique”. Botanique 1(1): 166 (1783). His protologue was:

11. Anémone oeil de Paon, Anemone pavonina.

Anemone foliis radicalibus profunde tripartitis, lobis cuneatis, incisis, dentatis; flore variegato [Anemone with basal leaves deeply divided into three parts, lobes wedge-shaped, incised and toothed, flowers variegated].

Anemone latifolia, pavo dicta major [broad-leaved anemone, called the greater Pavo]. Bauh. Pin. 176. No. 4, 5, 6. Anemone latifolia maxima versicolour [Large broad-leaved anemone with variously coloured flowers]. Bauh. Pin. 176. Tournef. 276.

A full description followed (translation from French): “This species, which has been cultivated for several years in the Jardin du Roi, produces flowers of a very pleasant appearance, of a form quite different from that of the Anémone des Fleuristes [i.e. his number 9. A. coronaria], and which blooms from the beginning of April. Its root is thick, tuberous, furnished with lateral fibres, and bears leaves which closely resemble those of the Sanicle officinale [Sanicula europaea L.]. These leaves are petiolate, deeply divided into three widened, wedge-shaped lobes, unequally incised, and ending in coarse teeth whose tips point in different directions. The stem is seven or eight inches high, a little hairy, and provided at two-thirds of its height with a small collar of three smallish leaves, two of which are very often simple, and the third a little divided. At the top of this stem is born a flower coloured with red and white, almost an inch and a half wide, composed of many oblong, slightly narrow, pointed petals, of which the interior ones are the smallest. These petals are veined longitudinally, slightly hairy on their reverses, whitish at their base, and a beautiful red towards their apex, and they have this remarkable feature that the exteriors are scarcely coloured, sometimes even entirely green, so that they appear to form a calyx contiguous to the corolla, as in the preceding species [i.e. his number 10. A. palmata]. It is evident, however, that these two plants have no other natural calyx except the small collar itself, which they carry a little below their flower: this part corresponds entirely to the small calyx of the hepatic Anémone [A. hepatica L. = Hepatica nobilis Schreb.], which is also a little distant from the corolla, a characteristic common to all the species of this genus.

The species I have just mentioned is, I believe, native to the Levant; and although I have only seen it with double flowers, there is no doubt that the natural plant which is typical of this species is very different from that which constitutes the Anémone des Fleuristes, n°. 9. It is cultivated in the gardens of the Curious: there it produces agreeable varieties (v. v.)”.

Lamarck initially recognised the existence of Anemone hortensis in the third volume of his “Flore Françoise” (Lamarck 1779), citing Linnaeus’s A. hortensis from the second edition of “Species plantarum” (Linnaeus 1762). Lamarck diagnosed his species number 18, Anémone des jardins, Anemone hortensis, with the sentence “Plus de sept petales, semences simplement laineuses [more than seven petals, seeds with simple woolly hairs]. He made no reference in this work to A. coronaria. He described the three-lobed digitate leaves, the collarette or involucre comprising three sessile, more or less incised leaflets, and the single flower composed of nine lightly purplish, narrow and veined petals which are a little velvety on their underside. He added that it grew in waste ground in Provence and that there were bright-coloured varieties of it grown in gardens (Lamarck 1779). Lamarck’s only other reference was Tournefort’s Anemone geranii rotundo folio, purpurascens, already cited above under Gérard (Tournefort 1700).

Four years later, in the first volume of his great work, the “Encyclopédie methodique. Botanique,” Lamarck placed both Anemone coronaria and his two new species, A. pavonina Lam. and A. stellata Lam., in the group of species he identified as having “semences chargees de duvet, mais non munies de longues queues plumeuses” [seeds covered with down but not provided with long feathery tails] (Lamarck 1783: 165). He included under A. coronaria two synonyms without additional descriptions: (a) Anemone hortensis, tenuifolia and (β) Anemone hortensis, latifolia. Following his description of A. coronaria, he also described two new species: Anemone pavonina and A. stellata (Lamarck 1783: 166).

Lamarck named the species the ‘eye of Paon’. According to Clusius, this was in reference to the orbs or eyes seen at the ends of the peacock’s tail feathers and was probably an allusion to the presence of a circle of gold or white formed from the pale bases of each of the tepals on some plants of the species. These paler zones surrounding the darker centre of the flowers of “Anemone hortensis latifolia pavo major 1” act as the eye’s pupil (Clusius 1601: 261). In Clusius’s own description of the flowers of “Anem. hort. latifol. pleno flore 1”, he stated (translated from Latin): “then emerging two inches above the aforementioned leaves [involucre], it supported a large and widely spread flower, consisting of numerous leaves [tepals], the outer and larger ten or twelve of which were mostly green in colour, their tips lightly and linearly stained with a scarlet colour, while the inner ones, smaller and narrower, shone with a more diluted red-purple, and those covering the middle flower were also of the same colour, reflexed to the navel, and as if clustered” (Clusius 1601: 261).

The Latin generic name Pavo L. is the name for peafowl (Linnaeus 1758). The famous “eyes” found on the tail feathers of the male peacock probably appertain to the legend of Zeus and his flirtation with Io, a mortal lover of Zeus, who was changed by him into a heifer in order to be concealed from his wife, Hera. Suspicious Hera consequently sent the all-seeing beast Argos Panoptes (or Argus) with his multitude of eyes to keep watch over Io. Gaia, goddess of the earth, had sympathy for Io and provided the red, purple and white flowers of the violet (Io’s flower) to the heifer as food. Zeus, angry with Hera for sending Argos to watch over Io, then sent Hermes to kill Argos, an act which inevitably enraged Hera, who consequently sent a stinging insect to pursue the escaped Io over land and sea (Ionian Sea). Io crossed the Bosporus (Oxen crossing) and was later transformed by Zeus back into her original mortal form, while Hera preserved Argos’s eyes for eternity locked inside the tail-feathers of peacocks.

Lamarck’s distinctions between A. coronaria and his A. pavonina correctly referred to the more finely divided and incised leaves of A. coronaria as opposed to the three broader wedge-shaped and lobed leaves of his A. pavonina having placed Linnaeus’s A. hortensis as a variety of A. coronaria as var. (β) Anemone hortensis, latifolia Lamarck 1783: 166). In addition, he alluded to the flowers of both the single and the double or multi-tepalled flowers of A. coronaria, a phenomenon which occurs in both species. In the second tome of Lamarck’s illustrated work “Tableau encyclopédique et méthodique” are illustrations by L. Fossier of both the single- and double-flowered forms of A. coronaria (Lamarck 1794: t. 496 figs a and b). It is of note that Lamarck did not include any illustrations of A. hortensis in that work. Nor was A. hortensis included by Poiret in the first part of the Supplement to the “Encyclopédie méthodique. Botanique,” even though the illustration of A. coronaria was specifically referred to (Poiret 1810). It is worthwhile noting that Poiret did not add the figure legends to Fossier’s illustrations of planche 496 in Lamarck’s original work until the publication of tome 3 of “Tableau encyclopédique et méthodique” (Poiret 1823).

Lamarck compared the rose, white, red, yellow, violet or blue coloured oval-oblong tepals of A. coronaria to the flowers of A. pavoninа, which he stated comprised many narrow tepals which are red with white at the base and which get smaller towards the centre of the flower. He indicated that he had seen living plants (v. v. = vidi vivam) and, significantly, that he had only seen plants with double flowers (Lamarck 1783).

The protologue of Anemone stellata Lam. (Lamarck 1783: 166) reads:

12. Anemone en étoile, Anemone stellata. Anemone foliis radicalibus tripartitis, lobis variè incisis, subtus venosis, petalis linearibus stellatim dispositis [Anemone with basal leaves divided into three parts, with lobes more or less incised, veined below, petals linear, arranged in a star-like pattern]. Anemone geranii rotundo folio, purpurascens [Anemone with rounded leaves like a Geranium L., slightly purplish] Bauh. Pin. 173. Tournef. 276. Anemone hortensis latifolia 3. Clus. Hist. 1. p. 249. Anemone 1. Dod. Pempt. 434. Anemone Hall. Helv. No. 1152. Anemone hortensis Lin. “Excluso Casp. Bauhini synonymo primo”.

Translation from French: “Although this species is very pretty, it is rarely cultivated in gardens; the preceding one [A. pavonina] is seen more often. This one has petals that are not as narrow and makes up a better-appearing flower, with more brilliance. Its root is tuberous, knotty, furnished with fibres, and has a slender, light and single-flowered stem. The basal leaves are borne on very long petioles, composed of three wedge-shaped leaflets, incised more or less deeply, veined and deflated, and provided at the end of their lobes with a small specific point. Some of these leaves are narrowly incised. The flower is terminal, either flesh-coloured, or red, or purplish, and composed of nine to fifteen narrow, linear petals, five to eight lines long [= 1.1–1.8 cm], coloured internally, whitish and a little hairy on their back, and which form a star shape by their arrangement. The collar is made up of three small narrow leaves, one of which is slightly cut. This plant grows in the stony and sterile places of Languedoc, Provence, Switzerland and Italy. It was communicated to me by Mons. Abbé Pourret. (v. s.). It flowers in March” (Lamarck 1783: 166).

Lamarck stated that he had received dried material of this taxon (v. s. = vidi siccam) from Pierre-André Pourret (1754–1818), who was born in Narbonne. After ecclesiastical studies, Pourret studied at the Jardin du Roi in Paris, which was then under the care of its intendant Georges-Louis Leclerc, Comte de Buffon (1707–1788). Thereafter, Pourret had maintained correspondence with the great naturalist scholars of his time, including Linnaeus, Jean-François Séguier in Nîmes, Philippe-Isidore Picot de Lapeyrouse in Toulouse and Carl Ludwig Willdenow in Berlin.

For the years 1787 and 1788, Pourret directed the natural history cabinet in Paris belonging to two brothers of the ancien régime, who both died in the Reign of Terror during the French Revolution (Bonnet 1916). The elder brother, Étienne Charles de Loménie de Brienne (1727–1794), was the Bishop of Condom and later Archbishop of Toulouse. His brother was Lieutenant-General Louis-Marie-Athanase de Loménie de Brienne (1730–1794).

In 1789, at the start of the French Revolution, Abbé Pourret left Narbonne and emigrated to Barcelona, where he was appointed director of the botanical garden and professor of natural history at the University of Barcelona. He became deputy director of the Botanical Garden of Madrid and then obtained a canonry at the Cathedral Church of Ourense in Galicia and canon-treasurer of the Metropolitan Church of Santiago de Compostela, where he gave lessons in botany. When he died in Santiago in 1818, he bequeathed his herbarium to the school of pharmacy in Santiago, which was then acquired by Complutense University in Madrid (MAF). There are no specimens of Anemone stellata among his collections in MAF (Paloma Cantó pers. comm.). There is, however, a specimen in Paris that was acquired from the “Collection de l’Abbé Pourret, extraite de l’Herbier légué de M. le Dr. Barbier 1847”. The surgeon Joseph-Athanase Barbier (1767–1846) had acquired the personal herbarium of the Abbé Pourret, which was initially conserved by Pourret at the château de Brienne for the brothers Loménie de Brienne. This was then moved to the cabinet of natural history in Paris (Bonnet 1916: 281, 283). The sheet, which has three flowering specimens, each with narrowly lanceolate tepals, as well as another specimen in fruit [P03181989], has two annotated labels on it and “Anemone decapetala L.” written on the sheet. The first label states, “Anemone stellata Lam. M. de Lamarck qui nous cite pour tenir cette plante de nous nous pavoir l’avoir malconnuè” [Mr de Lamarck who quotes us as having (received) this plant from us, (but) we must have misunderstood him]. The second has in Pourret’s hand “Anemone coronaria [crossed out] apennina [inserted instead] ex Seguiero et ex Gouano sola hortensis varietas cui synonymiam Tournefortii refert anemone geranii folio rotundo Tour. 276”.

When describing A. stellata Lamarck cited Linnaeus’s A. hortensis in his synonymy and moreover, did not refer to any specific material. The name Anemone stellata Lam. is therefore, not only illegitimate but it is also automatically typified by the type of A. hortensis (Art. 7.5). Haller’s description of Hal. 1152 which was also cited by Lamarck “Anemone seminibus lanatis, foliis radicalibus trilobis & multifidis, caulinis ovato lanceolatis” (Haller 1768) was described by Haller as having “petala novem” and “Flore miniato” and “purpureo flore, extus hirsuto”. It is clear he was describing variation within A. hortensis. There is a specimen collected by Haller conserved in G with the locality cited by Haller (1768) written on the label “Supra Moutru in dumetis” Hall. n. 1152. Anemone hortensis L.” [G00144586].

Lamarck had described his concept of A. hortensis in his earlier work (Lamarck 1779: 321) with the statement “La fleur est terminale, grande, lègérement purpurine, & composée de neuf pétales longs, étroits, marqués de quelques lignes & un peu velus en-dessous.” [The terminal flower is large, lightly purplish and composed of nine long, narrow petals marked with a few lines and the underside is a little velvety]. He went on to say that it grew in barren places of Provence and that some beautiful varieties are cultivated in gardens which have petals that are not as narrow [as the wild plant] and with much more vibrant colours (Lamarck 1779: 322).

In his protologue for the name Anemone pavonina var. fulgens Augustin Pyramus de Candolle (1778–1841) mentions “Gay ined.” (Candolle 1824: 18). This is a reference to Jacques Étienne Gay (1786–1864) a Swiss botanist from Prangins near Nyon c. 30 km from Geneva who worked extensively in Paris until 1848. Gay is known to have communicated with Candolle after the latter’s return there in 1816 and paid visits to Candolle in Geneva when visiting his family in Prangins (Candolle 1862: 411). There is, however, no extant correspondence specifically with respect to Anemone between Gay and Candolle in the archives in G, nor any extant specimens in G from Gay referring to this name (G. Barreira and P. Bungener pers. comm.). Candolle was uncertain of the status of this variety, describing it with a question mark and the statement “flores ampliores quam var a (v. s.)” [flowers larger than in var. a], the var. a being a reference to Lamarck’s A. pavonina i.e. a direct reference to plants with double flowers and the “v. s.” referring to the fact that he had only seen herbarium material (Candolle 1824: 18).

Candolle had already referred to the existence of both the single and double flowering states of this plant in his description of the double-flowered A. pavonina in an earlier work (Candolle 1818). In that he stated (translated from Latin): “It [i.e. A. pavonina] is rarely seen with a single flower; the variety with double-flowers is quite common in gardens under the names A. oeil de paon, Candiote, A. de Crète. Linnaeus referred to this in his synonymy, but in his herbarium, I found it among the varieties of A. coronaria; it is easily distinguished by the flower being surrounded by very acute sepals” (Candolle 1818: 198). Candolle included references to three collections “Hab. in vineis Vasconiae prope aquas-Tarbellicas (Dax) Thore; In Gallo-provincia circa Olbias (Ziz); Nicaeam Risso et verisimiliter in Oriente. ♃ fl. vere (v.s.sp. [vidi siccam spontaneum = I have seen it in a dried state from the wild] et v.c. [vidi cultam = I have seen it in cultivation])” (Candolle 1818).

The first collection mentioned by Candolle is to Jean Thore (1762–1823) a physician and botanist from Dax in the Landes. The locality cited by Candolle for Thore’s collection being vineyards near the Bronze Age hot spring settlement of the Tarbelli people, the foundation of the town of Dax. The second reference is to a collection by the German botanist Johann Baptist Ziz (1779–1829) from the archaeological site of Olbia, now known as Hyères, which sits on the coast between Marseille and Cannes in Provence. The third collection was by Joseph Antoine Risso (1777–1845) from near Nice.

Candolle in his later work described the species Anemone pavonina as having “sepalis 10 – 12 lanceolatis acutissimis” however, he also cited Morison’s illustration of a plant with multiple tepals (Morison 1680) and explicitly excluded two references “A. hortensis Thor. chl. land.: 238” and “A. pavonina Lois. not. 87” (Candolle 1824). Candolle then described his new var. fulgens with a question mark: “b var. fulgens foliis tripartitis, lobis cuneatis inciso-dentatis, involucralibus sessilibus oblongis integris subincisisve, sepalis oblanceolatis apice latioribus basi attentuatis [b var. fulgens with tripartite leaves, bases cuneate, lobes incised-dentate, involucral leaves sessile, oblong, entire or subincised, sepals oblanceolate, broader at the apex and attenuated at the base]. Candolle specifically included in var. fulgens the two references that he had excluded from A. pavonina above: “A. hortensis Thor. chl. land.: 238” and “A. pavonina Lois. not. 87” (Candolle 1824). He added v. v. et s. = vidi vivam et siccam “I have seen living and dried plants”.

The first reference is to Thore’s “Essai Chloris Landes” (Thore 1803) in which Thore states “Corol. d’un rouge très vif: poly-pétale.” The second is to Jean-Louis-Auguste Loiseleur-Deslongchamps (1774–1849) who included his concept of Anemone pavonina in “Notice sur les plantes à ajouter à la Flore de France” in which he describes the flowers as differing from Anemone hortensis by their larger size and that the corollas have 10 to 15 petals. He stated “Elle croît dans les vignes à St. Pandelon près de Dax, où elle a été trouvée par M. Thore qui me l’a communiquée; j’en ai aussi reçu des échantillons de M. Grateloup” (Loiseleur-Deslongchamps 1810). Loiseleur-Deslongchamps also mentioned that there are plants cultivated in gardens with double flowers (Loiseleur-Deslongchamps 1810). Both the descriptions by Thore and Loiseleur-Deslongchamps refer to plants with large, red flowers with 10 to 15 tepals and not to plants with multiple numbers of tepals.

Jean Charles Marie Grenier (1808–1875) and Dominique Alexandre Godron (1807–1880) together compiled and wrote the comprehensive “Flore de France” in three volumes from 1847 to 1856. In the first volume they included Anemone hortensis including three additional varieties (Grenier and Godron 1847). They defined A. hortensis with the description: “Calice à 10–12 sépales et plus, glabres extèrieurement, obovales ou lancéolés, plus ou moins aigus, ou enfin sublinéaires très-aigus” [Calyx with 10 – 12 sepals or more, externally glabrous, obovate or lanceolate, more or less acute or sublinear and very acute]. Their first variety var. a stellata they described as: “Sépales 8–10, lancéolés, obtus parfois apiculés [Sepals 8–10, lanceolate, obtuse or sometimes apiculate]. A. hortensis (de presque tous les auteurs). A. stellata Lam. Enc. 1. p. 166.” They stated the distribution for var. a was: “Grasse, Fréjus; Navarreins (Basses-Pyrénées); Dax”. Their var. β fulgens they described as: “Sépales 8–10, grands, obovales, en coin à la base, élargis au sommet obtus, parfois apiculés [Sepals 8–10, large, obovate, narrowed at the base, widened at the obtuse apex, sometimes apiculate]. A. hortensis Thore chl. land. 238; A pavonina Lois. gall. 1, p. 400; Rchb. l. c. f. 4650”. Distribution stated as: “across southern France and Corsica”. Their third var. γ pavonina was: “Sépales très-nombreux, lancéolés-linéaires, très aigus [Sepals very numerous, linear-lanceolate, apex very acute]. A. pavonina D C. fl. fr 5, p. 634; Dub. bot. 1 p. 5”. Distribution: “Dax, Saint-Sever; Grasse etc”.

Grenier and Godron when they published A. hortensis “α stellata”, were clearly distinguishing this variety from “the species”, i.e. from A. hortensis L. var. hortensis, indicated by their recognition of it as having a different number of sepals (Grenier and Godron 1847). Linnaeus in fact only alluded to numerous “petals” in most of his publications, mentioning “sepals nine” only once (Linnaeus 1771). Linnaeus’s type specimen however, consists of two flowering plants: one with ten tepals and the other with twelve. Grenier and Godron’s further statement “A. hortensis (de presque tous les auteurs [of almost all authors])” also indicated that they considered their var. stellata to be different from typical A. hortensis L. and their citation of “A. stellata Lam. Enc. 1, p. 166” can then be taken as explicitly excluding Lamarck’s synonym “Anemone hortensis. Lin.”, and thus the type of A. stellata. The name Anemone stellata Lam. being superfluous and illegitimate, is automatically typified by the type of A. hortensis (ICN Art. 7.5). Grenier and Godron’s A. hortensis var. stellata is based on a different type collected near Grasse as stated in the protologue. Original material was collected by Godron’s colleague Charles Joseph August de Baudot of Sarrebourg, and is designated as lectotype in this paper (Fig. 2A).

Grenier and Godron’s Anemone hortensis var. fulgens (DC.) Gren. and Godr. is a combination based on Candolle’s Anemone pavonina var. fulgens DC. (Candolle 1824). Although Candolle’s varietal name was not directly cited by Grenier and Godron (1847), it is nevertheless based on the same type according to their shared list of synonyms (see discussion and typification below).

A. hortensis var. pavonina (Lam.) Gren. and Godr. is also a new combination, although again it is only Candolle, and his contributing author Jean Etienne Duby, that were cited and not Lamarck. Both Candolle and Duby, however, included A. pavonina Lam. as a synonym in their own descriptions of A. pavonina, thereby permitting Grenier and Godron to validate the new combination (Candolle 1818, 1824; Duby 1828).

Theodor von Heldreich (1822–1902), director of the Natural History Museum and of Queen Amalia’s Royal Garden in Athens (now the National Garden) collected a small, white or very pale pink-flowered Anemone on the island of Crete in 1846. This was described three years later as a variety of Anemone stellata Lam. by the Swiss botanist Pierre Edmond Boissier (1810–1885) as A. stellata var. heldreichii Boiss. (Boissier 1849) It had to wait another 51 years until the Vienna based physician of Hungarian descent Eugen von Halácsy (1842–1913), also called Jenö Halácsy, who worked closely with Heldreich, correctly identified it as a variant of A. hortensis (Halácsy 1900). It was later recognised as a species in its own right Anemone heldreichiana by the French mycologist Michel Gandoger in 1916 until the Austrian botanist Karl Heinz Rechinger (1906–1998) demoted it to the rank of subspecies within A. hortensis in 1943, stating (translated from German) “As a geographical race it deserves the rank of a subspecies”.

It is a plant restricted to montane slopes of Crete and a few adjacent islands, exhibiting small ternate leaves, each of the three lobes much divided and incised and with pale midveins on the adaxial surface. The flowers are white occasionally pinkish, with narrow, elliptic tepals streaked with three blue veins. The anthers are also notably blue (Fig. 2D). This taxon, although known under the plethora of taxonomic ranks outlined here, has not been sampled adequately in molecular analyses to ascertain its correct taxonomic status.

The author has declared that no competing interests exist.

No ethical statement was reported.

No use of AI was reported.

No funding was reported.

The author solely contributed to this work.

All of the data that support the findings of this study are available in the main text.